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THE
KANSAS UNIVERSITY
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•DIVERSITY OF KANSAS SCIENCE BULLETIN
First Mailing of Volume XXIII — August 15, 1936
First Mailing OF Volume XXIV — February 16, 1938
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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
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DEVOTED TO
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UNIVERSITY OF KANSAS
Vol. XXIV
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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.
Dains et al.: Reactions of Thiazolidones VIII
43
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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.
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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,'<adenia atriplicifolin, Liatris aspera, Helianthus stnimosus, rig-
idus and Solidago canadensis gilvocanescens.
102 The University Science Bulletin
PLATE I
Photomicrographs of root cross sections. (X76)
1. SoHdago rigidiuscula.
2. Ambrosia trifida.
3. Actinomeris alternifolia.
4. Helianthus strumosus.
5. Silphium speciosiim (old root).
6. Mesadenia atriplicifoHa.
Mayberry: Hydrocarbon Secretions
PLATE I
103
2Q4 The University Science Bulletin
PLATE II
Photomicrographs of *tcin cross sections
1. Bidens frondosa X 76.
2. Helianthus rigidus X 150.
3. HeHanthus rigidus X 76.
4. Solidago rigidiuscula X 76.
5. Helianthus strumosus X 76.
6. Mesadenia atriplicifolia X 76.
Mayberry: Hydrocarbon Secretions
105
PLATE II
106 The University Science Bulletin
PLATE III
J'liotoniifrofiniphs of stem cross sections. (X"^)
1. Erigeron canadensis
2. Pyrrhopappus carolinianus.
3. Lepachiis pinnata.
4. Lactuca pulchella.
5. Ambrosia elatior L.
6. Sili)hiiim speciosum
Mayberry: Hydrocarbon Secretions
107
PLATE III
^•A
gW«"^'
?*:«?L?:^«M-
^' .- "^^j^^oiofHr
.-^r<'^ry^ v-^A
I
2
4- ^"jp^
's^aT''
rK^n-
.^^-^.
V; ^
5 6
108
The University Science Bulletix
PLATE IV
Photomicrographs of rhizome cross sections. (X 76)
1. Solidaso canadensis gilvocanescens.
2. Silphium speciosum.
3. HeUanthus rigidus.
4. Helianthus strumosus.
5. Liatris pycnostachya (corm).
6. Actinomeris alternifolia.
Mayberry: Hydrocarbon Secretions
109
PLATE IV
110 The University Science Bri.LK'iiN
PLATE V
Photomicrographs of leaf cross sections
1. Solidago canadensis gilvocanescens X "*j
2. Silphium laciniatimi X 76.
3. Amphiachyris dracunculoides X 76.
4. Silphium laciniatum X 150.
5. Helianthus salicifolius X 76.
6. Boltonia latisquama X 76.
7. Helianthus rigidus X 76.
8. Erigeron canadensis X 76.
9. Solidago rigidiuscula X 76.
Mayberry: Hydrocarbon Secretions
111
PLATE V
112 The University Science Bulletin
LITERATURE CITED
1. Col, A. 1899. A Quelques Rescherches sur 1' Appareil Secreteur des Com-
posecs. Journal de Botanique. 13 Annee. No. 7.
2. Col. A. 1904. Recherches sur 1' Appareil Secreteur Interne des Com-
posces. Journal de Botanique. 18 Annee. No. 5.
3. Lloyd, E. Francis. 1932. Mode of Occurrence of Caoutchouc in the
Guayule, Parthemium Argentatuni Gray, and its I\uictions. Plant
Physiology. Vol. 7. No. 1.
4. Moenike, Adalbert. 1924. Zur Fragc dcr Harzbildung Bei Umbelliferen,
Compositen and Aralisceenwurzeln. Botanisches Archiv. V Band.
5. Midler, N. J. C. 1866- "67. Untersuchungen iiber die Verteilung der Harze,
iitherischen Oele, Guuuni und Guninnharze, und die Stellung der secre-
tions Behiilter im Pflanzenkoiijer. Pringsheims Jahrbiicher fiir wissen-
schaftliche. Botanik. V Band.
6. Solereder, H. 1908. Systematic Anatomj' of the Dicotyledons. Vol. 1-2.
7. Tetley, Ursula. 1925. The Secretory System of the Roots of the Com-
positae. New Phytologist. Vol. XXIV. No. 3.
8. Transeau, E. N. 1904. (Jn the Development of Palisade Tissue and Resin-
ous Deposits m Leaves. Science, N. S. 19:866-867.
9. Triebel, — . 1885. Ueber Oelbehaltcr in Wiirzeln von Compositen. Nova
Acta der Kgl., Leop., Carol. Deutschen Akademie der Naturforscher.
10. Tschirch, A. 1906. Die Harze und die Harzbehalter mit Einschluss der
Milchsiifte. Vol. 1-2.
11. Tschnch, A. Die Harze. Band I. 1933.
12. Van Tieghem, — . 1885. Canauz Secreteurs des Plants. Annales des
Sciences Naturelles. Septieme Serie. Botanique. Tome Premier.
13. Vuillemin, — . 1884. Remarques sur la Situation de V A]>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
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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 .•<igitntu>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. p<dlid(i Osborn); Nesostclcs Kirkaldy (^ Auounana Distant);
Nesostclcs iuci.sus Matsunuira (:=z Euguathodus l(U't( us Baker, ^= Euguathodus
bisinuatiis DeLong, =^ Euynathodus pallidus Osborn, ^ Enguatlwdus bijurcatus
DeLong and Davidson, ^ Nesosteles arcolata Osborn, and ^ Nesosteles tutuil-
ann Ost)orn). Changes in generic assignment of sjiecies, other than that in-
dicated in the designation of genotypes and in the new synonymy: Jlulcria
affinis (Jsl)orn transferrctl to Arrugada; Scaphoidcus mexicnuus Osborn from
Purtanus to Osborucllus; Scaphoidcus boUine7}sis Baker, <S. hasemani Baker,
and *S'. hiiigifoniis Osborn to Portauu.^ Ball; Plat ymclnpius longiceps Berg to
Agudus Oman; Eutettix neociuctus Osborn, E. elegans Osborn, and Mcsamia
jnsciata Osborn to Me)wso7)ia Ball; Jassus palliditarsus Stal, Eutettix irroratus
Osborn, E. jemoratus Osborn, E. laticeps Osborn, and Aligia plena Van Duzee
to BaJiila Oman; Eutettix t(ss( lulus Osl)orn and TliaiNnotetlix hibatus Osborn
to Atanus Oman; Euscelis cjuadratulas Osljorn, E. pallida Osborn, Athysa)ius
digressus Van Duzee, Thamnotettix capieola Stal, Athysanus fusconervosiis
Motschulsky, Jassus t(u niaticeps Kirschbaum, Athysanus indicus Distant, A.
vanus Distant, A. atkiusoni Dislant, .1. siuiillini us Matsumura, and .4. traus-
versalis Matsumura to Exitianus Ball; Tha)nnot( ttix umazoiiensis Osborn to
Brazosa Oman; Agallia viulti punctata Osborn to Alaea Oman; Thamnotettix
Ivteosus Baker to Chlorotettix Van Duzee; Cicadula maidis DeLong and
AVolcott to Hiddnlus (Jman ; Euseclis (luadiiunu-ulata Osborn, and Thamnotet-
tix guaporoisis Oman to Neobala; Neocoelidia bimaculata Baker, N. 7nodesta
Baker, A'', inflata O.^^born, and N. eroceata Osborn to Coelidiana Oman; Neo-
coelidia smith ii Baker to Chiiuiia Bruner and Metcalf. Idiocerus Lewis is
placed in the Eurymelinae, Ionia Ball in the B3'thosco])inae and Chiuaia Bruner
and Metcalf in the Jassinae. P<u'h yojisis IThler is removed from synonymy.
Ionia Ball is consitlered as a subgenus of Aceralagidlia Kirkaldy, and Del-
topinus Ball, Cloatithus Ball, Nas:)itoidens Ball, and Convelinus Ball are con-
sidered as subgenera of Sea])li yt<>j)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
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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. Zo<ll. U. of Midi., No. 259, April 3, 1933,
pp. 12-13.
One specimen (No. 154) collected five miles southwest of Her-
mosillo was found coiled at night near an isolated rock mass. As
my light approached, the snake neither moved or rattled; in fact
the rattles were not sounded until it was being carried in tlie sack.
A second specimen (No. 224) was found coiled at night in low
mountains near La Posa.
These specimens have been studied by Mr. Howard K. Gloyd,
who, presumably, will publish the scale data taken.
TURTLES
Kinosternon sonoriense Le Conte
Kinusternnuvi sonoriense Le Conte, Proc. Acad. Nat. Sci. Phila., 18.54, p. ls4 (type de-
scription; type locality, Tucson, Ariz.).
Two specimens were taken, one consisting of a carapace and bones
of a dead specimen found in a dry arroj'o. Another specimen was
captured in the highway, fifteen miles north of Hermosillo. (No.
1168.1
Gopherus agassizii (Cooper)
Xerobates agassizii Cooper, Proc. California Acad. Sci., II, 1S()3, p. 120 (type de.-;cription ;
tjpe locality, mountains near Fort Mojave, Calif.).
This species was rather common in Sonora. Four specimens were
collected and several seen were not collected. Their usual hide-
outs during the day were small excavations around the base of
cliffs, and in the soft earth in old cliff caves.
I do not have material available from the tyjie locality, so the
association of these specimens with agassizii is done without due
critical examination.
The specimens are: No. 115, 40 miles south of Noria, June 22,
in a large rock crevice filled partly with refuse from a colony of
leaf-cutting ants; Nos. 237 and 237a, at base of cliff near La Posa,
June 29; No. 1167, twenty miles north of Noria, August 11, in road.
Taylor: Fauna of Sonora 499
ANNOTATED BIBLIOGRAPHY
Allen, Morrow J. 1933. Report on a collection of amphibians and reptiles
from Sonora, Mexico, with the description of a new lizard. Occ. Papers Mus.
Zool. Univ. of Mich., Xo. 259, Apr. 3, 1933, pp. 1-15.
Lists four amphibians, eleven lizards, six snakes and one turtle. One
lizard, Dipso-saurus dorsalis sonoriensis, is described as new. Plernohyla
jodicns and Gastrophrync texensis are rei)orted for the first time from
Sonora (Xoria).
Baird, S. F. 1858. Descriptions of new genera and species of North American
lizards in the musoinn of the Smithsonian Institution. Proc. Acad. Nat. Sci.
Phila., VII, pp. 253-256.
B.\ird, S. F., and Gir.ard, C. 1853. Catalogue of North American reptiles in
the museum of the Smithsonian Institution, part I. Serpents. Jan., 1853,
pp. 1-172. Washington. Crotalophonis Edwardsii (Sonora) ; Heterodon
nasicus Baird and Girard (Sonora) ; Piluophis bellona (Baird and Girard)
(Sonora); Ophibolus splendidus (Sonora); Salvadora grahamiae (Sonora);
Diadophis rcgalis (Sonora); Sonora semiannulata (Sonora).
There is doubt that the type localities of the new forms described actually
are in territory now included in Sonora.
1854. Characteristics of some new reptiles in the museum of the Smith-
sonian Institution. Proc. Acad. Nat. Sci. Phila., VI, 1852, pp. 125-129.
Barbour, Thomas. 1921. A new lizard from Guaymas, Mexico. Proc. New
England Zool. Club, 7, pp. 79-80.
Beldixg, L. 1887. Reptiles of the cape region of Lower California. West
Amer. Scientist, III, pp. 97-99.
Burt, C. E. 1931. A study of the tciid lizards of the genuis Cnemidophorus
with special reference to th(>ir 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 ■■<taN.^biiriaiia slcjin i/cri listed from Empalme.
1935a. Miscellaneous notes on Mexican lizards. Univ. Kan. Sci. Bull.,
XXII, April 15, 1935, pp. 119-155, pis. XXIII-XXV; text fig. 1. Phyllo-
dactylus homolepidurus is described as new from Hermosillo and Guaymas.
19351). Notes on some Mexican lizards of the genus Holbrookia, with
the descriptions of a new species. Univ. Kan. Sci. Bull., XXII, April 15,
■ 1935, pp. 185-201, pis. XXVIII, XXA'III. Notes on Holbrookia clcgans
thcrmopliila Barbour.
T.wLOK. E. H. 1933. New species of skinks from Mexico. Proc. Biol. Soc.
Wash., Vol. 46, (Jet. 26, 1933. i^p. 175-182. Eumeces parviawicidatus (Alamos,
Sonora).
1936. Desciiption of a new Sonoran snake of the genus Ficimia, with
notes on other Mexican species. Proc. Biol. Soc. Wash., Vol. 49, May 1, 1936,
pp. 51-54. Ficimia deserlorum (La Posa, 12 miles northwest of Guaymas).
Taylor: Fauna of Sonora 501
Van Denburgh, John. 1894. 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.
1922. The reptiles of western North America. Occ. Papers Cal. Acad.
Sci., X, Nov. 23. 1922, Vols. I and II, pp. 1-1028, pis. 1-128.
The following list from thi.s work, which purports to be complete to date,
1922, gives the known species of the state of Sonora : Coleonyx vanegatus
(Baird) (San Miguel de Horcasitas; Tepoca Bay); Dipsosnnrus dorsalis
doffialis (Baird and Girard) (northwestern Sonora) ; Snuro))iaulus townsendi
Dicker.son (Tiburon Is.; Guaymas) ; Crotaphytus collaris bnileyi (Stejneger)
(Hermosillo; Sierra Tule; Tiburon Is.); Crotaphytus wislizenii Baird and
Girai'd (northwestern Sonora; Tiburon Is.); Uma notata Baird (Tepoca
Bay); Calli.saurus imisitatus Dickerson (Tiburon Is.; Tepoca Bay; San
Pedro Bay; Guaymas; San Miguel de Horcasitas); Holbrookia maculata
approxiniaiif< (Baird) (Duros Millos; headwaters of the San Pedro river);
Holbrookia elegans Bocourt (Guaymas); Holbrookia lexana (?Sonora);
Uta ornata lateralis (Boulenger) (Guaymas; Tiburon Is.); LHa ornata sym-
metrica (Baird) (two miles south of Nogales; 32 miles south Nogales;
Duros Millos) ; I'ta .^tari.sburiana clcgans (Yarrow) (Tepoca Bay; San Pedro
Bay; Tiburon; Patos and Pelican Islands); Sceloporus scalaris Wiegmann
(no localities given) ; Sceloporus consohrinus Baird and Girard ("Sonora,"
Cope) ; Sceloporus jarroi^ii Coi)e (Pinetos Camp, 32 miles south of Nogales) ;
Sceloporus (iir(jii<iliis poiuscltii (Baird and Girard) ("Sonora"); Sceloporus
magister Hallowell (Tibui'on Is.) ; Scelopo)nis clarkii Baird and Girard (32
miles .south of Nogales; San Pedro Bay; Tiburon and San Pedro Nolasco
Islands) ; Phrynosoma douglassii her)iartdesi Girard (northern Sonora) ;
Phrynosoma ililuiarsi Stejneger (Sonora, a short distance south of the border
of Arizona) ; Pliryuosotnci solare Gray (Hermosillo) ; Phrynosoma goodei
Stejneger (Gulf Coast of Sonora) ; Phrynosoma m'ealUl (Hallowell) (Sonora) ;
Phrynosoma modcstum Girard (Sierra de la Narizo) ; Hclodcrma suspectmn
Cope (Guadalupe Caiion; San Bernardino; Niggerhead Mountain; San
Pedro Bay) ; Gerrhonotus kingii (Gray) ("Sonora," Cope) ; Cnemidophorus
gularis Baird and Girard (2 miles south of Nogales; Pinetos Camp 32 miles
south of Nogales; San Pedro Bay; Gua.ymas) ; Cnonidophorus mclaiioste-
thus Cope (Hermosillo; Tepoca Bay; San Pedro Bay; Guaymas; Tibu-
ron); Siagonodon humilis (Baird and Girard) (San Miguel do Horcasitas);
Constrictor imperator (no locality given); Diadophis regalis Baird and
Girard (San Magdalena) ; Heterodon nasicus Baird and Girard ("Sonora") ;
Coluber flagellum piceus (Cope) (Hermosillo; Tiburon Is.); Coluber semi-
lineatus (Cope) (San Pedro Bay); Salvadora hexalepis (Cope) (Guaymas;
Sierra Blanca; Tiburon Is.); Lampropeltis getulus yumcnsis Blanchard
(Sonora); Hypsiglena ochrorhynchus ochrorhynchus Cope (Guaymas);
Thamnophis eques (Reuss) (Guaymas) ; Thamnophi-s marcianus (Baird and
Girard) (Sonora) ; Sonora semianjiulaia (Baird and Girard) (Sonora) ;
Chilonieniscus cinctus Cope (Guaymas, t.ype locality); Oxybelis acuminatus
(Wied) (no definite locality given) ; Tnmorphodon lambda Cope (Guay-
mas); Micrurus euryxanthus Kennicott (Sonora); Hydrus platurus (Lin-
naeus) (Pacific coast, Guaymas) ; Kiiiosternon. sonoriense (Le Conte) (San
Pedro river; Cajon Bonita creek; San Bernardino river; Guadalupe Caiion;
Gopherus agassizii (Cooper) (Tiburon Is.; San Pedro Bay); Dermochelys
schlegelii (Garman) (seen at Guaj'mas).
502
The University Science Bulletin
PLATE XLIII
Fig. 1. Ficimia dc'icrtonnn Taylor. Photograph of type.
Fig. 2. Cnemidophorus burti sp. nov. Photograph of type.
Taylor: Fauna of Soxora
503
PLATE XLIII
^H i i ^
/.
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vol. XXIV.] July 15, 1936 [No. 20.
Notes on the Herpetological Fauna of the Mexican
State of Sinaloa
EDWARD H. TAYLOR, Department of Zoolog:>-, 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
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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()</( )i!js, j)()ins< llil, IIik ninlcyah's, j. jorrovii, j. minor, j. immucroimtus,
o. oriialus, o. cuienilcuf^, d. (hu/csu and (/. intcrrncdias. S. jcrrnriperczi Cope is
s^'nonyniized with S. t. lorqualus Wiegmann; <S. pleurolepis Giinthcr is synon-
ymized wit'h S. d. dugcdi Bocourt. S. gucntheri Stejneger is referred to the
spinofiut^ grouji. An analysis is presented of th(- phylogeny of the eleven i)ro-
posed groups (chri/sostictus, scalaris, t.>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<or\\.\\a\%
^" ^ nosol
supralabials
rostral
infrolabials
mental
obiomentols (outer rou^)
-postmeritals
_ lobiomcntals(inntriou)
-lortlobials
~ suboculor
mankal
__ infralabials
- w-'postmentols
labiomental (outer n
lobiom«ntals(inn»rr<
-— rostral
— internosols
--- subnosQl
--. nasal
■■- caatKols
"" frontonasals
"'prefrontals
"■ fronta(,Qnl:. part
""" supcrcillaries
frontal, post. part
frontoparietal
parietal
interparietal
Fig. 1. Head scales of Sceloporus torquatus jnclanogaster (Cope), showinj;
nomenclature adopted in thi.s paper. DHD & HMS 453, near San Tibiucio,
Zacatecas; actual head length, snout to occiput, 19 mm.
jpomRcttii the frontal is usually broken irregularly into several
scales. It is significant that in species in which the supraoculars
are in two rows, the frontal rarely or infrequently touches the in-
terparietal. The reverse is the case in species having a single row
of supraoculars.
Prefrontals. Two I'ather large scales preceding the frontal. They
may be in contact medially, or separateil either by the contact of the
median frontonasal with the frontal or by an azygous scale.
Frojitonasals. Three large scales preceding the prefrontals the
median of which is usually slightly the largest and may be in con-
tact with the frontal.
Internasals. Apparently two pairs of internasals is the ideal con-
dition, but this condition is seldom realized. The scales are usually
very irregularly divided.
544 The University Science Bulletin
Nasals. Small scales in which tiie nares are pierced. They are
never divided, and are always separated from the rostral. An ir-
regular series of small scales surrounds the nasals except below,
where a subnasal is present.
Si(praoculars. A series of large scales above the orbit, in some
species divided irregularly, or, in others, into two regular row's.
They are always separated from the median liead scales by a single
row of small scales varying in size and general shape according to
species (except in serrifcr, in which species the last supraocular may
not be separated completely from the median head scales). One,
two or three complete or incomplete rows of small scales separate
the supraoculars from the superciliaries, more or less according to
species or subspecies.
Superciliaries. Six superciliaries are always present, the first four
imbricating posteriorly, the fifth completely hidden below the fourth,
and the sixth o^•erlapping the posterior part of the fourth.
Canthals. Typically two, the posterior of which does not extend
far onto the superciliary region. The anterior canthal is, in some
species, frequently forced above the canthal ridge by contact of the
second (posterior) canthal with the subnasal.
Subnasal. A re]ati^■ely large scale immediately below the nasal,
in contact with the anterior canthal and loreal.
Loreal. A small scale below the canthals, in contact anteriorly
with the subnasal and posteriorly with the preocular. It may rarely
be divided into two or three scales. The first canthal occasionally
separates the loreal from the subnasal and contacts the rows of
scales above the supralabials.
Preocular. A small scale, witli a heavy keel near its upper pos-
terior edge, segmented from the anterior end of the suborular. The
preocular is, in some species, divided longitudinally.
Svbocular. A large, curved, elongate scale immediately below
the eye, with a heavy keel near its upper edge.
Postoculars. A'ariable, usually two. They follow the subocular,
curving posteriorly and upward at the edge of the orbit, and are
distinguished from the temporal scales by being heavily keeled and
usually somewhat larger.
Lorilabials. The small scales above the supralabials are so called.
On the sides of the head they are usually in two rows, sometimes
reduced to one row at some point below the suboculai'. One of
these rows invariably continues about the snout, passing immedi-
atelv above the rostral.
Smith: Genus Sceloporus 545
Svpralabials. A series of scales around the upper labial border,
excluding the scale at the tip of the snout. The scales of this series
are smooth and nonimbricating, and do not vary sufficiently to be
of assistance in distinguishing species and subspecies.
Infralabials. A series of scales around the lower labial border,
excluding the scale at the mandibular symphysis. The scales of this
series are similar in character to the supralabials.
Mental. A median anterior pentagonal or triangular scale border-
ing the lip.
Postmentals. A series of enlarged scales on each side posterior
to the mental. The scales are paired; those of the anterior pair are
always in contact medially, and those of the following pairs are
separated from each other by a varying number of scales. The
number of postmentals is irregular and of little taxonomic sig-
nificance.
Labiomentals. Two series of scales on each side between the post-
mentals and infralabials. In some species the anterior scale of the
outer row may usually contact the mental, and this condition is of
some taxonomic significance. When the anterior scale of the outer
row is separated from the mental, it is only by narrow contact of
the first infralabial with the first postraental.
The inner row of labiomentals never extends as far forward as
the outer row, and the position at which it terminates, in relation to
the infralabials, is of considerable taxonomic significance.
Auricular lobules. These are the scales on the anterior border of
the ear. Their number and relative size are of importance.
Lateral nuchal fold. A dermal pocket between the arm and the
ear. The lining of this pocket is without scales or only with small,
granular scales.
Postfemoral dermal pocket. A dermal pocket at the posterior
margin of the insertion of the hind leg. The lining is without scales
of any kind. It is not present in any species of the torquatus group.
Enlarged postanals. Two smooth, enlarged scales immediately
posterior to the anus in males. They are normally present in all
members of this group, but in jarrovii and its subspecies they are
occasionally lacking or very poorly developed.
Femoral pores. A longitudinal series of pores along the postero-
ventral border of the thighs. The number of pores in each series
is of much importance. They are always present in Sceloporus, and
35—6037
546 The University Science Bulletin
in the torquatus group are never less than nine, and the series never
approach each other closely on the median ventral line.
In a number of species of the torquatus group, the scales in the
preanal region are modified in such a manner that they appear pore-
like. They are not regularly so modified in any species of this
group, and never occur in most species.
General character of scales. The dorsal scales of the body in
species of the torqvxitus group are typically keeled, and with a
terminal mucrone. In a number of species the median dorsal scales
may lack terminal mucrones. The lateral scales are usually some-
what larger than the dorsal scales, and more strongly keeled and
mucronate. Spines or denticulations, when present, are more nu-
merous on the lateral scales than on the median dorsal scales. The
dorsal caudal scales are usually somewhat larger than the median
dorsal scales on the body. In ornatus the scales on the body have
terminal pits.
The ventral scales are always smooth, and in most species are
notched. The dorsal head scales are always pitted to some extent.
The dorsal scales of dugesii are remarkable in the fact that the
terminal mucrone arises distinctly within the free margin of the
scale. This is most evident on the lateral scales of the body, the
dorsal scales of the thigh and the temporal scales. Related species
show a tendency toward mucronation of this type. The extreme
development is in dugesii dugesii.
The dorsal scale rows are usually parallel, occasionally convergent
and rarely divergent. All three conditions may occur rarely in one
species. No species has divergent scale rows except as an aberrant
variation, while some species normally have convergent scale rows,
with occasional variants possessing parallel scale rows.
Lamellar formulae. The number of lamellae under the free parts
of the fingers and toes is of some importance in Sceloporus. The
range of variation in the torquxitus group is slight and of little signif-
icance. The lamellae are always tricarinate as in many other gen-
era of Iguanidae.
Nuchal collar. The most important characteristic for definition of
members of the torquatus group is the presence of a black, light-
bordered collar about the neck, which may or may not be complete
ventrally (only complete in males), according to species and age of
the individual. The light borders may be incomplete (most incom-
plete in a certain phase of jarrovii jarrovii) , but never absent. No
species not belonging to the torquatus group has such a collar. How-
Smith: Genus ScELOPORrs 547
ever, certain species of related groups (microlepidotus, formosus and
spinosus) may have black collars about the neck. These collars dif-
fer in lacking the light borders, and the species have other differen-
tial scale characters.
Ventral coloration. Females are light-colored below, and mark-
ings are confined, when present, to the gular region. Adult males
always have some type of ventral coloration. Usually the gular area
is bluish, as well as the sides of the abdomen. The blue areas on the
sides of the abdomen may be black-bordered. In some species the
entire ventral surface is blue.
Size. Maximum size is considered to be of as great importance
as many of the scale characters. The range in maximum size, in the
torquatus group, is from 75 mm. snout to vent {ornatus caeruleus)
to 143 mm. (cyanogenys) . S. ornatus ornatus and S. dugesii inter-
medius are approximately as small as ornatus caeruleus, while tor-
quatus melanogaster approaches closely the size of cyanogenys.
In the tables of measurements and scale counts, when no museum
is given for a certain specimen number, the Mexican collection at
Kansas University is referred to.
HISTORICAL DISCUSSION
A number of revisions of Sceloporus have appeared since the de-
scription of the genus by AViegmann in 1828. Six species were orig-
inally described in the genus — torquatus, spinosus, grammicus, pleu-
rostictus, aenev^, and scalaris, of which torquatus is the generic
type. In 1834 Wiegmann published his "Herpetologia Mexicana,"
in which he recognized nine species — torquatus, formosus, spinosus,
horridus, grammicus, microlepidotus, variabilis, aeneus and scalaris.
Dumeril and Bibron, in 1837 (Erpetologie generale), recognized ten
species, adding undulatu^ Latreille.
The next important work is that of Bocourt in the Mission
Scientifique, in 1874. This author recognized twenty-two species,
only two of which are of the torquatus group [torquatus and du-
gesii.). He also recognized poinsettii as a variety of torquatus.
S. ornatus, described in 1859 by Baird, and serrifer, described by
Cope in 1866, are mentioned without description.
In 1885 Cope published a synopsis of Sceloporus, in which he
recognized thirty-six species and subspecies, eleven of which belong
to the torquatus group. At that time only one other species of the
group had been described which Cope did not recognize — inter-
medins Duges (1877).
548 The University Science Bulletin
In the same year, the catalogue of the lizards in the British Mu-
seum, by Boulenger, appeared, and in this thirty-three species and
subspecies were recognized, nine of which are of the torquatus group.
In 1890 Giinther published a synopsis of Sceloporus in Biologia
Centrali-Americana, recognizing thirty species. Seven other de-
scribed forms are listed without comment as to validity.
In 1897 Boulenger presented his conclusions with regard to the
species of Sceloporus in his revision of the genus. He recognized
thirty-six species and subspecies, seven of which belong to the tor-
quatus group.
The last monograph of the genus is that of Cope, published in
1900, in "The Crocodilians, Lizards and Snakes of North America."
In this are recognized forty species and subspecies, eleven of which
belong to the torquatus group.
In this work, which deals with the torquatus group, I recognize as
valid sixteen species and subspecies. These include only ten of the
twelve recognized by Cope (1900), together with two previously
described.*
GROUPS IN THE GENUS
It appears that the genus is divisible into eleven fairly definite
groups, which I name as follows: torquatus, jormosus, microlepid-
otus, variabilis, scalaris, siniferus, spinosus, utiformis, undulatus,
pyrocephalus and maculosus. Of these eleven groups, the torquatus,
spinosus and undulatus groups are the largest. The species of the
spinosus group are well differentiated and bespeak an age greater
than that of the undulatus and torquatus groups. In these two
groups subspecies are numerous and their separation is compara-
tively difficult.
PHYLOGENY OF THE GENUS SCELOPORUS
Sceloporus is one of the most nearly ideal of living genera of
reptiles for the study of speciation and related phenomena. The
characteristics which it possesses and which are essential to an ideal
genus for such studies are :
1. A large number of living forms. One hundred and seven
species and subspecies have been described. Approximately eighty-
two of these are valid.
2. Prolificity. Where Sceloporus occurs, usually it is the most
common of all reptiles, or for that matter, of all vertebrates.
* It appears that Scelaporvs guentheri Stejneger (1918) does not belong to the torquntus
group, but probably to the spinosus group.
Smith: Genus Sceloporus 549
3. A large range, entirely contiguous. The genus occupies prac-
tically all of the United States, and occurs as far south as Panama.
4. Great adaptability. Species in this genus have adapted them-
selves to a considerable range of elevation — from below sea level
(Death Valley) to about 13,500 feet above sea level. They occur
in almost every conceivable terrestrial habitat — deserts, sand dunes,
forests, on rocks, trees, or ground, in grassy plains or heavy brush,
and even on houses, fences and other man-made structures.
5. Lack of obvious distinctive specific characters. Subspecies are
numerous and species not so well defined as in many other genera
of animals, and for this reason relationships may more definitely
be postulated.
These characters are indicative of a group of relatively recent
development.
It seems likely that Uta has been derived from Sceloporus. Uta is
undoubtedly more closely related to Sceloporus than any other
living genus. I assume this to be true because of the remarkably
close agreement of certain members of the variabilis group of
Sceloporus with certain members of the ornata group of Uta. I
present the following data in support of this view.
A postfemoral dermal pocket is known to be regularly present in
Uta ornata and its subspecies, U. caerulea, U. leiris, U. stansburiana
hesperis, and U. taylori, and is variable in certain other species and
subspecies.
In certain species of Sceloporus of the variabilis group, a vestigial
gular fold is present immediately in front of the arm which is
homologous with the gular fold present in Uta. In Sceloporus it is
most distinct in *S. couchii and S. merriami. In the former species,
there is present also a postfemoral dermal pocket immediately be-
hind the insertion of the hind leg. The dorsal scales are extremely
small for the genus, the laterals are minute and the size of the species
itself is small.
In Uta levis (as well as in the related U. ornata), the gular fold
is remarkably poorly developed. It is frequently interrupted medi-
ally, the scales of the gular region passing directly into the ventrals;
in many cases in which the fold is not thus completely interrupted,
small scales replace medially the tiny lateral granules. In U. levis
the lateral scales are uniform in size and very minute — practically
granular — but visibly imbricated. The enlarged dorsals are in some
specimens in several rows, gradually decreasing in size laterally and
550 The University Science Bulletin
grading into the lateral scales. Also, the dermal pocket is present
behind the insertion of the hind leg, as in S. couchii. Its size is
comparable with that of the latter.
It is not a far cry from the smaller, almost granular lateral scales
of Uta levis to the larger, but yet minute, laterals of S. couchii; nor
is it impossible to conceive of an increase in size of the lateral dor-
sals from a few rows of enlarged scales to several or many. The
vestigial gular fold of S. couchii is comparable with the partially
developed fold of Uta levis. A postfemoral dermal pocket is present
in each, and the sizes of the species are comparable.
It may be stated that the nearest point of contact between the
two genera, so far as it may be traced in living forms, is between
the species levis of Uta and couchii of Sceloporus. It cannot be said
that either gave rise to the other ; it is more likely that the two di-
verged from an ancestral type not greatly different from the above
two species. By this view the variabilis group (to which couchii
belongs) is the most primitive of the genus, while the torquatus
group is probably the most highly specialized.
It appears that the Sceloporus stock, early in its evolution, sepa-
rated into two divisions; from one came the large-scaled forms,
mostly of large size, and from the other came the small-scaled forms,
all of small size. From the latter division developed, among others,
the variabilis group.
The extensive range of Uta, although smaller than that of Scelop-
orus, and the relatively considerable amount of speciation, would
indicate that Uta is of considerable age. It would follow that the
variabilis group of Sceloporus and the ornata group of Uta are rela-
tively ancient, yet of much more recent development than the an-
cestral stock which gave rise to the other groups of Sceloporus. I
assume that the ornata group is primitive to other groups of Uta;
but it appears that the variabilis group is a specialized, divergent
group which has given rise, not to other species and groups of Scelop-
orus, but to the quite distinctive genus Uta.
The groups of the small-scaled division of Sceloporus are, for the
most part, difficult to allocate in a phylogenetic scheme, due to their
rather vague relationships. The following discussion may support
the tentative arrangement as here adopted.
Of the small-scaled forms, the chrysostictus group first became
stabilized. The variabilis group is undoubtedly closely related to the
chrysostictus group. S. cochranae (of the chrysostictus group) very
closely resembles variabilis variabilis, differing most markedly in the
Smith: Genus Sceloporus 551
absence of a postfemoral dermal pocket. The pyrocephalus group
possesses a species {gadoviae) which shows relationship with va-
riabilis by the possession of a postfemoral dermal pocket, and is
associated with the variabilis stem. The utijormis group also shows
direct relationship with the variabilis stem through S. merriami,
which possesses a prominent vestigial gular fold. Both members of
the utijormis group {utijormis and merriami) possess granular lat-
eral scales,
undutafus
torquatus
^Pfnosus
pyrocephalus
maculosus
• chrysostlctus
mictolepldotus- _ . ^ ^
^ ^ scalarls
SCELOPORUS
Fig. 2. Diagram illustrating the proposed phylogeny of the groups of
Scelopo7-iis. Generic names are in caps, group names in lower case.
The scalaris group has its closest relatives in the chrysostictus
group, through jalapae of the former group and ochoterenae of the
latter group.
The maculosus group, containing the single species maculosus,
has no very close relatives, and is difficult to place in a phylogenetic
tree. I assume, however, that its closest relationship is with the
pyrocephalus group, through S. nelsoni of the latter group. S. ma-
culosus certainly has no close relationship to the variabilis group,
lacking all specialized characters of the latter group; from the sea-
552 The University Science Bulletin
laris group it differs widely in possessing oblique rows of lateral
scales and smooth head scales; from the chrysostictus group it differs
in lacking keeled preanal scales in females, and rugose head scales.
The utijormis group is widely different from maculosus in the char-
acter of the lateral scales and the presence of a vestigial gular fold.
The remaining five groups {microlepidotus, formosus, spinosus,
undulatus and torquatus) apparently have no close relationship
with the other groups, and apparently diverged from a common
stock which early separated from the other. The microlepidotus
group is assumed to be the most primitive of these, largely because
of its very small scales. The habitus of the species of this group
and the fact that the species are ovoviviparous associate them defi-
nitety with the torquatus and jormosus groups. Certain species of
the microlepidotus group have large scales and closely resemble
forms of the jormosus group.
The formosus group appears to be near the common ancestral
stock of both the spinosus and torquatus groups, through acanthinus
of the former group and serrifer of the latter group. It may be added
that these two species are also the most closely related of the spi-
nosus and torquatus groups, respectively.
The undulatus group undoubtedly is most closely related to the
spinosus group, through undulatus of the former group and flori-
danus of the latter.
Two apparent discrepancies appear in the arrangement of the last
five groups discussed. First, all are of relatively large size except
the undulatus group; and second, three groups are ovoviviparous
{microlepidotus, torquatus and formosus) and the other two are
oviparous.
The relatively small size of the species of the undulatus group
must be assumed as a parallel development rather than a direct in-
heritance of the small size of the ancestor in the varibilis group,
for the close relationship of the spinosus and undulatus groups can-
not logically be disputed, nor is the close relationship of the spi-
nosus, torquatus and formosus groups doubtful. It is possible, or
even likely, that certain primitive species of the spinosus group,
such as acanthinus, may be ovoviviparous, and that this character
has been dropped in the further evolution of the group. It is well
known that both ovoviviparity and oviparity occur in closely re-
lated species of Phrynosoma.
Smith: Genus Sceloporus 553
DISCUSSION OF THE TORQUATUS GROUP
Habits and habitat. As a rule, species of this group are confined
to rocky habitats. So far as I am aware, only cyanogemjs tends
to live on or near the ground. S. jarrovii occasionally, and serrifer
perhaps usually, are found on trees. Individuals of most species are
extremely wary and difficult of acquisition. In central Mexico a
number of species are found on rock fences in great abundance.
Mating probably takes place in early spring, inasmuch as the
young are born in May or early June. Several species of this group
are known to be ovo viviparous, and the remainder are presumed
to be so.
Species included. The following species have been described:
S.bullen Boulenger (1894, pp. 729-730, pi. 48, fig. 3).
S. ornatus caeruleus Smith.
S.torquatus cyanogenys Cope (1885, p. 402).
S.dugesii Bocoiirt (1874, pp. 188-190, pi. 18, figs. 7, 7a, 7b).
S. ferrariperezi Cope (1885, p. 400).
S. jarrovii immucronatus Smith.
S.intermedius Duges (1877, pp. 29-34, pi. 1, fig.?. 21-32).
S. jarrovii Cope (1875, pp. 569-571, pi. 23, figs. 2-2d).
S. lineolateralis Smith.
S.melanogaster Cope (1885, pp. 400-401).
S. torquatus minor Cope (1885, p. 402).
S. torquatus rmicronatixs Cope (1885, p. 402).
S. omiltemanus Giinther (1890, p. 66, pi. 32, fig. A).
S. ornatus Baird (1859, p. 254).
S.jwiruseltii Baird and Girard (1854. pp. 126-127).
S.torquatus Wiegmann (1828, p. 369).
S.pleurolepis Giinther (1890, pp. 74-75, pi. 32, fig. B).
S.serrijer Cope (1866, pp. 124-125).
It is proposed herein that the following names should be recog-
nized:
y (S. bwWen Boulenger. S.mucronatus mucronatus (Cope).
S. cyanogenys (Cope). S. mucronatus omiltemamis (.GiJnther) .
H. S. dugesii dugesii Bocourt. \ S. ornatus ornatus Baird.
\ S. dugesii interm,edius (Diiges). S. ornatus caeruleus Smith.
\/ S. jarrovii jarrovii Cope. S.poinsettii Baird and Girard.
i. S.jarroini minor (Cope). S.serrijer Cope.
i S. jarrovii immucronatus Smith. S.torquatus torquatus Wiegmann.
y^^S. lineolateralis Smith. S.torquatus melanogaster (Cope).
554
The University Science Bulletin
I regard it necessary to synonymize Cope's ferrariperezi with t.
torquatus Wiegmann, and Giinther's pleurolepis with d. dugesii Bo-
court.
It appears that soon after the separation of the torquatus stock
from the other groups of Sceloporus, there was a separation into two
divisions, one of which exhibited a tendency to develop small scales,
jAffROVII
POIHSETTII
DUGESII
CYAHOGBNYS
UUCPONATUS
OUILTEUANUS
SERRIFBf?
Fig. 3. Diagram illustrating the geographical relationships and derivations
of species and subspecies of the torquatus group. Overlapping of ranges is not
indicated.
the other large scales. This small-scaled division is now composed
of species of smaller size, while the large-scaled division is composed
of species of larger size. In the latter, the maximum size of its
smallest species is greater than the maximum size of the largest
species of the small-scaled division. Each division contains certain
species having one row of supraoculars and others having two rows.
This character of the supraoculars I would consider to be of less
importance in distinguishing major divisions than maximum size of
species or size of scales on the body, because of the greater vari-
ability and more erratic appearance of this character in species not
closely related.
Smith: Genus Sceloporus
555
S. serrifer appears to be the oldest of the large-scaled species.
The postulation that this species, which is one of the larger ones of
the genus, and one having large scales, is nearest to the ancestral
type of the large-scaled division of the torquatus group may appear
to be contradictory to the postulation that Sceloporus is derived from
/ IMiUCROHATUS
' WNOn
I
SUPMOCUIAKS
I TWO RCMh
POmSCTVI '
CYANOCENrs I
5UPRA0CUlAOi,,\
ONE ROW \
S£RmFEf> K
/ sSUPgAOCUUKS
Fig. 4. Diagram illustrating the proposed phylogeny of species and subspecies
of the torquatus group. Group names are given in larger letters.
small species with small scales. However, my assumption seems to
be justified by the fact that serrifer occupies a southern position on
the periphery of the geographical area now occupied by the torqua-
tus group ; and by the fact that it shows its closest relationship with
the more primitive forms in the same area of the spinosus and
jormosus groups, which are obviously older, as indicated by the
clearer definition of the species associated wdth them, and the fact
that their present geographical distribution is peripheral to the gen-
556 The University Science Bulletin
eral area now occupied by the genus. The jormosus group has,
moreover, a discontinuous distribution also indicative of greater age.
Of the other species of the large-scaled division of the torquatus
group, cyanogenys, poinsettii and omiltemanus have scales smaller
than the remainder, and are for this reason considered to have di-
verged early from the restricted torquatus stem. S. mucronatus ap-
pears to be the nearest to the ancestral type of these three species,
despite the fact that it has larger scales than they. I so conclude
because of its centralized geographical position with relation to the
area occupied by the other three forms.
Of the small scaled species, it appears that lineolateralis is the
oldest living species. I am led to this belief because of the fact that
this species occupies a relatively isolated area between the area
occupied by most members of the small-scaled division and that
occupied by most members of the large-scaled division. Further, it
combines certain features of scalation more characteristic of species
in the large-scaled division with other features more characteristic
of species in the small-scaled division. S. jarrovii appears to be
most closely related to lineolateralis. From this species, or its
ancestors, the remaining species of the small-scaled division have
obviously been derived.
Key to Species
A. Supraoculars in a single series, with no scale divided.
B. Femoral pores, 8 to 14; dorsal scales, 28 to 35 S. serrijer Cope, page 558
BB. Femoral pores, more than 14 ; or, dorsal scales, more than 35.
C. Dorsal scales, 31 or less.
D. Nuchal collar divided on each side of neck, the area between
lighter in color ; dorsal color light, with dark and light spots ir-
regularly placed ; size large (maximum snout to vent measurement,
129 mm.) ; dorsal scales more strongly keeled and mucronate.
S. torquatus vielanogaster (Cope), page 571
DD. Nuchal collar broad, complete; dorsal color dark, without light
spots ; maximum snout-vent measurement, 98 mm. ; dorsal scales
more weakly keeled and mucronate.
S. torquatus torquatus Wiegmann, page 564
CC. Dorsal scales, more than 31.
D. Lateral scales about half as large as median dorsals.
S. lineolateralis Smith, page 617
DD. Lateral scales as large as or larger than dorsals.
E. Dorsal scales, 40 or more ; nuchal collar narrow, with light
borders broken ; a light line on side of head and another on
side of neck; each dorsal scale usually with a light medial
spot S. jarrovii jarrovii Cope, page 624
EE. Dorsal scales usually less than 40 ; nuchal collar broad, with
unbroken light borders ; no light lines on sides of head and
neck; no spots on dorsal scales. .. .<S. bulleri Boulenger, page 579
AA. Supraoculars in two series; or, if in one series, with one or more scales divided.
B. Lateral scales with the terminal mucrone arising well within the free posterior
margin; dorsal scales 41 or more.
C. Dorsal scales, 47 to 54; throat with very distinct, oblique, dark-blue
lines ; head scales not rugose (microscopically) ; oblique dark bands on
sides of body distinct ; maximum snout-vent measurement, 79 mm.
iS. dugesii intermedius (Duges), page 663
Smith: Genus Sceloporus 557
CC. Dorsal scales, 41 to 50; throat without or with very dim oblique lines;
head scales rugose (microscopically) ; oblique dark bands on sides of body
indistinct or absent; maximum snout -vent measurement, 87.5 mm.;
femoral pores, not over 13 S. dugesii dugesii Bocourt, page 657
BB. Lateral scales with the terminal mucrone arising at or very near the free
posterior margin; head scales not rugose (microscopically); no oblique, dark-
blue lines on throat.
C. Dorsal scales, 55 or more iS. ornatus ornatus Baird, page 647
CC. Dorsal scales, less than 55.
D. Dorsal scales, 47 to 53 S. ornatus caeruleus Smith, page 652
DD. Dorsal scales, less than 47.
E. Black nuchal collar six to eight scales broad; supraoculars
usually in two complete rows; irregular dark markings fre-
quently present on throat; dorsals, 34 to 46; series of dorsal
scales never with longitudinal light and dark lines; maximum
snout-vent measurement, 101 mm.
(S. jarrovii minor (Cope), page 631
EE. Black nuchal collar less than six scales broad.
F. Nuchal collar two or three scales broad; general ground
color blue, at least in males; males with the entire belly
black and blue; dorsal scales, 37 to 46; maximum
snout-vent measurement, 87 mm.
S. jarrovii immucronatus Smith, page 640
FF. Nuchal collar more than two or three scales broad;
maximum snout-vent measurement no less than 100 mm.
G. Tail with very distinct, broad, alternating dark
and light bands, most distinct toward tip of tail,
where they are complete ; supraoculars in two
complete rows ; head scales very irregular ; light
borders of nuchal collar broad ; a broad light band
across neck behind occiput ; preocular usually en-
tire; inner row of labiomental scales usually ter-
minating at a point posterior to suture between
second and third infralabials.
S. poinscttii Baird and Girard, page 606
GG. Tail without distinct alternating dark and light
bands of nearly equal width ; bands about tail
not complete toward tip ; dorsal scales, 40 or less.
H. Dorsal scales, 31 or less; two parallel dark
lines down middle of throat (except in adult
males) ; a broad median dark band usually
visible, composed of about five large dim
blotches.
S. mucronatxts mucronatus (Cope), page 583
HH. Dorsal scales, more than 31; or, if less,
without parallel lines down middle of throat ;
without a median dark band.
I. Light borders of nuchal collar usually
unbroken medially ; upper auricular
lobule much enlarged ; each dorsal scale
row usually with a longitudinal light
line ; general dorsal coloration dark ;
maximum snout-vent measurement, 114
mm S. mucronatus omiltemanus
(Giinther), page 591
IL Light borders of nuchal collar usually
broken medially ; auricular lobules not
well differentiated ; no lines on dorsal
scale rows ; general dorsal coloration
very light, usually of a shade of blue ;
maximum snout-vent measurement, 143
mm S. cyanogenys (Cope), page 599
558 The University Science Bulletin
Sceloporus serrifer Cope
(Text figs. 5 and 6; plate XLVII, fig. 2)
Sceloporus serrifer Cope, 1866, pp. 124-125; idem, 1885, p. 401; Boulenger, 1885, p. 221;
Cope, 1887, p. 37; Gunther, 1890, pp. XIII, 77; Boulenger, 1897, p. 488; Cope, 1900, p. 336;
Barbour and Cole, 1906, p. 150; Stejneger, 1918, pp. 91-92.
Sceleporus serrifer Werner, 1896, pp. 346, 352; idem, 1903, p. 344.
Sceloporus torquatus serrifer Stuart, 1934, p. 11 ; idem, 1935, p. 45.
Type locality. Yucatan. Type USNM 24868; apparently seven
paratypes, USNM 10298, 24869-74; all collected by Arthur Schott.
Diagnosis. A member of the torquatus group; dorsal scales, 28
to 35 from occiput to base of tail; femoral pores, eight to fourteen;
supraoculars in a single row, entire, the last occasionally in partial
contact with median head scales; tibia about equal to length of
shielded part of head; length of fourth toe usually greater than
snout-ear measurement; general dorsal coloration light; nuchal
collar complete, light bordered; parietals, interparietal and posterior
part of frontal each with a conspicuous light spot ; several light spots
or bands on neck; each dorsal scale with a black median streak
along the keel in females; throat bluish, reticulated or spotted with
white or cream; middle of belly and usually the chest cream or
white in males.
Description (from E.H.T & H.M.S. 2212, male). Head and
body not flattened; head scales pitted (especially in prefrontal and
internasal regions), not keeled, slightly convex; parietals single on
each side, subtriangular, about one fourth size of interparietal ; latter
hexagonal; frontoparietal single on each side, subrectangular, two
thirds the size of either parietal, separated medially by a small
azygous scale; posterior section of frontal about one fourth size of
anterior section; prefrontals in contact medially, their common
suture about one fourth their maximum length ; prefrontals narrowly
separated from second canthals, slightly less than one half size of
anterior section of frontal; median frontonasal shield-shaped,
slightly larger than either lateral frontonasal; two small, broad,
short scales in front of median frontonasal; these preceded by a
median pair of rather large, long internasals, separated from rostral
by a row of small scales ; nasal small, the naris pierced nearly in the
middle, leaving a narrow border, somewhat wider and tapering an-
teriorly; subnasal small; first canthal larger than second, not forced
above canthal ridge, not in contact with lorilabials, somewhat U-
shaped (probably fused with loreal) ; supraoculars 4-4, separated
from median head scales by a single row of small scales, from super-
ciliaries by a single row of very small scales; preocular divided; sub-
Smith: Genus Sceloporus
559
ocular followed posteriorly by two short, strongly keeled, well-dif-
ferentiated postoculars ; two rows of labiomentals, not reduced below
subocular; three and one half or four and one half supralabials and
five infralabials to a point below middle of eye.
Mental pentagonal, its labial border slightly more than half that
of rostral; mental followed by three or four pairs of postmentals,
the scales of the anterior pair in contact with each other; outer
row of labiomentals separated from mental by a narrow contact of
first postmental and first infralabial; inner row of labiomentals
terminating anteriorly even with the anterior part of third infra-
FiQ. 5. Head scales of Sceloporm serrifer Cope. UMMZ 71771, near Merida,
Yucatan; actual head length, snout to occiput, 15.7 mm.
labial; gular scales smooth, smallest anteriorly, largest laterally near
angle of jaws, with as many as three apical notches; notches more
numerous on posterior gular scales, absent on anterior gular scales.
Ear with four auricular lobules, smooth, rounded or pointed, not
well differentiated, smaller than preceding scales; scales between ear
and lateral nuchal fold subequal in size to those in temporal region,
but more strongly keeled and mucronate; a fold of skin between
nuchal pocket and a point below lower edge of ear, surmounted by
very strongly mucronate scales; dorsal scales weakly keeled, strongly
mucronate, with numerous lateral denticulations; lateral scales equal
to or somewhat smaller than dorsal scales, more strongly keeled,
mucronate and denticulate; ventral abdominal scales smooth,
slightly more than one half the size of median dorsals, each with
a single apical notch; median gular scales about two thirds size of
560 The University Science Bulletin
median abdominals, subequal in size to preanals; subcaudals smooth
except toward tip of tail; postanals enlarged, separated by two small
scales; dorsal caudals toward base of tail somewhat larger than
median dorsals on body.
Dorsal scales of foreleg keeled, mucronate, denticulate, those on
upper foreleg about two thirds size of median dorsals on body and
about one third larger than dorsal scales on lower foreleg; ventral
scales of foreleg smooth, slightly mucronate, those on lower foreleg
somewhat smaller than dorsal scales of same member, much larger
than ventral scales of upper foreleg; lamellar formula for fingers,
10-15-18-19-15 (10-15-20-20-15).
Dorsal scales of hind leg keeled, strongly mucronate, very weakly
denticulate, those on thigh two thirds the size of median dorsals on
back, those on shank slightly smaller than dorsals on body ; ventral
scales on shank smooth, rounded, slightly smaller than dorsals of
same member; ventral scales of thigh smooth, notched, those im-
mediately preceding femoral pores subecjual in size to preanal scales;
scales on posterior surface of thigh strongly keeled, strongly mucro-
nate, denticulate, nearly twice as large as preanal scales, abruptly
decreasing in size near femoral pores; no postfemoral dermal pocket;
lamellar formula for toes, 9-15-19-24-17 (9-15-19-23-17).
Color. Black nuchal collar broad, about four scales wide in
median dorsal line, continued across throat in males; collar bordered
on either side by a light band about one scale wide, the posterior of
which is complete and passes onto the foreleg near its insertion; the
anterior light border broken medially, terminating laterally at the
upper edge of the lateral nuchal fold; keels on dorsal scales black,
producing the effect of a narrow dark line down the middle of each
scale row; a transverse series of light spots across neck between
ears; a light spot in the center of the interparietal and each parietal;
a narrow light band across the middle of the orbits (may be repre-
sented by only a light spot on the posterior section of the frontal) ;
a light band across the prefrontals (very dimly visible in males) ;
tail dimly banded with light brown and olive.
Males uniform olive or brownish olive on back and limbs; throat
stippled heavily with blue and black, increasing in intensity toward
gular fold region, which is black; ventral surfaces of limbs, chest
and a narrow median abdominal band white; sides of belly caerulean
blue, tinged with purple; a rather broad black band (about three
scales wide) bordering lateral blue areas, involving groin, rarely
reaching to black of gular region.
Smith: Genus Scelopokus 561
Females brownish olive on back and limbs, with scattered ir-
regular dark-brown spots; ventral surfaces uniform whitish.
Variation. The variation in head scales of fifty-three specimens
is as follows: Parietal one third to one fifth size of interparietal;
frontoparietals divided on one side in one; frontal touches inter-
parietal in twenty-three ; frontoparietals contact medially in twenty-
one; an azygous scale separates frontoparietals medially in nine;
entire frontal longitudinally divided in three specimens; anterior
section of frontal divided longitudinally in two specimens; frontal
divided transversely in two places in one specimen; frontal entire
in six specimens; prefrontals contact medially in forty-two; median
frontonasal contacts frontal in six; an azygous scale separates them
in five; the supraoculars are entire in all, usually four on each side,
sometimes five; usually but a single row of scales separates the
supraoculars from the superciliaries ; in a few specimens one or two
extra scales of small size are present; the last supraocular is in con-
tact with the median scales on one side in nine specimens, on both
sides in two; canthals regularly 2-2; first canthal forced above the
canthal ridge in one specimen, in contact with lorilabials in six;
preocular divided on one side in three specimens, on both sides in
thirty-five; lorilabials reduced to one row below subocular on one
side in nine specimens, on both sides in twelve; the outer row of
labiomentals contacts the mental in four specimens; the inner row
•terminates at some point between the anterior part of the third in-
fralabial and the anterior part of the second infralabial; auricular
lobules three to five, relatively small.
The dorsal scales from occiput to base of tail vary between 28 and
35. The scales average somewhat larger in the specimens from
Yucatan. The specimens from La Primavera, Guatemala, have a
larger average number of scales. These specimens differ also from
the others examined in possessing a larger average number of
femoral pores, an azygous scale regularly present between the
frontoparietals, an azygous scale much more frequently present be-
tween the prefrontals than usual, and the inner row of labiomentals
terminating farther forward (even with the anterior part of the
second infralabial) than usual.
The femoral pores in the entire series vary from eight to fourteen ;
specimens from Yucatan have the lowest average number, while the
specimens from La Primavera, Guatemala, have the highest average
number.
A peculiar male specimen in the National INIuseum (No. 46861),
from Chiapas, Mexico, collected by Nelson and Goldman, is referred
3&— 6037
562
The University Science Bulletin
to this species. It is quite possible that it represents an iindescribed
species; however, since certain head scales are obviously abnormal,
it seems unwise to consider the specimen as representative of a new
species. Its characters are as follows: snout to vent, 77 mm.; tail,
112 mm.; snout to occiput, 15.9 mm.; snout to ear, 19.9 mm.; breadth
of head, 11.8 mm.; hind leg, 55.5 mm.; tibia, 15.5 mm.; fourth toe,
21.7 mm.; fifth toe, 12 mm.; lamellae fourth toe, 21-22; femoral
pores, 13-14; dorsals, 31; ventrals, 47; scales around body, 36.
Frontoparietals not divided ; parietal one fourth size of interparietal ;
frontal touches interparietal; frontal not divided; supraoculars 4-4,
Serrifer
Fig. 6. Distribution of Sceloporus serrifer Cope.
three divided on each side; one row of small scales between supra-
oculars and superciliaries; supraoculars separated from median head
scales; frontal contacts median frontonasal; latter separated on
each side from lateral frontonasals by a small scale; canthals 2-2,
normal; preocular divided; two rows of lorilabials, complete below
subocular; outer row of labiomentals separated from mental; inner
row of labiomentals terminating anteriorly even with posterior part
of second infralabial.
No distinct light marks on head, except pineal spot; ventral color-
ation exactly as in typical serrifer.
Habits and habitat. Stuart (1935, p. 45) records the following
notes on specimens collected at Zotz, Peten, Guatemala: "Many
were seen climbing about the rafters and thatched roofs of several
Smith: Genus Sceloporus
563
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564 The University Science Bulletin
deserted houses in the small clearing. One of the adults was taken,
ten feet from the ground, on the trunk of a large tree, and another
which was frightened readily ascended a tree to the height of about
forty feet. In actions the form appears much like Sceloporus magis-
ter of the southwestern United States; as an adult it is probably
largely arboreal." This arboreal tendency is of considerable interest.
So far as I am aware, no other species of the torquatus group shows
the same tendency.
Distribution. Along the eastern coastal plains of Mexico and
Guatemala south from southern Tamaulipas.
Locality records. Tamaulipas: Chocoy (M. C. Z. 17943, W. W.
Brown). Vera Cruz: Papantla (U. S. N. M. 47353, E. A. Gold-
man); four miles east of Encero (E. H. T. & H. M. S. 2210-2).
Yucatan: (Type locality) (Cope, 1866, 1885, 1887; U. S. N. M.
10298, 24868-74); Chichen-Itza (Barbour and Cole, 1906); near
Merida (UMMZ 72881-90, Edwin Creaser; UMMZ 71771 [6
specimens] ; MCZ 6269 [16] ) ; Tekanto (ANSP 8500). Guatemala:
Zotz, Peten (Stuart, 1935; UMMZ 74956 [12 specimens], L. C.
Stuart); La Primavera (MCZ 28153-9, A. W. Anthony, 2500 ft.).
Remarks. The records of Werner (1896) of serrifer from Guate-
mala and Honduras may be doubted. His record from Guatemala
was later (Werner, 1903) corrected.
Sceloporus torquatus torquatus Wiegmann
(Text Figs. 7 and 8; Plate XLVIII, Fig. 1)
(The following synonymy omits only those references to torquatus
which are quite certainly wholly applicable to other species. The
subspecies torquatus melanogaster has been included in much of the
material concerning which the following papers have been written,
but the uselessness of any attempt to allocate any of these with the
synonymy of melanogaster, without examination of the specimens,
is obvious.)
Sceloporus torquatus Wiegmann, 1828, p. 369; idem, 1834, p. 49, pi. 7, fig. 1; Fitzinger,
1843, p. 75; Miiller, 1865, p. 601; Sumichrast, 1873, p. 243; Bocourt, 1874, pp. 171-173,
pi. 18, figs. 9, 9a, 9b, 9c (part ?) ; ? Cope, 1875, p. 48; idem, 1879, p. 265; Sumichrast,
1882, p. 37 (part ?); Boulenger, 1885, p. 219; Cope, 1885, pp. 401-403 (part); idem, 1887,
p. 37; Duges, 1887, pp. 113-114; Carman, 1887, p. 14; Herrera, 1889, p. 331; Gunther,
1890, pp. XII, 66; Boettger, 1893, p. 64; Herrera, 1895, p. 18; Duges, 1896, p. 479;
Boulenger, 1897, pp. 477-481, text fig. 1 (part); Mocquard, 1899, p. 156; ? Cope, 1900,
pp. 336, 340, 347-350, text fig. 50; Herrera, 1904, p. 18; Gadow, 1905, pp. 194, 214 (part ?).
Tropidurus torquatus Wagler, 1830, p. 146.
Aga7na torquata Peale and Green, 1830, pp. 231-232; Garman, 1884, p. 17.
Tropidolepis torquatus Gray, 1831, p. 43; Dumeril and Bibron, Vol. IV, 1837, pp. 301-
303; Gray, 1839, p. 95; idem, 1845, p. 208; Aug. Dumeril, 1851, p. 77; Duges, 1870, p. 243.
Smith: Genus Sceloporus
565
Sceloporus ferrariperezi Cope, 1885, p. 400 (part) ; idem, 1887, p. 37 ; Gunther, 1890, pp.
XIII, 77; Cope, 1900, p. 335.
Sceloporus torquatus torquntus ? Cope, 1885, pp. 402, 403; idem, 1887, p. 37; Mertens,
1930, p. 158.
Sceloporus torquatus ferrariperezi Boulenger, 1885, pp. 220-221.
Type locality. Mexico.
Description. Head scales smooth; supraoculars in a single row,
rarely (about five percent of the specimens) with the outer third
or fourth of two or three separated off; two to three incomplete
rows of scales separating the supraoculars from the superciliaries;
one row of elongate to squarish scales between supraoculars, the
first supraocular rarely touching narrowly the median head scales
(2 percent) ; frontal normally contacting interparietal (91 percent) ;
Fig. 7. Head scales of Sceloporus torquatus torquatus Wiegmann. EHT &
HMS 3569, near Tres Cumbres (Tres Marias), Morelos; actual head length,
snout to occiput, 16.3 mm.
canthals normally two (96 percent), the first frequently (about 60
percent) above the canthus, the second canthal and subnasal con-
tacting; median frontonasal usually (66 percent) contacting the
frontal, or separated from it by a small median scale between the
two prefrontals ; the lateral series of scales intercalated between the
series of postmentals and the infralabials frequently (about 45 per-
cent) touching the mental anteriorly; lobules on anterior margin
of ear usually larger than those preceding, usually three in number,
the upper two largest; median dorsal scales 26 to 30 from occiput
to base of tail, in parallel series, smooth or weakly keeled, rounded
or weakly mucronate ; lateral dorsals somewhat larger than median
dorsals, more strongly keeled and mucronate; femoral pores, 14-23
(average 18) on each side, the two series closely approximated
medially; tibia approximately equal to length of shielded part of
head; length of fourth toe usually a little less than distance from
snout to posterior margin of ear, rarely slightly more.
566 The University Science Bulletin
Black nuchal collar always present, four or five scales long on
the middorsal line; light borders of black collar one scale wide;
posterior light border frequently narrowly interrupted medially, ex-
tending laterally onto the proximal end of the humerus; anterior
light border usually interrupted medially, extending laterally a
variable distance, sometimes to the posterior border of the ear, pass-
ing along the crest of the lateral nuchal fold, sometimes terminating
at a point dorsal and anterior to the insertion of the foreleg; the
anterior light border is sometimes horseshoe shaped, and extends
anteriorly on each side to a point above the ear, becoming quite dim
anteriorly; light borders variable in color, sometimes cream, some-
times blue or green, occasionally strongly tinged with orange ; a few
irregular light spots frequently present on neck; upper labial region
occasionally dimly barred; back uniform olive gray, drab, olive, or
blue-gray, varying according to the time elapsed since the last
ecclysis; spots on back present only in some very young specimens,
and when present, large and indistinct; anteroventral surface of
femur usually coarsely reticulated; dorsal surface of limbs rarely
banded, light bands dim and narrow when present, bluish in color;
posterior surface of femur usually traversed by a light band, usually
rather dim; tail with numerous, narrow, indistinct light bands,
bluish or cream in color.
Ventral gular and chin region irregularly spotted or reticulated,
all light areas disappearing and replaced by black or blue in adult
males; no longitudinal pair of dark lines near the midventral line
on the throat ever present. In some adult males, the entire ventral
surface of head, body, and tail is black, with the exception of cream-
colored areas on the tail and femur. In other large males the throat
is pale, iridescent blue, becoming azure blue toward the neck; the
neck is black, and this color is continuous with a broad, median
ventral black band, occupying the chest, a wide area in the middle
of the abdomen, and the groin. Usually some pale blue or cream-
colored areas are visible in the middle of the abdomen. The sides
of the belly are azure blue, with a generous tinge of companula blue.
In females, the belly may become drab-gray, usually darker
toward the sides, and the throat usually remains gray-blue with
numerous white flecks or reticulations.
Variation. The entire series is veiy uniform in scalation and
proportions. The more important variations are given in the diag-
nosis. In addition, the preocular is separated from the series of
scales above the supralabials in about thirty-seven percent of the
Smith: Genus Sceloporus 567
specimens; the anterior frontal is divided in about ten percent. In
one' specimen the frontal is not divided. Other obvious fusions of
head scales rarely occur.
The coloration is very constant, within limits of variation given
in the diagnosis.
Remarks. Bocourt (1874, pi. 18, fig. 9) shows the details of the
dorsal head scales of Wiegmann's type of torquatus. The supra-
ocular scales are exactly as in the specimens studied, in a single row,
with one complete and another incomplete row of scales between
them and the superciliaries. The frontal contacts the interparietal,
and there is an azygous scale between the two prefrontals.
Sceloporus torquatus has been recorded from the states of Chiapas
(Sumichrast, 1882), Chihuahua (Baird, 1859), Coahuila (Garman,
1887), Colima (Bocourt, 1874; Gadow, 1905), Durango (Boulenger,
1897), Guanajuato (Duges, 1870, 1896; Boulenger, 1897), Guerrero
(Boulenger, 1897; Gadow, 1905), Jalisco (Giinther, 1890; Duges,
1896; Boulenger, 1897; Mocquard, 1899), Mexico (Peale and Green,
1830; Duges, 1887, 1896; Herrera, 1889; Giinther, 1890; Boulenger,
1897; Gadow, 1905; Mertens, 1930), Nuevo Leon (Yarrow, 1883;
Cope, 1885), Oaxaca (Bocourt, 1874; Sumichrast, 1882), Sonora
(Yarrow, 1883), Tamaulipas (Baird, 1859), Vera Cruz (Sumichrast,
1882; Cope, 1885, 1887, 1900; Boulenger, 1897) and Zacatecas
(Duges, 1896). Cope (1888) records torquatus from San Diego,
Texas.
It is quite certain that some of these records are incorrect. Those
from Chihuahua, Coahuila, Durango, Nuevo Leon, Sonora and Ta-
maulipas are almost certainly of other species. Those from Chiapas,
Oaxaca and Zacatecas are doubtful, as are also some of those from
Colima, Guanajuato, Guerrero, Jalisco and Vera Cruz. Cope's rec-
ord from San Diego is obviously in error, probably referable to
'poinsettii.
Boulenger's (1897) records are obviously composites, as he
lumped nearly all the apparently related forms, recognizing only
poinsettii, and part of these specimens are of other forms. In his
table A (single row of enlarged supraoculars) those specimens from
Guadalajara, Guanajuato and Rio Santiago have characters of tor-
quatus melanogaster; the one from El Paso, collected by Forrer,
probably has incorrect locality' data (Forrer's specimens are notable
for incorrect locality data) ; and the remainder appear to be the
only torquatus torquatus he had (seven specimens). In table B, the
poinsettii from Duvall county, Texas, is peculiar in having such a
568
The University Science Bulletin
small number of scales from occiput to base of tail ; the locality may-
be incorrect. The specimen in table B from Mexico is possibly mu-
cronatus mucronatus. In table C, presumably consisting entirely of
poinsettii, only those from Duvall county, Texas, seem to have char-
acters of poinsettii; the others are very likely of other species, prob-
ably m. mucronatus and m. omiltemanus.
Habits and habitat. S. torquatus torquatus seems to be confined
to areas of high elevation. Specimens I have seen or collected have
been within the zone of evergreens.
Torcjuatus . . o
Melanoqaster . •
BuUcri ... A
<:'WJ^':^A:y:K-iM^'W'
Fig. 8. Distribution of Sceloporus torquatus torquatus Wiegmann, S. t. melano-
gaster (Cope) and S. bulleri Boulenger.
In the American Museum of Natural History are eight specimens
(Nos. 15540-7) which were "born in bag before May 18, 1919." The
female (or females) which gave birth to the young were collected
two miles wTst of Tacuba, D. F., by Paul D. R. R. Riithling, on
May 16, 1919.
Range. Central Mexico, including Hidalgo, western central Vera
Cruz, Mexico, Distrito Federal, northern Puebla, eastern Morelos,
southern Guanajuato and northern Michoacan.
Locality records. Hidalgo: Twenty-five miles south of Jacala
(EHT & HMS 601-604) ; Mexico: Two miles south of San INIartin
(EHT & HMS 3842) ; Valle de Mexico, 7,700 ft. (MCZ 33902,
W. W. Brown) ; TIalpam (USNIM 47769, E. W. Nelson) ; San Juan
Teotihuacan (MCZ 16063—6, E. R. Dunn; UMMZ 63937, H. B.
Smith: Genus Sceloporus
569
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Smith: Genus Sceloporus 571
Baker) ; Distrito Federal: (AMNH 15490, Riithling) ; Coyoacan
(MCZ 9551, 9597); Pedregal de San Angel (MCZ 16090, E. R.
Dunn) ; two miles west of Tacuba (AMNH 15540-7, Paul D. R.
Riithling) ; three miles west of Tacuba (AMNH 15582, Riithling) ;
San Juanico (AMNH 15552-4, 15592-3, 15592-7, 15603, 15605,
15608-9, 15612, Riithling) ; North of Guadelupe (AMNH 15604-5,
15607, 15610, 15611, 15613, Riithling) ; east of San Mateo Chalpa
(AMNH 15590, Riithling) ; M'exico City (USNM 12719, A. Her-
rera) ; west of Atzcoalco (AMNH 18473, Riithling) ; Michoacdn:
near Uruapan (EHT & HMS 3384-90, 3765-97) ; near Zinapecuaro
(EHT & HMS 3657, 3667, 3672-3) ; near Lake Cuitzeo (EHT &
HMS 3723); Querendaro (USNM 47162, 47233, E. W. Nelson);
three miles north of Maravatio (EHT & HMS 3733-39) ; Patzcuaro
(FMNH 996 [4 spec], 1399, S. E. Meek; USNM 20159, 20139-44,
P. L. Jouy; USNM 47222, E. W. Nelson; ANSP 15358, S. N.
Rhoads); Tupataro (USNM 10238, 10244, 10232, 10235, 10252,
Duges) ; Morelos: near K63, on highway between Mexico City and
Cuernavaca (EHT & HMS 1715, 1747, 1753) ; five kilometers south
of Tres Cumbres (Tres Marias) (EHT & HMS 1689, 1691, 1693-5) ;
near Tres Cumbres (EHT & HMS 616-624, 627-629, 3557, 3559-
3581, 5115-5118). Puebla: near Rio Frio (EHT & HMS 3349);
Atlixco (USNM 47034, E. W. Nelson). Vera Cruz: near Toxt-
lacuaya, about eighteen miles west of Jalapa (EHT & HMS 1860).
Indefinite localities: Mexico (USNM 12095; ANSP 8499 [type of
torquatus Peale and Green] ; AMNH 15601-2) ; no data (AMNH
15600).
Sceloporus torquatus melanogaster (Cope)
(Text Figs. 1 and 8; Plate XL VIII, Fig. 2)
Sceloponis melanogaster Cope, 1885, pp. 400-401; idem, 1887, p. 37; Duges, 1887, pp.
114-115, pi. 12, fig. 7; Herrera, 1889, p. 332; Gunther, 1890, pp. xiii, 77; Herrera, 1895,
p. 18; Cope, 1900, p. 335; Herrera, 1904, p. IS.
Sceloporus torquatus m,elanogaster Boulenger, 1885, p. 220; Boettger, 1893, p. 64.
Type locality. Cope (1885) says that the type was "probably
from Guanajuato." Duges (1887, p. 114) states: "Este esceloporo
no es de Guanajuato, como lo piensa el autor, porque yo fui el
que se lo remitio sin indicacion de localidad; todos los que he visto
venian de Mexico y la Noria (Michoacan) , 6 de Tupataro, cerca de
Cueramaro." Type USNM 9877, Duges collector.
Discussion. S. t. inelanogaster shows no important scale differ-
ences from torquatus torquatus. It is notable that the dorsal scales
of the latter form average very slightly smaller, are frequently al-
most smooth and rounded, and never more than weakly mucronate.
572 The University Science Bulletin
The supraoculars are smaller, being bounded laterally by never less
than two, and sometimes three, incomplete rows of scales, and
bounded medially by a row of scales more or less square in shape.
In torquatus melanogaster, on the other hand, the dorsal scales aver-
age slightly larger and are more strongly keeled and mucronate than
in t. torquatus. The supraoculars are larger, occasionally separated
from the superciliaries by only one row of scales. The series of
scales bounding the supraoculars medially are usually more elongate
than in t. torquatus. However, so much variation occurs in these
characters that they are useless for diagnosis ; they are, however, of
interest in general tendencies of variation, supplementing the other
more diagnostic features.
In maximum size attained, torquatus melanogaster approaches
more closely to cyanogenys than to any other form of the torquatus
group ; it reaches 129 mm. snout to vent, while t. torquatus reaches
only 98 mm.
Many very striking differences in coloration are apparent. The
name is quite misleading, as the belly is not as black in males of
t. melanogaster as in t. torquatus. The color variation is as follows:
The black nuchal collar is not over two or three scales wide; the
light borders are cream-colored, with sometimes a tinge of blue.
The posterior light border is not over one scale wide, and may be
quite indistinct in some specimens; it is not reduced, however, by
breaking into spots, but by general decrease in distinctness. The
anterior light border is quite variable, and does not extend laterally
beyond a point approximately even with a line drawn posteriorly
from the superciliaries. At this point it is usually continuous with
a light band about one scale wide which passes anteriorly to the
upper edge of the ear. This band, although sometimes quite irregu-
lar, is quite characteristic. At the point where this light band con-
nects with the black nuchal collar, the latter is narrowed consider-
ably and frequently completely broken. The area between these
two narrow places in the black collar may be of the same color as
the back, the collar confined to the shoulder region. A few light
flecks usually occur in the neck region. A dark band, usually
visible, passes from the upper margin of the eye to the upper margin
of the ear, and is bordered below by a light band which passes
underneath the eye.
The back is drab, drab-gray or glaucous-blue. A series of dim
dark spots are usually present on the back, either scattered irregu-
larly or arranged in two longitudinal rows, one on either side of
the middorsal line. A few dim light spots are usually scattered
Smith: Genus Sceloporus 573
irregularly over the back. The limbs are not or but very dimly
banded. The tail has dim, narrow, light bands, more distinct toward
the tip of the tail.
In males, the throat is blue-gray or indigo blue, with lighter blue
or whitish flecks scattered irregularly over the throat. The black
nuchal collar frequently is complete ventrally. When complete, its
color is continuous ventrally with a black area covering the chest,
occupying a broad median area of the abdomen and extending over
the groin and onto the ventral surface of the femur. When incom-
plete, the chest, median abdominal area and the ventral surfaces of
the femora are cream. A narrow black line begins on each side of
the belly opposite the axilla and extends posteriorly, covering the
groin. The sides of the belly are usually pale blue, becoming cam-
panula blue near the black borders. In lighter specimens the sides
of the belly are pearl-blue.
Females have the throat much as the males, but less brilliant; the
abdomen is usually uniform cream, but may have, in very large
specimens, faint traces of the markings of the male. A few dim,
bluish spots are usually present on the chest, and occasionally are
present on the abdomen.
The very young specimens have much the same dorsal coloration
of the adults, except that the pattern of dark and light spots is quite
distinct. The belly is cream, and the throat is coarsely, and usually
very distinctly, reticulated with bluish.
Remarks. The distinctness of melanogaster is quite striking, de-
spite its close relationship with t. torquatus. The incomplete black
nuchal collar, large size, light color and general spinosity give the
general impression of the spinosus group.
Specimens intermediate between t. melanogaster and t. torquatus
are available from a locality three miles north of Maravatio, Micho-
acan. In this series are two specimens which, if the black nuchal
collar were broader, would be typical t. torquatus; in the others, the
collar is broken or quite narrow laterally. In the adults of the
latter, however, the belly is much blacker than it is in typical
t. melanogaster. Specimens from other points in Michoacan seem
to have affinities for t. torquatus, and also some from parts of
Guanajuato, but the remainder are quite distinct.
It is notable that Boulenger's (1897) largest specimens of torqua-
tus from Mexico are either from areas of intergradation between
t. torquatus and t. melanogaster, or are well within the range of the
latter.
574 The University Science Bulletin
Two female specimens in the United States National Museum
(Nos. 46739-40) from Miquihuana, Nuevo Leon, collected by Nelson
and Goldman, are referred to this species despite certain differences
in coloration and femoral pore count. It is possible that these repre-
sent a race which has not been described, but for the present I prefer
to retain them under the name melanogaster until further material
is available. The specimens have the following characters (46739
and 46740, respectively): snout to vent, 79.0 and 73.5mm.; tail,
115 mm. (46739) ; snout to occiput, 16.3 and 15 mm.; snout to ear,
20.1 and 18.5 mm.; breadth of head, 11.8 and 11.2 mm.; hind leg,
46.5 and 46.2 mm.; tibia, 14 and 15 mm.; fourth toe, 17.8 and
17 mm.; fifth toe, 9.3 and 9 mm.; lamellae fourth toe, 19-19 and
19-19; femoral pores, 13-13 and 12-13; dorsals, 26 and 26; ventrals,
47 and 42; scales around body, 37 and 33; scales to head length,
6.3 and 5; parietal one third size of interparietal; frontal touches
interparietal; 3-4 and 2-3 small scales segmented off outer edges of
supraoculars; one row of scales between supraoculars and super-
ciliaries; none of supraoculars in contact with median scales; pre-
frontals in contact; two canthals, the first forced above the canthal
ridge on both sides in 46739; preocular divided in 47640; rows of
lorilabials reduced to one by a scale below subocular in 46739 ; outer
row of labiomentals separated from mental; inner row of labio-
mentals terminating anteriorly even with the suture of the second
and third infralabial.
The black nuchal collar is three or four scales broad, unbroken,
with light anterior and posterior borders about one scale wide,
broken medially; a few light flecks on nape of neck; a light line
from subocular region to ear; another light line from posterior cor-
ner of eye to nuchal collar; limbs with indistinct bands. Ground
color brown-olive, without darker markings in 46740; in 46739, the
back has about four indistinct, broad, dark crossbands; the tail has
narrow, distinct light bands, about nineteen in number.
Another specimen from Chapala, Jalisco (EHT & HMS 2832),
assigned to this subspecies, combines certain characters of melano-
gaster and dugesii dugesii. The dorsals from occiput to base of tail
are 38, the femoral pores are 12-13, and the lateral scales have the
terminal mucrone arising slightly within the free posterior border
of the scales ; in these characters it approaches dugesii dugesii. The
specimen ( ? ) measures 95 mm. from snout to vent and the supra-
ocular scales are entire ; in these characters the specimen approaches
melanogaster. The general dorsal coloration is more suggestive of
Smith: Genus Sceloporus 575
d. dugesii, while the ventral coloration and character of the nuchal
collar suggests melanogaster. It is quite possible that the specimen
represents a new species. However, because of the possibility of
hybridization, I prefer to consider the specimen an aberrant melano-
gaster, pending the accumulation of further material. Since typical
dugesii dugesii and typical torquatus melanogaster are available
from the same locality, it seems quite unlikely that this specimen
indicates intergradation.
Five specimens collected by W. B. Richardson in 1893 (INICZ) and
1895 (UMMZ) at Matagalpa, Nicaragua (MCZ 6443 [4 specimens] ;
UMMZ 65269), are referred to melanogaster, with the assumption
that the locality data accompanying the specimens are incorrect. A
very careful comparison of these specimens with specimens of mel-
anogaster from central Mexico has revealed no differences whatso-
ever in scale characters, with the exception that one specimen
(UMMZ 65269) has a greater number of scales from occiput to base
of tail (31) than normally occurs. In coloration, however, the
males differ from typical melanogaster in having the ventral mark-
ings confined to the throat and sides of the belly ; normally the chest
and middle of the belly have dark markings. A young male
(64 mm. snout to vent) has well-defined lateral abdominal mark-
ings. Normally no distinct markings appear in melanogaster of
that size. The ventral scales have slipped, however, and it is as-
sumed that this accounts for the prominence of the markings on the
sides of the abdomen. The adult male (UM:\IZ 65269 ; 89.5 mm.
snout to vent) has the lateral abdominal markings widely separated
from each other, a condition not observed in any other specimen of
melanogaster. It may be noted that this type of ventral coloration,
with regard to the lateral abdominal markings, is typical of serrifer
and related species. These specimens, however, are widely different
from serrifer in many respects.
Cope's five cotypes (USNM 9874, 9876, 9878, 9880, 9895) of
fcrrariperezi purport to be from Guanajuato. If they came from
that state, it seems likely that they are from the southern portion,
in an area in which melanogaster intergrades with torquatus. Since
Cope's specimens approach more closely in all characters to tor-
quatus than to the northern subspecies, ferrariperezi is here consid-
ered a synonym of torquatus. The type of melanogaster (USNM
9877) is quite typical of the northern subspecies.
The haste with which Cope examined his types of jerrariperezi is
indicated by the fact that one of them (USNM 9880) is a typical
female spinosus spinosus.
576 The University Science Bulletin
Habits and habitat. Mr. Dunkle and I found specimens of mel-
anogaster in abundance on rock fences in central Zacatecas. Certain
great areas in this region are level and devoid of cliffs and canyons
which the lizards probably would otherwise inhabit; as a result, the
occasional rock fences were almost overcrowded. At La Colorada,
Zacatecas, specimens were found in the long, deep canyons issuing
from the mountains west of the town. The adults were extremely
wary and difficult to approach within range of shot shells. Young
specimens, born in the spring, were much more frequently seen than
the adults.
Range. Northern Jalisco, west through all of Guanajuato except
the extreme southern part, north through central and southern San
Luis Potosi and Zacatecas. (See Fig. 8 for distributional map.)
Locality records. Aguascalientes: (FMNH 1654, C. H. T. Town-
send) ; ten miles south of San Francisco (DHD &. HMS 839).
Guanajuato: (USNM 9877, 26137, Duges) ; Celaya (FMNH 997
[2 spec], S. E. Meek) ; Santa Rosa (USNM 47780, Nelson & Gold-
man) ; three miles northeast of Santa Rosa (EHT & HMS 3760-5,
3773; near Acambaro (EHT & HMS 3967, 4000, 4002-6, 4008-29) ;
near San Felipe (EHT & HMS 4213-4227) ; near El Terrero (EHT
k HMS 4274, 4275). Jalisco: (USNM 58355, J. Hurtcr) ; Calotlan
(USNM 46936-41, Nelson & Goldman) ; ten miles west of Ojuelos
(DHD & HMS 1109) ; near Chapala (EHT & HMS 2831, 3064-9;
USNM 18971, 18975-6, P. L. Jouy) ; Ocotlan (FMNH 998a, 999,
S. E. Meek) ; Florencia (USNM 46931, Nelson & Goldman) ; Zapot-
lan (FMNH 1669 [3 spec], C. H. T. Townsend) ; near Jamay
(AMNH 17983, 18420-9, 18474-5, Paul D. R. Riithling) ; Juan-
acatlan (USNM 18972-4, P. L. Jouy) ; Atemajac (USNM 47891,
E. W. Nelson) ; Lagos (USNM 47039, Nelson & Goldman). Mich-
oacdn: Querendaro (USNM 47162, E. AV. Nelson). San Luis
Potosi: (MCZ 4548, Dr. Edw. Palmer) ; Charcas (UMMZ 77267-
77271, 77345 [27 spec], C. L. Lundell) ; near San Luis Potosi
(MCZ 4548 [2 spec]. Dr. Edw. Palmer) ; Morales (MCZ [1 spec],
W. W. Brown). Zacatecas: near Tiburcio (DHD & HMS 422-4,
430-454); near La Colorada (DHD & HMS 690-712, 826-836);
Berriozabal (USNM 47041-2, Nelson & Goldman) ; Valparaiso Mts.
(USNM 47871, Nelson & Goldman). Indefinite localities: Mexico
(AMNH 17963, 17986-7, 18452; USNM 11005; USNM 25364,
Duges); Guadelupe (MCZ 6807 [2 spec]. Comision geografico ex-
ploradora) ; Nochitongo Ditch (USNM 19004-5, J. T. Scovell & A.
J. Woolman).
Smith: Genus Sceloporus
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Smith: Genus Sceloporus
579
Several specimens (Nos. 410-415) in the American Museum of
Natural History purport to be from Real de Pinos, Gulf of Cali-
fornia, collected by L. Diguet. The locality is undoubtedly in-
correct.
Sceloporus bulleri Boulenger
(Text Figs. 8 and 9; Plate XLIX, Fig. 1)
Sceloporvs bulleri Boulenger, 1894, pp. 729-730, pi. 48, fig. 3; idem, 1897, pp. 481-482.
Type locality. La Cumbre de los Arrastrados, Jalisco, Mexico.
Diagnosis. A member of the torquatus group; supraoculars large,
in a single row; femoral pores, 14 to 19; dorsal scales, 33 to 41;
Fig. 9. Head scales of Sceloporus bulleri Boulenger. USNM 64655, San
Sebastian, Jalisco; actual head length, snout to occiput, 19.5 mm.
fourth toe slightly less than snout-ear measurement; tibia slightly
shorter than snout-occiput measurement ; nuchal collar three or four
scales wide, uninterrupted, with light anterior and posterior borders
about one scale wide, uninterrupted; nuchal collar complete about
neck in large males; sides of belly in males dark blue, black-edged.
Description (from USNM 64654, male). Head scales slightly
rugose; interparietal nearly straight-sided, pentagonal; parietal
single on each side, subtriangular, slightly less than one third size
of interparietal; frontoparietals rectangular, narrowly in contact
medially; frontal divided, the posterior section slightly more than
half the size of the anterior; prefrontals narrowly in contact medi-
ally; three frontonasals, the median almost twice as large as either
580 The University Science Bulletin
lateral; internasals irregular; nasal small, round, naris pierced in
middle; supraoculars five-five, separated from the superciliaries by
a single row of small scales, and from median head scales by another
row of small scales; canthals two-two, normal; subnasal present,
large; three small loreals on one side, one on other; preocular di-
vided; two postoculars, strongly keeled; two rows of lorilabials, re-
duced on one side to one row below posterior part of subocular; ap-
proximately four supralabials and five infralabials to a point below
middle of eye.
Mental more or less rounded in outline, its posterior extremity
pointed, with a labial border slightly more than half that of rostral;
inner row of labiomentals separated from mental by a narrow con-
tact of first postmental and first infralabial; inner row of labio-
mentals terminating anteriorly below anterior half of third infra-
labial; three pairs of well differentiated postmentals, the scales of
the anterior pair in contact medially; median and anterior gular
scales slightly smaller than lateral gular scales; median and pos-
terior gular scales with one, two or three notches on free border.
Three well-differentiated auricular lobules, smooth, the lower scale
pointed, the largest subequal in size to temporal scales; temporal
scales weakly keeled, weakly mucronate, about one third larger than
scales between ear and lateral nuchal fold ; a weak dermal fold pass-
ing from upper edge of lateral nuchal fold to lower edge of ear, sur-
mounted by strongly keeled, very strongly mucronate scales; skin
in nuchal fold without scales or granules.
Dorsal scales, except those immediately behind head, keeled and
with a moderately long apical mucrone and as many as four lateral
mucrones on each side of the apical mucrone; lateral body scales
more strongly keeled and mucronate than median dorsals, the larg-
est subequal in size to dorsals; dorsal scales between hind legs
slightly smaller than median dorsals; ventral abdominal scales
about one third size of median dorsals, smooth, rounded or with a
single apical notch; scales in gular fold region with as many as six
apical notches ; preanal scales about one third size of median ventral
abdominals.
Dorsal scales of foreleg keeled and mucronate, with several lat-
eral mucrones; dorsal scales of upper foreleg two thirds size of dor-
sal scales on body, about a third larger than dorsals of lower fore-
leg; scales on posterior surface of lower foreleg nearly as large as
dorsal scales of upper foreleg; ventral scales of lower foreleg sub-
equal in size to dorsals of same member, smooth, notched, about
Smith: Genus Sceloporus 581
twice as large as ventral scales of upper foreleg; scales in axilla
squamous; lamellar formula for fingers 9-7-18-17-13 (8-14-17-17-
13).
Dorsal scales of hind leg keeled and mucronate, those of thigh
about two thirds size of median dorsals on body, those of shank a
third larger than the former; ventral scales of tibia subequal in size
to dorsals; scales about tibiometatarsal joint, except ventral scales,
about one fourth size of dorsals on tibia; scales on anterior surface
of femur smooth, rounded, subequal in size to dorsal scales of same
member; ventral scales on femur about half as large as dorsal scales
of same member, subequal in size to preanal scales; scales on pos-
terior surface of femur subequal in size to preanal scales, strongly-
keeled and mucronate, strongly differentiated from dorsal scales of
femur, abruptly decreasing in size posterior to femoral pore series;
no postfemoral dermal pocket; lamellar formula for fingers 8-13-
19-20-15 (8-13-17-21-15).
Dorsal caudal scales but slightly larger than dorsals on body;
subcaudals smooth only near base of tail; enlarged postanals pres-
ent, separated from each other by two small scales.
Color. General ground color, olive, darker on sides of body; no
definite markings on back or limbs; nuchal collar three to four
scales broad, continuous about neck in males; light borders about
one scale wide, uninterrupted, the anterior border terminating at
upper margin of lateral nuchal fold; posterior light border passing
onto extreme proximal end of upper foreleg; no other markings on
neck nor on sides of head; throat slate, with central area light blue
in color in both males and females, but more distinct in the former;
sides of belly blue in males, bordered medially by black, which in
large males covers the chest; a small area in groin black; females
with sides of belly tinged with blue and with a few irregular black
areas on chest; in females a few white scales on throat; remainder
of belly and ventral surface of tail white or cream.
Variation. Only four specimens of this species have been avail-
able for study. The variation in scalation of these specimens is as
follows: parietals about one third size of interparietal; frontoparie-
tals divided in none, in contact medially in two; frontal touches in-
terparietal in two; frontal normally divided in all; supraoculars
entire in all, with one complete and sometimes another incomplete
row of scales between them and superciliaries (in FMNH 1670 the
inner, incomplete row has two relatively large scales on one side
and one on the other side) ; supraoculars four or five; prefrontals in
582
The University Science Bulletin
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Smith: Genus Sceloporus 583
contact in two, separated by contact of median frontonasal and
frontal in one, by an azygous scale in the other; canthals two, the
first forced above the canthal ridge in two specimens; preocular
divided on one side in one, on both sides in another; lorilabials re-
duced to one row below subocular on one side of one specimen ; outer
row of labiomental? separated from mental in all; inner row of
iabiomentals terminating anteriorly between the anterior part of
the third infralabial and the posterior part of the second; auricular
lobules three to five, not greatly enlarged. FMNH 1670 is aberrant
in having thirteen femoral pores on one side.
Boulenger (1897), who had eight specimens, states that the fron-
tal is "sometimes in contact with the interparietal, but more usually
separated from it by the frontoparietals." The dorsal scale rows
are in either convergent or parallel rows. Other variations are given
in the accompanying table.
Range. Known only from the western and southern parts of the
state of Jalisco. (See Fig. 8 for distributional map.)
Locality records. Volcano of Colima (FMNH 1670, C. H. T.
Townsend) ; La Laguna (USNM 64664, Nelson and Goldman) ; San
Sebastian (USNM 64654-5, Nelson and Goldman) ; Colonia Bri-
zuela (Boulenger, 1894, 1897) ; La Cumbre de los Arrastrados
(Boulenger, 1894, 1897); Hacienda de Jalisco (Boulenger, 1897).
Sceloporus mucronatus mucronatus (Cope)
(Text Figs. 10 and 11; Plate XLIX, Fig. 2)
Sceloporux torquatus ? Gravenhorst, 1837, pp. 7G2-764, pi. 66, figs. 6-12; ? Sumichrast,
1882, p. 37 (part), Cope, 1885, pp. 401-403 (part); idem, 1887, p. 37 (part); ? Giinther,
1890, pp. xii, 66 (part); Boulenger, 1897, pp. 477-481 (part); Cope, 1900, pp. 336, 340,
347-350, fig. 50.
Sceloporus torquatus mucronatus CTope, 1885, p. 402; Cope, 1887, pp. 37-38; Giinther,
1890, p. 77; Cope, 1900, p. 349.
Sceloporus torquatus torquatus ? Cope, 1885, pp. 402, 403; ? Cope, 1887, p. 37.
Sceloporus torquatus poinsettii Boulenger, 1885, p. 220 (part); idem, 1897, p. 481 (part).
Type locality. Mirador, Vera Cruz. Cotypes USNM 25074-9,
Sartorius collector. No. 25077 designated as lectotype.
Diagnosis. Dorsals, 27 to 30 from occiput to base of tail, weakly
keeled, weakly mucronate, but with several lateral mucrones; upper
lateral scales somewhat larger than median dorsals, more strongly
keeled and mucronate; dorsal scales of body somewhat smaller than
largest dorsal caudals; median ventrals about one third or one
fourth as large as dorsals; scales around body, 84 to 38; head scales
smooth ; frontoparietals always in contact medially, or separated by
an azygous scale; supraoculars always in two series; canthals ir-
regular, usually two, the first sometimes forced above the canthal
584 The University Science Bulletin
ridge, the second canthal and subnasal contacting; femoral pores,
11-17 on each side, averaging about 13; tibia about as long as
shielded part of head; length of fourth toe about equal to distance
from snout to posterior margin of ear; a black nuchal collar, about
four scales wide, continuous about throat in adult males; collar
bordered on each side by a light band one scale wide; the posterior
border sometimes continued over upper foreleg near insertion; a
broad median dark band usually visible, composed of about five
large, dim blotches, which are sometimes separated from each other
by narrow, somewhat lighter areas; two parallel dark lines, one on
either side of the median ventral line, extend from near the chin to
the gular fold region; tail dimly banded; dorsal ground color olive
to wood brown. In adult males, sides of the abdomen and area
anterior to gular fold region caerulean or cobalt blue; gular fold
region black; area anterior to this becoming darker with age, losing
all trace of markings; breast, median abdominal area and region
about groin black.
Description (from EHT & HMS 2092). Interparietal moderately
large, about three times as large as either parietal; a parietal on
each side, not much longer than broad; frontoparietals rectangular,
separated medially by a single azygous scale; posterior section of
frontal rectangular, about two thirds the size of the anterior section ;
prefrontals in contact medially; two rows of enlarged supraoculars,
the scales of each approximately of equal size and not much larger
than scales in the series separating the supraoculars from median
head scales; two irregular or incomplete rows of scales separating
supraoculars from superciliaries; median and lateral frontonasals
subequal in size; internasals quite irregular; nasals and internasals
separated from rostral by a single row of scales; naris pierced about
in the middle of nasal, its anterior rim slightly the wider; two
canthals on one side, one on other (first canthal fused with sub-
nasal) ; subnasal and loreal large, latter slightly the larger; pre-
ocular divided, the upper half about one and one half times larger
than lower, keeled; lorilabials in two complete rows below loreal,
preocular and subocular, not imbricating; three and three fourths
or four and one half supralabials to a point below middle of eye,
and about five lower labials to the same point; outer labiomental
series separated from mental by almost one half the length of the
first infralabial; inner labiomental series terminating below middle
of second infralabial on one side, below suture between second and
third infralabial on other side; about six pairs of well-differentiated
Smith: Genus Sceloporus
585
postmentals, the scales of the anterior pair in contact medially.
Scales of head, except in gular region, pitted, less so in the occipital,
temporal, supra- and interorbital regions.
Five smooth auricular lobules, the second and third largest; scales
in temporal region between eye and ear keeled, mucronate, about
equal in size to scales between ear and lateral nuchal fold, which are
more strongly mucronate; dorsal scales weakly keeled, moderately
mucronate, with two to four lateral mucrones, more numerous in
lumbar region; lateral scales similar to dorsals, some distinctly
larger than median dorsals; ventrals about one third or one fourth
Fig. 10. Head scales of Sceloporus mucronatus mucronatus (Cope).
EHT & HMS 2556, near Totalco, Vera Cruz; actual head length, snout
to occiput, 15 mm.
as large as dorsals, rounded in pubic region, with an apical notch
in the middle of abdomen and in anterior gular region, with several
notches on sides of abdomen and in posterior gular region; preanals
about two thirds as large as median abdominals, somewhat smaller
than scales on posterior surface of femur; dorsal caudals, slightly
beyond base of tail, nearly twice as large as median dorsals of body;
subcaudals smooth; postanals enlarged.
Dorsal scales of humerus somewhat larger than those on lower
foreleg, slightly smaller than median dorsals of body; ventral scales
larger on lower forearm than on upper, smooth, notched; lamellar
formula for fingers 9-14-17-16-13 (9-13-17-?-12).
Dorsal scales of tibia one and one half times as large as dorsals
of femur, and distinctly larger than median dorsals on body; ventral
scales of tibia larger than those on femur; lamellar formula for toes
8-12-18-18-13 (8-13-17-18-14).
586 The University Science Bulletin
Color. Head and neck slate; black collar four scales broad, with
a light border one and one half to two scales wide on each side;
anterior border terminating on each side between upper edges of ear
and lateral nuchal fold; black collar continuous about neck; pos-
terior light border crossing over proximal end of humerus; a series
of four large, confluent, slate-colored spots of indefinite outline on
middle of back, forming a broad band from light border of collar
to base of tail; sides of body light olive, without darker marks;
tail with narrow light bands at about every fourth whorl; limbs
apparently with narrow, indistinct light bands; throat navy blue,
almost black; posterior gular area, chest and a broad area down
middle of abdomen black, with irregular, splotched areas of white
appearing on posterior part of chest and in the median ventral line
on abdomen; sides of abdomen caerulean or cobalt blue, lighter to-
ward sides of body; median black border of lateral blue areas con-
tinuous to and covering a considerable area of the groin, and ex-
tending onto proximal anterior surface of femur; a few irregular
black spots in preanal area; ventral surface of tail immaculate.
Variation. The scales of the head are essentially the same as in
other species of the torquatus group. The supraoculars are very
regular, in two rows, the outer row composed of scales but slightly
smaller than those of the inner row. The scales between the en-
larged supraoculars and the median head scales are very large —
about half as large as the scales of the adjacent row of enlarged
supraoculars. The first large scale of the inner supraocular series
may fuse with a scale in the series adjacent medially, thus contact-
ing the median head scales (occurs on both sides in seven speci-
mens, one side in six). There are always at least parts of two rows
of small scales between the outer series of enlarged supraoculars and
the superciliaries. In some cases there are parts of three rows. The
preocular is usually not divided (single on both sides in four speci-
mens, one side in three). Usually the two series of scales above
the supralabials are not reduced to one below the subocular (re-
duced on both sides in seven specimens, on one side in two). The
medial of the two series of scales intercalated between the series of
postmentals and the infralabials usually begins at a point about
even with the suture between the second and third infralabial,
although in one specimen it begins even with the suture between the
third and fourth. The lateral series of these two intercalated rows
begins very near the mental, but does not quite touch it, separated
by less than half the length of the first infralabial. The first canthal
is forced above the canthal ridge, resulting in a contact between the
Smith: Genus Sceloporus 587
second canthal and subnasal, on both sides in two specimens, on one
side in two. The prefrontals are separated by an azygous scale in
three specimens; the median frontonasal is invariably separated
from the frontal.
Remarks. This species is related to torquatus, agreeing with the
latter in the character of the carination and mucronation of the
body scales, in the number of dorsals from occiput to base of tail,
and in a number of color characteristics. It differs widely, however,
in the following: Supraoculars in two rows; frontoparietals contact-
ing on the median line; median frontonasal invariably separated
from frontal; femoral pores fewer and the two series more widely
separated medially ; prefrontals rarely not in contact on the median
line; two parallel blue lines always present on the throat; and an
indistinct broad, dark band down the middle of the back. That
mucronatus cannot be a subspecies of torquatus is further indicated
by the fact that they exist side by side in the same locality, retain-
ing their identity without question. Specimens of both species are
in the collections from Toxtlacuaya, Vera Cruz.
The closest relationship of m. mucronatus is with m. omiltemanus
(Giinther), from which it differs in possessing usually a smaller
number of dorsal scales from occiput to base of tail, dorsal caudals
larger than dorsals on body, fourth toe usually shorter than distance
from snout to posterior margin of ear, supraoculars in two even
rows, and a different coloration.
Specimens of this species were found four miles east of Encero,
Vera Cruz, associated with Sceloporus serrifer.
Cope's description of mucronatus was based on six specimens
(USNM 25074-9) from Mirador, Vera Cruz, collected by Doctor
Sartorius. Unfortunately, the cotypes have characters intermediate
between omiltemanus and typical mucronatus. Typical specimens
of the latter are available from the region about Jalapa, and typi-
cal specimens of omiltemanus are available from the region about
Orizaba. Mirador is exactly between these two localities, and
Cope's cotypes are very nearly exactly intermediate between typi-
cal specimens of the two subspecies. It is my conclusion that the
characters of the series of cotypes as a whole approach more closely
to those of the northern subspecies than to those of the southern.
The color characters typical of the northern subspecies are not pres-
ent, but neither are the color characteristics of omiltemanus present.
The scale characters and proportions of the cotype correspond, on
the other hand, more closely to the northern than to the southern
form. Although a certain color pattern is one of the most distinc-
588
The University Science Bulletin
live features, I prefer to consider these cotypes as belonging to the
northern subspecies, since scale characters approach and since there
is a possibility that the types once had color and markings not now
visible which would also approach those of what I shall call typical
w.ucronatus.
As first reviser, I further designate USNM 25077, a male having
.scale characters most closely approaching the typical, as the lecto-
type of mucronatus.
Habits and habitat. Specimens were collected in western Vera
Cruz on rock cliffs at a rather high elevation above Jalapa.
Fig. 11. Distribution of Sceloporus mucronatus mucronatus (Cope) and
S. m. omiltemanus (Giinther).
UMMZ 65270 contains five well-developed young, removed from
the oviducts. No date is on the specimens.
Range. Parts of the states of Hidalgo, Vera Cruz, Puebla and
Mexico.
Locality records. Vera Cruz: Jalapa (Cope, 1885, et al.) ; Mira-
dor (Cope, 1885) (type locality) ; Las Vigas (USNM 47589-90, E.
W. Nelson) ; Toxtlacuaya, about eighteen miles west of Jalapa (EHT
& HMS 1795-7, 1859, 1861-2, 2092-8, 2130-1) ; Cruz Blanca (EHT
& HMS 2132) ; four miles east of Encero (EHT & HMS 2519) ;
near Totalco (EHT & HMS 2556, 2558, 2564, 2566-2568). Puebla:
Fifteen miles east of San Marcos (EHT & HMS 1854-1858).
Mexico: San Juan Teotihuacan (MCZ 1890, Dr. G. 0. Rogers) ;
? Bay south of Mexico City (USNM 2963, I. Potts). Hidalgo:
Smith: Genus Sceloporus
589
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Smith: Genus Sceloporus 591
(MCZ 31737) ; Tulancingo (USNM 47591, 47032-3, E. W. Nelson) ;
El Chino (USNM 47291, E. W. Nelson) ; Guerrero Mill (MCZ
10700-5, 17114-5, AV. M. Mann; UMMZ 65270 [2 spec.]); Velasco
(MCZ 13651-4; UMMZ 71439 [3 spec], 71453). No data: UMMZ
56487 (2 spec).
Sceloporus mucronatus omiltemanus (Giinther)
(Text Figs. 11 and 12; Plate L>
Sceloporus torquatus ? Gravenhorst, 1S37, pp. 762-764, pi. 66, figs. 9-12; Bocourt, 1874,
pp. 171-173, pi. 18, figs. 9, 9a, 9b, 9c (part); Sumichrast, 1882, p. 37 (part ?); Cope, 1885,
pp. 401-403 (part); Giinther, 1890, pp. xii, 66 (part); Boulenger, 1897, pp. 477-481, fig.
1 (part); Gadow, 1905, pp. 194, 214 (part).
Sceloporus poinsettii Bocourt, 1874, p. 173 (part).
Sceloporus torquatus torqriatm ? Cope, 1885, pp. 402, 403; ? idem, 1887, p. 36.
Sceloporus torquatus poinsettii Boulenger, 1885, p. 220 (part); idem, 1897, p. 481 (part).
Sceloporus omiltemanus Giinther, 1890, pp. -xii, 66, pi. 32, fig. A; Boulenger, 1890, p. 78.
Sceloporus torquatus omiltemanus Taylor, 1931, p. 131.
Type locality. Omilteme, Guerrero.
Diagnosis. A species of slightly more than moderate size, belong-
ing to the torquatus group; head shields smooth; supraoculars in two
irregular rows; frontal rarely contacting interparietal, usually sepa-
rated by an azygous scale, occasionally by contact of the two fronto-
parietals; laterals about one third smaller than dorsals, in oblique
series; dorsals weakly keeled, mucronate, in parallel or slightly con-
verging series, usually 30 to 38 from occiput to base of tail, rarely
less than 30; laterals a little more strongly keeled and mucronate
than dorsals; upper scale of auricular series extremely large, larger
than preceding scales and much larger than other auricular scales;
tibia about as long as shielded part of head; fourth toe as long as or
slightly longer than distance from snout to posterior margin of ear ;
ratio of hind leg to snout-vent measurement, 61.4 to 69.7; ratio of
fourth toe to snout-vent measurement, 22.3 to 26.4; black nuchal
collar about four scales wide, curving slightly posteriorly, and light
bordered; posterior light border complete, one to two scales wide,
extending onto humerus ; anterior border similar, but very short, not
extending beyond lateral nuchal fold and occasionally interrupted;
sides of belly cobalt blue in males, not or but narrowly and indis-
tinctly bordered with black laterally; gular fold region black, con-
tinuous with the black of the nuchal collar; area anterior to gular
fold region uniform cobalt or azure blue; dorsal caudals slightly
larger than dorsals on back.
Description (from EHT & HMS 3080, male). Dorsal head
shields smooth; enlarged supraoculars in two rows, the outer row
formed of scales about two thirds as large as those of the inner row;
592
The University Science Bulletin
supraoculars separated from superciliaries by one complete and an-
other incomplete row of small scales and bordered medially by a
series of small scales; superciliaries five; one large, heavily keeled
subocular, followed posteriorly by two small, strongly keeled post-
oculars; preocular scale strongly keeled, in contact with second
canthal and with the rows of scales above supralabials; first canthal
with a narrow downward extension which contacts the rows of scales
Fig. 12. Head scales of Sceloporus mucronatus omiltemanus (Giinther).
EHT & HMS 3080, two miles west of Acultzingo, Vera Cruz; actual head
length, snout to occiput, 18 mm.
above the supralabials, separating the loreal from the subnasal; not
quite half of second canthal forming a portion of the superciliary
series; two or three irregular rows of scales above supralabials, two
complete rows between the subocular and supralabials, and one row
continued around snout above rostral; rostral about two and one
half times as broad as high; interparietal much larger than either
parietal, separated from the frontal by the frontoparietals; latter as
large as parietals, in contact on median line; frontal divided, the
posterior section not quite half as large as the anterior; two pre-
frontals, in contact on median line; three frontonasals, the laterals
in contact with both canthals; internasals rather irregular, but
Smith: Genus Sceloporus 593
posterior pair distinct and separated from first canthal by a single
small scale; six scales surround the nasal; four supra- and five in-
fralabials to a point below the middle of the eye.
Mental pentagonal, its labial border four fifths that of rostal;
mental followed by a series of about five enlarged scales on each
side, the anterior in contact with its fellow on the median ventral
line; this series of scales separated from infralabials anteriorly by
a single row of elongate scales (except anterior scale, which is in
contact with the first infralabial), posteriorly by two rows, the
second row beginning below the junction of the second and third
infralabials and in contact with the series of postmental scales;
scales in temporal region keeled, mucronate in area near ear; an-
terior margin of ear with three smooth scales, the upper ver>^ large,
much larger than other auricular scales and distinctly larger than
preceding scales ; lateral cervical pouch present, a fold from about
its middle or upper margin to the lower margin of ear; this fold sur-
mounted by scales somewhat larger than the adjacent ventral scales,
but somewhat smaller than adjacent scales dorsal to the fold; scales
surmounting this fold strongly mucronate; dorsal scales in parallel
series (a small, short series intercalated on the median line) , weakly
keeled, mucronate, with one or two small lateral mucrones; lateral
scales in diagonal rows, about one third smaller than dorsals, more
strongly keeled and mucronate than dorsals, with many lateral
mucrones; dorsal scales about as large as or somewhat smaller than
dorsal caudals; median ventral scales about one third as large as
dorsals; breast scales somewhat larger than those in median ab-
dominal region; preanal scales somewhat smaller than the latter;
anterior scales in area in front of gular fold region about half as
large as the posterior scales; scales in area near angle of jaws but
little larger than those in median gular region.
Dorsal scales of foreleg keeled, mucronate, those of upper foreleg
about two thirds the size of the dorsals, those of lower foreleg a
little more than half as large as those on upper foreleg; dorsal
scales on hand weakly keeled or smooth, weakly mucronate; ven-
tral scales on foreleg smooth, mucronate, those of lower foreleg
about the same size as dorsals of the same member, those of upper
foreleg becoming smaller toward axilla ; one dorsal, two lateral and
one ventral row of scales around fingers and toes, all continuous to
tip except the two laterals which terminate behind the last scale of
the other series; lamellar formula for fingers, 9-13-15-16-13.
38—6037
594 The University Science Bulletin
Dorsal scales of hind limb strongly keeled, mucronate, less
strongly on foot; dorsal scales of tibia slightly smaller than dorsals
on body, those of femur somewhat smaller than those of tibia; ven-
tral scales of tibia somewhat smaller than dorsals of the same mem-
ber; scales on anterior surface of femur somewhat smaller than dor-
sals of same member, becoming progressively smaller toward fem-
oral pore series; scales immediately in front of femoral pores about
equal in size to preanal scales; scales immediately behind femoral
pores very small, much smaller than those immediately in front of
femoral pore series, becoming gradually smaller toward dorsal sur-
face; lamellar formula for toes, 9-12-18-19-13; subcaudals smooth
and rounded except toward distal end of tail; dorsal caudals rather
strongly keeled, strongly mucronate. A pair of enlarged postanal
scales.
Color. General ground color apparently drab; dorsum posterior
to nuchal collar with a pattern of light lines which follow a course
down the middle of each scale row, separated from each other by
narrow black lines, which include the edges of the scales. The black
nuchal collar is three or four scales broad, bordered anteriorly and
posteriorly by a distinct whitish band one or two scales broad. The
black extends over the anterior margin of the insertion of the fore-
leg and traverses the gular fold region ventrally. The posterior
light border is complete and extends over the proximal end of the
upper forearm. The anterior border is also complete and passes
laterally only to the upper edge of the lateral cervical fold. The
area anterior to this is blackish slate. The limbs are hair brown,
immaculate. The tail is of the same color, with lighter and darker
distal bands. The area anterior to the gular fold region is azure
blue; the sides of the belly are somewhat darker blue, with a faint
purple tinge; the chest and median abdominal region are suffused
with slate, becoming black near lateral abdominal blue areas; the
ventral surfaces of the forelimbs, femora and the preanal region are
suffused with slate. The ventral surface of the tail is light orange
toward the base, fading distally and replaced by gray.
Variation. The variation of the head scales of sixty-eight speci-
mens is as follows: The supraoculars are almost invariably in two
rows. A single young specimen from Omilteme, Guerrero (USNM
47738), is the only exception, having but a single row of enlarged
supraoculars. A peculiar variation frequently occurs, however, in
the series of scales normally separating the supraoculars from the
median head scales. In eight specimens one or more of the enlarged
Smith: Genus Sceloporus 595
supraoculars touch the frontal (first only in three ; first on one side,
first, second and fifth on the other, in one; first on one side, first
and second on the other, in one; first and second, in one; first,
second, third and fourth [parts of each], one). The frontal touches
the interparietal in five, the frontoparietals contact medially in ■
twenty-one; and a single median scale between the frontal and in-
terparietal is present in forty. The anterior part of the frontal is
longitudinally divided in six, the posterior part in one. The pre-
ocular is separated from the row of scales above the supralabials
on both sides in ten, on one side in four. The first canthal touches
the lorilabials on both sides in five, on one side in three. The second
canthal touches the subnasal (the first canthal above the canthal
ridge) on both sides in three, on one side in five. The posterior pair
of internasals contact the first canthal on both sides in seven, on
one side in six. The median frontonasal narrowly contacts the
frontal in two specimens, and is separated from frontal in the re-
mainder of the series.
The length of the labial border of the mental in relation to that
of the rostral varies from four fifths to a little over one half. The
series of scales above the supralabials are usually two below the
subocular, but occasionally reduced to one at some point below the
subocular. The series of scales between the series of postmentals
and the infralabials is much as in the type, except that the median of
the two series begins anywhere from a point even with the junction
of the first and second infralabial to a point even with the junction
of the third and fourth.
The upper ear lobule is invariably larger than the other auricular
scales, and larger than those preceding, except in six young speci-
mens. The dorsal scale rows vary from parallel series to slightly
converging rows.
The coloration of the adult males is much as in the specimen
described. Most characteristic is the pattern of light and black
longitudinal lines, which appear, so far as I am aware, in no other
species of the torquatus group except poinsettii. The anterior light
border of the nuchal collar is frequently broken, and the resultant
spots may enclose but a single scale. The posterior light border is
occasionally broken medially, and may be but one scale wide. It
always extends distinctly onto the proximal end of the upper fore-
leg. As the specimen described is one of the largest of the series,
the ventral coloration is darker than most of the other males.
596 The University Science Bulletin
Usually there is but little black ventrally, and the blue of the throat
and sides of the belly is more distinct.
The dorsal coloration of the females is much as in the males,
except that the longitudinal light and dark lines are less evident and
• are more frequently entirely absent. In these cases, small light and
dark spots may be scattered over the back. In a few specimens the
posterior light border of the nuchal collar is bordered behind by a
very narrow black band. The ventral surface is usually whitish,
sometimes with a general suffusion of blue.
In the young specimens very dim, dark bands broken medially
may traverse the back. Usually the back is irregularly spotted. In
only a few of the very young males are the longitudinal light and
dark lines visible.
Habits and habitat. The specimens collected near Acultzingo,
Vera Cruz, were found on rock cliffs on the abrupt edge of the
plateau, about three fourths of the distance from the bottom. Sev-
eral specimens collected on Cerro de San Luis in Oaxaca were at a
considerable elevation below the evergreens.
It is assumed that omiltemanus is ovoviviparous, as are several
other forms of the torquatus group, including mucronatus mucro-
natus.
Range. From central Vera Cruz in the region of Mirador south
and west to the edge of the plateau in Oaxaca and Guerrero. (See
Fig. 11 for distributional map.)
Locality records. Vera Cruz: two miles west of Acultzingo (EHT
& HMS 3073-3080, 3082-3102, 3171-3174, 3193-3195); Maltrata
(USNM 46823, Nelson & Goldman). Piiebla: near Tehuacan (EHT
& HMS 3338) ; vicinity of Puebla (AMNH 18815-20, Paul D. R.
Riithling) ; Los Reyes, near Santa Catarina (AMNH 18407, Paul
D. R. Ruthling); Atzitzintla (FMNH 1516 [16 spec]. Heller &
Barber, 8,500 ft.). Oaxaca: Cerro de San Luis, N. of Oaxaca,
Oaxaca (EHT & HMS 3839-43) ; lower part of descent into valley
of Miahuatlan (AMNH 17994-8, Paul D. R. Ruthling) ; Tlapen-
cingo (USNM 47832, Nelson & Goldman) ; Mt. Zempoaltepec
(USNM 47578-83, 47186-8, Nelson & Goldman); Tamazulapam
(USNM 47325-6, Nelson k Goldman). Guerrero: near Chilpan-
cingo (USNM 47830-1, Nelson & Goldman; MCZ 33903-4, W. W.
Brown) ; Omilteme (type locality of omiltemanus) (Giinther, 1890;
Boulenger, 1897; USNM 47738, Nelson & Goldman). Indefinite
localities: Mexico (AMNH 18400-4; USNM 31353, Boucard).
Smith: Genus Sceloporus
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Smith: Genus Sceloporus 599
Sceloporus cyanogenys (Cope)
(Text Figs. 13 and 14; Plate LI, Fig. 2)
Sceloporus torquatus cyanogenys Cope, 1885, p. 402; idem, 1887, p. 87 (part); Giinther,
1890, p. 77; Cope, 1900, pp. 349-350; Taylor, 1931, pp. 129-132; Burt, 1932, p. 158;
Stejneger and Barbour, 1933, p. 64 ; Mulaik, 1935, p. 156.
Sceloporus torquatus Baird, 1859, p. 5; Yarrow, 1883, p. 58 (part); Cope, 1885, p. 169
(part) ; idem, 1887, p. 37 (part).
Sceloporus torquatus mucronatus Boulenger, 1885, p. 220.
Sceloporus torquatus poinsettii (non Baird and Girard) Burt, 1932, p. 158.
Type locality. Monterrey, Nuevo Leon, Mexico. Cotypes USN
31373-7 and ? ANSP 11304-5; Cope collector.
Diagnosis. A member of the torquatus group of the genus Scelop-
orus; dorsal scales weakly keeled, mucronate, with several lateral
denticulations, 32 to 40 from occiput to base of tail; lateral scales
in oblique rows, somewhat larger than median dorsals; fourth toe
about equal to distance from snout to posterior border of ear; ratio
of hind leg to snout-vent measurement, 58.6 to 66.9; ratio of fourth
toe to snout-vent measurement, 20.3 to 25.2; supraoculars irregular,
not in two equal rows, rarely in a single row; median cephalic scales
not extremely irregular; preocular rarely divided; inner row of
labiomentals rarely terminating posterior to suture between second
and third infralabials; rows of lorilabials usually not reduced to
one at a point below subocular; femoral pores usually twelve or
more on each side. General ground color greenish blue (brownish
prior to shedding) ; nuchal black collar four or five scales wide,
sometimes continued on ventral surface, bordered anteriorly and
posteriorly by a light band one or one and a half scales wide, both
interrupted medially by a dark scale; a light spot in center of inter-
parietal and of each parietal; body without or with dim darker
markings; throat and sides of belly in males pearl blue to flax
flower blue or campanula blue; lateral abdominal blue areas in
males with a dark-blue medial border not encroaching on anterior
distal surface of femur; throat not barred or rayed.
Description (from EHT 1284 male, collected in hills north of Los
Olmos Bridge, three miles south of Rio Grande City, Starr county,
Texas, August 21-23, 1931) . Head and body not strongly depressed ;
head scales with small pits, all smooth except those between supra-
oculars and superciliaries, between supraoculars and median head
scales, and lorilabials; interparietal large, flat, shield-shaped, its
posterior border extending back of parietals about one third the
length of the scale; parietal single on each side, broadly wedge-
shaped, about one third the area of interparietal; frontoparietals
short, rectangular, in contact medially; frontal not divided (abnor-
600
The University Science Bulletin
mal) ; preoculars broadly in contact medially, nearly as large as
median frontonasal; frontonasals about equal in size, the laterals
contacting both canthals; a pair of rather large scales in front of
median frontonasal; preceding these a median pair of internasals,
separated from rostral by a row of small scales; nasal small, sepa-
rated from rostral, naris pierced almost in the middle; five supra-
oculars, in one complete row on one side, three divided on other
side; one complete and another incomplete row of scales between
Fig. 13. Head scales of Sceloporus cyanogenys (Cope). EHT & HMS
4929, Arroyo Los Olmos, three miles southeast of Rio Grande City, Stan-
county, Texas; actual head length, snout to occiput, 16.8 mm.
supraoculars and superciliaries, the scales keeled or rugose; row of
scales between supraoculars and median head scales keeled; two
canthals, normal; subnasal present; two loreals, the anterior some-
what larger than posterior; preocular divided; subocular followed
posteriorly by two small, strongly keeled postoculars; lorilabials
in two complete rows below subocular; four upper and five lower
labials to a point below middle of eye.
Mental with a labial border slightly more than half that of
rostral; outer row of labiomental scales separated narrowly from
mental by narrow contact of first postmental and first infralabial;
inner row of labiomentals terminating anteriorly below anterior
half of second infralabial; three or four well-differentiated post-
mentals on each side, the scales of the anterior pair in contact medi-
ally; scales in gular region approximately subequal in size, smooth,
Smith: Genus Sceloporus 601
those in the anterior part rounded, those in middle part with a single
notch, and the scales in the posterior part and toward angle of jaws
with two or three apical notches.
Four or five smooth, rounded auricular lobules, smaller than the
preceding scales; scales in temporal region keeled, mucronate, some-
what smaller than those between ear and lateral nuchal fold ; scales
surmounting lateral gular fold keeled, strongly mucronate; dorsal
scales weakly keeled, mucronate, with three to five lateral denticu-
lations; lateral scales somewhat larger than median dorsals, more
strongly keeled and mucronate, with as many as eight lateral mu-
crones; ventral abdominals about half as large as median dorsals,
all with a single apical notch (except those on anterior part of chest,
which have two or three) ; ventral scales on chest somewhat smaller
than those in middle of belly; preanal scales smooth, with one or two
apical notches, about half as large as median scales on belly; sub-
caudals smooth, mucronate, with several lateral mucrones; dorsal
caudals extremely large, about one and one half times as large as
median dorsals; dorsal scales between hind legs about two thirds as
large as median dorsals.
Dorsal scales of foreleg about three fourths as large as median
dorsals on body, those on lower foreleg about one fourth smaller
than dorsals on upper foreleg; ventral scales of foreleg smooth, mu-
cronate, denticulate, those on lower foreleg subequal in size to dor-
sal scales of same member, about half as large as ventral scales on
upper foreleg; lamellar formula for fingers, 9-13-16-18-12 (9-13-
16-17-12).
Dorsal scales of hind leg keeled, strongly mucronate, those on
tibia as large as median dorsals on back, those on femur somewhat
smaller; ventral scales of hind limb smooth, notched, those on tibia
about two thirds as large as dorsal scales of same member; ven-
tral scales on femur equal in size (near series of femoral pores) to
preanals; scales on posterior surface of femur about twice as large
as preanals, strongly mucronate, keeled, denticulate, abruptly de-
creasing in size immediately behind femoral pore series; lamellar
formula for toes, 9-13-18- ?- ? (9- ?-17-20- ?).
Taylor (1931) has given an excellent description of the dorsal
coloration:
"The color and markings of this form, while varj'ing in certain color char-
acteristics, maintained a striking uniformity as regards certain of the mark-
ings of the head and neck region. The typical markings are as follows:
Scales on the head, brownish olive with a cream-colored spot between the su-
praocular regions, one on the median parietal, and one on the enlarged lateral
scales adjoining the parietals; three spots on the small scales immediately
602 The University Science Bulletin
posterior to the parietal region; a broad black collar four to five scales wide
medially, but narrowing laterally, extends across the scapular region, and ter-
minates in front of the foreleg; the scales between the parietals and the pos-
terior median edge of the black collar are from fifteen to seventeen in num-
ber; the black collar is bordered posteriorly by a greenish-yellow to cream-
colored band one to one and one half scales wide which terminates laterally
at the base of the foreleg; this is, almost without exception, interrupted
medially by a single scale which is black or dark green; the black collar is
bordered anteriorly by an irregular band of cream, or greenish or bluish-
yellow, which does not extend as far on the sides as the black collar.
"The neck in front of the bands is greenish, bluish or bluish-brown with
from five to seven cream or bluish-cream spots, each covering usually about
two scales; the sides of the head and neck are greenish, bluish or brownish;
a black spot on the head immediately behind the superciliary scales is bor-
dered below by a lighter, usually a cream-colored spot; a cream-colored line
passes from the loreal region below the eye back across the auricular region,
appearing very indistinct posteriorly.
"The general body color is usually a brilliant greenish blue (often showing
metallic iridescence) or (prior to shedding) brownish. On the back the
markings are variable; sometimes there are a series of light ocelli or dark
blackish or brownish spots; more or less distinct transverse blotches or series
of spots may be present.
"Two individuals, apparently about to shed, show the brownish coloration
without or only with a trace of the greenish-blue coloration. The fore and
hind legs are similar to the general body color, the hind legs frequently show-
ing some darker mottling; the coloration of the tail varies remarkably from
dull brown to bluish-black with a series of 16-18 indefinite lighter bands
usually one scale wide which vary from whitish to ultramarine in color; some-
times the bands are almost obsolete."
The ventral coloration of the males is most brilliant and quite
different from that of other species of the torquatus group. The
sides of the belly and the area anterior to the gular fold region are
pearl to flax flower or campanula blue. In young males the blue of
the sides of the belly are bordered with a narrow band of pale blue,
and the center of the throat toward the gular fold region is also
pale blue. In the oldest males the blue borders of the lateral ab-
dominal patches become marine blue, and these borders expand
posteriorly, covering the region of the groin. The middle of the
belly is whitish, not suffused with any darker color in even the
largest males. This median band is about eight scales wide. The
nuchal collar extends across the gular fold region ; the centers of the
scales in this region may be azure blue.
The females have no markings whatsoever on the ventral surfaces.
The ventral surface of the tail in both sexes is immaculate and
whitish.
Smith: Genus Sceloporus 603
Variation. Details of cephalic scalation in forty-one specimens
are as follows: The frontoparietals contact in six, are separated by
contact of frontal and interparietal in seven, by an azygous scale in
twenty-seven, and by a median pair of scales in one. The supra-
oculars are extremely irregular in form; in one specimen the supra-
oculars are entire on both sides; in another they are entire on one
side ; in two specimens there are two complete rows ; in the remainder
they are variously divided or fused. The anterior frontal is longi-
tudinally divided in four specimens; in two the frontal is quite
irregular; in another the anterior frontal is fused with one pre-
frontal. The prefrontals contact medially in all but two, in which
they are separated by contact of the median frontonasal and the
frontal. The canthals are two in all, the first not forced above the
canthal ridge by contact of the second canthal and subnasal with
one exception ; the first canthal touches the rows of scales above the
supralabials on both sides in four specimens, on one side in ten.
The two rows of scales between the subocular and the supralabials
are reduced to one at some point on both sides in seven, on one side
in five specimens. The first pair of postmentals are separated in
one specimen. The inner of the two rows of scales intercalated on
each side between the series of postmentals, and the infralabials
terminate even with the suture between the first and second infra-
labial in five cases (two cases per specimen), at the anterior half of
the second in ten cases, at the middle of the second in ten, at the
posterior half of the second infralabial in sixteen, at the suture be-
tween the second and third in thirty-three, and at the anterior half
of the third in eight.
The auricular scales are small, the upper not greatly enlarged,
occasionally slightly larger than the preceding scales.
The median dorsal scales are weakly keeled, weakly mucronate,
usually in slightly or rather strongly converging rows, but occasion-
ally in parallel rows. The laterals are slightly more strongly keeled
and mucronate, and are larger than the dorsals at a point halfway
between and slightly above a line from the axilla to the groin. The
laterals elsewhere, toward the ventral surfaces and toward the axilla
and groin, are smaller than or no larger than the dorsals. The
latter are definitely smaller than the dorsal caudals near the base
of the tail. The femoral pores vary from eleven to seventeen.
Habits and habitat. Taylor (1931) gives the following notes on
habits:
604
The University Science Bulletin
"While collecting in Southern Texas in 1930 I kept an eager outlook for
this subspecies, which resulted in its discovery. I encountered it first near Rio
Grande City in the hills, one half mile to the north of the Los Olmos bridge.
The specimens were extremely numerous, as many as ten or fifteen might be
seen at one time running over the face of the outcropping rock which caps
the hills. The largest males seemed most wary and would disappear in deep
holes in or under the rocks (rather than in cracks and crevices) ; the larger
females and the younger specimens were less wary and instead of disappear-
ing to safety would frequently hide from sight behind a jutting rock and
then expose their heads to view a moment later and allow me to approach
close enough to kill them with a .22-caliber rifle using small-shot shells. It was
usually necessary to wait a considerable time before the males would re-
appear, only to disappear again for a longer period if they chanced to see or
hear me. A series of seventeen specimens were taken at this locality.
"At Arroyo El Tigre, west of Rio Grande City, the species was encountered
on the dry earth banks where no rocks were to be found. They took shelter
in cracks in the earth, where they were routed by causing portions of the bank
to cave in. Five adult specimens were taken here.
"Eighteen miles south of Laredo I took a single specimen at Arroyo Salado.
At Eagle Pass, in the cliff on the immediate edge of the town, another speci-
men was found. This habitat was rocky, much the same type as at Rio
Grande City, but this and a single young specimen which escaped were the
only specimens observed in half a day's collecting."
It is assumed that cyanogenys is ovoviviparous, as are other
species of the torquatus group.
Range. Southern Texas from Devil's river southeast to Starr
county, Texas, south through northern Mexico to central Tamaulipas
in the east and Santa Catarina (west of Monterrey) on the west.
Fig. 14. Distribution of Sceloporus cyanogenys (Cope) and S. poinsettii
Baird and Girard.
Smith: Genus Sceloporus
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606 The University Science Bulletin
Locality records. United States: Texas, Starr Co.: Arroyo Los
Olmos, three miles southeast of Rio Grande City (Taylor, 1931;
EHT & HMS 4800, 4801, 4803, 4883-5, 4925-31, 4991, 5051-53;
EHT 1 spec; MCZ 33549) ; Arroyo El Tigre, west of Rio Grande
City (Taylor, 1931; KU 8176-8) ; Rio Grande City (KU 8197-8210,
15010-21; EHT 2 spec.) ; five miles east of Rio Grande City (LMK
25239-40, S. Muliak) ; twenty miles northwest of Rio Grande City
(KU 15022-3) ; near San Ignacio (near Rio Grande City) (EHT 4
spec). Zapata Co.: near Zapata (EHT & HMS 4719-21); four
miles east, one half mile south of Zapata (UMMZ 74752 [2 spec],
Helen Monroe). Webb Co.: Arroyo El Salado, eighteen miles
south of Laredo (Taylor, 1931; KU 8180-1). Maverick Co.: Eagle
Pass (Taylor, 1931; KU 8179; USNM 47587-8, W. Lloyd & C. P.
Streator). Mexico: TaynauUpas: three miles south of Nuevo
Laredo, June 7, 1932 (EHT & HMS 256-264) ; four miles southwest
of Nuevo Laredo (Burt, 1932) ; Mier (USNM 47715-7, 47457, W.
Lloyd) ; between Laredo and Camargo (USNM 2959, A. Schott) ;
Rancho El Plato, thirty-eight miles, by road, southeast of Remosa
(USNM 95182, Edgar Bowles) ; Tamaulipaca (UMMZ 69228 [2
spec], 69229 [2 spec], 69230 [3 spec], 69231 [2 spec], 69232 [7
spec], 69241, L. R. Dice) ; San Jose (UMMZ 69233-5, L. R. Dice) ;
Garza Valdes (FMNH 1287). Nuevo Leon: six miles south of Sa-
binas Hidalgo, June 10, 1932 (EHT & HMS 447-451) ; Monterrey
(type locality) (Cope, 1885, 1887, 1900; USNM 31373-7 [cotypes] ;
ANSP 11304-5 [probably cotypes; no locality given]); Caderita
(MCZ 5940) ; Santa Catarina (USNM 4107, Lt. B. Couch) ; Pes-
quieria (USNM 2925, Lt. B. Couch).
Sceloporus poinsettii Baird and Girard
(Text Figs. 14 and 15; Plate LI, Fig. 1)
Sceloporus poinsettii Baird and Girard, 1854, pp. 126-127; Baird, 1859, p. 5, pi. 29, figa.
1-3; Cope, 1875, p. 48; Coues, 1875, p. 595; Yarrow, 1875, p. 573; Yarrow & Henshaw,
1878, p. 1645; Cope, 1880, p. 17; idem, 1883, p. 12; Garman, 1884, p. 17 (part); idem,
1887, p. 14 (part); Duges, 1887, p. 114; Giinther, 1890, pp. xii, 65; Herrick, Terry and
Herrick, 1899, pp. 123-125, pi. xv; Bailey, 1913, p. 34.
Sceloporus torquatus ? Yarrow, 1883, p. 58; Garman, 1887, p. 14; Cope, 1888, p. 397.
Sceleporus poinsettii Miiller, 1865, p. 602.
Tropidolepis poinsettii Duges, 1870, p. 243 (very doubtful).
Agama torquata ? Garman, 1884, p. 17 (part).
Sceloporus poinsetti Yarrow, 1883, p. 58 (part).
Sceloporus torquatus poinsettii Boulenger, 1885, p. 220 (part) ; Cope, 1885, p. 402 ; idem,
1886, p. 283; idem, 1887, p. 37; Boulenger, 1897, p. 481 (part); Cope, 1900, pp. 340. 349,
350-352, text fig. 51 (part); Brown, 1903, p. 546; Stone and Rehn, 1903, p. 31; Bailey,
1905, p. 42; Ditmars, 1907, pp. 130, 140-141, pi. 45; Strecker, 1909, p. 13; Strecker, 1915,
p. 20; Stejneger and Barbour, 1917, p. 56; Van Denburgh, 1922, pp. 326-329; Stejneger and
Barbour, 1923, p. 59; Van Denburgh, 1924, pp. 206-207; Strecker, 1927, p. 14; idem, 192S,
p. 12; Taylor, 1931, pp. 130-131; Mosauer, 1932, pp. 7-8; Stejneger and Barbour, 1933, p
65; Strecker, 1933, p. 78; ide7n, 1935, p. 32.
Smith: Genus Scelopoeus 607
Type locality. "Rio San Pedro of the Rio Grande del Norte, and
the province of Sonora." Cotypes USNM 2948 (2 spec.) and 2952
(2 spec.) ; J. H. Clark, collector.
Diagnosis. A member of the torquatus group of the genus Sce-
loporus, possessing a broad, black nuchal collar two and one half to
three scales broad, uninterrupted medially, with broad, light-colored
borders about two scales wide; tail very strongly banded, the dark
bands wider than the light bands and extending completely around
tail near tip; general color, straw to reddish yellow; back frequently
with broad bands; dorsals, 31 to 41 (average 35.7) from occiput to
base of tail, nearly smooth, not or but weakly mucronate; supra-
oculars in two complete rows; median cephalic scales usually very
irregular; the anterior frontal usually divided; preocular rarely di-
vided; inner row of labiomental rarely terminating anterior to
suture between second and third infralabials ; lorilabials usually re-
duced to one row at some point below subocular; femoral pores
usually not over 12; maximum snout-to-vent measurement, about
120 mm.
Description (from DHD & HMS No. 105, male). Head some-
what compressed; dorsal and lateral cephalic scales rather strongly
pitted; parietal single on each side; interparietal pentagonal, about
three times as large as either parietal; frontoparietals divided; an
azygous scale separating frontoparietals medially; posterior section
of frontal about one half the size of anterior section of frontal; a
small scale between frontal and frontoparietals on one side ; anterior
section of frontal longitudinally divided ; prefrontals separated medi-
ally by an azygous scale, which is wedged deeply between either
half of anterior section of frontal; median frontonasal wedged deeply
between prefrontals, separated on one side by a small scale from
lateral frontonasal; scales in internasal region quite irregular; nasal
(and internasals) separated from rostral by a row of small scales;
anterior rim of nasal much broader than posterior rim; supraoculars
about six, in two rows, the scales subequal in size; supraoculars sepa-
rated from median cephalic scales by one complete row of scales,
and separated from superciliaries by one complete and another in-
complete row of scales; two canthals, the anterior not forced above
canthal ridge; one subnasal, small; loreal single, somewhat larger
than subnasal; preocular not divided; subnasal short, followed by
two small, strongly keeled postoculars; lorilabials in two rows, re-
duced to one at a point near posterior end of subocular; supralabials
about four, infralabials six, to a point below middle of eye.
608
The University Science Bulletin
Mental more triangular than pentagonal in shape, with a labial
border almost equal to that of rostral, postmentals poorly differ-
entiated, about five pairs discernible, the scales of the anterior pair
in contact; outer row of labiomentals narrowly separated from
mental by a narrow contact of first postmental and first infralabial;
inner row of labiomentals terminating below middle of third in-
fralabial; scales in gular region about half as large as median ven-
tral abdominals, about equal in size to preanals; gular scales en-
FiG. 15. Head scales of S. poimettii Baird and Girard. DHD & HMS 1142,
30 miles west of Monterrey, Nuevo Leon ; actual head length, snout to occiput,
17 mm.
tire anteriorly, with a single median notch posteriorly, and with two
notches laterally at posterior part of throat.
Three or four well-differentiated auricular lobules, smooth, acu-
minate, smaller than preceding scales; scales in temporal region
weakly keeled, weakly mucronate, with a few lateral mucrones;
lateral nuchal fold not extremely deep, surmounted by very long,
strongly acuminate, keeled scales; dorsal scales of neck and body
smooth or very weakly keeled, not acuminate, but with several
mucrones at free end; lateral scales weakly keeled, mucronate, with
numerous lateral denticulations, arranged in oblique rows; median
lateral scales distinctly larger than median dorsals; scales between
bases of hind legs one half to one third as large as median dorsals
on back; dorsal caudal scales weakly keeled, strongly mucronate,
Smith: Genus Sceloporus 609
those near base of tail distinctly larger than median dorsals on back;
median ventral scales one fifth or one sixth as large as largest later-
als, with a weak median notch; scales at extreme anterior portion
of breast larger than scales posterior; subcaudals smooth; postanals
distinctly enlarged, separated by two small scales.
Dorsal scales of foreleg subequal in size, those of hmnerus some-
what the larger, all slightly smaller than median dorsals of back,
weakly keeled, rather strongly mucronate, with several latei-al
denticulations; ventral scales of lower foreleg subequal in size to
dorsal scales of same member, about three or four times as large as
ventral scales of humerus; ventral scales of foreleg smooth, acumin-
ate; lamellar formula for fingers, 9-13-15-17-11 (8-12-15-16-12).
Dorsal scales of tibia somewhat larger than those on femur, sub-
equal in size to dorsal scales on body; dorsal scales of hind leg
weakly keeled, strongly mucronate, with several lateral mucrones;
ventral scales of hind leg smooth, rounded or notched, those on tibia
slightly smaller than dorsals of same member, those on femur much
smaller, the smallest about equal in size to preanal scales; scales
on posterior surface of femur four times as large as preanals, keeled,
strongly mucronate, abruptly becoming quite small immediately
posterior to femoral pore series; no postfcmoral dermal pocket;
lamellar formula for toes, 9-11-16-18-13 (8-11-15-17-13).
Color. General ground color, above, straw yellow; head reddish-
brown above; black nuchal collar about three scales wide, bounded
on either side by a light band two scales wide, each very narrowly
interrupted medially ; anterior light border terminating on each side
between ear and foreleg; posterior border terminating at axilla, not
crossing onto shoulder; a light band about two scales wide across
neck halfway between nuchal collar and occiput, terminating at
upper edge of ear; a broad light line down each dorsal scale row,
one separated from the other by a narrow black line involving the
lateral edges of each scale row; these lines terminating at base of
tail; about five faint dark bands across back, separated from each
other by slightly narrower lighter bands; these bands very distinct
on tail, the darker bands (reddish-brown in color) about two and
one half scales wide, the lighter bands (straw yellow in color) about
one and one half scales wide; dark bands completely encircling tail
near tip; limbs straw yellow, with no evidence of markings.
Chest cream, with dark gray flecks; black nuchal collar complete
across ventral surface of neck, but very narrow; throat uniform
eaerulean blue, not barred; sides of belly pale to caerulean blue,
bordered medially by a black band which posteriorly involves a
39— €037
610 The University Science Bulletin
considerable area in groin and passes onto proximal portion of femur,
and anteriorly becomes narrower and disappears about one fourth
of the distance from the axilla to the groin ; preanal region pale blue,
with flecks of black; ventral surfaces of limbs pale yellow.
Variation. Detailed data on the character of the head scales of
fifty-two specimens from all parts of the range of the form show the
following variation: Frontal never contacting interparietal; fron-
toparietals contacting medially in fourteen ; an azygous scale separat-
ing frontoparietals medially in thirty-five (frontoparietal and fron-
tal scales indistinguishably broken up in the other three specimens) ;
frontoparietals divided into two on both sides in twenty specimens,
and on one side in thirteen ; the anterior frontal is divided longitudi-
nally in thirty-eight specimens; the posterior frontal is divided
longitudinally in five specimens, broken into three sections in seven
specimens, into four sections in four, and into five sections in one;
the parietal and frontoparietal of one side are fused in one specimen;
the supraoculars are usually in two rows, the outer composed of
scales smaller than the inner; in seven specimens the outer row is
represented by two or three scales segmented from the inner row,
and do not form a complete outer row as in the others ; one enlarged
supraocular narrowly contacts the median head shields on one side in
three specimens, on both sides in one; the row of scales separating
the supraoculars from the median head shields is supplemented by
the addition of one, two or three small scales, forming an incomplete
inner row, in eight specimens; the prefrontals contact medially in
thirty-seven specimens, are separated by contact of the median fron-
tonasal with the frontal in two specimens, and are separated by an
azygous scale in twelve; the preocular is separated from the rows of
scales above the supralabials on both sides in four specimens, on one
side in four; the two canthals are present in all, the first forced above
canthal ridge by contact of second canthal and subnasal on both sides
in three specimens, on one side in four; the anterior canthal touches
the rows of scales above the supralabials on both sides in five speci-
mens, on one side in two; the posterior canthal touches these rows in
one specimen ; the rows of scales below the subocular and above the
supralabials are reduced to one at some point on both sides in thirty-
eight specimens, on one side in four; the inner of the two series of
scales intercalated between the rows of mentals and the infralabials
continues forward usually to about even with the anterior half of the
third infralabial (on one side in nineteen, on both sides in twenty-
two) ; on one side in two specimens it extends to the middle of the sec-
ond infralabial; on one side in two, to the posterior part of the sec-
Smith: Genus Sceloporus 611
ond; on one side in six, to the suture between the second and third;
on both sides in one and on one side in five, to the middle of the third;
on one side in three, to the posterior part of the third; on one side
in six and both sides in one, to the suture between the third and
fourth; on one side in eight and both sides in one, to the anterior
part of the fourth ; on one side in one, to the middle of the fourth ;
on one side in one, to the middle of the fifth.
The dorsal scale rows are sometimes parallel, sometimes converg-
ing (two or three rows lost in eleven specimens, one in ten) . In two
the rows are actually diverging, a row being intercalated at about
the middle of the back and continuing to the base of the tail. In
several (four noted) the rows diverge anteriorly and converge pos-
teriorly, a short median row being intercalated.
The median lateral scales and the caudals near the base of the
tail are distinctly larger than the median dorsals in all.
Preceding the anus males frequently have several modified scales
appearing somewhat porelike.
The scales on the anterior margin of the ear are usually smaller
than those preceding them; only rarely is the upper scale slightly
larger.
The coloration is quite uniform and very characteristic. In no
other species of the torquatus group are the dark and light bands
about the tail so distinct. Toward the tip of the tail the bands are
visible upon the ventral caudal surface (not in cyanogenys) . Other
characteristics of the doreal coloration are, likewise, distinctive. The
nuchal collar is always complete, and about three scales wide. It
is bordered anteriorly and posteriorly by a light band about two
scales wide, the anterior of which extends laterally to the lateral
nuchal fold, and the posterior to the upper margin of the insertion
of the foreleg, but not passing onto the leg. Either or both light
borders rarely may be broken medially, but are never broken into
spots or appear as narrow bands. The black collar may extend
across the gular fold region, uniting ventrally. Anterior to the
nuchal collar is a light band about two scales wide across the pos-
terior part of the head, passing laterally to the upper edge of the
ear. This band may be broken into spots, but usually is distinct
and complete. A light band, usually visible, passes from the upper
labial region to the anterior margin of the ear.
Frequently, more often in males, there are light bands which pass
down the back along the middle of each longitudinal scale row, sepa-
rated from each other by narrow black bands. This type of color-
612 The University Science Bulletin
ation occurs also in mucronatus omiltemanus. The females fre-
quently lack this and may also have irregular and indistinct dark
spots on the back.
In males, the sides of the belly and the gular region are caerulean
blue, becoming darker with increased age. The sides of the belly
may become somewhat tinged with purple, and the gular region
marine blue, which increases in intensity posteriorly toward the
gular fold region. The blue of the sides of the belly is bordered
medially by a rather narrow black band, which expands posteriorly
and covers the entire area of the groin. The median abdominal
area is whitish, suffused with black in older males.
The gular region in females is irregularly marked with black.
Habits and habitat. This species seems to be confined entirely
to a rocky habitat. In Texas and New Mexico they are found on
limestone bluffs in considerable numbers, and are difficult to cap-
ture. Not only does their color, except for the brilliantly banded
tail, blend well with the color of the rocks, but they are exceedingly
wary, diving into cracks or under rocks at the first sign of danger.
Once hidden in cracks, it is only with the greatest difficulty that
they can be routed. Rarely are they discovered before they have
hidden under rocks or in cracks.
In Durango and Chihuahua specimens were secured on red gran-
ite ledges, where the lizards assumed a reddish hue quite similar
to the color of the rocks.
The species is ovoviviparous, as is indicated by a female in the
Museum of Zoology of the University of Michigan (No. 49846).
This specimen, collected May 31, 1916, contains ten well-developed
embryos. The young are presumably born during the early part of
June.
Remarks. The characters which distinguish poinsettii from cya-
nogenys are numerous, although for the most part minor. No in-
tergradation between the two forms has been observed in the speci-
mens examined, specimens of each form remaining quite distinct in
the areas of closest approach of their respective ranges. The type
locality of cyanogenys is Monterrey, Nuevo Leon, and specimens of
typical poinsettii are available from a locality thirty miles west of
Monterrey. In Texas, specimens of poinsettii are available from the
mouth of the Pecos river, Valverde county (and the type locality, in
part, is Devil's river), and specimens of cyanogenys from Eagle
Pass, Maverick county, not over seventy miles away.
Aside from color differences, which are quite striking and constant
both in live and preserved material, poinsettii is to be distinguished
Smith: Genus Sceloporus 613
from cyanogenys by the following points: supraoculars usually in
two complete rows, the inner row seldom incomplete and never lack-
ing; median cephalic scales usually very irregular, the anterior
frontal usually divided; the preocular rarely separated from the
rows of scales above the supralabials (10 in 96 in poinsettii, 60 in
78 in cyanogenys) ; the medial of the two rows of scales between
the postmentals and infralabials terminating farther posteriorly (in
9 counts of 96 in poinsettii the row extends anterior to the suture be-
tween the second and third infralabials; in cyanogenys, only 6 in 78
have the row extending posterior to that suture) ; the rows of scales
between the subocular and supralabials rarely not reduced to one
at some point (84 out of 96 in poinsettii have the rows reduced to
one; in cyanogenys, 16 in 76); femoral pores usually fewer. The
maximum size is also much greater in cyanogenys, the latter reach-
ing 143 mm. snout to vent, poinsettii, 118 mm. (KU 8185).
Range. Southern New Mexico east to central Texas', south
through western Nuevo Leon and southern Coahuila to central
Durango. (See Fig. 14 for distributional map.)
Locality records. United States: New Mexico: Eddy Co.:
Guadalupe Mts. (Mosauer, 1932; KU 14996), Guadelupe Mts.,
forty-four miles northwest of Carlsbad (AMNH 46389, Barnum
Brown), Dark Canyon, Guadelupe Mts. (MCZ 31728-9) ; Grant Co.:
Santa Rita del Cobre (Cope, 1900; Van Denburgh, 1922), Hachita
(Van Denburgh, 1922), Gatton's Park (LMK 24280-3, D. 0. Scott) ;
Lincoln Co.: Forks of Ruidoso Creek (Van Denburgh, 1922);
Otero Co.: Alamogordo (Van Denburgh, 1922), Sacramento Mts.
(ANSP 15163, Rehn & Viereck) ; Sierra Co.: Chloride, Grafton
(Van Denburgh, 1922); Socorro Co.: Magdalcna Mts. (Herrick,
Terry and Herrick, 1899; Van Denburgh, 1922), Socorro (Cope,
3 883; Van Denburgh, 1922). Texas: Bexar Co.: (Strecker, 1927) ;
Helotes (Taylor, 1931; Strecker, 1933; KU 8174-5); San Antonio
fAMNH 44411, R. B. Parks). Brewster Co.: (Strecker, 1909);
Chisos Mts., 6,000 ft. (Bailey, 1905) ; fifty miles south of Marathon
(Bailey, 1905) ; Glass Mts., five miles south of Marathon (KU 15009,
15001-4); three miles southwest of Alpine (Taylor, 1931; KU
14993-4, 15131, 15133, 15137-40) ; six miles west of Alpine (Taylor,
1931) ; Alpine (KU 8182-94) ; fourteen miles north of Terlingua
(Taylor, 1931, KU 8166-8168; EHT 2 spec). Cidberson Co.: Gua-
dalupe Mts. (Bailey, 1905; Mosauer, 1932). Brown Co.: San Saba
river, south of Brownwood (H. M. Smith). Duvall Co.: (Boulenger,
1897); San Diego (Cope, 1900; Strecker, 1915). Jeff Davis Co.:
614 The University Science Bulletin
six miles east of Valentine (KU 14998-15000) ; Davis Mts. (KU
18422), Davis Mts., 5,700 ft. (Bailey, 1905); Phantom Lake (UM
MZ 49846, H. T. Gaige). Kerr Co.: Japonica (Bailey, 1905).
Pecos Co.: Pecos (Stone and Rehn, 1903; ANSP 15768-84, A. E.
Brown) ; fifty miles southwest of Pecos (Brown, 1903) ; Fort Stock-
ton (Bailey, 1905). Presidio Co.: Paisano (Bailey, 1905). Real
Co.: West Frio Canyon (Strecker, 1935). Reeves Co.: near Toyah
(Bailey, 1905); Waihnacht's Draw, Pecos Valley (MCZ 31745).
Terrell Co.: Between Dryden and Sanderson (KU 15005-8). Val-
verde Co.: Rio San Pedro of the Rio Grande del Norte (now Devil's
river) (Baird and Girard, 1854; Yarrow, 1883; Cope, 1900; USNM
2948 [2 spec], J. H. Clark [cotypes]) ; five miles east of mouth of
Pecos river (KU 14995) ; near mouth of Pecos river (KU 14997).
Mexico: Chihuahua: (Cope, 1887, 1900; USNM 14233, 14243) ;
Ojo Caliente (Yarrow, 1883; Cope, 1900; USNM 2958) ; Chihuahua
City (Cope, 1886) ; ten miles south of Moctezuma, June 20, 1934
(DHD & HMS 104-113) ; eighteen miles north of Escalon, June 25,
1934 (DHD & HMS 246-256) ; Colonia Garcia (ANSP 1, Dr. W. E.
Hughes) ; Sierra Madre (USNM 47419-21, Nelson and Goldman) ;
Balleza (USNM 47417, Nelson and Goldman) ; Casas Grandes
(FMNH 1655, C. H. T. Townsend) ; Samachique (FMNH 11841-7,
15724 [18 spec], R. M. Zingg) ; Arroyo del Alamos (USNM 42873,
E. C. Erdis) . Coahuila: Monclova (Carman, 1887 ; USNIM 46699,
Nelson & Goldman) ; Santa Caterina (Cope, 1900) ; fifteen miles
east of Saltillo, July 31, 1934 (DHD & HMS 1141) ; Sierra Guade-
lupe (USNM 47591-2, Nelson and Goldman) ; Jaral (FMNH 1547,
Heller & Barber). Durango: (MCZ 16030, P. Townsend; USNM
23992, Dr. E. Palmer); Ciudad (Glinther, 1890; Boulenger, 1885,
1897) ; six miles northeast of Pedriceha, August 27, 1932 (EHT &
HMS 4361, 4362, 4364-4371) ; near Pedricena, August 29, 1932 (EHT
& HMS 4456-4470) ; near Pasaje, August 28, 1932 (EHT & HMS
4456); Coyotes (FMNH 1510 [10 spec], Heller & Barber); Du-
rango (USNM 46844-5, Nelson & Goldman). Nuevo Leon: thirty
miles west of Monterrey, July 31, 1934 (DHD & HMS 1142).
The above locality records appear to be reasonably correct. There
are many more which do not so appear. They are from Arizona
(Coues, 1875; Yarrow, 1875, 1883; Yarrow and Henshaw, 1878;
Cope, 1900; MCZ 6831 [Flagstaff?]); California (Boulenger, 1895;
KU 11383) ; Colima (Bocourt, 1874) ; Guerrero (Boulenger, 1897) ;
Jalisco (Duges, 1870) ; Mexico (Duges, 1887) ; Michoacan (Duges,
1887) ; Oaxaca (Bocourt, 1874) ; San Luis Potosi (Carman, 1887) ;
Smith: Genus Sceloporus
615
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Smith: Genus Sceloporus 617
Sonora (Baird and Girard, 1854; Baird, 1859; Yarrow, 1883; Gar-
man, 1884; Cope, 1900; Van Denburgh, 1922; USNM 2970, Col.
Graham; USNM 2952 [2 spec], J. H. Clark [cotypes]) ; Tamaulipas
(Burt, 1932) ; and Vera Cruz (Boulenger, 1885, 1897).
Sceloporus lineolateralis Smith
(Text Figs. 16 and 17; Plate LII)
Sceloporus lineolateralis Smith, 1936, pp. 92-95.
Type locality. Six miles northeast of Pedricena, Durango, Mex-
ico. Holotype EHT & HMS 4323; paratypes EHT & HMS 4321-2,
4324-6, 4332-40, 4342-3, 4363, 4372a and b, 4471-6.
Diagnosis. A species of moderate size belonging to the torquatus
group; dorsal head scales smooth; lateral scales about half as large
as median dorsals, but not strongly differentiated from them ; ventral
scales about one third as large as laterals; longitudinal rows of dor-
sals converging toward median line; thirty-eight to forty-seven
scales from occiput to base of tail; length of tibia about equal to
length of shielded part of head; length of fourth toe about equal to
distance between snout and posterior margin of ear; hind limb
reaches to ear; fourteen to twenty femoral pores, not extending onto
preanal region; enlarged supraoculars in a single row; two canthals;
a narrow, uninterrupted black collar on neck, arising from shoulder,
with a light, sometimes medially interrupted, posterior border; sides
of belly in males china blue, very slightly darker medially; gular
region very pale blue.
Description of type. Dorsal head scales smooth; a single row of
enlarged supraoculars, bordered medially by a single row of small
scales, and separated from the superciliaries by one complete and
another incomplete row of scales; seven superciliaries, the last over-
lapping the preceding two (the sixth superciliary is entirely over-
lapped by the fifth) ; one large, heavily keeled subocular, followed
posteriorly by a series of seven scales which curves upward about
the orbit, contacting the parietal; lower two postocular scales
strongly keeled; the series of scales between supraoculars and fron-
tals continued downward between postoculars and margin of orbit,
terminating at the second postocular (from subocular) ; preocular
scale strongly keeled, in contact with second canthal and with the
rows of scales above supralabials; loreal larger than subnasal, smaller
than preocular, in contact with both canthals; two rows of low,
elongate scales above supralabials, the lower row continuing around
snout, passing above rostral, the other terminating anteriorly below
the subnasal; two canthals, the second with but a very small por-
618
The University Science Bulletin
tion entering the superciliary series; rostral low, about three times
as broad as high; interparietal at least twice as large as parietal,
separated from frontal by a pair of frontoparietals in contact medi-
ally; a single large parietal on each side; frontal divided trans-
versely, the anterior section about one half larger than posterior; two
prefrontals, broadly in contact on median line, larger than anterior
section of frontal; three frontonasals, the median about as large as
either prefrontal ; the lateral frontonasals somewhat smaller, in con-
tact with both loreals; three irregular pairs of internasals, the pos-
FiG. 16. Head scales of Sceloporus lineolateralis Smith. EHT & HMS 4323,
six miles northeast of Pedricena, Durango; actual head length, snout to occiput,
15 mm.
terior pair the largest and in contact with the frontonasals, the an-
terior pair smallest and in contact with the row of scales behind
rostral; a single small scale separates the posterior internasals from
anterior loreal, but does not enter series about nasal ; latter broadly
in contact with subnasal, narrowly in contact with row of scales
above rostral, and surrounded above by four small scales; four
supra- and five infralabials to a point below the middle of the eye.
Mental pentagonal, its labial border about two thirds that of
rostral; mental followed by a series of four or five enlarged scales
on each side, the anterior in contact with its fellow on the median
ventral line; this series of scales separated from infralabials an-
teriorly by a single row of elongate scales (except anterior, which
Smith: Genus Sceloporus 619
is narrowly in contact with anterior infralabial) , posteriorly by two
rows, beginning below the middle of the third infralabial; scales on
dorsal and lateral surfaces of head (including temporal region)
more or less pitted; scales in temporal region keeled, mucronate in
area near ear; four or five rows of scales on neck behind occiput
nearly smooth, not or but weakly mucronate; anterior margin of
ear with four smooth, pointed scales, the upper largest and larger
than preceding scales; lateral neck fold low, relatively shallow, a
weak fold from its upper margin to lower margin of ear; the scales
surmounting this fold no larger than those adjacent, much smaller
than those in the lateral gular region ; median anterior gulars smaller
than posterior median gulars; lateral gulars in region of angle of
jaws much larger than median gulars; scale rows on neck parallel,
abruptly differentiated from the scale rows arising near the insertion
of the foreleg and passing obliquely upward onto back; median
dorsals weakly keeled, mucronate, usually with not over one lateral
mucrone, except on neck, where there may be as many as two;
median dorsals about twice as large as laterals, gradually differ-
entiated from them ; laterals three or four times as large as median
vcntrals midway between fore and hind limbs; laterals more
strongly keeled and mucronate than dorsals, with as many as three
lateral mucrones; scales in axilla very small, smooth, rounded; scales
following axilla gradually increasing in size and becoming more
strongly mucronate, but smooth for about one third the distance
from axilla to groin.
Dorsal scales of upper and lower foreleg keeled and mucronate, the
latter somewhat smaller than dorsals on upper foreleg; dorsal scales
on hand and toes smooth or very weakly keeled, the former slightly
mucronate, some of the latter bi- or tricuspid; one dorsal, two lateral
and one ventral row of scales around fingers and toes, all con-
tinuous to tip except the two laterals, which terminate behind the
last scale of the other series ; ventral scales of upper and lower fore-
leg smooth, the latter about twice as large as former; scales of palm
and sole weakly keeled and mucronate; ventral lamellae with three
or five mucrones, about equal in size; lamellar formula for fingers,
9-13-16-16-14 (9-13-16-15-14).
Dorsals of tibia and femur strongly keeled, mucronate, about the
size of median laterals; anterior ventrals of femur and ventrals on
tibia subequal in size; ventrals on femur decreasing in size toward
femoral pores; lamellar formula for toes 7-12-19-21-18 (8-12-18-
?-17) ; breast scales much larger than median ventral abdominal
scales; median ventrals increasing in size laterally, merging with
620 The University Science Bulletin
laterals ; anterior preanal scales about equal in size to those anterior
to series of femoral pores ; a pair of enlarged postanals ; dorsal scale
rows converging on base of tail ; scales behind femoral pores, except
one or two rows immediately adjacent to femoral pores, keeled, with
three mucrones, of about the same size as those preceding the pore
scries; dorsal scale rows of tail somewhat larger than median dor-
sals on body, more strongly keeled and mucronate.
Color. Back, dorsal surface of limbs, and temporal region of head
glaucous to pale blue ; sides of body tinged with gray ; a narrow black
collar two or three scale rows wide on neck, arising almost straight
from shoulders and not continued onto ventral surfaces; anterior
and posterior borders of collar parallel, the latter with a light edge
one or two scales wide; a few indistinct light spots on neck; sides of
belly from axilla to groin china blue, lighter toward axilla, darker
posteriorly ; a small black area in groin, median edges of lateral blue
areas very slightly darker, with no less than five rows of cream-
colored scales separating them; gular region and breast very pale
blue, the breast suffused with gray; ventral surfaces of limbs and
tail cream.
Variation. Enlarged supraoculars invariably a single series; in
three specimens two incomplete, as well as one complete, rows of
small scales between supraoculars and superciliaries, latter usually
six in number; the row of postoculars from the posterior border of
the subocular to the temporal is irregular, but always two rather
strongly keeled scales follow the subocular; the series of scales in-
tercalated between the supraoculars and frontal is continued around
the posterior border of the orbit in all specimens; the loreal con-
tacts both canthals in all but one specimen (one side only) ; the
preocular invariably touches the second canthal, but in ten speci-
mens the lower portion is broken off to form an extra scute separat-
ing the remainder of the preocular from the rows of scales above
supralabials; these latter rows of scales irregular, sometimes but a
single scale separating medially the subocular from the supralabials ;
in all but one specimen one of the rows (apparently the upper one
as a rule) is continuous around the snout above the rostral; in this
exception the median scale above the rostral is apparently fused with
the latter, the anterior pair of internasals thus contacting the ros-
tral; twenty-one specimens have the frontoparietals separated, the
frontal and interparietal in contact; the prefrontals and frontonasals
are as in the type in all specimens ; the internasals and scales about
the nasals, however, are too irregular for description of variation.
Smith: Genus Scelopoeus 621
although very frequently (perhaps normally) they are as in the
type ; character of scales on body and limbs uniform throughout.
The coloration of the males is practically identical with that of
the type. A light line, barely visible in the type, passes from the
upper labials directly through the ear to the black nuchal collar,
passing above the lateral cervical fold; this line is more distinct in
the young males, and is bordered above by a narrow black line
arising on the posterior margin of the orbit and passing through the
extreme upper edge of the ear and thence to the black collar. This
line is variable and is most distinct in the young. In ventral color-
ation there is but little variation. The smallest male (44 mm. snout
to vent) has very distinct lateral abdominal areas of blue, but the
gular region and breast are white. In a male 54 mm. from snout
to vent the gular region is as in the type, but with small scattered
white flecks. The breast and median abdominal regions become
grayish with increased size.
The females are markedly different from the males in coloration
of the back. A light (whitish) line passes from above the insertion
of the foreleg along the side of the body to the groin; above this is
an irregular black line formed by the fusion of large black spots,
which project upward from the black line; the latter arises from the
upper margin of the insertion of the foreleg, separated from the black
neck collar by the light posterior border of the latter; below the
lateral hght line is a narrow, indistinct and incomplete black line;
on each side of the back is a row of large black spots, about six on
each side; the ventral surface of the body, with the exception of
the region anterior to the gular fold, is grayish.
Of the other described species of the torquatus group, only ;. jar-
rovii, torquatus, melanog aster, bulleri, serrifer and lineolateralis have
the supraoculars large, in a single row. S. guentheri may be dis-
tinguished by having a much longer fourth toe. S. serrifer, torquatus
and melanogaster have much larger dorsal scales, with the lateral
dorsals larger than the median dorsals. S. j. jarrovii differs not
only in having the dorsal scale rows parallel, but in having the lat-
eral scales larger, much shorter hind legs, different coloration, etc.
S. bulleri has shorter hind legs, larger scales on the body, a broad
neck band and much different ventral coloration.
It is of interest that in at least five of these species (;. jarrovii, tor-
quatus, melanogaster, serrifer, and lineolateralis) the normal or very
frequently occurring condition is for the frontal to contact the inter-
parietal. This condition may occur in guentheri (not stated in de-
622
The University Science Bulletin
scriptions), but in most of the forms with two rows of supraoculars
it occurs rarely (frequently in dugesii).
In body proportions, the narrow nuchal collar, the shape of the
tail, and in various relationships of the head scales, lineolateralis
suggests formosus. The nine species of this group (formosus Wieg-
FiG. 17. Distribution of Sceloporus lineolateralis Smith, S. jarrovii jarrovii
Cope, S. jarrovii minor (Cope) and S. jarrovii immiicronatus Smith.
mann, irazuensis Giinther, lunaei Bocourt, malachiticus Cope, sal-
vini Giinther, schmidti Jones, smaragdinus Bocourt, taeniocnemis
Cope and viviparus Cope) which have been described, are not, how-
ever, comparable by description to the species here described, nor
has direct comparison with many specimens of the formosus group
from many localities in Central America and Mexico shown any
close relationship to these species with lineolateralis.
Remarks. In some respects lineolateralis resembles jarrovii, a
species which is closely associated geographically. It is possible
Smith: Genus Sceloporus
623
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624 The University Science Bulletin
that Boulenger's (1897) and Gunther's (1890) specimens from
Ciudad, Durango, are actually of lineolateralis and not of jarrovii.
Range, Known only from Durango.
Locality records. Six miles northeast of Pedricena, Durango
(EHT & HMS 4321-6, 4332-4340, 4342-3, 4363, 4372a and b; four-
teen miles northeast of Pedriceiia, Durango (EHT & HMS 4471-6) ;
Guadelupe, Mexico (MCZ 6807 [3 spec], Comision geografico ex-
ploradora).
Sceloporus jarrovii jarrovii Cope
(Text Figs. 17 and 18; Plate XL VII, Fig. 1)
Sceloporus jarrovii Cope, 1875, p. 48; idem, 1875, pp. 569-571, pi. 23, 2-2d; Yarrow,
1883, p. 57; Garman, 1884, p. 17; Cope, 1885, pp. 396, 403; idem, 1887, p. 38; Van Den-
burgh, 1896, p. 342; Cope, 1900, pp. 335, 341, 345-347, fig. 49; Stejneger, 1902, p. 150;
Stone, 1911, p. 227; Bailey, 1913, p. 34; Van Denburgh and Slevin, 1913, pp. 392, 403;
Stejneger and Barbour, 1917, p. 54; Van Denburgh, 1922, pp. 321-326, pi. 26; Stejneger and
Barbour, 1923, p. 56; Van Denburgh, 1924, p. 207; King, 1932, p. 177; MacCoy, 1932,
pp. 19-21; stejneger and Barbour, 1933, p. 62.
Sceloporus yarrowi Coues, 1875, p. 595-6, pi. 23, figs. 2-2c.
Sceloporus yarrovii Boulenger, 1885, pp. 223-224; Giinther, 1890, pp. XIII, 69; Boulenger,
1897, pp. 483-485 (part); Ditmars, 1907, pp. 129, 135.
Sceloporus torquatus poinsettii Cope, 1900, pp. 350-352 (part).
Type locality. Southern Arizona. Cotypes USNM 8494 (2 spec.)
and 8495.
Diagnosis. A Sceloporus of the torquatus group; dorsal scales,
38 to 46 from occiput to base of tail; dorsal scales denticulate,
weakly carinate; largest lateral scales somewhat larger than median
dorsals; ratio of hind leg to snout- vent measurement, 54.1 to 65.8;
ratio of fourth toe to snout- vent measurement, 18.8 to 25.4; enlarged
supraoculars in a single series, bordered laterally by three rows of
scales, two or three of which are incomplete; the inner of these
three rows is composed of the largest scales; frontal usually in con-
tact with interparietal, or separated by an azygous scale; prefrontals
usually in contact; first canthal seldom forced above canthal ridge
by contact of second canthal and subnasal; preocular usually not
divided; usually two complete rows of scales below subocular. Black
nuchal collar three or four scales wide, with a narrow light border
sometimes not well defined; a suffusion of black confluent with the
collar may occur over the neck and back; a light line two scales
broad passes from the nuchal collar to the temporal region; scales
on rest of dorsal surface black, with a light spot in the center of
each scale.
Description (from K. U. 13145, male). Head scales smooth,
strongly pitted in prefrontal and internasal regions; head not
strongly depressed; interparietal large, fully four times the size of
either parietal; parietal single on each side, subtriangular ; fron-
Smith: Genus Sceloporus
625
toparietals single on each side, separated medially by contact of
frontal and interparietal; posterior section of frontal about one
third smaller than anterior section; prefrontals in contact medially;
median frontonasal one third larger than either lateral frontonasal;
nostril pierced slightly posterior to middle of nasal scale; anterior
part of nasal much narrower than part surrounding naris ; nasal and
internasals separated from rostral by a row of small scales; scales
in internasal region quite irregular; five-six enlarged supraoculars,
separated from median head scales by a row of small scales;
supraoculars separated from superciliaries by three rows of small
Fig. 18. Head scales of Sceloporus jarrovii jarrovii Cope. KU 12371,
Montezuma Canyon, Huachuca Mts., Arizona; actual head length, snout to
occiput, 17.5 mm.
scales, one complete and two incomplete; two canthals, the anterior
smaller than the posterior; second canthal contacting subnasal on
one side; subnasal somewhat larger than loreal; preocular not di-
vided; subocular followed posteriorly by two small, strongly keeled,
well-differentiated postoculars; two rows of lorilabials on sides of
head, not reduced to one row^ at any point below subocular; four
supralabials and five infralabials to a point below middle of eye.
Mental with a labial border about two thirds that of rostral;
mental pentagonal, followed by a series of postmentals, only the
anterior two or three of which are well differentiated; scales of
anterior pair of postmentals in contact; outer row of labiomentals
feparated from mental by narrow contact of first infralabial and
40—6037
626 The University Science Bulletin
first postmental; inner row of labiomentals terminating anteriorly
at a point below posterior part of second infralabial; gular scales
smooth, gradually increasing in size posteriorly; anterior gular
scales rounded, not strongly imbricate ; posterior gular scales strongly
imbricating, with a single apical notch.
Three or four well-differentiated auricular lobules, the upper
largest, rounded, the others pointed; scales in temporal region weakly
keeled, weakly mucronate (more strongly mucronate near ear),
somewhat smaller than largest scales between ear and lateral nuchal
fold; scales in latter area keeled and very strongly mucronate;
lateral nuchal fold relatively shallow; dorsal scales weakly keeled,
weakly mucronate, those on neck and between shoulders without a
well-differentiated apical mucrone; dorsal scales denticulate, their
posterior edges with a somewhat truncate instead of rounded con-
tour; lateral scales of body more strongly keeled, more strongly
mucronate, more denticulate and not truncate ; ventral scales slightly
more than half the size of median dorsal scales, those on breast equal
to or slightly smaller than those in middle of abdomen; preanal
scales slightly more than half the size of scales in middle of ab-
domen; subcaudals smooth at base of tail; postanals enlarged, sepa-
rated by four small scales; dorsal scales between hind legs some-
what smaller than preceding scales, one third or one fourth as large
as scales on base of tail.
Dorsal scales of foreleg mucronate and weakly keeled, those on
upper foreleg somewhat larger than those on lower foreleg and
slightly smaller than median dorsals; ventral scales of lower fore-
leg subequal in size to dorsals of same member, smooth, mucronate;
ventral scales of upper foreleg smaller than those of lower foreleg,
smooth, notched; lamellar formula for fingers 9-13-15-15-12 (8-13-
15-15-11).
Dorsal scales of hind leg keeled and mucronate, those on shank as
large as or slightly larger than median dorsals on body, and dis-
tinctly larger than those on thigh ; ventral scales of shank somewhat
smaller than dorsals of same member, smooth, mucronate; ventral
scales of thigh subequal in size to preanal scales, smooth, notched;
scales on posterior surface of thigh somewhat larger than preanal
scales, keeled, weakly mucronate, abruptly decreasing in size im-
mediately posterior to femoral pore series; no postfemoral dermal
pocket; lamellar formula for toes 8-12-16-19-14 (8-12-17-18-14).
Color. General dorsal ground color black; head reddish-brown,
with a light yellow-brown spot on nearly every scale ; black nuchal
collar three or four scales broad, with a broad median extension
Smith: Genus Sceloporus 627
about six scales wide from its anterior border to occiput; lateral to
this black band a pale-blue line two scales wide, passing from an-
terior edge of black collar to posterior corner of eye ; lateral to this,
an indistinct dark band confluent with black collar posteriorly,
passing to eye anteriorly; this is followed laterally by pale blue,
which color continues on sides of head and neck; black nuchal
collar continuous around throat; no distinct light posterior border
for collar; each scale on back and sides with a large median pale-
blue spot; limbs with narrow, indistinct dark bands; tail dimly
banded.
Ventral surface of head pale blue, somewhat darker than sides
of head; middle of abdomen cyanine blue, the color extending onto
preanal region and ventral surface of femur, becoming somewhat
lighter; no light median abdominal band, the blue color of the sides
fusing medially and leaving isolated median areas lighter in color;
a black patch in groin; a small area in immediate area about anus
cream-colored; ventral surface of tail pale blue, with an indefinite
suflusion of cream medially.
Variation. The variation in head scales of thirty-six specimens
is as follows: The frontal contacts the interparietal in sixteen, and
is separated by an azygous scale in eleven, by contact of the two
frontoparietals in eight (one specimen injured) ; the anterior frontal
is longitudinally divided in one; the median frontonasal contacts
the frontal in five, is separated by an azygous scale in two, and by
contact of the prefrontals in twenty-nine ; the second canthal touches
the subnasal on both sides in six, on one side in seven; the first
canthal is always present; the supraoculars are always in a single
series, always with a few small scales segmented off the series of en-
larged scales, but not approaching the size of the latter; scales be-
tween supraoculars and superciliaries in three incomplete rows; row
of scales bounding supraoculars medially always complete, the length
of each scale about equal to its width; preocular scale separated
from the two rows of scales above supralabials in ten (one side only
in two) ; subocular separated from supralabials by two complete rows
of scales, occasionally (in nine) reduced to one by one scale; pos-
terior fourth of subocular segmented off on both sides in two, on one
side in two; outer row of labiomentals touching mental in two,
separated by a partial contact of first infralabial and first postmental
in remainder; inner row of labiomentals extends anteriorly to a point
even with the suture between the second and third infralabials,
never extending farther than the middle of the second infralabial,
628 The University Science Bulletin
not beginning farther back than the middle of the third infralabial.
The second pair of postmentals contacts medially in one specimen.
The femoral pores vary between thirteen (one specimen, one side)
and eighteen, and the dorsal scales from occiput to base of tail are
usually forty to forty-six; three specimens from Moctezuma, Chi-
huahua, have a higher average (47, 48, 52).
Specimens from various localities in the mountains of western
Chihuahua have a peculiar variation in color pattern which I have
not observed in specimens from other localities. The back and neck
are strongly suffused with black, which is continuous with the black
nuchal collar. This in turn has lost its light borders medially. Ap-
parently no other differences accompany this color variation. Cer-
tain specimens from the same localities have a normal color pattern
similar to that found in Arizona specimens. As a general rule the
pattern so characteristic of Arizona material, of a light spot on each
scale on the back, is usually not evident on Mexican specimens.
Stejneger (1902, p. 150), however, remarks that under certain en-
vironmental conditions the spotted pattern is not evident even in
Arizona specimens.
Habits and habitat. Ecologically, jarrovii jarrovii seems to be
confined for the most part to areas of high elevation. The three
specimens collected near Moctezuma, Chihuahua, are the only ones,
so far as I am aware, which have been found at a low elevation.
These were found in a semiarid region, on low, barren hills capped
by large, granite boulders. They were in company with S. poinsettii.
Range. From central Arizona east to western New Mexico, and
south through Chihuahua and western Sonora to extreme western
Zacatecas and extreme northern Tepic. (See fig. 17 for distributional
map.)
Locality records. Arizona: Cochise Co.: (Yarrow, 1883, Van
Denburgh, 1896, 1922; Boulenger, 1897; Cope, 1900; Stejneger, 1902;
Stone, 1911; Van Denburgh and Slevin, 1913; K. U. 6885-6919, 7018-
7039, 7091-7093, 7311-7336, 7521-7533, 10836, 11840-11872, 12349-
12429, 12431-12441). Pima Co.: (Van Denburgh, 1922; King,
1932). Santa Cruz Co.: (Cope, 1900; A^an Denburgh, 1922; Mac-
Coy, 1932; K. U. 13143-13156). Navajo Co.{f): (Yarrow, 1883;
Cope, 1900). New Mexico: (Bailey, 1913) ; Hidalgo Co.: Animas
Peak and Big Hatchet Mts. (Van Denburgh, 1924). SoxXora:
Pinetos Camp, thirty-two miles south of Nogales (Van Denburgh,
1922). Chihuahua: Ten miles south of Moctezuma (DHD &
HMS 98-100); Samachique (FMNH 15723 [6 spec], 11829-39,
Robert M. Zingg) ; Colonia Garcia (FMNH 1658, C. M. Barber;
Smith: Genus Sceloporus 629
USMN 46660-1, Nelson & Goldman; MCZ 7476, C. S. Brimley) ;
Sierra Madre (USNM 47422, 47234-8, Nelson & Goldman) ; San
Luis Mts. (USNM 47015, E. A. Goldman; USNM 21032, E. A.
Mearns; Guadelupe (USNM 47066, Nelson and Goldman) ; Meadow
Valley (USNM 26600, Brimley) ; Madera (MCZ 17526-7, W. W.
Brown, 4,400 ft.); Pacheco (MCZ 15610-4, W. W. Brown). Du-
RANGo: Hiiasamota (USNM 46930, Nelson & Goldman) ; El Salto
(USNM 46638 Nelson & Goldman); Coyotes (FMNH 1511 [19
spec]. Heller & Barber); Cuidad (Boulenger, 1885, 1897; Cope,
1887; Giinther, 1890). Zacatecas: Sierra Madre (USNM 46934,
Nelson & Goldman). Tepic: Santa Teresa (USNM 46630, Nelson
& Goldman).
Remarks. A specimen reputed to be from Monclova (probably
meant for Coahuila) (MCZ 4557), collected by Palmer in 1880, ap-
pears to have incorrect locality data. It was in a jar with a speci-
men presumably from the same locality, but which possesses a tag
stating "San Luis Potosi," and which is typical jarrovii minor. The
untagged specimen is well preserved and not faded, while the other
is shrunken and faded. Until further collecting indicates the pres-
ence of jarrovii in this region of Coahuila, I assume the locality data
to be incorrect.
Six specimens are present in Field Museum (384), collected by
E. S. Walton in August, 1897, at Fort Bliss, Arizona (Texas?).
The validity of the report of jarrovii from "North of Rio San-
tiago, Jalisco," cannot bfe definitely accepted because of the con-
fusion by Boulenger of dugesii dugesii and jarrovii, and because of
the possibility of erroneous locality data. The table of characters
given by Boulenger (p. 484) indicates that the specimen not form-
ing the type of pleurolepis may be jarrovii, but the pleurolepis type,
from the same locality, almost certainly is dugesii dugesii. Boulen-
ger has other specimens from Duvall Co., Texas (locality incorrect,
or the specimen is poinsettii) , Catorce, San Luis Potosi (probably
jarrovii minor), and from La Cumbre de los Arrastrados, Jalisco,
Ixtlan, Jalisco, and La Venta, Guadalajara, all of which specimens
are very probably d. dugesii.
S. jarrovii seems to be most closely related to d. dugesii. However,
the two forms are sufficiently different that it seems unwise to place
them as subspecies until intergradation is definitely established as
a fact.
Some of the differences which may be pointed out are: supra-
oculars entire in jarrovii, partly divided in dugesii; prefrontals usu-
630
The University Science Bulletin
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ally in contact in jarrovii, usually separated in dugesii; rows of
scales between subocular and supralabials usually reduced to one
at some point in dugesii, usually two complete rows in jarrovii; dor-
sal head scales smooth (although pitted) in jarrovii, rugose (micro-
scopically) in dugesii; dorsal scales from occiput to base of tail
usually more numerous in dugesii (41-50) than in jarrovii (40-46
recorded by Van Denburgh [1922], average 43; the three specimens
of jarrovii from Chihuahua are exceptional and have 48, 47 and
52) ; femoral pores fewer in dugesii (9-13) ; posterior frontal not re-
duced in jarrovii as in dugesii. The light spot in the center of each
scale is missing in dugesii.
Sceloporus jarrovii minor (Cope)
(Text Figs. 17 and 19; Plate LIII, Fig. 2)
Tropidolepis poinsettii 7 DugSs, 1870, p. 243.
Sceloporus torquatus minor Cope, 1885, p. 402; idem, 1887, p. 38; idem, 1900, pp. 353,
349.
Sceloporus poinsettii ? Duges, 1887, p. 114; Garman, 1887, p. 14.
Sceloporus yarrovii Boulenger, 1897, pp. 483-485 (part).
Type locality. Zacatecas. Cotypes USNM 26166-7 ; Duges, col-
lector.
Diagnosis. A species of moderate size belonging to the torquatus
group; dorsal head scales smooth; largest lateral scales subequal in
size to, or somewhat larger than, dorsals, ventrals much smaller
than (about one fourth as large as) dorsals; femoral pore series
short, not extending onto preanal region, sometimes with an ex'tra
short series; dorsals 34 to 46 from occiput to base of tail; dorsal
scales smooth or very weakly keeled, veiy weakly mucronate, in
parallel or very slightly converging rows ; tibia as long as or longer
than shielded part of head; fourth toe from base of fifth equal to
or greater than distance from snout to ear; supraoculars usually in
two complete rows, never in one row; black collar continuous around
gular region, on dorsal surface of neck very broad, covering from
six to eight scale rows; collar with narrow light borders, sometimes
interrupted ; adults with a ground color of uniform seal brown, with
distinct, narrow light bands on distal portion of tail, not encroach-
ing upon ventral surface; males with sides of abdomen from axilla
to groin pale blue, sometimes with a purple suffusion, the median
edges and area in groin black or marine blue; throat pale blue,
sometimes spotted.
Description (EHT & HMS 4207, San Felipe, Guanajuato). Dor-
sal head shields smooth ; enlarged supraoculars in two rows, bordered
medially by a single row of small scales, and separated from the
632
The University Science Bulletin
superciliaries by one complete and another incomplete row of scales;
six superciliaries, the last overlapping the preceding two (the fifth
superciliary is entirely overlapped by the fourth) ; one large, heavily
keeled subocular, followed posteriorly by two elongate, heavily
keeled postoculars; the series of scales between supraoculars and
frontals continued downward, bordering posterior margin of orbit,
and contacting the upper (posterior) postocular; preocular scale
strongly keeled, in contact with second canthal and with the rows of
Fig. 19. Head scales of Sceloponts jarrovii minor (Cope). EHT & HMS 4207,
near San Felipe, Guanajuato; actual head length, snout to occiput, 17.5 mm.
scales above supralabials ; loreal very slightly larger than subnasal,
somewhat smaller than preocular, in contact with both canthals;
two irregular rows of scales above supralabials and in contact with
subocular, preocular, loreal and subnasal, one row continuous
around snout above rostral; two canthals, the second with but a
very small portion entering the superciliary series; rostral low, at
least three times as broad as high; interparietal much larger than
either parietal, roughly pentagonal, the posterior edge straight, the
sides nearly parallel; a single parietal on each side, but little larger
than frontoparietal; a pair of frontoparietals, in contact on median
line and separating frontal from interparietal; frontal transversely
divided; two prefrontals, in contact on median line; three fronto-
Smith: Genus Sceloporus 633
nasals, the lateral in contact with both canthals; two pairs of in-
temasals, the posterior pair somewhat larger than the anterior, the
former separated from the first canthal by a small scale; six to seven
scales, including the subnasal, surround the nasal; four supra- and
five infralabials to a point below the middle of the eye.
Mental pentagonal, its labial border about two thirds that of
rostral; mental followed by a series of five or six enlarged scales on
each side, the anterior in contact with its fellow on the median
ventral line; this series of scales separated from infralabials an-
teriorly by a single row of elongate scales (except anterior, which is
narrowly in contact with anterior infralabial) , posteriorly by two
rows, the second row beginning below the middle of the third in-
fralabial and in contact with the series of postmental scales; scales
in temporal region keeled, mucronate in area near ear; anterior
margin of ear with five smooth, pointed scales, the upper two larger
than the preceding scales; lateral cervical pouch rather shallow; a
fold of skin from about its middle to the lower margin of the ear,
surmounted by enlarged, pointed scales; scales adjacent to this
fold and gular scales in region of angle of jaws subequal in size;
dorsal scales in parallel rows, the scales near the median line very
weakly keeled or smooth, rounded or very weakly mucronate; scales
on sides very weakly keeled, with one to five weak mucroncs ; scales
between axilla and groin smooth, but with five or seven mucrones;
largest dorsal scales on body about half as large as largest dorsal
caudals; scales in anterior median part of region in front of gular
fold smaller than scales near gular fold; the latter scales but slightly
smaller than those near angle of jaw; scales on breast largest of
ventrals on body; median abdominal scales somewhat smaller than
lateral abdominals.
Dorsal scales of body somewhat larger than those on dorsal surface
of upper foreleg, these somewhat larger than those on dorsal surface
of lower foreleg; dorsal scales of foreleg weakly keeled and mu-
cronate, those on hand nearly smooth; ventral scales of foreleg
smooth, mucronate, those on lower foreleg as large as or larger than
the dorsals of the same member, those on upper foreleg much
smaller than those on lower foreleg; one dorsal, two lateral and one
ventral row of scales around fingers and toes, all continuous to tip
except the two laterals, which terminate behind the last scale of the
other series; lamellar formula for fingers ?-13-18-17-12 (8-15-18-18-
14). Dorsal scales of hind limb keeled, mucronate, those on tibia as
large as largest dorsals on body, those on femur somewhat smaller;
ventral scales of tibia and femur smooth, rounded, those on tibia
634 The University Science Bulletin
somewhat smaller than dorsal scales of same member; scales on
anterior surface of femur smooth, large as dorsals of same member;
ventral scales of femur, near femoral pores, about equal in size to
preanals; scales on posterior surface of femur, immediately behind
femoral pores, much smaller than preanals; scales in median area of
posterior surface of femur about four times as large as those im-
mediately behind pore series; lamellar formula for toes 10-14-18-
20-14 (9-14-20-21-16); a pair of enlarged postanals; subcaudals
smooth near proximal end of tail, becoming weakly keeled toward
distal end ; dorsal caudals more strongly keeled than scales on body,
mucronate.
Color. Ground color above uniform olive to Prout's brown; no
markings on back or limbs ; neck traversed by an extremely broad,
black band passing over the shoulder and dimly crossing gular fold
region ; the band covering seven or eight scales in the median dorsal
line; collar light-bordered, the posterior border narrow (one scale
wide), the anterior border broader (two or parts of two scales wide) ;
a broad, light band about two scale rows wide between upper edge of
ears, separated by a dim, bluish-gray band from the light anterior
border of neckband; posterior border of neckband very strongly con-
vex, the median part rather angular, the anterior border straight;
head clove to seal brown; sides of belly from axilla to groin pale
blue, with a narrow median border of black; a small area in groin
black; breast, a median abdominal band about four to six scale rows
wide, ventral surface of limbs, preanal region and ventral surfaces of
tail whitish; region of gular fold gray; area in front of this gray
intermingled with blue, and with small, scattered white flecks.
Variation. The following variation of fifty-four specimens was
noted: The frontoparietals contact medially in ten; they are sepa-
rated by an azygous scale in forty-one, and by contact of the frontal
with the interparietal in three. The frontoparietals are divided into
two on both sides in four, on one side in three. The anterior frontal
is divided longitudinally in ten specimens. The posterior frontal is
divided into four scales in one specimen, into three in six, and into
two scales in three specimens. An azygous scale separates the pre-
frontals in three; they are separated by contact of the median
frontonasal and the anterior frontal in five; in the remainder they
contact medially. These primary head shields vary but little in
comparison with poinsettii; fusion and irregular segmentation do not
frequently occur, and never to the degree common in the latter form.
The supraoculars are almost always in two complete rows ; in four
or five specimens the outer row is short, composed of but two or
Smith: Genus Sceloporus 635
three scales. Usually there is not a great difference in the size of
the scales in the two rows. There is no irregular segmentation of
the scales as is the rule in cyanogenys and immucronatus. The first
canthal is forced above the canthal ridge by contact of the second
canthal and subnasal on both sides in six specimens. In two of these
it appears that the first canthal may be gone entirely (perhaps
fused) , but in the others it is obvious that it has been merely pushed
up to take a place in the series of small scales about the nostril.
The first canthal touches the series of scales above the supralabials
on both sides in five specimens, on one side in one. The preocular is
separated from these rows of scales on both sides in eight specimens,
on one side in ten. The two rows of scales between the subocular
and supralabials are reduced to one row at some point on both sides
in twenty-nine specimens, on one side in ten.
The medial row of the two rows of scales intercalated between the
postmentals and infralabials extends anteriorly to the anterior half
of the second labial in one count (two counts per specimen) ; to the
middle of the second in six; to the posterior half of the second in
ten; to the suture between the second and third in forty-eight; to
the anterior half of the third in thirty-four; to the posterior part
of the third in one; and to the suture between the third and fourth
in two.
Much variation occurs in the arrangement of the scales between
the frontonasals and rostral. There are never more than two pairs
of internasals, although one or both pairs of internasals may be
broken into two or more smaller scales, or fused together in part.
The rows of scales above the rostral and anterior to the first pair
of internasals is present in the entire series, with usually four in the
series between the middle of the nares (two in one specimen, three
in two, five in two and six in one).
The dorsal scale rows are distinctly converging in eleven speci-
mens; in the others they are either parallel or weakly converging
(weakly converging is used in description of cases in which a single
scale row drops out somewhere on the back, and the other rows show
some evidence of converging to accommodate this loss; in some a
row may be dropped out, and the scales of the adjacent rows en-
larged immediately posterior to this point, with the result that no
convergence is apparent).
The basal caudal scutes are from one and one third to two or
more times larger than the median dorsals.
The postanal scales are but slightly enlarged in some males; in
the young males they are not visibly enlarged. In two male speci-
636 The University Science Bulletin
mens the femoral pore series is double near the median end, increas-
ing the pore count from a maximum of 17 to a maximum of 23
(20-22 in one specimen, 21-23 in the other) .
The males vary somewhat in ventral coloration. The throat, an-
terior to the gular fold region, is usually uniform pale blue. The
sides of the belly are also pale blue in some specimens; in older
males the black neckband is continued very distinctly across the
gular fold region ; in the largest the black borders of the lateral blue
abdominal areas are expanded and meet on the median ventral line,
and some black spotting is present on the preanal region. The light
band between the upper margins of the ear are present in all adult
males, but is either absent or broken into spots in young males and
in females.
The dorsal coloration of the females is much like that of the
males. The light band across the neck between the upper margins
of the ear is either absent or indicated only by irregular light spots.
In most of them the back is uniform, as in the males, but in some
there are indistinct dark and light areas scattered over the back.
The black spots tend to form in rows, and down each side of the mid-
dorsal line, and the light spots border posteriorly each dark spot.
In most specimens in which these spots are visible, however, they
are quite indistinct and irregular in their disposition. The young
specimens are more frequently so spotted than the larger females.
The sides of the belly in the females are grayish ; the area anterior
to the gular fold region is also grayish, with irregular light spots, or
light lines converging toward the middle. The ventral surfaces of
the tail, limbs and the preanal region are cream to white.
The tail, which is broken in the type, is Vandyke or seal brown
toward the distal end, and has fairly distinct, narrow bands of
whitish, usually one scale wide. These bands do not encroach upon
the ventral surface of the tail, although the darker color may be
visible toward the tip.
Rernarks. S. j. minor occupies the northern central plateau region
of Mexico, and presumably intergrades with immucronatus to the
east. All specimens are quite uniform in character except few speci-
mens in a large series from Charcas, San Luis Potosi. In three speci-
mens the dorsal scales from occiput to base of tail are 35-35-34. This
represents an extreme, from one locality, much lower than that from
any other locality. Thirty-six scales are present in two specimens
from San Felipe, and all the remainder have no less than thirty-
eight. Another specimen from farther north than Charcas (El
Smith: Genus Sceloporus 637
Salado) has characters typical of minor, including scale counts, form
of supraoculars and general character of other head scales.
The characters which differentiate minor and cyanogenys may be
summarized. In coloration the differences are striking. The black
nuchal collar is much broader in minor, the light anterior borders
are rarely divided (more frequently in the Charcas and El Salado
specimens), and the general coloration more brownish and darker.
In scale characters, minor has a higher average number of scales
from occiput to base of tail ; the supraoculars are rarely not in two
complete rows in minor, and rarely in two rows in cyanogenys; the
preocular is usually not separated from the rows of scales above the
supralabials in minor, usually separated in cyanogenys; and scales
to head length (snout to occiput) fewer in cyanogenys.
Cyanogenys also reaches a much greater maximum snout-to-vent
length than minor (143 mm. in the former, 100.5 in the latter).
Range. Northern parts of the states of Queretaro and Guana-
juato, north through western Zacatecas, including most of San Luis
Potosi, to southern Coahuila. (See Fig. 17 for distributional map.)
Locality records. Guanajuato: (Cope, 1887) ; near San Felipe
(EHT & HMS 4198-4212, 4228, 4246, 4248-4262); three miles
northeast of Santa Rosa (EHT & H]\IS 3756-3759) ; near El Ter-
ero (EHT & HMS 4276-4278). Zacatecas: (Cope. 1885, 1900;
USNM 26166-7) (type locality); Valparaiso- Mts. (USNM 47868-
70, 47816-7, E. A. Goldman). San Luis Potosi: (USNM 4557,
Palmer); near San Luis Potosi (MCZ 4548 [2 spec], Dr. Edw.
Palmer) ; Morales (MCZ 20020-5, W. W. Brown) ; Alvarez (MCZ
20062, 28265-28325 [+45 duplicates], W. W. Brown; UMMZ
67692 [25 spec], W. W. Brown); Alvarez or Morales (MCZ 4
spec.) ; Concordia (MCZ 4569 [3 spec], Dr. Edw. Palmer) ; Charcas
(UMMZ 77275-77282 [71 spec], C. L. Lundell) ; Jesus Maria
(USNM 47297, 47299, E. W. Nelson); Villar (USNM 47300-1,
47161, E. W. Nelson) ; Ahualuko (USNM 47298, E. W. Nelson) ;
Santa Inez Mine, Charcas (USNM 17656-9, P. L. Jouy; Cope,
1900); thirty miles north of El Salado (EHT & HMS 4170).
Coahuila: Sierra Guadelupe (USNM 46700, 46702, 47493-5, Nel-
son & Goldman). Nuevo Leon: Miquihuana (USNM 46741-2,
Nelson & Goldman) .
638
The University Science Bulletin
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640 The University Science Bulletin
Sceloporus jarrovii immucronatus Smith
(Text Figs. 17 and 20 ; Plate Llll, Fig. 1)
Sceloporus torquatus poinsettii ? Boulenger, 1885, p. 220 (part); ? idem, 1897, p. 481
(part).
Sceloporus jarrovii ivimucronatus Smith, 1936a, pp. 223-227.
Type locality. Ten miles north of El Pinalito, Hidalgo, Mexico.
Holotype EHT & HMS 500; paratypes EHT & HMS 498-9, 501-4,
506-8, 510-1, 605-615; E. H. Taylor and H. M. Smith, collectors.
Diagnosis. A species of moderate size belonging to the torquatus
group; head shields smooth; laterals in diagonal rows, weakly
keeled, very weakly miicronate; the median laterals somewhat
larger than, to one and one half times as large as, median dorsals;
dorsals in parallel longitudinal rows, very weakly keeled, but
slightly mucronate, 37 to 46 from occiput to base of tail ; supraocu-
lars usually in two rows, the inner complete, the outer incomplete;
canthals regularly two; tibia as long as or slightly longer than
shielded part of head; fourth toe from base of fifth slightly longer
than distance from snout to posterior margin of ear; ratio of fourth
toe to snout-vent measurement, 23.8 to 28.6 (in adults) ; ratio of
hind leg to snout-vent measurement, 64.0 to 71.3; femoral pores,
12-19; ground color, cobalt blue; collar black, complete, narrow (2
or 3 scales wide), extending across gular fold region in males; region
anterior to gular fold and sides of belly cobalt blue ; ventral surfaces
of limbs and tail glaucous blue or pale blue ; median ventral surface
of belly in old males black, the color extending onto proximal por-
tion of ventral surfaces of limbs.
Description of type. Dorsal head shields smooth; enlarged supra-
oculars in two rows, the outer row formed of scales about one half
as large as those of inner row; supraoculars separated from super-
ciliaries by one complete and another incomplete row of scales (the
latter present on one side only), and bordered medially by a series
of small scales ; superciliaries six, the last overlapping the preceding
two (the fifth superciliary is entirely overlapped by the fourth) ; one
large, heavily keeled subocular, foUow^ed posteriorly by two small,
strongly keeled postoculars, these in turn followed by a series of
scales passing upward and contacting the parietal; the series of
scales between the supraoculars and frontals continued downward
between the former series of scales and the margin of the orbit,
terminating at about the middle of the posterior postocular; pre-
ocular scale strongly keeled, in contact with second canthal and with
the rows of scales above supralabials; first canthal in contact with
rows of scales above supralabials, separating the loreal from the pre-
Smith: Genus Sceloporus
641
ocular; about three fifths of the length of the second canthal forming
a part of the superciliary series ; rostral about three times as broad as
high ; two irregular rows of scales above supralabials and in contact
with subocular, preocular and subnasal, one row continuous around
snout above rostral; interparietal much larger than either parietal,
more triangular than pentagonal in shape; a pair of frontoparietals,
separated on the median line by the contact of frontal and inter-
FiG. 20. Head scales of Sceloporus jarrovii immucronatus Smith. EHT
& HMS 500, ElPinalito, Hidalgo; actual head length, snout to occiput,
16.5 mm.
parietal ; frontal transversely divided ; two prefrontals, in contact on
median line; three frontoparietals, the lateral in contact with both
canthals; three pairs of small internasals, the posterior largest, the
anterior smallest; a single small scale, not entering into series sur-
rounding nasal, separates the posterior internasal from the first can-
thai; five scales, including the subnasal, surround the nasal; four
supra- and five infralabials to a point below the middle of the eye.
Mental rather U-shaped, its labial border almost equal to that of
rostral ; mental followed by a series of five or six enlarged scales on
each side, the anterior in contact with its fellow on the median
ventral line; this series of scales separated from infralabials an-
teriorly by a single row of elongate scales (except anterior scale,
41—6037
642 The University Science Bulletin
which is in contact with the first infralabial), posteriorly by two
rows, the second row beginning below the junction of the second
and third infralabials and in contact with the series of postmental
scales; scales in temporal region keeled, mucronate in area near ear;
anterior margin of ear with five smooth, pointed scales, on one side
the upper two much larger than the others, on the other side all
fi.ve approximately equal in size, the upper two but little larger
than the others and smaller than the preceding scales (subequal on
one side) ; lateral cervical pouch present, a fold from about its
middle to lower margin of ear; this fold surmounted by scales more
strongly mucronate than those adjacent and somewhat smaller than
those in region near angle of jaw; dorsal scales in parallel or weakly
converging rows, very weakly keeled, weakly mucronate; lateral
scales more strongly keeled and mucronate, somewhat smaller than
dorsals and in diagonal rows; laterals with as many as seven
mucrones; dorsal scales a little more than half as large as largest
caudals; ventrals relatively large, about one half the size of dorsals;
breast scales no larger than those in median abdominal area; ventral
scales in anterior region, near chin, smaller than gular scales nearer
gular fold region; the latter scales as large as those in region near
angle of jaw; median gular scales about as large as abdominals;
preanals about as large as gulars; ventrals emarginate.
Scales on dorsal surface of upper foreleg somewhat smaller than
those on back, those on dorsal surface of lower foreleg considerably
smaller; all dorsal scales of arm, except hand, keeled and mucronate;
ventral scales of lower forearm somewhat smaller than dorsals of
the same member, those on ventral surfaces of upper forearm much
smaller; all ventral scales of arm, except hand, smooth; one dorsal,
two lateral, and one ventral row of scales around fingers and toes,
all continuous to tip except the two laterals, which terminate behind
the last scale of the other series; lamellar formula for fingers 9-13-
18-19-14; dorsal scales of hind limbs keeled, mucronate, less so on
foot; dorsals of tibia of about the same size as dorsals on body, those
on femur somewhat smaller; ventral scales of hind limb, except foot,
smooth; ventrals of tibia somewhat smaller than dorsal scales of
the same member; scales on anterior surface of femur smooth, be-
coming progressively smaller toward the femoral pore series; scales
immediately anterior to femoral pore series somewhat smaller than
preanals; scales on posterior surface of femur keeled, mucronate,
smallest near femoral pores, and becoming progressively larger
toward dorsal surface ; scales immediately posterior to femoral pore
series about one third to one half as large as those immediately
Smith: Genus Sceloporus 643
anterior to pore series; lamellar formula for toes 9-14-20-23-16; a
pair of distinctly enlarged postanals; caudals strongly keeled and
mucronate except those on ventral surfaces.
Color. General dorsal ground color cobalt blue of varying degrees
of brilliance; this color uniform, without markings, over dorsal sur-
faces of tail, limbs and back as far forward as neck. A narrow
black neckband is present, two or three scales wide, bordered be-
hind and in front by relatively broad pale-blue bands about two
scales wide; the black collar passes onto shoulder, widens there some-
what, then becomes narrow and is complete across the gular region.
The dorsal surface of the neck and head anterior to the nuchal collar
is darker blue (berlin or indigo blue), with irregular pale-blue spots.
A broad pale-blue band begins on the rostral and passes over the
supralabials, including the subocular, broadens in the temporal
region and passes through the ear, covering the entire opening. This
band unites with the light anterior border of the nuchal collar. The
area anterior to the gular fold is uniform ultramarine blue; a broad
central band down the belly is grayish, with very dark-blue edges,
which in turn border on lateral lighter blue areas of slightly darker
blue than the dorsal ground color. The ventral surfaces of the limbs
are light blue, a narrow edge of each scale usually white; the ventral
surface of the tail is pale blue, with a suffusion of white in some
areas.
Variation. The outer row of enlarged supraoculars (the smaller
of the two rows) is frequently fused in part with the inner row, but
never is there but a single row. The outer row is formed of scales
of varying sizes, sometimes almost as large as the scales of the inner
row, sometimes much smaller. In four specimens the frontal is in
contact with the interparietal ; in one the two frontoparietals are in
contact; in the remainder of the series a single, small, median scale
separates the frontal from the interparietal; of the specimens with
the latter arrangement, there are three in which the posterior section
of the frontal is divided transversely, and the posterior scale of these
two is divided longitudinally. In all paratypes the loreal is in con-
tact with the subnasal, the first canthal never (except in holotype)
contacting the series of small scales above the supraoculars ; the pre-
ocular is separated from these latter series of scales in seven speci-
mens; the first canthal is always distinct, never confined to the area
above the canthal ridge. The posterior pair of internasals is in con-
tact with the first canthal in three specimens; there are three pairs
of internasals, as in the type, in only three paratypes; in nine others
there are two distinct pairs, and in the remainder of the series the
644 The University Science Bulletin
scales are irregular. The median frontonasal contacts the frontal
in one specimen, and is separated from the frontal in another speci-
men by a small scale intercalated between the two prefrontals.
The two rows of scales between the subociilar and superior labials
are reduced to one at one point on both sides in nine specimens, on
one side in four. The medial of the two rows of scales intercalated
between the postmentals and inferior labials extends anteriorly to
the anterior half of the second infralabial in one count (two counts
per specimen) ; to the middle of the second in one ; to the posterior
half of the second in five ; to the suture between the second and third
in fourteen; to the anterior half of the third in nineteen; to the
middle of the third in one; to the suture between the third and
fourth in four; and to the anterior half of the fifth in one.
The dorsal scale rows are distinctly converging in seven speci-
mens. In the remainder they are either parallel or very slightly
converging (one row dropping out).
The dorsal coloration of the males is almost identical wdth that
of the holotype. The light borders of the nuchal collar are in some
specimens broken; the posterior border may be broken medially,
and the anterior may be broken into a series of large light spots.
The general bluish coloration of the throat is hardly developed in
small specimens, but distinct in those about 70 mm. from snout to
vent and larger. The blue of the sides of the belly is continuous
with the dorsal and lateral ground color. The neckband is complete
around the gular fold region in specimens 70 mm. from snout to vent
and larger.
The dorsal coloration of the females is essentially like that of the
males. The ground color, however, is grayish instead of blue, and
there are present in all but the largest specimens indistinct black
spots scattered over the dorsal and lateral surfaces of the body.
They are larger on either side of the middorsal line and tend to
form two longitudinal rows. The tail is indistinctly banded. The
ventral coloration is light, sometimes suffused with bluish in the
lateral abdominal regions.
The most characteristic feature of the coloration is the brilliant
blue ground color, which is varied only with black. The blue is less
distinct in females than in males, but even in these blue forms an
essential element of the coloration.
Remarks. S. j. immucronatus is most closely related to cyanogenys
and minor. No specimens have been examined which show inter-
gradation with either, however, and it is rather difficult to determine
Smith: Genus Sceloporus 645
from which it has been derived. I have concluded for the present
that immucronatus is derived from minor. Geographical association
points toward such a conclusion, as well as general form, maximum
size, character of the preocular and average scale count from occiput
to base of tail. It conforms more closely to cyanogenys in color,
character of the supraoculars and number of femoral pores.
Summarizing the points of distinction between minor and immu-
cronatus, the following may be mentioned: Ratio of hind leg to
snout-vent measurement greater in immoucronatus (64.0 to 71.3,
average 66.0 in the latter; in minor, 61.2 to 67.5, average 63.4) ;
supraoculars irregular in immucronatus (as in cyanogenys) ; femoral
pores usually more numerous (rarely more than 14 in minor, rarely
less in immucronatus) ; enlarged postanals present and well de-
veloped in males of immucronatus, variable in minor. In color
immucronatus differs by having a distinctly narrower black nuchal
collar, a general ground color of blue, and much black in the ventral
coloration of males.
From cyanogenys, immucronatus differs in possessing a larger
number of scales from occiput to base of tail (37 to 46 in immucro-
natus, 32 to 40 in cyanogenys) , preocular usually not divided (usu-
ally divided in cyanogenys) , ratio of hind leg to snout-vent measure-
ment greater (58.6 to 66.9, average 64.6), and lesser maximum size
(86.0 mm. snout to vent in immucronatus, 143 mm. in cyanogenys) .
In color, the males of immucronatus differ by having much black in
the ventral coloration; cyanogenys does not.
Habits and habitat. The specimens from Hidalgo were collected
on granite rocks on the crest of the plateau edge. The males, with
their brilliant blue coloration, were conspicuous from a considerable
distance.
A number of very young specimens were collected with the adults,
and it is assumed from this that immucronatus is ovoviviparous, as
are other members of the torquatus group.
Range. Western Queretaro through Hidalgo to southern central
Vera Cruz. (See fig. 17 for distributional map.)
Locality records. El Pinalito, Hidalgo (EHT & HMS 498-504,
506-508, 510-511) ; twenty-five miles south of Jacala, Hidalgo
(EHT & HMS 605-615) ; Presidio, north of Montzorongo, Vera Cruz
(MCZ 21093, W. W. Brown) ; Pinal de Amoles, Queretaro (USNM
47776, Nelson & Goldman).
646
The University Science Bulletin
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Smith: Genus Sceloporus 647
Sceloporus ornatus ornatus Baird
(Text Figs. 21 and 22; Plate LIV, Fig. 1)
Sceloporus ornatus Baird, 1859, p. 254; idem, 1859a, pp. 5-6; Muller, 1865, p. 602; Cope,
1875, p. 48; Yarrow, 1883, p. 57; Carman, 1884, p. 17; Boulenger, 1885, p. 224; Cope, 1885,
p. 403; idem, 1887, p. 38; Boulenger, 1890, p. 78; Giinther, 1890, pp. xii, 72; Boulenger,
1897, pp. 485-486; Cope, 1900, pp. 335, 340, 344-345, Fig. 48; Strecker, 1915, pp. 19-20;
Stejneger, 1916, pp. 227, 228.
Type locality. Patos, Coahuila. Holotype USNM 2845, Lt. B.
Couch, collector.
Diagnosis. A member of the torquatus group; dorsal scales 55 to
63 from occiput to base of tail; supraoculars quite irregular or in
two rows; posterior section of frontal reduced in size, broken in
small scales or absent; lorilabials usually reduced to one row below
subocular; two canthals, normal in position; inner row of labiomen-
tals terminating below third infralabial; femoral pores 16 to 20
(rarely 14 or 15) ; ratio of hind leg to snout-vent measurement, 61.4
to 65.5; ratio of fourth toe to snout-vent measurement, 20.2 to 24.4;
tibia longer than snout-occiput measurement; length of fourth toe
usually somewhat less than snout-ear measurement. Neckband
narrow, four to seven scales wide, with narrow light borders two
scales wide; back dimly banded; throat uniform white in females,
pale blue in males, without darker markings; males with sides of
belly, from axilla to groin, caerulean blue, bordered medially by a
hyacinth blue line ; the blue area on the sides of the belly are sepa-
rated by eight to twelve scale rows.
Description (from EHT & HMS 4420, male). Head and body
strongly depressed; scales of head pitted, the scales in prefrontal
and internasal region more strongly; cephalic scales smooth; inter-
parietal pentagonal, four times as large as either parietal; parietals
single on either side, subtriangular; frontoparietals single on either
side, rectangular, about two fifths the size of parietal; frontopa-
rietals separated medially by an azygous scale; posterior section of
frontal split irregularly into three scales; prefrontals as large as
lateral frontonasals, in contact medially ; frontonasals approximately
equal in size; internasals irregular, separated from rostral by a
single row of small scales; nasal small, subcircular, separated from
rostral; supraoculars irregularly divided, small; a row of scales
separating supraoculars from median head scales, and one complete
and another incomplete row between supraoculars and supercili-
aries; two canthals, the first smaller than the second, second can-
thai not in contact with subnasal; first canthal not touching lori-
labials ; loreal rectangular, somewhat larger than subnasal, preocular
completely divided on one side, partially on the other; subocular
648
The University Science Bulletin
followed by three small, well-differentiated, keeled postoeulars;
lorilabials in two complete rows below siibocular, reduced to one at
a narrow point on one side; three and one half siipralabials and four
and one half infralabials to a point below middle of eye.
Mental with a labial border about two thirds that of rostral;
mental pentagonal, followed by several pairs of postmentals, the
anterior two pairs well differentiated, and the scales of the anterior
pair in contact; outer row of labiomentals separated from mental by
partial contact of first postmental and first inf ralabial ; gular scales
Fig. 21. Head scales of Sceloporus ornatus ornatus Baird. EHT &
HMS 4422, near Saltillo, Coahuila; actual head length, snout to occiput,
12 mm.
smooth, rounded except in extreme posterior part of the gular region,
where they are weakly notched; gular scales smallest in middle of
throat, largest toward angle of jaws.
Three or four auricular lobules, smooth, the lower one or two
pointed, the upper two rounded and nearly twice as large as preced-
ing scales; temporal scales very weakly keeled and very weakly
mucronate, becoming more strongly keeled and mucronate toward
ear; temporal scales subequal in size to scales between ear and lat-
eral nuchal fold ; a fold of skin between upper edge of lateral nuchal
fold and lower edge of ear, surmounted by strongly keeled, very
strongly mucronate scales; nuchal fold or pouch moderately deep;
dorsal scales not or but very weakly keeled, not or but very weakly
mucronate, not denticulate; lateral scales keeled, mucronate, weakly
Smith: Genus Sceloporus 649
denticulate, the largest slightly larger than median dorsal scales;
ventral scales smooth, rounded, about two thirds the size of median
dorsal scales; ventral scales on chest slightly larger than median
ventral abdominals; scales in preanal and interfemoral regions
slightly smaller than preceding scales; postanal scales enlarged,
separated by two small scales; subcaudals smooth except toward tip
of tail; dorsal caudals near base of tail about twice as large as
median dorsals on body.
Dorsal scales of foreleg as large as or larger than median dorsals
on body, those on upper foreleg somewhat larger than those on lower
foreleg, all keeled and mucronate ; scales on anteroventral surface of
lower foreleg smooth, rounded or weakly mucronate, somewhat
smaller than dorsal scales of same member; scales on posteroventral
surface of lower foreleg somewhat larger, keeled, mucronate; ventral
scales of upper foreleg smooth, rounded, about one third size of
dorsals of same member; scales in axilla not granular; lamellar
formula for fingers, 8-12-16-16-12 (7-12-16-16-13).
Dorsal scales of thigh about as large as median dorsals of body,
keeled, mucronate; dorsal scales of shank nearly twice as large as
median dorsals on body, keeled, strongly mucronate; ventral scales
of shank about two thirds size of dorsal scales of same member,
smooth, rounded; scales on ventral surface of femur near series of
femoral pores subequal in size to preanal scales, smooth, rounded or
weakly notched; scales on posterior surface of femur somewhat
larger than preanal scales, keeled, mucronate, abruptly decreasing in
size toward series of femoral pores; no postfemoral dermal pocket;
lamellar formula for toes, 8-12-19-20-15 (8-12-18-20-15).
Color. The dorsal coloration is practically identical in all. A
very distinct black band crosses the neck and passes onto the
shoulder. It is four to seven scales long on the median dorsal line,
and is bordered on both sides by a broad light line, covering about
two scale rows, of a whitish or iridescent pale-blue color, sometimes
with a tinge of orange laterally. Both light lines may be complete
or either one or both broken on the median line. The area between
the neck band and frontal region, as well as the temporal region, is
variously marked; in some females no markings are present there,
while in males this area is spotted with whitish or iridescent pale
blue, sometimes forming an indistinct light band across the neck. .
The back is sepia in females, more or less black in males. About
seven indistinct, broken, narrow light bands traverse the back, dis-
appearing on the sides, which are lighter in color than the median
dorsal area. The sides are usually pale blue, suffused to a varying
650
Ti-iE University Science Bulletin
degree with orange, but in some females are brownish, somewhat
lighter than the back. In some males the lighter color of the sides
encroaches on the median darker area, resulting in a pattern of a
series of paired dark spots down the middle of the back. The tail
is dimly barred with usually rather narrow, light bands alternating
with much broader dark bands.
In males, the throat and gular regions are pale blue; the sides of
the belly, from axilla to groin, are caerulean blue, bordered medially
Fig. 22, Distribution of Sceloporus omatus ornatus Baird, S. o. caeruleus Smith,
S. dugesli dugesii Bocourt and S. d. intermedius (Duges).
by a hyacinth blue line; the blue areas on the sides of the belly are
separated by eight to twelve scale rows.
Variation. Only eight specimens have been available for a study
of variation. Parietal one fourth to one fifth size of interparietal;
frontoparietal divided on both sides in one, on one side in two; fron-
tal never touches interparietal; frontoparietals contact medially in
two; an azygous scale separates frontoparietals in six; frontal vari-
ously divided, normally divided in two specimens; supraoculars
divided in all; prefrontals contact in all; canthals 2-2 in all, normal
in relationship to other scales; preocular divided on both sides in
two, on one side in one; scales below subocular not reduced to one
row at a point below subocular on one side in one specimen; outer
labiomental row separated from mental in all; inner row of labio-
mentals terminating at a point even with the anterior half of third
Smith: Genus Sceloporus
651
infralabial (middle of third infralabial on one side of one and on
both sides of another).
Auricular lobules three to five, usually the latter number, the
upper three lobules largest; femoral pores 16 to 20 (14-15 in one
specimen) ; dorsals from occiput to base of tail 55 to 63.
Remarks. The species is apparently confined to semiarid locali-
ties. The specimens collected near Saltillo were found by Edward
H. Taylor near the crest of a high hill at the north edge of the city.
Range. Known only from southeastern Coahuila.
Locality records. Coahuila: Patos (USNM 2845; Lt. Couch);
near Saltillo (EHT & HMS 4300, 4418-4422, 4598) ; four to five
miles north of Gomez Farias (EHT & HMS 4301) ; Jaral (FMNH
1548, Heller & Barber) . Published records are available from Red-
mond's Pass, Texas (Cope, 1900) ; Nuevo Leon (Boulenger, 1890,
1897; Giinther, 1890); Duvall Co., Texas (Boulenger, 1897).
Stejneger (1916) states that the records from the latter two locali-
ties are based upon Sceloporus disparilis. Cope's specimen cannot
now be found. Garman (1884) gives the locality "Sonora" — un-
doubtedly incorrect.
Measurements and Scale Counts of Sceloporus omatus ornatus Baird
Museum
EHT
&
HMS
4422
EHT
&
HMS
4598
EHT
&
HMS
4420
EHT
&
HMS
4421
EHT
&
HMS
4418
USNM
Number
2845
Snout to vent
63.5
67.0
82-1-
12.0
16.0
43.0
13.0
15.0
7.6
18-19
16-17
55
65
57
9.5
64.1
22.3
9
72.5
109.0
13.2
18.0
47.5
16.2
17.0
8.7
19-19
18-19
63
72
57
13.0
65.5
23.4
73.5
74.0
82.5
Tail
Snout to occiput
12.0
16.0
?
?
15.2
7.2
20-19
16-16
56
60
54
10.5
13.5
17.5
18.0
9.2
19-18
20-19
55
74
54
11.0
24.4
13.0
17.0
43.0
14.0
15.0
8.2
19-19
17-18
58
70
57
10.5
61.4
20.2
9
15 2
Snout to ear ....
19 4
Hind leg
51 4
Tibia
16 5
Fourth toe
17.6
Fifth toe
9 6
Lamellae, fourth toe
20-20
Femoral pores
15-17
Dorsals
60
Ventrals
76
Scales around body
61
Scales to head length
11 0
Ratio, hind leg to snout-vent
62 4
Ratio, fourth toe to snout- vent
Sex
23.9
9
21.3
cf
652 The University Science Bulletin
Sceloporus ornatus caeruleus Smith
(Text Figs. 22 and 23 ; Plate LIV, Fig. 2)
Sceloporus ornatus caeruleus Smith, 1936a, pp. 227-230.
Type locality. Five miles south of San Pedro, Coahuihi, Mexico.
Holotype DHD & HMS 350; paratypes DHD & HMS 348-9, 351-
361; David H. Diinkle and H. M. Smith, collectors.
Diagnosis. A member of the torquatus group of Sceloporus; dor-
sal scales, 47 to 53 from occiput to base of tail; posterior section of
frontal reduced or variously divided; supraoculars in two rows, the
scales of the outer row somewhat smaller than those of the inner
row; lorilabials reduced to one row at a point below subocular;
two canthals, normal in position; inner row of labiomentals termi-
nating below third infralabial; femoral pores 12 to 17 (average
13.7); ratio of hind leg to snout-vent measurement, 66.6 to 74.5;
ratio of fourth toe to snout-vent mesaurement 22.7 to 27.4; length
of tibia greater than snout-occiput measurement; length of fourth
toe usually slightly greater than snout-ear measurement, sometimes
equal or slightly less. General ground color bluish; nuchal collar
four to six scales wide, light bordered; ventral surfaces of males en-
tirely blue, without darker markings; a series of about seven dark
blotches more or less evident on middle of back.
Description of holotype. Head and Body somewhat compressed;
scales of head not rugose or keeled ; scales in prefrontal, frontonasal
and internasal regions rather strongly pitted; a few pits on other
head scales; parietals single on either side, subtriangular, about one
third size of interparietal; interparietal pentagonal, the posterior
edge rounded and confluent with posterior edges of parietals; fron-
toparietals small, square, separated medially by a small azygous
scale; another small scale between frontal and right frontoparietal;
posterior section of frontal slightly less than one third size of an-
terior section; prefrontals in contact medially, but slightly smaller
than anterior section of frontal ; lateral frontonasals subequal in size
to prefrontals; median frontonasal somewhat larger; scales in inter-
nasal region irregular; nasal small, the anterior border about nostril
somewhat wider than posterior border; nasal separated from ros-
tral; supraoculars in two rows, the scales of the outer row about half
as large as scales of inner row; one complete and another incomplete
row of scales between supraoculars and superciliaries; one row of
rather broad scales between supraoculars and median head scales;
six superciliaries, the fifth entirely concealed by the fourth, which
is abnormally divided into two; two canthals, normal in position, the
first somewhat smaller than the second; subnasal smaller than loreal,
Smith: Genus Sceloporus
653
which is very long and narrow; preociilar not divided; siibociilar fol-
lowed by two small, well-differentiated, strongly keeled postociilars;
lorilabials reduced to one row by two scales below subociilar.
Mental subtriangular (tending toward pentagonal), with a lateral
border slightly more than one half that of rostral ; mental followed
by about four pairs of postmentals; scales of first pair of postmen-
tals in contact medially; outer row of labiomentals separated from
mental by partial contact of first postmental and first infralabial;
inner row of labiomentals terminating at a point even with anterior
half of third infralabial ; median gular scales about one third smaller
than lateral gular scales; scales in anterior part of gular area with
a terminal pit, those in extreme posterior part of gular area notched.
Fig. 23. Head scales of Sceloporus omatus caeruleus Smith. DHD & HMS
355, five miles south of San Pedro, Coahuila; actual head length, snout to
occiput, 11.5 mm.
Auricular lobules three or four, distorted on one side; on the
other side the median two are much larger than the outer two,
rounded, not keeled, subequal in size to scales in temporal region;
scales between ear and lateral nuchal fold one third or one fourth
the size of scales in temporal region; latter scales keeled, denticu-
late, mucronate; a weak skin fold between upper edge of lateral
nuchal pocket and lower edge of ear, surmounted by small, very
strongly keeled and mucronate scales; median dorsal scales smooth,
sometimes rounded, usually with one or two terminal pits or den-
ticulations, rarely with a terminal mucrone; lateral scales somewhat
smaller than median dorsal scales, keeled, mucronate, denticulate,
with terminal pits; median abdominal scales about one third to one
half the size of median dorsal scales, somewhat larger than preanal
scales ; scales on chest slightly larger than those in middle of belly ;
scales on abdomen and on chest smooth, rounded; enlarged postanals
654 The University Science Bulletin
present, separated by two scales; subcaudals smooth except toward
extreme tip of tail; dorsal caudal scales near base of tail slightly
larger than median dorsals on body.
Dorsal scales of foreleg somewhat smaller than dorsal scales on
body, keeled, mucronate, with terminal denticulations and pits;
scales on lower foreleg somewhat smaller than those on upper;
ventral scales of foreleg smooth, notched or with terminal pits, those
on lower foreleg somewhat smaller than dorsal scales of same mem-
ber; ventral scales on upper foreleg about half as large as ventral
scales of lower foreleg; lamellar formula for fingers, 10-14-17-18-13
(9-14-18-18-13).
Dorsal scales of hind leg keeled, mucronate, denticulate and with
terminal pits; dorsal scales of tibia somewhat larger than dorsals on
body, those on femur slightly smaller; ventral scales of hind leg
smooth, rounded or notched; ventral scales of shank slightly smaller
than dorsal scales of the same member; ventral scales in front of
femoral pore series subequal in size to preanal scales, gradually in-
creasing in size toward anterior face of femur; scales on posterior
surface of femur strongly keeled, strongly mucronate, slightly larger
than preanal scales, abruptly decreasing in size immediately pos-
terior to femoral pore series; no postfemoral dermal pocket; lamellar
formula for toes 7-13-17-20-16 (7-13-18-20-16).
Color. General dorsal color pale blue, with a greenish suffusion;
a series of large black blotches, about six in number, of indefinite
outline, on middle of back, separated from each other by one row
of light scales; nuchal collar four scales wide, with light anterior
and posterior borders about two scales wide; posterior light border
extending onto proximal end of forearm ; anterior border terminating
on sides of neck slightly above and posterior to lateral nuchal fold;
a light bar across neck between ear and foreleg, terminating on sides
of neck about at a level with upper edge of ear; an indistinct, broken
light line across neck between upper edges of ear; a few light-colored
scales in temporal region; limbs dimly banded.
Black nuchal collar continuous about neck; a suffusion of black
in groin, extending somewhat on ventral surface between groins;
entire ventral surfaces of other parts, including head, limbs and tail,
bluish; the sides of the belly, lower surfaces of hind limbs, median
gular area and basal region of tail grey smalt blue; lower surfaces of
forelegs and distal part of tail suffused with pale blue; median
anterior abdominal and anterior gular regions suffused with irides-
cent apple green; ventral surfaces of hands and feet white.
Smith: Genus Sceloporus 655
Variation. The dorsal coloration of all paratypes is much as in
the holotype. In females the general ground color is less brilliant.
The ventral coloration of the males is much as in the holotype. In
younger males the ventral surface of the tail, hind limbs and an
indistinct, narrow median area in middle of abdomen are whitish.
Females are whitish below, without dark markings, sometimes with
a general light suffusion of pale blue.
The parietals are from one third to one sixth the size of the
interparietal; the frontoparietals are divided on one side in one, on
both sides in another (in the latter specimen, the posterior part of
each divided frontonasal is fused with the interparietal) ; the frontal
touches the interparietal in two; the frontoparietals contact medially
in one; an azygous scale lies between the frontoparietals in nine;
the posterior section of the frontal is divided into two scales in five
specimens ; in one the posterior section is absent ; in seven the frontal
is noiTQally divided, the posterior part always considerably smaller
than the anterior; the supraoculars are always in two rows, the
scales of the outer row smaller than those of the inner; the row of
scales separating the supraoculars from the median scales is always
composed -of broad, not elongate, scales; all supraoculars separated
from median head scales; prefrontals in contact in all; invariably
two canthals, normal in position and relationships with other scales;
preocular divided in eight; lorilabials reduced to one row at a point
below subocular in all; outer row of labiomental terminating below
anterior half of third infralabial in all but two; in one of these it
terminates below the posterior half of the third, in the other it
terminates below the anterior half of the fourth.
Auricular lobules three to five, usually with two much larger than
the others; dorsal scales 47 to 53, average 49.7; femoral pores 12 to
17, average 13.7. Other characters of scalation approximately as in
holotype. Variations in proportions and certain scale characters are
given in the accompanying table.
Habits. Specimens of this subspecies were found on large rocks
on the north exposure of a range of low, arid hills a short distance
north of the road from Torreon to Saltillo, about five miles south
and slightly east of San Pedro. Peculiarly enough, the species was
not found on a somewhat higher range of hills a short distance south
of the road; it is possible that it was not sufficiently warm when Mr.
Dunkle and I collected there. It was about 11:00 a. m. when the
lizards were encountered in relative abundance. They were ex-
tremely wary, and could be collected successfully only by sighting
656
The University Science Bulletin
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Smith: Genus Sceloporus 657
them at a considerable distance and then approaching as cautiously
as possible. The brilliant ventral coloration of the males was very
conspicuous as they protruded their heads and breasts above the
edges of rocks.
Range. Known only from the type locality. (See fig. 22 for dis-
tributional map.)
Sceloporus dugesii dugesii Bocourt
(Text Figs. 22 and 24; Plate LV, Fig. 1)
Sceloporus dugesii Bocourt, 1874, pp. 188-190, pi. 18, figs. 7, 7a, 7b; Garman, 1884, p.
18; Boulenger, 1885, pp. 224-225; Gunther, 1890, p. 71.
Sceloporus pleurolepis Gunther, 1890, pp. xii, 74-75, pi. 32, fig. B.
Sceloporus yarrovii Gunther, 1890, pp. xii, 69 (part); Boulenger, 1897, pp. 483-385 (part).
Type locality. Colima.
Diagnosis. A member of the torquatus group of the genus
Sceloporus; dorsal scales, 41 to 50 from occiput to base of tail;
lateral scales considerably larger than median dorsal scales, in
oblique rows, and each with an apical mucrone arising within the
free margin of the scale; ratio of hind leg to snout- vent measure-
ment, 52.5 to 61.7; ratio of fourth toe to snout- vent measurement,
17.4 to 24.1; fourth toe shorter than distance from snout to posterior
border of ear; femoral pores, 9 to 13; head scales rugose (micro-
scopically); posterior section of frontal reduced; supraoculars in
two irregular rows; frontoparietals rarely in contact medially; black
nuchal collar narrow, three or four scales wide, with a light posterior
border not over a scale wide, passing onto shoulder; anterior light
border of collar absent or indistinct; a dim, dark line from eye
through upper part of ear to collar, bordered above and below by
a narrower light line; back uniform gray or with indistinct darker
spots; throat not conspicuously barred. Maximum snout-to-vent
measurement, 87.5 mm.
Description (from EHT & HMS No. 2759, male). Head and
body flattened; head scales rugose and pitted; a single rounded
parietal on each side ; interparietal pentagonal, about three and one
half times as large as either parietal; a single frontoparietal on
each side, rectangular, about two thirds as large as either parietal;
frontal touching interparietal, separating frontoparietals medially;
posterior section of frontal about two fifths the size of anterior sec-
tion; prefrontals slightly more than half the size of anterior section
of frontal, separated medially by narrow contact of frontal and
median frontonasal; latter separated on each side from lateral
frontonasals by a small, elongate scale; a pair of rather large scales
preceding median frontonasal, in turn preceded by a pair of large
42—6037
658
The University Science Bulletin
internasals separated from rostral by a pair of small, narrow scales;
supraoculars six or seven, divided, separated from median head
scales by a row of small scales, from superciliaries by one complete
and another incomplete row of scales; nasal small, separated from
rostral, the portion anterior to naris strongly tapered but much
broader than part posterior to naris; subnasal present, moderate in
size; a single loreal on each side, about equal in size to subnasal;
preocular large, not divided ; two canthals, the first not forced above
Fig. 24. Head scales of Sceloporus dugesii dugesii Bocourt. EHT & HMS
2761, near Magdalena, Jalisco; actual head length, snout to occiput, 15.7
mm.
canthal ridge nor contacting lorilabials; subocular normal, followed
posteriorly by two well-differentiated, strongly keeled postoculars;
rows of lorilabials reduced to one at a point below subocular; about
four infra- and supralabials to a point below middle of eye.
Mental pentagonal, with a labial border about two thirds that of
rostral; about three pairs of well-differentiated postmentals, the
scales of the anterior pair in contact medially; outer row of labio-
mental scales narrowly separated from mental by partial contact of
first infralabial and first postmental ; inner row of labiomental scales
terminating anteriorly below the posterior part of second infralabial;
gular scales smooth, rounded except in extreme posterior part of
gular region, where they have a single, small, apical notch.
Smith: Genus Sceloporus 659
Three or four auricular lobules, the upper ones smooth and
rounded, the lower ones acuminate, all smaller than preceding scales;
temporal scales keeled, not mucronate; scales between ear and lat-
eral nuchal fold somewhat larger than in temporal region, keeled
and strongly mucronate; lateral nuchal fold strongly oblique; dorsal
scales weakly keeled, not mucronate ; lateral scales in oblique rows,
nearly twice as large as median dorsals, keeled, rather strongly
mucronate, with the apical mucrone of each arising within the free
edge of the scale; ventral scales rounded or very weakly notched,
about one third the size of median dorsals; preanal scales as large
as or slightly larger than median ventral abdominals; scales between
hind legs about half the size of preanals; scales on anterior part of
chest slightly larger than the scales posterior; subcaudals smooth
except toward tip of tail; postanals enlarged, separated by a pair
of small scales ; dorsal caudals near base of tail about twice as large
as median dorsals on body.
Dorsal scales of foreleg subequal in size, about equal to dorsals
on body, keeled, weakly mucronate; ventral scales of foreleg very
weakly keeled or smooth, weakly mucronate, those on lower foreleg
about one third larger than those on humerus and somewhat smaller
than dorsals of foreleg; lamellar formula for toes, 8-12-16-17-12
(9-13-17-16-11).
Dorsal scales of hind limb keeled, strongly mucronate, those on
femur subequal in size to dorsals on body and somewhat smaller
than those on tibia; ventral scales of hind leg smooth, rounded,
those on tibia about one half size of dorsals of same member; scales
preceding femoral pore series smaller than preanals; scales on pos-
terior surface of femur about three times as large as preanals,
strongly keeled, strongly mucronate, abruptly decreasing in size im-
mediately behind femoral pore series; scales on posterior surface of
femur near insertion surrounded and separated slightly from each
other by granules; no postfcmoral dermal pocket; lamellar formula
for toes, 8-12-16-21-13 (9-11-16-19-14).
Color. Dorsal ground color brown-gray ; black nuchal collar nar-
row, about three scales wide, darker posteriorly and merging with
gi'ound color anteriorly; collar with a light, narrow, posterior border
about one scale wide, passing onto humeinis; a few irregular dark
and light markings on neck ; a dark line about one scale wide from
posterior edge of orbit through upper part of ear to nuchal collar;
this line bordered above and below by a light line about one scale
wide; back uniform, without markings; a faint dark line from a
point above axilla to groin; below this line, sides of body darker;
660 The University Science Bulletin
limbs uniform or very dimly banded; tail with narrow, very dim
bands toward tip; throat cobalt blue, lighter anteriorly; chest
cream, blue encroaching a short distance on the sides anteriorly;
sides of belly azure blue laterally, cobalt blue medially, bordered
internally by a black band three or four scales broad, terminating
anteriorly a short distance from axilla, and posteriorly involving
the region of the groin and encroaching a short distance upon an-
terior proximal surface of femur; ventral surfaces of limbs, tail and
middle of belly cream, immaculate.
Variation. Variation in essential scale characters of the body
and in measurements is given in the accompanying table. The
variation of the head scales is as follows: Enlarged supraoculars
always divided, the scales of the inner row larger than those of the
outer row, which is usually incomplete; supraoculars always sepa-
rated from the median scutes by at least a single series of small
scales, frequently by another incomplete series also; one fronto-
parietal divided in one specimen, both divided in two; posterior
frontal contacting interparietal in thirteen specimens, separated by
an azygous scale in fourteen, and separated by contact of fronto-
parietals in four; posterior frontal transversely divided in four
specimens, broken into small scales (three or four) in four; anterior
frontal one and one half to three and one half times as large as
posterior frontal; median frontonasal contacting frontal in ten speci-
mens, separated by an azygous scale in thirteen, and by contact of
the two prefrontals in twelve; second canthal contacts subnasal,
forcing first canthal above the canthal ridge, on both sides in three
specimens, and on one side in three; two complete rows of scales
separating subocular from superior labials on both sides in seven
specimens, and on one side in two; lateral row of the two rows of
scales intercalated between infralabials and postmentals separated
from the mental by a partial contact of first postmental and first
infralabial in all but one specimen ; inner of these two rows termi-
nated under second sublabial on both sides in ten specimens, on one
side in three, under the third labial on both sides in five, on one side
in three, under the suture between the second and third sublabial on
both sides in fifteen, on one side in four.
The dorsal head scales are usually distinctly although not strongly
rugose, both in young and in old specimens. Occasional specimens
have tliis rugosity confined to the internasal region and the scales
about the supraorbitals.
Smith: Genus Sceloporus 661
The coloration is more or less as given by Bocourt (1874). The
ner-kband is narrow, three or four scales wide at the widest point.
The two rows of spots down the back, one on each side of the mid-
dorsal line, are not regularly present, nor are the spots on the sides
usually present. Usually no spotting whatsoever is visible on the
back. In one specimen the spots on the sides are arranged in longi-
tudinal series, not in oblique series diverging posteriorly. Some
specimens have very light-bro"^Ti spots scattered over the dorsum.
The lateral ventral abdominal areas of males are azure to cobalt
blue, bordered internally from opposite the axilla to the groin by a
black band some four scales wade. A median whitish abdominal
area is six scales wide at the narrowest point; the throat is largely
cobalt blue, with some lighter areas sometimes visible as oblique
lines. The females are whitish beneath, with irregular blue mark-
ings on the throat; sometimes these markings are arranged in ob-
lique lines converging posteriorly.
Habits and habitat. Specimens of this species were collected in
Nayarit and Jalisco on rock fences. Some were collected as early
as June 10, and at this time the newly born young were running
about the fences. It is assumed that the form is ovoviviparous.
Remarks. Boulenger's specimens of Sceloporus dugesii described
in his monograph of 1897 are not dugesii dugesii, but dugesii inter-
medius. However, it appears very probable that the specimens he
included with jarrovii from north of Rio de Santiago, Jalisco (ex-
cept one) ; La Cumbre de los Arrastrados, Jalisco; Ixtlan, Jalisco;
and from La Venta, Guadalajara, are all dugesii dugesii. Gunther's
type of pleurolcpis is in this series. The description of jarrovii given
by Boulenger, thus considered, is a composite and not descriptive
of either species alone. In the table, however, are given a number
of rather illuminating data. The femoral pores are fewer in the
southern specimens, the fourth toe shorter, the tibia shorter and the
snout-ear measurement shorter — all points of difference between
jarrovii and dugesii dugesii. Gunther's description of pleurolepis,
moreover, is entirely applicable to dugesii dugesii — much less to
jarrovii. The very large lateral scales, which he stresses, are much
larger in dugesii dugesii than in jarrovii. Aside from the shape of
the frontal, the depiction of which may not necessarily be accurate,
the figures accompanying the description of pleurolepis illustrates
well enough the form dugesii dugesii. The femoral pores are 12 in
pleurolepis, never less than 13 in jarrovii, but 9 to 13 in dugesii
662
The University Science Bulletin
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Smith: Genus Sceloporus 663
dugesii; and the supraoculars are not divided, or are not so nearly
equally divided in jarrovii as in dugesii.
The single specimen of Boulenger's jarrovii from Rio Santiago
which does not form the type of pleurolepis appears actually to be
jarrovii, judging from the data given in his table. If the locality
is correct, then jarrovii and dugesii overlap each other in their dis-
tribution.
In the agglomeration of Boulenger's table for jarrovii are also two
specimens from San Luis Potosi which have the characters of minor,
and another from Duvall county, Texas, which has the characters of
'poinsettii.
Range. Along the coastal ranges of western Mexico from south-
ern Nayarit to Colima. (See fig. 22 for distributional map.)
Locality records. Jalisco: near Magdalena (EHT & HMS 2758-
2772) ; near La Quemada (EHT & HIMS 2806-2824) ; near Chapala
(EHT & HMS 2833); north of Rio Santiago (type locality of
pleurolepis) (Giinther, 1890; Boulenger, 1897); La Venta, Gua-
dalajara (Boulenger, 1897) ; Itzatlan (USNM 47887-90, Nelson and
Goldman) ; Jacala (USNM 64658-60, Nelson and Goldman) ; west
of Orendain (AMNH 18454, Paul D. R. Riithling) ; El Aguilar
Mine, Hostotipaquillo (AMNH 15493, Paul D. R. Riithling).
Colima: (Bocourt, 1874; Garman, 1884) (type locality of dugesii).
Nayarit: Ixtlan (Boulenger, 1897).
Sceloporus dugesii intermedins (Duges)
(Text Figs. 22 and 25; Plate LV, Fig. 2)
Tropi dole-pis intermedins Dugos, 1870, p. 243 (nomen nudum).
Sceloporus intermedius Duges, 1877, pp. 29-34, pi. 1, figs. 21-32; Giinther, 1890, p. 71.
Sceloporus dugesii (non Bocourt) Cope, 1879, p. 265; idem, 1885, p. 403; idem, 1887,
p. 38; Duges, 1896, p. 479; Boulenger, 1897, p. 486; idem, 1898, p. 915; Cope, 1900, pp.
336, 341-343; ? Werner, 1903, p. 344.
Type locality. Noria, near Zamora, hacienda of D. Epifanio
Jimenez (Michoacan) .
Diagnosis. A member of the torquatus group of the genus (Sce-
loporus; dorsal scales, 47 to 54 from occiput to base of tail; femoral
pores, 10 to 15; fourth toe shorter than (rarely about equal to) dis-
tance from snout to posterior border of ear; ratio of hind leg to
snout-vent measurement, 54.7 to 61.1; ratio of fourth toe to snout-
vent measurement, 19.5 to 22.1; lateral scales in oblique rows, the
median laterals distinctly larger than median dorsals, and with
terminal mucrones arising within free edge of scales; head scales
not rugose; frontoparietals usually in contact; posterior section of
frontal reduced; supraoculars in two rows; a series of dark spots
664
The University Science Bulletin
down back on either side of middorsal line, with a dark line extend-
ing obliquely posteriorly from each; black nuchal collar narrow,
about three scales wide, with a narrow, light, posterior border;
anterior light border indistinct; throat with distinct, oblique, blue
bars; sides of abdomen blue, bordered medially by a broad band of
black; middle of belly and chest cream; black nuchal collar not
confluent on ventral surface; ventral surfaces of limbs and tail
cream. Maximum snout-vent measurement, 79 mm.
Description (from EHT & HMS No. 3690, male). Head some-
what depressed; dorsal and lateral head scales with numerous pits,
Fig. 25. Head scales of Sceloporus dugesii intermedins (Duges). EHT &
HMS 3690, near Acambaro, Guanajuato; actual head length, snout to occiput,
11.5 mm.
otherwise smooth; a small, rounded parietal on each side; inter-
parietal about four times as large as either parietal ; frontoparietals
single on each side, rectangular, about two thirds size of parietal,
broadly in contact medially; frontal nomially divided, the posterior
section slightly less than half the area of the anterior section; pre-
frontals in contact medially; median frontonasal as large as an-
terior section of frontal, wedged between the two prefrontals; lat-
eral frontonasals about one third the size of median frontonasal, in
contact with the latter and both canthals ; four intemasals, the pos-
terior pair in contact with median frontonasal, the anterior pair
separated from rostral by a row of small scales; nasal small, sepa-
rated from rostral, the part anterior to the nostril about one fourth
the diameter of naris; subnasal small; loreal elongate, subequal in
Smith: Genus Sceloporus 665
size to subnasal; preocular large, entire; two canthals, the anterior
smaller, not forced above canthal ridge nor in contact with lori-
labials; subocular followed by one well-differentiated, strongly
keeled postocular; supraocular scales in two rows, the scales of the
median row somewhat larger than those of the lateral row; one
row of small scales between supraoculars and median head scales;
one complete row, and another incomplete row on one side, of scales
between supraoculars and superciliaries; lorilabials reduced to one
row at a point below subocular near its posterior end; four supra-
and five infralabials to a point below middle of eye.
Mental with a labial border about two thirds that of rostral, fol-
lowed b}^ about three pairs of well-differentiated postmentals, the
scales of the anterior pair in contact medially; outer row of labio-
mental scales narrowly separated from mental by a narrow con-
tact of first postmental and first infralabial; inner row of labiomen-
tals terminating below anterior half of third infralabial; gular scales
smooth, mostly entire, those on posterior part of throat weakly
notched at apex.
Three auricular lobules, smooth, rounded, the upper longest, the
middle largest, somewhat larger than preceding scales; scales in
temporal region weakly keeled, weakly mucronate; scales between
ear and lateral nuchal fold somewhat larger than temporal scales,
those on crest of nuchal fold rather strongly keeled and mucronate;
lateral nuchal fold not extremely deep; dorsal scales very weakly
keeled, not mucronate; lateral scales of body more strongly keeled
and mucronate, with numerous lateral denticulations; mucrones of
lateral scales not arising at tip of scale, but a short distance within
the free margin, leaving a denticulate flange about the posterior
edge of the scale; lateral scales considerably larger than dorsals, in
oblique rows; ventral scales about half as large as dorsals, smooth,
rounded or very weakly notched at apex; preanal scales somewhat
smaller than median ventral abdominals; scales on anterior part of
chest larger than scales posterior; subcaudals smooth at base of
tail, keeled and mucronate distally; postanals enlarged, separated
by two small scales; dorsal caudals largest just back of base of tail,
about twice as large as median dorsals on back.
Dorsal scales of foreleg subequal in size, those on humerus some-
what larger and about equal in size to median dorsals of body;
dorsal scales of foreleg keeled, mucronate, the mucrones arising
within the edge of the scale; ventral scales of foreleg keeled and
mucronate (except some on lower foreleg), those on lower foreleg
666 The University Science Bulletin
equal in size to dorsals of same member, about two or three times
as large as scales on ventral surface of humerus; lamellar formula
for fingers, 9-12-16-16-13 (?-ll-16-16-?).
Dorsal scales of hind leg strongly keeled, strongly mucronate
(more so on tibia) , those on femur about equal in size to those on
middle of back, those on tibia much larger, slightly larger than
median lateral scales on body; ventral scales on tibia smooth, mu-
cronate, about two thirds as large as dorsals of same member; ven-
tral scales of femur smooth, rounded or weakly notched at apex,
smaller toward femoral pores, the smallest about equal in size to
preanals; scales on posterior surface of femur keeled, mucronate,
somewhat larger than preanals, abruptly decreasing in size im-
mediately posterior to series of femoral pores; scales on posterior
surface of femur near tail surrounded and separated slightly from
each other by granular scales; no postfemoral dermal pocket; lamel-
lar formula for toes, 8-12-17-18-14 (7-12-17-18-14).
Color. General ground color brownish-gray to light brown (pale
blue in specimens having shed the scales) ; black nuchal collar nar-
row, three or four scales wide, with a narrow, posterior light border,
about one scale wide, interrupted medially; light posterior border
passing onto humerus, with the part on body narrowly bordered
posteriorly with black; light anterior border of black collar broken
into spots; an indistinct light line from upper labial region through
upper portion of ear across neck to black nuchal collar; a few light
flecks on neck anterior to collar; a series of seven small, rounded
black spots of indefinite outline on each side of middorsal line, the
spots not regularly paired; about an equal number of narrow dark
bands on sides of body, one or two scales wide, passing obliquely
laterally and posteriorly, each originating at or near one of the dark
spots on back; a few scattered light flecks on back; limbs dimly
barred or spotted with black or dark gray; tail distinctly banded,
the dark bands about two scales wide, the light bands about one;
each band darker distally than proximally.
Throat with a pale blue ground color, becoming white near chin;
several narrow, oblique, dark blue lines on throat converging pos-
teriorly ; extreme anterior part of chest with two bands encroaching
medially a short distance from the sides ; chest, a narrow band down
middle of abdomen four to six scales wide, ventral surfaces of limbs
and tail cream; sides of belly azure blue, bordered medially by a
broad black band about five scales wide; this black band abruptly
Smith: Genus Sceloporus 667
terminating a short distance posterior to axilla, slightly increasing in
width in region of groin, encroaching slightly upon anterior proximal
surface of femur.
Variation. The variation in head scales is as follows: The supra-
oculars are always in two rows, or one row and part of another.
Occasionally there is but a single scale divided. The frontoparietals
contact medially in thirty ; an azygous scale separates them in four ;
and the frontal contacts the interparietal in nineteen. The frontal
is transversely divided into three scales in three ; the anterior frontal
is longitudinally divided in two. The posterior portion of the frontal
is always smaller than the anterior, varying from two thirds to one
fourth the size of the latter. The frontoparietals are divided into
two scales on one side in two specimens. The median frontonasal
contacts the anterior frontal in twenty-five; it is separated by an
azygous scale in one; the prefrontals contact medially in twenty-
seven. The first canthal touches the rows of scales above the supra-
labials in three specimens ; the preocular is separated from these rows
in three; the second canthal contacts the subriasal, forcing the first
canthal above the canthal ridge, on both sides in sixteen, on one side
in six. The lateral row of the two rows of scales intercalated be-
tween the infralabials and the series of postmentals is invariably
separated from the mental by a partial contact of the first post-
mental and first infralabial.
Other details of scutellation and proportions are given in the ac-
companying table.
The coloration is much as described by Duges (1877), and quite
different from that of d. dugesii, when specimens of both forms are
compared side by side. In dorsal coloration there is but little if any
sexual dimorphism. A narrow black nuchal collar about five or six
scales wide (maximum) is present; its anterior border either blends
more or less insensibly into the ground color of the neck or is
marked by a few small whitish spots; its posterior border is always
distinct and delimited by a narrow white line one and one half to
two scales wide, sometimes broken medially into a series of spots.
Behind this on the back follows a double series of about seven,
usually rounded, black spots of indefinite outline. On the sides of
the body is a corresponding number of oblique black bars, one to
one and one half scales wide, divergent posteriorly, each originating
from or near one of the median black spots on the corresponding
side. These bars and spots are not easily visible on specimens which
668 The University Science Bulletin
have shed their scales, but on all others are easily discernible,
although the black is variable in intensity. The tail is distinctly
banded, the lighter olive bands narrower than the darker olive or
brownish bands.
In males the lateral ventral abdominal areas are usually lavender,
sometimes azure blue ; these areas are bordered medially by a black
band about four scales wide, extending from opposite the axilla to
the groin. The medial abdominal area, ventral surfaces of limbs,
chest and sometimes the throat and gular regions are cream. The
throat region is occasionally pale blue. Oblique, narrow, dark-blue
lines traverse the throat. These lines are always quite distinct, both
in young and in old.
The ventral surfaces of the limbs and abdomen of females are
usually uniformly cream-colored; in some very large specimens a
faint indication of the markings of the male is evident. The gular
region is marked as in the male, except that the ground color is al-
ways cream, not bluish.
Habits and habitat. S. d. intermedius was found most frequently
on rock fences, occasionally on limestone ledges. They are not wary
and sometimes could be caught by hand.
No females have eggs in the oviducts. Very young specimens
were frequently collected in August. In the early part of June I
collected diigesii dugesii on the western coast of Mexico, and at this
time the females had no eggs or young in the oviducts. Very small
individuals were numerous on rock fences. Although positive evi-
dence is lacking, it is assumed from these data that both subspecies
of dugesii are ovoviviparous, and that they must give birth to their
young during May.
Remarks. As may be discerned by comparison of the above de-
scription of intermedius with that of dugesii, there is ample justi-
fication for the recognition of both forms. They differ obviously
in the average number of dorsals from occiput to base of tail; the
median lateral scales are proportionately larger than the median
dorsals in dugesii; the head scales are definitely rugose (micro-
scopically) in dugesii, smooth in intermedins; the frontoparietals
rarely contact medially in dugesii, but usually do so in intermedium ;
the maximum size is greater in dugesii (87.5 mm., snout to vent)
than in intermedius (79 mm.). The oblique dorsal abdominal bars
are absent in dugesii; the gular region is rarely rayed in dugesii as
in intermedius, and never distinctly so.
Cope (1900) remarks that dugesii {d. intermedius) is most closely
related to omatus. The additional material of the latter species in
Smith: Genus Sceloporus
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670 The University Science Bulletin
these collections makes still more evident this relationship. There
are but few points of difference in scutellation. The supraoculars in
intermedius are larger, and the posterior section of the frontal is
less reduced and never entirely suppressed as in ornatus. Limb
proportions are quite different in the two species, ornatus possessing
considerably longer limbs; the fourth toe is also longer. Differences
in coloration are apparent, ornatus lacking the convergent lines on
the throat and the diagonal lines on the sides of the body.
Range. Known only from Guanajuato and northern and central
Michoacan. (See fig. 22 for distributional map.)
Locality records. Guanajuato: (USNM 9904-5, 9886, 9893, A.
Duges; Duges, 1870; Cope, 1879, 1885, 1887); near Acambaro
(EHT & HMS 3651, 3652, 3683-3697, 3940-3948, 3950-3966, 3968-
3972, 4041). Michoacan: (Boulenger, 1898); Zamora (USNM
47791-2, Nelson & Goldman); La Noria (Duges, 1877, 1896; Bou-
lenger, 1897) ; Arroyo Zarco (Duges, 1896) ; near Zinapecuaro (EHT
& HMS 3662-3666) ; Patzcuaro (FMNH 1005 [5 spec], C. E. Meek;
USNM 20145-58, P. L. Jouy; USNM 47219-21, 47223, E. W. Nel-
son). Jalisco: Ocotlan (FMNH 999 [4 spec], S. E. Meek).
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the species occurring north of the isthmus of Tehuantepec, with references.
Bull. Essex Inst. XVI :46 pp.
1887. Reptiles and batrachians from Texas and Mexico. Essex Inst.
Bull. 19:20 pp.
Gravenhorst, J. L. C. 1837. Beitriige zur genauem Kenntniss einiger Eidech-
sengattungen. pp. 713-784, pis. 54-56.
Gray, J. E. 1831. A synopsis of the species of the class Reptilia. Appx. to
Griffith's "Cuvier's 'Animal Kingdom,'" pp. 1-110.
1839. "Tropidolepis torquatus" in Richardson, Zoology of Beechey's
Voyage to the Pacific, p. 95, pi. 30, fig. 2.
1845. Catalogue of the specimens of lizards in the collection of the
British Museum, pp. 1-289.
GtJNTHER, Albert C. L. G. 1890. Biologia Centrali-Americana. Reptilia and
Batrachia. xx, 326 pp., 76 pis.
Herrera, Alfonso L. 1889. Notas acerca de los Vertebrados del Valle de
Mexico. La Naturaleza (2nd Ser.) 1:299-342.
1895. Catalogo de la coleccion de reptiles y batracios del Museo Na-
cional. 66 pp.
1904. Catalogo de la coleccion de reptiles y batracios del Museo Na-
cional. Segunda Edicion. 65 pp.
Herrick, C. L., John Terry, and H. N. Herrick. 1899. Notes on a collection
of hzards from New Mexico. Bull. Sci. Lab. Denison Univ. 11 (6):117-148,
pis. 14-24.
King, F. Willis. 1932. Herpetological records and notes from the vicinity
of Tucson, Ariz., July and August, 1930. Copeia, 1932 (4) : 175-177.
MacCoy, Clinton V. 1932. Herpetological notes from Tucson, Ariz. Occas.
Papers Boston Soc. Nat. Hist. 8:11-24.
Mertens, Robert. 1930. Bemerkungen liber die von Herrn Dr. K. Lafrentz
in Mexiko gesammelten Amphibien und Reptilien Abh. u. Ber. a. d. Mus.
fiir Natur. u. Heimat. u. d. Naturw. ver. 6(2) : 153-161.
MocQUARD, M. F. 1899. Reptiles et batraciens recueillis au Mexique par M.
Leon Diguet en 1896 et 1897. Bull. Soc. Philom. de Paris (9th ser.) 1:154-
169, 1 pi.
Mosauer, Walter. 1932. The amphibians and reptiles of the Guadelupe
mountains of New Mexico and Texas. Occas. Papers Mich. Univ. Mus.
Zool. 246:18 pp., 1 pi.
Mulaik, Stanley. 1935. Tail regeneration in Coleonyx brevis Stejneger.
Copeia, 1935 (3): 155-156.
MtJLLER, J. W. VON. 1865. Reisen in den Vereinigten Staaten, Canada und
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Peale, T. R., and J. Green. 1830. Description of two new species of Linnean
genus Lacerta. Journ. Acad. Nat. Sci. Phila. 6 (1) :231-234.
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Sceloporus from Mexico. Proc. Biol. Soc. Wash. 49:87-96.
1936a. Two new subspecies of Mexican lizards of the genus Sceloporus.
Copeia, 1936 (4) :223-230.
Smith: Genus Sceloporus 673
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• 1916. A new lizard of the genus Sceloporus from Texas. Proc. Biol.
Soc. Wash. 29:227-230.
1918. De.-cription of a new snapping turtle and a new lizard from
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5-126 pp.
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of portions of southern New Mexico and western Texas. Proc. Pliila. Acad.
Nat. Sci., 1903, pp. 16-34.
Strecker, John K. 1909. Contributions to Texan herpetologj\ Reptiles and
amphibians collected in Brcw.stcr county, Texas. Baylor Univ. Bull. 12
(1):11-16.
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1928. Common Engli.'<li folk names for Texas amphibians and reptiles.
Baylor Univ. Mus. Contr. 16:21 pp.
• 1933. Collecting at Helotes, Bexar county, Texas. Copeia, 1933 (2) :
77-79.
1935. Notes on the zoology of Texas. Bajdor Univ. Bull. 38 (3) :viii,
69 pp.
Stu.\rt, L. C. 1934. A contribution to the knowledge of the herpetological
fauna of El Peten, Guatemala. Occas. Papers Mus. Zool. Univ. Mich.
292:1-18, 1 fig.
. 1935. A contribution to a knowledge of the herpetology of a portion
of the savanna region of the central Peten, Guatemala. Misc. Publ. Univ.
Mich. Mus. Zobl. 20:56 pp., pis. 1-4, map.
Sumichr.ast, Fr.ancois. 1873. Coup d'oeil sur la distribution geographique
des reptiles au Mexique. Arch. Sci. Phj^s. Natur. 46:233-250.
1882. Eniuneracion de las especies de reptiles observados en la parte
meridional de la republica Mexicana. La Naturaleza 6:31-45.
Taixor, Edward H. 1931. The discovery of a lizard Sceloporus torquatus
cyanogenys Cope in Texas, new to the fauna of the United States. Proc.
Biol. Soc. W^ash. 44:129-132.
Van Denburgh, John. 1896. A list of some reptiles from southeastern Ari-
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known from that state. Proc. Calif. Acad. Sci. (4th ser.) 13 (12) :189-230.
43—6037
674 The University Science Bulletin
and Joseph R. Slevin. 1913. A list of the amphibians and reptiles of
Arizona, with notes on the species in the collection of the Academy. Calif.
Acad. Sci., Pioc. (4th ser.) 3:391-454, pis. 17-28.
Wagler, J. 1830. Natiiriiches System der Amphibien . . . pp. 1-354.
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von Centralamerika und Chile, sowie einiger seltenerer Schlangenarten.
Verb. K. K. Zool.-Botan. Ges. Wien 46:344-365, pi. 6.
1903. Ueber Reptilien und Batrachier aus Guatemala und China in der
zoologischen Staats-Sammlung in Miinchen nebst einem Anhang iiber sel»
tene Formen aus anderen Gegenden. Abh. der Iv. Bayer. Akad. Wiss. II
KL, XXII Bd., II Abt., pp. 343-384, 1 col. pL, 4 figs.
WiEGMANN, A. F. A. 1828. Beitriige zur Amphibienkunde. Isis 1828 (3) :364-
383.
1834. Herpetologia Mexicana. vii, 55 pp., 10 pis.
Yarrow, H. C. 1875. Report upon the collections of batrachians and reptiles
made in portions of Nevada, Utah, California, Colorado, New Mexico and
Arizona, during the years 1871, 1872, 1873, 1874. Geog. Geol. Surv. W. 100th
Mer. 5:509-584, pis. 16-25.
1883. Checklist of North American reptilia and batrachia. Bull. U. S.
Nat. Mus. 24:vi, 249 pp.
-, and H. W. Henshaw. 1878. Report upon the reptiles and batrachians
collected during the years 1875-1877 in California, Arizona, and Nevada.
Ann. Rept. Chief of Engineers to Sec. War. for 1878 (Part 3) Appx. L, in
Appx. NN, pp. 1628-1648.
676
The University Science Bulletin
PLATE XLVH
Fig. 1. Sceloporus jarrovii jarrovii Cope. KU 13151, male. Santa Cniz
county, Arizona. Snout-to-vent mea.*urenient. 80 mm.
Fig. 2. Sceloporus serrifer Cope. UMMZ 72890, male. Near Merida, Yuca-
tan. Snout-to-vent measurement, 101 nun.
Smith: Genus Sceloporus
677
PLATE XLVII
678 The University Science Bulletin
PLATE XLVIII
Fig. 1. Sceloporiis torquatus torquatus Wiegmann. EHT & HMS 3785,
male. Near Uruapan, Michoacan. Snout-to-vent measurement, 101 mm.
Fig. 2. Sceloporus torquatus melanog aster (Cope). DHD & HMS 691, fe-
male. La Colorada, Zacatecas. Snout-to-vent mcastnement. 119 mm.
Smith: Genus Sceloporus
679
PLATE XLVIII
680 The University Science Bulletin
PLATE XLIX
Fig. 1. Sccloporus bullcri Boulenger. USXM 64664, female. La Lagima,
Jalisco. Snout-to-vent nieaf^urcnient, 83.5 nun.
Fig. 2. Sccloporus mucroiuifus mucwnaliis (Cope). EHT & HMS 2096, fe-
male. Near Toxtlacuaya, about 18 niile.-^ west of Jalapa, Vera Cruz. Snout-to-
Aent measurement, 95 mrn.
Smith: Genus Sceloporus
681
PLATE XLIX
#^
f.?*^
682 The University Science Bulletin
PLATE L
Fig. 1. Sceloporus mucronatus omiltemanus (Giinther). EHT & HMS
3094, male. Two miles west of Afultzingo, Vera Cruz. Snout-to-vcnt measure-
ment, 86.5 mm.
Fig. 2. Sceloporus mucronatus omiltemanus (Giinther). MCZ 33904, male.
Chilpancingo. Guerrero. Snout-to-vent measurement, 95 mm.
Smith: Genus Sceloporus
683
PLATE L
684 The University Science Bulletin
PLATE LI
Fig. 1. Sceloporus poinseitii Baird and Girard. LMK 24283, female. Gat-
ton's Park, Grant county. New Mexico. Snout-to-vent measurement, 91 mm.
Fig. 2. Sceloporus cyanogemjs (Cope). EHT & HMS 5053. male. Arroyo
I>os Olmos, three miles southeast of Rio Grande City, Starr county, Texas.
Snout-to-vent measurement. 132.5 mm.
Smith: Genus Sceloporus
685
PLATE LI
686 The University Science Bulletin
PLATE LII
Fig. 1. Sceloporus lineolateralis Smith. EHT & HMS 4332, male. Six miles
northeast of Pedricefia, Durango. Snout-to-vent measurement, 71 mm.
Fig. 2. Sceloporus UneoIateraUs Smith. EHT & HMS 4363. female. Six
miles northeast of Pedriceila, Durango. Snout-to-vent measurement, 73 mm.
Smith: Genus Sceloporus
687
PLATE LII
G88 The University Science Bulletin
PLATE LIII
Fig. 1. Sceloporus jarrovii immucronatus Smith. EHT & HMS 500, male.
Near El Pinalito, Hidalgo. Snout-to-vent measurement, 86 mm.
Fig. 2. Sceloporus jarrovii 7ninor (Cope). EHT & HMS, 4207, male. Near
San Felipe, Guanajuato. Snout-to-vent measurement, 97 mm.
Smith: Genus Sceloporus
689
PLATE LIII
isF ■
^•^iil^
z^, Y
-V
-*;,^2 . ■'^.
«.^;^:^
.S.^^^'^
44—6037
690 The University Science Bulletin
PLATE LIV
Fig. 1. Sceloporus ornatus ornatus Baird. EHT & HMS 4420, male. Near
Saltillo, Coahuila. Snout-to-vent measurement, 72.5 mm.
Fig. 2. Sceloporus ornalus caeruleus Smith. DHD & HMS 350, male. Five
miles south of San Pedro, Coahuila. Snout-to-vent measurement, 75 mm.
Smith: Genus Sceloporus
691
PLATE LIV
692 The University Science Bulletin
PLATE LV
Fig. 1. Sceloporus dugesii dugesii Bocourt. EHT & HMS 2770, female.
Near Magdalena, Jalisco. Snout-to-vent mea.sui-ement, 79 mm.
Fig. 2. Sceloporus dugesii intermedium (Duges). EHT & HMS 3961, fe-
male. Near Acambaro, Guanajuato. Snout-to-vent measurement, 68 mm
Smith: Genus Sceloporus
693
PLATE LV
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