UNI iou
HARVARD UNIVERSITY
LIBRARY
OF THE
Museum of Comparative Zoology
| aay MOS. eee mon |
buen <}- ; a a hiv!
| é ¢ oe
; JAN 1.7 1087
eo JAN ii
sy « PA A Ga | ed
| UNIVERSITY OF KANSAS 4_Jiiitisity
_ SCIENCE BULLETIN
UNIVERSITY OF KANSAS PUBLICATIONS
- University of Kansas Science Bulletin - Vol. XXXVIII - Part I
December 20, 1956
Lawrence, Kansas
—
ANNOUNCEMENT
The University of Kansas Science Bulletin (continuation of the
Kansas University Quarterly) is issued in parts at irregular inter-
vals. Each volume contains from 300 to 700 pages of reading mat-
ter, with necessary illustrations. Exchanges with other institutions
and learned societies everywhere are solicited. All exchanges should
be addressed to
Tue Universrry or Kansas Scrence BuLLeTIN, —
Liprary or THE University or Kansas,
Lawrence, Kan.
PUBLICATION DATES
The actual date of publication (i. ¢., mailing date) of many of the
volumes of the University of Kansas Science Bulletin differs so
markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
time as the whole volume.
Vol. Vol.
XX—October 1, 1932. XXXI, Pt. I—May 1, 1946.
XXI—November 27, 1984. . Pt. II—Nov. 1, 1947.
XX1II—November 15, 1935. XXXII—Nov. 25, 1948.
XXIII—August 15, 1936. XXXII, Pt. I—April 20, 1949.
XXIV—February 16, 1938. Pt. I[—March 20, 1950.
XXV—July 10, 1939. XXXIV, Pt. I—Oct. 1, 1951.
XXVI—November 27, 1940. : Pt. Il—Feb. 15, 1952,
XXVII, Pt. I—Dec. 80, 1941. XXXV, Pt. I—July 1, 1952.
XXVIII, Pt. I—May 15, 1942. Pt. I1—Sept. 10, 1953.
. Pt. Il—Nov. 12, 1942. Pt. III—Nov. 20, 1953.
XXIX, Pt. I—July 15, 1943. XXXVI, Pt. I—June 1, 1954.
Pt. Il—Oct. 15, 1943, Pt. [1—July 15, 1954.
XXX, Pt. I—June 12, 1944. XXXVII, Pt. I—October 15, 1955,
Pt. —June 15, 1945. XXXVII, Pt. [1—June 29, 1956.
HOWOE © Epwarp H. Taytor
Editorial Board .............. Epwarp H. Tayior, Chairman
CHar.es MicHENER
Pau. Roorr
Dwicur J. Muurorp
Wortme H, Horr
Parke H. Wooparn, Secretary
US. GUMP, 09) |
[YAN 1 7 1957
HARVARD
GnuveRsiT
i
ee]
ERRATA
UNIVERSITY OF KANSAS SCIENCE BULLETIN
vol. XXXVIITI, pt. I. Dec. 20, 1956.
Page 3, line 9; p. 6, line 11; p. 12, line 3; p. 297, line 13; p. 306, line
40; p. 308, line 1; p. 309, line 1 for brachypodus read brachypoda.
page 441, line 10, insert “that” after “structures”
page 443, line 33, for Petiloglossa read Ptiloglossa
page 464, line 10, for Anthopora read Anthophora
page 474, line 2, for (teunomia) read (Dieunomia)
page 488, line 6, for Fimbriata read fimbriata
page 468, line 26; p. 469, line 6; p- 470, lines 7, 8, 9, 20; p. 490, lines
4, 5, 6, for mellifica read mellifera.
page 582, line 27, for 582, read 634.
page 589, line 27, for 633-34, read 634-641.
page 608, line 3, for “amounts,” read “mounts.”
page 616, line 22, for 616, read 614.
page 919, line 28; p. 987, line 42, for “scent gland opening” read
“metathoracic spiracle.”
page 1028, line 45, for “are” read “is.”
THE UNIVERSITY OF KANSAS
SCIENGE BULLETIN
Votumr XXXVIII Parr I
DepicaTEp TO
HERBERT BARKER HUNGERFORD
Eprror 1933-1940
+(e ae His editorship, volumes XXI through XXVI were
Issued. In 1945-46, in the absence of the then editor in Asia,
Doctor Hungerford edited volume XXX, part 2, and volume
XXXI, part 1. Again in 1947 he likewise edited volume XX XII
during the absence of the editor in Central America. Due to
an oversight his name was omitted as editor from this latter
volume.
Through the years Doctor Hungerford has been an active
Contributor to the Screncr Butuetin, the first of his articles
appearing in volume VIII in 1914. Since that date he has
Published eleven other papers in the Screncr BULLETIN, two
of them studies of monographic extent dealing with cosmopoli-
tan families of insects and involving the greater part of two
Whole volumes. Other studies on his part are in progress at
the present time.
Lawrence, Kansas Jung, 1956
HERBERT BARKER HUNGERFORD
pee carEER of Herbert Barker Hungerford, spanning forty-five
years at the University of Kansas has been one of inspirational
teaching and productive research in his chosen field of entomology.
His adult life, devoted almost entirely to one institution, can be
measured by the renown and esteem which he has brought to the
school whose destiny he has helped to mold. She in turn must
acknowledge him as one of her great, one who has added prestige
and honor to her accumulated distinction. The name of Hunger-
ford should be rightfully classed with those of Snow, Bailey,
Williston, Moore, Stevens, and others, as great Kansans in the field
of science.
His is the story of a true son of Kansas. He was born at Mahaska,
August 30, 1885, of pioneer stock, the son of Artemus Manwarring
and Bertha Estelle (Barker) Hungerford. Early life in a rural
community undoubtedly engendered an interest in the living world
which remained an abiding passion throughout a distinguished and
productive career as educator and man. His early education was
obtained in the local Kansas schools and at the Kansas State Normal
School (now Kansas State Teachers College) at Emporia. At the
onset, family plans anticipated that the son would follow in the
father’s pattern as landholder and grain operator. Advisers pre-
vailed upon him, however, to validate his teaching certificate with
a year’s experience, as was then required by State Board of Educa-
tion regulations. This idea was not entirely foreign to his thinking,
for the boy had long been an admirer of his maternal grandfather,
George William Barker, who had been principal of the Genoa
Academy in Cayuga County, New York state. The necessary year
of experience was lengthened into several through jobs as teacher
and principal in the school system of Haddam (1904-06), and later
as superintendent of schools at Enterprise (1906-'09).
On August 30 (his birthday), 1905, Hungerford married a col-
lege friend, Mary Frances Kenney, of Paola. Their only child, a
daughter, Helen Estelle, graduated as an honor student from the
University of Kansas in 1930. She is now Mrs. Andrew Benedict
Hamilton of St. Petersburg, Fla.
As a teacher Hungerford attended the University of Kansas at
Lawrence for the summer sessions and during these enrollments
developed his earlier interest in entomology. Giving up his teach-
[1]
ing he enrolled at the University as a full-time student in the fall
of 1909, receiving his bachelors degree in 1911. During his senior
year he was elected to Phi Beta Kappa in recognition of his scho-
lastic achievement. Later (1948-50) he served this chapter as its
president.
Upon receipt of his baccalaureate degree, Hungerford was ap-
pointed as an instructor in entomology at the University of Kansas
and progressed rapidly up the academic ladder as assistant pro-
fessor (1913), associate professor (1917), and professor (1920);
becoming head of the department of entomology in 1924, a position
which he held for twenty-five years. In 1949 he relinquished his
administrative duties for what to him was a more interesting life
as research professor of entomology, and continued in this capacity
until his retirement in 1956.
Doctor Hungerford’s continuous service at the University of
Kansas was twice interrupted. In 1916 he was granted a year’s
leave of absence to continue graduate studies at Cornell University
where he received his Ph. D. degree in 1918. — Note on Pilea,
Bull. Brooklyn Ent. Soc., vol. 23, no. Sip 182,
——— Some South American Corixidae,
Bull. Brooklyn Ent. Soc., vol. 23, no, 4, pp. 174-179; 2 plates.
1929, Two new species of Hemiptera in the collections of the Museum
National of Paris (Ranatra wagneri and Velia conata),
Bull. du Mus, Paris, 2° serie, tome 1, no. 8, pp. 198-200; 1 plate,
3 figures,
~——— The European elm seale (with George A. Dean),
Kansas Ent. Commission Circ, 9, 8 pp.; 1 plate, 3 figures,
———— A new Velia from Peru. (Hemiptera, Veliidae. )
Ent. Tidskr., vol. 50, no. 2, pp. 146-147,
—Some new semiaquatics from North America with a record of stridu-
latory devices. (Veliidae Velia. )
Jour. Kans, Ent, Soc., vol. 11, no. 38, pp. 50-59; 2 plates.
“—— Concerning Velia inveruglas Kirkaldy and related forms. ( Veliidae-
Hemiptera).
Ann. Ent. Soc. Amer., vol. 23, no. 1, pp. 120-125; 1 plate, 13 figures.
~~ Three new Velia from South America,
Jour. Kansas Ent, Soc., vol. 3, no. 1, pp. 23-27; 1 plate, 11 figures.
~——— A new Velia from Arizona with notes on other species. (Hemiptera-
Veliidae, )
Ann. Ent. Soc. Amer., vol, 22, no. 4, pp. 759-761; 4 figures,
——— Searching for types in Europe’s Musty Museums.
Graduate Magazine (K. U.), vol. 27, pp, 18-14: 4 photographs.
— A new genus of semiaquatic Hemiptera,
Bull. Brooklyn Ent. Soc., vol, 24, no. 5, pp. 288-291; 1 plate.
— Concerning two of Guerin-Meneville’s types in the National Museum
of Paris (Hemiptera; Notonectidae and Corixidae ).
Pan-Pacific Emits, VOle 00.2, pp. 78-775 figures.
~——— Report on the nomenclature of some neotropical Notonecta with the
descriptions of some new species (Hemiptera-Notonectidae ),
Bull. Brooklyn Ent. Soc., vol. 25, no, 8, pp. 138-143; 2 figures.
a
es
Se
Se
[1x]
An unusual nest of Vespula.
Ent. News, vol. 41, pp. 329-330; 1 plate, 2 figures.
1930. New Snow Hall takes its place on the campus.
Graduate Magazine (K. U.), vol. 28, pp. 7-10.
New Corixidae from western North America (Hemiptera).
Pan-Pacific Ent., vol. 7, no. 1, pp. 22-26; 1 plate.
Two new water bugs from the western U.S.A. (Nepidae and Noto-
nectidae).
Canadian Ent., vol. 62, no. 10, pp. 216-218.
Concerning the egg of Polystoechotes punctatus Fabr. (Neuroptera).
Bull. Brooklyn Ent. Soc., vol. 26, no. 1, pp. 22-23; 1 figure.
A new Velia from Trinidad. (Hemiptera-Veliidae. )
Annals and Magazine of Nat. Hist., ser. 10, vol. 7, pp. 174-175.
1931.. The Hydrometridae of the Hungarian National Museum and_ other
studies in the family. (Hemiptera) (with N. W. Evans).
Harvath, Hungary. Ann. Musei Nat. Hungarici, vol. 28, pp. 31-112;
12 plates.
1932. New Notonecta of the N. mexicana A. and S. Group.
Jour. Kansas Ent. Soc., vol. 5, no. 2, pp. 53-55.
Concerning a fossil water bug from the Florissant (Nepidae ).
Univ. Kansas Sci. Bull., vol. 20, pt. 2, pp. 327-330; 1 plate.
A new Platygerris with notes on P. caeruleus Champion (Gerridae ).
Bull. Brooklyn Ent. Soc., vol. 27, no. 4, pp. 178-182.
The male of Notonecta compacta Hungerford.
Bull. Brooklyn Ent. Soc., vol. 28, no. 3, pp. 185.
A new Potamobates (Gerridae).
Bull. Brooklyn Ent. Soc., vol. 27, no. 5, pp. 228-230.
Report of some insect outbreaks of the past season.
Biennial Rept. Kansas State Hort. Soc., vol. 41, pp. 89-41.
1933. Some aquatic and semiaquatic Hemiptera from Sumatra.
Miscellanea Zoologica Sumatrana, vol. 75, pp. 1-5.
Report upon the aquatic and semiaquatic Hemiptera collected by E. P.
Creaser in Yucatan and Campeche in 1932,
Publication Carnegie Institute 1933, pp. 145-150.
Concerning some aquatic and semiaquatic Hemiptera from Australia.
Bull. Brooklyn Ent. Soc., vol. 29, no. 2, pp. 68-73,
[A new Hydrometra from British Honduras] (Hemiptera-Hydrometri-
dae).
Jour. Kansas Ent. Soc., vol. 6, no. 4, pp. 142-143,
1934. A new Notonecta from Mexico (ft lemiptera ).
Jour. Kansas Ent. Soc., vol. 7, no. 3, pp. 97-98.
1933. The genus Notonecta of the World.
Univ. Kansas Sci. Bull., vol. 21, pp. 5-195; 17 plates.
1934. The oriental fruit moth in Kansas.
Bien. Rept. Kansas State Hort. Soc., vol. 42, pp. 57-59.
1935. A termite new to Kansas.
Jour. Kansas Ent. Soc., vol. 8, no. 1, p. 24.
The genus Bacillometra Esaki including the description of a new
species from Peru.
Revista de Entomologia, vol. 5, fasc. 2, pp. 117-124; 2 figures,
[x]
Aquatic and semiaquatic Hemiptera collected in Yucatan and Cam-
peche,
Carnegie Institute Washington, publ. no. 457, pp. 145-150; 1 figure.
1986. The Mantispidae of the Douglas Lake, Michigan Region with some
biological observations.
Ent. News, vol. 47, pp. 69-72, 85-88; 1 plate.
A new Potamobates from Peru.
Bull. Brooklyn Ent. Soc., vol. 31, pp. 178-180.
———- The male of Notonecta arabiensis Hungerford (Notonectidae, Hemip-
tera),
Jour. Kansas Ent. Soc., vol. 9, pp. 101-102; 1 text figure.
——— Recent information concerning some approaching plant pests.
Biennial Rept. Kansas Hort. Soc., vol. 48, pp. 111-115.
1987, A second new Potamobates from Peru, S. A., with notes on other
species (Hemiptera-Gerridae ),
Bull. Brooklyn Ent. Soc., vol. 82, no. 4, pp. 144-147; 1 plate.
A new Potamobates from Mexico ( Hemiptera-Gerridae ).
Jour, Kansas Ent. Soc., vol. 10, no. 2, pp. 63-65.
~——— Pseudomasaris occidentalis (Cresson) in Kansas.
Jour. Kansas Ent. Soc., vol. 10, pp. 133-134,
———— A new Notonecta from Mexico ( Hemiptera-Notonectidae ).
Pan-Pacific Ent., vol. 13, no. 4, pp. 180-182; 1 figure.
1988. A new Hydrometra from New Caledonia and Australia,
Pan-Pacific Ent., vol. 14, pp. 81-83.
——— A new Graptocorixa from Mexico ( Corixidae-Hemiptera ).
Jour. Kansas Ent. Soc., vol. 11, no. 1, pp. 28-80,
~—— A third new Potamobates from Peru, 8. A. (Hemiptera-Gerridae ),
Jour, Kansas Ent. Soc., vol. 11, pp. 85-87.
~—— Report upon some water bugs from Mexico collected by Mr. Meldon
Embury.
Pan-Pacific Ent., vol. 14, pp. 76-81; 1 plate.
~—— Mesoveloidea williamsi Hungerford—A note on its distribution.
Bull. Brooklyn Ent. Soc., vol. 33, no. 5, p. 218,
~—— Insects in the affairs of men.
Graduate Magazine (K.U.), vol. 36, pp. 6-8.
1989; Two new genera of Hydrometridae from the Marquesas Islands (He-
miptera ).
Pac. Ent. Survey Publication 8, art. 25, pp. 217-220; 1 figure. (B. P.
16 Bishop Mus. Bull. 142.)
38. A new species of Neocorixa ( Corixidae-Hemiptera )
Bull. Brooklyn Ent. Soc., vol. 33, no. 4, pp. 170-172; 1 plate.
~—— Some new Graptocorixa from Mexico and other notes (Corixidae-He-
miptera ),
Jour. Kansas Ent. Soc., vol. 11, no. 4, pp. 134-141; 1 plate,
1939. A note on Sigara griffini (Kirk.).
Jour, Kansas: Ent, Soc., vol. 12, no. 2, p. 72.
~——— A Corixid from deep water,
Ann. Ent. Soc, Amer., vol. 82, no. 3, pp. 585-586.
~———— A report on some water bugs from Costa Rica.
Ann. Ent. Soc. Amer., vol. 32, no. 3, pp. 587-588.
[x1]
Two new Corixidae from Bolivia, South America.
Jour. Kansas Ent. Soc., vol. 12, no. 8, pp. 97-99; 1 plate.
Two new Corixidae from Mexico.
Jour. Kansas Ent. Soc., vol. 12, no. 4, pp. 123-125; 1 plate.
Oberea bimaculata Oliy. injuring perennial asters.
Jour, Econ, Ent., vol. 82, no. 4, p. 596,
—_—— A note on Mantispidae.
Bull. Brooklyn Ent. Soc., vol. 34, no. 5, p. 265.
———— A new Corixid from Mexico,
Jour. Kansas Ent. Soc., vol. 12, no. 4, pp. 133-134; 1 figure.
1940. New Corixidae from China, Manchuria and Formosa.
Jour. Kansas Ent. Soc., vol. 13, no. 1, pp. 8-14; 2 plates.
_ A new Enithares for Australia (Notonectidae-I femiptera ).
Jour. Kansas Ent. Soc., vol. 13, no. 4, pp. 130-131; 1 plate.
Results of the Oxford University Cayman Islands Biological Expedition
of 1938 (Aquatic Hemiptera ).
Ent. Month. Mag., vol. 76, pp. 255-256.
1941. A new Corixid from China.
Jour. Kansas Ent. Soc., vol. 14, no. 1, pp. 20-21; 1 plate.
A remarkable new Naucorid water bug.
Ann. Ent. Soc. Amer., vol. 34, pp. 1-4; 1 plate.
_____. New distributional note on Notonecta borealis B. & H.
Jour. Kansas Ent. Soc., Vole, nove, pod!
——— Concerning Trichocorixella Jacz. (Hemiptera-Corixidae ).
Jour, Kansas Ent. Soc., vol. 15, no. 2, pp. 62-63; 1 plate.
1942, Three new Corixidae from the Southern States.
Bull. Brooklyn Ent. Soc., vol. 37, no. 4, pp. 127-131; 1 plate.
Coleopterocoris, an interesting new genus of the subfamily Potamoco-
rinae (Naucoridae: Heteroptera).
Ann. Ent. Soc. Amer., vol. 35, no. 2, pp. 135-139.
___. New record for Notonecta borealis Bueno and Hussey.
Bull. Brooklyn Ent. Soc., vol. 87, no. 2, p. 61.
___ A new Corixid from Minneosta (with R. I. Sailer).
Bull. Brooklyn Ent. Soc., vol. 37, no. 5, pp. 179-180; 1 plate.
1943. The tropical rat mite in Kansas.
Jour. Kansas Ent. Soc., vol. 16, no. 4, p. 154,
Relation of entomology to the war effort.
Trans. Kansas Acad. Sci., vol. 46, pp. 303-308.
1944, Synonymic notes in the genus Glaenocorisa.
Bull. Brooklyn Ent. Soc., vol. 89, no. 1, pp. 82-84,
—_—— Some Venezuelan aquatic Hemiptera.
Zool. Sci:, Contributions of the New York Zool. Soc., Vol ees pi os
Dy AZ;
1945. ‘The sweet potato leaf beetle Typophorus viridicyaneus (Crotch).
Jour. Kansas Ent. Soc., vol. 18, no. 4, pp. 154-155.
1946. A new genus and species of Notonectidae.
Jour, Kansas Ent. Soc., vol. 19, no. 2, pp. 59-62; 1 plate.
1945. On the status, under Article 25 of the International Code of Specific
Names published with descriptions but without comparison with
allied species.
Bull, Zool. Nomencl. 1945, p. 102.
[xu]
———— Book Review. Argosidae of North America.
Ann, Ent. Soc. Amer., vol. 37, p. 13.
1946. Bed Bug.
Encyclopaedia Britannica, vol. 3, p. 295.
-— Bug;
Encyclopaedia Britannica, vol. 4, p. 345.
——— Chinch Bug.
Encyclopaedia Britannica, vol. 5, p. 555.
——— Harlequin Bug.
Encyclopaedia Britannica, vol. 11, p. 198.
~——— Hemiptera.
Encyclopaedia Britannica, vol. 11, pp. 416-420,
——— Water Boatman.
Encyclopaedia Britannica, vol. 23, p. 407.
———— Water Scorpion.
Encyclopaedia Britannica, vol. 23, p. 480.
1947, A new genus of Corixidae.
Jour. Kansas Ent. Soc., vol. 20, no. 3, p. 98.
~——— A new species of Cymatia from Australia (Hemiptera, Corixidae ).
Jour. Kansas Ent. Soc., vol. 20, no. 4, pp. 154-157; 1 plate.
1948. The eggs of Corixidae (Hemiptera).
Jour. Kansas Ent. Soc., vol. 21, no. 4, pp. 141-146; 2 plates.
——— The Corixidae of the Western Hemisphere (Hemiptera).
Univ. Kansas Sci. Bull., vol. 32, pp. 5-827; 19 figures; 112 plates (in-
cludes paper by Reece I. Sailer, The genus Trichocorixa [Corixidae,
Hemiptera], pp. 289-407; plates 47-54),
Two new generic names,
Jour. Kansas Ent. Soc., vol. 23, no. Se ore any,
~——— On the distribution of Notonecta petrunkevitchi Hutchinson (Hemip-
tera-Notonectidae ).
Jour. Kansas Ent. Soc., vol, 23, no. 8, p. 98.
~— Recent advances with insecticides,
Biennial Rept. Kansas Hort, Soc., vol. 50, pp. 54-57,
A new Mesovelia from Mexico and Guatemala.
Jour. Kansas Ent. Soc., vol. 24, pp. 32-34; 1 figure.
~—— A new Metrobates from Brazil, South America (Hemiptera-Gerridae),
Jour. Kansas Ent. Soc., vol. 24, pp. 72-73; 1 figure.
~——— A new Hydrometra from Mauritius.
Jour. Kansas Ent, Soc., vol. 24, pp. 109-111; 1 plate.
“~—— Concerning some Hydrometra from Africa (Hemiptera ).
Psyche, vol. 58, no. 2, pp. 65-72; 1 plate.
~~—— An interesting new gerrid from Madagascar.
Jour. Kansas Ent. Soc., vol. 24, pp. 131-133; 1 plate.
A new Agraptocorixa from New Guinea ( Corixidae-Hemiptera ).
Jour. Kansas Ent. Soc., vol. 26, pp. 39-40; 1 plate.
A TOV. Agraptocorixa from Australia.
Jour, Kansas Ent, Soc., vol. 26, pp. 43-44; 1 plate.
~—— Concerning Charmatometra bakeri (Kirkaldy ).
Ent. News, vol. 64, pp. 172-175.
“—— Concerning Rheumatobates rileyi Bergroth.
Ent. News, vol. 64, pp, 91-92; 1 text figure.
1950,
1951,
1953,
[xu]
Concerning Mesovelia douglasensis Hungerford.
Jour. Kansas Ent. Soc., vol. 26, pp. 76-77.
1954. The genus Rheumatobates Bergroth (Hemiptera-Gerridac ).
Univ. Kansas Sci. Bull., vol. 86, pp. 529-588; 9 plates.
____ First Florida record for Hydrometra consimilis Barber.
Jour. Kansas Ent. Soc., vol. 27, p. 80.
Paul Bowen Lawson.
Jour. Kansas Ent. Soc., vol. 27, no. 3, pp. 81-83.
1955. A subaquatic light trap for insects (with Paul Spangler and Neil A.
Walker).
Trans. Kansas Acad. Sci., vol. 58, no. 3, pp. 387-407.
A new Limnometra from Sumatra.
Jour. Kansas Ent. Soc., vol. 28, pp. 67-68.
1956. A new Cenocorixa from the northwestern United States.
Jour. Kansas Ent. Soc., vol. 29, no. 2, pp. 39-41; 1 plate.
UNIVERSITY OF KANSAS
SCIENCE BULLETIN
DEVOTED TO
THE PUBLICATION OF THE RESULTS OF
RESEARCH BY MEMBERS OF THE
UNIVERSITY OF KANSAS
VOLUME XXXVIII, Parr I
University or KANSAS PUBLICATIONS
Lawrence, DecemBeR 20, 1956
PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1104,
2.
10.
Contents of Volume XXXVIII, Part I
Number
Ale
A Review of the Lizards of Costa Rica.
Edward H. Taylor,
Selective Sensitivity of the Formed Elements of the Blood
of the White Rat as Affected by Radium Chloride In-
jected Intraperitonedlly. 9.5... ... Paul G, Roofe,
Studies on Kansas Mosses II.
R. L. McGregor and E. L. Hartman,
. Notes on the Vegetation of Southeastern Mount Desert Is-
WAT C MAING a ee A. W. Kiichler,
Biological and Systematic Studies of Two Species of
Cheyletid Mites, with a Description of a New Species
(Acarina, Cheyletidae ).
Robert E. Beer and David T. Dailey,
Comparative Studies of the Thoracic Musculature of Bees.
Alvaro Wille,
Catalogue of the Types in the Snow Entomological Mu-
seum. Part I (Hymenoptera) ....Wallace E. LaBerge,
A Revision of the Bees of the Genus Melissodes in North
and Central America. Part II (Hymenoptera, Apidae ).
Wallace E. LaBerge,
The Biology and Morphology of Hydrometra martini
Ita coe oe ey Isabelle Baird Sprague,
A Taxonomic Study of the Genus Rhagovelia (Hemiptera,
Veliidae ) of thé Western Hemisphere. .John A. Bacon,
. A Supplementary Taxonomic Study of the Genus Rhago-
velia (Hemiptera, Veliidae) of the Western Hemi-
sphere. A Deductive Method....... Ryuichi Matsuda,
Concerning the Relationship of Certain Neotropical Gek-
konid Lizard Genera, with Comments on the Micro-
scopical Structure of Their Glandular Scales.
Edward H. Taylor and A. Byron Leonard,
PAGE
915
1019
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVI, Pr. I] December 20, 1956 [No. 1
A Review of the Lizards of Costa Rice
BY
Epwarp H. Taytor *
Apsstracr. The lacertilian fauna of Costa Rica is reviewed on the basis of
collections made by the author on four Costa Rican expeditions. The fauna
is discussed and listed. The following forms are described as new: Anolis
humilis marsupialis, Anolis woodi attenuatus, Anolis biscutiger, Anolis aquaticus,
Anolis achilles, Ameiva festiva occidentalis, Letolopisma cherriet lampropholis,
Mabuya brachypodus. Altogether seventy-nine species and subspecies are
treated. Each form is described and for the most part illustrated by photographs.
CONTENTS
PAGE
MUP UOO a orem cans : 6
Distribution of Costa Aloan. gard families and WOTCUH cess necks. 7
History of collections and literature.............. Sle ae er agatt els 8
axonomy of Costa Rican Sautin, «oe 12
IWOVMLOeMRILMGA: pata th os a we a Ae ee ere Oe
Diperamily Gekkonoidea......,........... eee ata ei tee 13
I hagillhye Wjhfoitcrole(enakdlalcte a Weed ee a brie Gt alee
Genie (GOleOn ve Inve a 14 att ia ncth wae cbek Gece fo ‘peu itae it 15
Coleonyx mitratus (Peters)........... ‘pee Ohi 16
Family Sphaerodactylidae ....... aes Piadoer te ie Le
Key to the genera of Sphaerod: abtyBdae. TERRE SES radi cas SoHoe a 20
Genus Lepidoblepharis Peracca .. . . ; Gis asidaths 20
Lepidoblepharis xanthostigma Noble. 20
(Coehici1Gieyeuziicolal pda li abohet:in a Oaae cies ae wee ata ae aa ee diy ie 27
Gonatodes fuscus Hallowell 27
Genus Sphaerodactylus Wagler ..... be RT Sa ee ake, BY
Key to the species of Sphaecrodactylus .......... aera = : 33
Sphaerocactylus pacttious Stejneger...............-...: 34
Sphacrodactylus homolepis Cope....... 38
Sphaerodactylus lineolatus Lichtenstein and von Me artet ise D
Sphaerodactylus imbricatus Anderson pes eat Aa 45
Sphaerodactylus millipunctatus Hallowell............... 49
Sphaerodactylus sp. ....... Par ie ip beh se eia NO
Family Gekkonidae .......... atc ee re ee a 54
* Department of Zoology, University of Kansas.
(3)
Tse Universiry SCIENCE BULLETIN
PAGE
Genws -Theondaowvits Cimon. yi ee a oe a na 54
Thecadactvius tapicaudus HOuttivns 0) ayia oa) es 55
Genus PhivllodaciyitseGiaye «acre aw isu ees see 58
Phyllodactylus eduardofisheri Mertens................... 59
Pamilyelguanidae. 9 is eo eas 1 es eae 63
Key to the genera of Costa Rican Iguanidae..............1....4 OE
Gens Anolis’ Dauditiey ey sii orev ieee ca One® Win Cire 64
Key tothe Costa Rican species of Anollg igo. ks ye 64
AMOS NNCTOUs COPE as. ceiy sia piers sy Men es 68
Avolig pentapiion:, Cone.at. citi ats oy kerk is ne Be btn fa
AMOlig MUlpUlesCOUs, (CONS yc tie ae ns eras ed ey 75
AMOUG. (iislomiscGOne sy ios tr es i gr 79
ANOUS DISGUE GEN Bi), NOVA: whe Gree Poa i era ia 81
AOS MITONGZ OOO es. kati fs ed pans ke doce § 84
Amolis-polvlepis# PSters.t ey i a ee 87
ATOMS INIA POLGrS oe 840, 7h ee yin ee ey een ioe 91
AMOS MINS Huts SUCNS. ir Hi eke ee nian et 91
AMOS MUMS MATSUDIAMS BUDS; MOVr 1 iy sis cre. 9
AMOlis-lemmuniiis LOMUnMUs GOW ty sy Vie ey a4 eh 100
AMOS AMSHINC CUS Peters se tw er ee ee a) 106
AMOS Opals HALLOW GU 6 Uae an ees ad Pires ney 110
Galli Bei npee eer aetae Maran | Moar, Mar Oran mn ere ore ger tee rer 113
(Ariolig altrec iin Beer pc ok hs eae oye 113
AMOS WOOG WOOdW IDUMs 4a os Phat ies cee eis 11
Amolis Wood! attentiatiis subsp, MOV,....6.0).v0.+..9. 2. 118
Anolis townsend Steewers 52.4 wee 123
Anolis capito. (Peters) nat co ee ten ee ee 126
Anolis; bipordatts Wiestnamn 3.7.0, .51 0) el ly 133
ANGUS TONOUs, GODS i ects ete rn ee 137
Anolis dquaticus si. MOV eit ee 141
Anolis DACHYPUS CONE 462) cieire eee eva ee 14!
Anolis tropidolepis boulenGets. 625 cs ee 146
Anolis: Gupte BOUlENGeL 65 ic ay Pee ee es 151
ANOS Aces BP; MOVE seiei Vel ed beni) dee ce tas 153
Atolls Bodmin WOWeNweI. s+, 4s ee 156
Gents: Poole CUVIEt yl rev Ce hte rr, 15
POW GUMS CIULINOSIIS) DELLHOLA nce baie etic o,f 15
Genus Cotvthoniines BOG: (4.5 ial ences eon ce hee 16
Corythophanes cristatus (Merrem) ...................5 161
Wells Beeuisclis sattrenvi.: jk cs eee 1¢
Key to species of Basilisous in Costa Rica......7............ 45 167
Basilicous. luimiimons Gopec7, 0h icts i et. 167
Basilisous -vittats Wega 6.4650. es oe 171
Basilisous Wasilisous (lites). 45 hs 00s es sc ear Nyy
Basilisaus. galerie 4, IDUMISHY ipsa ys cee ors, 177
Gents Ctenosaurus Wieemann , fos. scr, eh cena 179
Ctenosaurus similis similis Gray..................
Review or Lizarps or Costa Rica
Genus SCElOMOMIS Vie ANn soce naan vis ory aden er etc
Sceloporus vatiabilis olloporus Smith ...................
Sceloporus malachiticus malachiticus Cope..............
em AM GU bed scp net ester Yuu aa liocmlel warner: » aac bee als
Key to the Costa Rican genera of the Anguidae....................
Genus IiploglossuseWieginerty sei ne) ee
Key to Costa Rican species of Diploglossus................. se...
IDIDlOBIGSSUsTmOMOMOpIs GONUL) aes av ee ees hn
Diploglossus bilobatus O’Shaughnessy ..................
Genie: -GelestusoGravn nity. ran eee as oa te
Key to the Costa Rican species of Celestus.......0..04..0..00,
LOL cl juhl pepe AAG Oar Renews, Paliaeiee Corey tae reac ReMi 8 206 Sota eet Hie
Gelestus cyanochlons Gone sca evi ete ts cs as
GOMTS DESIG IRV ey ee ere ek ee | ees ee
IDRIISIE IMO MULOWIO, GONG. is i ei a ee cal
Pero NEMvUSICAGtan iy wine ee wee eee ci aieere ce kt
Genma epicOmuvinac wy WO MINOTH eee, la, eae A acs g
Key to the Costa Rican species of Lepidophyma.................
Lepidophyma reticulatum Taylot ....................
Repo whyine OOSOUTIn er OOUn 4
Lepidophyma ophiophthalmum Taylor .................
MOLLY ENO adh el Seles be git NEL arene. \ compa eee «tac meal 4.
Genus, Gyimnopithealmus,.Merrem: .. ee ics tes cs
Gymnophthalmus speciosus speciosus Hallowell ..........
Gems Lenosomd Spin: (city ee SAGE Rare ad
Key to forms of Leposoma in Gosta Rion, 6... 02.4... 6. ses
Iheposome, southis orientalis’ Tavlor |... 40... 3.
Enis ECV GUOGLOSSIIS: (BOULEN War ture iets te Cac teses aia)
REVOMOMLORSUIS: TUCHLUS WLBVIOL). 200, ck Oe
USGI ATCT EN Vie age ihe tread Grae oe iets a hou
WeVetG LOTS or Amecinedt Goste IGA. 44 rales ee a cies
Anadia. metallica metallica: Gope ......7) 01. ....4.2.0.,
Anadia metallica arborea Taylor... 6.4. ss sees esas
Anadia metallica attenuata Taylor. 6... .cc. ccc vas
Genus Amelva Guvier (Oe tat eee ya
Key to species of Ameiva in Costa Rica... 0... cee
AMGIWAMIODLODUEVS GONG, eaves cL lGun ah ying hoes
Ameiva festiva Lichtenstein and yon Martens............
Ameiva festiva occidentalis subsp. nov. .................
Ameiva undulata parva Barbour and Noble..............
Ameiva quadrilineata Hallowell 05.00 0......00......5.
Genus Cremidoplioris Wagler (te. seta le ie.
Ghemidoplioris aliaronis Gope ...48...7.........4....
Cnemidophorus deppei deppei Wiegmann ..............
Cnemidophorus lemniscatus lemniscatus Linnaeus
Tue Universitry SCIENCE BULLETIN
PAGE
Fanttly Sinead. aie ccs Verein fay ered |g Sinn Narn eneee vis fr. 282
Key to the genera of Costa Rican Scincidac.. ii... 283
Ibetoloptstma (Dummer aid ilbrOn a) %) tse ss 288
Key to Costa Rican subspecies of Leiolopisma cherriei........ .. 284
etolopisina cleric: cherie: Gope i 827s 284
Leiolopisma cherriei lampropholis subsp. nov. ........... 287
Gels “Himmeces Wierintita it. 2,59 i a et PIO et 291
Eumeces managuae Dunn .......... SoA ee ees 262
Genus Mabuya Fitzinger ......... PAS re anpsahey Lay! , 29D
Key to the species of Mabuya in Costa Rica........... Bri 297
Mabuva- Stachvirocdtis Sp, tov, 4). 1 he a08
Wabnve alinced One wis ese ee haa Ps hes 298
Mabuya unimarginata Cope ...... NES. Sil Re Aigo th es 301
BiplO stati eee ea ee ee phe Pee he eS Stes
INTRODUCTION
Of the fourteen families of lizards now recognized in the West-
ern Hemisphere eight are known in Costa Rica. These are the
Eublepharidae, Sphaerodactylidae, Anguidae, Teiidae, Gekkonidae,
Xantusiidae, Iguanidae, and Scincidae. These differ considerably
in the number of genera and species represented, varying from
three species (one genus) in the Xantusiidae to 36 forms and seven
genera in the Iguanidae. Some of the families are confined chiefly
to the lowlands, while cthers—Iguanidae and Anguidae—may have
representatives up to the tops of the highest mountains.
The Western Hemisphere families, absent in Costa Rica, are
the Anelytropsidae, Bipedidae, Aniellidae, Xenosauridae, and Helo-
dermatidae. The Amphisbaenidae, sometimes placed in the Sauria,
are absent. All occur in Mexico, but the number of representatives
have dwindled and they may be regarded as relicts. If they have
ever occurred in Central America or in the Eastern Hemisphere
they have become extinguished there.
The following families of the fourteeu mentioned above occur
outside of the Western Hemisphere: Eublepharidae, with repre-
sentatives in Asia and Africa; Anguidae, with legless forms in Asia,
Indonesia, Europe and Africa; Iguanidae with a few forms in
Madagascar, and the Fiji and Friendly Islands. The Gekkonidae
and Scincidae are widespread in the Eastern Hemisphere, and com-
prise numerous genera and species.
Judging by the present distribution of the Teiidae we presume
that this family has developed and evolved chiefly in South Amer-
Review or Lizarps oF Costa Rica 1
ica. A few of its numerous genera have reached Costa Rica while
two of its more progressive genera have reached northern Mexico.
One of these, Cnemidophorus, has become widespread in the
United States,
Distribution of Costa Rican Genera of the Sauria
i : Bastern
United sxicg | Guate- | Costa |p... | South api
States Mexico mala Rica Panama America Seen
».4 X oS Petey Es RA Rye
ae tet x x x eae
x X X Pe PO ee
x xX x Oe Eerie Sleds
X xX xX ees palin Hee
x x x x ».«
x x xX eee A Fea thd
aie xX x Be (Eas wae
x x Xx eet cdl hisses kee
x x x Barer w inca
X xX x BF liccstapne ca
x xX OE NG ara pee ky
x x x Pee Tulane ore
x MN eda neans | i eakberieliiauty e oa 4
x MSA dlc iseral wie Hohl eal ee eRdEliN gia cca eA
x x ae IR Wake Sal cd vey
».4 Xx x * Sa peg eee
elec Gyre AR pepe aie iaa Lara ieAleer Oi) qe toe Xx x x
PACE etre ac esea les Mee Guticad | dari eestev | uacnets ales vis aie xX Xx xX
I UY CHUBIOBSUB TV re cir rire iw ci Pvingey Fain eae X Xx x
Ban hick Re ee a x x xX xX xX
Chemidophorus a ee ee x x x x x x
BPLOLOMIBSEN . Gusyisciti once Baas acs xX xX X xX x haere
UNMGOGE ak ocr tere ec x X PG ae gee ees ek
BOUVET edifices tech clk yet xX X x
That South American genera other than the ones listed may be
represented in Costa Rica is a strong probability. This is es-
pecially true for certain Teiid and not impossibly Iguanid groups.
Echinosaura and Scolecosaurus, Teiid genera, are known to occur
in Panama.
The most striking feature of the lizard fauna is the remarkable
Series of species and subspecies that has developed in the genus
Anolis, These vary from small inconspicuous terrestrial forms, t¢
large showy arboreal creatures. Because of the large number of
Species inhabiting Costa Rica, one is convinced that the genus as
now generally regarded is a composite, one actually composed of
Several genera. In certain cases these species-group differences
have prompted workers to propose generic or subgeneric names for
them—i. e. Draconura, Coccoéssus, Gastrotropis, etc. While even-
tually this great unwieldly composite genus must be broken up,
one must examine in some considerable detail the entire series of
Teal and nominal species before it can be attempted. It would be
Preferable that the task be done by other than an amateur worker—
one who can give the problem adequate study in the field.
S
Tur Universtry SCIENCE BULLETIN
As in the case with other herpetological groups, the data avail-
able on Costa Rican Sauria are scattered in the literature. There
is some confusion owing to the fact that a single species may have
been described by two or more persons either due to inadequate
literature or inaccessibility to the literature. Our knowledge of the
fauna has been of slow growth. Often the species has been de-
scribed from a place outside of Costa Rica and the discovery in
the latter country may have happened at a much later date.
I have previously reported on certain parts of the collections from
Costa Rica (see bibliography). However, only two of these (Tay-
lor 1949 and 1955) deal with lizards.
HISTORY OF COLLECTIONS AND LITERATURE
Of the eighty species and subspecies of Sauria treated in this
work, Cope, as early as 1871, listed eight species collected in Costa
Rica. Anolis insignis, Anolis macrotus and Anolis trochilus [=
Anolis limifrons Cope] were described as new. Besides these the
following names also appeared: Anolis nannodes var. Cope, Anolis
hoffmani Peters, Sceloporus malachiticus Cope, Cyclura acanthura
Wiegmann, and Phyllodactylus (not identified as to species). This
collection, obtained from Dr. Van Patten, was derived from the
country in the neighborhood of the city of San José, capital of the
Country, several specimens perhaps coming from the southern and
western slopes of Volcan Irazi.
A second and more considerable collection sent to the United
States was made by William Gabb—‘“under the auspices of the gov-
ernment of Costa Rica, was obtained in the southern portion of the
region of Costa Rica which lies east of the elevated mountain range,
which traverses that country, and at different elevations on the range
itself.”
Certain of the specimens were obtained on Pico Blanco, a peak
rising to an elevation of (circa) 11,800 feet. Cope states that most
of the lizards were taken at Sipurio, a point on Rio Sixaola about
eighteen air miles from the nearest coast, close to the Panama
border, and having an elevation of some 200 feet.
In this paper (Cope 1875) the following species were described
as new: Anolis pachypus, Anolis oxylophus [ = Anolis lionotus
Cope], Basiliscus plumifrons, Mabuia alliacea [| = Mabuya alliacea
Cope], Chalcidolepis metallicus new genus, new species | = Anadia
metallica metallica Cope]; Amiva gabbiana | = Ameiva quadri-
lineata Hallowell]. Aside from the new forms, the following were
listed, those appearing in the previous list are preceded by an
Review or Lizarps or Costa Rica 9
asterisk: Coleonyx elegans Gray [ = Coleonyx mitratus Peters],
*Phyllodactylus sp., Sphaerodactylus glaucus Cope [ = ?], Theca-
dactylus rapicaudus Houttuyn, *Anolis microtus Cope, *Anolis in-
signis Cope, *Anolis trochilus Cope [ = Anolis limifrons Copel],
Anolis intermedius Peters, Anolis capito Peters, Anolis tessellatus
O'Shaughnessy [ = Anolis intermedius Peters], Polychrus multi-
carinatus Peters | = Polychrus gutturosus Berthold], Iguana rhino-
lopha Wiegmann, Cyclura acanthura Wiegmann [ = Ctenosaurus
similis similis Gray], Basiliscus vittatus Wiegmann, Sceloporus ma-
lachiticus Cope | = Sceloporus malachiticus malachiticus Cope],
Diploglossus monotropis Kuhl, Gerrhonotus fulvus Bocourt [ =
Barisia monticola Cope], Amiva festiva Lichtenstein and von Mar-
tens [ = Ameiva festiva festiva Lichtenstein and von Martens],
Mocoa assata Cope | = Leiolopisma cherriei lampropholis subsp.
nov. |, Mabuia cepedei [ = ?].
Cope omitted two names, Anolis nannodes var. [ = Anolis inter-
medius Peters] and Anolis hoffmani [ = Anolis cupreus Hallowell],
appearing in his first list. Cope, in 1877, described the form listed
above as Gerrhonotus fulvus Bocourt, as Barisia monticola sp. nov.
Cope’s Catalogue of Batrachia and Reptiles of Central America
and Mexico appeared in 1887 and in it are listed thirty-one forms.
A few changes are made in the names.
Mabuia cepedei Gray appears in the synonymy of Mabuia agilis
Raddi =(?). Anolis nannodes Cope [ = Anolis intermedius Peters |
and Anolis hoffmannii [ = Anolis cupreus Hallowell] are restored
to the list, and two forms described by Boulenger in 1886 are added
—Anolis tropidolepis and Anolis godmani. Cope’s species Chalci-
dolepis metallicus is reported under Leposoma metallica Cope.
Celestus bilobatus O'Shaughnessy (in Bocourt) is listed, and Scelo-
porus squamosus, and Basiliscus americanus Laurenti [| = Basiliscus
basiliscus Linnaeus] also appear.
Thus seven names are added to the Costa Rican list, and one is
placed in synonymy making a net gain of six.
In 1893 Cope published on a collection made in southwestern
Costa Rica by Mr. George K. Cherrie. In this were described as
new, the following species: Mocoa cherriei [ = Leiolopisma cher-
nei cherriei Cope], Amiva leptophrys | = Ameiva leptophrys Cope],
and Anolis longipes [ = Anolis capito Peters].
The following also were listed: Gymnophthalmus laevicaudus
Cope [ = Gymnophthalmus speciosus speciosus Hallowell], Amiva
quadrilineatus [| = Ameiva quadrilineatus Hallowell], Amiva festiva
10 Tue Universtry SCIENCE BULLETIN
Lichtenstein and von Martens [ = Ameiva festiva occidentalis
subsp. nov.], Celestus Steindachneri [ = Celestus ?], Ctenosaurus
yg; Iguana tuberculata Linnaeus, Anolis intermedius Peters, Anolis
trochilus Cope [ = Anolis limifrons Cope], Basiliscus vittatus Wieg-
mann.
In 1894 Cnemidophorus amivoides [| = Ameiva undulata parva
Barbour and Noble], Cnemidophorus alfaronis and Celestus cyano-
chloris were described by Cope and in a posthumous paper (1899,
edited by J. Percy Moore), he described Anolis purpurescens from
Colombia.
Wilhelm Peters (1863) described two Costa Rican species col-
lected by Dr. C. Hoffman, Anolis hoffmannii and Anolis capito, and
two Panamanian species, Anolis intermedius and Anolis humilis, later
to be reported from our territory. Later the same year, he described
a specimen collected by Dr. A. von Frantzius as Brachydactylus mi-
tratus [ = Coleonyx mitratus (Peters)]. In 1869 he described
Polychrus (Chaunolaemus) multicarinatus [ = Polychrus gutturo-
sus Berthold]. In 1873 (or 1874) Peters described several Pana-
manian forms, among which were Anolis polylepis and Anolis obtusi-
rostris [ = Anolis biporcatus Wiegmann] later to be found in Costa
Rica. These names appeared in Cope’s papers.
Albert Giinther, 1885-1893, in the Biologia Centrali-Americana:
Reptilia and Batrachia, listed thirty forms most of which had ap-
peared in Cope’s papers. The following, however, had not been
listed: Ameiva surinamensis Laurenti [ = ?], Ameiva undulata | =
Ameiva undulata parva Barbour and Noble]; Anolis cupreus Hal-
lowell, and Anolis intermedius Peters. This work was largely based
on collections made in Central America by Salvin and Godman
chiefly in Mexico and Guatemala.
George Albert Boulenger, 1885-1886, described or reported from
Costa Rica, Anolis godmani* [ = ?] and Anolis tropidolepis. In
1893 Anolis curtus was described.
Emmet Reid Dunn added the following names to the known
fauna: Anolis humilis Peters, Anolis stigmosus Bocourt, Anolis altae
sp. nov., Anolis lionotus Cope, Anolis limifrons Cope, Anolis penta-
prion Cope. He also described Anolis woodi from Panama (here
reported for the first time from Costa Rica). These descriptions
were based on material collected by him for the Harvard Museum.
Charles Burt, 1931, reports Sceloporus variabilis [= Sceloporus
* Regardless as to whether the description of Anolis godmani in the Biologia Centrali-
Americana: Reptilia and Batrachia (dated October 1885) appeared before that in Bou-
lenger’s Catalogue (1885) is entirely immaterial since his placing of the Boulenger reference
(as yet unpublished) following the name, gives conclusive evidence of authorship.
Review or Lizarps or Cosra Rica dd
variabilis olloporus Smith] and Cnemidophorus deppei deppei and
in the same year shows the presence of Cnemidophorus lemniscatus
lemniscatus occurring in Costa Rica in his distributional map.
Burt has also reported Epaphelus sumichrasti (in synonymy ) from
Costa Rica.*
My studies on the Costa Rican fauna began in 1947 when, accom-
panied by my son, Richard Clark Taylor, I made my first Costa
Rican sojourn. This was followed by a second journey in 1951, a
third in 1952, and a fourth in 1954, the last three being undertaken
alone. The four expeditions allowed for a total of ten months in the
field during which I acquired, for the collections of the University
of Kansas, more than ten thousand specimens. These, together
with specimens collected by Richard Clark Taylor, have been avail-
able for my studies. A fair proportion of the collections are repre-
sented by Sauria. I have examined only a part of the Costa Rican
specimens in other collections of the United States, since much of
the critical material was not available to me at the time this work
was being prepared.
Very much work still needs to be accomplished before a com-
plete report on the Saurian fauna of this region will be written.
However, since no work purporting to cover this fauna has appeared
since Boulenger’s catalogues 1885-1886, a period of seventy years,
I offer this resumé as a contribution to our present knowledge. I am
aware of numerous forms that, with adequate material, will eventu-
ally be designated as subspecies. And doubtless there are others
that have not been seen by me. Others already described now
appearing in synonymy may be revived, and some now recognized
may be again relegated to the oblivion of synonymy. New species,
too, will surely be discovered.
T have published two papers that include descriptions of Costa
Rican saurians; one (Taylor, 1949) describes Alopoglossus plicatus
[= Ptychoglossus plicatus| and Leposoma bisecta [= Leposoma
southi southi Ruthven and Gaige]; a second (Taylor, 1955) de-
Scribes Anadia metallica attenuata, Anadia metallica arborea, Lepo-
soma southi orientalis, Lepidophyma reticulatum, Lepidophyma
anomalum |= Lepidophyma obscurum Barbour] and Lepidophyma
ophiophthalmum. Eumeces managuae Dunn is reported for the
first time in Costa Rica.
In the present work the following species are described as new:
* Phe 7 . ‘ *. : : ”
a This reference is in error, since Dr. Burt has seemingly considered the presence of the
scription of this species in Cope’s 1875 paper, in a footnote, as evidence of its occurrence
™ Costa Rica.
12 THe Universiry SCIENCE BULLETIN
Anolis humilis marsupialis, Anolis woodi attenuatus, Anolis biscuti-
ger, Anolis aquaticus, Anolis achilles, Ameiva festiva occidentalis,
Leiolopisma cherriei lampropholis, Mabuya brachypodus.
The following are presumably here reported from Costa Rica for
the first time: Gonatodes fuscus Hallowell, Sphaerodactylus homo-
lepis Cope, Sphaerodactylus millipunctatus Hallowell, Phyllodacty-
lus eduardofischeri Mertens, Anolis purpurescens Cope, Anolis poly-
lepis Peters, Anolis lemurinus lemurinus Cope, Anolis woodi woodi
Dunn, Basiliscus galeritus A, Duméril, Gymnophthalmus speciosus
speciosus Hallowell, Mabuya unimarginata Cope.
In this report I am including numerous figures to aid in the
identification of species, since many have not been figured, and of
those that have, many are in books that are not readily accessible.
Many of my figures are satisfactory save for color; others less so,
but even these I believe are better than no figure at all.
Costa Rica, together with much of western Panama, constitutes
a landmass that in former times may have formed an ancient island.
The low land of the Canal Zone and of the Nicaraguan border may
well have been below sea level, thus isolating it from landmasses
northward and eastward. The Cordillera Volcanica and the Cor-
dillera Central are separated by the relatively high Meseta Central
(about 1500 meters at its highest part) from the Talamanca Range
that extends into western Panama. The Meseta does not completely
separate the faunas in the two areas. Several Panamanian species,
as yet unknown in our territory, may be discovered eventually.
While many parts of the country have been visited and partially
explored, much work still must be accomplished, especially in the
southern mountains at elevations between 800 and 2,800 meters,
There are two other regions—the Peninsula de Nicoya, Western
Guanacaste, and the Peninsula de Oso, Puntarenas Province, that
have escaped exploration, These two areas may once have been
detached from the mainland and their faunas may have undergone
change as a result of isolation.”
TAXONOMY OF COSTA RICAN SAURIA
Kry TO THE FAMILIES OF SAURIA IN Costa Rica
1. Body and top of head covered with small granules and/or scales but
lacking typical symmetrical plates on top of head................ P)
Body covered with granules or scales but top of head covered with sym-
asteiuees shields or shina PaO ev eS ee Seek ee ee 4
* A discussion of the geogre hale of Costa Rica has appeare " in a previous paper, See
Taylor, Univ. Kansas Sci. Bull., vol. 34, pt. 2, 1951, pp. 12-16
Review or Lizarps oF Costa Rica 13
2. Vertebrae amphicoclous, “spectacle” covering eye, no fovea; postanal
sacs and bones present; femoral and preanal pores may be present
or absent (absent in endemic forms in Western Hemisphere),
Gekkonidae
Vertebrae procoelous; eye-covering variable....................0.. 8
3. True upper and lower eyelids present, without “spectacle”; parietal
bones united; pupil vertical; postanal sacs and bones present; typical
DIGATIAW POLES imeMiale, iki yer 9 sh cen Cees te, eeener ae Eublepharidae
“Spectacle” covering eyes, without true eyelids; postanal sacs and bones
absent; parietals paired; pupil rounded (in Costa Rican forms),
Sphaerodactylidae
4 No eyelids; éves petmmnently Open... 1... es i ic, 5
Eyelids present, movable; scales generally cycloid, with or without
keels, more or less similar above and below.................. 8
2. Inner finger absent; scales cycloid in about thirteen rows about middle
of body; nasal plates separated (Gymnophthalmus)........ Teiidae
Inner finger present; scales variable; three or four supraoculars...... . 6
6. Body covered on dorsum and sides with granules; venter with six to ten
longitudinal rows of enlarged scutes; a few enlarged scutes on arms
and legs (Ameiva, Cnemidophorus).............. 000005, Teiidae
Body not covered with granules, or if granular, intermixed with tuber-
cles and the enlarged supraoculars lacking...................... 7
7. Body covered with scales on dorsum, sides, and venter; supraoculars
present (Leposoma, Ptychoglossa, Anadia)............... Teiidae
Body tubercular or granular, intermixed with larger tubercles; no en-
larged supraoculars; the frontal divided mesially and occupying
supraocular area; lower labials and chinshields fused. . Xantusiidae
A lateral groove or fold with fine granules strongly differentiated from
enlarged: ventral scales (Barisia),...................5. Anguidae
ING Brooveron folg) along side Of boy. «i ie din ts. ew 9
9. Only one scale touches frontal anteriorly; two rows of supraoculars
(outer row may consist of only one or two scales) (Celestus, Diplo-
RIDERS Breen eit whet as eit ae OM) a skh Oe Cie anus Anguidae
Two or three scales touch frontal anteriorly, or frontal not distinguish-
able; with more than 15 scales on top of head................ 10
10. Fewer than 20 enlar ged scales on top of head; two or three scales er ar
frontal anteriorly; scales on dorsal, lateral and ventral surfaces
cycloid, subequal, in longitudinal rows................ Scincidae
More than 20 seales on top of head; scales variable, large imbricate
scales, or minute juxtaposed scales or granules on dorsum, sides and
VERT Sh ee ey ue eG epi ya ce ea Iguanidae
SUPERFAMILY GEKKONOIDEA
Until further researches are undertaken it seems wise to follow
the classification of the gekkonoid lizards offered by Garth Under-
Wood.,*
Following this arrangement Costa Rican forms will be arranged
as follows:
* Proc, Zool. Soc. London, vol. 124, pt. 8, Nov. 1954, pp. 469-492.
Tue Universiry ScreNceE BULLETIN
14
Family—Eublepharidae
Genus—Coleonyx Gray
Coleonyx mitratus Peters
Family—Sphaerodactylidae
Genus—Lepidoblepharis. Peracce
Lepidoblepharis xanthostigma Noble
Genus—Gonatodes Fitzinger
Gonatodes fuscus Hallowell
Sphaerodactylus Wagler
Sphaerodactylus pacificus Stejneger
Sphaerodactylus homolepis Cope
Sphaerodactylus lineolatus Lichtenstein and von Mar-
tens
Sphaerodactylus imbricatus Andersson
Sphaerodactylus millepunctatus Hallowell
Sphaerodactylus sp. (tail).
Family—Gekkonidae
Subfamily—Gekkoninae
Genus—Thecadactylus Cuvier
Thecadactylus rapicaudus Houttuyn
Genus—Phyllodactylus Gray
Phyllodactylus edwardofischeri Mertens
Genus
The presence in Costa Rica of a species of Aristelliger (subfamily
Diplodactylinae fide Underwood) is problematical. Boulenger
(Catalogue of the lizards in the British Museum, vol. 3, Addenda
to vol. 1) gives Nicaragua as representing a part of the range of
the genus. If this data is correct, this genus may extend into
Costa Rica. As yet I have learned of no specimen that has been
found in the latter country.
Imported species of the genus Hemidactylus, although occurring
both north and south of Costa Rica, seemingly have not entered
the country.
Key ro Faminies oF THE GEKKONOIDEA IN Costa Rica
1. Vertebrae amphicoclous; spectacle covering eye; no fovea; three or
four pairs of sternal ribs; postanal sacs and bones present; femoral
and/or preanal pores present or absent (absent in endemic forms in
the Western Hemisphere) (subfamily Gekkoninae)....GEKKONIDAE
Vertebrae procoelous; eye-covering variable ....................0 2
2. True upper and lower eyelids present without spectacle; parietals
united; pupil vertical; postanal sacs and bones present. Typical
Preanal pores tiNale Gt et 4 e EUBLEPHARIDAE
Spectacle covering eye; without true eyelids; postanal sacs and bones
absent; parietals paired; pupil rounded in Costa Rican forms.
SPHAERODACTYLIDAE
Review or Lizarps or Costa Rica 15
Famity EvsBLeEPHARIDAE Boulenger
Eublepharidae Boulenger, Ann. Mag. Nat. Hist. ser. 5, vol. 12, 1883, p. 308;
and ser, 5, vol. 14, 1884, p. 119.
Vertebrae procoelous; parietal single; body covered with small
scales or granules; eyes with vertically elliptic pupil and thick mov-
able upper and lower eyelids; males with typical preanal pores; digits
Straight, slender, lacking lamellae; postanal sacs and bones present.
Two eggs are laid. A voice mechanism present.
Boulenger writing of this group (1885 Catalogue, vol. I, p. 229)
States: “Three species are native of Central America, one of the
southern parts of the United States, two of Southern Asia, and one
of West Africa; the genus Eublepharis occurs in America as well as
in Asia. This extraordinary distribution seems to indicate that a
few representatives of this small family are the remnants of some
ancient, more generally dispersed group; it never-the-less remains
a matter of wonder how forms, now so widely separated, have re-
tained so great a resemblance, not only in structure, but also in the
pattern of coloration.”
While the concept of the genera and species belonging to the
group has changed somewhat, Boulenger’s comment is still worthy
of repetition.
Underwood recognizes the following genera as members of the
family Eublepharidae:
Aeluroscalabotes Giinther. Malaya, Borneo, Sumatra.
Eublepharis Gray. Southwest Asia, including Hainan and Riu
Kiu Islands.
Hemitheconyx Stejneger. W. Africa and Somaliland.
Holodactylus Boettger. Somaliland.
Coleonyx Gray. Southwestern U. S. to Panama.
Only the latter genus occurs in the Western Hemisphere. En-
demic American members of the Gekkonidae and of the Sphaero-
dactylidae seemingly have not developed true preanal and femoral
Pores.* True preanal pores are present in this family.
Genus CoLEonyx Gray
Coleonyx Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 16, 1845, p. 162 (type of
Senus Coleonyx elegans).
Description of genus: Body covered with granular or tubercular
Scales, intermingled with larger tubercles; digits slender short with-
Out widened lamellae; abdominal scales strongly imbricate, with a
Series of preanal pores in males; no femoral pores.
* oy ¢ a ‘ eo asad. .
once” Presence of Hemidactylus in the Western Hemisphere is, I believe, due to im-
aon in historic time by man,
16 Tue Universtry SCIENCE BULLETIN
This genus (as recognized in Klauber’s revision, 1945) ranges
from the southern part of Texas, west to California (including Utah
and Nevada), through northern and western Mexico, south into
Durango, Sinaloa and Baja California. After a considerable break
in Mexico in the known distribution, a southern group begins in
Central Veracruz and Colima, and following the lowlands, extends
south through Central America to Panama. I am not aware that
there is contact anywhere between the two groups.
These two groups are distinct, and seemingly the differences are
of such a character as to warrant a separation into two genera.
The name Coleonyx properly belongs to the southern group. It
consists of two species, Coleonyx elegans with two subspecies, and
Coleonyx mitratus with two forms mitratus and dovii. Only a single
form, Coleonyx mitratus Peters, is known to occur in Costa Rica.
Coleonyx mitratus (Peters )
(Fig. 1)
Brachydactylus mitratus Peters, Monatsb. Akad. Wiss. Berlin, 1863, pp. 41-44
(type locality, Costa Rica).
Coleonyx elegans Bocourt, part., Mission Scientifique au Mexique et dans
l'Amérique Central; Etudes sur les Reptiles, livr. 2, 1873, p. 49 (includes
mitratus in synonymy); Cope, Journ. Acad. Nat. Sci. Philadelphia, 2nd ser.,
vol. 8, 1876 (1875) p. 118; Proc. Amer. Philos. Soc., June 20, 1879, p. 271;
Wettstein, Sitzungsb. Akad. Wiss. Wien, Math-natur. Klasse, Abt. 1, Bd.
143, Heft 1, 2, 1934, pp. 19-20 (Bebedero; Central Plateau in the vicinity
of San José).
Eublepharis dovii Boulenger, Catalogue of the lizards in the British Museum
(Natural History), vol. 1, 1885 (type locality Panamé); Giinther, Biologia
ee ree Batrachia and Reptilia, 1890, Feb., pp. 83-84, pl. 31,
g. 18.
Coleonyx dovii Stejneger, North Amer. Fauna, no, 7, 1893, p. 163.
Coleonyx mitratus Schmidt, Field, Mus. Nat. Hist. Zool. Ser., vol. 12, 1928,
no. 16, p. 194; Klauber, San Diego Soc. Nat. Hist., vol. 10, no. 11, Mar. 9,
1945, pp. 199-203, map; Mertens, Abh. Senckenb. naturf. Ges. No, 487,
1952, pp. 37-38, pl. 11, fig. 63 (El Salvador).
Diagnosis: Rostral bordering the lip for a greater distance than
mental, the first infralabial in contact with mental; femoral pores
present, usually 5-9, 7 or 8 being most frequent; scalerows on venter,
27-29; nostril wholly in nasal scale, not bordered by the internasal;
tips of claws strongly visible.
Description (From KUMNH No. 34180 Tenorio, Las Canas,
Guanacaste, Costa Rica): Rostral large, its width (2.2 mm.)
greater than its height (1.7 mm.), occupying a longer border on
lip than mental; the first lower labial not in contact with rostral
when mouth is closed; two internasals border rostral, their narrow-
est width is only slightly more than half their greatest length; nostril
included in a single nasal; nasal followed by two small and four
Review or Lizarps or Costa Rica 17
minute scales; nine scales along the posterior borders of internasals:
about ten irregular scalerows between nostril and eye; scales on
snout larger, somewhat hexagonal with a few smaller irregular
Ficure 1. Coleonyx mitratus Peters (juvenile), KUMNH
34180 (young); Tenorio, Las Cafias, Guanacaste, C, R.; ac-
tual total length, 96 mm.
No,
Scales; between the orbits the larger scales are reduced in number
and there
are more numerous smaller ones; some granules between
Scales:
between the edges of eyelids at middle of orbit about 55
SCales: ea. . P . . .
cales; scales along outer edges of eyelids a little enlarged; eyelids
2—3378
18 Tue Universrry ScieENCE BULLETIN
distinct, functional, the upper separated from the interorbital area
by two very minute scalerows in which some scales are absent; 22
scales in a row across snout between the second labials; ear-opening
large, nearly twice as long as wide normally; 7-7 supralabials, last
very small; 6-6 infralabials; mental rather rounded posteriorly, fol-
lowed by four scales, one separated from the others because of
minute intercalated granules.
Dorsal squamation granular, interspersed with tubercles similar
to those between orbits, but tubercles are a little more elevated and
a little larger, arranged in about 24 irregular rows across back, and
from 22-26 in a longitudinal line from vent to axillary level; scales
on venter larger, imbricate, arranged in 24-26 rows, all larger than
scales on chin and neck; eight preanal scales bearing pores; edge
of vent without enlarged scales; openings of the postanal sacs dis-
tinct; a single large anal tubercle on each side of vent; tail segmen-
tation indicated by whorls of eight tubercles, usually distinct at
base, but less distinct distally where the tubercles are reduced in
number; each segment with six rows of scales (rarely part of an-
other present); on ventral part of tail larger scales alternate with
one or two pairs of smaller scales.
Arms and legs relatively short, the tip of longest toe not reaching
elbow of adpressed arm; fingers short, covered below with thick-
ened, widened scales, 12 under fourth fingers; tips with a sheath
formed of three scales that fail to cover completely the retracted
claw, leaving much of claw exposed; 18 scales under fourth toe;
arm covered largely with flat imbricate scales without distinct
tubercles; legs with flat imbricate scales except on dorsal side of
tibia and dorsal and posterior face of thigh.
Color in life: Brownish to brownish black above, with the curved
and transverse bands on body whitish yellow dorsally, becoming
cream laterally; lips spotted; chin and throat cream; flesh-white on
venter and on light areas under tail; laterally the lighter ventral
color encroaches on the black blotches; dark mark on top of head
with some light brown areas evident.
Measurements in mm.: Snout to vent, 50; tail, 46; axilla to groin,
24.3; snout to foreleg, 22; snout to ear, 12; head width, 9; head
length, 15.
Remarks: A second specimen, obtained at San Isidro del General,
is larger. The brown markings of the head are much lighter and
the lateral encroachment of the light color on the dark blotches
has proceeded farther.
Review or Lizarps or Costa Rica 19
Records of the species for Costa Rica are as follows: San José,
Province: San Isidro del General, Tableland near San José, San
José. Guanacaste Province: Bebedero, an island in the Gulf of
Nicoya, and Tenorio. Cartago Province: Turrialba.
Most of the localities for the species are in the Pacific drainage
area. The exception is Klauber’s (1945) record for Turrialba. In
my collecting at Turrialba, and in the Turrialba area, I have not suc-
ceeded in finding it.
Don Jose Zeledén reported to Cope that the species was to be
found in ant-hills near San José. If this is a natural habitat, it may
account for the apparent rarity,
It is called perro sonpopo and is regarded as dangerous. Needless
to say it is a completely harmless animal. The specimen described
was found by John Baker under a small board in a forested area.
FamMiIty SPHAERODACTYLIDAE Underwood
Sphaerodactylidae Underwood, Proc, Zool. Soc. London, vol. 124, pt. 8, Novem-
ber 1954, p. 476,
The recent proposal of Underwood to distinguish a group of small
saurian genera under the above name is followed here. His defini-
tion of the family is:
“Spectacle covering eye; second branchial arch nearly complete;
interruption between base—and epibranchial; vertebrae procoelous;
clavical perforated or not. Four or five pairs of sternal ribs; digital
pads, if present single, terminal; temporal fovea. Postanal sacs
and bones absent; no femoral or preanal pores; parietals paired;
Pupil round, elliptical or straight vertical.”
To the above characters may be added: no voice mechanism
Present; a single egg produced; in Costa Rican genera, males with
a larger or smaller series of differentiated glandular scales in preanal
ventral region and often with similar series on femora, rarely in
Postanal region; no eyelids, but extrabrillar fringe continuous, sur-
Tounding eye, becoming more extensive over the anterior part of eye.
The Costa Rican genera belonging to this family are Gonatodes,
Lepidoblepharis and Sphaerodactylus. Two other genera, Pseudo-
Sonatodes * and Coleodactylus,* recognized as members of the fam-
ily are South American.
All specimens of the Costa Rican genera examined have a rounded
Pupil, and the males are equipped with specialized glandular scales
Probably subserving the same function as femoral and preanal pores
and not impossibly the ancestral type of such pores,
* ON
lhe Presence of glandular scales have not been demonstrated in these genera,
Tue University SCiENCE BULLETIN
Key to rue Costa Rican GENERA OF THE SPHAERODACTYLIDAE
1. Claws not retractile into a sheath; distal portion of digit compressed
and forming an angle with the more proximal parts ... .Gonatodes
bo
Claws tettactilednto.a sheathns 300. a. epi et ay
2. Each digit terminating in an asymmetrical lamella,* at the proximal
side of which the claw is retractile into a sheath ..Sphaerodactylus
Rach digit terminating in a symmetrical sheath composed of six scales
into which the claw is withdrawn............... Lepidoblepharis
Genus LepmosLePHaris Peracce
Lepidoblepharis Peracca, Boll. Mus. Torino, vol. 12, no. 300, 1897, p. 1 (type,
by monotypy, Lepidoblepharis festae Peracca ).
Lathrogecko Ruthven, Occ. Papers Mus. Zool. Univ. Michigan, no. 21, 1916,
p. L (type, Lepidoblepharis sanctae-martae Ruthven).
Description of genus: “Digits short or moderate, cylindrical, the
distal joints forming an angle with the basal portion, with smooth
transverse lamellae inferiorly; claw retractile vertically into a large
compressed sheath composed of six scales—a pair of elongate infero-
laterals in contact inferiorly, a pair of elongate supero-laterals, an
elongate median dorsal separating the latter, and a small terminal
lying in the angle between the tips of the supero-laterals. Head
and neck covered with small granular scales; body depressed, tail
round; ventral and inferior caudal scales smooth, imbricate; dor-
sals granular, tubercular or imbricate. Pupil round; eyelid well
developed above. Rostral shield large, with a U-shaped impres-
sion on its upper surface. Clavicle moderately dilated, not per-
forated. Male without femoral pores.” Parker, 1926, redefinition.
To the above characters must be added: males with a series of
specialized “glandular” preanal and (frequently) femoral scales.
Vertebrae procoelous. One egg produced.
This genus of small gekkoes ranges from southern Central Amer-
ica into northwestern South America; at least seven species are cur-
rently recognized, only one of which has been reported from Costa
Rica. This is a small terrestrial gekko living for the most part under
rotting logs or trash, usually in moist situations.
Lepidoblepharis xanthostigma Noble
(Figs. 2,3)
Lathrogecko xanthostigma Noble, Proc. Biol. Soc. Washington, vol. 29, April 4,
1916, pp. 87-88 (type locality, Zent, near Puerto Limon, Costa Rica, H. S.
Blair collector); Amer. Mus, Novitates, no. 4, Mar. 10, 1921, pp. 3, 6, fig. 3
(skeletal characters ).
Lepidoblepharis xanthostigma Dunn, Occ. Papers Boston Soc. Nat. Hist., vol. 8,
June 7, 1933, p. 75 (El Valle de Anton, Panama’); Proc. Acad. Nat. Sci.
Philadelphia, vol. 92, 1940, p. 112 (Costa Rica: Suretka, Monteverde,
Guiapiles, Carillo, “La Castilla farm on the lower Reventagén sic (= Revent-
* This scale is described in the literature as spherical. At least this is usually untrue,
Review or Lrzarps oF Costa Rica vA
azon). Panama: “Cerro Campana”, about 3,000 ft., El Valle); Parker,
Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, pp. 291-301 (relationship);
Underwood, Proc. Zool. Soc. London, vol. 124, pt. 3, Nov. 1954, p. 476
(member new family Sphaerodactylidae ).
Diagnosis: Diminutive gekkoes, probably not exceeding 38 mm.
in snout-vent length; rostral dorsally flattened with an inverted
U-shaped ridge; toes symmetrical, the claws retractile into sheaths
composed of six scales; “escutcheon” consisting of numerous glandu-
lar scales; femora with or without some similar scales; fringe of eye
“spectacle” forms a shelflike projection bordered by much-enlarged
scales,
Description of species (From KUMNH No. 38926, Limén ): Head
a pointed oval, the eyes moderately prominent; rostral large, flat-
tened on top, the flat area bordered by an inverted slightly elevated
U-shaped ridge that extends onto the internasals, the flattened por-
tion cut by a suture, no part of scale extending posterior to middle
of nostril; internasals enlarged, wider than long, separated by two
scales, one behind the other; nostril surrounded by rostral, inter-
nasal and one or two postnasals border rim; snout without a canthus
rostralis, rounded transversely, covered with juxtaposed flat scales
diminishing in size from the internasals to between eyes; first supra-
labial very large, more than twice as long as high, more than twice
the size of second, highest near the middle of scale; this followed
by four differentiated supralabials, rapidly diminishing in size; re-
Mainder of mouth-border edged with minute granular scales; rostral
large, flattened posteriorly with a labial border considerably greater
than that of mental; first infralabial only a little larger than second,
Which in turn is three to four times size of third; latter followed by
three other small infralabials diminishing in size; mental bordered
behind by five scales, the three median largest; following this row
the scales diminish rapidly in size and the chin and anterior part
of throat are covered with small nearly equal-sized juxtaposed flat
Sranules,
Eye surrounded by a “spectacle” and fringe which on upper an-
terior three fourths forms a shelflike projection, bordered by en-
larged scales; posteriorly these are flattened pointed scales, while
anteriorly there are three enlarged ciliary scales, the very large
median scale double the size of the others and as wide but not
quite so long as second supralabial; below this row of scales is a
horizontal row of subciliary scales also bordering the edge of “spec-
tacle”; on lower edge of orbit, touching the supralabials but lying
Mostly within the orbit, is a series of five differentiated scales, dark
Fic. 2. Lepidoblepharis xanthostigma Noble (left to right). No. 1, KUMNH 34641; snout-vent length,
37 mm., tail, 41 mm.; Isla Bonita, Heredia Prov. No. 2, 38937; snout-vent length, 34 mm., tail regenerated;
Palmar, Puntarenas Prov. No. 3, 34626; snout-vent length, 32 mm., tail, 37 mm.; Bataan, Limon Proy. No. 4,
34653: snout-vent length, 31 mm., tail, 36 mm.; Los Diamantes, Guapiles, Limon Prov. No. 5, 34651; snout-
a length 31 mm., tail, 39 mm.; Los Diamantes, Guapiles, Limon Prov. No. 6, 34643; snout-vent length,
ASB xem... cath. 39 mr - ~ Turriallba, Cartage Prov.
TH [,
a
AIG KLISUAAIN ()
HONG
NILATIAG
Review or Lizarps or Costa Rica 93
in color; about 23 scales across snout between the posterior edges
of the first supralabials. Ear opening a third larger than an inter-
nasal scale.
On back the scales are small, elevated, juxtaposed, the granules
growing a little larger and somewhat more conical on sides; these
are abruptly differentiated from the large imbricating scales of the
venter; granules on occiput scarcely half as large as those on dor-
Fic. 3. Lepidoblepharis xanthostigma Noble. — Pat-
tern of escutcheon scales of venter and femora (scales
without pigment); much enlarged.
‘um; ventral scales in about 20 longitudinal rows; scales on posterior
Part of neck and breast similar in character to other ventral scales
but somewhat smaller. Posteriorly on venter, scales are somewhat
larger than elsewhere; dorsal and anterior part of upper arm, an-
terodorsal region and part of underside of forearm, front and ven-
tral part of thigh, and underside of tibia with enlarged imbricating
scales, while remaining parts generally are covered with small scales
Or granules,
24 Tue University ScieNCE BULLETIN
Dorsal and lateral surfaces of tail covered with enlarged imbri-
cating scales, more rounded than those on venter; on ventral sur-
face, beginning somewhat behind the vent, a median series of about
50 enlarged subcaudals, the scales alternating larger and smaller;
tail rounded in cross-section.
Digits of hand and foot with their distal part forming an angle
with the basal part, all with claws vertically retractile into a sheath
composed of six scales; under surface of digits with widened flat
imbricating scales, one or two at base and terminal scales usually
divided; scales on palm and sole small, flat, juxtaposed.
On posterior ventral surface an escutcheon consisting of a group
of about 100 modified “glandular” scales, distinguished from adjoin-
ing scales by an almost complete absence of surface pigment ex-
cept at extreme edges of individual scales, the pigment forming <
slightly wider border where these scales adjoin the normal scales.
On distal femoral area, three scales show a partially or completely
similar modification; none are present in the postanal area.
Color in life: Above dark brownish lavender with very indistinct
lighter areas; head somewhat less dark, with an indefinite light band
partly divided mesially by a dark line, and bordered fore and aft
by darker bands; top of head lighter than body, the snout and supra-
labials rather brownish; a very indefinite light line behind eye,
bordered by darker color above and below; infralabials dark blackish
brown with cream spots; chin and throat yellowish with brown
dots, the lines or reticulations extending back to near breast; venter,
except for glandular scales, nearly uniform light lavender; tail
slightly lighter than body, dorsally and laterally with indefinite
lighter markings; under surface lavender; glandular scales paper
white; limbs similar to body in color, somewhat lighter beneath;
back surface of thigh dark, enclosing dull cream spots.
Measurements in mm.: Snout to vent, 31; tail, 35.5; total length,
66.5; arm, 8.9; leg, 12.2; width of head, 4.8; length of head, 9.
Variation: The Costa Rican material available consists of 95
specimens, the following from the Caribbean drainage area: Limon
Province: KUMNH Nos. 38926-38934, Limén; 38935, Mountain Cow
Creek, near Limén; 34617-34631, Bataan; 34656, 34655, 38925, La
Lola; 38000-38013, 38923-38924, Suretka; 34609-34614, 34645-34655,
34664, 38944-38947, Los Diamantes, Giapiles. Heredia Province:
Nos. 36272-36273, Puerto Viejo; 34661, Cariblanco; 34641-34642,
Cinchona, 4,500 feet.
From the Pacific drainage area: Guanacaste Province: Nos.
Review oF Lizarps or Costa Rica 25
38936, 34666-34668, Tenorio, Las Camas; 36261-36371, Tilardn.
Puntarenas Province: Nos. 34669, 38937-38943, Palmar; 34634,
Golfito.
In the series of 95 specimens now in the collection there is con-
siderable variation, concerned with sexual dimorphism, color pat-
tern, scale characters, size, and age. A group of specimens from a
locality may exhibit almost as much variation in certain characters
as specimens from the Pacific lowlands differ from those in the
Caribbean area. The group of specialized scales in the abdominal
region of the male and on the distal part of the femur are wanting
in the female, and the surface pigmentation of the scales is un-
differentiated in the area.
The ventral scales of females are perhaps a little more rounded
posteriorly, and since the skin may be distended a little more, the
scales of the longitudinal rows seem actually larger having wider
visible contact with the following and preceding scales. The num-
ber of enlarged ventral rows varies between 16 and 20, those from
the Caribbean drainage having nearly the same average as those
from the Pacific drainage. In snout-vent length the largest speci-
men measured 38 mm. This was from the northernmost point
where the species was found in the Caribbean drainage. Specimens
measuring 37 mm. were taken at Tenorio, the northernmost locality
in the Pacific drainage, and at Cinchona, the point of highest ele-
vation where specimens were found.
The specialized scales on the venter show decided variation in
the number involved. Thus, in the escutcheon series from Limon
specimens, I count from 60 to 110 “glandular” scales in adult and
old specimens; but only 28 to 46 in a juvenile and a young adult.
The differentiation of the scales, as to absence of surface pigment,
begins posteriorly. It is probable that other scales are actually
already differentiated, but these will not assume their typical ap-
pearance until later. Even in some of the older specimens, scales
adjoining the differentiated scales may show areas in which no
Surface pigment is present. There usually are three modified
glandular scales on the distal part of the femur but these may be
indistinct or absent; on the other hand there may be two or three
in a second row making a total of five or six of the specialized
glandular scales,
The shape of the mental varies in that the median rim may be
sharply truncate or may be irregular followed usually by three,
Sometimes two or very rarely as many as five scales. In some col-
lections the median of the three scales is largest.
26 Tue Universiry ScrencE BULLETIN
The spectacle and fringe are difficult to describe. However, they
are continuous and closely applied about the eyeball.
The two large horizontal ciliary and subciliary scalerows border-
ing the shelflike expansion (one row above, the other below) vary
and the posterior pointed scales may be blunt. One of the scales
is usually nearly as wide as the shelflike expansion. The relative
sizes of the labials do not remain completely constant in the series.
In two specimens the internasals are broadly in contact along the
median line; in other specimens the scales are smaller and may be
separated by a single large scale or by as many as three tiny ones.
On the sides of the base of the tail there is a distinct shallow de-
pression apparent in practically all of the specimens.
The scales on the tail are not arranged in transverse rows but
rather in diagonal rows. Only two or three specimens of the 95
have a complete tail, the others being wholly or partially regener-
ated. The subcaudals are enlarged, a larger wider scale alternating
with one somewhat smaller and narrower, the two together marking
a caudal vertebra.
The character of the ear-opening in the preserved material is
somewhat deceptive and may be represented by a line or may be
oval or nearly circular at times.
The somewhat curved lighter band across the occiput is usually
discernible. This may be bordered anteriorly and posteriorly by a
narrow darker band, one of which may have a median dark process
partially dividing the light band. The infralabials and mental bear
large dark spots surrounded by cream or yellow color. The mark-
ings on the chin are usually present as spots or lines, the latter
evident especially in the area below and somewhat posterior to
eye. In nearly every group of specimens there is one or more that
have distinct dorsolateral cream lines that originate behind the
eyes and join together on the base of the tail, the junction usually
preceded by a dark area. These are usually males, but not neces-
sarily so. One specimen from Turrialba is light with dorsolateral
lines and a clearly differentiated pattern of dark median transverse
spots, about ten in number. The dorsolateral cream line is bordered
below by a discrete dark line continuing on the tail as a broken line.
The lighter sides have numerous dark specks while the chin and
throat have a series of diagonal lines. The posterior surface of the
thigh is cream with a black median stripe on one side and irregular
spots on the other. The venter and subcaudal regions are flecked
with darker.
Review oF Lizarps oF Costa Rica 27
Specimens from Palmar (a few meters above sea level) in south-
western Costa Rica have the occipital cream band more distinct
and the sides of the body have a series of small ocellated cream
spots running along the side of the body and tail.
It is not improbable, that with material from more northern and
southern areas of the range of the species that it will be possible
to differentiate subspecific populations.
Genus Gonatopes Fitzinger
Gonatodes Fitzinger, Systema Reptilium, 1848, pp. 18, 91 (type of genus,
Gymnodactylus albogularis Duméril aad Bibron),
Description of genus: Digits more or less cylindrical, the distal
portion compressed forming an angle with more proximal parts;
the lower surface of all digits with a single row of plates or scales;
body slightly depressed, granular above; tail not compressed; pupil
circular. Males without typical femoral or preanal pores, but with
a large series of special glandular scales on posterior part of venter
and sometimes on ventral surface of femora and behind vent on
base of tail,
This genus formerly included numerous Asiatic and African
Species now recognized under the genus Cnemaspis. Seemingly
there is but a single species represented in Costa Rica,—Gonatodes
fuscus Hallowell. It occurs along the entire eastern coast, and per-
haps also on the west coast. It was a common species at and near
the port of Puntarenas. It has also been reported from Bebedero,
Guanacaste Province. I have taken it at Liberia in the same prov-
ince.
Gonatodes fuscus Hallowell
(Fig. 4)
Stenodactylus fuscus Hallowell, Journ. Acad. Nat. Sci. Philadelphia, ser, 2,
vol, 3, 1955, p. 33 (type locality, by restriction of Smith and Taylor, Rama,
Nicaragua); Cope, Proc. Acad, Nat. Sci. Philadelphia, 1874, p. 68 (Nic-
Girenedechilie fuscus A. Duméril, Arch. Mus. vol. 8, 1856, p. 477; Bocourt,
lission Scientifique au Mexique et dans l’Amérique Centrale; Etudes sur
les reptiles, livr, 2, 1873, pp. 48, 49, pl. 10, fig. 5 a-c.
Goniodactylus braconnieri O'Shaughnessy, Ann. Mag. Nat. Hist. ser. 4, vol.
16, 1875, p. 256 (fide Boulenger; type locality, Baranquilla, Colombia),
Gonatodes albogularis fuscus Boulenger, Catalogue of the lizards in the British
Museum, 2nd Ed., vol, 1, 1885, pp. 59-60 (Colombia; Panamé); Wettstein,
Sitzungsb. Akad. Wiss. Wien, Mathem.-natur. Kl. Abt. 1, Bd. 148, Heft,
1 and 2, 1934, p. 19 (Bebedero, Costa Rica).
Gonatodes albogularis Giinther (part.) Biologia Centrali-Americana; Reptilia
, and Batrachia, Feb, 1890, p. 80 (Nicaragua and Panami).
Gonatodes fuscus Stejneger, Proc, U. S. Nat. Mus., vol. 53, L917, pp. 264-265;
Parker, Ann. Mag. Nat. Hist., ser. 9, vol. 17, May 1926, p. 549 (Gorgona
Island, Colombia); Gaige, Hartweg, and Stuart, Occ, Papers Mus. Zool.
Tue Universiry SCrENCE BULLETIN
Univ. Michigan, no. 357, 1987, p. 8 (Nicaraguan localities); H. Smith,
Copeia, 1944, p. 187; Evans, Copeia, 1947, no. 3, Sept. 12, p. 168 (Corozal,
Canal Zone). Smith and Taylor U. S. Nat. Mus. Bull. 199, 1950 pp. 45-
46 (Chiapas, Mexico).
Diagnosis: Basal phalanges of digits cylindrical; strong sexual
dimorphism, the males much darker than females. Snout obtusely
pointed a little longer than the diameter of orbit. Gular region of
male orange or yellowish. A median whitish line bifurcating on
chin.
Description of species (From KUMNH No. 34577; Limon, Costa
Rica): Head short, the snout length a little greater than the diam-
eter of the orbit (skeletal), not or slightly greater than the greatest
depth of the head in the occipital region; rostral scute large, the
part visible above somewhat flattened medially, with a median
suture (not reaching down to lip); nostril surrounded by rostral,
first labial, supranasal, and (on the posterior rim) three postnasals;
supranasals large, separated by a depression occupied by a single
small scale anteriorly and two small scales posteriorly; six supra-
labials, first large, nearly twice area of second; second and third
of nearly same size; fourth, fifth, and sixth diminishing in size, the
suture between last two lying below the middle of pupil; posterior
part of mouth bordered by eight scarcely differentiated scales; a
spectacle, with an extrabrillar fringe surrounding eye, widest an-
teriorly; anterodorsally there is a ply or fold in the fringe; outer
anterodorsal edge of fringe with a series of enlarged somewhat com-
pressed conical scales and anteriorly these are preceded by other
somewhat enlarged flattened scales.
Snout covered with tubercular or pyramidal scales, often some-
what compressed, larger than other dorsal scales; on back about 22
in a distance equal to that between middle of eye and nostril and
26 across snout between the second infralabials; mental very large,
its labial border equal to that of rostral and four fifths of that of
first labials; first infralabials separated by a distance about equal
to width of one labial; two enlarged postmentals; sach first infra-
labial bordered by three chin scales, one of which is the post-
mental; five differentiated infralabials, the first double the size of
second; behind these, others, diminishing in size, angle of mouth
bordered by about five small scales; two or three somewhat en-
larged rows of scales on chin following postmentals; other scales
on chin and throat granular, juxtaposed, the transition to the
rounded imbricate scales of venter, abrupt. Scales on back, conical
tubercles, slightly larger on sides; on tail, dorsal and lateral scales
Fic. 4. Gonatodes fuscus Hallowell. From left to right, KUMNH No. 345652, Maribella Hotel,
mouth of Rio Barranca, Puntarenas Province; actual snout-to-vent measurement, 43 mm., tail, 40 mm.;
346082, La Lola, Limén Province; snout to vent, 33 mm., tail, 41 mm.; 345862, Limon, Limon Provy-
ince; snout to vent, 42 mm.; 345804, Limdn, Limon Province, snout to vent, 42.8 mm., tail 51 mm.;
345542, Maribella Hotel, Puntarenas Province.
YY VSO) AO SGUVZITT AO AVATAR
Vv
30 Tue Universrry SCIENCE BULLETIN
flat, subimbricate, forming nearly straight transverse rows, smaller
dorsally, growing larger on sides; the median subcaudal row about
three times width of adjoining scales; the posterior two thirds of
tail regenerated; in this part, dorsal scales are smaller than those
on original part of tail, the rows not normally transverse, the median |
subcaudals shorter but considerably wider than original subcaudals;
scales under thighs and tibia similar to ventral scales; on front of
thigh a group of still larger scales.
Limbs well developed, the toes of adpressed limbs reaching to
axilla; digits all with claws; basal part of toes slightly widened,
covered with enlarged scales; distally digits somewhat compressed,
under fourth toe eight larger basal scales and 15 scales on distal
joints. On anterior face of upper arm and anterior face of forearm
enlarged imbricating scales; on front of thigh and underside of
tibia, scales likewise enlarged and imbricating; a slight axillary |
depression filled with minute, granular scales.
No typical femoral or preanal pores, but lying anterior to vent
and extending forward more than one third length of the body, a
group of about 200 specialized “glandular” escutcheon scales; on
underside of each femur a group of about fifty similar scales ar-
ranged in five rows; on base of tail certain similar scales including
one or two of the enlarged median subcaudals.
Color in life: Head generally orange brown to orange, with black
spots below eye and mouth angle; body bluish black; tail on top
and sides similar except tip which is dirty flesh-white. Under sur-
face somewhat magenta. Chin and throat orange-red with a median
yellow dividing line anteriorly, and with traces of darker and lighter
olive-yellow parallel lines; belly blackish with the specialized
escutcheon scales dirty flesh; sides of neck somewhat olive.
In preservative (formaldehyde and alcohol), head grayish brown,
slightly lighter on snout and temporal regions; body and tail black
with numerous grayish granules intermixed; chin and throat dirty
white; underside of hands and feet light; specialized glandular
scales light gray; tail black. A more or less discrete black area
covering angle of mouth and the labial region below eye, extending
to the second labials; digits dark, flecked with light.
Measurements in mm.: Snout to vent, 43.5; tail (reg.), 48.2; width
of head, 8; length of head, 13.1; axilla to groin, 18.2; arm, 16; leg, 20.
Variation: The Costa Rican specimens in the collection are from
the following localities: Puntarenas Province: KUMNH Nos, 34554-
Review or Lizarps or Cosra Rica 31
34573, 37033, Maribella Hotel (mouth of Barranca River), Limén
Province: Nos. 34574-34604. Limon; 34603-34606, Mountain Cow
Creek; 34607-34608, La Lola. Guanacaste Province: two unnum-
bered specimens from Liberia. The species has been reported from
Bebedero.
Sexual dimorphism in color is striking, the females differing in
having the ground color gray or brownish gray. There are two dark
spots on the rostral above, three spots on the frontal region, and
a broken horseshoe-shaped dark mark beginning behind the eyes
Partly surrounding several indefinite dark marks on the occiput,
There is a pair of dark spots on neck just in front of the shoulders
and five transverse rows of indefinite dark spots on the body with
an indefinite lateral row of rounded lighter spots. A pair of dark
spots on the sacral region is followed on the dorsal surface of the
tail by several indefinite dark spots sometimes with lighter cream
centers. The sides of the body are darkly mottled in blackish and
dull cream. The chin has a median light line, and there are some
dim darker longitudinal lines that are lost on the neck. The mental
is whitish. The median part of venter has an indefinite lighter line,
and the underside is lighter than the sides of the tail. The labials
are clouded with darker, the color usually partly surrounding the
‘ream spots. The toes are banded with dark and light and the
limbs have dark and light spots. In old females the dark color is
more diffuse, the pattern as described is less distinct, the entire
ventral surface is lighter, and the chin markings may be obsolete.
There is some variation in the exact number of the specialized
“glandular” scales but specimens from both coasts show these in
“pproximately the same number and arrangement save that there
are usually some basal subcaudal scales obviously present in speci-
mens taken along the western coast, that are usually if not always
absent in eastern specimens.
Remarks: In Costa Rica the species is known in low areas es-
pecially near the seashore. It was very common in the port city of
Limon. Many specimens were observed in the trees of the city
Park, in fact specimens could be observed in almost any sizeable
tree in the city or its environs, Occasionally the gekkoes were seen
about houses. Two newly-hatched young taken at Liberia, Guana-
caste were found in the doorway of a hotel.
The females lay a single egg. One egg measured 7.1 x 6.15 mm.;
the total length of an unhatched embryo was 33 mm,
THe Universrry SCIENCE BULLETIN
Genus SpHaERopActyLus Wagler
Sphaerodactylus Wagler, Naturliches system der Amphibien mit vorangehender
Classification der Saugetiere und Vogel, 1830, p. 143 (Sphaerodactylus sputa-
tor Cuvier, type of genus ).
Description of genus: Digits narrow, slender, free, each with an
asymetrical terminal lamella, the terminal portion transverse and
in a different plane from the narrow proximal portion; scales under
digits simple, transverse, usually undivided; all digits with claws
retractile into a sheath on the proximal side of the terminal lamella;
body scales may be granular, flat, or keeled, and either juxtaposed
or imbricating. The “eyelid” is a spectacle with an extrabrillar
fringe closely applied to eyeball and widened about upper anterior
part of eye. No preanal or femoral pores, but in males a series of
glandular scales on venter, anterior to vent, and in some species
similar scales extending onto femoral region. Pupil usually, if not
invariably round or very nearly so. A soft pointed spine is above
the eye.
There are probably seven species of this genus in Costa Rica,
including one on Cocos Island. It seems wise to recognize the fol-
lowing described forms: Sphaerodactylus homolepis Cope, 5. lineo-
latus Lichtenstein and von Martens, S. imbricatus Andersson, S.
millepunctatus Hallowell, S. pacificus Stejneger. Aside from the
described forms there is evidence of the occurrence of another spe-
cies in southwestern Costa Rica discussed elsewhere.
Sphaerodactylus glaucus Cope,* described from near Mérida,
Yucatan, has been reported from near Sipurio as follows: “Variety
with dark bordered interscapular and sacral transverse Spots. it
purports to be a specimen from the William Gabb Collection. The
proper allocation of this specimen is in doubt. It is not impossible
that a smooth-scaled form, still undescribed, inhabits southeastern
Costa Rica. On the other hand it may be a young specimen Of a
described form. I await more material before I shall regard 5S.
glaucus a member of the Costa Rican fauna. It would not be sur-
prising, however, if it did occur since lineolatus occurs in the state
of Veracruz, Mexico.
The genus occurs in the West Indies, southeastern Mexico, Cen-
tral America, and northwestern South America.
The species are for the most part diminutive, and arboreal. Only
a single egg is laid. This is placed in some relatively dry place
* Journ. Acad, Nat. Sci, Philadelphia, ser. Q,. voli. 8° 1876: (1875), pie Ts:
Review oF Lizarps oF Costa Rica 83
under bark or debris, usually in company with the eggs of other
individuals. So far as is known there is no voice mechanism in these
gekkoes.
The presence and distribution of glandular preanal and femoral
scales, the so-called “escutcheon” * scales in the males of Sphaero-
dactylus offer very important characters in delineating species.
These have been completely overlooked by many workers in this
group.
Kry to Costa RicAN SpEciEs OF SPHAERODACTYLUS
1. Subcaudal scales forming a single enlarged median series. Scales under
regenerated tail much widened. Males with “escutcheon’’ scales
of the preanal region continuous with the femoral series kn)
Subcaudal scales not forming a single enlarged median seri
of regenerated tail not strongly widened. Males with escutcheon
scales forming a group in preanal ventral region, none on femora... — 3
9
2. Snout more elongate; a somewhat depressed area on median dorsal
part of rostral; a crural spot outlined in cream often with cream area
following; a white line on back of thigh bordered above by a brown
stripe. In male, escutcheon scales are somewhat pigmented around
edges, with two rows extending to near knee. Head of male reticu-
lated in black, dotted below; dorsal scales imbricate, keeled. Female
with five or seven narrow lines on head............... lineolatus
Snout shorter without depressed area on rostral. Scales on snout larger
than in lineolatus, strongly keeled, as are scales on back. Escutcheon
scales cream to bluish white without surface pigment, the femoral
scales in three rows proximally, two distally; a dark and light band
on back of thigh; male lacking heavy black reticulation on head
BRCM cee Ne ec aur eee osu arte millepunctatus
3. Body and tail banded with darker, the young with smooth scales, adults
With: scales somewhat keeled) aviv cca eG homolepis
Inkoidhy sahocalnzhill voyeledorumcekctal yykdal Uhlig oe ee 4
4. Head of males heavily striped or reticulated with black, similar area on
chin and throat; females with obsolete head lines but with dim
lineation on neck extending over shoulders, breaking up into in-
definite rows of dots on body; no crural spot; back of thigh brown
enclosing one or two cream spots; scales on body strongly imbricate
and keeled. Smaller, 28 to 30 mm. snout to vent........ imbricatus
Males without heavy black lineation or reticulation on head. Gen-
erally light lavender, with darker lavender clouding tending to form
bands on neck; preanal escutcheon scales 10-16; scales on dorsum
ridged, juxtaposed, becoming somewhat flattened, imbricate and dis-
tinctly keeled on flanks. Larger species, 47 mm. snout to vent,
pacificus
*So named by Chapman Grant.
Tue Universiry SctENCE BULLETIN
Sphaerodactylus pacificus Stejneger
(Figs, 5,6)
Sphaerodactylus pacificus Stejneger, Proc. Biol. Soc. Washington, vol. 16, 1903,
pp. 3-4 (type locality, Cocos Island, Pacific Ocean west of Costa Rica,
Prof. Biolley, collector. USMN No. 31057); Barbour, Mem. Mus. Comp.
Zool. Harvard College, vol. 47, no. 38, Dec. 1921, pp. 236-237, pl. 1, fig. 3,
pl. 18, figs, 14,
Diagnosis: A rather large species of the genus (maximum size
known, 47 mm. snout to vent); head and dorsal scales keeled,
juxtaposed, small on middle of back, about 22 to 24 scales in a
distance equal to that between tip of snout and center of eye,”
smaller (about 36 in same distance) on occiput; about 12 on median
part of venter; supranasals large, separated by two or three scales;
snout somewhat wedge-shaped, flattened; mental truncate pos-
teriorly, usually followed by two scales and these in turn by four;
tail slightly longer than head and body; ear-opening not, or but
rarely horizontal; a group of specialized glandular scales on venter
in male anterior to vent (possibly subserving the function of pre-
anal or femoral pores). A pointed scale on supraciliary border.
Description of the species (From KUMNH No, 38958): Head
somewhat triangular, sloping strongly from interorbital region to
tip of snout, leaving snout rather wedge-shaped; rostral large, the
length of the part visible above, 1.2 mm., notched behind with a
short median groove running forward nearly half length of scale;
a pair of enlarged supranasals, separated by two small granular
scales, with a third crowded into the rostral notch; four large supra-
labials anterior to the median point of eye, at which point the line
of the mouth curves upwards abruptly, bordered by two small
supralabials, making a total of six supralabials; nostril surrounded
by rostral, supralabial, and a small “post nasal,” the first labial
being excluded from contact; ** movable part of “eyelid” nearly sur-
rounds eye, the anterior and superior parts widest, covered with
six to eight rows of granules; above the middle of eye a small erect
spine on edge of the “lid” arising from the last of a series of larger
scales; sixteen scales across snout between the second supralabials;
frontal scales keeled, those on snout flat; mental with a larger labial
border than rostral, truncate behind, bordered by three small scales;
three infralabials to a point below middle of eye, the first longer
than the two following; two very small posterior infralabials; ear-
opening small, diagonal but not horizontal; scales on snout larger
* This “standard distance” used by Barbour loc, cit., is useful as a basis of comparative
measurement,
** This differs from the statements in other descriptions.
Review or Lizarps or Costa Rica 35
than other head scales; on occiput 35 granules in a distance equal
to that from tip of snout to middle of eye. On dorsum near middle,
9 ay; ; .
22-24 granular scales; on throat, 30 in same distance. Scales granu-
lar, juxtaposed with a keel visible on those of the sides and dorsum,
5. Mic. 5. Sphaerodactylus pacificus Stejneger. Inner figure, KUMNH No.
38958; actual length 128.5mm. Middle figure, No. 38953; length, 121 mm.
Outer figure, No. 388961; length, 129.5. All from Cocos Island, Pacific Ocean,
West of Costa Rica.
the scales tending to form straight longitudinal rows; scales on
venter much larger than those on back and sides, subcycloid, im-
bricate; on sides and dorsal parts of tail, fourteen straight transverse
rows in standard distance; four or five rows of subcaudal scales
36 Tue Universiry SCuENCE BULLETIN
are larger, irregular, not arranged in straight rows; two well-defined
hemipenial swellings behind vent, with three slightly enlarged
tubercular scales lateral to and somewhat in advance of the
swellings; tail circular in cross-section; in advance of the vent a
group of about 15 specialized “escutcheon” scales, lacking pigment;
limbs overlap a distance equal to length of longest toe; digits
terminating in an unsymmetrical flat digital disc, on inner side
of which the retractile claw is directed inward; granules on upper
posterior part of limbs smaller than those on the anterior faces;
subdigital lamellae broad, flat, single, save for those immediately
following the terminal lamella.
J & ae
——-s
imm.
Fic. 6. Sphaerodactylus pacificus Stejneger.
Escutcheon scale pattern. Cocos Island (topo-
type); much enlarged.
Color in life: Above generally light lavender with darker lavender
clouding and markings tending to form narrow bands across neck;
yellowish on venter and under tail.
_ Measurements in mm.; Snout to vent, 42.2; total length, 88; tail,
45.8; width of head, 7; head length, 12.2; snout to ear, 10.8; arm
and hand, 12.2; leg and foot, 15.4; snout to eye, 4.6; diameter of
eye, 2.
Variation: A series of 21 specimens (KUMNH Nos. 38953-38973 )
taken at the same time as the described specimen, vary considerably
in the character of the dorsal markings. This is shown in the figures
here given. In some of the specimens the lighter markings are
frosted silvery flecks. Younger specimens are somewhat rose-colored
under the tail.
Review or Lizarps or Costa Rica 37
Compared with the type description there are certain differences
that are evident. Thus in my series the first labial is excluded from
the nostril. The ear opening is varied and in none is the ear-opening
horizontal. In some it is somewhat moon-shaped, in others the
Opening is nearly circular. From the type description it is obvious
that the tail of the type is a reproduced one, since in the original
tail the scales form very straight rows across the tail and are almost
completely uniform in size except ventrally. Moreover, the ventral
caudal scales although a little larger are not widened except where
the tail is reproduced. Following the mental there are usually only
two scales and these followed by four; behind these, the scales be-
come smaller and merge into the minute granules of the throat.
Specimens are noted with five supralabials to below center of eye
and at least one has four infralabials on one side to the level of the
middle of eye. The modified glandular scales on the median pos-
terior part of the venter have a very thin loose epidermis under-
neath which is a flat cake of soft friable presumably noncellular
material which may be loosened and removed with the touch of a
needle; the bottom of the cuplike cavity shows a pigmented surface.
The number of these scales in pacificus varies from ten to sixteen.
They are contiguous but the group is not necessarily symmetrical.
Remarks: The gekkos were found for the most part close to the
shore, under debris, and especially about the bases of coconut trees
where the outer corky bark had grown over the surface of the aerial
Toots. They were captured when the bark was torn or chopped
away. The eggs of this species were found hidden among these
roots, usually several inches above the surface of the ground. Eight
or ten empty shells might be in the same nook or live eggs might
be mixed with the empty shells. In one place six live eggs were
taken. Two of the females in the series are gravid each having
only a single oviductal egg. It is presumed that this represents the
Normal clutch. Most of the females (but none of the males) have
@ goiterlike swelling on each side of the neck. An examination of
Preserved specimens shows, within the swelling, a sac filled with a
calcareous deposit, connecting with the ear cavity. The deposit is
pure white in color and of about the hardness of crayon. It is not
or scarcely evident in the younger specimens. This goiterlike con-
dition is also indicated in the figure given by Barbour (loc. cit. pl.
1, fig. 3),
Stejneger described this lizard from a small series of four speci-
mens collected on Cocos Island by Prof. P. Biolly in 1902. A speci-
men of Anolis townsendi was also taken, a species described by
38 Tue Untiversiry SCIENCE BULLETIN
Stejneger in 1900 (Bull. Mus. Comp. Zool. Harvard Col., vol. 36,
no. 6, Nov. 1900, pp. 163-64), and possibly the only other lizard on
the island. I have examined the types in the U. S. National Museum.
Sphaerodactylus homolepis Cope
(Fig. 7)
Sphaerodactylus homolepis Cope, Proc. Amer, Philos, Soc, vol, 23, no, 122,
Noy. 1886, p. 277 (type locality, Nicaragua; type USNM No. 14207);
U.S. Nat. Mus, Bull, 32, 1887, p. 27; Boulenger, Catalogue of the lizards
in the British Museum, 2nd ed. vol, 3, addenda, p. 491; Giinther, Biologia
Centrali-Americana, Reptilia and Batrachia, Apr. 1893, p. 82 (no speci-
Ps ct a homolepis carinatus Andersson, Medd, Goteborgs Mus, Zool,
Af: 1916, p. 5 (type locality, Costa Rica. C. Bovallius coll.; Dunn,
Herpetologica, vol. 1, no. 7, Jan. 29, 1940, p. 189-190).
Sphaerodactylus lincolatus (part.) Barbour, Mem, Mus, Comp, Zool,, Harvard
Col., vol. 47, no. 8, Dec., 1921, pp. 238-240, pl. 4, fig. 1.
Diagnosis: Young not keeled above; adults with dorsal scales
keeled; subcaudals similar to dorsal caudal scales, i.e. not greatly
enlarged on median row.
Head with seven longitudinal lines which may be distinct or
obsolete. Four dark lighter-bordered cross-bands on body, one on
nape, one on shoulder, one on middle of body, one in lumbar region.
Tail with four dark bands separated by four lighter bands each
more than double width of light bands,
Description of species: “Scales of upper surfaces small, flat, not
granular nor keeled, a little smaller than those of the abdomen.
Rostral plate large. Labials 5/4, first inferior labial corresponding
to three superior labials. Muzzle a little longer than distance from
eye to auricular meatus, and one and two-thirds times the length
of the eye’s diameter. Scales of lower surface of normal tail similar
to those of upper surface,
“Brownish cream color with dark brown bands, longitudinal on
the head, and transverse on neck, body and tail. There are seven
lines on the head, one median, and three on each side. The inferior
is short and is anterior to the auricular meatus; the second extends
from the end of the muzzle through the eye to the neck, and the
third runs backwards from the superciliary region to an equal length.
The crossbands are not so wide as the spaces between them. One
is at the nape, one crosses shoulders, one the middle of the body
and one on the groin. There are four complete annuli on the tail.
“This species is of very small size. Total length m. .024; of head
and body, .016; of head to auricular meatus, .004. No, 14207.
“This Sphaerodactylus is nearest the S. sputator of Cuba.
Review or Lizarps or Costa Rica 39
“In that species the scales are smaller, there are subcaudal scutella
and the head bands are less numerous and distinct.” Cope, 1886.
(The specimen one learns from the context is from Nicaragua
sent by Dr. J. F. Bransford, U.S. N.)
Remarks: Cope described this species from a small juvenile speci-
men, reported as being 24 mm. in total length, the head and body
Fic. 7. Sphaerodactylus homolepis Cope. Figure
from Barbour, “Sphaerodactylus,” pl. 4, fig. 1.
16mm. The length of the tail is thus only one half that of the head
and body, since it has banded color it must be presumed that it is a
“normal,” original tail. Barbour (loc. cit, pl. 4, fig. 1), figures a
specimen of this species from Zent, Costa Rica which he regards as
the young of Sphaerodactylus lineolatus Lichtenstein and von Mar-
tens. The markings in the figure and his description agree rather
closely with Cope’s original color description as far as can be ascer-
40 Tae University SCIENCE BULLETIN
tained. In this the tail is shown to be nearly as long as head and
body.
Andersson reports a specimen, described as Sphaerodactylus
homolepis carinatus nov, var. that is a large adult specimen 53 mm.
in length of which the head-body length is 26, the tail 27. Thus it
appears that the banded pattern is not merely a juvenile coloration
but a characteristic of the species. Barbour has regarded it pre-
sumably as a juvenile character.
A specimen that I believe to have been of this species, also an
adult, was seen at Los Diamantes, Guapiles, Limon Province, Costa
Rica. It was on the buttress of a large tree amply protected from a
collector by wasps from several very active nests. The specimen
escaped into a nest of large ants under wasp protection.
The chief differences between Andersson’s carinatus specimen
and the type are the presence of distinctly keeled scales, and a longer
tail, both of which might be accounted for by the juvenile character
of the type, since in certain other species the younger specimens
having a relatively shorter tail than adults; and if keeled, the keels
may be indistinct or even seemingly absent in the very young, The
general similarity of color pattern also lends weight to their being
the same form.
Sphaerodactylus lineolatus Lichtenstein and von Martens
(Figs. 8, 9)
Sphaerodactylus * lineolatus Lichtenstein and von Martens, Nomenclator Rep-
tilium et Amphibiorum Musei Zoologici Berolinensis, Berlin, 1856, p. 6 (type
locality, Veragoa = Veragua, Panama’); Bocourt, Mission Scientifique au
Mexique et dans l’Amérique Centrale; Etudes sur les Reptiles, livr. 2, 1873,
pp. 46-47 (a description of a type specimen loaned by W. Peters); Boulen-
ger, Catalogue of the lizards in the British Mus. 2nd ed. vol. 1, 1885 pp.
221-222 (part.) (two varieties listed one from an unknown locality, the
other based on two specimens, one from Santa Cruz and one from Chontales,
Nicaragua); Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
Apr. 1893, p. 82 (part.) (reported from Mexico, Nicaragua, British Hon-
duras, Panama and Guiana); Barbour, Mem. Mus, Comp. Zool., Harvard
Col., vol. 47, no. 3, Dec. 1921, pp. 238-240 (part.) (lower Central Amer-
ica, Guatemala, Nicaragua, Costa Rica, Panama; Noble and Klingel, Amer.
Mus. Novitates, no. 549, Aug. 11, 1932, p. 14; Gaige, Hartweg, Stuart, Occ.
Papers Mus. Zool., Univ. Michigan, no. 357, 1937, p. 8 (Nicaraguan locali-
ties); Dunn, Herpetologica, vol. 1, 1940, pp. 189-190 (part.)).
Diagnosis: A median row of enlarged subcaudals; dorsal scales
small, equal, lightly carinated; brownish fawn with five or seven
dim longitudinal dark lines on head all disappearing on anterior
part of body, except median, which does not pass beyond occiput;
pectoral and abdominal scales cycloid or somewhat diamond-
shaped.
* Spelled Sphaeriodactylus.
Review or Lizarps or Costa Rica Al
Description of species: The original type description of this spe-
cies is very brief. It follows:
“Squamis dorsi aequalibus, granularibus. Rufescens, capiti supra
linia mediana et utrinque tribus lateralibus nigricantibus ornato,
dorso fusco-vermiculato. Long. a rostro ad ca ud. bas 1k’,
caud, 1",)”
Fortunately one of the cotype specimens was later redescribed by
Bocourt (loc. cit.) so that little doubt remains as to the application
of the name. I designate this specimen described by Bocourt as the
type (lectotype) of the species.
Bocourt’s description (rough translation) reads:
Upper part of the body brownish fawn with five very distinct
longitudinal lines which arise on the head passing over the neck,
and disappearing on the anterior part of the trunk. Scales of the
back small, equal, lightly carinated, those of the venter larger,
smooth, thin, and imbricated. Pupil rounded, claws relatively long.
This small species resembles Sphaerodactylus fantasticus Cuvier
somewhat in the general form of the head, the rostral and mental
scales, the labials and the ear-opening. Nevertheless it may be
distinguished by the following characters: scales on the muzzle
larger than those covering the temporal and occipital regions; on
the neck, the back and the superior parts of the limbs they are small,
of equal size and lightly carinated; under the neck they are granu-
lar; pectoral and abdominal scales much larger, diamond-shaped,
smooth, thin, and imbricated. Tail mutilated, circular at its begin-
ning, and probably of moderate length; above covered with imbri-
cating scales arranged transversely; below there is a longitudinal
Series of large scales of nearly equal diameter and hexagonal form.
Digits slender, their inferior surfaces bearing transverse scales, and
having at their extremities a small smooth lamella, circular, and
bearing a relatively large claw placed on the proximal edge of the
lamella.
Length from snout to vent, 26 mm.
The upper parts of the body are brownish gray; five longitudinal
lines of a darker color arise on the head. The shortest one is mesial
on top of head and does not pass beyond occiput; the other two
Pairs begin one above the other below the eye and disappear at the
beginning of the sides. Lower surfaces pale yellowish, the throat
finely punctate with brown forming small linear designs slightly
undulating. It is to the courtesy of Prof. Peters that the Paris
Museum is acquainted with this small saurian, which comes from
Veragua, Central America.
TH J,
e
4,
Z
Z
=
L
Z
NILATIOG
Fic. 8. Sphaerodactylus lineolatus Lichtenstein and von Martens. KUMNH No. 40504, actual
length snout to vent, 27 mm., total, 45 mm.; 40502, snout to vent, 30 mm.; 36123, snout to vent, 28.5; 40503,
snout to vent, 30; tail, 53. All from Tenorio, Las Cafias, Guanacaste, except 36123, from Tilaran, Guana-
caste.
Review or Lizarps or Costa Rica 43
Specimens of lineolatus agreeing so far as can be ascertained
with Bocourt’s description are in the collection. These are KUMNH
Nos. 40501-04 from Tenorio, Las Cafas, Guanacaste, Costa Rica.
The specimens from Tenorio were found on the buttresses of a
large forest tree ensconced in growing plants. The general color
was a light tan, approaching fawn color, with narrow blackish lines
on the head, extending back to body. The chin and venter were
bright yellow, or in one case cream-white on chin and venter. The
under side of the tail was pinkish. The toes are banded indefinitely
in dark and light. There is dark brown or blackish crural mark
which may be bordered with cream or there may be a cream area
preceding and following this mark.
The head markings are very narrow black lines arranged as fol-
lows: a black line from nostril to eye; a median line from back
level of eyes to occiput; two lines from eye, the upper running back
and tending to curve inward somewhat while the other runs straight
back towards shoulder, the area between them showing a dimly
distinct lighter line. Two other lines beginning at lower part of
eye, the upper passing above the auricular opening, the other below
it. A rather distinct white line bordering the darker dorsal color
laterally has an indefinite darker border below it. There is a
sharply defined white stripe on the back of the thigh that is con-
tinuous with a whitish bar across the lower dorsal part of the tibia.
Above this line on the thigh is a transverse dark-brown stripe.
There is a cream dark-edged spot on the knee, and a small ocel-
lated spot above the thigh near the body. The arm has a cream
mark on the elbow and one on the forearm.
A male specimen, 40505, taken at Tenorio is also referred to this
species. The color, different from that of the females, is as follows:
above tan (nearly fawn) with minute blackish or blackish brown
dots, slightly larger and somewhat elongate on the basal part of the
tail; the head is black, the reticulations. enclosing cream spots and
areas and tending to form three black and three cream lines pos-
terior to eye. Most of the black markings terminate shortly behind
Occiput. The rostral, mental, and labials are cream, with distinct
black spots, the chin and throat cream being strongly punctate with
black. The venter is whitish with a fine scattering of brownish pig-
ment. The glandular scales (figured) have a narrow border of
brownish pigment. There is no black spot on the sacrum. The
arms and legs have faintly distinct cream spots bordered by a
darker color and a cream line on back of thigh. The underside of
the tail is pinkish with some brownish pigment. There are some
44 THe Universtry SCIENCE BULLETIN
indistinct light bands on the digits. In preservative the cream has
become white and the pink shade has disappeared.
In all these specimens (except those with regenerated tails) there
is a strongly defined median series of subcaudals. The alternate
scales are slightly smaller than the scales which precede them.
Fic. 9. Sphaerodactylus lineolatus Andersson. KUMNH No. 40506; Tenorio,
Las Cafias, Guanacaste. Pattern’ of escutcheon scales, much enlarged.
The snout is rather long and there is a flattened depressed area
on the part of the rostral scale exposed above. The scales are rather
large on the back and sides, there being 42-44 strongly keeled scale-
rows. The rows of cycloid scales on venter vary between 19 and
22 thus making a total of approximately 65 scales around the middle
of body.
Measurements of Sphaerodactylus lineolatus
INEMDCY ya ce eaGr A oaths Cia | SOO LO e 40501 9 40502 9 40503 2 40505
WOGHUUY rose Peper at Tilaran Tenorio Tenorio ‘Tenorio Tenorio
Snout to vent..... a 28.2 30.5 29.2 29 26.5
WEA pie cae ss ata Data eninge k wee ura Ver aR Mara 12.8* 22.5* 18.5*
ON Ms Of MOA seit vies 9 8.5 8.6 8&
WV MON OOH yen itt ney) pat 4.9 5.3 §.1 5,2 6
PMOUt1O Ati ise) stele 1] WA} 11.3 alent 11
ASING $6 SPO cee eae 13.2 15.2 14.5 14.6 12,2
Nha ve ace Ves ve A Ey ki vf 7.3 wad 7.3 7
HOC Mee trikes we vs Pe eal he NEM een 8.7 9.2 9 8.8 8.3
Snout to eye... . ee 4 4 4 3.7
* Incomplete.
Review or Lizarps oF Costa Rica 45
Remarks: Barbour’s treatment of lineolatus probably involves
more than one species. He specifically mentions enlarged sub-
caudals in the specimen he describes from Saboga Island, Panama
Bay, but examination shows that they are not in a single median
line in the specimen. The only data in the literature regarding the
glandular preanal and femoral scales is a statement by Noble and
Klingel (1932). They do not give the source of their lineolatus and
it may or may not be that of Lichtenstein and yon Martens. They
state that S. lineolatus has a band of hypertrophied scales on the
thigh, closely resembling S. cinereus. The latter is described as
having the median group [of hypertrophied “escutcheon” s sales |
small, and a narrow band of thickened scales extending down the
ventral surface of the thigh for its entire length. The band is three
scales wide at the proximal end of the thigh and two scales wide at
the distal end.
Sphaerodactylus imbricatus Andersson
(Figs. 10, 11)
Sphaerodactylus lineolatus imbricatus Andersson, Med. Goteborgs Mus. Zool.
Afd, 9, 1916, p. 5 (type locality Costa Rica, C. Bovallius collector ).
Sphaerodactylus Daeolans (part.) Dunn, Herpetologica, vol. 1, no. 7, 1940,
pp. 189-190.
This species originally described from Costa Ric: ( presumably
from the eastern lowland) was synonymized with S. lineolatus by
Dunn who states that this form, together with several other named
species or subspecies (S. homolepis carinatus Andersson, S. homo-
lepis Cope, 8. argus Meerwarth [non Gosse] and possibly S. mille-
punctatus Hallowell), and “all other Sphaerodactylus material I have
seen from Panama, Costa Rica, and Nicaragua” are conspecific with
lineolatus.
Diagnosis: Scales about body in 70-73 rows, about 20 of which
are smooth, cycloid, ventral; escutcheon scales confined to preanal
region, none appearing on femora; strong sexual dimorphism; small
species probably not exceeding 30 mm. snout to vent, dorsal scales
flat, strongly imbricate, keeled.
Description of species (From KUMNH No. 40506, Bataan, Limon
Proy., Costa Rica): Snout much narrowed at tip; rostral large,
without a depressed flattened dorsal area, notched posteriorly and
with a forward-directed median suture or groove, the most pos-
terior part of the median dorsal portion of the scale not r saching
front level of nostril; a pair of large supranasals bordering rostral,
separated anteriorly by a single tiny scale, in the middle by two, and
46
Tue UNtversiry SCIENCE BULLETIN
posteriorly by four scales; nostril surrounded by rostral, supranasal,
first labial, while only a single small scale borders the posterior rim;
six supralabials, the first a little smaller than second, the series then
diminishing in size posteriorly, the last, minute, lying behind level
of median part of pupil; behind this, mouth bordered by five or six
Fic. 10, Sphaerodactylus imbricatus Andersson. Left figure,
KUMNH No. 40507 9; actual length snout to vent, 31.2, total,
45.1 mm.; right figure, 40506 ¢@; snout to vent, 28.8, total length,
46 mm. Bataan, Lim6én Province, C. R. Female has calcium de-
posits on side of neck.
small granular scales; scales on snout somewhat enlarged, flattened,
keeled, about 19 between posterior borders of third labials; “eyelid”
continuous, surrounding eye, widest above and in front of anterior
half of orbit; this part bordered by a row of somewhat larger keeled
scales, the posterior developed into a soft pointed spine; below
these, on under surface of lid is a row of larger scales, lacking me-
Review or Lizarps oF Costa Rica 47
dian keels; pupil circular. Four infralabials, the first very large,
equal to three and a half supralabials; fourth lies below sixth supra-
labial and is of about same size; mental with a larger labial border
than rostral, its posterior part reaching back mesially beyond level
of the mental-infralabial suture, and bordered by three somewhat
enlarged chin scales. On chin and throat small juxtaposed some-
what cycloid granules continued to narrowest point of neck, at
which point the scales abruptly become larger, cycloid and defi-
nitely imbricating and continue back over breast and venter to anus,
interorbital scales keeled, a little larger, and more elongate than
keeled scales on occiput; dorsal scales ridged, juxtaposed, those on
:
ee
Fic, 11, Sphaerodactylus imbricatus Anders-
son. KUMNH No. 40506. Escutcheon scales
on venter (much enlarged).
sides and flanks becoming a little larger and somewhat imbricate.
Scales on the tail forming transverse rows but the segments are not
clearly marked; on under surface of tail the scales of several rows
are larger than lateral and dorsal scales but there is no median
enlarged series present; the ventral scales laterally pass gradually
into the smaller laterals so that an exact count of dorsals and ventrals
is difficult, but there are approximately 53 dorsal and 20 ventral
rows around widest part of body. The specialized glandular “es-
cutcheon” scales in the preanal part of venter number 60; none are
present in the femoral region.
Scales on dorsal and anterior part of upper arm and forearm
48 THe University Science BULLETIN
larger, imbricating, other scales smaller, juxtaposed, granular; legs,
except back face of femur, with larger cycloid imbricating scales.
Digits asymmetrical terminating in an enlarged lamella, irregular
in shape, the distal, somewhat oval portion lies transversely and is
in a slightly different plane from the posterior narrower portion;
claws are retractile into a sheath lying along the inner (proximal )
edge of lamella; subdigital scales transverse, single (rarely one or
more divided), eleven or twelve under longest toe. No postanal
sac or bones.
Color: Entire head and extreme anterior part of neck black and
cream-white arranged in more or less irregular black and white
longitudinal stripes on sides of head; one black line beginning on
rostral continues behind eye; the median lines are irregular and
somewhat reticulate; labials with black spotting; chin and throat
heavily reticulated in black and cream-white, the color extending
back to the narrow point of neck, where it ends abruptly; remainder
of neck and body nearly uniform brownish with a few indefinite
brownish flecks on sides and posterior part of dorsum, each fleck
not exceeding size of a scale; venter light brown, somewhat yellow-
ish in life, Tail lighter tan on dorsum and sides with brownish flecks
larger than those on body; under side of tail whitish; a few whitish
dark-bordered spots on leg and foot; back face of thigh with one
or two indefinite light areas. There is no evidence of darker lines
on neck and shoulders.
Measurements in mm.: Snout to vent, 28.8; total length (part of
tail missing), 46; axilla to groin, 11; length of head to ear, 7.1; width
of head, 5; arm, 7.3; leg, 9; snout to center of eye, 4; center of eye
tO Cat, 3.0,
Variation: Three female specimens referred to this species are
KUMNH No. 40507 from Bataan, Limon Province and 40508-40509
from Lake Bonilla, Tunnel Camp, Peralta, Cartago Province. These
lack the heavy black marking and reticulation on the head, as well
as the glandular preanal scales.
The scalerows around the widest part of the body are from 72
to 74 of which 20 or 22 are ventral, cycloid, smooth, and imbricate,
The subcaudal series agrees with that of the described male, as does
the keeling and general characters of body squamation.
The color markings are different. The head is a little lighter than
body, and beginning at the nape there is a series of six or eight
very dim nuchal darker-brown lines extending onto shoulders then
breaking up into very numerous smaller brown flecks over the body.
Review oF Lizarps or Costa Rica 49
There is a light darker-edged spot on back of thigh, one at the ankle
and knee, and one or two light flecks on the dorsal surface of the
thigh. The arm is indefinitely reticulated with brown, enclosing a
slightly lighter area. On the head there is a canthal line and two
indefinite darker lines behind the eye.
In the Lake Bonilla specimens the nuchal lineation is more pro-
nounced, and the dark flecking on the body is more in evidence.
Remarks: The described specimen was captured on a tree at
Bataan. Andersson (1916) mentions having three specimens. The
type presumably is designated by No. Re. ex. 1326. However, the
description is something of a composite and should the number
represent a lot number I propose that one of the specimens with
the black and white head be designated the type. It is, of course,
not impossible that two forms are represented in Andersson’s ma-
terial,
Sphaerodactylus millepunctatus Hallowell
(Figs. 12, 18)
Sphaerodactylus millepunctatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia,
Oct, 1860, pp. 480-486 (type locality, Nicaragua fide Cope. Hallowell
states, “abdominal scales carinated.” Cope [P. A. N. S.P. 1861, p. 499]
regards Dr, Hallowell’s statement concerning carinated scales as being in-
correct); Cope, Proc. Acad. Nat. Sci. Philadelphia, Dec, 1861, pp. 494-500
(Nicaragua); Troschel, in von Miiller, Reisen in den Vereinigten Staaten,
Canada und Mexico, vol. 3, p. 599; Bocourt, Mission Scientifique au Mexique
et dans Amérique Centrale; Etudes sur les reptiles, livr. 2, 1878, pp. 45-46
(no specimens ).
The discovery of a specimen of Sphaerodactylus at Tenorio, Las
Cafias, Guanacaste, Costa Rica, 70 km. south of the Nicaraguan
border, that differs from lineolatus and seems to agree with the
published characters of millepunctatus, leads me to believe that
previous authors who have placed this form in the synonymy of
S. lineolatus may have done so in error. The type description of
the species is very brief:
Special characters: “Dorsal scales very small, unicarinate; color
reddish with numerous small brownish spots; under parts white;
length of head and body, 11 lines.
Description: “Scales upon muzzle larger than those upon the
vertex; those upon body remarkable for their small size, . . .
Abdominal scales carinated, much larger than those upon throat
and chin; color reddish-brown above, with numerous brown spots,
intermingled with very minute white points; under parts white.
Two specimens.”
Cope (1861, loc. cit.) adds some pertinent data, stating that the
3—3378
50 Tue University SCIENCE BULLETIN
occipital scales are granular, the frontal scales keeled, the muzzle
short and the auricular orifice smaller than the digital expansion;
the color is light brown with a postaxillary, a loreal, a median oc-
cipital, and two postocular brown lines. A brown, transverse,
crural spot, bordered posteriorly with white. From Nicaragua.
Fic. 12. Sphaerodactylus millepuntatus Hal-
lowell. KUMNH No. 405044; Tenorio, Las
Cafias, Guanacaste, C. R.; actual total length of
figure, 35.4.
Diagnosis: A small species, the scales strongly keeled on frontal
and dorsal body regions; reddish or grayish brown above with
numerous small darker spots; white below; muzzle short; a brown
crural spot; in males a large group of glandular scales on posterior
part of venter continuous with two series extending onto thighs;
strongly enlarged series of subcaudals, each alternate scale a little
smaller than preceding.
Review or Lizarps or Costa Rica ‘OL
Description of species (From KUMNH No. 40504, Tenorio,
Las Cafias, Guanacaste, Costa Rice ): Snout short with a relatively
small rostral scale having a small entrant suture posteriorly; dorsal
part of rostral without a small depressed posterior portion; a
Pair of internasals separated by a single scale anteriorly, by two
posteriorly; nostril surrounded by rostral, supranasal, first labial,
While the posterior rim is bordered by two small scales, the larger
upper one of which appears to be segmented from internasal; scales
on snout juxtaposed, much larger than those on occiput, each with
smooth surface but bearing a clearly defined keel; about 17 scales
ma transverse row across snout between the second labials; five
supralabials, four to a point below pupil, the first largest, the series
diminishing in size posteriorly. Mental larger than rostral, with
a larger labial border and with two postmentals touching it pos-
teriorly; five infralabials, the first larger in area than the four suc-
ceeding labials; posterior part of mouth bordered above and be-
low by minute granules. Scales following postmentals somewhat
enlarged, juxtaposed, gradually becoming smaller to narrowest
Point of neck then rather quickly transforming to cycloid imbri-
Cate smooth scales which continue on venter to vent; on sides
the transition to the ventral body scales is rather gradual, about
22 longitudinal ventral rows can be distinguished at middle of body;
‘ales on neck tubercular, juxtaposed; dorsal and lateral body scales
imbricate, distinctly keeled; ventrals smooth, Large imbricate
scales on dorsal and front face of upper arm, on part of dorsal
and front face of forearm; remainder covered with smaller granu-
ar scales; legs with large imbricate keeled scales save on posterior
dorsal and posterior parts of femur and upper surfaces of tibia
Which are covered with smaller scales or granules; toes with smooth
transverse lamella, irregular, the outer part asymmetrically trans-
verse and in a different plane from the proximal portion, which
Is differently colored from the distal part.
On the posterior part of the venter and on the ventral surface of
the thighs there is a symmetrical group of specialized glandular
Scales, whitish in color, and almost without surface pigment. The
Scales total approximately 91. The ventral (preanal) group consists
of about 30 scales from the posterior borders of which are extended
Outward three linear series of scales, two of which reach the ex-
tremity of the femoral region. On the right femur there are 29, on
the left 39 scales.
A series of greatly enlarged, transversely widened, somewhat
hexagonal, subcaudal scales (the posterior part of tail missing )
52 THe University SCIENCE BULLETIN
each alternate scale a little larger than one following; on tail the
transverse rows of lateral and dorsal scales not clearly marked; how-
ever, a diagonal arrangement is clearly defined; a row of scales
bordering subcaudal series somewhat enlarged; dorsal scales on
base of tail distinctly keeled; ear-opening moderate, somewhat ver-
tical, smaller than a terminal digital lamella.
Fic. 13. Sphaerodactylus millepunctatus Hallowell. KU
MNH No. 40504, Escutcheon scales (much enlarged). Te-
norio, Las Cafias, Guanacaste, C. R.
Color: The dorsal coloration generally is reddish to gray brown
above, the head only slightly lighter than dorsum; a short median
line barely discernible on occiput. A diagonal line from eye across
occiput intersects one from opposite side on middle of neck; second
line from eye -parallels the first, the area between a little lighter
than elsewhere. Body with very numerous dark spots covering
two to four scales, the spots more or less arranged in rows; a barely
distinguishable crural spot with a somewhat lighter area behind it;
the tail with a dim but more or less regular pattern; pink under
tail; chin and belly dirty white, pigment scattered equally on
scales, but each scale somewhat darker on borders. No pigment
on surface of the glandular scales of the escutcheon; line of de-
marcation between dorsal and ventral coloration not strongly
marked. An indistinct transverse cream line on back of thigh con-
tinuous with cream mark on ankle; light spot just below knee.
Review or Lizarps or Costa Rica 53
Measurements in mm.: Snout to vent, 27.2; tail (broken), 8.2;
snout to middle of eye, 3.8; head length, 7.3; snout to arm insertion,
10.25; axilla to groin, 11.8; arm, 6.8; leg, 8.7.
Remarks: This specimen taken in the same general area with
What I regard as typical Sphaerodactylus lineolatus seemingly dif-
fers from them in having a distinctly shorter head, shorter rostral
Without a flattened portion; a scale divided from internasal back
of nostril; larger strongly keeled frontal scales; diagonal or chev-
ton-shaped arrangement of the dorsal and lateral caudal scales;
absence of a marked delimitation of dorsal color by a light line
from axilla to groin; the terminal lamella of the digits proportion-
ally larger than in specimens of lincolatus. The male does not have
® specialized color of black and cream.
Whether this specimen is referable to Hallowell’s millepunctatus
may be questioned. Boulenger placed millepunctatus in the syn-
Onymy of S. lineolatus. Barbour disregarded the name entirely.
Tt would appear that no reviser, except possibly Cope, has seen the
type specimen. Cope in his review of the genus (1861) gives data
on Nicaraguan specimens presumably the types and corrects Hal-
lowell’s statement that the ventrals are keeled. He recognized it
as a distinct species but at the same time was seemingly unfamiliar
with S. lineolatus of Lichtenstein and von Martens.
Unless the type can be re-examined and more detailed data made
available as to the character of the subcaudals, the number of scale
tows about body and the character of the “excutcheon” scales the
Status of millepunctatus and of the described specimen will remain
obscure.
Sphaerodactylus sp?
A specimen of a small sphaerodactyl captured by John Baker, at
Palmar, Puntarenas Province, Sept. 2, 1952, either escaped from a
faulty collecting bag or was eaten by a small Leptodeira taken at
the same time. Only the tail of the gekko remained when the bag
teached camp.
I believe that the tail belongs to a species of Sphaerodactylus
that differs from the recognized forms at hand in being banded with
six lavender brown bands separated by five, distinctly narrower
cream bands. On the under side of the tail the dark bands are of
4 somewhat lighter shade. A row of 38 enlarged, somewhat hex-
agonal scales are present on the medial ventral surface, each alter-
Nate scale a little smaller than its predecessor. The adjoining lat-
Tue Untiversrry SCIENCE BULLETIN
54
eral scales are somewhat enlarged. The lateral and dorsal caudal
scales are small, imbricating, unkeeled, and rather unequal in size.
They form straight transverse rows, four to each vertebra. There
are, seemingly, 19 vertebrae as indicated by faintly distinct lines.
FAMILY GEKKONIDAE Gray
Underwood's recent redefinition of this family is as follows:
“Spectacle covering eye; vertebrae amphicoelous; clavicles per-
forated or secondarily imperforated; three or four pairs of sternal
ribs; digits with or without pilose pads; no fovea; male commonly
with preanal and/or femoral pores and postanal sacs and bones.”
He separates the family into two subfamilies. as follows:
A. Pupil vertical with straight margins. Diplodactylinae
B. Gekko-type pupil or secondarily circular. Gekkoninae
The subdivision into subfamilies is as yet based on rather meagre
characters, but subsequent study may strengthen their status.
One of the extraordinary facts which has not been pointed out
previously is that none of the Gekkonidae endemic in the Western
Hemisphere have developed preanal or femoral pores. They are
known to occur only in the imported genera, Hemidactylus and
Peropus. The two cosmopolitan genera with endemic species Tar-
entola and Phyllodactylus, are without femoral or preanal pores
throughout their range in the Western Hemisphere so far is is
known. Presumed endemic forms of Hemidactylus should be re-
examined. Underwood has shown that Gymnodactylus, sensu stictu
is a South American genus without pores and that Old World forms
formerly regarded as Gymnodactylus belong in one or more different
genera,
Genus THecapactryLus Cuvier
Thecadactylus Cuvier, Régne Animal, vol, 2, p. 48 (type of the genus Gekko
laevis = rapicauda, by monotypy).
Description of genus: Digits half webbed; tail subcylindrical with
imbricate scales; digits with two series of lamella divided by a
groove; claws present on all digits, retractile into a groove in a ter-
minal sheath (sometimes absent on inner digit). Body covered with
juxtaposed scales, those on venter imbricate; no femoral or preanal
pores, and likewise lacking specialized glandular scales on venter
and femora; pupil vertical; “eyelid” incomplete, not entirely sur-
rounding eye.
Only a single species is recognized in the genus. This is a wide-
spread form, occurring from Yucatan in Mexico to northern South
America. It is also found in the West Indies.
Review or Lizarps or Cosra Rica 55
Thecadactylus rapicaudus (Houttuyn )
(Fig. 14)
Gekko rapicauda Houttuyn, Verh, Zeeuwsch, Genootsch, Wet. Vlissingen, vol.
9, 1782, p. 823, pl. 3, fig. 1 (type locality, American Islands, restricted by
; Smith and ‘Taylor to Chichén Itz, Yucatan).
Gekko laevis Daudin, Histoire naturelle des reptiles, vol. 4, p. 112-114 (type
: locality, “Amérique Meridionale” ).
Gekko Surinamensis Daudin, idem, vol. 4, pp. 126-129 (type locality, Surinam).
Lacerta rapicauda Gmelin, Systema Naturae, vol, 1, p. 1068.
Thecadactylus laevis Gray, Ann, Philos, ser, 2, vol, 10, p. 198; Cope, Journ,
Acad, Nat, Sci. Philadelphia, ser, 2, vol. 8, 1876 (1875), p. 119 (Costa
_,, Rica “Mouth of Estrella or north River, East Coast”). :
l'hecadactylus rapicaudus Gray, Catalogue of the specimens of lizards in the
collections of the British Museum, 1845, p, 146; Boulenger, Catalogue of
the lizards in the British Museum, vol. 1, 1885, pp, 111-112; Giinther,
Biologia Centrali-Americana; Reptilia and Batrachia, April, 1893, p. 81;
peor and Taylor, U. S. Nat. Mus. Bull, 199, 1950, p. 49 (restriction of type
_ locality),
Platydactylus theconyx Duméril and Bibron, Exrpetologie Générale, vol. 3, 1836,
p. 306, pl. 33, fig. 2.
Description of species (From KUMNH No. 34112): Head
rather triangular, very distinct from neck. Rostral enlarged, its
width nearly two and one-half times its length, bordered behind
by a pair of large internasals that are wider than long, and form
@ common median suture; nostril lying posterior to suture between
rostral and first labial, surrounded by rostral, internasal, and first
labial, its posterior rim bordered by four small scales; scales on
snout only slightly larger than those on back, about fifty in a line
across snout between the third labials; eyeball nearly as large as
orbit; spectacle covering eye, the fringe incomplete, extending across
top of eye and half way down in front of eye, and for a short distance
behind posterior edge of eye; latter portion beset with a series of
eight or nine enlarged spinelike scales; scales on anterior part of
fringe somewhat enlarged, but not spinelike; a small fold on antero-
dorsal part; 12-12 supralabials, behind which there are six to eight
very small undifferentiated scales bordering mouth; ear-opening
relatively small; mental small, about half the width of rostral and
bordered behind by a pair of somewhat elongate scales (one of
these is abnormal in the described specimen), and on each side a
Series of five or six sublabials bordering the infralabials, diminish-
mg in size posteriorly; scales on chin and neck small, granular,
changing abruptly into the larger flat imbricate scales that cover
-reast, venter, the front part of upper arm, front and underside
of femur and tibia; laterally the flat scales pass gradually into the
somewhat elevated juxtaposed scales of the sides of the body;
about 125 scales about body; scales on base of tail larger than other
dorsal scales, equal or slightly larger than ventral scales, becoming
Fic, 14. Thecadactylus rapicaudus Houttuyn. Es-
quinas Forest Preserve (between Palmar and Golfito
on railroad), Puntarenas Province, C, R., actual total
length, 200 mm.
Review or Lizarps or Costa Rica 57
still larger posteriorly and reducing again in size near tip; caudal
scales forming transverse imbricating rows; tail slightly wider than
deep.
Hands and feet about half webbed; digits with two series of
lamellae separated by a deep groove anteriorly; claws. retractile
through this groove into a sheath; on dorsal side of digit the terminal
joint forms a dorsal elevation but is not free from the widened tip
of digit,
Coloration in life: Ground color brownish lavender; supralabials,
Save for dark upper edges, cream; snout variegated brown; a more
or less definite blackish stripe begins behind eye and runs back
above the ear towards the shoulder where it becomes somewhat
diffuse, the upper border of which is more intensely dark throughout
its length; above this a lighter stripe, also black-edged above, runs
back on the sides to near the middle of the body, the edges very
irregular posteriorly and interrupted by darker indefinite spots; an
"regular dark pattern on occiput; on back and across shoulders
tregular dark or black marks tending to form six or seven trans-
verse bands between neck and level of hind limbs; tail a little lighter
than body with very irregular dark marks tending to form transverse
blotches or bands but interrupted dorsally or laterally except for
one broad complete band near the end of tail; a terminal band,
Separated narrowly from the preceding by a narrow cream band, is
less dark; yenter light lavender, clouded with some darker pigment;
arms and legs brownish lavender, spotted or blotched with black;
the under side of hands and feet lighter on sole and palm, the outer
Part of digits blackish. In preservative the lavender appears brown-
ish,
Measurements in mm.: Snout to vent, 113; tail, 87; width of head,
22.2; length of head, 30.2; axilla to groin, 49; arm, 31; leg, 39.
Remarks: Costa Rican specimens examined include the following
numbers: Guanacaste Province: KUMNH Nos. 34108-34110, Las
Flores, Tenorio, Las Cafas, Puntarenas Province: 34111-34112
Esquinas Forest, between Palmer and Golfito, 34553 Palmar, Limén
Province: R. C. T. Coll. No. 1367, Los Diamantes, Guapiles.
Variation: There seems to be but little observable sexual di-
Morphism save in the region of the base of the tail. In the males
there are two strong swellings, each bearing a large slightly flat-
tened, somewhat shovel-shaped tubercle (the latter scale also pres-
eres in females but smaller and less conspicuous ). I am not able to
distinguish any significant color or pattern differences in the sexes
58 THe Untiversrry SCrENCE BULLETIN
save that females may be somewhat lighter. The geographical
variation between those from the Caribbean drainage and Pacific
drainage areas is scarcely greater than may be found in individuals
from the same localities.
In a young specimen from Tenorio, the bands on the anterior part
of the tail are more complete, irregular in size, and separated by
bands of lighter color.
The regenerated tail is usually greatly thickened at the base,
tapering more quickly than the original tail. It is grayish lavender
with very numerous short black longitudinal streaks. The scales
are smaller dorsally on the greater part of the tail, while those on
the latter fifth are larger than in the original tail.
Genus PuyiLopacryius Gray
Phyllodactylus Gray, Spicileg. Zool., 1830, p. 3, pl. 3, fig. 1, (type of the genus
Phyllodactylus pulcher, Tropical America? ).
Description of genus: All digits clawed, unwebbed, moderately
slender, terminating in a pair of widened symmetrical lamellae, be-
tween which is a groove into which the terminal claw is retractile
into a sheath; transverse lamella or scales present on under surfaces
of digits; upper surfaces of body covered with juxtaposed, more or
less uniform granules intermixed with larger tubercles; scales on
venter usually cycloid, imbricating; no preanal and/or femoral
pores; no specialized glandular scales in preanal and femoral re-
gions; a well-defined elongate lateroventral ridge or fold.
With the exception of Tarentola this genus is presumably the
only cosmopolitan genus of this family in the Western Hemisphere
antedating historical time. It is a populous one, widespread, the
species extending from the extreme southwestern United States,
western and southern Mexico, south through Central and South
America. Species are known in the West Indies and the Galapagos
Islands.
Collections I have made in Costa Rica have been notably lacking
in specimens of this genus. However, I collected a series at Tenorio,
Las Cafias, Guanacaste, in 1952, and in 1954 a few specimens
were taken in the general region about Tilaran, Guanacaste. I have
seen certain specimens of Phyllodactylus from Costa Rica, now in
the National Museum; but these specimens are in such a poor state
that I hesitate to attempt a specific allocation.
The genus has several species in Mexico, three or four of which
may occur in the same locality in the wetter southern part of the
country. It is unlikely that the genus should be reduced in Costa
Review or Lizarps oF Costa Rica 59
Rica to a single species, confined to the drier part of the country,
particularly since there are wet jungle forms farther southward.
So I conclude that there is a strong probability that at least another
form awaits discovery or recognition in the eastern drainage areas.
A species of the genus described by O'Shaughnessy, as Phyllo-
dactylus ventralis purported to be from Jamaica. However the
Species was purchased from a dealer and the provenance of the
Specimen may be questioned. I have seen two specimens of this
Species from Managua, Nicaragua and it is possible this species also
Occurs in Costa Rica. It may be easily diagnosed by the reduced
number of enlarged ventrals, the number being about 16.
The specimens I have collected are referred to Phyllodactylus
eduardofischeri Martens, described from Salvador. Seemingly they
are adapted to a relatively dry rocky habitat, rather than to wet
jungle,
Phyllodactylus eduardofischeri Mertens
(Fig. 15)
Phyllodactylus eduardofischeri_ Mertens, Zool Anz., Bd. 148, 1952, p. 88-89
(type locality, “Westufer des Rio Chilama, bei la Libertad, Dept. La Liber-
cad ; El Salvador); Abh, Senckenberg, naturf. Ges., no. 487, 1954, p. 39, pl.
5 HIG, OS,
Phyllodactylus tuberculosus Bocourt, (nec Wiegman), Mission Scientifique au
Mexique et dans l’Amérique Centrale; Etudes sur les reptiles, livr. 2, 1873,
p. 43 (La Unidén, Salvador); Werner, Mitt. Nat. Mus. Nat. Hist., vol. 12,
1928 p. 194 (Divisadero, Dept. Morazin, El Salvador); Wettstein, Sit-
zungsb, Akad. Wiss. Wein, Math-naturw, Kl. Abt. 1, Bd. 148, Heft 1, 2,
1934, p. 19 (Bebedero, Guanacaste Proy., Costa Rica).
Phyllodactylus (indet.) Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871, p. 205
near San José, Costa Rica); Cope, Journ. Acad, Nat. Sci. Philadelphia,
Ser. 2, vol. 8, 1876 (1875) p. 118 (presumably same specimen mentioned
supra); U.S. Nat. Mus. Bull. 32, 1887, p. 28 (part).
tis entirely probable that certain other references to Phyllodactylus tubercu-
osus in Central America, apply to this species. ]
Diagnosis: A medium sized species (to 64 mm, snout to vent)
the complete tail equal to or a little longer than head and body;
ventral scalerows usually 32-34; usually 22-24 granules between
middle of eyes; usually 14-16 rows of enlarged dorsal tubercles;
Supranasals in contact behind rostral; tail segments with several
transverse rows of small scales intermixed with larger somewhat
keeled flattened scales, the largest marking the posterior row of
the vertebral segment; a ventrolateral fold covered with cycloid
scales; ventral cycloid-type scales reaching laterally to the outer
rows of tubercles.
Description of species (From KUMNH No, 34090 Tenorio, Las
Cafias, Guanacaste): Snout oval, the head widening behind eyes;
neck distinct; rostral 1% times as wide as high, lowest on median
60 Tue Universiry ScieENCE BULLETIN
line, the scale with an entrant suture from posterior border, equal
in length to half the height, bordered behind by a pair of supra-
nasals forming a median suture, the suture in a deep valleylike
depression; nostril surrounded by rostral, supranasal, first labial
and two postnasals, the upper large, differentiated; snout covered
Fic, 15. Phyllodactylus eduardofischeri Mertens. Left figure, KUMNH
No. 34090; actual total length, 135 mm.; right figure, No. 34101; total
length, 119 mm., (tail regenerated). Both from Tenorio, Las Cafias,
Guanacaste, C. R.
Review oF Lizarps or Costa Rica 61
with moderately large elevated juxtaposed tubercles, largest in
front of upper part of orbit; about 28 scales across snout between
the posterior end of the third labials. Scales on occiput granular,
intermixed with larger scales; sixth supralabial below middle of
eye, followed by two diminutive supralabials and three undif-
ferentiated granules; mental with a wider labial border than
rostral, narrowed, forming an angle posteriorly, and bordered by
a pair of chinshields; a row of five larger chin scales border chin-
shield, the second row with ten scales; seven infralabials followed
by two or three granular scales; scales on chin and front part of
neck small, rounded, juxtaposed; eye with spectacle; vertical pupil;
ear diagonally vertical.
Granules on dorsum and sides small, intermixed with large
trihedral tubercles arranged in about 14 irregular rows at the
widest part; the two median rows straight, separated by three or
four rows of small granules; below, beginning on back part of neck,
scales cycloid, flat, imbricating, about 33 across widest part of
venter and extending on sides to enlarged lateral tubercles, about
64 in a row from neck to vent. A ventrolateral fold on which the
flattened scales become more elevated. Scales on tail variable,
imbricating, intermixed with larger keeled scales, largest being
the posterior one of each series of whorls, indicating vertebral
Segments; a median series of transversely widened subcaudals
Paired at base of tail and then alternating larger and smaller, two
to each segment. On latter third of the tail alternate scales may be
Paired,
Dorsal and anterior aspect of upper arm and thigh, anterior and
ventral aspect of forearm and tibia covered with larger imbricating
scales; trihedral tubercles on outer face of forearm and upper
Posterior face of thigh; posterior part of upper arm and_ thigh
Covered with small granules.
Digits unwebbed, palm and sole covered with imbricating scales
below, with usually transverse lamella or scales under digits, termi-
nating in a pair of widened lamellae separated by a groove; above
these widened lamallae, a sheath covered with specialized scales
into which the claw is withdrawn. The lamellae following the
terminal widened lamellae paired, 15 under longest toe.
Color: Above fawn, with a paired series of small dark-gray spots
or flecks. Other similar darker marks on sides, limbs, and head.
Posterior half of tail with about seven dark transverse bars growing
black toward tip and surrounding tail. Ventral surface of body, chin,
and limbs yellowish, Digits indistinctly banded.
62 Tue University ScieENCE BULLETIN
Measurements in mm.: Total length, 136; snout to vent, 64; tail,
72; width of head 14; length of head 18; snout to middle of eye, 9.5.
Variation: The length of the original tail varies in the young and
adults. In the youngest specimens with a complete tail, the head
body length is 36.2mm.; the tail 34mm. In an adult female the
same measurements are 64 and 72.
The supranasals are in contact mesially in all specimens, the com-
mon suture being in a valleylike depression, The mental, which al-
ways has a larger labial border than the rostral, extends back usually
to a line drawn across chin from points where the common suture of
the first two labials touch the chinshields. A few specimens may
extend to a point one millimeter in front or behind this line.
The scales on the regenerated tail are nearly uniform in character.
They lack the enlarged tubercular scales and are not arranged in
transverse series. The mesial subcaudal scales are usually shorter
but distinctly wider than those on the original tail. The dark and
light banding is absent, and replaced by small black spots, each
covering several scales, The frontal depression is more pronounced
in old specimens. I count 24 vertebrae in the tail.
The general dorsal coloration consists of two series of more or less
regularly paired dark-gray spots on a fawn ground color beginning
on the neck and continuing to the tail, A few specimens show a
more or less distinct median lighter line. The ground color may
have a lavender shade or it may be a dirty gray color. Below, the
chin, venter, and under surface of the limbs are light yellow. In
daylight the fawn areas are darker, nearly gray in color.
Remarks: The series of specimens taken at Tenorio (KUMNH,
Nos. 34090-34103) is remarkable, in that not a single male is rep-
Measurements of Phyllodactylus eduardofischeri Mertens. Tenorio, Las Canas,
Guanacaste
eo Tail Head Tead Ventral I ce Rows
hed x length width scales ya te
vent eyes tubercles
34090....... 64 72 18 14 33 23-+2 14
SaLOL yy ali hs 64. 55 19 14.2 32 22 +2 15
34098. [hfue mead Dre oeee aarr 19 14 32 24+2 14
34091 . 63 61 Led 13.2 32 24-+-2 16
34100. 63 67 17.4 14 34 2442 17
34097. 63 34 ee 14 32 23'+ 2 16
34094. 58 55 16 13 32 2242 16
34093... 59 55.5 17 12.8 32 23+2 16
34096... 59 59 17 12.8 34 2442 16
34095... 57 50.2 16 12.3 32 24+ 2 16
340984. 67 55 15.5 12 30. 2212 14
34092.. Ppa ing arate 13 9.2 ? ?
34103.. 36.2 28 12 8 30, 23 +2 16
84102..,.; BO ey arises 10 ve 31 24+2 i
Review or Lizarps or Costa Rica 63
resented, It suggests that males are either present in smaller pro-
portion than females or that they are more wary than females.
Most of the gekkoes were taken at night at a corral, bordered by
a high rock fence partly covered with cactus. Many other speci-
mens seen, escaped. Some specimens were taken on the walls of
an adjacent shed. Four young and one half-grown male (KUMNH
Nos, 35756-35760) were captured in daytime on boulders in an open
field between Tilaran and Los Angeles de Tilaran. The two eggs
produced are plastered on a rock, usually on the under surface
of a loose rock lying on top of another, or in a covered crevice. Only
the empty shells were seen.
Famity IGUANIDAE
Tguanidae Gray, Phil. Mag., ser, 2, vol. 2, 1827, p.
This large and populous family, with the exception of certain
forms in Madagascar, and members on a few Pacific Islands, is re-
stricted to the Western Hemisphere. In our territory seven genera
have representatives. These are: Sceloporus, Ctenosaura, Corytho-
phanes, Anolis, Polychrus, Basiliscus, and Iguana. Most members
of the genera Sceloporus and Ctenosaura occupy territory north of
Costa Rica. The-other groups occur both north and south of this
country,
Key To IGUANID GENERA IN Costa RIGA
to
PH GmOte) poree Hisene Im NOL SEXES cc rity tha aia ae
Demoval pores present, Cistinot (eMAleS Ae yuy i bis ie se 4
Proximal part of digits with widened lamellae; distal part narrow;
Len EW ale) send neleleret Olen aetat tate nia tea monn etait ein ete 5 Anolis
Proximal part of digits not widened; infradigital lamellae keeled... .. . 8
Head with two ridges uniting posteriorly to form a high crest; toes
NOCWOMATRO Nah Oele: cares pene opiate ely Corythophanes
No head ridges; a dorsal crest on head and dorsum; sometimes a caudal
CUS LHe LOGS oO DELO ny ites, is nee aie aN ret lickers cee I hs Basiliscus
Third and fourth toes equal in length; tail very long, prehensile; body
strongly compressed; no head or body crest; a row of elevated scales
Ivamechint me OF CHIN ye oi kev tees ey eee Polychrus
Fourth toe longer than third; tail not prehensile................... 5
A large, nondilatable gular appendage; a large sitbeloeutes: sents below
tympanum; very large lizards (5 ft.)........ Caen ee Ce be Iguana
‘i large gular appendage or sac; no large scales below the tympanum, 6
A dorsal crest; body not or scarcely compressed; large arboreal lizards
(COREG nee iy cre r Cn eet tan eae Ctenosaurus
No dorsal crest; body OE ae or somewhat flattened; small lizards (5-7
tok yeep ee Urata ae ay Via tea sas ......Sceloporus
J
~
64 Tue Universiry ScreNCE BULLETIN
Genus ANnouis Daudin
Anolis Daudin, Histoire naturelle générale et particuliére des reptiles, vol. 4,
An. X (1802), pp. 50-55.
Description of the genus (From Boulenger, 1885): “Tympanum
distinct. Body more or less compressed, cylindrical, or feebly de-
pressed, covered with juxtaposed or imbricate scales or granules.
Dorso-nuchal crest slightly developed or absent. Male with a gular
appendage. Digits more or less dilated, inferiorly with smooth
transverse lamellae, the distal joints slender and raised above the
basal ones. No femoral or preanal pores. Tail long, cylindrical or
compressed, not prehensile. Lateral teeth tricuspid; pterygoid teeth
generally present. No sternal fontanelle. Abdominal ribs.”
The genus as now accepted is one of the most populous of all
reptilian genera. It is confined to tropical and subtropical areas in
the Western Hemisphere only one species reaching the United
States. The presence of so large a number of species (more than
20 in Costa Rica) suggests strongly that Anolis is a composite genus.
Jertain species formerly in the genus have already been separated
into acceptible genera of their own. Other generic or subgeneric
names have been proposed that are not at the present accepted.
The problem of making a key for species of the genus Anolis has
its difficulties, since the variation in various populations of a species
may be considerable. I submit the following key based on usual or
average external characters, believing that it will be an aid
in identifying most Costa Rican species. However, once the pre-
sumed identity has been established by use of the key, it should
be verified by comparison with the description here given, as well
as with other descriptions in the literature, if available.
Variation in color pattern is great, with the possible exception
of that of the dewlap (throat fan). This usually has a fixed pattern
until preserved. Then even the color and markings of this area
may be lost. In using size as a key character, it is well to remember
that the young or immature specimen of a large form may fall into
the category of the lesser species, in which case, if you are in doubt
as to age, try the comparable larger category.
Kry TO THE SpeciES OF ANOLIS IN Costa Rica
1, Ventral scales smooth, unkeeled........... eee ae eee 2
Weave ee eC cs ieee Bice Wee ee ie rinks Ba a's 9
2. Large lizards at least 75 mm.-145 mm. snout-vent length.......... 3
Small lizards not more than 55 mm. snout-vent length............. 6
oO
Review oF Lizarps oF Costa Rica
No ridge bordering an occipital depression; four subcaudal scale rows
enlarged, each with a longitudinal, somewhat serrate keel; one
dorsal serrate keel; tail compressed; dorsals small, unkeeled; a small
nuchal crest, and a large cherry-red dewlap in males; ear small,
double size-ob nostilie7S mime: oy eo i pentaprion
Ridges bordering two sides of an occipite al ‘depression not meeting
posteriorly in midline; two enlarged subcaudal scalerows with
Strong keels} one. dorsal: caudal keel. 3. ely. els.
Ear very small, about size of nostril; dorsal, lateral, and ventral scales
small, subequal in size; hind leg reaches jaw angle; 108 mm.,
microtus
Ear larger, three, to many, times larger than nostril; dorsal, lateral,
Gnd Ventralscnles not subequal im SiZe 0 i.e.
A trace of two frontal ridges, not connecting with the circumorbital
semicircles; dewlap very large, white or cream in color; medial
dorsal scalerows keeled; ear-opening nearly equal to eye-opening;
leg reaches in front of eye; ventral scales twice size of dorsals;
Wate Mm Ae) he ee ee a es Ae a rie purpurescens
No trace of frontal ridges; dorsal and lateral scales rather smooth ex-
cept for four somewhat enlarged median rows which are sub-
conical; leg reaches almost to ear; ear-opening about half size of
eye-opening; dewlaps moderate; 145 mm............. insignis
Circumorbital semicircles separated by one scale row; outer postanal
scales greatly enlarged; two suborbitals in direct contact with supra-
libel occipital scale twice (or more) size of auricular opening,
separated from semicircles by three scale series; 40 mm.,
biscutiger
Circumorbital semicircles separated by two or more scalerows; post-
BIBS Werle bles so Ca
Head longer than tibia; leg ** reaches to “potnt be tween eye and nostril,
Head a little shorter or equal to tibial length; leg reaches end of snout
or a little beyond; head scales keeled, four scalerows between cir-
cumorbital semicircles; the scales of semicircles not larger than ad-
joining scales separating them; occipital scale smaller than ear;
postanals not enlarged; 47 mm. pachypus
Circumorbital semicircles separated by three scalerows; suboculars
rather large, in contact with supralabials; dewlap small, dull
greenish white; occipital scale about equal to ear; smaller, 40 mm.,
limifrons
Semicircles separated by two or three scalerows; suboculars sepa-
rated from labials by one scalerow; dewlap larger, bright orange;
occipital a little longer but less wide than ear; separated (usually )
by three scalerows from semicircles; larger, 55 mm... ... . polylepis
Distinct axillary pockets present; dorsal scales larger than ventrals
No distinct axillary pockets, although an axillary depression may or
may not be discernible; dorsal scales variable
*
* Measurements given are snout-vent feats
“(Except in trochilis believed to be a synonym of limifrons. )
—~I
THE UNIVERSITY SCIENCE BULLETIN
10. Axillary pocket smaller with an opening about as wide as the depth
of pocket; the walls of pocket not scaled or pigmented; size smaller,
AQATHIM Mess oe Mn penatch ele itched sepa e teeter s
Axillary pocket very large, more open, its diameter usually as great
as its depth; walls of pocket partly scaled and pigmented; 12-14
dorsal scalerows enlarged, the two median larger than the adjoining
rows, all however equal to or (mostly) larger than ventrals; low
nuchal crest; size larger, 44 mm.............. humilis marsupialis
11. Dorsal eight or ten scalerows enlarged, keeled, the two median rows
or parts of the rows, usually smaller than adjoining rows, without
trace of dorsal or nuchal crest; dorsal scales considerably (one third
or more) larger than ventrals; dewlap deep red orange or red,
edged with yellow, without a purple spot at base; 38 mm.,
humilis humilis *
Dorsal eight or ten scalerows about equal in size to ventrals; a bright
red dewlap with a purple spot at base; 40 mm.,
humilis uniformis **
12. Hind leg reaches to a point between front of eye and tip of snout... .
Hind leg reaches to a point between shoulder and eye...........
13. Head scales somewhat multicarinate; tail about twice snout-vent
Lcsnkedsln eet, Ore PRs Sotto USP ker aa aca wee Den Toor dR SING einer TN gi ENG COINS aie
[ead scales largely multicarinate; tail not twice snout-vent length,
Hot Cistinctly compressed 2 6.5 cance ers ek eh ens
14, Head high, shorter than tibia; 16-18 dorsal scalerows generally hex-
agonal, juxtaposed, smooth, the median rows not larger than
others; no nuchal crest in males; a large axillary depression;
posterior supraciliary region with two or three rows of larger
scales; upper temporal scales larger than lower temporals; semi-
circles keeled or roughened, separated normally by two scalerows;
about 108 scales around body; tail about double snout-vent length;
dewlap smalls: 90 int, fyi08 14.0 Miers wy eee ees capito
Head, not high relatively, its length equal to or longer than tibia;
tail 2 2.3-2.4 times snout-vent length, compressed; scales not hex-
agonal; an axillary depression; a small nuchal crest in males; upper
and lower temporal scales subequal. il)
Dewlap large,-dark olive to blackish, with magenta scales; body lz argely
brick-red with blackish markings; about 140 scales about body;
scales bordering occipital not especially enlarged; 83 mm.,
woodi attenuatus
Dewlap large, pink orange on edge, then a band of amber-yellow,
while at base an area of bluish white; many scales on dewlap
whitish; body generally olive, with indistinct rusty spots; about
125 scales about body; occipital bordered by two much enlarged
BCulege Vo Mit, seis ey Wy Late Way Grysey Phones woodi woodi
ci
al
13
15
* Boulenger’s (1885, p. 83) description of Anolis humilis is based on h. uniformis from
Guatemala.
** Anolis quaggulus from Nicaragua may represent a population separable from humilis
humilis, and humilis uniformis and it may actually be the proper name to apply to this
population. This cannot now be decided by me.
Ne
18.
21
Review or Lizarps or Costa Rica
Suboculars separated from labials by two scalerows; semicircles com-
posed of relatively small scales, a part of which are multicarinate,
the series separated by from five to seven scales; no enlarged post-
anals; occipital smaller than ear, separated from semicircles by
five or six scales; leg reaches nostril; 148 scales around body;
DR MNT ee, on ee Ue lee Ct Ne ee EL Lae env te curtus
Suboculars larger, two in contact with labials; semicircles of larger
scales multicarinate, separated by from three to five scales; occipi-
tal about one third of the size of ear, separated from semicircles
by four scales; no enlarged postanals; leg to anterior border of
orbit; about 160 scales and granules around body; 50 mm.,
tropidolepis
Occipital scale surrounded by relatively minute scales, separated
from the semicircles by from six to nine rows; two median scale-
rows enlarged, keeled, larger than adjoining rows, but some lateral
scales may equal the median in size; tail compressed, serrate above,
banded; suboculars separated from labials by two scalerows; occipi-
tal about half as large as ear; about 140 scales around body; 65
PID ie plone waver Le A ed Cee Heats vee ac ecw NRA ELL 5 ee ER ee aquaticus
Occipital scales not especially small, separated from semicircles by
four or less scales; suborbitals touching labials or separated by
Oly OMG TOW: OLIRUAIESY Jy wats soy a ons seas wee a es
Dorsal scales enlarged in 20-24 rows, usually hexagonal or juxtaposed
not or scarcely keeled; largest about equal to ventrals; tail dis-
tinctly compressed, not or but dimly banded; postanals enlarged;
a shallow axillary depression; occipital scale about equal to ear;
about 125 scales around body: 75 mim) is. lionotus
Dorsal scales not hexagonal, juxtaposed, smaller than ventrals; tail
MOU GOmpressed: postatials VANADIC; fv..55 issn eeu is ices
Very large species (102 mm, snout to vent); longest toe to near ear;
dorsals small, keeled, ridged or moundlike; the two or four median
rows discernibly enlarged; semicircles well defined, nearly smooth,
separated by two scalerows; chin usually streaked with brown;
body green in life; about 100 scales around body; 102 mm.,
biporcatus
Smaller species 41 mm. to 63 mm, all with more than 100 scales
about body, none green in life, none with chin streaked with
MEOW eee eck) er Ba ees Sa Soe Gee
Heel with enlarged scales two to three times larger than surrounding
scales; seven or eight median scalerows slightly enlarged, none
half size of ventrals; leg to ear; about 130 scales and granules
around body; occipital scale two and one-half times size of ear-
Opening; semicircles separated by one scalerow; head _ scales
Neer ai: Mine er eee es Ge eats fae achilles
No especially enlarged heel plates; dorsal scales and other characters
VATIOIDIGe secriginsae srr viotice harmless MRA cst we aay SETS
Rostral touching anterior nasal scale; suborbitals touching labials;
mental divided by a groove; ventral scales equal to or larger than
19
20
21
Tue Universrry SCIENCE BULLETIN
the two largest median dorsals; about 112 scales around body; small
nuchal orest? Cocos Island; 69. mim, 01) ero ee, townsendi
Rostral separated from nasal by at least one scale; other characters
VETTE eek eee eh gh SEE asinine ely 22,
22, Scales of semicircles flattened, unkeeled but sometimes with some
mugosities: small dewlapes. 0) ee. ou Wipe pe tes 23
Scales of semicircles with single or multiple keels; dewlap large in
Cipretigy imino Wi a ae en ee Pe ed 24
23, Smaller; two or three suboculars touch labials; no frontal depression;
four or five scales between anterior canthals; four loreal series;
two rows of median enlarged temporals; four or five scales between
upper nasals; largest dorsals nearly size of ventrals; leg to between
shoulder and ear; around body about 126 scales and granules; semi-
circles in contact mesially; 33 lamellae and scales under fourth
foes 202M ee Leg eee ie hie a. une intermedius
Larger; suborbitals separated from labials (or in contact at one point);
eight or ten scales between anterior canthals; semicircles some-
what elevated, separated by one scalerow; rarely a trace of keels;
shallow frontal depression; 38 scales and lamellae under fourth
toe; about 160 scalerows around body; 63 mm.,
lemurinus lemurinus
24, Two faint prefrontal ridges converging anteriorly; head scales uni-
carinate or flat to concave; semicircles separated by two scalerows;
large supraoculars touch semicircles; six loreal rows; occipital larger
than ear, separated from semicircles by two scales; no prefrontal
Hesse ie witty 16a a pa Lage PEER Sa aia; . .altae
Dewlap attachment extending to middle of abdomen; semicircles sepa-
rated by two scalerows; occipital scale double size of ear, separated
from semicircles by three (often) tricarinate scales; small; 42
rites ee A TAGE ear ee cupreus
Anolis microtus Cope
(Fig. 16, figure on upper right)
Anolis microtus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1871, p. 214 (type
locality, San José, Costa Rica, Dr. Van Patten, coll. U.S. N. M. No. 31,282);
Journ. Acad, Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 119, pl.
24, fiz. 2; Boulenger, Catalogue of the lizards in the British Museum 2n
ed., vol. 2, 1885, p. 62 (Irazi, Costa Rica, 1 male and 1 female, F. D. God- {
man, coll.); Dunn, Proc. New England Zool. Club, vol. 12, Aug. 7, 1930, I
p. 21 (reference to type); Barbour, Bull. Mus. Comp. Zool., Harvard Col.
vol. 77, no. 4, June 1934, p. 142 (“Another of the big rainforest forms which
we have from La Palma, Costa Rica, collected with Anolis insignis”); Dunn,
Proc. New England Zool. Club, vol. 15, Jan. 1, 1937, pp. 6, 8 (Boquete,
Panama, 4500 ft. elev., 2 specimens, color changes given).
Diagnosis: A large species, reaching a snout-vent length of 108
mm.; body triangular in cross section; ear small, scales of supraor-
bital area smooth; scales of the interorbital ridges elevated, rough,
separated by two rows of flat scales (one single scale of a third row);
a pair of indistinct ridges on snout; a deep occipital depression
bordered by occipital ridges; mental partly divided, bordered by
Review oF Lizarps or Costa Rica 69
Fic. 16. Upper left, Anolis oxylophus Cope, total length, 217 mm. Upper
middle, Anolis insignis Cope, actual total length, 440 mm. Upper right, Anolis
Microtus Cope, total length, 310 mm.; lower left, Anolis pachypus Cope, total
ength, 185, Lower right, Chalcidolepis metallicus Cope, total length, 165.
types, from Cope (1875), pl. 24.
70 Tue Universtry SCIENCE BULLETIN
four scales; ear minute (.8 mm. in diameter); hind leg reaches to
5 mm. behind ear; dorsal scales smooth, not or but little smaller
than smooth ventrals.
Description of species: Body somewhat triangular in cross
section, with a rather sharp dorsal edge; scales of the interorbital
rings elevated, very rough, with transverse ridges or tubercles; sep-
arated by two rows of flat scales (three at one point) that like-
wise have some roughened tubercles; canthal series not forming
a sharp edge, but are roughly rounded, extending close to rostral;
snout with two indefinite ridges; frontal depression marked, the
scales covering it smooth; occipital depression bordered by rela-
tively high ridges, which do not unite posteriorly, but leave a notch;
the surrounding scales small, rough; rostral moderate, much wider
than high, its labial border less than that of mental; mental with
a suture entering from behind and extending somewhat more than
halfway across scale; mental bordered behind by four scales; six
rows of scales between the nasal rings; large supraciliaries absent;
supralabials, 9-9, low, smooth, not separated from the subocular
series; three or four scale rows in the loreal series; nine infralabials
and two or three rows of somewhat enlarged smooth sublabials,
the two anterior largest; supraocular area rather small, with one
or two larger scales surrounded by two rows of five somewhat en-
larged scales, and these by three rows of minute scales all of which
are smooth; auricular opening (.8 mm.) not larger than nostril;
seven scales between canthi at middle of muzzle; a rather promi-
nent temporal ridge.
Limbs rather short and stout; fingers reaching the tip of snout
and posteriorly not quite reaching groin; longest toe reaching to
near jaw angle (5 mm. back of auricular opening); the lamellae
under the penultimate joint of digits distinctly widened.
Scales of back and sides small, pavementlike, equal (20 in a dis-
tance of 10 mm.), not or scarcely larger than ventrals, all unkeeled;
tail depressed at base, cylindric distally; “scales on tail and foreleg
three or four-keeled”; no enlarged postanals; throat fan large, ex-
tending 8 millimeters behind the level of arm insertion.
Color; Generally a rich yellow-brown; a deep-brown band cover-
ing eye extends to shoulder, where it is marked with pale-centered
ocelli. It is separated above by a narrow paler band from a large
dark-brown patch that covers the nape and scapular regions; cross
bands of large paler ocelli crossing sides behind axilla, at middle
and on rump; ocelli of middle band distinctly larger than others;
Review or Lizarps oF Costa Rica cal
limbs and tail banded with brown; belly and throat whitish, un-
spotted,
Measurements in mm. (Type)
Motel lengtheiy ire ss Cs eens 810
BOULOGNE cee aan ae seer aie ea en 108
MHOUL HOMOMD ete oe ea Pes 15
SHOU tO OH 1a ela ge es oy coe 30
SrOUb TOTO ce a as vey 95
Nh ee ar Soe aa ere a sn ce een cnnyes 202
WiHeltlisOl GAC oni: Senco ene ers Any
Teen Cth OP HGRG ec) ete oe
Remarks: 1 have seen only the type which is in a good state.
It is USNM No. 31282 from “near” San José, Costa Rica. It has
been suggested that the specimen may have been obtained at
higher elevation than San José. The description given here in-
corporates data from the type description and from notes made on
the type by me.
Boulenger’s specimen is from “Irazu,” presumably somewhere
on the voleano. Dunn has identified two specimens from Boquete,
Panama from 4500 feet elevation, as belonging to this species. I
have not seen these specimens, Dunn mentions that the dewlap
of his male specimen was pink with white scales.
Anolis pentaprion Cope
(Fig. 17)
Anolis (Coccoéssus) pentaprion Cope, Proc, Acad Nat. Sci. Philadelphia, 1862,
p. 178 (type locality, near Rio Truando, New Granada, Colombia).
Anolis pentaprion Cope, Proc. Acad, Nat. Sci. Philadelphia, 1874, p. 68 (Nica-
ragua [probably southern, since the specimen was collected on the Nica-
raguan Surveying Expedition of 1873]; “the most northern locality yet
discovered for this Anolis.” ); Giinther, Biologia Centrali-Americana; Reptilia
and Batrachia, Oct. 1885, p. 47 (Panama; Ecuador); Boulenger, Catalogue
of the Lizards in the British Museum, 2nd Ed., vol. 2, 1885, p. 64 (Guay-
aquil, Ecuador) ( étotal length, 170 mm.); Dunn, Proc. New England Zool.
Club, vol. 12, Aug. 7, 1980, p. 20 (Nicaragua to Colombia); Schmidt,
Smithson. Misc. Coll., vol. 89, no. 1, Mar. 16, 1938, p. 10. _(Bohio,
Panama [locality now covered with water fide Barbour loc. cit.]); Barbour,
Bull. Mus. Comp. Zool., Harvard College, vol. 77, no. 4, June, 1934, p. 145
(places sulcifrons Cope [Colombia] and panamensis Boulenger in. syn-
onymy); Wettstein, Sitzungsb. Akad. Wiss. Wien, Math.-natur. KI, Abt. 1,
Bd. 148, heft, 1-2, 1934, p. 22 (Porto Jimenez, Costa Rica); Gaige, Hart-
weg and Stuart, Occ. Papers, Mus. Zool. Univ. Michigan, No. 841, p. 10
(Rio Siquia 7 miles above Rama),
Diagnosis: A medium-sized species; body triangular in cross-
section, rugose, but unkeeled; the circumorbital scales forming
relatively low ridges, one scale of the series much larger than the
others; semicircles separated by a single scalerow; four labials in
contact with four suboculars; mental almost completely divided,
its labial border longer than that of rostral; dorsal scales small, not
72 Tue University ScreENCE BULLETIN
keeled; ventrals very slightly larger, unkeeled; a nuchal fold or
crest in males; two median dorsal rows somewhat enlarged, becom-
ing slightly elevated posteriorly; tail compressed with a low serrate
crest; two ventral subcaudal rows much enlarged, keeled; adjoin-
ing scalerow somewhat smaller, likewise with strong keels; lateral
caudal rows smaller; limbs with small scales, few or none keeled;
ear small, a little larger than nostril. Males with a large cherry-
red dewlap; 22 widened lamellae under fourth toe; about 186 scales
and granules around middle of body.
Description of species (From KUMNH No. 34254, Palmar, Costa
Rica): Head relatively long, much longer than wide, the inter-
orbital region a little more elevated than the flat occipital, or frontal
regions; rostral wide transversely, about three and one-half times
as wide as high, bordered behind by two labials and seven scales;
nasal scale separated from the first labial by one scale, from rostral
by two, from its fellow by eight scales; snout rather flattened above;
head scales rough but not keeled; enlarged scales of circumorbital
rings not strongly defined from other scales; behind orbit they be-
come more or less continuous with a row of post- and suboculars.
Mental much larger than rostral, with a longer labial border than
rostral, its width two and one-fifth times its height, its posterior bor-
der irregular, bordered behind by eight scales; supralabials, 12-18,
infralabials, 10-11, the posterior of each series much reduced; rows
of scales bordering infralabials enlarged anteriorly; four rather elon-
gate scales of suborbital series in contact with labials; larger supra-
ocular scales in about five rows, flat, pentagonal; four or five rows
of much smaller scales on outer part of orbit; occipital large, more
than twice size of auricular opening, separated from circumorbital
rings by two scales; three relatively small supraciliaries on anterior
supraciliary border; posterior border with two rows of scales dis-
cernibly larger than surrounding granules; auricular opening. tri-
angular, small; temporal ridge poorly defined; a small nuchal crest,
the two median rows slightly enlarged; scales on body all smooth,
unkeeled, scarcely differentiated in size, the dorsal scales very
slightly larger than laterals, but discernibly smaller than the small,
unkeeled ventrals; approximately 188 scales and granules in a
row around middle of body; body rather flattened, not compressed;
legs with relatively small scales, all smooth except a very few larger
ones on front of forearm and front of tibia, which may have low
keels; scales on dorsal surface of digits smooth; toes widened con-
spicuously except on distal part; lamellar formula for hand: 4, 14,
21, 25, 12; for foot: 8, 14, 25, 32, 15; fingers reach nostril but fail
mrQ
Review or Lizarps or Cosra Rica 13
Fic. 17. Anolis pentaprion Cope. KUMNH No. 34254,
Palmar, Puntarenas Province, C. R.; actual length, snout-to-vent,
71 mm.; total length, 163.
74 Tue Universiry ScreNCE BULLETIN
to reach groin; toe fails to reach ear-opening by six millimeters;
large throat fan (dewlap) of male reaches back slightly beyond
level of arm insertion, its width (distended) 20 mm. Tail com-
pressed, with the four enlarged subcaudal scalerows bearing four
strong serrate keels; no enlarged postanals; dorsal scales of tail
forming a keel.
Color: In life, grayish, generally with a wash of yellow on venter
and thighs; laterally an indistinct light line above shoulder, the
sides with numerous larger or smaller lighter whitish-gray areas;
arms and legs speckled with lighter above; very indefinite darker
dorsal areas and darker and lighter flecks evident on median dorsal
line; tail indefinitely banded darker and lighter gray; males with
a bright dark-red dewlap.
Measurements of Anolis pentaprion
INV Cc ee Type* 34254 34255
Grout. to Vets 1 ee es 63.52 75 WO!
a Ae a ee ee 84.68 97 92
Head: width: <7 ee ihe Lé Wey
Head lencth | tice oe 22 21.8
Snowe fo Cat yi rs 16.96 19 19
Snout to foreleg.) 03. aks nae $1.5 29
Axa tO CTO a eee os 81 29
ANNO ee a ee 28 25
DOO er ee ene corte) Iie 29.04 40.5 40
Rihana ee ee fa L5 15)
Variation: Cope states in the type description that the hind leg
scarcely reaches to the auricular orifice. In these specimens it fails
to reach the ear by several millimeters. Moreover, there are four
loreal rows of scales instead of three. As is typical of many species
of Anolis the color and markings vary considerably. The general
scheme of color and markings of No. 34255 is as follows: general
color above dark gray, with a dark indefinite blotch at nape, one on
the shoulder, one across the back behind shoulder and some other
irregular smaller dark areas on the posterior part of back. A median
series of small fawn marks on the median line. The sides with
numerous fawn spots, some subcircular, others more elongate. The
tail is indistinctly banded beyond its basal widening. The head is
variegated dark gray above, and the labials have light gray mark-
ings. The auricular region is light gray and some indefinite darker
areas are on the sides of head. The chin is a dirty gray while the
venter shows considerable pigment. The under side of arms and
limbs and under surface of the base of the tail is gray-white. The
* The type measurements were given in inches and lines, These are here changed to
their metric equivalents.
Review or Lizarps oF Costa Rica ffs)
widened lamellae become darker distally and the limbs have indefi-
nite light brownish marks. The throat fan is cherry-red, while most
of the individual scales on its sides are orange.
The bright coloration of the throat fans of anoles usually disap-
pears largely in formalin or alcohol; but in this species, after nearly
a year of preservation in formalin (then transferred to alcohol),
the red coloration has been retained in its entirety,
Remarks: Two male specimens were captured low on the bole of
a tree near the bank of Rio Diquis at Palmar. No others were seen.
I have examined a specimen, USNM No. 67349 from “Colombiana
C. R.” and two in Harvard, one from Barro Colorado, C. Z. and one
from Balboa, C. Z. These seemingly agree with my specimens in
essential characters.
I suspect that this form is most closely related to the preceding
microtus,
Anolis purpurescens Cope
(Fig. 18)
Anolis ?? reticulatus Cope, Proc. Sete Nat. Sci. Philadelphia, 1862, p. 356
part. (Truando, Darien, 4321, Turbo 4313),
Anolis purpurescens Cope, The Philadelphia ee Sci. Bull., No. 1, May
26, 1899, pp. 7-8 (type Ha ao T ruando River, New Granada [= Colombia],
US: N. M. No. Cope states “The only specimen of the A. pur-
purescens which I tee seen is No. 4821 . .” thus suggesting that
No. 4318, noted supra, re of bana species Jp Barbour, Bull. Mus. Comp.
Zool., Harvard Col, vol. no. 4, June 1934, p. 147, part. (“common on
Barro Colorado. Island.” Buca (infra) believes specimens regarded as
purpurescens by na ig are really frenatus ); Dunn, Proc. New England
Zool. Club, vol. 16, Jan. 1, 1937, p. 9 (“still known only from type”).
Anolis longipes Burt and Burt, Trans. Acad. Sci. St. Louis, vol. 28, no. 1, June 1,
1933, p. 18 (part.).
Diagnosis: A very large species of Anolis, snout-vent length, 125
mm,
eo)
a)
>
Fic. 21. Anolis polylepis Peters. First figure, KUMNH No. 34351, Palmar, C. R.; actual length, snout to
vent, 48 mm., total length, 127 mm. (tail regenerated); second figure, 40666, Agua Buena, Cafias Gordas, actual ee
length, snout to vent, 51.5 mm., total, 158 mm.; third figure 34354, Palmar, actual length, snout to vent, 49 mm.,
total length, 154; fourth figure 40658, Agua Buena, Cafias Gordas, actual length, snout to vent, 47.5 mm., total
length, 139 mm.; all from Puntarenas Province, C. R.
90 Tur Universiry SCIENCE BULLETIN
Color in life: Nearly uniform light olive brown above and on
sides, cream-white on ventral surface. Dewlap bright orange; tail
brownish above and below; under a lens almost every granule
has a tiny black spot; legs with dim bands of brown; an indefinite
dark line passes above arm and continues on the side for some
distance; a few darker flecks on chin and faint indication of radiating
lines below eye; tail nearly uniform fawn color.
Measurements in mm.: Snout to vent, 52; tail, tip regenerated, 92;
snout to ear, 13.4; snout to arm insertion, 21.2; axilla to groin, 20.8;
head length, 15.8; head width, 8; arm, 23; leg, 43; tibia, 14.6.
Variation: Color variation occurs, as is to be expected. Many of
the females have a lighter median stripe which may be generally
fawn with a median darker center and bordered laterally by a broad
band of deep brown which begins behind the eye. A few speci-
mens have a series of blackish brown median spots connecting with
the brownish lateral stripe by scarcely discernible brown chevrons.
A black spot is often present on the neck, and the tail is more or
less distinctly banded. In some females the median stripe may be
brownish black bordered laterally by narrow fawn lines. In such,
the dark brown lateral line is present and the tail is dark brown
without bands. One female with brown dorsal spots has a series
of rounded gray spots nearly white-edged alternating with the
brown spots. This continues on to the tail.
Many of the specimens show a dark, rather indistinct band across
the head and many of the females have a rather heavy ventral pig-
mentation.
Remarks: Dunn, loc. cit., suggests that the species is related to
Anolis pentaprion. However, the reduced limbs, the presence
of a nuchal crest, fewer loreals, reduced ear-opening, the contact
of suboculars and labials, the very different character of the “penta-
prion” tail all point to a form very widely separated from polylepis.
Moreover, the ventral scales are in about fifty rows while in poly-
lepis the number is less than half this number. The character of
the superciliaries are quite unlike those of polylepis and the dorsal
squamation of head, especially of the supraorbital region, differs
almost as much as can be found between two species within the
genus.
Specimens usually found on low shrubs, are easily taken.
All the specimens in the collections at hand were obtained in
southwestern Costa Rica. The following localities are represented:
Puntarenas Province: KUMNH Nos. 34348-34358, Palmar; 34400-
Review or Lizarps or Costa Rica 91
34408, Esquinas Forest Reserve between Palmar and Golfito; 40648-
40671, Agua Buena, Cafias Gordas. San José Province: Nos.
34359-34371, San Isidro del General and vicinity (including points
10-15 km WSW).
Anolis humilis Peters
This species is represented in Costa Rica by three recognized sub-
Species. The most common form h. humilis occupies territory on
the northern and eastern lowland areas, ascending to some 4000
ft. elevation at least at some points; h, marsupialis, here described
4s new, is seemingly confined to the southwestern lowlands, while
the form here identified as h. uniformis is from the northwestern
lowland area.
With the exception of Anolis limifrons, A. humilis is the species
Most frequently encountered in forest. Limifrons however, is less
confined to forested areas and in the abaca plantations at Bataan it
Was especially numerous. In the same place humilis was rare save
in areas where there were forest remnants standing nearby.
Kry ro Susspecirs or ANouis Humiis tin Costa Rica
its
Dewlap bright red, with a purple spot in its basal part; eight or ten
dorsal scalerows enlarged, about equal to ventral scales in size;
head scales of semicircles nearly undifferentiated from the scales
adjoining them; no nuchal crest in males: axillary pouch rather
SiiallaO ints es ce ee humilis uniformis
2. Twelve to fourteen enlarged dorsal scalerows, the two median rows
larger than adjoining rows, and equal in size to ventral scales; a
low nuchal crest and often a trace of a dorsal crest in males;
lateral scales small with distinct keels pointing upwards and back-
wards; axillary pouch large, the sides of pouch at least partly
scaled and pigmented; size larger, 44 mm... . humilis marsupialis
Dorsal eight or ten scalerows enlarged, the two median rows usually
smaller than adjoining scales; larger dorsal scales one third (or
more) larger than ventral scales; no dorsal or nuchal fold or crest
in males; axillary pouch smaller, the side walls not scaled or pig-
IMemteds sie sinallen soumitny . sy 703, ek humilis humilis
Anolis humilis humilis Peters
(Fig. 22)
Anolis humilis Peters, Monatsb. Akad, Wiss. Berlin, 1862 (1863), pp. 138-139
type locality, Veragua, Panaméa, collected by von Warszewicz); Bocourt,
fission Scientifique au Mexique el dans lAmérique Centrale; Btude sur les
reptiles, livr. 2, 1873, pp. 105-106, pl. 13, fig. 7, 7a; pl. 16, fig. 31; Cope,
U.S. Nat. Mus. Bull. 32, 1887, p. 30 (listed); Boulenger, Catalogue of the
lizards in the British Museum (Natural History), 2nd Ed., vol. 2, 1885,
92. Tur Universitry ScreNCE BULLETIN
pp. 82-83 (Verapaz, Guatemala; probably Anolis humilis uniformis); Dunn,
Proc, New England Zool. Club, vol. 12, Aug. 7, 1930, pp. 16, 22, 24.
?Anolis quaggulus Cope, Proc. Amer. Philos. Soc., vol. 22, July 29, 1885, p. 391
(type locality, Rid San Juan, Nicaragua ).
Diagnosis: A small terrestrial species living in forest, maximum
snout-vent size about 38 mm.; a deep axillary pocket, its walls
without scales; dorsal eight or ten scalerows enlarged, keeled, the
two median usually a little smaller than adjoining rows, without
trace of dorsal or nuchal crest in males; dorsal scales one-third, or
more, larger than ventrals; dewlap red or red-orange edged with
yellow, without a purple spot at base.
Description of species (From KUM NH No. 40960, Suretka, Limon
Prov. Costa Rica): Rostral with a narrower labial border than
mental, about four times as wide as high, bordered by two supra-
labials and eight postrostrals; two scales between rostral and nasal
ring; eight carinate scales between nasals; a slight frontal depres-
sion; circumorbital semicircles somewhat larger than other head
scales, somewhat elevated anteriorly, separated anteriorly by three
scales, mesially by two scalerows; no occipital or interorbital de-
pression; five or six enlarged supraoculars, two scales usually larger
than others, but all separated from semicircles by an irregular row
of smaller scales; head scales keeled, the edges of many scales
somewhat upturned causing the scales to appear tricarinate; four
canthal scales lying somewhat diagonally, their keels not forming < |
continuous line; two anterior supraciliaries much elongated, over-
lapping; posterior supraciliary region with small granular scales;
occipital very small, about one fourth to one fifth of the size of ear,
and surrounded by scales many equally as large or larger than the |
occipital itself, separated from the semicircles by three scales; }
loreal scales in six or seven rows; scales of upper temporal region
larger than lateral temporals, two enlarged rows of temporals be-
gin behind eye; eight or nine supralabials at least part of them
keeled; suboculars separated from the supralabials by a row of
small scales, but they may be in contact at a single point; about
seven infrabials (on one side scales fused); five scalerows bor- |
dering the infralabials enlarged; mental partly divided by a groove :
and bordered behind by two infralabials and six irregular scales.
About ten dorsal scalerows enlarged, keeled (sometimes. tri-
carinate ), most of them larger than ventrals, the scales of the two
median rows very often smaller than those in adjoining rows; lateral
scales granular; no dorsal or nuchal crest in males; about 84 s rales
WON] WLSOr) AO SQUVZITT AQ AAATAAYT
Fic. 22. Anolis humilis humilis Peters, First figure, KUMNH No. 40882 (with ovarian eggs) actual length,
snout to vent, 39 mm.; second figure, 48852 2; third figure, 40847; fourth 40859¢: fifth figure, 408234; actual
length, snout to vent, 36 mm. All from Moravia de Chirripé, Limon Province, C, R.
6
94 Tur Universtry SCIENCE BULLETIN
and granules about middle of body; caudal scales large, keeled, the
median dorsal row only a little larger than adjoining scalerows and
not or but slightly serrate; two subcaudal rows enlarged with con-
tinuous keels; no enlarged postanals.
Scales on dorsal part of upper arm and the front and dorsum of
forearm enlarged but smaller than dorsals on body; scales on front
and upper side of thigh, and those on front and upper surface of
tibia large, some as large as dorsal body scales; lamellae under
proximal joints of toes only slightly widened; scales and lamellae
under fourth finger 19; under fourth toe, 25; hind leg reaches to
middle or front of eye. Dewlap moderate, distended, its depth
a little less than head length.
Color: Above and on sides of body rather dull brownish olive;
venter light flesh with some pigment; yellowish-white line under
tail, throat fan reddish with a yellow border; supraorbital and
interorbital area a little darker than remainder of head; some small
yellow spots on back of thigh.
Measurements, in mm.: Nos. 40960 g¢ and 40961 ¢ : Snout to
vent, 33,37; tail, 58, 55; total length, 91, 92; snout to arm insertion,
15.3, 15.1; axilla to groin, 12.6, 15; length of head, 11, 11.2; width of
head, 6.2, 6.8; arm, 15, 15; leg, 26, 26.5; tibia, 9, 9,2.
Variation: In the large series of this species available to me there
is considerable variation. The largest specimen measures 40 mm.
in snout-vent length. The character of the dewlap remains nearly
constant as to size and color.
In color and pattern there was a marked range, some specimens
being dark olive above and rather creamy or whitish below; others
were dirty brown and rusty olive below. In females there were
often somewhat distorted dorsal chevrons, or distinct spots, some-
times large, sometimes small; some specimens had the dark mark-
ings arranged longitudinally in two or four rather indistinct lines.
None of the females seemed to develop a median cream stripe such
as occurs in many species of the genus.
Remarks: This species described from Panama, is one of several
forms having the strongly developed axillary pocket, which has
seemingly been overlooked by previous students of the group.
There are several species or subspecies recognized beginning with
tropidonotus and uniformis in the north,* perhaps quaggulus in
* Smith and Kerster have recently described a form, Anolis compressicauda Ss Oaxaca,
Mexico, related to tropidonotus and uniformis. Herpetologica, vol. 11, 1955, pp. 193-198,
figs. A and
Review or Lizarps or Costa Rica 95
Nicaragua, humilis in Costa Rica and Panama, and marsupialis in
southwestern Costa Rica (perhaps also in Panama).
In the Costa Rican material of humilis I find a number of female
specimens with dorsal markings suggesting those of quaggulus.
However Cope’s description of that species mentions a large throat
fan showing its male sex. The golden green color and the vertical
lateral lines dividing the sides into separate areas, and the yellow-
ish head, suggest that-it may represent a separable population,
Since the type locality is Rio San Juan, on or near the Costa Rican
border, such specimens should be looked for in the northern part
of Costa Rica. Cope described quaggulus in the same paper with
uniformis and states that it is allied to that species and to tropi-
donotus Peters.
The name humilis is well chosen since it is indeed an earth anole.
I have found it on the ground, in piles of trash or brush, on low
plants, or on fallen logs. To escape, it seeks refuge on the ground
rather than ascending to higher elevation.
The functional significance of the axillary pouch is difficult to
explain, It is ample enough to hold ten or fifteen larval chigger
mites. Certain pouches contain as many as ten; others may contain
a smaller number, while some lack the parasites.
Four specimens harboured, below the skin, larvae of an insect still
unidentified. These formed conspicuous cysts:
Specimens in the KUMNH collections are from the following
localities: Heredia Province: Nos, 40779-40791, 40897-40900, 40933,
Isla Bonita (Cinchona); 40901-40904, Cariblanco. San José Prov-
ince: ?40926, La Carpintera Mt.; 40927-40929, Tapanti. Cartago
Province: Nos. 40775, 40798-40800, 40819-40822, 40905-40908,
40930-40931, Turrialba; 40879-81, Tuts; 40777, Santa Cruz; 40776-
40778, Morehouse Finca, 4-5 mi. from Turrialba; 40793, Navarro.
Limén Province: 40792, 40920-21, Los Diamantes, Guapiles; 40794-
40797, 40823-40878, 40882-40888, Moravia de Chirripé; 40801-40818,
Bataan; 40909, Limén; 40910-40916, Mountain Cow Creek; 40933-
40949, 40952-40958, 40963-40965, Suretka; 40959-40961, Volio (near
Suretka ys
Anolis humilis uniformis
Anolis uniformis Cope, Proc. Amer, Philos, Soc., vol, 22, no, 4, July 29, 1885,
pp. 391-392 (type locality, Guatemala and Yucatén (restricted to 2 mi.
N Santa Teresa (EI Petén, Guatemala), by Smith and Taylor); Boulenger,
Catalogue of the British Museum, vol, 2, 1885, p, 84; Stuart, Occ, Papers
Mus, Zool, Univ. Michigan, No, 464, 1942, Dp, L-2.
Anolis humilis uniformis Stuart, Misc, Publ, Mus, Zool, Univ, Michigan, No,
69, 1948, p. 48; Smith and Taylor, U. S. Nat, Mus, Bull, 199, 1950, p. 60;
and Univ. Kansas Sci, Bull., vol, 33, pt. 2, no. 8, Mar, 20, 1950, p. 818,
96 Tue University SCIENCE BULLETIN
Anolis ruthveni Stuart, Occ. Papers Mus, Zool, Univ. Michigan, no, 310, 1935,
pp. 1-4 (type locality 2 mi. N Santa Teresa, El Petén, Guatemala).
Diagnosis: A small form 40 mm. or under in snout-vent length; a
well-developed axillary pocket; eight or ten dorsal scalerows en-
larged, the two median smaller than adjoining scales about equal
or larger than ventral scales; about 100 s rales and granules around
body; scales of the semicircles a little larger or not than the adjoin-
ing scales, usually slightly elevated; occipital scale less than a
third of ear-opening, not larger than other scales in occipital area,
which occupy all the upper temporal region. Dewlap with a purple
spot on base of red.
Description (From KUMNH No. 40893, Tenorio, Guanacaste,
Costa Rica): Rostral very low its width more than five times its
height, touched posteriorly by two labials and seven postrostrals;
eight (or seven) scales between the tiny nasal scales; circumorbital
semicircles consisting of about eight scales, the two anterior largest,
separated from the canthal edge by a large scale, all with a single
strong keel and certain other rugosities, sepa rated mesially by three
scale rows; many anterior scales in the shallow frontal depression
and on the snout have the outer edges upturned suggesting a tri-
‘arinate condition; about four canthals, the anterior small the pos-
terior larger and contiguous with anterior of two large supraciliaries;
posterior supraciliary edge with only slightly enlarged scales; area of
occipital less than a third of ear-opening distinctly smaller than
scales in the outer occipital region; scales of upper temporal region
larger than those on lower part, seven sup ralabials, low, elongate,
keeled at lower edge, the posterior end of sixth below middle of
eye; eight infralabials; five rows of scales below infralabials enlarged,
keeled; mental wider but a little higher than rost ral, nearly severed
by a median groove, bordered by two labial and six postmentals;
suborbitals separated from supralabials by a row of scales; supra-
oculars in two or three rows of larger scales, separated from the
semicircles by a row of smaller scales.
Eight rows of dorsal scales enlarged, more or less keeled, the
two median smaller than the three scalerows on each side of them;
the row bordering these enlarged scales may itself be abruptly
larger than the elevated lateral granules; a pair of enlarged scale-
rows begins at occipital region, at first larger than anterior granules
of neck; in middle of nuchal region there is a small group of scales
usually discernible larger than surrounding enlarged series; ventral
scales scarcely more than a half as large as the larger dorsals, the
Review or Lizarps or Costa Rica 97
scales keeled, and elevated posteriorly suggesting a leaning pyramid
in profile; (no enlarged postanals in males); dorsal scales on arm
and leg enlarged, keeled, some of them approaching size of dorsals;
scales above digits, polycarinate; subdigital lamellae widened on
basal joints; lamellae and scales on distal joint 28 under fourth toe;
leg extended, the longest toe reaches orbit.
Color: Dark dirty olive brown with traces of two dark longitudi-
nal stripes bordering the outer enlarged scalerow; head olive, with
a trace of a line between eyes; venter brownish white; throat pinkish
in females (in males dewlap red with a purple basal spot).
Measurements in mm.: Snout to vent, 38; tail, 47; snout to ear,
10; snout to arm insertion, 15; axilla to groin, 16.5; length of head,
11; width of head, 7; arm, 15; leg, 29.
Remarks: Four specimens are in the collection from Tenorio,
Las Cafias, Guanacaste Province.
There is some question as to the wisdom of regarding these north-
western Costa Rican specimens under this name. With the excep-
tion of the curious coloration of the dewlap, there seems no char-
acter that cannot be duplicated in at least some specimen in the
series at hand that have been referred to humilis humilis.
I was unaware that the species involved such a problem and as
sometimes happens where the species is common, variable in colora-
tion and tiny, the proper field notes were not always taken. The
necessity for a complete review of this species from Mexico to
Panama is indicated,
Anolis humilis marsupialis* subsp. nov.
(Fig. 23)
Type: KUMNH No. 40893, collected about 15 km. WSW of San
Isidro del General along the Dominical Road, July 14, 1954; Edward
H. Taylor, Coll. (Probably in Puntarenas Province, or near the
boundary of this province in San José Province. )
Paratypes: KUMNH Nos. 40889-40892; 40894-40896, same data.
Diagnosis: Similar to humilis humilis but larger (to 44 mm.);
the enlarged dorsal scales in ten to fourteen rows, the two median
tows elevated into a suggestion of a dorsal crest and continuous
With a similar nuchal crest, and larger than adjoining scalerows;
dorsal scales about equal to or a little larger than ventral scales;
axillary pocket much enlarged, the opening nearly as great as its
depth, the side walls usually partly scaled; ventrals keeled.
98 Tuer Universitry SCIENCE BULLETIN
Description of subspecies: Rostral low, about five times as wide
as high, bordered by two labials and seven small postrostral scales,
separated from nasal scales bordering nostril by one scale; supra-
orbital semicircles well defined, the scales large, moderately ele-
vated, terminating in frontal area, separated between orbits by i,
three or four scalerows, anteriorly by four scales; the scales on the
row bordering the semicircles much larger than other head scales,
and equally as large as the scales composing the ridge; none or only
Fic. 23. Anolis humilis marsupialis subsp. nov. Left figure, KOUMNH No.
408899, actual length, snout to vent, 46 mm.; middle figure, 40892 ?, actual
length, snout to vent, 88 mm.; right figure, 40893 4, type, actual length, snout
to vent, 43 mm., tail, 68.2. All, 15 km. WSW San Isidro del General, (prob-
ably) in Puntarenas Province, C, R.
Review or Lizarps or Costa Rica 99
& vague occipital depression; a well-defined but shallow frontal
depression; two elongate supraciliaries reach back of level of the
middle of eye and are continuous with the canthal edge, which
Consists of four large scales; supraoculars in two rows, the inner
the larger, separated from the supraorbital ridges by a row of small
scales; most dorsal head scales with one or (more usually) three
keels or ridges, the outer ridges formed by the upturned edges of
the scale; six scales between dorsal nasal scales; occipital scale a
half (or more) larger than adjoining scales, separated from the
semicircles by three scalerows; about six loreal scalerows; a row
of enlarged suboculars separated from the labials by a row of
scales; nine or ten supralabials and about the same number of
infralabials; back part of mouth bordered by granules; mental
border on mouth very slightly larger than the rostral border, the
scale three-fourths divided by a median groove, and bordered be-
hind by eight scales, none especially enlarged; five scalerows below
infralabials somewhat larger than other chin scales; a well-devel-
oped dewlap extending onto breast; about eleven or twelve rows
of enlarged keeled dorsal scales, the two median largest and not
in a groove but somewhat elevated above other dorsal scales;
lateral scales keeled, the keels pointing upwards and backwards;
ventral scales enlarged, the largest equal to or at most slightly less
than dorsals; scales and granules around body about 82 of which
about 23 are to be regarded as enlarged ventrals; scales on upper
part of arm, front of forearm, on front face of femur, and around
tibia, as large as dorsal scales, the keels forming straight lines di-
tected outward from body; a postanal groove but no enlarged
Postanals. Fingers and toes very moderately widened with twelve
Or thirteen subdigital lamellae under widened part of the two
longest fingers, and eight scales under the distal segment; nineteen
lamellae under wide part of fourth toe, with nine scales under
the distal section; scales on. tail larger than dorsals, the single
tow of median dorsals slightly serrate; about thirty tail segments
all somewhat compressed except at base of tail; height of tym-
panum equal to about three fifths of eye-opening; opening of the
axillary pouch about as large as eye-opening, and about equal to
its own depth; head longer than tibia.
Color in life: Above generally dark olive brown with a vague
Suggestion of lighter transverse markings between somewhat darker
areas; belly light brown with some metallic reflections; dewlap dark
reddish with a yellow border.
100 Tue Universiry SCIENCE BULLETIN
Measurements in mm.: (Nos. 40893 and 40889): Snout to vent,
43, 45; tail, 68.2, —; total length, 110.7, —; head length, 13, 14.9;
head width, 7.3, 7.6; snout to ear, 12.2, 12.5; snout to arm insertion,
19.2, 21; axilla to groin, 18, 18.8; arm, 17.5, 19.2; leg, 34, 36; tibia,
Maldsg. Py
Remarks: Variation in the small series is not great. It is pre-
sumed that the form may intergrade in the Pacific lowlands in
Panama with the more typical humilis humilis. If this is not proved
it may be necessary to regard it as a distinct though related species.
Anolis lemurinus lemurinus Cope
(Figs. 24, 25, 26)
Anolis (Gastrotropis) lemurinus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1861,
p. 213 (type locality Veragua, New Granada [= Veragua, Panama] Barbour,
loc. cit., suggests Cucuyos as probable locality ).
Anolis lemurinus Roulen gr Catalogue of the lizards in the British Museum,
2nd Ed., vol. 2, 1885, pp. 86, 87; Giinther, Biologia Centrali-Americana;
Reptilia ‘and Batrachia, Oct. 1885, Dr Da) Schmidt, Smithson. Misc. Coll.,
vol. 89, Mar. 16, 1933, p. 8 (Mt. Pirri, Darien); Barbour, Bull. Mus. Comp.
Zool., vol. 77, no. 4, June, 1934, p. 137 (Anolis (Dracontura) vittigerus
Cope, and Anolis palpebrosus Peters, Anolis frontatus Thominot, Anolis
biporcatus Peracca (part.), Anolis binotatus Barbour, (part.) in synonymy ).
Anolis lemurinus lemurinus Stuart, Misc. Publ. Mus. Zool. Univ. Michigan
no. 91, June 6, 1955, pp. 18-19.
Anolis biporcatus Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
Oct. 1885, p. 52, (part.).
Diagnosis: A medium-sized Anolis with the anterior scales of the
semicircles much enlarged, the series separated mesially by one row
of scales, or occasionally they may be in contact at narrow points;
dewlap small, indicated also in females; no enlarged postanals; no
axillary pockets; mental divided by a groove; one or no suboculars
touch labials. Twenty median dorsal scalerows slightly enlarged
but smaller than ventrals; approximately 160 scales and granules
in a row around body; supraoculars in about four irregular rows,
the two median largest, keeled; tail subcircular or slightly com-
pressed distally; a series of large blackish or brownish marks along
dorsum, or a broad median golden stripe from occiput to tail (pres-
ent in some females); rarely diamond-shaped marks are present;
roughness of head scales variable.
Description of species (From KUMNH 34270, Turrialba, C. R.):
Rostral three times as wide as high, bordered behind by two labials
and seven postrostrals; one scale between rostral and nasal;
scales between nasals; the semicircles consist of seven to nine scales
one of which, lying anteriorly is 2% mm. long, and separated from
its fellow by two to four scale rows; the series separated mesially
Review or Lizarps or Costa Rica LOL
Fic. 24. Anolis lemurinus lemurinus Cope, KUMNH No, 34270;
Turrialba, Cartago Province, C. R.; actual length, snout to vent, 63.5
mm.; total length, 165.5 mm.
102 Tue Universtry SCIENCE BULLETIN
by one scalerow; occipital scale irregular, larger than ear-opening,
separated from semicircles by three or four scale series; supra-
oculars in four series the two median largest, somewhat keeled or
roughened, the inner row separated, at least partially, from. the
semicircles by a row of granules; frontal depression very shallow;
scales on snout lightly keeled; four or five canthals, each with a
sharp keel; two small longitudinal rows of temporals; one much-
enlarged supraciliary almost completely overlapping the second
(posterior ) one; six loreal series; six supralabials to below middle of
eye; five or six infralabials to same point; six or seven enlarged
suboculars the largest alone touching the labial series; three or four
series of scales lying along infralabials somewhat enlarged; mental
three-fourths divided by a groove, bordered behind by two labials
and six postmentals, the outer pair enlarged; much-enlarged scales
on each side of dewlap; scales of posterior occipital region very
minute; about twenty enlarged dorsals, the four median a little
larger than others; 72 scales on median line between level of axilla
and groin; lateral granules small with a few scattered, slightly
enlarged scales low on side near groin; ventrals subround, usually
pointed slightly, distinctly keeled; dorsal caudal row much enlarged
at base, keeled, but not serrate; two ventral series enlarged; all
caudal scales keeled; (no enlarged postanals in males); digits wid-
ened; 28 scales and lamellae under the third and fourth fingers;
38 under fourth toe; scales on front and dorsal part of upper arm,
on upper surface of forearm, on front of thigh, and ventral surface
of tibia much enlarged, equal to or larger than the ventrals, all
keeled; head longer than tibia; leg reaches eye; arm fails to reach
groin.
Color in life (at night): Pinkish gray with large blackish quad-
rangular blotches with somewhat grayish centers on dorsum; sides
gray; pinkish-white below on venter, under tail, and limbs.
Measurements in mm.: Snout to vent, 63.5; tail broken, partly
regenerated, 102; snout to ear, 16; snout to leg insertion, 24.2; axilla
to groin, 28; length of head, 18.3; width of head, 11; arm, 26; leg,
46.
Variation: The specimens having the dorsal black blotches are of
both sexes and are from the following localities: Limén Province:
KUMNH Nos. 40583-40584 Sipurio. Cartago Province: 34269,
34970 Turrialba; 34275 Tunnel Camp, near Peralta; Guanacaste Prov-
ince: 34271, Las Flores, Tenorio, Las Cafias; 4 to 5 km. ENE
Tilaran. Specimens having the dorsal stripe are all females. Car-
Fic. 25. Anolis lemurinus lemurinus Cope. KUMNH
No. 342739; Golfito C.R.; actual length, snout to vent,
60 mm.; tail 125 mm.
104 Tue Universiry SCIENCE BULLETIN
tago Province: No. 34269, Morehouse Finca. Puntarenas Proy-
ince: 34272-34274, Golfito.
The specimen from Sipurio, No. 40583 ¢ , differs in having the
dorsal blotches rather dim and series of small orange spots forming
dorsolateral rows, with small orange spots on thighs and near base
Fic. 26. Anolis lemurinus lemurinus Cope. KUMNE
No. 40585, 4-5 km. ENE Tilaran, Guanacaste Province,
C. R.; actual length, snout to vent, 59.2 mm.; tail, 97 mm.
of tail. The dewlap has a depth of about 7 mm.; in preservative
it is reddish orange in color on outer parts, the basal part white,
the scales stippled with black. There are no enlarged postanal scales.
In life the specimen was olive with slight yellowish wash, the
frontal area and snout brownish. A light indistinct band was pres-
Review or Lizarps or Cosra Rica 105
ent across the head involving the front part of the upper eyelid.
The axilla was dull yellowish. A distinct dorsolateral row of light
orange spots were present. The scales on the side of the dewlap
were pure white in life.
A second specimen taken from the stomach of Anolis biporcatus
that was captured on Mt. Mirador near Suretka, showed the dorsal
surface of head and body with bright varied metallic iridescence.
On the head there are numerous small black spots; the occipital
area is almost solid black and there is a black line from the eye
meeting its fellow on the anterior nuchal region. The venter and
throat are light with greenish or bluish metallic reflections. How-
ever there are a few blackish flecks on the chin and a brown spot
on the belly. The dewlap is red, the temporal area copper-colored.
The females from Golfito are generally greenish gray on the
sides of the head and body. A broad golden band begins just
Posterior to the occipital scale, widens and continues onto tail
Where it is gradually lost; it is bordered by narrow indefinite
brownish lines. The banding on the tail is not distinct. There is
a large brownish blotch in the middle of the belly extending onto
the sides,
A female from the Morehouse Fincé , Turrialba has a broad golden
Stripe, the edges sinuous and forming at intervals four lateral
angles. The head and the sides of the neck are deep brown; No.
34273 has a dim hair-fine median golden line bordered by narrow
black stripes. The venter is uniform light. The stippling on the
scales is scarcely discernible under a lens. There is some difference
in the rugosity of the head scales; the semicircle scales in some
Specimens continue on behind the eye; in others the number is re-
duced and they are not distinguishable back of the eye.
The Golfito specimen (No. 34272) measured 58 mm. snout to
vent. The tail, complete, is 123 mm. in length, thus more than
twice the head-body length.
Remarks; These animals are rather slow moving. Some have
been taken at night seemingly asleep at the outer tip of small
branches. The female from near Turrialba has three ovarian eggs,
two much larger than the third.
A specimen KUMNH 40505 from 4.5 km. ENE of. Tilaran.
Guanacaste Province, is gray with diamond-shaped blackish marks
(figured),
Tue Universiry SCrENCE BULLETIN
Anolis intermedius Peters
(Figs, 27, 28)
Anolis intermedius Peters, Monatsb, Akad, Wiss, Berlin, 1863, p, 143 (type
locality, Veragua, Panama); Bocourt, Mission Scientifique au Mexique et
dans ’Amérique Centrale; Etudes sur les reptiles, livr, 2, 1873, pp. 69-71,
pl. 15, fig. 4, (redescription of the type); Cope, Journ, Acad, Nat. Sci.
Philadelphia, ser, 2, vol, 8, 1876 (1875), p. 124 (Costa Rica); Boulenger,
Catalogue of the lizards in the British Museum, 2nd Ed, vol. 2, 1885, pp,
78-79 (part.) (places nannodes Cope, and tessellatus O'Shaughnessy in syn-
onymy); Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
Oct. 1885, p. 49 (part.) (Irazi, Costa Rica); Cope, U. S. Nat. Mus, Bull.
32, 1887, p. 30, Proc. Amer. Philos, Soc, Dec, 28, 1893, p, 344 (San José
C. R.); Boettger, Ber. Senck, Natur. Ges, Frankfort, 1894, p. 27; idem, ibid,
1895, p. 36 (San José. C. R.); Dunn, Proc, New England Zool, Club,
vol. 12, 1936, p. 18 (Santa Rosa, Costa Rica); Wettstein, Sitzungsb, Akad.
Wiss. Wien, Math-natur. KI. Abt. 1, Band 143, Heft, 1-2, 1934, p. 21 (La
Carpintera, C. R.); Barbour, Bull, Mus, Comp, Zool. Harvard Col., vol. 77,
no, 4, 1985, p. 185 (San Jose),
Fic. 27. Anolis intermedius Peters. Left figure, KUMNH
No, 342839; length, snout to vent, 49 mm., Cartago, Cartago
Province; middle figure, 34286 ? , length, snout to vent, 47.3 mm.;
right figure, 34284@, length, snout to vent, 44 mm.; same loc.
Review or Lizarps or Costa Rica 107
Diagnosis: A medium-small species, the median semicircles com-
posed of few large scales of which one, often two, are in contact on
middle line; head scales slightly rugose or smooth; four rows of
loreal scales; ventrals keeled, a little larger than median dorsals;
three of subocular series bordering labials; occipital larger than ear;
frontal depression scarcely discernible; four postmentals, the two
outer enlarged; two longitudinal series of somewhat enlarged tem-
porals; a row of somewhat enlarged postanals; hind leg reaches half-
Way between arm insertion and ear.
Description of species (From KUMNH No. 34283 Cartago, Car-
tago Prov. C. R.): Rostral about three times as wide as high, bor-
dered behind by two labials and five postrostrals; four scales between
tiny supranasals; three or four enlarged canthal scales; anterior
smallest; the semicircular rows are not clearly distinguished; a pair
of very large scales lies between the supraorbital areas completely in
contact; these preceded by a pair of scales nearly as large but
separated mesially by a single scale; these are separated from can-
thals by a single smaller scale; behind the central scales there are
three of the semicircular series curving behind supraorbital area;
occipital larger than the ear-opening, separated from semicircles
by two seale series; behind occipital region two somewhat swollen
areas each covered with rather large scales in three or four series
but separated by granules from a double line of somewhat en-
larged temporal scales; five scales across snout between first can-
thals; two irregular series of supraoculars, the inner three scales
largest, separated from semicircles by a row of regular granules;
four rows of loreal scales; four suboculars enlarged, three touching
labials, only one scale between rostral and nasal; eight supralabials,
the seventh below middle of eye; eight infralabials; mental larger
and wider than rostral, divided by a groove, bordered behind by
two labials and four postmentals, the outer enlarged; three scale
series bordering the infralabials somewhat enlarged, one larger
than others.
Ventral scales keeled, in about 26 rows, merging gradually to
fine granules of sides; dorsolaterally scales become gradually larger,
the dorsal fourteen rows nearly same size; however the median
three or four are slightly largest but smaller than ventrals; (males
with a slight nuchal fold or crest, and the median dorsal scales may
be slightly elevated); a few scattered larger scales among granules
on sides preceding groin; about 126 rows of scales and granules
about middle of body; digits except first dilated, 13 widened
108 Tue Universitry SCIENCE BULLETIN
lamellae and seven scales under third and fourth fingers; no axil-
lary pocket; 33 lamellae and scales under fourth toe; hind leg very
short reaching about halfway between arm insertion and ear;
scales on upper side of upper arm, front of forearm, front face of
thigh, and the back and under portion of tibia with large keeled
scales equal or in places larger than ventrals. Tail nearly cylindri-
cal ‘at base, more compressed distally; the single median dorsal
series not serrate.
Fic. 28. Anolis intermedius Peters. KUM
NH No. 34282, Cartago, Cartago Province,
C. R.; actual length, snout to vent, 50.2 mm.,
tail (regen. ), 62 mm.
Review or Lizarps or Cosra Rica 109
Color in life: Very changeable. Darker median stripe covering
about twelve scalerows bordered by a dim line of fawn on which
are discernible four pairs of somewhat triangular brown spots, the
last between hind limb insertions breaking continuity of fawn line;
narrow brown line on head between eyes; occipital region blackish;
a few silvery flecks on the scattered enlarged scales in region an-
terior to groin; tail banded in dirty fawn and brown, dirty whitish
below, with some brownish flecks and minute black stippling;
metallic reflection on belly and under limbs.
Measurements in mm.: Snout to vent, 48; tail broken and regen.
58; snout to arm insertion, 19; snout to ear, 14; axilla to groin, 23;
length of head, 15; width of head, 8.4; arm, 18; leg, 29; tibia, 10.
Variation: The arrangement of the large scales on the head varies
and ‘scarcely any two specimens having the same exact arrangement
can be found in my small series. In one other Cartago female
(specimen No. 34284), there are seven scales in the semicircles,
only one pair, between eyes is in contact. In this case however
the large median pair is transversely segmented and a small median
scale separates the anterior moieties; the scales on head are slightly
More rugose; the supraoculars are fewer and larger, one or two
touching the semicircles. The color pattern is practically the same
as the preceding.
In a male from Cervantes, the head scales have shallow depres-
sions giving the appearance of being very rugose; two large pairs
of scales are in contact between eyes. There is a distinct nuchal
fold or crest and an indistinct dorsal crest. The mental is followed
by only two large postmentals. There are five or six enlarged
postanal scales present. The dewlap is small, its depth probably
not half as great as head length. There are seven or eight large
Supraoculars largely covering supraocular region.
A specimen from higher elevation in the National Forest agrees
in color with the described specimen. In this, a male, the postanal
Scales are a little smaller; a female specimen 47 mm. long has a
Complete tail measuring 76 mm. This specimen is generally gray-
ish with a faint rose wash and some blackish dorsal flecking or
shading. The supraocular region is brownish and the interorbital
and occipital area gray while the tip of the snout is darker; there
is a faint lavender tinge on the white of the venter. The dewlap
of males is usually white with sometimes a faint greenish or rosy
Wash,
Remarks: This species seems to persist after the removal of forest.
They thrive in gardens in the cities, on hedges, or on fences with
110 Tue Universrry ScreNcE BULLETIN
live posts. They seem to be curious and will perch on a branch of
a shrub watching children playing close by. They are not much
afraid and one may often approach them slowly and grasp them with
little difficulty. The dewlap of the males may be distended as the
animal remains otherwise quiet, within a few feet of the beholder.
The appendage is small and not especially conspicuous.
Anolis cupreus Hallowell
(Fig. 29)
Anolis cupreus Hallowell, Proc. Acad, Nat. Sci. Philadelphia, 1860, pp. 481-
482, (type locality, Nicaragua); Cope, Proc. Acad, Nat. Sci. Philadelphia,
1874, p. 68 (Nicaragua); Boulenger, Catalogue of the lizards in the Brit-
ish Museum, Qnd Ed., vol. 2, 1885, pp. 80-81 (Guatemala and “neighbor-
hood of Irazu [Vole4n], Costa Rica”; Giinther, Biologia Centrali-Americana;
Reptilia and Batrachia, Oct. 1885, p. 50; Boettger, Katalog der Reptilien-
Sammlung im Museum der Senckenbergischen Naturforschenden Gesell-
schaft in Frankfurt am Main, I Teil. Aug. 1893, p. 59 (San José); Dunn,
Proc. New England Zool. Club, vol. 12, 1930, p. 17 (San José, Costa Rica);
Schmidt, Field Mus. Publ. Zool., vol. 12, Nov. 21, 1928, p. 195 (El Salva-
dor); Barbour, Bull. Mus. Comp. Zool., Harvard Col. vol. 77, no. 4, 1934,
p. 129-130 (includes hoffmani Peters from.Costa Rica); Stuart, Misc. Publ.
Mus. Zool. Uniy. Michigan, no, 91, June 6, 1955, pp. 15-16 (places macro-
phallus Werner in synonymy ).
Anolis hoffmanni Peters, Monatsb. Akad. ‘Wiss. Berlin, 1862 (1863?) p. 142
(Costa Rica); Bocourt, Mission Scientifique, au Mexique et dans Amérique
Centrale; Etudes ‘sur les reptiles, livr. 2, 1873, p. 86, pl. 15, figs. 15, 16;
Cope, U. S. Nat. Mus. Bull. 32, 1887, p. 31 (Costa Rica and Guatemala ).
Diagnosis: A diminutive species, (40-42 mm.) the median dor-
sals (fifteen to twenty rows) larger than lateral granules; four
median rows largest, keeled, but much smaller than the ventral
scales which are rhémboidal, imbricating, and distinctly keeled;
gular appendage of male relatively very long longitudinally, ex-
tending to middle of belly, not or scarcely indicated in female;
head slightly longer than tibia; no enlarged postanals; scales of
supraorbital semicircles large, separated by one or two series of
scales; occipital larger than ear-opening; six loreal rows.
Description of species (From KUMNH No. 40751 Las Flores,
Tenorio, Las Cafias, Guanacaste Province, C. R.): Rostral about
three times as wide as high, bordered by two labials and seven
postrostrals; nine scales between nostrils; nasal a tiny ring about
nostril separated from rostral by one scale; supraorbital semicircles
well defined, not or scarcely elevated into ridges, each scale with
one sharp keel and other sculpturing; scales on frontal area keeled,
the edges of the scales somewhat elevated with some fine tubercu-
lation; semicircles separated by one or two scale rows; two large
supraoculars and four smaller ones, all keeled, the larger ones sep-
arated from the semicircles by a row of granules; occipital en-
Review oF Lizarps or Costa Rica aia
larged, nearly double size of ear-opening, surrounded by three
series of scales, with their edges elevated appearing to be tricari-
nate, and separated from the semicircles by three scales; two
elongate supraciliaries, the anterior overlapping the posterior; back
part of supraciliary edge without enlarged scales; canthus formed
by three enlarged sharp-edged scales; six or seven supralabials and
Fic. 29. Anolis cupreus Hallowell. KUM
NH No. 40739, Maribella Hotel, mouth of Rio
Barranca, Puntarenas Province, C. R.; actual
length, snout to vent, 41 mm.
112 Tue Universtry SCIENCE BULLETIN
seven or eight infralabials; seven loreal series; three or four enlarged
suboculars separated from labials by one row of scales; five or six
scalerows below infralabials somewhat larger than median series
on chin; mental slightly wider than rostral, divided by a groove,
the parts nearly triangular, about 2s wide as deep, followed by two
infralabials and six keeled postrostrals; five median scalerows on
back larger than others, the edges being upturned they appear
tricarinate; they are polygonal, chiefly pentagonal; the next eight
or ten rows gradually merge into the small lateral granules; low on
sides the keeled granules gradually enlarge; ventrals much larger
than the dorsals, with a single keel, imbricating and somewhat
rounded posteriorly; a faint trace of a dewlap in female repre-
sented by a slight fold on neck; about 125 scales and granules in a
row about the middle of body; caudal scales hexagonal, large,
keeled, the median dorsal row enlarged as are the two ventral
rows; tail thickened at base, slightly compressed to tip; about ten
pointed scales bordering vent anteriorly; (no enlarged postanal
scales in males ); arm short not reaching more than halfway between
eye and nostril; leg reaches eye; digits widened somewhat; 16
lamellae and scales under the fourth finger; about 19 widened lamel-
lae under fourth toe (distal joints of both fourth toes incomplete and
scales not counted); enlarged scales on dorsal part of upper arm,
front face of forearm, front face of thigh and ventral part of tibia,
some about the size of ventrals; ear-opening small, 1.1 mm. x .7
mm.; head longer than tibia.
Color: A median cream line covering two scalerows; this bor-
dered laterally by a grayish stripe of varying width bordered on
both sides by brown, the outer edge sinuous or forming angles at
intervals; head nearly uniform olive-brown; a faint brown mark
begins behind eye and runs back across temporal region; below
light on chin and venter.
Measurements in mm.: Snout to vent, 41; tail, 71; snout to ear,
10.5; snout to leg insertion, 17.3; axilla to groin, 17; length of head,
12.2; width of head, 6.8; arm, 15.5; leg, 29.5; tibia 9.3.
Variation: Specimens in the collections are from the following
localities: Guanacaste Province: KUMNH Nos. 40743, 40752-
40763, 4 to 5 km. ENE Tilar4n; 40747-40751 Las Flores, Tenorio,
Las Canfas. San José Province: 40741 San José; 5-10 km. E Sarchi.
Puntarenas Province: 40739, 40740, 40742 Maribella Hotel, mouth
of Rio Barranca.
Review or Lizarps or Costa Rica 113
Males have a gular dewlap which is elongate, extending back to
the middle of the belly. The color varies from cerise to magenta,
enclosing an area of orange or of orange mixed with black.
The body of males appears more compressed. Often the median
part of dorsum bears an indefinite dark stripe. One or two of the
specimens have a dim brown spot on the shoulder. The female
specimen figured has a yellow median line and dim paired black
spots in preservative.
Remarks: This species, which somewhat resembles limifrons in
size and behavior, was taken only on the Pacific slopes of the coun-
try. Specimens were seen most frequently in high grass which was
their permanent dwelling. I have not found the species in typical
forest,
[Anolis sp.]
(Fig. 80)
A specimen KUMNH No. 10649 from near Esparta, Puntarenas
Province, differs from the typical cupreus in being significantly
larger (49 mm. snout to vent, the tail regenerated ); six median
Scalerows enlarged, smaller than ventral scales; postanals enlarged;
about 132 scales and granules around body; occipital much larger
than ear-opening; semicircles separated by one scalerow; suborbital
elongate and except for last, separated from labials; six loreal
Series; posterior supraciliaries small; dewlap attachment elongated
to middle of abdomen, as in typical cupreus.
The color is olive green with a large black shoulder spot followed
by three other lateral spots less strongly distinguished. Whether
this specimen represents a distinct species, a subspecies of cupreus,
ris only an aberrent specimen of cupreus cannot be decided now.
Anolis altae Dunn
Anolis altae Dunn, Proc. New England. Zool. Club., vol. 12, Aug. 7, 1930,
bp. 22-23 (type locality, Acosta Farm, about 7000 ft, elev. Volean Barba,
Costa Rica),
Diagnosis: An Anolis with keeled ventrals; dorsals much smaller
than ventrals, granular to keeled, gradually enlarged to middorsal
region, but none much so; two scales between semicircles; tail
Cylindrical with even large keeled scales.
Description of type: “Two faint frontal ridges diverging, and two
ainter prefrontal ridges converging; head scales unicarinate or
at to concave; rostral low and wide, lower than mental; semi-
Circles separated by two scales; occipital larger than ear, separated
from semicircles by two scales; supraocular disks of three large
5—3378
f
Tue UNiversiry ScreENCE BULLETIN
_ Fic. 30, Anolis sp.P KUMNH No. 40632, 6 mi. I
Esparta (on cliff along highway), Puntarenas Province,
C. R.; actual length, snout to vent, 49 mm.
Review or Lizarps or Costa Rica 115
scales and smaller ones, flattish and unicarinate, in contact with
semicircles; canthus rostralis evident only for a short distance just
anterior to eye; three angulate scales, in front of which is a uni-
carinate scale, the most anterior of the semicircles; the posterior is
on the eyelid; posterior edge and half of eyelid granular; six rows
of loreal scales; six supralabials; temporals granular; fingers and
toes not much dilated; sixteen lamellae under phalanges I and II
of toe IV; tail almost cylindrical, scales large, imbricate and uni-
carinate; ventrals much larger than dorsals, keeled; dorsals grad-
ually increasing to last row, which is keeled but not much enlarged.
Color: “Brown, a broad (16 scale wide) tan dorsal band; whitish
below, Dewlap not noticeable (a female ).”
Measurements in mm.: “Total length, 142; head and body 52;
tail, 90.”
Remarks: Dunn regards this species as being closest to concolor.
Anolis woodi woodi Dunn
(Fig. 81)
Anolis woodi Dunn, Proc. Acad. Nat. Sci. Philadelphia, vol. 92, 1940, pp.
110-111 (type locality, El Volean, Chiriqui, Panama).
Diagnosis; Rather large form, 72+ mm. snout to vent; ear-opening
four fifths of the length of eye-opening; suboculars separated by
One scalerow from the supralabials; a large shallow axillary de-
Pression, scaled and pigmented; a slight nuchal fold or crest, and
4 slight median dorsal ridge; the 20 median scalerows larger than
laterals; dorsal scales distinctly smaller than ventrals, which are
keeled and strongly imbricating, rather oval in posterior outline,
the tips slightly elevated; postanals distinctly enlarged; supra-
Orbital semicircle consisting of elevated enlarged scales, separated
by two or three scalerows; the canthal scales and those bordering
them have from two to five fine keels or ridges directed outward
and backward: occipital scale, flanked by two enlarged scales, is
less than half size of ear-opening; tail 2.3 times length of head and
body,
Description of species (KUMNH 40633, 1 mi. W Cafias Gordas,
Puntarenas Province): Greatest total length known, 235 mm.
Rostral nearly four times as wide as high, bordered behind by eight
Tough postrostral scales and two labials; separated from nasal by
ne scale; eight scales between nasals, with high ridged keels;
Supraciliary scales overlapping the larger ones beginning behind
the middle of eye, extending forward, and joining the canthal
Series; latter with sharp canthal edge extending to below nostril,
about six scales in the series, the anterior ones small; seven loreal
Tue Universiry ScueNCE BULLETIN
Fic. 31. Anolis woodi woodi Dunn. KUMNE No. 40633, Canas
Gordas, Puntarenas Province, C. R.; actual length, snout to vent, 71
mim., tail, 164 mm.
Review or Lizarps or Costa Rica ABS
Series; supraorbital semicircles composed of large scales somewhat
elevated, each with two to five fine brown keels or ridges directed
variously, the series separated by three rows (the median of which
may be interrupted); a shallow frontal depression floored with
rather large scales; supraocular region with about four rows of en-
larged keeled scales, the inner series largest, the outer little larger
than the granules of outer part; largest series separated from the
semicircles; occipital distinct, pointed behind, flanked by two much-
enlarged scales; the occipital separated from semicircles by four
Tows of scales; eleven or twelve supralabials, the ninth below
middle of eye; seven or eight rather large suboculars separated
from the supralabials by an intercalated series of smaller scales;
mental wider than rostral, about three times as wide as high with
two short posterior projections and a groove that divides it for more
than two thirds of its height, bordered behind by a large outer
nultiple-ridged scale and three smaller ones, as well as by two
labials; eleven or twelve infralabials, the posterior part of the
mouth bordered by small granules; five or six rows below infra-
labials somewhat enlarged; occipital area with several rows of
Scales bordering occipital, all somewhat enlarged; granules behind
these smaller than those on dorsal part of body; about 125 scales
around middle of body; a very slightly elevated nuchal crest, more
or less continuous with a discernible dorsal crest (in males) which
Continues onto tail; body distinctly triangular in cross-section; tail
triangular at base, distally compressed; dorsal caudal scales en-
larged, not or but slightly serrate; caudal scales im whorls; tail ends
in a slightly widened black tip.
Ventral scales much larger than dorsal (dorsals, eleven in 5 mm.
distance, ventrals, eight in same distance ); granules on side small,
the twenty dorsal rows considerably larger than laterals; dorsals,
laterals, and ventrals keeled or ridged; postanal groove deep,
arched, with a single median and two lateral s rales, all distinctly
enlarged; paired subcaudals larger than adjoining scales.
Dewlap large, extending back of level of axillae for seven milli-
Meters; its greatest length (depth) about equal to head length;
length of tibia equal to head length.
Arm brought forward the fingers reach snout tip; seventeen
lamellae under widened part of the two middle fingers; a large
distinct depression in axillary region, not forming a typical pocket.
“eg extended, the fourth toe reaches to a little in front of orbit.
Wenty-four scales under widened part of fourth toe and ten on
118 Tue Universrry SCIENCE BULLETIN
distal phalanx making a total of 34. Tail 2.3 times snout-vent
length.
Color in life: Generally olive with some indistinct rusty spots;
tail rusty brown, indistinctly barred with olive; venter olive with
much rusty brown in evidence; leg olive, narrowly barred with
rusty brown; eyelids with a brown band, and several fine trans-
verse lines; a dark area in occiput and some radiating brown bands
below eye; a dark light-edged nuchal spot.
Dewlap (throat fan) large, pink-orange on edge, then a band
of amber-yellow, while at base an area of bluish-white; many
scales on sides of dewlap whitish. Chin pinkish; rusty orange
spots under base of tail; lips lighter than sides of head. The char-
acteristic markings of the preserved specimens are indicated in the
figure given.
Measurements in mm.: Snout to vent, 71; tail, 164; total length,
935; snout to ear, 174; snout to arm insertion, 28.5; length of head,
20; width of head, 12; height of tympanum, 2.3; arm, 33; leg, 58;
tibia, 20; axilla to groin, 30.
Remarks: This specimen was the only one of this species that
was seen. It was caught on a low bush in the daytime.
The female type is the only other specimen that has been reported.
The type description is somewhat inadequate and some important
data are omitted. Dunn suggests that the color and size is similar
to Anolis capito and perhaps they may be related. Such relation-
ship is indeed remote and no pertinent similarities are in evidence.
The size of the type female is only two millimeters larger than
the male here described, one millimeter in snout-vent length, and
one in tail length.
The specimen differs markedly in coloration, as there is no trace
of the white lobulate dorsal band occurring in the female, and the
number of scales and lamellae under the fourth toe is greater (34
instead of 29).
Anolis woodi attenuatus * subsp. nov.
(Fig. 82)
Type: KUMNH No. 34257, collected at Isla Bonita (American
Cinchona Plantation) southeastern slope Volcan Pods, elev. circa
5,500 ft., July 29, 1947, Edward H. Taylor, coll.
Paratypes: Richard C. Taylor, No. 668 and KUMNH No. 34256,
topotypes.
* From Latin attenuatus = drawn out, tapered, in reference to the greatly elongated tail.
Review or Lizarps oF Costa Rica 119
Diagnosis: A medium-large species 78-83 mm. snout to vent;
the tail 183-201 mm.; mental about 2/3 divided with smaller labial
border than rostral; six scales (beside labials) border rostral; a pair
of semicircular ridges not reaching canthus, outlining a frontal and
an interorbital depression; trace of a pair of low frontal ridges;
occipital depression rather shallow; head scales keeled: about
twenty rows of larger scales on back, but more or less gradually
merging into the granules on sides; keeled ventral scales perceptibly
larger than dorsals; postanals more or less enlarged; leg reaches be-
yond eye; arm reaches groin; tail compressed, approximately 2.4
times body length; supralabials usually 10; infralabials 12-18; ven-
tral scales keeled; head longer than tibia.
Description of type: A medium-sized species reaching a known
total length of 283 mm., snout-vent length 83 mm.; body not or but
slightly compressed, the tail definitely so (about 1.4 times as high as
wide ); head rather large, its length slightly less than twice its width,
its height (11 mm.) slightly less than half its length; canthus ros-
tralis not pronounced, consisting of four enlarged scales which extend
little more than halfway from orbit to nostril; rostral about four
times as wide as high, bordered by two labials and six other scales;
snout elevated, covered with somewhat elongate bi- or tricarinate
scales, the median largest; a pair of interorbital ridges running for-
ward, diverging; trace of two parallel frontal ridges that enclose a
slight depression, which is continuous with interorbital and occipital
depressions, all of which are rather shallow; scales on floor of frontal
depression longer than wide, arranged in longitudinal lines, bi- or
tricarinate; scales of the semicircles have two or three keels on their
Outer faces, and two rows of scales separate them between orbits;
Supraorbital areas narrow, with four rows of keeled scales, some-
What enlarged, the inner row for the most part directly in contact
With the semicircles; two rather short supraciliaries anteriorly, not
reaching halfway across the supraciliary front; other scales of the
Supraciliary edge not especially differentiated; occipital area with
& small occipital plate, less than one third of the area of the auricular
Meatus, surrounded by small scales, about four rows of which
Separate the occipitals from the scales of the semicircles; two low
tidges bordered by swollen areas border occipital region, which
is somewhat depressed; none of the scales of the semicircles much
elongated: eight rows of scales in loreal region; scales of the sub-
ocular series strongly enlarged, not continuous behind eye with
the semicircles, separated from the supralabials by a row of narrow
Tse Universrry Science BULLETIN
Fic. 32. Anolis woodi attenuatus subsp. noy. Type, KUMNH No. 34257;
Isla Bonita (Cinchona), Heredia Province, C, R.; actual length, snout
to vent, 82 mm.; total length, 283 mm.
Review or Lizarps or Costa Rica 121
scales; supralabials 10-10, the posterior reduced; 13-13 infralabials,
the posterior small; mental divided for about .6 to .75 of its height
by a suture entering from behind, its labial border shorter than that
of the rostral; eight rows of somewhat enlarged scales border the
infralabials; two labials and six other scales bordering mental.
Throat fan large, when distended about 30 mm. long and 26 mm.
wide. Arms long, when extended they reach two or three millime-
ters past the groin; scales on top and front of upper arm and front
face of forearm much larger than dorsal or ventral scales, keeled;
16 scales and lamellae under basal joints of middle finger, the digits
moderately but distinctly widened; a depression, but no distinct
axillary pocket behind arm insertion; leg long, slender, the tip of
the longest toe r aching to a point beyond eye one third of distance
between orbit and nostril; 27 scales and lamellae under middle toe
(not counting those extending onto the sole); ten scales under distal
joint; scales on front of thigh and tibia larger, keeled. A pair of
somewhat enlarged median dorsal scalerows; adjoining eight rows
distinctly larger than the small lateral granules into which they
Sradually merge; ventral scales a little larger than median dorsals,
strongly keeled; about 140 scales and granules around body; median
Postanal scales somewhat enlarged, a little larger than scales that
follow them, but much larger than scales in the depression in front
of them; crest of basal part of tail with median paired scales (for
15 mm.) followed by a single row, broken occasionally by single
Pairs, all scales keeled. Scales forming rows on sides of throat
fan are contiguous; tail compressed.
Color: In life, general color dull brick-red with variegated black
Markings, Throat fan very dark olive to blackish with magenta
scales; under part of body salmon to pink, with some areas tending
toward the dorsal coloration; areas on head, above eyes, greenish.
In preservative, generally fawn-brown to darker brown; head
flecked and variegated with darker brown, and with minute black
Pigment spots; the greenish color in supraorbital areas persists with
Some darker transverse tracings; three darker spots below eye,
and an indefinite short stripe behind eye; occipital region some-
what black; a strong blackish band on shoulder in front of arm,
hearly meeting its fellow on neck, preceded by an area somewhat
lighter; a somewhat lighter indefinite band on back behind arms,
°ne at middle of body that is somewhat divided by lighter tan,
and another indefinite band on sides of rump; arms with four or
five indefinite bands, those on legs very dim; some indefinite cream
‘pots on back of thighs; tail very indistinctly banded. Below,
122 Tue University SCrENCE BULLETIN
Measurements in mm. and scale data of Anolis woodi attenuatus subsp. nov.
Nome i see ae ene 668 84257 84256
GO a i Ge eee ee é ry) é
Total Venous aici sf ee teesmetee in 260* 283 261
Shout iG Vent a en ena eer ae 83 82 78
Mena os Re ie ne Pes ye a 201 183
Snoul tO Cate Cae ee 21 21.2 20
Snoutto foreleg a eens ais 81.5 82.5 80
ASMA HO CREOIN tc oie a aan ee Oe 88.2 85
Head length to jaw angle ............ 23 23.4 22.2
Hidad Wicthieredtest se ey i 12.6 13.2 12
AAD a Aico ie AEE temic a eee 40 388 39
ee os he dq re ie ie eee ie cen? 69.2 dak 69.1
Pitt een et a 21 21 21
ROC ai (OG8 ek kar en ye 31 81 29
Scales between ridges ........ es 2-3 2-2, 2.2
Scales between ridge and occipital scale, 3-4 4-4 4-4
Labials and subocular series separated, yes yes no™”
Longest toe extends beyond eye ......4 mm. 3% mm. 5mm.
Avo to beyond Groin Gv. ee yes yes yes
Nichal crest-présent |. 6...8...5. yes yes yes
Occipital scale smaller than ear ...... yes yes yes
Scales behind rostval 7.5.5... ..... 6 6 6
Scales behind, mental .....%. 40... -- 6 6 6
LiOHGH | TOW Siero Mek ot oer te Re esse) 8 8
Stpralabials..... SHEN outers cn eve beh 10-9 10-10 10-10
Intvalabiale: 0 5 ee 12-12 13-18 12-12
* Tip missing.
** Two scales are in contact.
chin with two broken transverse marks; belly and under limbs
light fawn, with some scattering of brownish flecks. Throat-fan
deep black with dark grayish scales; digits banded to tips.
Variation: Most of the variation not recorded in the table has
to do with the distinctness of the color patterns. R.C.T. No. 665
shows the banding more distinct on the thighs; in KUMNH
34256, the markings are less distinct, the banding on limbs and
digits scarcely discernible.
Remarks: These three specimens were taken at the American
Cinchona Plantation at Isla Bonita on the southeastern slopes of
Volcan Pods. They were rather brightly colored and conspicuous.
Richard Taylor who observed a pair on the flattened top of a tree
fern, made the following note:
“Two anoles, a male and female were observed on the top of a
tree fern from the bank of a rivulet from about the same level as
the top. When first seen I presumed them to be fighting. The
male with dewlap extended would attempt to bite the female.
Then the female would retreat to another frond, and shortly would
return to attack the male. At one time in the procedure they
came face to face with mouths open. The female retreated fol-
lowed by the male, who at all times had the throat fan extended
even up to the time I captured it. As I approached closer they
Review or Lizarps or Costa Rica 123
became aware of my presence, and stopped, alert and motionless.
The male was captured, but the female es caped. It is possible
that this was a demonstration of courtship.”
The length of the tail in this form is considerable and is equalled
by the length of the tail in Anolis longicauda Hallowell. Boulenger
in his Catalogue disregarded this latter species (mentioned in foot-
note, vol. 2, p. 11) and Barbour, Bull. Mus. Jomp. Zool. Harvard
Coll. places it as a synonym of Anolis sericeus. I would suspect
that the two species, longicauda and this species are related. The
type of longicauda is presumably lost, and the type description is
inadequate. Only the acquisition of more material from the type
locality of longicauda can settle completely the problem of the
relationship of the subspecies.
Anolis townsendi Stejneger
(Fig. 33)
Anolis townsendi Stejneger, Bull. Mus. Comp. Zool. Harvard Col., vol. 36, no. 6,
Noy. 1900, pp. 163-164 (type locality, Cocos Island, Costa Rica, Chas. Hy,
Townsend collector); Proc. Biol. Soc. Washington, vol. 16, Feb. 21, 1903,
p. 3 (mention only).
Diagnosis: A small Anolis characterized by having rostral touch-
ing anterior nasal scale, bordered behind by two labials, two nasals,
and five small postrostrals; eight scales between the nasals; supra-
Orbital semicircular ridges moderately well defined, separated by
two or three rows of small scales; occipital scale larger than ear-
Opening, separated from the semicircular ridges by three rows of
Scales; a very small nuchal crest in males; keeled dorsals larger
than laterals but smaller than the keeled ventral, A white lateral
line from shoulder to groin bordered below by a black stripe; a
Tow of postanal scales, somewhat enlarged,
Description of species: Rostral more than three times as wide as
high, bordered behind by two labials, two nasals and five other
Scales; semicircles of enlarged scales begin somewhat behind eye
and curve around supraorbital area continuing on to frontal area,
Separated by two rows of scales at narrowest point and anteriorly
>y five or six rows; supraoculars in about four rows, the inner row
with the largest scales, but separated from ridges by a row of small
Scales; outer rows diminishing in size; occipital scale much larger
than surrounding scales, larger than ear-opening but much smaller
than eye-opening, separated from semicircles by two or three scale-
rows; all head scales keeled; supraciliary scales elongate sharp-
edged, overlapping; first canthal followed by a widened scale: five
"ows of frenals; a row of ocular scales, beginning close to a canthal
124 Tuer Universiry SCUENCE BULLETIN
scale, passing below eye, at least three of the scales in contact with
supralabials; scales bordering the occipital somewhat enlarged;
seven supralabials, the anterior rather elongate and keeled; seven
or eight infralabials, anterior ones nearly smooth, posterior ones
Fic. 33. Anolis townsendi Stejneger. Left figure, KUMNH No. 36493 4;
actual length, snout to vent, 45 mm.; middle, No. 36501 ¢, length, snout to
vent, 48.2 mm.; right, No. 36500 4, length, snout to vent, 46.5 mm, Cocos
Island, C. R.
Review or Lizarps or Cosra Rica 125
keeled; four or five outer longitudinal rows of chin scales some-
what larger than median series; mental divided by a groove into
two moieties; bordered behind by eight scales the outer distinctly
largest. A well-developed dewlap beginning on breast, its depth
a little less than head length.
Scales on dorsum variable, the two median rows largest, the three
or four adjoining also somewhat enlarged, all distinctly larger than
the lateral granules; ventral scales sli ghtly larger on the average than
the median dorsal rows, all keeled, the keels directed inwards and
backwards; scales and granules around middle of body about 112;
posteaudal groove somewhat chevron-shaped and bordered behind
by an arched row of scales at least four of which are enlarged; a
single row of dorsal caudals, not or but little larger than adjoining
scales, not serrate; tail rather cylindrical, rather indistinctly seg-
mented, the scales forming whorls.
Arm laid forward the fingers reach snout tip; scales on front part
of upper arm, and front face of forearm, larger than ventral body
scales; the scales on front face of thigh, and front and dorsal parts
of tibia similarly enlarged; fingers and toes widened except on
ultimate joint; seventeen widened lamellae on two longest fingers,
the distal joint with eleven scales; 28 widened lamellae under longest
toe with ten scales under distal portion. No axillary depression or
Pouch; tympanum small in size, 125 mm. x .8 mm.
Color in life: Above, generally uniform olive with a golden yellow
stripe beginning on shoulder in front of arm insertion, running back
to groin, becoming somewhat diffuse posteriorly, bordered with
blackish above and below. The dewlap is a deep amber, with a
faint suggestion of olive; venter lighter, with a yellowish wash and
Metallic reflections.
When preserved one notes certain small median dorsal brownish
Spots; the dorsolateral region grayish, variegated with darker and
lighter shades; legs more or less barred with dark and light, and
With a few lighter golden flecks; chin, belly, and base of tail whitish.
Measurements in mm, KUMNH Nos. 36500 and 36501: Snout to
vent, 47, 49; tail, 98, —; total length, 140, —; snout to ear, 13, 13.9;
Snout to arm insertion, 21, 21; length of head, 14, 15; width of head,
7.8, 8; axilla to groin, 19, 20; arm, 21, 23; leg, 39, 41; tibia, 13, 13.
Remarks: 1 collected KUMNH Nos. 36489-36508 on Cocos Island,
June 14-17, 1954. The species was quite common along the forest
edge near the seabeach. | found them in piles of trash, under palm
leaves, and about drift on the beach where there was shade. When
126 Tue Universiry SCIENCE BULLETIN
exposed they usually escaped to trees running up on the trunk
a short distance where they were easily taken.
Only one other lizard is known on Cocos Island, a smaller species,
Sphaerodactylus pacificus Stejneger.
Anolis capito Peters
(Figs. 34, 35, 36)
Anolis (Draconura) capito Peters, Monatsh. Acad. Wiss. Berlin, 1862, pp. 142-
3 (type locality, Costa Rica. Dr. C. Hoffman coll., restricted to Palmar,
Puntarenas Prov. by Smith and Taylor).
Anolis capito Bocourt, Mission Scientifique au Mexique et dans VAmérique
Centrale; Etudes sur les reptiles, livr. 2, 1873, pp. 101-103, pl. 16, figs. 27,
27a (figure of the head and tip of the tail of the type specimen ); Cope,
Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 124 (Old
Harbor [ = Puerto Viejo] Limén Proy.); Boulenger, Catalogue of the liz-
ards in the British Museum, 2nd Ed., vol. 2, 1885, p. 94-95 (Guatemala;
Tabasco to Costa Rica); Giinther, Biologia Centrali-Americana; Reptilia
and Batrachia Oct. 1885, p. 52; Cope, Amer. Philos. Soc., vol. 23, 1886,
p. 276 (Nicaragua); U. S. Nat. Mus. Bull. 32, 1887, p. 30 (places A. carneus
Cope in synonymy); Dunn, Proc. New England Zool. Club, vol. 12, 1930,
p. 18 (Corona, western Panamaé); Barbour, Bull. Mus. Comp. Zool. Har-
vard Coll., vol. 77, no. 4, June 1934, pp. 126-127 (refers Cope’s longipes
to the synonymy of this species); Gaige, Hartweg, and Stuart, Occ. Papers
Mus. Zool. Univ. Michigan, no. 357, Oct. 26, 1937, pp. 8-9 (Rio Mico,
10 mi. above Recero, Nicaragua); Stuart, idem. ibid. no. 471, May 17, 19438,
p. 18 (Guatemala); Misc. Publ. Mus. Zool. Univ. Michigan, no. 69, June
12, 1948, p. 47 (Guatemala localities); Smith and Taylor, U. S. Nat. Mus.
Bull. 199, 1950, p. 65 (Mexican records); Stuart, Misc. Publ. Mus. Zool.,
Michigan no. 91, June 6, 1955, pp. 13-14.
Anolis carneus Cope, Proc. Acad. Nat. Sci. Philadelphia, 1864, p. 169 (type
locality, Lower Verapaz forest, Guatemala ).
Anolis longipes Cope, Proc. Amer. Philo’. Soc., vol. 31, sec. 23, 1893, p. 343
(type locality, Palmar and Boruca, Costa Rica); Dunn, Proc. New England
Zool. Club, vol. 12, Aug. 7, 1930, p. 21 (confuses this species with Anolis
purpurescens stating that they are the same): Burt, Trans. Acad. Sci. St.
Louis, vol. 28, no. 1, Aug. 1, 1933, p. 18 (includes purpurescens as a syn-
onym, possibly on Dunn’s authority ).
Diagnosis: Width of mental equal to or less than width of rostral;
head short, high; semicircular ridges distinct with a deep frontal
depression; suboculars separated from labials by a single scalerow;
six loreal series; two median dorsal caudals on base of tail, then a
single median, the series not serrate; dorsal and lateral scales flat,
smooth, subequal, juxtaposed, hexagonal, all smaller than the
strongly keeled ventrals; leg reaches to eye, tip of snout, or beyond;
occipital scale much smaller than ear, separated from semicircles
by two or three scale series; semicircles at middle separated by
two to four series; about 40 scales and lamellae under fourth toe;
males with small dewlap, not or scarcely indicated in females.
Description of species (From KUMNH No. 34726, Esquinas For-
est Reserve between Palmar and Golfito, Puntarenas Proy. ); Rostral
three times as wide as high, bordered by two labials and six post-
Review or Lizarps or Costa Rica
capito Peters. KUMNH No. 84726, Esquinas Forest Pre
(between Palmar and Golfito), Puntarenas Province, C. R.: actual length,
s ;
Nout to vent, 90 mm.; total length, 261 mm.
128 Tue University SCiENCE BULLETIN
rostrals, the two median largest, and separated from nasal by two
scale series; nostril surrounded by a single narrow scale; frontal
area excavated, declivous; semicircles extend from posterior supra-
ciliary border around the supraocular areas to frontal region, the
anterior scutes much enlarged, the ridge tending to bifurcate near
front of supraocular area, one or two scales separating the series
between eyes; supraorbital area nearly covered with supraoculars,
arranged in about four irregular rows and totaling about 24 scales
of varying size; three supraciliary scales overlapping, the anterior
large, thickened; a double row of smaller scales are along the poste-
rior supraciliary border; canthal scales about four, the two anterior
small; the third is much elongated with a sharp lateral edge; six
rows of loreals, the lowermost continues back and separates the
suboculars from the labials; a small occipital depression; the occip-
ital scale small, separated by three scale series from the semi-
circles, about one fourth area of ear-opening; the scales surround-
ing occipital irregular, the scales lateral to these on the swollen
occipital areas also large, hexagonal; two horizontal rows of tem-
porals; lateral temporals smaller than body scales; a suggestion
of a transverse ridge immediately behind the eyes scarcely indi-
cated mesially; two vertical rows in front of ear noticeably en-
larged; twelve supralabials the tenth below middle of eye, all very
rugose, with the posterior and lower edges of each elevated; about
fifteen infralabials; four or five scale series along infralabials some-
what enlarged; mental a little less wide than rostral, bordered by
two labials and six or seven postmentals; gular dewlap small in
males not or but scarcely indicated in female; scales on chin and
neck much smaller than ventrals; dorsals, nuchals and lateral scales
subequal, a little larger in rump area, all polygonal, usually hexag-
onal, juxtaposed, flat, smooth; head scales, except temporal and lat-
eral occipitals, keeled and somewhat rugose; about 108 scales and
granules in a row around middle of body; ear-opening about five
sixths of eye-opening; arm reaching to insertion of femur; fourth
finger longest, all except first widened, with eighteen widened la-
mellae and 14 scales on distal joint; on distal joint the proximal scutes
are double or triple; leg elongate, the fourth toe reaching slightly
beyond tip of snout; fourth toe with 28 lamellae and 15 scales; on
foot following first toe, a series of enlarged scales extending to heel;
dorsal and anterior scales on upper arm, on dorsal part of lower
arm, on front of thigh, and front and ventral surface of the tibia,
enlarged, keeled. Other scales of limbs may be keeled or in some
parts smooth. Tail with two median ventral series largest; basal
Review or Lizarps oF Cosra Rica
a Anolis capito Peters. KUMNH No. 34721; Los Diamantes,
Guapile , Limén Province, C. R.; actual length, snout to vent, 83.5 mm.: total
€ngth, mm.
130 Tue University SCIENCE BULLETIN
part of tail with two median dorsal scalerows, varied more pos-
teriorly, then continued as a single row; all scales of tail keeled but
the dorsal part is not serrate.
Color (female): Beginning behind occiput a broad dorsal golden-
brown stripe covering 16 median rows of scales, bordered by
blackish brown stripes that unite anteriorly on occiput, each bor-
dered by a golden line on upper edge continuing onto tail;
the sides are greenish yellow, more or less uniform; belly cream to
yellow-white, grayish in gular region; bronzy brown on back of
thigh; a fawn band on arm; side and top of head olive brown; tail
brownish, not barred; after preservation some indistinct darker
streaks are evident on the sides.
Measurements: See table.
Variation: The pattern of males and many females consists of
an olive to olive-brown ground-color on which one discerns three
transverse body blotches and one at the base of tail, bordered
anteriorly by a fine dove-tailing line of black. There is usually
a black or brown chevron-shaped head-band crossing front part
of supraocular regions; another band across frontal area, and a
dim one across snout. The young specimens show that these bands
continue diagonally on the sides of head, the posterior crossing
the eye and terminating near mouth angle; the next forward crosses
lips and continues under chin. The underside of chin shows two
transverse cream lines separated by darker bands; this banding
‘an also be discerned in several half grown and in some adults. In
the younger specimens the tail may be banded in brown and fawn.
The male has a diminutive dewlap not very conspicuous in pre-
served specimens.
A specimen from Golfito (No. 34724), a young male, 72 mm.
snout to vent has the typical head markings scarcely discernible,
and the dorsal blotches almost completely obsolete. There are nine
scales between nostrils and the mental is distinctly smaller than
the rostral.
The most puzzling color variant is a young specimen from Moravia
de Chirripé (No. 34725). There is a hair-fine median golden line
bordered on each side by a narrow irregular brownish lavender
stripe two to three scales wide, with some dark irregular longitudinal
streaks; below this a broken blackish-brown stripe darker along
its borders. The sides have three other series of discontinuous
streaks, while low on sides the markings, more numerous and
shorter are continued to middle of venter. The head marks differ,
Review oF Li S OF ysTA Rica
Fic, 8
6. Anolis capito Peters. KUMNH No. 3 Atirro, Cartago Province,
C. R.; actual length, snout to vent, 72 mm.; tail, 120 mm.
182 THe Universiry SCtENCE BULLETIN
the posterior covering most of supraocular area and the point of
the chevron almost touches occipital. There is a clay-white mark
across the jaw angle and in front of ear-opening. The variation
is puzzling.
Remarks: Specimens are at hand from the following localities:
Cartago Province: KUMNH Nos. 34717-34718, Morehouse Finca,
5 km. SE Turrialba; 34720, Peralta. Limén Province: 34721, Los
Diamantes, Guadpiles; 40572-40576, Suretka; 34722, 34725, Moravia
de Chirrip6. San José Province: 34723, Atirro. Puntarenas Prov-
ince: 84774, Golfito; 34726, Esquinas Forest Preserve between
Palmar and Golfito. The species has been reported in Mexico,
British Honduras, Guatemala and Nicaragua.
Cope described Anolis longipes from Palmar and Boruca. It
would appear that the specimen described is very similar. At first
glance this specimen seems very different from specimens from
eastern Costa Rica. However the particular color pattern (a broad
median golden or golden-brown stripe) occurs in the females of
many species of Costa Rican Anolis.
Cope’s species, Anolis carneus described from “Lower Verapaz
forest Guatemala”, agrees in considerable detail with eastern Costa
Rican specimens, especially as concerns the dorsal coloration.
Seemingly it is a larger specimen than I have seen, having a snout-
vent length of approximately 95 mm. The tail is somewhat shorter
(160 mm.) than the specimen here described. The smallest speci-
men examined measures 30 mm. snout to vent.
A round worm about 1 centimeter in length was found bur-
rowed beneath the skin of No. 10568.
Measurements in mm. and scale data on Anolis capito
INGUDGT EP i yes ya eee as 84726 | 34720 | 34721 34719 | 34724 | 34723 | 84725 | 34718 | 40573
BOX 4 ce ks Vali ae fagl wot RT || eel Hosea HE vee if
TOtal lengtltercr tas isa eo 236 231 222 206 192 120 ae 205
BOUL UO Vent 56 ws Li ern 90 85 83.5 | 82 74 72 50 51 74
LN niin sei ews, Oraiergt emia ra igh anya! 151 147.5 |140 132 120 70 76 131
BOW. tOrGAl cris ai cele rin 25 22.5 | 22 21.2 | 19.5 | 18 14 14.7 | 18.6
Snout to foreleg...... eal oU 33 34 82,3 | 28 27 23 22.5 | 29
AAS, CO BPO i on esa aes .| 38 39 35 35 29 31.5 | 20 22
ELA LOM, waa res eae 26.3 | 24 24 23 22 10.81 15 16 20
Head width.......... esse Ut ya [aL Oneee | Ong | UO WO LO sO lbs O | el) 13.6
Height of head...... 14.2 | 12 Li lodge?) 10,0 | 18 8.2 9 10.3
Petts ci Racha eed 40 44 35 BH 31 33 21 28 36
Leg... 84 80 70 76 60. 61 39 AO, 68.5
ELATED i e52 by 26 23 24 24 20 20 12.6 | 18.4 | 24
Foot and t pevirh ? 35 31 28.5 | 28 24 25 15.6 | 13 26
Scales between ridges........| 2 3 2 2 1-2 2 3—2 2 2
Seales between oc¢cipital and
ridges.... Ely Mehr Gre Vek 3-38 3-3 2+3 3-4 2-2 3-3 3-3 38-3 3-3
Postrostral scales . : y 6 8 6 7 us v6 6 ve
Postmental scales . 6 8 5 5 6 6
Supralabials..... Rare es 12-18 | 12-12 | 12-18 | 10-10 | 12-18 | 11-12 | 12-13 | 18-12
TOPO Bis) y da Princes 5 | 15-15 | 14-15 | 14-15 | 11-18 | 16-16 | 15-15 | 14-14 | 14-14
ey, 106 05 100
Review or Lrzarps or Costa Rica 133
Anolis biporcatus Wiegmann
(Fig. 87)
Anolis biporcatus Wiegmann, Herpetologia Mexicana 1834, p. 47 (type locality
“Mexico”; by restriction Piedra Parada, Chiapas).
Anolis copei Bocourt, Mission Seientifique au Mexique et dans Amérique Cen-
trale, Etudes sur les Reptiles, livr. 2, 1873, pp. 77-78, pl. 15, figs. 10, 10a
(type locality, Santa Rosa de Pansos, Guatemala); Cope, Journ, Acad. Nat.
Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875), p. 121 (83 specimens from Old
Harbour, Costa Rica. “Color in life blue oa brown. This animal haunts
sunny spots on the edge of the forest.”); Boulenger, Catalogue of the lizards
in the British Museum, 2nd Ed., vol. 2, 1885, p. 65-66, (Verapaz, Alta
Verapaz, in Guatemala; Costa Rica, W. Ecuador [one of the types of A.
fraseri]; spelled copii); Giinther, Biologia Centrali-Americana; Reptilia and
Batrachia, Oct. 1885, p. 47 (Guatemala and Costa Rica, [spelled copii] );
Cope, Proc. Amer. Philos. Soc., vol. 23, 1886, p. 276 (Nicaragua); Boulenger,
Proc. Zool. Soc. London, 1898, p. 111 (Paramba, Ecuador [spelled copii] );
Dunn, Proc. New England Zool. Soc., vol. 12, 1930, p. 19 (places obtusirostris
and brevipes in synonymy); Barbour, Bull. Mus. Comp. Zool., Harvard Col.
vol. 77, no. 4, 1934, p. 128; Gaige, Hartweg and Stuart, Occ. Papers Mus.
Zool, Univ, Michigan, no, 357, 1937, p. 9, (Rio Siquia, Nicaragua, 7 mi.
above Rama); Wettstein, Sitzungsb. Akad. Wiss. Wien, Abt. 1, Bd. 148,
Heft 1-2, 1934, p. 21 “Tilleran” C. R. identification questioned by author.
Anolis fraseri. (part.) Giinther, Proc. Zool. Soc. London, 1859, p. 407 (one
specimen of A. copei included in the types from the “Andes of western
Ecuador,” This is figured by Bocourt loc. cit. plate 15, fig. 12, [not fig. 10
as stated by Barbour loc. cit.]; locality uncertain); Boulenger, Catalogue of
the lizards in the British Museum, 2nd Ed., vol. 2, 1885, p. 65.
Anolis brevipes Boettger, Cat. Rept. Senck. Mus., vol. 1, 1893, p. 57 (type lo-
cality, Cairo plantation, La Junta near Limdén, Costa Rica); Ber. iiber
Senckb. Naturforsch. Ges, in Frankfurt am Main, 1893, p. XXVIII and 1893,
p. LXXIIL (mention of type of brevipes).
Anolis obtusirostris Peters, Monatsb. Akad. Wiss. Berlin, Jan. (or Feb.) 1874,
p. 741 (type locality, Chiriqui Prov. Panama); Boulenger, Catalogue of the
lizards in the British Museum, 2nd Ed., vol. 2, 1885, p. 95.
Anolis petersii Schmidt, Smithsonian Misc. Coll., vol. 89, Mar. 16, 1933, p. 9,
satun, Canal Zone.
Diagnosis: Large species (reaching 102 mm. snout to vent), some-
what triangular in cross section; scales on back and sides small,
convex, moundlike, ridged or slightly keeled on back and _ sides,
nearly uniform save for two slightly enlarged median dorsal series;
venter with scales two to three times as large as dorsals, imbricate,
strongly keeled as are the adjoining small scales low on sides; groove
does not completely divide mental; suboculars separated from labials
by a row of scales (or except for one or two points of contact);
head scales generally unkeeled except on snout; leg reaching just
anterior to arm insertion or to ear; head longer than tibia; arm three
fourths of axilla-to-groin distance; occipital depression shallow, dimly
outlined by the occipital elevation or ridges; frontal depression shal-
low; postanals enlarged.
Description of species (From KUMNH No. 34249 Los Dia-
mantes, Guapiles, Costa Rica); Body compressed but little, head
elongate; rostral at least four times as wide as high, bordered be-
134 THe University SCIENCE BULLETIN
hind by five postrostral scales and two labials; ten scales between
nostrils, thick, keeled; frontal depression shallow, continuous with
the interorbital and occipital depressions, its floor covered with flat
scales; circumorbital semicircles forming a low ridge, not or scarcely
discernible on the frontal region, the scales without or with very
Fic. 37. Anolis biporcatus Wiegmann. Tronadero, Lake Arenal, Guana-
caste Province. Left figure, KUMNH No. 40567; actual length, snout to vent,
93 mm.; tail, 213 mm.; right figure, 40568, actual length, snout to vent, 92 mm.;
tail, 200 mm.
Review or Lizarps or Costa Rica 135
indefinite keels, the anterior ones enlarged; behind eye this series is
continuous with the subocular series that terminates in the loreal
region and is separated from the labials by an intercalated row of
scales (that is continuous, save for two points on the right, one point
on the left side, thus permitting contact with the labials at these
points ); four enlarged scales forming a sharp canthus; two elongate
Supraciliary scales followed by two smaller scales; scales of cir-
cumorbital semicircles separated by three (two) scalerows; occipi-
tal depression bordered by two ridges or moundlike elevations, the
area covered with small scales; a transverse row of somewhat larger
Scales continued back of the eye; supraocular region with three
tows of irregularly enlarged scales bearing low keels that are sepa-
rated from the semicircles by a row of smaller scales; nine scales
between canthi in front of the frontal depression; twelve scales be-
tween the first canthal scales; eleven supralabials to middle of eye,
followed by two smaller scales; twelve infralabials; mental with a
wider lip border and greater height than rostral, divided for about
four fifths of its length by an entrant suture and bordered behind
by two labials and eight postmental scales, the two outer not en-
larged; six scalerows bordering infralabials enlarged, heavily keeled;
the other gular series smaller, thickened, and keeled; seven loreal
Series; occipital scale about size of the auricular meatus, separated
from the semicircles by three scales; keels on labials longitudinal; no
enlarged scales on front edge of ear; scales on dorsum and sides
juxtaposed, nearly uniform, save for certain slightly enlarged median
tows, the scales thick, rounded, with a ridge or keel; about 100
scales and granules around body; males with a slight nuchal ridge
Or crest; leg reaches to front of ear; scales of arms keeled, those on
front face of forearm distinctly larger than others; scales on back
of thigh large; on tibia the larger scales are on both front and back
edges, all more or less keeled; ventral scales more than double size
of dorsals, keeled, often mucronate, imbricating; tail slightly com-
pressed with a dorsal series of enlarged scales at top, keeled but
Not spiny or serrate; subcaudals, including those under base of tail,
eeled; postanal depression in males with a chevron-shaped series
of enlarged postanal scales, some larger than others. Broad scales
and lamellae of fingers (excluding distal phalanx) with the follow-
mg formula: 4-8-17-21-12 for foot 4-15-24-35-21; throat fan (dew-
lap) in males small, scarcely reaching breast; its length less than
head length. A throat fan indicated in females.
Color; In life, brilliant yellow-green above, yellowish white be-
low except for a darker area on throat; tail darker, banded with
136 Tue Untversiry SCIENCE BULLETIN
light fawn. In preservative, nearly uniform violet-brown, the ven-
ter lighter gray and fawn with brown longitudinal fine lines on
throat; small flecks and spots of brown on venter and underside of
thighs; the tail dark brown, the narrow light banding only slightly
indicated; a slightly lighter area on back of head, the median and
frontal parts lead color.
Variation: Many of the significant variations are evident in the
data given in the appended table. The scales generally, in the supra-
orbital areas, vary somewhat in number and size. In one specimen
three scales are double the size of adjoining scales.
When specimens are submerged in clear liquid, darker spots are
in evidence. These dark spots on the body differ, and in No.
34249 they seem to coalesce to produce four indefinite dorsal bands,
that extend on sides to venter. No, 34250 has four lighter dots
on the back, the underside of the chin and venter showing but
little pigment. The light bands on the tail vary in their distinctness.
Males have a small throat fan barely reaching the breast. In life
the fan, or dewlap, is white or slightly greenish but preserved speci-
mens show considerable pigment in the fan. The canthi are dis-
tinct in all * and a slight nuchal crest may be indicated. Bocourt
states that the type has the dorsal scales feebly keeled. The keels
are not strong, but very faintly indicated in all cases in Costarican
specimens examined. The hind leg varies somewhat in its reach
but probably never reaches farther than the eye. Color changes
in life are made with considerable rapidity.
Remarks: The specimens in the KUMNH Collection are from the
following localities: Cartago Province: KUMNH Nos, 34247, 34251,
Morehouse Finca near Turrialba. Limdén Province: Nos. 34248,
34249, 34250, Los Diamantes, near Gudpiles, No. 34252, La Lola,
No. 34253, Tunnel Camp near Peralta, No, 40570 Suretka. Guana-
caste Province: Nos. 40564, 4 to 5 km. ENE Tilaran; 40566-40569,
Tronadero, Tilardn, 40571, 4 km. ESE Los Angeles de Tilaran, It
has also been reported from Limon, Costa Rica. Outside of Costa
Rica it has been reported from Guatemala, Nicaragua, Panama,
and Ecuador,
Remarks: When captured No. 40570 had just swallowed a speci-
men of a smaller species, Anolis lemurinus lemurinus. The hind
legs were sticking out of the mouth and it is doubtful that it could
have been completely engulfed until it had been partially digested.
* Dunn (loc. cit. 1930), referring to the type of copei states, “Four scales between
semicircles; no canthus rostralis; small scales in semicircles; weak ventrais,”” all statements
differing from Bocourt’s statements and figure.
ry
Review or Lizarps oF Cosra Rica 187
When removed from the digestive tract the specimen showed no
deterioration due to digestion. The A. capito measures 88 mm.
snout to vent, while the A. 1. lemurinus is 50 mm. snout to vent.
The thick basal portion of the tail, still attached, adds 14 mm. more
to the length,
Anolis lionotus Cope
(Fig. 38)
Anolis (Dracontura) lionotus Cope, Proc, Acad, Nat, Sci. Sci, Philadelphia,
Aug. 1861, pp. 210-211 (type locality, Cucuyas [== Cucuyos] de Veragua,
New Granada [ = Panama]; an abandoned mine on the Rio Santiago, fide
Barbour),
Anolis lionotus Boulenger, Catalogue of the lizards in the British Museum, vol.
2, 1885, p. 70 (repeats Cope’s description ); Dunn, Proc. New England Zool.
Club, vol. 12, 1930, p. 18 (Suretka and La Hondura, C, R.; places rixii and
oxylophis in synonymy); Burt, Bull, Amer, Mus, Nat. Hist., vol. 61, June
11, 1931, p. 260 (Columbia); Barbour, Bull, Mus, Comp. Zool., Harvard
Coll., vol, 77, no, 4, 1935, p. 140-141; Gaige, Hartweg, and Stuart, Occ.
Papers Mus, Zool, Uniy. Michigan, no, 357, 1937, pp, 9-10 (a variant men-
tioned from Nicaragua), ‘
Anolis oxylophus Cope, Journ, Acad, Nat, Sci, Philadelphia, ser, 2, vol. 8, 1876
(1875), p. 123, pl. 24, fig. 4, pl. 28, fig. 5 (SE Costa Rica by inference,
presumably from Sipurio, Limén Proy.); Boulenger, Catalogue of the lizards
in the British Museum, vol, 2, 1885, p. 85; Giinther, Biologia Centrali-
Americana; Reptilia and Batrachia, Oct, 1885, p. 51 (Costa Rica); Cope,
Proc, Amer, Philos. Soc., vol. 23, 1886, p, 276, (Nicaragua); U. S. Nat.
Mus, Bull, 32, 1887, p. 31 (the species lionotus is omitted from this work);
Dunn, Proc. New England Zool. Club, vol. 12, 1930, pp. 18-19; Barbour,
Bull, Mus, Comp, Zool., Harvard Coll, vol. 77, no, 4, June 1934, p, 140-141
(Nicaragua, Costa Rica; western Panama to Canal Zone).
Anolis rixii Boulenger, Proc. Zool. Soc. London, 1894, p. 727, pl. 48, fig. 1
(type locality, Chontales Mine, Nicaragua).
Diagnosis: Medium large, semiaquatic species, characterized by
about 20 rows of enlarged pentagonal or hexagonal dorsal scales,
Occasionally oval or subcircular in shape, with or without slight keels
On some scales; occipital about size of ear-opening; semicircular
Series separated by from one to three scalerows; nine loreal series;
suboculars separated from labials by one scalerow; mental much
wider than rostral, its depth (height) scarcely more than twice
width; a large gular fan (dewlap); ventrals as large as largest dor-
Sals, strongly keeled; postanals somewhat enlarged; usually a large
shallow axillary depression evident; about 115 scales around middle
of body; a cream stripe from eye through ear, passing above arm to
Near groin, bordered below by a brownish-olive stripe and above by
a broad dark olive band, narrowest just behind eye; dorsum faintly
blotched; tail compressed and strongly banded; leg reaches near
to orbit,
Description of species (From KUMNH No. 34157 Turrialba C.
R.): Rostral twice as wide as high, the sides truncate, bordered
behind by two labials and eight postrostrals; about ten scales be-
Fic. 88. Anolis lionotus Cope. Left figure, KUMNH No, 34158, Rio
Reventazon, Turrialba; actual length, snout to vent, 68 mm.; right figure, No.
34157; actual length, snout to vent, 76 mm,
|
\
R
Review or Lizarps oF Costa Rica 189
tween nasal scales which consists of a narrow rim; latter separated
from the rostral by a single scale; supraorbital semicircles composed
of about ten scales forming a low ridge beginning behind eye near
supraciliary border and terminating in frontal area, separated
mesially by three series of scales, anteriorly at termination by six
series; only slight interorbital and frontal depressions; supraocular
area with a clearly delimited group of six much-enlarged keeled
scutes, with four or five smaller scales, separated from semicircles
by small granules; occipital (usually touched by two enlarged
scales) longer than wide, about equal in area to the ear-opening,
separated from the semicircles by three scale series; a large supra-
ciliary followed by one or two smaller ones, scales on posterior
supraciliary border not enlarged; canthal series of four scales; nine
loreal series; suboculars few, enlarged, separated from labials by
one or two rows of scales; nine enlarged supralabials; eleven in-
fralabials the posterior borders of mouth with small granular scales;
a sinuous double series of relatively small median temporal scales;
mental border much wider than that of rostral, its length (height)
in its width, two and one-eighth times; scale divided by a groove
for three fourths of its length, and followed by two labials and nine
postmentals; several scale series bordering infralabials larger than
the median chin series; a large gular dewlap, its depth equal to
about four-fifth times the length of head. About twenty scalerows
on dorsum enlarged, variable in shape from polygonal to subcircular,
many with dim keels or rugosities; outer rows gradually merging
into granules on sides which are generally smooth but become
keeled as they approach the enlarged keeled ventral scales; about
125 scales and granules around body; postanal scales somewhat en-
larged; tail compressed strongly, the scales keeled throughout; the
median dorsal row enlarged, not serrate; two median ventral rows
also enlarged; only a very faint elevation of the two median nuchal
rows; nine series of loreal scales, keeled; scales on snout not keeled
but the edges may be slightly elevated; a large deep postaxillary
depression; digits not greatly widened; 16 lamellae under wider part
of fourth finger, ten scales under its distal joint; arm not reaching
groin; leg, laid forward, the longest toe reaches to edge of orbit;
scales on dorsal part of upper arm, front and dorsal part of forearm,
front and dorsal part of thigh, and most of the scales on tibia much
enlarged and strongly keeled; ear small, 1.2 mm. x .9 mm.
Color in life: Above, nearly uniform olive with faint indication of
& series of dorsal blotches; a darker lateral olive-brown band on
which are some hieroglyphlike marks and an indefinite series of
140 Tue Universiry SCIENCE BULLETIN
distant small ocelli, which extend from neck to near groin; a cream
line from eye to near groin; a small brown line from ear to mouth
angle; labials light bronze. All under surfaces nearly uniform light
cream with a yellow wash; dewlap light orange. Tail banded in
dark and light olive; regenerated part of tail blackish; limbs barred
with fawn.
Measurements in mm.: Snout to vent, 71.5; tail (partly regen. ),
92; snout to ear, 18; snout to arm insertion, 30.5; axilla to groin, 33;
length of head, 20; width of head, 11.6; arm, 29.5; leg, 53; tibia, 20.
Variation: This species is somewhat variable in color but the fe-
males do not seem to develop the median linear markings so com-
mon in many groups of Anolis. However the character of the dorsal
blotching, and the length of the lateral light line is variable, but
not necessarily a sex character. In many young specimens, espe-
cially, there is a series of black hieroglyphs in the brown lateral
stripe. One just above arm insertion may have a deltoid shape;
others may be similar to the letter O; with these there is a series
(not always discernible) of small ocellate cream marks which may
persist in preserved specimens. The tail banding may be brown
and fawn, without olive. A specimen from Isla Bonita (40579 ¢ )
has the dorsum uniform gray-olive, and the brown lateral band
(almost black on neck) has become partly broken up into a series
of irregular dark-edged spots; the tiny ocelli occupy spaces between
the brown spots. One from Turrialba (No. 34159 9? ) has the
band broken (or partially so) into eight spots. These are con-
nected across the neck, back, and base of tail by as many narrow
brown chevrons, the angles directed backward, tiny ocelli are like-
wise present between spots. Specimens from Guanacaste (Tilaran
and Tenorio) have the dark ventrolateral area strongly punctate
with cream. The head markings usually show a median lighter-
olive area on snout with darker olive on each side, a transverse
chevronlike band across head, and other marks behind eyes failing
to meet mesially. A specimen from San Isidro del General has a
bright cream spot running under the eye to ear. Variation in the
character of the supraoculars, which range in number from four
large subequal keeled scales to ten of variable size does not seem
to be geographical. The degree of keeling or rugosity on the dorsal
scales, likewise variable, may or may not be due to geography. It
seems to differ some with age. The ventral scales are always
strongly keeled. The lateral white line may reach the groin or it
may terminate shortly behind the level of the axilla. The axillary
|
Review or Lizarps or Costa Rica 141
depression is strongly evident in fully adult specimens, but not
necessarily so in young specimens.
Remarks: Cope described Anolis lionotus from Cucuyos, Veragua
Province, Panama in 1861, and Anolis oxylophus from Southeastern
Costa Rica either from lowlands or the sides of Pico Blanco, in
1875, Certain subsequent authors have regarded the forms as dis-
tinct; others (Dunn and Barbour) have placed oxylophus as a
Synonym of lionotus.
Unfortunately I do not have available Panamanian material. The
differences are seemingly significant, the most important being, that,
as described, lionotus has smooth dorsals, smooth ventrals, twelve
instead of twenty rows of enlarged scales and the longer hind leg
reaching to the nostril; while in the other form it scarcely touches
the orbit. While I am here tentatively regarding the two as syn-
onyms, I consider the matter still unsettled.
The habitat of the species is usually along pools or streams. In-
dividuals may be seen perched on rocks amidstream or at the edge
of the water. When disturbed on land they run to the water, swim
to some rock if one is near or dive under the water and emerge at
some point some distance away.
The Costa Rican material available to me consists of the follow-
ing specimens. Limén Province: KUMNH Nos. 34163-34166, 34259-
34260, Los Diamantes, Guapiles; 40581-82, Suretka; 34155, 34261-62,
34268, Moravia de Chirripé6; 34162, 34265, La Lola. Cartago Prov-
ince: 25652-54, 34258, La Suiza; 34153-54, 34157-61, Turrialba. He-
redia Province: 25656-59, 34156, 40579, Isla Bonita. Guanacaste
Province; 40580, 4 to 5 km. ENE Tilaran; 34167, 4 mi. NW Tenorio,
Las Cafias. San José Province: 25655, 2 mi. SW San Isidro del Gen-
eral; 34263, 9 mi. west San Isidro del General on Dominical road
(possibly in Puntarenas Province ).
I propose to fix or restrict the type locality of A. oxylophus Cope
to Sipurio, Lim6én Province, C. R.
Anolis aquaticus* sp. nov.
(Fig. 89)
Type: KUMNH No. 34276, Palmar, Puntarenas Province, Sept. 3,
1953; John Baker, Coll.
Paratypes: KUMNH Nos. 34277, 34278, Golfito, Puntarenas Prov-
ince; Edward H. Taylor Coll.
Diagnosis; A whitish line on upper lip running back and above
tympanum meeting its fellow on nuchal line; a light line from mouth
* From Latin aquaticus, living in or near water.
142 THe Universtty SCIENCE BULLETIN
angle to groin; tail strongly compressed, triangular in cross-section;
a median row of enlarged serrate scales on tail; two larger median
rows of scales on back; ventral scales strongly keeled; semicircular
ridges strongly pronounced, the enlarged scales reaching canthus,
but the ridges do not; the suboculars separated from the supralabials
by two (or three) scalerows; three or four scalerows between
Fic. 89. Anolis aquaticus sp. nov. Left figure, KUMNH No. 84277 ¢4,
Golfito, Puntarenas Province, C. R.; actual length, snout to vent, 61 mm.; right
figure, type, Palmar, Puntarenas Province, C.
the supraorbital semicircles; three irregular rows of keeled supra-
oculars; ear-opening large, only slightly smaller than eye-opening;
a strong axillary depression; tail annulated to tip; body with lateral
dark brown bars, dim on median line, separated by lighter tan
transverse marks; male with large orange-colored dewlap.
Review or Lizarps or Cosra Rica 148
Description of type: Rostral about three times as wide as high,
its labial border distinctly less than that of mental, and bordered
behind by eight small scales, separated from scales surrounding
nostril by two scalerows; five small scales around nostril; seven
supralabials, each with a longitudinal keel; circumorbital semi-
circular ridges curved posteriorly, then running nearly straight for-
ward, terminating on forehead, but large scales of series continuing
to canthal edge, these often with several transverse ridges or keels;
Occipital scale a nearly perfect oval bearing “eyespot,” about half
size of tympanum, separated from enlarged scales of ridges by about
six scalerows, the area surrounding scale flattened and on about
the same level as the depression between ridges which continues
to beginning of elevated part of snout; scales on snout ridged or
keeled, on forehead, less so; supraoculars in about three irregular
rows, the inner row largest, all with one or more keels, the group
Surrounded by fine granules but only one row separating inner
Supraoculars from ridges.
Canthal series begins above eye, the first two scales very small,
followed by a distinct row of four large keeled canthals, the
keels forming nearly a straight line, terminating some distance
behind nostril; a series of eight suboculars bordering lower orbital
tim, separated from supralabials by two or three small scalerows;
eight enlarged supralabials terminating below middle of eye; pos-
terior border of mouth bordered by minute granules; mental a
little more than twice as wide as long with a median suture enter-
ing from behind, almost completely segmenting the scale; infra-
labials nine, extending back a little farther than supralabials; ten
or eleven rows of scales in loreal area; four or five rows of scales
lying along infralabials, slightly enlarged but no distinguishably
enlarged row; six scales border the mental besides the labials, the
two outer a little larger than others.
Occiput covered with fine granules, smaller than those surround-
ing occipital; two median rows of scales somewhat enlarged, the
20 succeeding lateral rows smaller or some may be larger laterally,
all distinctly larger than the granules on sides; about 140 scales
about body; ventral scales much larger than dorsal scales, imbricate,
keeled, the largest ones on sides of dewlap; no enlarged postanal
Scutes; two median rows of subcaudals largest, strongly keeled;
Median dorsal caudal row enlarged, not or but slightly serrate; all
Caudal scales keeled; tail segments number 29, eight whorls of
Scales to a segment; largest scales on arms and legs about size of
Median dorsals; 16 widened lamellae under the two longest toes;
144 THE Universiry SCIENCE BULLETIN
under the terminal joints, ten or eleven; 23 widened lamellae under
longest toe; eleven lamellae under terminal joint of same toe, total-
ling 34 lamellae in all. Tympanum vertically oval, 2.2 mm. x 1.4
mm.; depth of dewlap equal to length from tip of snout to arm
insertion; tail compressed, triangular in cross-section; length of tibia
equals or slightly exceeds head length; no enlarged scales on eyelid.
Color in life: Above olive to olive-brown; back with a series of
seven dark transverse bands, laced with cream, which become less
distinct dorsally; a median dorsal row of light marks; head olive
brown; a broad cream line from snout tip, involving most of tym-
panum, joins its fellow on median nuchal line; a light cream stripe
from jaw angle above arm to groin, and bordered below by a series
of narrow blackish diagonal lines bearing a few cream dots, and
separated by diagonal cream lines; arms, legs, and digits banded
with fawn or cream; chin, and most of venter cream-white with
some little yellow; dewlap orange. Tail banded with broad brown
bands separated by olive, the dark bands growing wider posteriorly;
a dark band on the infralabials; under side of limbs cream white.
Measurements in mm. (KUMNH Nos. 34276 ¢, 342773,
34278 9 respectively): Snout to vent, 65, 61, 54; tail, 97, —, 90;
length of head, 18.2, 17, 15; width of head, 10.5, 10.2, 9.8; snout
to ear, 16, 15, 14; snout to arm insertion, 25, 24, 21; axilla to groin,
28, 26.7, 24.4; arm, 29, 29, 27; leg, 58, 51, 44; tibia, 18.5, 17.2, 14.5.
Remarks: The narrow diagonal dark lines low on the sides are
separated by narrow cream lines of equal width and the dark lines
themselves have rows of small cream dots. The female has no
dewlap. The axillary depression is less conspicuous in the two
paratypic specimens.
The type is from Palmar, the other two (34277-78) are from
Golfito. All were taken at night, presumably asleep, clinging to boul-
ders in rivulets just above the water. Certain others seen during the
day escaped by diving and hiding below water. The figures are of
the two male specimens; that on the right is the type.
One suspects that the relationship of the form is with lionotus.
They agree in being semiaquatic in habit, each has a cream lateral
line, and an orange dewlap. The differences are that lionotus has
dorsal scales that are larger (nearly double the size) and instead
of being keeled are smooth; the dorsum is flattened rather than
compressed, and the tail is less strongly compressed than in the
form at hand.
From Cope’s Anolis oxylophus there is also some similarity in
t
i
{
|
Review or Lizarps oF Costa Rica 145
the lateral line extending from the scapular region to near the
groin and in having the dorsal scales keeled. In the middle of
the back there are certain median rows a little larger than the others
(equal to ventrals) but the adjoining 16 rows are larger than the
lower lateral scales with considerable irregularity among them-
selves. Differences in coloration markings are striking. It would
appear that both are semiaquatic. The auricular opening is ap-
proximately only half as large as in the described form.
Anolis pachypus Cope
Anolis pachypus Cope, Journ, Acad, Nat, Sci, Philadelphia, ser, 2, vol. 8, 1876
1875), pp. 123-124, pl. 24, fig. 3, (type locality “slope of Pico Blanco,”
Limén Province, Costa Rica, William Gabb, coll.); Boulenger, Catalogue of
the lizards in the British Museum, 2nd Ed. vol. 2, 1885, pp. 63-64 (copied
description); Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
Oct., 1885, p. 47 (no specimens); Dunn, Proc. New England Zool, Club,
vol. 12, 1930, p. 16 (Volcin Barba C. R.; places A. curtus and A. tropi-
dolepis in synonymy); Barbour, Bull, Mus, Comp. Zoology, Harvard Col.,
vol. 77, no. 4, 1935, pp. 144-145; Wettstein Sitzungsb, Akad, Wiss., Wien,
Math-natur. KI., Abt. 1, Band, 143, Heft, 1 and 2, 1934, p. 20 (Volcan Irazu,
Volein Pods, Bebedero, San José, La Carpintera. [The report from Bebe-
dero, near sea level is surprising for this species]. )
I have no authentic specimen representing this species, in the col-
lections at hand. The type, USNM No. 30683, is in bad condition,
since it has been dried and somewhat rotted, the legs are broken
off, the mental is missing, and consequently it does not serve for a
redescription. Hence I append the data from the original descrip-
tion.
Description of species (Data from type, or type description ) :
Head rather short, its length equalling that of the tibia and one-half
greater than its width; supraorbital semicircles composed of scales
no larger than the fine series of scales that separate them; no ridges
but a depression in occipital region continued between orbits to a
rather deep frontal concavity; scales in frontal concavity a little
smaller than those which separate them from canthal row; six can-
thal and supraciliary scales extending to near middle of supraciliary
area; supraorbital region covered with projecting granules except
a single row of three or four small keeled supraoculars near the
Supraorbital border; four rows between supraorbital series and
Occipital scale; eight loreal rows; one series of very small infra-
labials, the gulars running up to them in longitudinal lines; ear-
Opening larger than the occipital scale.
Scales of abdomen smooth, those of sides very small, on the back
gradually enlarging to two or three largest and faintly keeled rows
6—3378
146 Tue Universtry SCIENCE BULLETIN
on the median line which are much’ smaller than those on venter.
Scales on top of head subequal in front; keeled; smaller and angular
behind, covering a three-sided area behind the orbits, which is
abruptly distinguished from the minute granules of the temples
and nape.
The feet are stout and clumsy; the wrist reaches to the front of
orbit; third toe of hind limb reaches end of muzzle. The digital
dilations are little or not more expanded distally than proximally
and the distal joints are covered with wide keeled scales above;
gular fan well developed.
Tail cylindrical with equal scales, swollen at base.
Color: Emerald-green with a broad brown band from orbit to
middle of side, and a narrower one on each side of sacrum, A green
band from orbit to below tympanum, and a dark-brown band from
orbit below canthus rostralis; a dark-brown cross-band between the
supraciliary borders across frontal area; a deep brown spot behind
occipit, and a smaller one on the nape; femur with a longitudinal
black band behind, and an oblique one across the superior face;
tibia and forearm with a broad cross-band.
Measurements in mm.: Total length, 135; snout to vent, 47; snout
to axilla, 20; snout to tympanum, 11; arm, 22; hand, 9; leg, 43; foot,
19; tibia, 12.
Remarks: Until sufficient new material from the Pico Blanco
area is available there will be some doubt about the validity of the
distinction of this southern form. The type specimens from an
unknown elevation on Pico Blanco differ from most Costa Rican
specimens available in having the ventrals smooth, and the third
toe of the hind foot reaching the end of muzzle. The supracili-
aries [ = semicircular series] not larger than the scales of the five
rows which separate them; a single row of three or four small
keeled supraoculars. Its relationship would seem to be with Anolis
tropidolepis Boulenger.
Anolis tropidolepis Boulenger
(Fig. 40)
Anolis tropidolepis Boulenger, Catalogue of the lizards in the British Museum,
2nd Ed., vol. 2, 1885, p. 53 (type locality Irazti [Voleén?]); Giinther,
Biologia Centrali-Americana; Reptilia and Batrachia, Oct. 1885, p. 46, pl.
27, fig. C. (figure of type).
Diagnosis: Ventrals keeled; longest toe not reaching nostril or
tip of snout; scales of semicircles distinct, polycarinate, separated
by from three to five scales; supraoculars numerous in two or more
_*
147
w oF Lizarps or Costa Rica
ni
Revie
“WU GPT [e}0} “WU GP ‘JUSA 0} NOUS “Y4SUa] [eNjoe “vourTg vIvA IvoU ‘UOps|9Z
eourg “CEgoP ‘eInsy YWnoF “uu QFT ‘YSUeT [e}o} “WU GF ‘UeA 0} NOUS ‘Y}ZUe] [eNjoOR ‘vouLTG vIeA W TW E
“6O0LOF ‘1NSYy psy} “Wu QET “YSUe]T [e}0} “UIUT YE ‘JUBA 0} NOUS “Y{SUET [eNJoe ‘equeg uvo[oA jo edojs oyloeg uo
jezureg eeu
€
SLOOP ‘eInSy puoodss “WUT ZZT “YIBusT [eo] “WU TE “UeA 0} JNOUS “YIBUaT JenJOR “YY “D “eoulAOIg
vipsisy{ “eourlg eiVA | WE € ‘TLLOF ON HNINAY ‘omsy ysiq “lesuet~nog sidajopido.y syouy ‘OF “O14
148 Tue University SCIENCE BULLETIN
rows; scales of area anterior to supraorbital region with polycarinate
scales (two to five keels); head longer than tibia.
Description of subspecies (From KUMNH No. 40730 near Finca
Zeledon, near pass between Volcan Pods and Volcin Barba): Ros-
tral three and one-fifth times wider than high, bordered behind by
seven rugose postrostrals and two labials; ten scales between nasals,
the latter separated from rostral by two scale series; semicircular
series begin near inner level of supraocular region, run forward to
the canthal edges, separated mesially by four scale series; about 16
scales between canthals; scales of frontal and snout areas with from
one to five keels, sometimes more on the semicircle scales anteriorly;
three larger, and four or five smaller supraoculars; five rather broad-
ened canthals, the keels not forming a continuous line, continuous
with two elongate superciliaries almost completely overlapping each
other; these in turn are followed on the posterior supraciliary region
by three series of slightly enlarged supraciliaries, the median series
largest; nostril surrounded by a single narrow rim strongly over-
hung by a supranasal; about four enlarged suboculars, one or two
touching the labials (usually completely separated from labials );
seven or eight supralabials to below middle of eye; nine infralabials;
five scale series below infralabials, larger than median rows on
chin; eight loreal rows; mental wider than rostral about two and
one-half times as wide as high, bordered behind by two labials
and six postmentals the two outer largest; scales on sides of throat
only slightly enlarged; occipital scale about one third of the size
of ear-opening separated from semicircles by four scale series;
occipital ‘area slightly depressed bounded behind and laterally by
two swollen areas covered with fine granules; scales of temporal
region very small, the scales of the upper temporal border not
noticably enlarged, ventral scales rather rounded behind, keeled,
the scales not flat; scales on side granular but on dorsum becoming
somewhat larger; four largest along median line only about half
as large as ventrals and rather distinctly keeled; about 160 scales
in a row around body; tail broken and partly regenerated. (In
another specimen, tail widened at base; tapering suddenly, cylin-
drical, attenuated; median dorsal scalerow not or scarcely larger
than other caudal scales; no enlarged postanals; the postanal de-
pression shallow; gular dewlap in male shorter than head); dewlap
not developed in female; ear-opening relatively large, vertically
oval, about 1.5 mm. x 1.1 mm.; fingers scarcely widened distally;
toes noticeably widened; about twenty-two lamellae and scales
Review or Lizarps oF Costa Rica 149
under fourth finger; thirty-six under longest toe; arm reaches groin
or a little farther; leg reaches to anterior border of orbit; scales
on dorsal part of upper arm smaller than those on upper surface of
forearm; scales on front face of thigh equal or larger than scales
On posterior and ventral surfaces of tibia.
Color: Above generally olive with three or four indefinite narrow
brown chevrons across back; a lateral yellowish-brown stripe vary-
ing in width and at places marked with dim lighter brown mottling
or dim diagonal lines; a greenish-olive stripe from supraciliary
, Fic. 41. Anolis tropidolepis color var. Left figure, KUMNH No. 40727,
S mi. E Vara Blanca; actual length, snout to vent, 50 mm.; right figure, 40672,
Boquete samp (2,000 m.) west slope Cerro de la Muerte; actual length,
Snout to vent, 43.5 mm.
150 Tue Universiry SCIENCE BULLETIN
border back onto neck; a similar diagonal line’ from eye across
angle of mouth to foreleg; a greenish-olive mark below eye more
or less continuous with similar area along chin below infralabials;
venter bronzy yellowish washed with brown; greenish flesh under
thigh; chin dirty white; thigh and tibia with a diagonal brown
lighter-edged band. Gular appendage in male orange to reddish,
somewhat narrow at base; a brown band across head, angular
mesially, with a lighter center; front of snout olive.
Measurements in mm.: Nos. 40724 9 and 40704 ¢. Snout to
vent, 50; 46; tail, reg., —; 78; snout to ear, 18.7; 11.8; snout to leg
insertion, 19.4; 18; axilla to groin, 22; 19; head length, 16; 14; head
width, 9.8; 8.8; arm, 24; 21; leg, 42; 39; tibia, 12; 11.8.
Variation: The color changes are rapid. In preservation the
specimens show considerable variety of patterns. Females often
have a very narrow dark line or stripe on the median dorsal line
connecting with a black occipital spot; when this is present there
are no head bands and the lateral brown stripe may be obsolete.
Bodies of males are distinctly compressed and the dewlap is
usually purplish red in life. There is considerable variation in the
character of the scales of the semicircular series. Rarely one
subocular may touch the labials. Specimens from the northern
part of the Talamanca range (along the Pan-American Highway )
the semicircles are largest and most strongly keeled. The series
from Heredia Province north of the Cordillera Central have the
scales small with a lesser amount of keeling, and separated by more
scale rows,
Remarks: Specimens occur only at relatively high elevation. 1
have taken them from about 5,000 to 8,000 feet. They are likely
to be on the ground or near the ground on tree trunks. In Heredia
Province many were found about or on stumps six to eight feet high
on which many plants were growing.
The following specimens have been collected. Cartago Province:
KUMNH No. 40674, Quebrada Onda; 40681, 40683, Pacayas; 40687-
89 Finca Quirazi. San José Province: 40714-40719, La Carpintera;
40681, 40682, near Empalme, south of Cartago; 40675-40676, Na-
tional Forest on Pan-Amer. Highway south of Cartago, 6,000-7,000
ft. elev.; 40672, Boquete Camp 2,000 m. west slope Cerro de la
Muerte, 40677, south slope Irazii northeast of San José. Heredia
Province: 40698-40713, 40722-40725, 40726-40734, Vara Blanca and
vicinity; 40794-40797, Finca Zeled6n near pass between Volcan Pods
and Volcan Barba; 40678, near Carizal; 40679-80, Finca Monte-
christo.
Review oF Lizarps or Costa Rica 151
Anolis curtus Boulenger
(Fig. 42)
Anolis curtus Boulenger, Proc. Zool. Soc. London Noy. 29, 1898, p. 919, pl. 55,
fig. 2, 2a (type locality La Estrella, Cartago, Cartago Province, Costa Rica,
C. F. Underwood Coll. ).
Diagnosis: Head a little longer than tibia; frontal ridges low, or
obsolete, the scales of the semicircles rather small, separated mesially
by from five to seven scale series; eight loreal series; occipital smaller
than ear-opening, separated from semicircles by six series of scales;
gular dewlap very small; mental divided by a groove; two rows of
scales between small suboculars and supralabials; two bands of
brown across head (in males); tail much inflated at base, then sud-
denly slender, cylindrical; length, snout to vent, 51 mm,
Description of species (From KUMNH No. 4837, Finca Monte-
christo, Volein Barba, circa 5,800 ft.): Rostral not or scarcely visi-
ble from above, about four to five times as wide as high, followed
by two supralabials and eight small scales; rostral separated from
nasal by three scale series; nine scales between nostrils; no dis-
tinct ridges, the semicircles composed of somewhat enlarged multi-
keeled scales, larger however than surrounding scales, the series
reaching canthus; anteriorly they are separated by about ten series
of scales at narrowest part by five or six scale series; supraciliaries
rather small except two anterior; these preceded by six narrow
subequal canthal scales, their outer edges not forming a continuous
edge; about eight loreal series; a series of six or seven small sub-
Ooculars, separated from the labials by two rows of scales; ninth
Supralabial and eighth infralabial below middle of eye; supraoculars
Small, irregular in size in about four rows; all smaller than adjoining
Circumorbital séries, and separated from those scales by a row of
granules; occipital enlarged but smaller than the auricular opening,
Separated from the semicircles by five or six scale series; occipital
area very slightly depressed; temporal area covered with very small
granular scales; five or six somewhat enlarged scale series bordering
the infralabials; mental divided by a deep groove into two trian-
Sular moieties, the rostral border greater than that of mental; two
labials and six postmentals border mental; auricular opening verti-
cally oval (1.4 mm. x .8); no nuchal crest or fold; about four dorsal
scalerows larger than the laterals but there is no strong line of
demarcation, since they gradually merge into the fine granules of
the sides: approximately 148 rows of scales and granules about
body; ventral scales small, thin, larger than largest dorsal scales,
keeled, imbricating, in about 22 rows, however there is no defined
Tue Universitry SCrENCE BULLETIN
Fic. 42. Anolis curtus Boulenger. KUMNH No. 4837, Finca
Montechristo, Volcan Barba, circa 5,800 ft. Actual length, snout
to vent, 49 mm.; total length, 126 mm.
Review or Lrzarps or Costa Rica 158
line of demarcation; caudal scales keeled, the median dorsal row not
serrate, the two ventral rows larger than the median dorsal row;
postanal depression without distinctly enlarged postanals; scales on
top of upper arm, front of forearm, and front of thigh much enlarged;
Scales of under side of tibia larger than other tibial scales; longest
toe reaches to near nostril; postanal region strongly inflated, be-
hind which the tail tapers suddenly; thirty three lamellae and scales
under fourth toe; under middle fingers 22-24.
Measurements in mm.: No. 4837 and Type. Snout to vent, 49,
51; tail, 77, 56 regenerated?; total length, 126, 107; snout to ear, 12.2,
—; snout to foreleg, 21.6, —; length of head 15.2, 15; width of head,
8.7, 9; axilla to groin 20.5, —; arm 22, 21; leg, 40, 39; tibia 12.2,
Color in life: Above generally brown lacking a stripe but with a
frosted silvery region on dorsum from occiput to tail with some
fine black flecks, more distinct on tail (type has black marks all
along median line of back); top of head silvery with metallic re-
flections, lighter in the depressed area of head and on tip of snout;
labials yellowish or cream; chin and gular appendage dark olive
brown; belly heavily pigmented and with bluish or greenish metallic
reflections; tail gray brown with some lighter flecks along the dorsal
part; extreme tip regenerated, deep black; an indefinite dark mark
on thigh and a band on tibia.
Remarks: This specimen taken in the same area with Anolis tropi-
dolepis shows some similarities to that species. The differences
however, appear to outweigh the similarities. Boulenger named this
form curtus, but I strongly suspect the tail of the type specimen
is regenerated, thus incomplete—hence the apparent difference in
actual length. However the reduced size of the suboculars sepa-
tated by two scalerows from labials instead of one, six instead of
four canthals that are distinctly narrower, five to seven instead of
three or four scale series between semicircles, smaller and more
Numerous supraoculars, all point to a species or subspecies differing
from tropidolepis.
Anolis achilles * sp. nov.
(Fig. 43)
Type. KUMNH No. 40634 La Palma, San José Province, C. R.,
June 20, 1954. Edward H. Taylor, Coll.
Diagnosis: A small anole characterized as follows: sixth or seventh
Supralabial and sixth infralabial below middle of eye; five suboculars
: Referring to the Greek hero, Achilles who had a specialized heel,
154 Tue Universrry SCIENCE BULLETIN
in contact with the supralabials; six loreal series; tail with a dorsal
row of moderately enlarged supracaudals, not serrate, the tail very
slightly compressed; head much longer than tibia; longest toe reach-
ing ear; about 130 scales and granules around middle of body; semi-
circles consisting of seven or eight large scales reaching canthal
edge, separated mesially by one row of scales; only largest supra-
ocular touches the semicircle, in contact with two or three scales;
posterior supraciliary scales not differentiated; seven or eight
median dorsal scalerows slightly enlarged, but not half as large as
ventral scales; a few much enlarged scales on heel, two to three
times larger than adjoining scales.
Description of type: Rostral two and one-half times as wide as
high, bordered behind by two labials, four postrostrals and seem-
ingly by the two nasals; six scales between the two nasals on snout;
semicircles consist of seven scales the fourth normally largest, the
series terminating anteriorly with the anterior canthal, the individual
scales of the series keeled, and anteriorly they are also somewhat
rugose, between orbits separated by a single series of small scales,
anteriorly separated by seven scales.
Scales on snout keeled, the scale edges are often elevated sug-
gesting a tricarinate condition; three keeled canthals relatively
wide and rugose, the outer keel forming a rather sharp edge; anterior
supraciliary scarcely longer than median canthal, but overlapping
a small second supraciliary; about half of supraciliary border cov-
ered with fine granules; occipital scale about two and one-half times
area of ear-opening, separated from semicircles by two scalerows;
supraoculars in two rows, the inner curved, consisting of four scales;
all except largest supraocular separated from the semicircles by
small granular scales; second row of three scales; six (or five ) loreal
series; five suboculars strongly enlarged, the largest almost touch-
ing the labial border; seven (or eight) supralabials, the sixth or
seventh below middle of eye; the scales elevated on their lowe
border; seven infralabials, the sixth below middle of eye; mental
with a considerably larger labial border than the rostral, divided
by a groove for three fourths of its height, the height in width two
and one-half times; the whole scale bordered by two labials and
six postmentals, the two outer enlarged; four rows of scales below
infralabials somewhat enlarged, rugose or keeled; ear-opening sub-
triangular; ventral scales dimly keeled in about twenty rows,
gradually merging into small lateral granules which, as they ap-
proach the dorsal line, are again slightly enlarged, the two median
Review or Lizarps or Cosra RIcA 155
discernibly the largest; no medial ridge; no axillary pockets; body
slightly compressed; relatively large calcium deposits present be-
hind ear (possibly somewhat abnormal); limbs short: the lamellae
under digits widened, except on distal part; lamellae and scales
under fourth finger, 14 + 7; under fourth toe, 24 +- 8; scales on
Fic. 43. Anolis achilles sp. nov. KUMNH No. 40634
2, type; La Palma, San José Province, C. R.:; actual
length, snout to vent, 45 mm.; tail, 77 mm.
oO
"pper arm, on front and dorsal surfaces scarcely more than half as
arge as those on the front of the forearm, where they are as large
on ventrals; scales on front of thigh essentially as large as those on
tibia often with two, rarely more, keels; scales on under side of
ree] two to three times the size of adjoining scales bearing four to
SIX low keels: longest toe reaches ear; hand brought forward, the
Tue Universiry SCUENCE BULLETIN
longest finger fails to reach the tip of the snout; dorsal caudal row
somewhat enlarged, not serrate; two ventral rows larger, all keeled;
tail subcircular or slightly compressed.
Color: The female specimen preserved in formalin is, save for a
few brownish flecks, nearly unicolor dorsally; on the side, however,
there is a distinguishable lighter stripe bordered above and below
by a brown line; the forearm is dimly banded with brown and the
hind limb shows some darker brown flecks on the dorsal surface,
and there is slight evidence of darker and lighter areas on the latter
half of the tail; under side of tail lavender; ventral surfaces of body
and head dull white with scattered pigment; occipital region black-
ish.
Measurements in mm.: Snout to vent, 45; tail, 77; total length,
122; snout to ear, 11; snout to arm insertion, 19; axilla to groin, 19;
head width, 7.3; head length, 13.1; arm, 18; leg, 31; tibia, 10.
Remarks: Only a single specimen was taken. The presence of
the widened heel plates has not been discerned in other species of
the genus. The presence of the enlarged visible auricular deposits
of calcium is of common occurrence in the Gekkonidae and Sphaero-
dactylidae. It is of less frequent occurence in Costa Rican [guani-
dae.
Anolis godmani Boulenger
Anolis godmani Boulenger, Catalogue of the lizards in the British Museum,
Ind Ed., vol. 2, 1885, pp. 85-86 (type locality Guatemala* and Iraztii Costa
Rica [here restricted to Volcin Irazii, Costa Rica]); Cope, U. S. Nat. Mus.
Bull, 32, p. 31 (listed); Dunn, Proc. New England Zool. Club, vol. 12,
Aug. 7, 1930, p. 20 (seemingly regarded this as a synonym of Anolis limi-
frons, regardless of its keeled ventral scales).
Diagnosis: Ventrals keeled; head slightly longer than tibia; uppet
head scales very small, uni- or tricarinate; semicircles separated
by two to four scales; occipital larger than ear-opening, separated
from semicircles by from three to five scales; loreal rows six to
eight; dorsal scales very small, subrhomboidal, keeled; ventrals
larger, roundish-hexagonal, imbricate; hind leg reaches nearly to
end of snout; tail cylindrical, nearly twice head-body length; »°
enlarged dorsal caudal series; upper lip yellow, bordered above
by a sharply defined brown band passing through eye and above
ear; crossband between eyes.
Description of species (From type description): “Head once and
three fourths or twice as long as broad slightly longer than tibia;
* Stuart (1955) seemingly disregards Boulenger’s record of the species for Guatemala.
Review or Lizarps or Costa Rica 157
snout rather pointed; forehead and interorbital region concave;
frontal ridges short and divergent; upper head scales very small,
uni- or tricarinate; scales of the supraorbital semicircles enlarged,
Separated by two to four series of scales; a few enlarged, keeled
supraocular scales; occipital larger than the ear-opening, separated
from the supraorbitals by three to five series of scales; canthus
Tostralis sharp, canthal scales three or four; loreal rows six to eight;
six upper labials to below the centre of the eye; ear-opening mod-
erate, vertically oval; gular appendage moderate; gular scales
keeled. Body not or scarcely compressed; no dorso-nuchal fold;
dorsal scales very small, subrhomboidal, keeled, graduating into
the minute granules of the sides; ventrals larger, roundish-hex-
agonal, imbricate, keeled. Limbs slender; the adpressed hind limb
teaches nearly the end of the snout; digits feebly dilated; thirteen
or fourteen lamellae under phalanges II and III of the fourth toe.
Tail cylindrical, nearly twice as long as head and body, without
vertebral series of enlarged scales. Greenish-brown above; upper
lip yellowish, bordered above by a sharply defined brown band
Passing through the eye and above the ear, vanishing behind the
shoulder; a dark cross band between the eyes; sometimes a few
small distant brown spots on the vertebral line.” Total length, 107
mm.; head, 12; width of head, 6; body, 26; forelimb, 16; hind limb,
34; tibia, 11; tail, 69.
Remarks: Boulenger had four Guatemala specimens (male and
halfgrown ), and three from Irazi, Costa Rica (male and female),
in his type series of A. godmani. The species agrees fairly well
with limifrons in many of its characters. One presumes that it is
mountain form while limifrons is a lowland form probably not
Securring much above 2,000 ft. elevation. The exact elevation from
Which the types of godmani were obtained is not recorded but it is
safe to believe that they came from above 5,000 ft. This species
has been omitted from the key,
I propose to restrict the type locality to Volesn Irazu, C. R.
Genus Potycurus Cuvier
Polychrus Cuvier, Régne Animal, vol. 2, Ist ed. 1817, p. 40 (type of genus
Lacerta marmorata Vy
Diagnosis: Body compressed, lacking both nuchal and dorsal
Crests; head scales somewhat enlarged; body covered with small
imbricating scales; no gular fold; gular sac in males; tympanum
distinct; femoral pores present in both sexes; digits compressed with
Subdigital. lamellae :carinate; four large scales near the base of the
158 Tue Universiry SCIENCE BULLETIN
claw; tail much elongate, nearly circular in cross-section. Ptery-
goid teeth present; lateral teeth tricuspid; abdominal ribs present.
A single species is recognized in Costa Rica, present on both the
Caribbean and Pacific slopes of the country. It remains rare in
collections.
Polychrus gutturosus Berthold
(Fig. 44)
Polychrus gutturosus Berthold, Abh. Ges. Wiss. Gottingen, vol. 3, 1847, p. 5,
pl. 1, fig. 1 (type locality, Colombia); Boulenger, Catalogue of the lizards
in the British Museum, 2nd Ed. vol. 2, 1885, p. 100 (Colombia and Costa
Rica); Cope, Journ, Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875),
p. 124; U. S. Nat. Mus. Bull. 32, 1887, p. 32: Giinther, Biologia Centrali-
Americana; Reptilia and Batrachia, Oct. 1885, p. 53.
Polychrus (Chaunolaemus) multicarinatus Peters, Mon. Akad. Wiss. Berlin,
1869, p. 786 (type locality, Costa Rica); Bocourt, Mission Scientifique
au Mexique et dans l’Amérique Central; Etude sur les Reptiles, livr. 2 and 3,
1873, 1874," p. 112, pl. 17, fig. 8.
Diagnosis: Lateral scales smaller than dorsals or ventrals; no
longitudinal serrated ridge on front of throat; ventral scales multi-
carinate (two to six) smaller than the gulars; semicircular supra-
orbital series larger than other supraorbital scales, but smaller than
scales on snout.
Description of species (From KUMNH No. 25170): Body com-
pressed, somewhat triangular in cross-section; head angular, with
a sharp canthus rostralis and a sharp ridge above and following eye;
rostral low, its width about two and one-half times its height;
bordered behind by two median striated scales, two labials and one
scale in front of nasal; semicircular series in supraorbital region
consist of nine scales on left, ten on right side, separated mesially by
a single row of scales; supraoculars in about five rows, the inner
scales larger than outer; eight supraciliaries; five larger scales on the
temporal ridge; nostril in a single nasal separated by a single scale
from rostral, equidistant between eye and tip of snout; a supra-
nasal; frontonasal broken into three parts; ten or more scales in the
frontal area, somewhat symmetrical; occipital scales smaller than
those on snout, the “occipital” scale somewhat elongated bearing
an indistinct “eye”; scales on snout and supraorbital region striate,
while in occipital region scales almost or completely lacking stria-
tions; three canthals between eye and nostril; six or more scales in
loreal region; preocular, subocular, and postocular series continuous,
small, totaling 14 scales; a second larger subocular series just above
labials; about eight rows of scales in temporal series; supralabials
* The second page of this description was published in the third livraison which
appeared in 1874,
Review or Lizarps or Costa Rica 159
9-9; infralabials 8-8; mental having as wide a labial border as rostral
but smaller; a suture enters mental from behind; a pair of enlarged
postmentals followed by four scales; scales of the gular pouch
smooth, separated by soft granules; nostril equidistant between eye
and snout-tip; scales on head, chin and lips with numerous striations
or low irregular keels; scales of middle ten rows of body larger
than laterals, with one or more keels; lateral scales imbricating, oval
Fic. 44, Polychrus gutturosus Peters. KUMNH No. 25170, Turrialba,
nti Province, C. R.; actual length, snout to vent, 163 mm., total length,
Oo mm,
C
160 Tue Universitry SCIENCE BULLETIN
or pointed posteriorly, usually with a single keel and usually pointing
back and somewhat downward; lower on side scales with more
than a single keel; ventral scales usually with two to five keels, larger
than the lateral scales; scales on posterior face of femur with a single
keel, elsewhere with two or more; scales of the tail strongly imbri-
ating, larger than body scales, with a single sharply defined keel.
Limbs slender, the digits compressed, the third and fourth fingers
nearly of equal length as are the third and fourth toes; claws angu-
lar; base of claw surrounded by a large dorsal and three other
scales; subdigital lamellae or scales widened, with numerous keels.
Hind leg brought forward, the longest toes reach axilla or slightly
beyond; anal flap semicircular, its scales a little larger than the
scales that precede them. Femoral pores in female small, the
inner ones somewhat enlarged, the ten outer scarcely perforating
the scales; tympanum distinct; auricular opening usually smaller
than eye opening; a few projecting lobules in front of auricular
opening.
Color in preservative: Head and body purplish brown but darker
on sides of head below and behind eye; a white spot preceded by
a darker spot on upper lip; a cream diagonal line from eye to angle
of mouth and a cream spot below mouth angle; a cream stripe
low on shoulder passes onto the base of arm. A broad irregular
cream stripe from axilla to groin, continued on front of thigh as a
line or series of cream flecks; a cream spot or stripe barely indicated
at base of tail, but none on back of femur; skin between gular
scales bluish (?); venter purplish brown; tail light yellowish-tan
with about twenty indistinct darker bands suggested.
Measurements of Polychrus gutturosus
10660 hes
25170
Moller adn page Peau nae 685 583 —_—
Grout to Vent es ere 163 133 153
Tail ees Ua ree 522 450, 338+
Length of litad ..00 38 32 85
Width of head ....... a ale de Plat PUM by}
A ee iad 61 — 63
Lee ee 85 oe 19
TN ee ei ota ey tn) — —
AU NC OH ey ire ited 16 — 14
Q1 — 20
MUNGO) oe cia a
* Type of Peter’s Polychrus (C.) multicarinata. KUMNH No. 25170, Turrialba, Costa
Rica; USNM No. 70660, Tilaran, Guanacaste, Costa Rica.
Variation: The small number of specimens that have been ex-
amined tells little regarding variation. The color markings of this
specimen described (Turrialba, Costa Rica) differs from “multi-
Review or Lizarps oF Costa Rica 161
carinata” of Peters in lacking the stripe of yellow on the posterior
side of the femur, and the olive yellow venter. It does have a
broad irregular cream stripe from axilla to groin on the outer
ventral surface, with a broad purplish-brown median area from
chin to vent.
A specimen in the U.S. National Museum from Tilaran, Guana-
caste has a cream stripe six scales wide, from occiput to tail. It is
brownish on the sides, and below this a bluish olive. The tail is
light with elongate darker spots alternating on each side.
In the latter specimen there are 32 lamellae under the fourth
finger, 44 lamellae under fourth toe. The specimen figured is a
female containing four ovarian eggs in the right, five in the left
ovary, These aré nearly spherical and measure 12 mm. in diameter.
The tail is 3.2 times the head-body length probably exceeding
that of other lizards in proportionate length.
The stomach contents consist chiefly of insect remains with a
meager portion of vegetable matter ingested, perhaps accidentally.
Corythophanes Boie
Corytophanes (sic) Boie in Schlegel, Isis von Oken, vol. 20, pt. 3, 1826, p. 290
(type of genus Agama cristata Merrem).
Chamacleopsis Wiegmann, in Gray, in Griffith, The Animal Kingdom ey
by Baron Cuvier . . . vol. 9, 1831, Synopsis, p. 45 (type, Chamaeleop-
sis hernandesii Wiegmann ).
Diagnosis: “Tympanum distinct. Head with two sharp ridges
uniting posteriorly into an elevated bony crest directed upwards
and backwards; a bony supraorbital arch formed by the prae- and
postfrontal. Body compressed, covered with imbricate smooth or
keeled scales; a nuchal crest, attached to the cranial crest; a low
dorsal crest. A gular pouch and a transverse gular fold. Infra-
digital lamellae keeled. No femoral pores. Tail long, feebly com-
pressed, Lateral teeth tricuspid; pterygoid teeth. No sternal fon-
tanelle. No abdominal ribs.” [from Boulenger, Cat. vol. I, p. 101.]
A single species of this genus is known in Costa Rica, its range
extending from Mexico to Panama.
Corythophanes cristatus Merrem
(Fig. 45)
Agama cristata Merrem, Tentamen systematis amphibiorum ..., Marburg
1820, p. 50 (type locality “Ceylon”, ex errore; restricted to Orizaba, Vera-
_ eruz, by Smith and Taylor).
Corythophanes cristatus Gray, in Griffith, the Animal Kingdoms 1, by
Baron Cuvier, Synopsis 1831, p. 55 (Cytophanes (sic) and Cyrtophanes
(sic); Gravenhorst, Acta Acad, Leop.-Carol., vol. 16, pt. 2, 1838, p. 938,
pl. 65, figs. 6-10; Wiegmann, Herpetologia Mexicana 1834, p. 15; Duméril
and Bibron, Erpétologie Générale, vol. 4, 1837, p. 174; Fitzinger, Systema
Reptilium, 1843, p. 52; Gray, Catalogue of the specimens of lizards in the
162 THe Universiry SCIENCE BULLETIN
collection of the British Museum, 1845, p. 194 (“inhab. Mexico”); Duméril,
and Duméril, Catalogue Méthodique, Museum Paris, 1851, p. 60 (Peten
specimens); Duméril, Arch, Mus, Hist. Nat. Paris, vol. 8, 1856, p, 517;
Bocourt, Mission Scientifique au Mexique et dans l’Amérique Centrale;
Etudes sur les reptiles, livr. 3, 1874, pp. 118-119 (Peten specimens); Cope,
Journ. Acad, Nat. Sci. Philadelphia, ser. 2, vol. 8, 1876, (1875) p. 124
(Sipurio, Costa Rica); idem. ibid., p. 156 (Buhio Soldado, Panama);
Giinther, Biologia Centrali-Americana; Reptilia and Batrachia, Oct. 1885,
p. 53 (Guatemala, Costa Rica, Panama); Boulenger, Catalogue of the lizards
in the British Museum, 2nd Ed., vol. 2, 1885, pp. 101-102 (Guatemala to
Costa Rica); Cope, U. S. Nat. Mus. Bull. 32, 1887, p. 33 (East Costa Rica;
Gaige, Hartweg and Stuart, Occ. Papers Mus. Zool. Michigan, no. 357, 1937,
p. 10 Rio Mico, 10 mi. above Recero, Nicaragua); Smith and Taylor, U. S.
Nat. Mus, Bull. 199, 1950, p. 69 (Corytophanes) (Veracruz to Costa Rica).
Corythophanes mexicanus Cope, Proc. Amer. Philos. Soc., 1884, p. 183 (fide
Cope).
The figure designated Lacerta ceilonica cristata et pectinata in
Seba’s Thesaurus, vol. 1, p. 147, pl. 94, fig. 4, served Merrem as the
type of this species. Merrem presumably saw no specimen. It is
obvious that the locality, Ceylon, is erroneous. Smith and Taylor
have restricted the type locality to Orizaba, Veracruz, Mexico.
Corythophanes percarinatus A. Duméril is reported (with a ques-
tion) from Ecuador, by Boulenger. The species is certainly known
from Guatemala. It should be looked for in southern Central
America.
Description of species (From KUMNH No. 34308, Golfito, Costa
Rica): Head and body compressed; a fine rather indistinct fold
from jaw angle, curving above arm along side to groin; a lateral
fold low on the sides from axilla to groin; skull casquelike with
sharp canthal ridges continued above and behind eye until they
meet and then continue on to the tip of the bony casque; from this
high elevation a nuchal crest descends diagonally to the body and
is continuous with the low dorsal crest on the body.
Head longer than wide, the snout narrowed and rather blunt; ros-
tral about twice as wide as high, bordered behind by two labials, and
a pair of postrostrals, separated mesially by a small scale; nostril in
an enlarged nasal, the opening directed nearly backward, situated
about equidistant from orbit and tip of snout, separated from the
rostral by postrostral and two vertical series of scales, from canthals
and labial each by a row of small scales; scales on snout, frontal,
and occipital areas irregular, not keeled; semicircular supraocular
series differentiated, each consisting of ten or eleven scales, sepa-
rated mesially by at least two irregular rows of small scales; three
or four of the supraocular scales are larger than the others; canthal,
supraciliary, and postocular ridges continuous; beginning with the
anterior elevated canthal scale above nasal, there are twenty scales
Review or Lizarps oF Cosra Ric
_ Fic. 45. Corythophanes cristatus Merrem. KUMNH No. 36121, Suretka,
pumén Province, C. R.; actual length, snout to vent, 104 mm., total length,
30%
325 mm.
Tue Universiry SCIENCE BULLETIN
164
on each lateral crest anterior to their junction, beyond which there
are eight scales to the summit of casque; area enclosed by the lateral
ridges depressed, the supraorbital areas not elevated; about eighteen
scales in loreal region in about five rows; supralabials, 12-15, the line
of mouth behind eye level, bending down somewhat; infralabials
12, the posterior ones small; all labials somewhat tubercular; the
scales below infralabials definitely keeled with from one to three
keels; eye-opening diagonal to line of mouth, about size of tym-
panum; eyelids continuous around eye, covered with very nu-
merous small granules; scales on temporal region and on the sides
of the elevated part of casque irregular in size, without keels except
three or four enlarged scales that bear tubercles or short keellike
ridges; tympanum a vertical oval, twice as high as wide; a distinct
nuchal fold in front of shoulders; scales on chin forming parallel
longitudinal rows, separated by minute granules; median row form-
ing a low serrate crest; gular pouch present, inconspicuous; scales
on side of body somewhat irregular, unkeeled except for more or
less distinct vertical series of larger scales that bear a low keel ter-
minating in a slightly elevated tubercle or spine; scales below second
lateral fold are larger, forming transverse rows, strongly keeled,
mucronate.
Scales on limbs enlarged and keeled; preanal scales only slightly
differentiated; caudal scales larger than ventrals, monocarinate;
basal part with vertical series of larger scales marking whorls; no
caudal crest. Limbs moderate, the arm extended forward, the
wrist reaches the nostril; the leg extended, the longest toe reaches
tip of snout; digits clawed, the base of claw surrounded by two
enlarged scales; lamellae under digits with single mucronate keels;
scales on palm and sole each with a pearly tubercle or mucrone;
no femoral or preanal pores.
Color in life: Top of head greenish as are most of the scales on
ridges and crests; on sides of crests a few yellow-green or brick-red
scales; body generally blackish, with single scales or groups of
yellow scales low on sides continuous with yellow spots on sides of
venter; these spots with darker brown edges. Arms and_ legs
greenish or yellow-green with a yellow-cream spot under upper
arm, and a very distinct white spot or spots at elbow; a series of
dark lines more or less radiating from eye, crossing line of mouth,
separated by broader interspaces that are greenish above and
brown. below, except the last one which is all green. Tympanum
surrounded by greenish scales; clay-white irregular spots under
thighs; chin brownish black, the median serrate series tipped with
Review or Lizarps or Costa Rica 165
yellow; tail brown with eleven magenta-red blotches not forming
bands; a brick-red spot in the postfemoral region extending on to
tail; some very indefinite darker banding on sides barely suggested,
a black x-mark on head between eyes.
Measurements of Corythophanes cristatus
34808 34306 tee 34305 34807 34309
Total Length ae ees 358 367 345 261 167
SUOUU TO VONy ee kLe 4 oat 118 107 104 104 84 56
ee eer ee 240 260 221 239 ide bila
Length of head to behind jaw 82 4 83 81 23 16
Length to end of casque..... 56 55 55 53 34 30
Width of head—greatest.... 20 6 19.8 22 16 13
Height of casque from
throat level 4 70 a ae 44 — 43 Al 29 28
CMa ei ait aaltepet cake 51 53 50 48 4] 31
DEG ae se) RY ay 109: 110 100 98 78 56
Ulotini(dar Wehayeyeil ee 15 ANGE 13.6; 2-13 12 9
OUR EOG ota ee en: 25 25 24 22 20 15
Variation: A female from Turrialba (Caribbean drainage, eleva-
tion 1900 feet), shows some squamational differences from the speci-
men described. The rostral scale length is one and one-half times its
height, (not twice ); the mental has a wider labial border than rostral
(smaller); bordered by two scales (by three); serrated scale series
on chin smaller, less conspicuous; radiating lines from eye wanting,
or not now discernible; and the reddish spots on tail are dim, scarcely
discernible,
A specimen taken at Suretka, near the Panama border differed
most from the described specimen. The top of the head is lined
and flecked with black, and the area about the eye is black with
the radiating lines forming angles at lips, and not meeting on the
chin and throat, A series of three cream spots are present on the
curving lateral fold above the arm. There are few or no enlarged
scales below the lateral fold in the anterior half of body, but numer-
ous on the posterior part, and bluish in color. Many of the scales
on limbs and on the base of tail are dark edged. The entire side
and crests have very numerous black dots or flecks. The yellowish
Spots on the venter are obsolete. The markings on the limbs are
typical. The spines at the junction of the nuchal with dorsal body
crest are very small and likewise small on latter fourth of body. The
light dorsal marks on tail are not evident. Light cream areas below
the eye between the black radiating lines are strongly defined,
A young specimen (KUMNH No. 34309, 55 mm. snout to vent)
has the following markings: very light olive-green becoming very
light brown on body; dim radiating darker lines from the eyes
166 THe Universiry SCIENCE BULLETIN
meeting on the throat or chin. A lighter area followed by a darker
area on the sides of the casque above the arm, The sides have a
series of five wide bands, the first and third with a black spot. Large
reddish-cream areas about the base of the tail extending slightly
onto the back of the femur. The tail is faintly banded, the anterior
edge of each band being darker, the band becoming a little lighter
posteriorly. The dorsal reddish blotches are indistinct or absent;
the venter is more or less banded. Typical light spots are present
on arms and legs.
Remarks: The described specimen is a female taken September
9, 1952. When first observed she was depositing her eggs in a cavity
dug in the hard soil of a forest path. Five eggs had been deposited
and a sixth was laid after capture. When laying her eggs she was
seemingly quite oblivious to my presence and was picked up with-
out resistance and made no attempt to escape. A second female
was discovered by my friend John de Gooijen of San José, Costa
Rica, depositing eggs in a similar cavity dug in the hard earth in a
forest path. Six eggs had been laid. This: specimen, captured alive,
was killed and partially destroyed, by. a forest rat that gained ad-
mittance to the laboratory-through the ceiling.
On Aug. 13, 1951, I captured a specimen (KUMNH No, 34305)
at the I. A.J. A. Farm at Turrialba, Cartago Province. I was hunt-
ing salamanders in a rotten log on the ground and this lizard
descended the truck of a nearby tree and came to rest, head down,
at a point eight feet from the ground where it remained motionless,
watching my movements. After a time I arose quietly, approached
the tree slowly, then reached up and apprehended the lizard. It
made little or no struggle when captured.
The color changes began immediately and were rather rapid,
going from green and olive to brown and blackish. The ventral
and caudal markings and the spots on the arms and thighs are evi-
dent in the preserved specimen.
The specimens at hand are. distributed; as follows: Limén
Province: KUMNH No. 36121, Suretka. Cartago Province: 34305-
34306, 34309, Turrialba. San José Province: 34307 San Isidro del
General. Puntarenas Province: 34308, Golfito.
Cope (1875) reports a specimen from Sipurio, Limén Province
and I have examined a specimen in the Museo Nacional de Costa
Rica from Jiménez, Puntarenas Province.
My specimen, taken at Suretka, had an egg case of a spider
plastered to the posterior part of the dorsal concavity in the head,
leaving the head with the appearance of being flat in that region.
|
|
Review or Lizarps or Costa Rica 167
Genus Basiniscus Laurenti
Basiliscus Laurenti, Specimen medicum, exhibens synopsin reptilium . ‘
Viennae 1768, p. 50 (type of genus Basiliscus americanus Linnaeus, by
monotypy ); Stejneger, Copeia, 1936, No. 1, Novy. 15, 1936, p, 136.
Diagnosis; Large arboreal lizards with strong sexual dimorphism;
tympanum distinct; no femoral pores; body compressed, covered
with small juxtaposed and imbricating scales; males with a head-
crest or “helmet”, and usually with a dorsal and caudal crest
Supported by bony rays or spines, the crest however may be re-
duced to a serrate ridge; gular pouch small or wanting; one or two
transverse gular folds; tail long, compressed; compressed toes bor-
dered by a scaled dermal expansion or fringe; lateral teeth tricuspid;
pterygoid teeth present; no abdominal ribs.
Four species of this genus are known to occur within the confines
of Costa Rica. The name americanus used by certain authors for
this species was a revival of a pre-Linnaean name given by Seba
(Thesaurus, vol. 1, pl. c. fig. 1, “Draco arboreus volans Americanus
sivi Basiliscus”. It would appear that the figure in Seba served
as the type of the Linnaean Lacerta basiliscus. The specimen
itself was later presented to the Paris Museum and served as the
type for Basiliscus mitratus Daudin (Rept., vol. 3, 1803, p. 310, pl.
42),
Kry To THE SpEcIES OF BaAstuiscus Iv Costa Rica
Ventral scales keeled, less distinctly so in males than in females; a
single triangular head crest, as high as long; dorsal crest of male low
(less than 10 mm,); usually a cream or reddish line from behind eye
to a point on shoulder, or may continue to rump. . Basiliscus vittatus
VOTraleeuales GINOOth co 1 ek ee ee
Crest on head consisting of two parts, the anterior smaller; a high
dorsal crest, and a rather high crest on tail, the upper edges of which
are scalloped; one or two rows of lateral yellowish or golden spots
may be present on sides; eye red........... Basiliscus plumifrons
IMOHG Ghect not Givided itd TWO DATES . 409.69. ks ce. 8
A high dorsal crest and a high caudal crest, the upper edges serrate
but not scalloped, the crests covered with thin, somewhat enlarged
SCRIGS | erates Glin eal: cg oo Meee te ere Basiliscus basiliscus
No high dorsal crest but a serrate series of compressed trihedral tu-
bercles, touching or separated by smaller scales; a low, serrate ridge
OU NOE iv ci meny nl 6 Naan nes Basiliscus galeritus
i)
to
Basiliscus plumifrons Cope
(Fig. 46)
Basiliscus plumifrons Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8,
1876 (1875) pp. 125-127, pl. 25, fig. 1 (type locality, Sipurio, Costa Rica);
Boulenger, Catalogue of the lizards in the British Museum, 2nd Ed. vol. 2,
1885, pp. 108-109; Giinther, Biologia Centrali-Americana: ‘Reptilia and
168
THe UNIversiry SCIENCE BULLETIN
Batrachia, Oct. 1885, p. 55 (no specimens); Ahl, Aquar. Berlin, 1930,
pp. 133-136 (photo); Wettstein, Sitzungsb. Akad. Wiss. Wien, Math-naturw.
Kl. Abt. 1, and Bd. 143, Heft 1-2, 1934, p. 22 (Finca Hamburg, Rio
Reventaz6n; and Finca Waldeck); Gaige, Hartweg, Stuart, Occ. papers Mus.
Zool., Univ. Michigan, no. 357, Oct. 26, 1937, p. 10 (Rio Siquia, 7 mi.
above Rama; and Rio Escondido, near Rama, Nicaragua).
Fic, 46, Basiliscus plumifrons Cope. Figure (1 and lad) (from Cope,
Journal of the Academy of Natural Sciences, Philadelphia, ser. 2, vol. 8, 1876
[1875], pl. 25). Presumably type, presumably from Sipurio, Limén Province,
Costa Rica (figure 2, upper left is B. mitratus); actual length, 655 mm.
Review oF Lizarps or Costa Rica 169
Diagnosis: A large arboreal lizard; eye red; head with a small
Crest arising from between posterior part of eyes, and more or less
continuous at its base with a high crest which arises over occiput
and terminates about middle of neck; a.dorsal and caudal crest; two
nuchal folds; the dorsal crest begins at level of arm insertion and
continues to above level of femora, supported by 17 bony spines,
first low, the highest equal to depth of body; caudal crest continuous
but partly separated from dorsal by a deep notch, the upper edge
of caudal crest s scalloped.
Description of species (From R. C. Taylor Collection No. 491,
Turrialba, Cartago Prov.): Rostral large, nearly twice as wide as
high, its labial border a little greater than that of mental, bordered
behind by a single pair of enlarged postrostrals and the first labials;
nostril in a single nasal, the latter separated from rostral by two
Scales, from first labial by only one scale, from its fellow by eight
scales; supraorbital semicircles differentiated, not extending behind
orbit; separated by from two to three rows of scales mesially;
Supraoculars small, in twelve or thirteen rows; scales in median
Supraorbital area a little larger than those surrounding it; most of
the dorsal head scales with single keels or the larger with several
he ridges or keels; “occipital” scale small, immediately preceding
the anterior cranial crest; three canthal scales; about twelve supra-
Ciliaries, the anterior horizontal but not sharp-edged; frontal scales
similar to those on snout; loreal region vertical, the scales in about
four tows; pre- and subocular series enlarged somewhat, the sub-
Sculars the largest, separated from suborbital labials by a row of
small scales, and larger than these labials; supralabials eight-nine,
the seventh below middle of eye; eight infralabials, the mouth angle
above and below with small granules. Mental bordered behind by
pair of chinshields; three anterior pairs of chinshields in contact
With infralabials; scales under chin small, slightly larger on the small
ular sac,
Anterior head crest begins at back level of eyes, its base reaching
back to anterior level of tympanum; at this point the larger cranial
crest begins, rising diagonally rather than vertically the upper part
extending farther back than its base which terminates in a small
distinct transverse fold about middle of neck; upper parts of crests
Covered with larger scales than the top of head; dorsal crest begins
a little behind the cranial crest, rises gradually to a height more
Or less equal to the height of body and supported by bony spines
18 or 19 in number; the upper edge of crest somewhat scalloped;
170 Tue Universiry SCIENCE BULLETIN
caudal crest more or less continuous with dorsal but a deep notch
between,:near posterior level of femur; caudal crest confined to
basal third. of tail only, its upper edge scalloped, supported by
eighteen or:nineteen bony spines directed up and backwards; tail
strongly compressed; crests of back and tail with thin scales, not, or
only slightly, imbricate, pointing up and somewhat backwards; occi-
put with a fleshy elevation; below mouth angle a fleshy swelling;
tympanum smaller than eye-opening, strongly diagonal, the upper
part forward.
Scales on temporal,area, sides of neck and. body variable in size
but all small, definitely smaller than ventral scales; scales on outer
part of arm, and, scales on front of femur, above tibia and on foot
larger than those,gp posterior parts of femur, and all keeled; scales
on tail (not on gyests ) keeled; no femoral pores; preanal and post-
anal scales smooth; digits compressed, the toes on their outer edges
with a narrow,-scaled fringe present on both sides of outer toe;
about 40 subdigital lamellae under longest toe, variable in character,
some ridged or with ventral knobs. The hind limb laid forward, the
longest toe reaches to tip of snout.
Color: Above generally greenish, becoming bluish on the crests;
eye bright red; a row of golden spots beginning on the head con-
tinue to groin along the sides somewhat below the base of the dorsal
crest. A second similar row more or less distinct low on sides; tail
with lighter and darker bands, not indicated on distal third, Venter
somewhat lighter.
Measurements in mm.: Total length, 714; snout to vent, 194; tail,
520; snout to arm insertion, 85; axilla to groin, 75; length of head, 58:
width of head, 32; arm, 90; leg, 181; longest toe, 52; anterior head
crest from anterior base to distal point, 33; occipital crest from be-
ginning to most distant point, 50; height of dorsal crest, 40; of long-
est spine, 45; height of caudal crest, greatest, 37.
Remarks: This species is not uncommon about Turrialba and
most of the specimens were obtained through the courtesy of Mr.
Curtis Good in the immediate environs of his home at Finca La
Dominica, located in the eastern edge of Turrialba. The lizards
there have become almost domesticated so that often one could
approach to within a few feet of specimens in shrubs or on fences.
The yellow or golden spots vary somewhat in their distinctness.
Females lack the dorsal and caudal crests but they may have some
distinct dark markings across the back, and a tiny head crest with
even the small anterior segment in evidence.
Review or Lizarps or Cosra Rica 171
Basiliscus vittatus Wiegmann
(Figs. 47, 48)
Basiliscus vittatus Wiegmann, Isis von Oken, vol. 21, p. 373 (type locality,
Mexico, restricted by Smith and Taylor to Veracruz, Veracruz, Mexico );
Herpetologia Mexicana, 1834, p. 40, pl. 15; Bocourt, Mission Scientifique
au Mexique et dans lAmérique Centrale; Etudes sur les reptiles, livr. 8,
1874, pp. 129-181, pl. 17, fig. 3 (skull; see this work for early synonymy
and literature); Cope, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8,
1876 (1875), p. 125 (Nicaragua and Costa Rica); Boulenger, Catalogue
of the lizards in the British Museum 2nd Ed. vol. 8, 1885, pp. 109-110
(synonymy); Giinther, Biologia Centrali-Americana; Reptilia and Batrachia,
Oct. 1885, pp. 55, pl. 28 (Mexico to Ecuador); Ahl, Aquar. Berlin, 1930,
pp. 133-136 (photo); Fink, Aquar. Berlin, 1931, pp. 97-100; figs.; Mertens,
Abh. Senck. Natur. Gesel., no, 487, 1952, pp. 45-46, pl. 6, figs. 21-22; pl. 12,
: fig. 72 (El Salvador localities ).
Cristasaura mitrella Gray, Ann, Mag. Nat. Hist., ser. 2, vol. 10, 1852, p. 489
(type locality, Honduras ).
Draconura bivitatta Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1860, p.
482, (type locality, Nicaragua, restricted by Smith and Taylor to Greytown,
Nicaragua ). :
Basiliscus (Cristasaura) nuchalis Cope, Proc. Acad. Nat. Sci. Philadelphia,
1862, p. 181 (type locality, Greytown, Nicaragua ).
Diagnosis: A smaller member of the genus. Scales on venter
strongly keeled; nasal touching one, often two, supralabials. Head
crest on males subtriangular rising from a point between anterior
borders of tympanum. Dorsal crest very low, supported by short
bony spines; caudal crest a serrated ridge; a longer or shorter lateral
longitudinal reddish or yellowish band usually present; black marks
on back usually visible.
Description of species (From KUMNH No. 10152): Body strongly
compressed; head rather narrow; rostral large, bordered behind by
.
first supralabials, a scale preceding each nasal, and a pair of post-
rostrals which meet mesially; an elongate supranasal separated from
its fellow by five scales; two canthals continuous with the supra-
Ciliaries of which there are seven; supraocular “semicircles” but little
curved, consisting of seven-eight larger scales, separated mesially
by three rows of smaller scales; scales on snout and frontal area
Variable in size, rugose but not keeled; supraocular scales in about
eight rows, the inner ones larger than outer, bearing an irregular
keellike ridge; “occipital” scale slightly enlarged, bearing “eyespot,”
lying at the beginning of the cranial crest; nasal large, touching one
or two supralabials; nostril vertically oval, directed outward; two
Postnasals; loreal region vertical, covered with four or five scale-
tows, the upper the largest; two or three preoculars and one or two
Suboculars, at least one touching the supralabials; eleven differenti-
ated supralabials to mouth angle, the sixth below the eye; about
nine infralabials the sixth below middle of eye; mental with a
*(punorsypeq ) opewuisy ‘( punorBd10}) spew “gz “jd
‘eIporneg pure eyndey “euRoLIOWY-I[eRqUaT) vIsojorg “reyquny WOIF SINSIY “UULUISETA\ snjnjj10 SnNOsSIUSDG "LP “OIA
BULLETIN
SCIENCE
‘a
5
5
Zz
—_
r
|
i
|
|
Review or Lizarps or Cosra Rica 178
labial border at least as great as the rostral, bordered behind by two
short series of chinshields, only the first pair in contact and these
likewise touching first infralabial; scales between chinshield rows
subequal forming slightly diagonal rows the free edges of the scales
directed mesially; a fold in front of breast, and a slight fold indicated
in front of this; gular pouch very small or wanting; auricular open-
ing probably as large as eye-opening, oval, diagonal; scales in the
temporal region in twelve to thirteen rows with an indication of a
dim ridge running back from eye.
A bony or cartilaginous ridge arises from occiput forming a
“helmet” and is continued as a crest terminating at middle of neck;
immediately behind this point a dorsal body crest begins, at first
represented by a single row of serrate s sales then becoming ele-
vated and supported by bony spines; its greatest elevation scarcely
greater than greatest diameter of the tympanum; the dorsal crest
more or less continuous with the caudal crest, that extends two thirds
of the length of the tail, somewhat lower than the dorsal crest, but
likewise serrate on its upper edge; tail strongly compressed.
Scales on the sides of the crest larger than other lateral body
scales, directed upward and backwards; most of the scales on limbs
enlarged, equal to or larger than ventrals, which in turn are larger
than lateral scales. Scales on limbs strongly keeled; scales on upper
Surface of hands, feet and digits with from two to five keels; digits
Compressed with a fringe of scales on outer edge of digits, usually
touching the subdigital lamellae; subdigital lamellae irregular, each
scale bearing a dark-colored knob or tubercle except those near the
claws; 35-36 lamellae under fourth toes; scales on the ventral sur-
face of the body dimly keeled (strongly in females); subcaudals
more strongly keeled. Limbs strong, the leg laid forward, the fourth
toe reaches a little beyond tip of snout; the knee and elbow do not
touch.
Color in life: Dorsum generally brownish or olive-brownish above,
somewhat variegated in shade; tail lighter brown with dim indica-
tion of banding on distal portion; below, dull purplish gray with
some cream mottling under limbs. Each ventral seale has a little
more pigment in center than at edges; a light, slightly reddish brown
stripe begins behind eye and continues back along side of neck
to shoulder; a second line indicated on side of neck back of tym-
panum.,
Measurements in mm. of KUMNH Nos. 40977 and 40979: Snout to
vent 126, 113; tail, 372, broken; total length, 498, 297; snout to
Tue Universitry SCIENCE BULLETIN
Fic. 48. Basiliscus bivitatus Wiegmann. RCT No, 8789, 5 mi. E. San
Isidro; about natural size.
t
i
I
Review or Lizarps oF Cosra Rica 175
forearm, 53, 48; axilla to groin, 50, 46; length of head, 35, 31; width
of head, 21, 19; from anterior base of cranial crest to distal tip, 39;
14; height of dorsal crest, 8, 3; height of caudal crest, 35, 2; arm,
52, 49; leg, 119, 106.
Variation: Sexual variation is especially great since the crests are
poorly developed in females. The ventral keels on scales of the fe-
males are"miich better developed than in the male. In one speci-
men the postrostral scale is broken and the rostral touches a total
of seven scales including labials, ete.
The coloration of fernales differs also, The ground color is olive
brown but a seriés of black quadrangular dorsal spots is present
on the sides of the back, sometimes fused mesially or alternating;
a light reddish-brown band from eye can be traced to above hind
leg, less distinct posteriorly; below this another series of quad-
Tangular dark spots. The lighter intervals between dark spots are
mottled in darker and lighter, many of the scales being clay white.
Chin and venter dirty cream with numerous black flecks; the legs
purplish or lavender below with a mottling of cream, Caudal band-
ing is somewhat evident on basal part of tail (distal part missing ).
The lips have narrow dark lines between the supralabials which ex-
tend down to the infralabials.
Remarks: This species has a wide distribution from Mexico, to
Panama. Cope described the southern form as nuchalis believing
it to be smaller than the northern form. He later abandoned this
idea and synonymized it with vittatus. All specimens I have at
hand came from the Caribbean drainage areas; and all specimens
are from localities less than a thousand feet in elevation.
’ Basiliscus basiliscus (Linnaeus )
Lacerta basiliscus Linnacus, Systema Naturae vol. 1, p. 336 (type locality un-
34334... fou 72 oe 11 16.8 | 14 23 | 38 20.5 27 | 26 | 6-6 yes | 54 | a. se
34336 fo 72 ee 10.8 16 | 42.7 23 35 21 24 27 6-6 yes 55 22 e
7 pie a ems 2 ee: Loe dese bee 15 i ia 25 42 [2k OE Be 28.5 66 | yes a te 22
34338...... 2 ot 1 Seng MaMa aaa 11 5 13 23.4 | 33.5 20 26 28 | 5-5 no* 55 | 1
34339...... fa 2 Te os Bes Se 11.6 16 13.3 24.6 38 | 20 25 28 6-6 | no* 58 | 2-2
MO 3 ees: ou 71 Cis ie = ee 15 13 | 26 35 22 | 25 28 | 6&6 yes 55 | 2-2
ie)
eS
—
302 Tue Untversiry SCIENCE BULLETIN
side bronzy olive; chin ivory, belly greenish white with metallic
reflections; tail generally olive with darker flecks on its dorsal sur-
face; a few black flecks on occiput; preanal scales dirty white; arms
and legs olive-brown, speckled with black; under part of hands and
feet, black, speckled olive above.
The ventral color of this specimen changed in preservation; pig-
ment has moved to edges of scales, thus leaving a series of dis-
tinguishable longitudinal lines of ultramarine separated by lines
of gray or bluish white, the edges of these lines indefinite.
Measurements in mm.: Snout to vent, 88; tail, 158; head width,
12.8; head length, 17; snout to ear 15.2; snout to arm insertion, 30;
axilla to groin, 49; arm, 23; leg, 28.
Variation: A table of measurements and scale data here presented
gives the chief variation that occurs in the specimens. The follow-
ing KUMNH specimens are at hand: Cartago Province: Nos, 34326-
27, 34335, Turrialba; 34342, 34347, Lake Bonilla, near Tunnel Camp.
Limén Province: No. 34328, Los Diamantes, Guapiles; 34328-34334,
Bataan; 34336-34338, Limén; 34339-34340, La Lola.
The general color pattern is maintained in all of the specimens.
Sometimes the outer dorsal black stripe is not solid and the black
color covers only the outer part of the scalerows leaving a lighter
streak down through the middle, thus one may count six dark
dorsal lines.
Remarks: The species is ovoviviparous. A specimen, (No, 34339 )
contains several nearly full-time embryos, each showing the typical
color pattern. Ahl (loc. cit.), describes a specimen from Finca Ham-
burg on the lower Rio Reventazon, near Siquirres, Limén Province.
Dunn (loc. cit.), had specimens from several localities in Costa Rica
but possibly he has confused more than one species under a single
name.
Mabuya unimarginata Cope
(Fig. 75)
Mabuya unimarginata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1862, pp. 186-
187 (type locality, “Panama” ).
Mabuya aurata Boulenger, Catalogue of the lizards in the British Museum
(Natural History), 2nd Ed., vol. III, 1887, pp. 189-190 (part.) .
Mabuia agilis Giimther, Biologia Centrali-Americana; Reptilia and Batrachia,
Oct. 1885, p. 33 (part.).
Mabuya mabouya mabouya Dunn, Proc. Acad. Nat. Sci. Philadelphia, vol. 87,
1935 (Feb. 11, 1936), pp. 537-546 (part.); Ahl, Sitzungsb. Ges. naturf.
Freunde Berlin, 1940, p. 247; Mertens, Abh. senckenb. naturf. Ges Frankfort
a. Main, No. 487, 1952, p. 58.
Mabuya mabuya mabuya Slevin, Proc, California Acad, Sci., vol, 23, no, 82,
June 25, 1942, p. 467 (Boquete, Panama).
Review or Lizarps or Costa Rica : 303
Cope’s original description of this species (loc. cit.) is as follows:
“Form slender, head as wide as the neck to the orbits, the muzzle
not elongate; scales (in the specimen described ) in thirty-two rows.
Supranasals in contact; frontal elongate, not truncate anteriorly;
4 postnasal. Supraoculars four; superior labials eight, eye over
sixth. Adpressed limbs overlapping. Seven marginal anal plates.
From muzzle to tympanum 6 lines; from muzzle to vent 2 in. 7 lin.;
from vent to end of tail 3 in. 10 lin.; anterior extremities 9 lin.
“Above rather dark brown; a narrow black band extends from
the end of the muzzle to the groin, which has a bright yellow in-
ferior border. Beneath this the sides are brown. Abdomen yellowish.
“Habitat: Panama.
“This species resembles the cepedii and aenea.”
IT am not wholly convinced that this name is definitely applicable
to the population dealt with here. The type is presumably lost.
Oreover I have no information as to the probable exact type
locality unless the habitat “Panama” refers to the city. I am, how-
ever, using this name, well aware that future Panamanian collections
may demand that another name be used for the population here
considered. The character, and an important one, in which the
type description differs, is in color pattern, no mention being made
of the black flecking that occurs on the dorsal scales of all specimens
examined,
Diagnosis: Above, the color on head and body brown with very
numerous black flecks scattered or tending towards forming a reticu-
ation; a dim dorsolateral light line sometimes evident; a lateral
stripe beginning at eye nearly solid black to groin, bordered below
by a bluish-white line beginning on tip of snout; below this a dark
Stripe on neck and sides of body that has very numerous tiny
lighter flecks; greenish white below. A large species, 88 mm.
snout to vent; scalerows usually 30 (rarely 28 or 32); frontonasal
Separated from rostral (100 percent); eye above sixth labial, (82
Percent) above fifth labial (18 percent) (only four out of 31 with
five on both sides). Scales, parietals to above vent ¢ 54-56, 9 56-58;
frontonasal does not touch rostral (100 percent); frontonasal touches
frontal (96 percent) one or two pairs of chinshields touch labials
1 percent, 29 percent) (only three out of 31 having two on
both sides); one pair of nuchals (71 percent) 2 pairs (29 percent).
Description of species (From KUMNH No. 36306. Agua Buena,
Puntarenas Province, Costa Rica): Rostral well visible above, its
height (1.6 mm.), one half its width (3.22 mm,); supranasals
Fic. 75. Mabuya unimarginata Cope. KUMNI
No. 86306, Agua Buena, Cafias Gordas, Puntarenas
Province; actual length, snout to vent, 81 mm., total
length, 214 mm.
Review or Lizarps or Costa Rica 805
broader anteriorly than posteriorly, forming a strong median suture;
frontonasal broader than long (3.7 mm. x 2.7 mm.) separated from
the rostral but very narrowly in contact with the frontal, touching
anterior frenal; prefrontals large, bordering both loreals equally,
touching first supraciliary and first and second supraoculars; frontal
longer than wide (4.2 mm. x 2.8 mm.) somewhat diamond-shaped,
the posterior point slightly rounded, touching a single supraocular
and both interparietals; latter scales touch two posterior supraocu-
lars; four supraciliaries; interparietal slightly longer than wide,
enclosed by the parietals; latter scales much wider than long, touch-
ing two postoculars, two temporals, and nuchal; nostril pierced in
Posterior part of a single nasal, with an entrant suture from upper
tim of nasal; postnasal present; two loreals, the upper fused to a
small upper preocular (abnormal?); lower preocular followed by
a single presubocular; supraciliaries separated from palpebrals by
about three rows of granules; lower eyelid with a transparent disc,
bordered above by infrapalpebrals, and from subocular by seven
tows of small granules; eight supralabials, fifth below eye, sixth
with a scale segmented from its upper part, seemingly making three
instead of two anterior temporals. Three rows of small postocular
scales, the upper of each series largest; temporals 3 4-3 -- 4; about
eleven scales surround ear-opening, at least two overlap the front
edge of the opening; seven infralabials; mental more than twice as
wide as long, with a labial border greater than that of rostral;
azygos postmental touching two labials; a single pair of chinshields
touching labials, in contact mesially; second pair separated from
labials by one scale, the third pair by two scales, the second and
third pairs scarcely differentiated; arm and leg overlapping when
adpressed; two middle fingers with 14 lamellae, the longest toe
with 18; sole and palm covered with juxtaposed granules, that at
heel enlarged; eight preanal scales, four median the larger; body
scales smooth with a faint trace of two striae; 29 scales around neck;
30 around middle of body; 108 subcaudals; 57 from parietals to
above vent; the axillary pit barely evident, much reduced in size
and depth.
Color in life: Above, brown on head and body, the head un-
Spotted, the dorsum heavily spotted or flecked with black; a broad
black lateral stripe beginning at nostril and extending to groin,
the upper edge of which has a series of small light-olive dashes
Orming an indefinite discontinuous row; a line beginning on tip
of snout is cream on labials becoming bluish white in front of ear
1]—-3878
306 THe Universiry SciENCE BULLETIN
and continuing to groin, the edges irregular; below this on chin
and sides of body a brownish black line covering about three scale
rows with small dashlike marks of brownish cream or olive-green;
these are continued along tail to tip; below light bluish- to greenish-
white, with metallic reflections; (in preservation, the color below
is light ultramarine, each scale with a somewhat lighter center );
the upper part of supralabials dark brown; a series of darker marks
on the sutures of the infralabials; under-side of sole, palm, and digits
black; subcaudal areas colored like venter; arms and legs brownish
black, with dashlike lighter marks.
Measurements in mm.: Snout to vent, 79; tail, 137; head width,
11.8; head length, 17.5; snout to ear, 14; snout to arm, 26; axilla to
groin, 43; arm, 21; leg, 27.
Variation: The table of measurements and scale data accompany-
ing this description details much of the variation. The degree of
overlap in the adpressed limbs may be ascertained by comparison
of the combined length of the limbs with the axilla to groin meas-
urement. The variation is due in part to sex, the adult females
usually having a slightly longer axilla-to-groin measurement than
males; moreover the limbs tend to grow but little in old age and
younger specimens have longer legs proportionally. I suspect that
on the whole females reach a larger size than males. There is no
reduction in the number of the supraoculars. In the supraciliaries
the last may be divided so that a count of five results occasionally.
A fusion of the first two in two cases give a count of three (on one
side only). In a few cases there are three temporals in the first
row instead of the expected two.
Remarks: The specimens of this species at hand are as follows:
San José Province: Nos. 25608-25618, 34318-34325, environs of San
Isidro del General. Puntarenas Province: Nos. 36310-86311, 15-18
km. WSW San Isidro del General; 36300-36309, Agua Buena, Cafias
Gordas. Guanacaste Province: No. 36314, 4 km. ENE Tilardn.
The finding of the two species of Mabuya in the same region
(about 7 miles apart and at about the same elevation) speaks
strongly that two species (not subspecies or lesser categories) are
involved.* While this species is to be expected on the Caribbean
Coast I have obtained no specimens there. Cope believed he had
two species from southeastern Costa Rica collected by William
Gabb.
* This other species is Mabuya brachypodus. See following description.
Measurements and scale data on Mabuya unimarginata Cope
Snout aie at | z = pee Fabil | Sabo] Seca ai — ghia gs a
“= S to . ea ea, Snout Snout | Axilla | et | parietal | Scale | nasa nasa shie = 7
Number | Sex vent Tail | width length | to ear to arm | to groin Arm Leg | below — to above| rows touch touch touch | Nuchals
| | | Becaes beteees ee | rostral | frontal | labials |
ae : | : | ‘as
Q 81 15.2 16 14 27 42 21 29 66.43 | 58 30 no yes. 1-1 a-t ring
2 63 10 14 12 30 35 16 21.2 | 6-6 | 57 30 no yes | 1-1 Zt =
Eg? 68 10.3 15 13 24.5 35 20 Zt | 6-6 SF. oO no yes tt 1-2 =
roi 52 8 11.5 10.6 19.5 28 14.2 | 20 | 66 |. 54 30 no yes =. Lt ei
Q 88 12 18 14.2 28.5 49 22.5 | 28 5-6 |- 58 30 no yes 1-2 <
fou 77 13 i 15 28 40 21 28 a-6. fF: 54 28 no yes EI 2-1 ~
2 85 11.8 16.2 14 re § 44 20 5} ZF 5-6 58 30 no yes rt Es =
fof 65 : 10 14 12 20 35 1 21:5 |}.6-6, 2: 55 30 no yes 3: 1 a
1 56 ee 8.9 14 11.8 20.2 28 15.2 | 21 t 62654525 54 30 no yes | 1-1 1-1 =
g 62 111 10 14.6 12 22 35.5 16 | 23 6-6 112 54 30 no yen LS 12 N
Q 64 p 10 14 $2 20.5 37 18 | 22 PGB is oce se: 57 32 no yes | — 2-1 px
Sv ae 95 13 |} 10.8 20 28.8 16 19 6-6 j 111 56 30° | no yes tt 5 =
of 84 12 18.2 13.5 29 45 21.3: | 29 pra fe | 56 30 no yes 2-2 2-2 is)
Sf 12 18 16 ae Pee 20 28 Ge ee eee no yes _ 2 a
re 72 11.5 16 14 28 39 18 23 6-6 | 56 30 no yes +. 1-2 fo)
roi 73 10.8 16 14 22.6 37 19 26 6-6 | 56 30 no yes 3 ee 2-2 ies]
2 56 85 + 8.5 13.2 11 19 30 14 19.5 | 5-5 | 102+ 56 30 no | yes. | st LS
ep ee to es 16-2 14 25 39.2 | 21 27.4 | 6-6 | 57 | 30 no yes | 1-0 24 ‘©.
Q 61 105 9.6 13.5 11 19 32 16 22 6-6 113 57 29 no yes | Et zs e)
ou 73 122 il 15 13 24 4] 19 26 6-6 108 55 32 no yes it i-1 =
2 57 93 9 13 11.8 20 30 16 21 6-6 | 107 60 32 no yes 1-1 i-l >
fot Si pe 9 12.2 11.2 20.6 30 15.6 | 20 6-655: 54 | 30 no yes rE 1-1t
ayo 59 8.8 13 11.3 21 30.5 16 20 5-5 | 55 30 no yes bs 1-1 zs)
Bee, ee Pee 8.2 13 il 19 28 16 19 6-6 58 32 no yes LE ae je ~
2 61 103 9 13.2 12 2E.2 33 if 22.2 | 6-6 108 57 30 no yes be, Ue. 2
ou 64 113 9.8 14.2 | 12 23 33 18.2 | 24 6-6 105 56 31 no yes 1-1 | 1-1
2 af Re 8.1 25°) 20 Ze 15 19 ale eee 56 30 no yes 2-1 1-1
ee Eee 137 11.8 17.5 14 26 43 21 27 | 5-5 | 108 57 30 no yes +t j-1
fou 60 esol 9 13.6 11.8 21 31 17 22 |6-6 | 55 30 no yes Dee! es |
Pry 63 9.8 14.4 12 22 34 17.6.5-23 6.1.66): p22 25 55 29 | no yes 2-2 2-2
36309...| a2 10.4 16 14.2 24.4 35.3 21 Se et ee oe 55 30 | no yes] 1-1 1-0
36314...| o ee Bic aes 9 12.3 11.2 21 29 15.3 | 20 5-5 54 30 no yes | 22 tf
LOS
Tue Universiry ScieNCE BULLETIN
Mabuya brachypodus sp. nov.
(Fig. 76)
Type: KUMNH No, 36258; collected 4 km, ESE of Los Angeles
de Tilaran, Guanacaste, Aug. 15, 1954 by Edward H. Taylor.
Paratypes: No. 34326, Palmar; Nos. 36312-36313, Barranca; Nos.
34310-34313, Maribella Hotel, at mouth of the Rio Barranca at
sea level,—all from Puntarenas Province. Nos. 34314-34317, Tenorio,
Las Canas, Guanacaste Province.
Diagnosis: Head and dorsum brownish, with none or only a dim
dorsolateral light line; a black stripe from eye along side to groin,
bordered below by a bluish or greenish white line beginning on tip
of snout; the light line bordered by a dotted black line below.
When limbs are adpressed, they fail to meet by a distance equal
or greater than length of hand; fifth (subocular) labial below eye;
one pair of nuchals; parietals normally in contact; usually two pairs
of chinshields touch labials; scales from parietals to above vent,
59 to 61; scale rows usually 30 (sometimes 28 or 32),
Description of the type: Rostral rather narrowly visible above,
more than twice as wide as high; a pair of supranasals, wider an-
teriorly than posteriorly, in contact on median line; frontonasal much
wider than long (2.3 mm. x 3.5 mm.), touching first loreal laterally,
but excluded from contact with rostral and frontal; prefrontals
large, in contact on median line, touching two loreals laterally, and
(normally ) first supraciliary, first supraocular, and frontal (on right
side the first and second supraoculars are abnormally fused and the
right prefrontal is likewise fused along a part of its suture with the
frontal); frontal minutely less than twice as long as wide; fronto-
parietals larger than interparietal; latter with a strongly convex
suture with the frontoparietals; parietals touching two postoculars
and two temporals laterally, much wider than long (5.2 mm. x 2.8
mm.), in contact behind interparietal; one pair of nuchals; scales
following nuchals enlarged (fusion of two body scales? ); nostril
a diagonal oval, pierced in posterior part of nasal, without an en-
trant suture from upper rim; a small postnasal; first and second
loreals subequal; one (or two) preoculars and a presubocular below
it; two rows of postoculars, the upper of each series largest; tem-
porals 2 + 2+ 3, 2+ 3 + 3; eight supralabials, the fifth under eye;
eight infralabials; mental with a larger labial border than rostral; a
large postmental bordered laterally by two infralabials; two pairs
of chinshields in contact with labials, the first pair touching each
other mesially, the second pair separated by a single scale; third
Review or Lizarps or Costa Rica 309
Fic, 76. Mabuya brachypodus sp. nov. KUMNH No, 36258,
4 km, ENE Tilardn, Guanacaste, C, R.; actual length, snout to
vent, 82 mm.
Tue Universitry ScieNCE BULLETIN
510
chinshields enlarged, separated from labials by a single scale; ear
surrounded by about 14 scales; lower eyelid with a transparent
disc, bordered above by a row of palpebrals, separated from sub-
ocular labial by about five rows of granular scales; four supracili-
aries, second elongate, separated from the palpebrals by two rows
of granular scales.
Arms short, with a well-defined axillary pit, the surface of which
is covered with small erect scales; two middle fingers each with
twelve lamellae; longest toes with 16 lamellae; vent bordered by |
eight scales, the four mesial scales somewhat enlarged; scales
around neck, 28; around middle of body, 30; sole and palm covered
with small juxtaposed granular scales, those at wrist and heel but
little larger than adjoining granules; scales from parietal to above
vent, 59.
Color in life: Above olive brown to brown, the head and body
immaculate save for a very few flecks on the rump and tail base;
a very indefinite olive-brown line borders the edge of the dorsal
coloration tending to be less conspicuous after preservation; the
black stripe from eye along side of neck and body is not of solid
color, but the black covers only part of each scale; below this a
bluish-white line from snout-tip to groin covering supralabials and
surrounding ear-opening; below this light line an olive band, the
upper edge of which has a row of small triangular black flecks.
Chin and venter nearly white, remaining so in preservation; under
side of hand and foot dirty white; some brown flecks on infralabials
below eye.
Measurements in mm.: Snout to vent, 82; tail (regenerated );
width of head, 12; length of head, 16; snout to ear, 13; snout to arm
insertion, 25.4; axilla to groin, 41; arm, 16; leg, 23.5,
Variation: Two problematical specimens captured at Barranca
show a dark lineation through each scalerow. The larger of the
two is a female with uterine young. Except for this color pattern
difference they agree with the remainder of the specimens of this
species. It is, of course, possible that these represent a form worthy
of taxonomic recognition occurring on the Peninsula of Nicoya an ,
area that in the past may have been isolated by water from the main-
land. For the present I am associating these specimens with the
form here discussed.
This species seemingly follows up the Pacific coast at least as far
as San Salvador. Mertens (1952)* reports on El Salvador speci-
* Mertens, Abh, Senckenb, naturf,, Ges., No. 487, 1952, p, 58, pl. ll, fig. 59,
Measurements and scale data on Mabuya brachypadus sp. nov.
RQ : Seales | Fronto-| Fronto-| Chin-
‘et a — Tait | Head | Head Snout.; Snout | Axilla | 4) ips Parietal = parietal | — seed — Scale x
cae width | length | toear | to arm | to groin separate | “eve Pa — a = rows 2
poh | is
36312... g 71 9 13 11.8 24.7 41 16 23 no 5 61 | yes yes 2-2 30 .
30316. =| 56 90+} 8.8 13 10 20 29 14 | 20.5 no | 5 ie nee 11) 30 re
34326... J ER ts ee 9 13.2 12 23 36 18 | 19.2 yes 6 59 no yes 2-1 30 E
34310... ro 82 12.2 17 12.8 26 46 a9 24.5 | no 5 61 yes yes 2-2} 30 fs
34311. :| of 68 10 15 12 24 38.5 | 16 20.3 no 5 60 yes yes 2-1 | 32 =
34313... j-¢ 67 9 13 11.2 22 37.5 15.8 | 20 no 5 61 no no 2-2 32 2
Ste a Ga oh. ®. 10 14 11.8 | 23 35 17 (A no 5 60 no no 2-2 | 30 a
36258... 2 >: patent ee cee an 12 16 13 25.4 41 16 23.5 no 5 59 no no 2-2 | 30 g
MG es le ie Pes Pat itech A 16 | 21 no 5 59 abe eno tpt ee | >
34315... Q (a See Pees 9.2 14 12 13 40 16.5 | 20.2 no 5 61 yes no: (f- 2-4 30 z
34314... ou 71.5 rE ? ? ? 42 16 7. ON fe ee 5 60 yes no 2-2 28 S
34317... of 50 71 7.9 12.2 10.2 18.6 25 12 16 no 5 59 yes yes 2-2 28
(ee)
—
~_
312 THe University Scrence BULLETIN
mens under the name Mabuya mabouya mabouya and gives an
excellent figure. Differences between his specimens and those here
reported are small, the dorsolateral line being a little more strongly
evident and the flecking on the limbs a little more pronounced, The
limbs when adpressed to the body fail to meet.
Remarks: It is probable that reports on specimens of this species
have appeared in the literature under other names, the one just
mentioned (supra) being such a case. My chief reasons for believ-
ing Cope’s unimarginata is inapplicable to this form is that that
species has overlapping limbs and the sixth labial is below the eye.
In this species these facts are not true.
This form resembles more the northern mabuyas in Mexico. It
may be that the Mexican form, now regarded as alliacea will prove
to be a smaller subspecies of this form.
Four subcaudal counts are available, 111, 111, 110, 109; The
nuchals are invariably 1-1; the parietals are invariably in contact
and the frontal touches the frontonasal in all except a single instance.
BIBLIOGRAPHY
ANDERSSON, Lars GABRIEL.
1916. Notes on the reptiles and batrachians in the Zoological Museum at
Gothenburg (with an account of some new species), Meddel.
Géteborgs Musei Zool. Afd., vol. 9, pp. 1-40.
AHL, ERNST.
1930. Basilisken. Aquar. Berlin, 1930, pp. 133-186, figures.
BAILEY, JOHN WENDELL.
1928. A revision of the lizards of the genus Ctenosaura. Proc. U, S. Nat
Mus., vol. 78, art. 12, no. 2733, 1928, pls. 1-30.
Batrp, SPENSER F,
1858. Description of the new genera end species of North American
lizards in the museum of Smithsonian Institution. Proc. Acad, Nat:
Sci. Philadelphia, 1858, pp. 253-256.
BERTHOLD, ARNOLD ADOLPH.
1847, Verzeichnis der aufgestellten Reptilien. Abh. Ges. Wiss. Gottingen
vol. 3, nos. 8-10, 1846 (1847), p. 5, pp. 1-28, pl. 1, fig. 1.
Barsour, Tomas
1905. The vertebrata of Gorgona Island, Colombia. Bull, Mus, Comp.
Zsol., Harvard Col., vol. 46, no. 5, 1905, pp. 98-102.
1921. Sphaerodactylus. Mem. Mus, Comp. Zool. Harvard Col., vol. 47;
no. 3, 1921, pp. 217-278, pls. 1426;
1923. Notes on reptiles and amphibians from Panama, Occ. Papers Mus:
Zool., Univ. Michigan, no, 129, Jan. 25, 1928, pp. 1-6.
1925. Two noteworthy new lizards from Panama. Proc. New England
Zool. Club, vol. 10, Jan. 4, 1925, pp. 7-10.
1928. Reptiles from the Bay Islands. Proc. New England Zool, Club.
vol. 10, July 26, 1928, pp. 55-61.
Review or Lizarps or Costa Rica 313
1934. The anoles II. The mainland species from Mexico southward.
Bull. Mus. Comp. Zool. Harvard Col., vol. 77, no. 4, June 1934,
pp. 119-155.
Barnoun, Tomas, and Nosir, G. KincsLey.
1915. A revision of the lizards of the genus Ameiva. Bull. Mus. Comp.
Zool., Harvard Col., vol. 59, no. 6, Oct. 1915, pp. 415-479.
Barnour, Tomas, and SHREVE, BENJAMIN.
1984. A new race of rock Iguana. Occ. Papers Boston Soc. Nat. Hist.,
vol. 8, Nov. 30, 1934, pp. 197-198.
Bocourr, FERMIN.
1872-1882. Itudes sur les reptiles. Mission Scientifique au Mexique et dans
VAmérique Centrale. Pt. 3 Sauriens, pp. 28-1000, [issued as fol-
lows] livr. 2, 1873; livr. 3, 1874; livr. 4, 1874; livr. 5, 1878; livr. 6,
1879; livr. 7, 1881; livr. 8, 1882, pls. 10-22J (actual total of
35 pls.).
1873. Notes erpétologique. Ann. Sci. Nat., ser. 5, vol. 17, art. 2, 1873
(not paged).
Borrrcrr, Orro.
1898. Katalog der Reptilien-Sammlung im Museum der Senckenbergi-
schen Naturforschenden Gesellschaft. Teil. 1., 18938, pp. I-X, pp.
1-140.
1898a, Geschenke und Erwerbungen fiir die Reptilien—und_ Batrachier-
sammlung. Ber. Senckenb. nat. Ges. Frankfort am Main, 1893,
pp. XXVITII-XXXI.
1893b. ibid, 1893, pp. LXXII-LXXIV.
1894. ibid, 1894, pp. XXVII-XXXIV.
1895. ibid, 1895, pp. XXXII-XXXVII.
Boutencer, Grorce A,
1883. Remarks on the Nyctisaura. Ann. Mag. Nat. Hist., ser. 5, vol. 12,
1883, p. 308.
1884. Descriptions of new reptiles and batrachians in the British Museum,
Part II. ‘Ann. Mag. Nat. Hist., ser. 5, vol. 18, 1884, pp. 396-398.
1884a. Synopsis of the existing families of the Lacertilia. Ann, Mag. Nat.
Hist., ser. 5, vol. 14, 1884, pp. 117-122.
1885-1887
Catalogue of the lizards in the British Muesum (Natural History),
2nd Ed., vol. I, 1885, pp. I-XII, 1-436, pls. 1-32; vol. I, 1885,
pp. I-XII, 1-497, pls. 1-24; vol. II, 1887, pp. I-XII, 1-573, pls.
1-40.
1890, First report on additions to the lizard collection in the British
Museum (Natural History). Proc. Zool, Soc. London, 1890, pp.
77-86, pls. 8-11.
1891. Remarks on an Iguana with a reproduced tail. Proc. Zool. Soc.
London, 1891, pp. 466-467.
1894, Second report on additions to the lizard collection in the Natural-
History Museum. Proc. Zool. Soc. London, Dec. 4, 1894, pp. 721-
736, pls. 47-49.
1897, Zool. Record for 1896, no. 33, Reptilia and Batrachia, 1897, p. 22.
1897a. A revision of the lizards of the genus Sceloporus. Proc. Zool. Soc.
London, May 18, 1897, pp. 474-522, pl. 32.
314 Tue Universiry Scrence BULLETIN
1898. An account of the Reptiles and Batrachians collected by Mr. W.
I’, H. Rosenberg in western Ecuador. Proc. Zool. Soc, London,
Feb. 15, 1898, pp. 107-126, pls. 10-18.
1898a. Third report on additions to the lizard collection in the Natural-
History Museum. Proc. Zool. Soc. London, Nov. 29, 1898, pp.
912-928, pls. 55-57.
BroncersMa, L. D.
1946. Some notes on species of the genera Bachia and Scolecosaurus.
Zool. Meded. Leiden, vol. 26, 1946, pp. 237-246, figs.
Burr, CHarues FE,
1931. A study on the teiid lizards of the genus Cnemidophorus with
special reference to their phylogenetic relationships. U. S. Nat.
Mus. Bull. 154, 1931, pp. 1-8, 1-286, figs. 1-37.
1931a. South American lizards in the collection of the American Museum
of Natural History. Bull. Amer. Mus. Nat. Hist., vol. 61, art. 7,
June 11, 1931, pp. 227-895.
1935. Notes on a collection of lizards from western Mexico and tropical
America. Trans. Micr. Soc., vol. 54, no. 2, Apr. 1935, pp. 167-178.
Burt, Cuarves E., and Burr, May DANHEIM.
1933. A preliminary check list of the lizards of South America. ‘Trans.
Acad. Sci. St. Louis, vol. 28, no. 1, issued June 1, 1933 (or Aug. 1,
1933) (both dates recorded), pp. 1-104.
Burt, Cuarites E., and Myers, Grorce S,
1942. Neotropical lizards in the collection of the Natural History Museum
of Stanford University. Stanford Univ. Publs. Uniy. Ser, Biol. Sci.,
vol. 8, no. 2, 1942, pp. 1-52.
Corr, Epwarp Drinker.
1861. Notes and descriptions of anoles. Proc. Acad. Nat. Sci. Phila-
delphia, 1861, pp. 208-215.
186la.On the genera Panolopus, Centropyx, Aristelliger and Sphaero-
dactylus. Proc. Acad, Nat. Sci. Philadelphia, 1861, pp. 494-500.
1862. Synopsis of the species of Holcosus and Ameiva, with diagnosis
of new West Indian and South American Colubridae. Proc, Acad.
Nat. Sci. Philadelphia, 1862, pp. 60-82.
1862a. Contributions to neotropical saurology. Proc, Acad, Nat. Sci.
Philadelphia, 1862, pp. 176-188.
1862b, Catalogues of the reptiles obtained during the explorations of the
Parana, Paraguay, Vermejo and Uruguay Rivers by Capt. Thos. J.
Page, U. S. N., and of those procured by Lieut. N. Michler, U. S.
Top. Eng., commander of the expedition conducting the survey of
the Atrato River. Proc. Acad, Nat. Sci. Philadelphia, 1862, pp.
346-359,
1864. Contributions to the herpetology of tropical America. Proc. Acad.
Nat. Sci. Philadelphia, 1864, pp. 166-181.
1864a, On the characters of the higher groups of Reptilia Squamata, and
especially of the Diploglossa. Proc. Acad, Nat. Sci. Philadelphia,
1864, pp. 222-231,
1866, Fourth contribution to the herpetology of tropical America. Proc.
Acad, Nat. Sci. Philadelphia, 1866, pp. 128-132.
1868.
1869.
Wis wally
1874,
1883.
1885,
1885a.
1886,
1886a.
1887,
1893,
1894,
1899,
L900,
:
Crawrorn,
Daupin, F,
1803,
Review or Lrzarps or Costa Rica 615
An examination of the Reptilia and Batrachia obtained by the Orton
Expedition to Ecuador and the Upper Amazon, with notes on other
species. Proc. Acad. Nat. Sci, Philadelphia, 1868, pp. 96-140.
Annual Report of the Trustees of the Peabody Academy of Science
for 1869 (1870). Nos. 2, 8, 1871, p. 80.
Ninth contribution to the herpetology of tropical America. Proc.
Acad. Nat. Sci. Philadelphia, 1871, pp. 200-224, pl. 17.
Description of some species of reptiles obtained by Dr. John F.
Bransford, Assistant Surgeon United States Navy, while attached to
the Nicaraguan Survey Expedition in 1873. Proc. Acad. Nat. Sci.
Philadelphia, 1874, pp. 64-72.
On the Batrachia and Reptilia of Costa Rica. Journ. Acad. Nat. Sci.
Philadelphia, ser. 2, vol. 8, 1876 (1875), pp. 93-157, pls. 23-28.
Tenth contribution to the herpetology of tropical America. Proc.
Amer. Philos. Soc., vol. 17, Aug. 15, 1877, pp. 85-97.
Eleventh contribution to the herpetology of tropical America. Proc.
Amer. Philos. Soc., vol. 18, Aug. 11, 1879 (1880), pp. 261-277.
Notes on the geographical distribution of Batrachia and Reptilia in
western North America. Proc. Acad. Nat. Sci. Philadelphia, 1883,
pp. 10-35.
Twelfth contribution to the herpetology of tropical America. Proc.
Amer. Philos. Soc., vol. 21, 1884 (Mar. 7 or July 30, 1885), pp.
167-194, pl. 1.
A contribution to the herpetology of Mexico. Proc. Amer. Philos.
Soc., vol. 22, 1885, pp. 379-397.
Jatalogue of the species of Batrachians and Reptiles contained in
a collection made at Pebas, upper Amazon, by John Hauxwell.
Proc. Amer. Philos. Soc., vol. 28, no. 121, 1886, pp. 94-108.
Thirteenth contribution to the herpetology of tropical America.
Proc. Amer. Philos. Soc., vol. 23, no. 122, April 1886, pp. 271-287.
Jatalogue of the batrachians and reptiles of Central America and
Mexico.” U. S. Nat. Bull, 82, 1887, pp, 1-100,
Second addition to the knowledge of the Batrachia and Reptilia of
Costa Rica. Proc. Amer. Philos. Soc., vol. 81, Nov. 17, 1898, pp.
333-347.
Third addition to the knowledge of the Batrachia and Reptilia of
Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. 194-206.
Contributions to the herpetology of New Granada and Argentina
with description of new forms. The Philadelphia Museum’s Sci.
Bull., no. 1, May 26, 1899, pp. 1-29, pls. 1-4.
Crocodilians, lizards and snakes of North America. Ann. Rept.
U. S. Nat. Mus. 1898 (1900), pp. 1-18; 155-1294, pls. 1-36, text
figs. 1-847,
STANTON C,
II Field keys to the lizards and amphibians of British Guiana. Ann,
Carnegie Museum, vol. 21, no. 1, Nov. 1931, pp. 11-42.
M.
Histoire naturelle générale et particuliére des reptiles. vols. 8
(an X), pp. 1-452, numerous plates; vol. 4 (an X), pp. 1-397,
numerous plates.
316 Tue Universtry ScrieNCE BULLETIN
Dumurit, A. M. C.,, and Brsron, G.
1837-1839, Erpétologie: générale on histoire naturelle compléte des reptiles.
vol. 4, 1837, pp. 1-572, pls. vol. 5, 1839, pp. 1-855, pls.
Dumirim, M. C., and Dumitrnin, Auc.
1851, Catalogue méthodique de la collection des reptiles. Mus, d’Hist.
Nat. Paris, Livr. 1, 2, 1851, pp. 1-224.
Dunn, Emmer Ret.
1930. Notes on Central American anoles. Proc. New England Zool.
Club, vol. 12, Aug. 7, 1980, pp. 15-24.
1932. Reptiles and amphibians from Honduras. Proc. Acad. Nat. Sci.
Philadelphia, vol. 84, Mar. 22, 1932, pp. 21-32.
1933.’ A new lizard from Nicaragua. Proc. Biol. Soc. Washington, vol. 46,
Mar, 24, 1933, pp. 67-68.
1933a. Amphibians and reptiles from FE] Valle de Anton, Panama. Occ.
Papers Boston ‘Soc. Nat. Hist., vol. 8, June 7, 1933, pp. 65-79.
1934. Notes on Iguana. Copeia, 1934, no. I, Apr. 24, pp. 1-4, figs. 1, 2.
1936. Notes on American mabuyas. Proc. Acad. Nat. Sci. Philadelphia,
vol. 87, 1935 (Feb. 11, 1986), pp. 533-557.
1937. The giant mainland anoles. Proc. New England Zool. Club,
vol. 16, Jan. 1, 1937, pp. 5-9.
1937a. Notes on some Colombian reptiles, Proc, Biol, Soc. Washington,
vol. 50, Feb. 28, 1937, pp. 11-14.
1939. Zoological results of the George Vanderbilt South Pacific Ex-
pedition of 1937. Part II. The lizards of Malpelo Island,
Colombia. Notulae Naturae, no. 4, May 26, 1939, pp. 1-8.
1940. New and noteworthy herpetological material from Panama. Proc.
Acad. Nat. Sci. Philadelphia, vol. 92, Nov. 18, 1940, pp. 105-122,
pli 2.
1940a. Notes on some American lizards and snakes in the Museum of
Goteborg. Herpetologica, vol. 1, no. 7, Jan. 29, 1940, pp. 189-194.
1944, The lizard genera Anadia and Ptychoglossus in Colombia. Caldasia,
vol. 3, no. 11, July 20, 1944, pp. 63-68.
Dunn, Emmet Resp, and Saxe, L. H.
1950. Results of the Catherwood-Chaplin West Indies Expedition of
1948. Part V. Amphibians and reptiles of San Andres and Provi-
dencia. Proc. Acad. Nat. Sci. Philadelphia, vol. 102, Dec. 29,
1950, pp. 141-165.
Evans, Howarp E.
1947. Notes on Panamanian reptiles and amphibians. Copeia, 1947,
no, 3, (Sept. 12), pp. 166-170.
Fink, H.
1931. Zuchterfolge bei Basiliscus vittatus. Aquarium Berlin, 1931, pp.
97-100, text figs.
Firzincer, Leopoip JosEerH
1826. Neue Classification der Reptilien . . . Wien, 1826, pp. I-VIII;
1-66, I pl.
1843. Systema Reptilium, fasc, primus, 1843, pp. 1-106, and I-VI.
Gaicr, Heten T., Harrwec, Norman, and Sruarr, L. C,
1937. Notes on a collection of amphibians and reptiles from eastern
Nicaragua. Occ. Papers Mus, Zool., Univ. Michigan, no, 357,
Oct. 26, 1937, pp. 1-18.
Review or Lizarps or Costa Rica SLi
Gravennorst, Jouann, Lupwic, Curistran.
1833. Uber Phrynosoma orbicularis, Trapelus hispidus, Phrynocephalus
helioscopus, Corythophanes cristatus und Chamaeleopsis hernandesii.
Nova Acta Acad, Leop.-Carol., vol. 16, pt. 2, Aug. 5, 1833, pp. 909-
958, pls. 63-65.
Gray, Joun Epwarp.
1825. A synopsis of the genera of reptiles and amphibia, with a descrip-
tion of some new species. Ann. Philos., (New Ser.) vol. 10, pp.
193-217.
1829-1830, Specilegia zoologica or original figures and short systematic
/ descriptions of new and unfigured animals. 1829-1830 Nos. I, II,
pls. 1-11, London.
1831, A synopsis of the species of the class Reptilia (Appendix pp. 1-100)
in Griffith, The Animal Kingdom by Baron Cuvier, vol. 9, Class
Reptilia, vol. 9, pp. 1-480, 1-100.
1839. A catalogue of the slender-tongued saurians, with descriptions of
many new genera and species. Ann. Mag. Nat. Hist., vol. 1, 1838,
pp. 274-283; 388-395, vol. 2, 1839, pp. 287-293; 331-337.
1845. Description of a new genus of night lizard from Belize (Coleonyx).
Ann. Mag. Nat. Hist., ser. 1, vol. 16, 1845, pp. 162-163.
1845a. Catalogue of the specimens of lizards in the collection of the British
| Museum, 1845, pp. I-XXVIII; 1-289.
| 1852. Description of several new genera of reptiles, principally from the
collection of H. M.S. Herald. Ann. Mag. Nat. Hist., ser. 2, vol. 10,
1852, pp. 437-440.
1863a. Descriptions of two new genera of lizards (Holaspis and Poriodo-
gaster A. Smith, M.S.). Proc. Zool. Soc. London, 1863, pp. 152-
155, pls. 20, 21.
Ginrner, ALBERT C, L. A.
1885-1902. Biologia Centrali-Americana; Reptilia and Batrachia. 1885-
1902, pp. I-XX; 1-326, pls, 1-76.
HaLvows.n, Epwanp.
1860. Report upon the Reptilia of the North Pacific Exploring Expedi-
tion under command of Capt. John Rogers, U.S.N. Proc. Acad.
Nat. Sci. Philadelphia, 1860, pp. 480-510.
Hourruyn, Martin.
1782. Het onderscheid der Salamanderen van de Haagdissen in’t alge-
meen, en van Gekkoos in’t bysonder. Verhandl. d. Zeeuwsch.,
senootsch. Vlissingen, D. 9, 1782, pp. 305-336, pl. 3, fig. 1.
KLAuBER, LAWRENCE M,
1945. The geckoes of the genus Coleonyx with descriptions of new sub-
species. Trans. San Diego Soc. Nat. Hist., vol. 10, no. 11, pp. 133-
216, 2 maps.
Kum, Hirnicr.
1820. Beitrage zur Zoologie und Vergleichende Anatomie. 1820, Frank-
furt am Main.
Tue Universitry ScueNCE BULLETIN
318
LAuRENTI, JosepH NIcOLAI.*
1768. Specimen medicum, exhibens synopsin reptilium emendatam cum
experimentis circa venena et antidota reptilium Austriacorum, Vien-
nae. 1768, pp. 1-214, 5 pl.
Lovermnce, ARTHUR. :
1929. A new Anadia from Colombia with remarks on other members of
the genus. Proc. Biol. Soc. Washington, vol. 42, 1929, pp. 99-102.
Licurenstem, Henri, and von Martens, Dr.
1856. Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolin-
ensis, 1856, pp. 1-48.
Mertens, Roser.
1952. Neues iiber die Reptilienfauna yon El Salvador. Zool. Anz., Bd.
148, 1952, pp. 87-93.
1952a. Die Amphibien und Reptilien von El Salvador, auf grund der
Reisen von R. Mertens und A. Zilch. Senckenb. Nat. Ges., no. 487,
1952, pp. 1-120, pls. 1-16,
Merrem, BLAstus.
1820. Tentamen systematis amphibiorum. 1820, Marburg.
MirrLeman, M. B.
1950. The generic status of Scincus lateralis Say, 1923. Herpetologica,
vol. 6, no. 2, 1950, pp. 17-20.
Mttier, F.
1878. Mittheilungen aus der herpetologischen Sammlung des Baslers
Museums. Verh, naturf. Ges. Basel, vol. 6, 1878, pp. 389-427,
pls. I-III.
Nose, G. KINncsLey.
1916. Description of a new eublepharid lizard from Costa Rica, Proc.
Biol. Soc. Washington, vol. 29, Apr. 4, 1916, pp. 87-88.
1921. The bony structure and phyletic relations of Sphaerodactylus and
allied lacertilian genera, with description of a new genus. Amer.
Mus. Novitates, no. 4, Mar. 10, 1921, pp. 1-16, figs. 1-8.
Nosie, G. Kincsiry, and Kiincen, G. C.
1932. The reptiles of Great Inagua Island, British West Indies. Amer.
Mus. Novitates, no. 549, Aug. 11, 1932, pp. 1-25,
O’Suaucunessy, A. W. E.
1873. Herpetological notes. Ann. Mag. Nat. Hist., ser. 4, vol. 12, pp.
44-48,
1875. List and revision of the species of the Anolidae in the British Mu-
seum collection, with descriptions of new species. Ann. Mag, Nat.
Hist., ser. 4, vol. 15, 1875, pp. 270-281.
Parker, H. W.
The neotropical lizards of the genus Lepidoblepharis, Pseudo-
gonatodes, Lathrogecko and Sphaerodactylus. Ann. Mag. Nat.
Hist., 9th ser., vol. 17, 1926 (March), pp. 291-301, text figs. 1-9.
Prracca, M. G.
1896. Descrizione di un nuovo genere e di una nuova specie di Teiidae
raccolta nel Darien dal Dott. E. Festa. Bol. Mus. Torino, vol. 11,
no. 235, 1896, pp. 1-4, figs.
wes ‘ ta Stejneger (1936 foctnote). Suggests that this work was actually written by
inter},
Review or Lizarps or Costa Rica 819
1897. Viaggio del Dr. Enrico Festa nell’Ecuador e rigioni vicini Rettili.
Bol. Mus. Zool. Univ. Torino, vol. 12, no. 300, 1897, pp. 1-20.
Perens, W.
1863. Eine mittheilung iiber einige neue Arten der Saurier-Gattung
Anolis. Monatsb. Akad. Wiss. Berlin, 1862 (1863), pp. 135-149.
1863. Uber einen neuen Gecko Brachydactylus mitratus aus Costa Rica.
Monatsb. Akad. Wiss. Berlin, 1863, pp. 41-44.
1869, Eine mittheilung tiber neue Saurier Polychrus (Chaunolaemus)
multicarinata, Tropidolepisma Richardi und Gymnodactylus Steud-
neri) und Batrachier (Cyclorhamphus fasciatus und Hyla_ gra-
cilenta). Monatsb. Akad. Wiss. Berlin, 1869, pp. 786-790.
1878. Uber neue Saurier aus Central America, Mexico und Australien.
Monatsb. Akad. Wiss. Berlin, 1873 (1874? ), pp. 738-747, 6 pls.
1874, Extract of a letter relating to Poriodogaster grayii. Proc. Zool.
Soc. London, May 19, 1874, p. 807.
1874a. Eine Mittheilung iiber neue Saurier Spaeriodactylus (sic.), Anolis,
Phrynosoma, Tropidolepisma, Lygosoma, Ophioscincus aus Cen-
tralamerica, Mexico und Australien. Monatsh. Akad. Wiss. Berlin,
1873 (between Jan. 3-Feb. 10, 1874), pp. 738-747.
Ruan, Rupoupno.
1952. Revisionary studies of some South American Teiidae. Bull. Mus.
Comp. Zool. Harvard Col., vol. 106, no. 11, June 1952, pp. 475-529,
figs. 1-10.
Ruraven, ALEXANDER.
1916. A new genus and species of lizards from Colombia with remarks on
the genus Pseudogonatodes. Occ. Papers Mus. Zool. Univ. Michi-
gan, no. 21, Feb. 10, 1916, pp. 1-8, pl. 1.
1922, The amphibians and reptiles of the Sierra Nevada de Santa Marta,
Jolombia. Misc. Publ. Mus. Zool. Univ. Michigan, no. 8, Sept.
15, 1922, pp. 1-69, pls. 1-12, charts and map.
Ruruyven, ALEXANDER, and Gaicr, HeLen T.
1924. A new Leposoma from Panama. Occ, Papers Mus. Zool. Univ.
Michigan, no. 147, Apr. 26, 1924, pp. 1-3.
Scumupr, Karu P.
1928. Reptiles collected in Salvador for the California Institute of Tech-
nology. Field Mus. Nat. Hist., Zool. Ser., vol. 12, no. 16, Nov. 28,
1928, pp. 193-201, 1 text fig.
1933. Amphibians and reptiles collected by the Smithsonian Biological
Survey of the Panama Canal zone, Smithson. Mise. Coll., vol. 89,
no. 1, Mar. 16, 1938, pp. 1-20.
1947, Notes on amphibians and reptiles of Michoacan, Mexico, Fieldi-
ana, Zool. Chicago Nat. Hist. Mus,, vol. 31, no. 9, Feb, 20, 1947,
pp. 63-85,
Sureve, BENJAMIN.
1947, On Venezuelan reptiles and amphibians collected by Dr. G, Kugler,
Bull. Mus. Comp. Zool, Harvard Col., vol. 99, no, 5, 1947, pp.
517-587.
THe Universiry SCIENCE BULLETIN
320
SuEvIN, JoserpH R.
1942. Notes on a collection of reptiles from Boquete, Panama, with the
description of a new species of Hydromorphus, Proc, California
Acad, Sci., 4th ser., vol. 23, no. 32, June 25, 1942, pp. 463-480, pls.
39-42,
Smiru, Hosarr Murr.
1935. Miscellaneous notes on Mexican lizards. Univ, Kansas Sci. Bull.,
vol, 22, Nov. 15, 1935, pp. 119-155, pls, 23-25,
1936. Descriptions of new species of lizards of the genus Sceloporus
from Mexico. Proc, Biol. Soc. Washington, vol. 49, July 3, 1936,
pp. 87-96.
1937. A synopsis of the variabilis group of the lizard genus Sceloporus
with descriptions of new subspecies, Occ, Papers Mus, Zool.
Univ. Michigan, no, 358, Oct. 30, 1937, pp. 1-14.
1939. The Mexican and Central American lizards of the genus Sceloporus.
Zool, Ser, Field Mus, Nat, Hist, vol. 26, July 27, 1939, pp, 1-397.
1939a, Mexican herpetological novelties. Proc. Biol, Soc, Washington,
vol, 52, December 15, 1939, pp, 187-196.
1941, A new race of Lygosoma from Mexico. Proc. Biol, Soc, Washington,
vol, 54, Dec. 8, 1941, pp. 181-182.
1944, Herpetological notes. Copeia, 1944, no, 3, Sept. 10, p. 187.
1944a. Additions to the list of Mexican amphibians and reptiles in the
Carnegie Museum, Ann. Carnegie Mus, vol. 30, pp. 89-92.
Smiru, Hopartr Murr, and Laurr, Leonarp FE.
1946. A summary of Mexican lizards of the genus Ameiva, Univ.
Kansas Sci, Bull,, vol. 31, pt. 1, no. 2, May 1, 1946, pp, 1-69, pls.
1-2.
1949. Miscellaneous notes on Mexican lizards, Journ, Washington Acad.
Sci., vol. 39, 1949, pp. 34-43, photos.
Smiru, Hoparr Murr, and Taytor, Epwarp.,
1950, An annotated checklist and key to the reptiles of Mexico, exclusive
of the snakes. U.S. Nat. Mus, Bull, 199, 1950, pp, 1-253.
SmitH, MALCOLM.
1935, Fauna of British India, including Ceylon and Burma, Reptilia and
Amphibia, vol. II, Sauria, London, Feb, 7, 1935, pp, 1-13; 1-440,
pl. 1, text figs, 1-93,
Spix, Jean.
1825. Animalia nova siva species novae lacertarum . . . Brasil,
1825.
STEJNEGER, LEONARD.
1900. Descriptions of two new lizards of the genus Anolis from Cocos
and Malpelo Islands. Bull. Mus, Comp, Zool, Harvard Col., vol. 36,
no. 6, Nov. 1900, pp. 159-164, 1 pl.
1903. Description of a new species of gecko from Cocos Island, Proc.
Biol, Soc, Washington, vol. 16, Feb, 21, 1908, pp, 3-4.
1907. A new gerrhonotine lizard from Costa Rica. Proc, U, $, Nat. Mus.,
vol, 32, 1907, pp. 505-506.
1936. The types of the amphibian and reptilian genera proposed by
Laurenti in 1768, Copeia, 1936, no. 3, pp. 183-141,
Review or Lizarps or Costa Rica 821
Snyper, Ricuanp C.,
1949, Bipedal locomotion of the lizard Basiliscus basiliscus. Copeia, 1949,
no. 2, June 80, pp, 129-137, pl. 1.
Stuart, Lawrence C,
1939, A description of a new Gymnophthalmus from Guatemala, with
notes on other members of the genus, Occ, Papers Mus, Zool.
Univ. Michigan, no, 409, 1939, p. 1.
1940. Notes on the “Lampropholis” group of middle American Lygosoma
(Scincidae ), with descriptions of two new forms. Occ. Papers Mus.
Zool. Univ, Michigan, no, 421, Nov, 8, 1940, pp. 1-16.
1943, Comments on the herpetofauna of the Sierra de los Cuchumatanes
of Guatemala, Occ, Papers Mus, Zool., Univ, Michigan, no, 471,
May 17, 1948, pp. 1-28, pl. 1.
1948. The amphibians and reptiles of Alta Verapaz, Guatemala, Misc.
Publ, Mus. Zool, Univ. Michigan, no, 56, pp. 1-33, pls, 1-2.
1955. A brief review of the Guatemalan lizards of the genus Anolis.
Misc, Publ. Mus. Zool, Univ, Michigan, no, 91, June 6, 1955, pp.
1-81.
Sumicurast, FRANCISCO.
1882. Enumeracién de las especies de reptiles observados en la parte
meridional de la Republica Mexicana, La Naturaleza, ser. 1, vol.
6, entrega, 2, 3, 1882, pp. 31-45.
TAyLor, Epwarp H,
1936. A taxonomic study of the cosmopolitan scincoid lizards of the genus
Eumeces with an account of the distribution and relationship of its
species, Univ. Kansas Sci. Bull., vol. 23, 1935 (Aug. 15, 1936),
pp. 1-643, pls, 1-43, text figs. 1-84,
1949. Two new Teiid lizards from Costa Rica. Univ. Kansas Sci, Bull.,
vol, 88, pt. 1, no. 5, Apr. 20, 1949, pp. 271-278, text figs. 1-2.
1955. Additions to the known herpetological fauna of Costa Rica with
comments on other species. No, II, Univ, Kansas Sci, Bull., vol.
87, pt. 1, no, 18, Oct. 15, 1955, pp. 499-575, figs. 1-9.
Trin, Jor.
1949. The genera of gerrhonotine lizards. Amer, Mid. Nat., vol. 41,
no. 38, Mar. 1949, pp. 579-601.
1949, A review of the lizard genus Barisia. Univ. Kansas Sci. Bull.,
vol. 83, pt. 1, Apr. 20, 1949, pp. 217-256,
1954, Gerrhonotine lizards recently added to the American Museum col-
lection, with further revisions of the genus Abronia. Amer. Mus.
Novitates, no. 1687, Nov. 4, 1954, pp. 1-26.
Troscuen, Ir’, Huco.
1865, [List of Reptiles] in von Miiller, Reisen in der Vereinigten Staaten,
Canada und Mexico, vol. 8, 1865, pp. 595-620.
NDERWOop, GARTH.
1954. On the classification and evolution of the geckoes. Proc. Zool. Soc.
London, vol. 124, pt. 3, Nov. 1954, pp. 469-492.
Wacuxr, JouANN.
1830. Naturliches System der Amphibia mit vorangehender Classifica-
tion der Siiugetiere und Vogel Miinchen. 1830, I-VI, 1-354,
U
822 Tue University SCIENCE BULLETIN
WALTERS, VLADIMIR.
1953. Notes on reptiles and amphibians from El Volcan de Chiriqui,
Panama. Copeia, 1953, no. 2, May 29, pp. 125-127.
WERNER, FRANZ.
1925. Zeits. wiss. Zool., Leipzig, Bd. 125, 1925, pp. 533-556, 1 map,
1 fig.
WETTSTEIN, Orro.
1934. Ergebnisse der osterreichischen biologischen Costa Rica-Expedition.
1930; die Amphibien und Reptilien. Sitzungsb. Akad. Wiss. Wien,
Math-Naturw. KI., Abt. 1, Bd. 143, Heft 1-2, 1934, pp. 1-39, text
figs. 1-9.
Wirecmann, A. F.
1828. Beitrige zur Amphibienkunde Isis, von Oken, v. 21, pp. 364-383.
1834, Herpetologia Mexicana. Berolini, 1834, pp. 1-54, pls. 1-9.
1837. Herpetologische Notizen. Arch. f. Naturg., vol. 3, pt. 1, 1837,
pp. 123-186.
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Von. XXXVIII, Pr. I] Dercemper 20, 1956 [ No. 2
Selective Sensitivity of the Formed Elements of the Blood
of the White Rat as Affected by Radium Chloride
a4 . ‘ *
Injected Intraperitoneally
BY
Pau G. Roorr
Department of Anatomy, University of Kansas,
Lawrence, Kansas
Assrracr: The total white-cell count is lowered appreciably by giving
tadium chloride (in varying dosages over a short period of time) intraperi-
toneally to young male albino rats. The polymorphs are slightly more sensitive
than lymphocytes. Blood platelets are also sensitive, reacting a little more
slowly, but when they do they show a pronounced drop.
Until more accurate data are available concerning the rate of phagocytosis
by liver, spleen, and bone marrow, as well as irradiation effect upon phagocytes,
Specific information concerning irradiation effects upon the formed elements of
the blood must wait. Also, uniform sampling must be used for differences
€xist between various neural regions. For example, the white-cell count of
blood from the heart may well be different from that of the periphery.
INTRODUCTION
This paper is concerned primarily with the sensitivity of the
formed elements of the blood to radiations (chiefly alpha-particles )
emitted by radium and its disintegration products. Ninety-six per-
cent of total energy emitted by radium in total disintegration is as-
Sociated with the alpha-particle emission. Of the remaining, 4 per-
Cent, only a fraction is absorbed in the body. Hence, the effects of
the other radiations will be negligible (Hoecker & Roofe *49, “49a).
There are numerous papers in the literature dealing with the
effects of radiation upon the peripheral blood. It is not the aim
Of this paper to present a review of this field. Jacobson & Marks
(°47) emphasize the need for more quantitative work “to determine
and evaluate incipient damage to personnel from radiation exposure
ke * Done in accordance with a contract between the Office of the Surgeon General, U. S.
“My, and the University of Kansas, Contract No, W49-007-MD466.
(323)
324 Tue Universiry SCIENCE BULLETIN
in the ‘tolerance range’.” Their paper carries an excellent bibli-
ography.
Bloom’s “Histopathology of Irradiation” (’48) does not deal with
the peripheral blood directly but extensively discusses aud surveys
the work relating to the irradiation of the hemopoietic organs, which
is most essential for the understanding of the peripheral picture.
Suter (48) has shown that the lymphocyte is more sensitive to
massive doses of radiation than the neutrophil. This is contrary
to what we have found for internal radiation from radium.
MATERIALS AND METHODS
Twenty-five young-adult white (Sprague-Dawley) rats were in-
jected intraperitoneally with radium chloride in groups of five,
the groups receiving the following dosages: 10, 20, 30, 40, 50 pg
respectively. Ten animals served as controls. All were placed on
a well-balanced diet and were kept in an air-conditioned room at
72-75°F. Extensive blood-cell counts were made prior to radium
chloride treatment to determine the average counts. One animal
from each group was sacrificed at three-day intervals up to and
including the fourteenth day. This latter period was short one day.
Two controls also were sacrificed on those days. All counts were
done on the day of sacrifice. Samples of blood for all counts were
obtained from the tails of the rats which had been cut just previous
to the sampling. Several drops of blood were allowed to escape
before the blood was drawn into the pipette or a drop used for a
smear. Counts of both red and white blood cells were made by
standard techniques using hemacytometer diluting Thoma pipettes
and Spencer's bright-line Neubauer counting chambers. All counts
were made in duplicate. Differential white-blood-cell counts were
made using Wright's stain. For the platelet counts, the blood was
diluted in red-blood-cell pipettes; the platelet solution containing
brilliant cresyl-blue, sodium citrate, and formalin, The normal
count and the experimental-group count (before treatment ) had
an average of 20,255 white cells per cubic millimeter, This was
used as a base line. Each point represents the count on a single
animal.
OBSERVATIONS
Red Blood Cells
During the period of this experiment there was no significant
change in the red-cell count. On 164 counts of normal animals, the
average red-cell count was 7,920,000 and the experimental did not
Wuitrt Rar BLoop AFFECTED BY RapruM CHLORIDE 325
deviate enough from this average (significantly) to be held statis-
tically valid.
Total White Cells
Observable effects produced by the radiations (chiefly alpha-
particles) from radium and its disintegration products present in
the blood or in the hemopoietic tissues, appear to depend upon
the amount of radium injected and the time elapsed after injection.
The extent of this dependence is indicated in Figures 1 and 2. Al-
TOTAL WHITE COUNT
“80
{80% —-
Ww
.o)
Wd
bs
z
3
)
WwW
2)
ag DOSAGE
% 10 pGMS —e
y 200" anno
9 30
40
ie
z 50
WwW
Yy
f
0% il i I \ J
6 9 12 14 1
TIME IN DAYS
TOTAL WHITE COUNT
KILLED
3RD °
20% aaa
z as
4 1 O%-4 aTH = ae
« (Chee nase
ow L n I rl J
a 10 20 40 50
DOSAGE IN MICROGRAMS 2
though the same dates have been plotted in both figures, two sets
of curves have been used in order to show more clearly the manner
in which the percentage decrease in total white cells depends upon
the two controlled variables, time and radium dosage.
In Figure 1, five sets of data each representing a different radium
dosage, have been plotted to show the effect of time on the per-
centage decrease in total white count for each given dosage. A
326 Tue Universitry SCIENCE BULLETIN
saturation effect is evident after the fifth day following injection of
radium. The effect of the radiation increases rapidly during the
first five days but reaches its maximum and no further increase is
observed after the sixth day for the four higher doses, 20, 30, 40, and
50 wg. The 10 pg curve is somewhat erratic, but rises more slowly
and does not appear to reach saturation until near the end of the
14-day period.
This saturation effect is difficult to explain but certainly must be
the result of several biological and physical factors. The purely
biological factors are probably complex. Some of the factors,
which must be in part responsible, are the rapid elimination of the
radium from the blood stream and the soft tissues, including the
bone marrow, through actual elimination from the body through
deposition in the bone tissue where the hemopoietic tissues
are no longer accessible to the alpha-particles. Another factor
that has not been dealt with quantitatively is the rate of phagocytosis
by the liver, spleen, lymph nodes, and bone marrow.
The same data are plotted in Figure 2, with percentage decrease
in total white cells as ordinate but with dosage in micrograms as
abscissa; the different sets of data now represent different times of
sacrifice. The configuration of these curves is very similar to that
of the curves discussed above, and indicates a dosage saturation
effect. The rats killed on the third day after injection show a very
marked dependence of white cell decrease on dosage. This might
reasonably have been expected since the alpha-particle tissue dosage
undoubtedly increases with the amount of radium injected. How-
ever, the remaining four curves representing the rats sacrificed on
the sixth, ninth, twelfth and fourteenth days, do not show a de-
pendence on radium dosage greater than 20 yg. This is an un-
expected dosage saturation effect which appears to indicate that
the effect of radium which has been in the body for approximately
six days or longer does not depend appreciably on the amount in-
jected above 20 pg.
These observations lead to the conclusion that within a few days
after injection the larger amounts of injected radium produce
markedly greater effects in decreasing the total white cells, and that
small doses of radium produce large decreases in white cells only
after longer periods of time. The combined effect of both time
and radium dosage does reach a maximum of about 80 percent de-
crease in white-cell count, and as this maximum is approached,
either by increasing the initial quantity of radium or by allowing
greater time to elapse, the other factor becomes much less effective.
Wuitt Rar Bioop Arrecrep By RApruUM CHLORIDE 827
In order to compare the RBE (relative biological effectiveness )
of alpha particles with that of X-rays, the data of Suter (’48) on
white cells have been plotted in a manner similar to that used in
Figures 1 and 2. Rises are much steeper and his data show evi-
dence of a recovery effect after the maximum drop in white-cell
TOTAL WHITE COUNT
”
+4 80% +
io
cs)
ty 70% +
&
o °
= 60% +
Z
50% +
Ww
3 DOSAGE 300r
ty 40% -- DATA FROM
a RESPONSE OF HEMATOPOIENC
o SYSTEM TO X-RAY
Q 30% MDDG- 02 4
ay
m GEORGE M SUTER MO.
E 20%+4
to
bd
fi 10% +
b
0% . 4 4 4 J
QO 3 6 9 12 14
TIME IN DAYS 3
Count is reached, These differences might well have been expected
because of different rates of tissue irradiation. The X-ray dose was
Probably administered within a few minutes after which no more
irradiation was received by the tissues. It is well known that the
effectiveness of X-rays is greater for a given dose as the time of
administration is shortened.
Blood Platelets
The data of Suter (’48) showing the percentage decrease in plate-
lets for different X-ray doses have been plotted versus time in
Figure 4. Our own data for different quantities of injected radium
PLATELETS PLATE LETS
90% + [DOSAGE a
200 Orece e
500 ana, -
Ndi Lo) mye
Ei eng eee a Pe ies WEAR ALS Nehaa a eae
70% 4 ; ‘ - :
ie -
rf . .
[Opa Gs ee anes ee aay at foe a
« : f Peet
5 50% a4 5,
s lf o
z
z
< 40% 4 Py
% ind
< ”
a $$$ Q
mn RESPONSE OF HEMATOPOIETIC. i
SYSTEM TO. X-RAYS* a
Sy MDDC- 824 0
~ ey “
ren,
5 10% 4. GEORGE M, 8U) 4
©
oy w =
0% 44 rt
uae
"0 awn ae |
10% Loe =20%® L 1 i 4 t 1 : 1
6 12
DAY KILLED AFTER INJECTION
328 Tue Universitry SCIENCE BULLETIN
have been plotted in Figure 5, in a smilar manner in order to show
the striking similarity in the effects of X-rays and radium on the
platelet count. The curves are almost identical in form rising
abruptly after three days to a maximum at very nearly nine days
after injection. This apparent similarity in the behavior of the
platelet count may furnish a basis for comparing the RBE of alpha-
particle irradiation with that of X-rays.
It is observed in both figures 4 and 5 that the maximum of the
curve is reached at approximately nine days. The curve for the
500 r X-ray dose reaches a maximum of 75 percent decrease in
platelet count. From figure 5, it is seen that a 75 percent decrease
would have been produced by a 40 microgram injection of radium
in the same time.
Differential Sensitivity
Cantril, Jacobson & Nickson (’46) have reported a higher sensi-
tivity of lymphocytes over polymorphs and they further state that
this seems to be a well-established fact both for acute and chronic
eno Tea.
PHS
KILLED 3rd DAY KILLED 6th, oF Ay | WILE ED ath. DAY KILLED lath DAY KILLED 14th DAY
com
x 100
Radium
of Cells after
of Cells
Number
Numer
Lisiseal
10 20 30
DOSAGE l0Opg. DOSAGE 209 DOSAGE saa DOSAGE 40jq. DOSAGE S5Oypq.
TbtAL wire SOUT
tLyMpwacyre
5¢! Pouvmonrns... of "0
(’ 7
|
|
\ |
i\ |
‘AG
|
eone eye eee ny i hina,
3.6 9 12 14 3 6 9 Ie I4 3.6 9 (2 f4
Day Killed after Injection
}
|
Wuite Rar BLoop AFFECTED BY RAprUM CHLORIDE 829
" experiments. Graphically, Figures 6 and 7 do not illustrate this in
our material, In Figure 6, time is constant and dosage variable.
The opposite is true in Figure 7. With the exception of the third
day, polymorphs show a slightly more sensitive condition than the
lymphocytes when time is the constant factor. Where dosage is
constant and time variable, the sensitivity of the polymorphs is even
more pronounced, There are two slight rises of 201 percent and
169 percent for the polymorphs on the third day with dosages of 20
wg and 40 pe respectively. The average normal differential count
for each rat was arbitrarily fixed as 100 percent. Any variation
from this is expressed in percentage of the normal.
DISCUSSION
It seems to be agreed upon by all workers that the formed ele-
ments of the blood and their precursors are very sensitive to ionizing
radiation. The degree of sensitiveness among them is different and
different workers report conflicting results as to their degree of
sensitivity. It is quite possible that the conditions of the experi-
ments are not the same, especially the physiological state of the
animals. A large factor must be considered when studying the loss
of cells in the circulating blood, This factor is the rate of phagocy-
tosis by the liver, spleen, lymph node, and bone marrow. Ac-
curate studies must be carried out to determine the degree of in-
jury of the white cells by radiation necessary to result in their phag-
ocytosis. When these studies become standardized we can express
a little more clearly the degree of sensitivity of any one of the
formed elements of the blood. Until then we can only express
quantitatively what is found in the periphery. The phagocytes of
the body no doubt are injured also in the process of irradiation.
This study in itself is a large field that should be investigated.
Knisely (’48) and his co-workers have paved the way in the study
of rates of phagocytosis under other experimental conditions.
Another factor for consideration is the source of blood for study.
Quimby, Saxon, and Goff (’48) have shown that the peripheral blood
of the rat has three times as many white cells as heart blood. Roofe
(50) et al., have shown also that this is true for the guinea pig. In
this same paper, it was pointed out that the red-cell count was also
higher at the periphery.
Tue Untiversiry SCrENCE BULLETIN
330
CONCLUSIONS
The total white-cell count is lowered appreciably by giving radium
chloride (in varying dosages over a short period of time) in-
traperitoneally in young male albino rats. The polymorphs are
slightly more sensitive than lymphocytes. Blood platelets are also
sensitive, reacting a little more slowly, but when they do they show
a pronounced drop.
LITERATURE CITED
Bioom, WILLIAM
1948. Histopathology of Irradiation from External and Internal Sources.
McGraw-Hill, New York.
Cantrii, S. T., L. JAcosson and J. T. Nickson
1946. The Effects of Irradiation on the Blood-forming Tissues. U. S.
Atomic Energy Commission, MDDC 991.
Horcker, FRANK E., and P. G. Roorr
1949. Structural Differences in Bone Matrix Associated with Metabolized
Radium, Radiology, vol. 52, pp. 856-864.
1949a. Metabolism and Distribution of Radium in Bone. Research Re-
views, vol. 1, pp. 10-24.
Jacosson, LrEon O., and E. K. Marks
1947, The Hematological Effects of Ionizing Radiations in the Tolerance
Range. Radiology, vol. 49, pp. 286-298.
KniseLy, M. H., E. H. Buocu, and L. WARNER
1948. Selective Phagocytosis I. Det Kongeliqe Videnskabernos Selskab,
Biologiske Skrifter, Bind IV, Nr. 7, pp. 1-93.
Qumpy, F. H., P. A. Saxon, and L, G. Gorr
1948. Total White Cell Counts of Peripheral and Heart Blood of the Rat.
Science, vol. 107, p. 447.
Roorr, Paut G., H. B. Latimer, M. Mapison, M. Marre, and P, WiLKINSON
1950. Comparison of Peripheral Blood with Heart Blood in the Guinea
Pig. Science, vol. 111, p. 337.
Suter, Georce M.
1948. Response of Hematopoietic System to X-Rays. U. S. Atomic
Energy Commission, MDDC 824.
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVIII, Pr. I] December 20, 1956 [No. 3
Studies on Kansas Mosses II
BY
R. L. McGrecor and E. L. Hartman
University of Kansas, Lawrence
Asstract: Eight taxa of mosses are added to the known flora of Kansas.
This increases the total number of species to 154. The presence of the alpine
and northern-inhabiting Grimmia teretinervis in Kansas is verified, and_ its
remarkable distribution is discussed.
Field and herbarium studies made during the past four years
have provided additional data concerning mosses little known in
our flora, or not recorded in previous works. The more important
taxa form the basis of this report.
Brotherella recurvans (Mx.) Fleisch. This moss, previously un-
known from Kansas, has been collected on moist sandy banks in
Leavenworth and Woodson counties. We have it from Leaven-
worth county, as Hartman 118, and from Woodson county as Mc-
Gregor, Hartman, Showalter, and Roth, March 21, 1954, and
Hartman 90. Grout (1932) gave the range of this taxon as from
Newfoundland to Manitoba, Minnesota, Wisconsin, south in the
Alleghanies to Georgia. Gier (1955) lists the taxon from Missouri
as an unverified report from Shannon county in southeast Mis-
souri, It thus appears that our collections extend the known range
of this species considerably westward.
Diphyscium foliosum (Hedw.) Mohr. We have specimens of
this taxon, as McGregor 11054, from Woodson county in southeast
Kansas. The plants were found on barren banks in an open
wooded sandy habitat dominated by Quercus stellata, Grout (1938 )
reports this taxon as common on shaded banks east of the Rocky
Mountains. However, the nearest known station in our area was
reported by Gier (1955) in east-central Missouri. Our collec-
tions extend the known range some 300 miles west in the plains
area,
(831)
Tue University ScteNCE BULLETIN
332
Grimmia olneyi Kindb. has been previously listed for Kansas as
noted by McGregor (1950). However, specimen evidence was
not known until March 21, 1954, when numerous clumps of the
moss were found on sandstone outcrops in Woodson county. These
are: Showalter 137 and Hartman 76, Grout (1933) gave the
range as Nova Scotia to Ontario, south to Georgia. Gier (1955)
listed the species from Missouri on the basis of a literature record,
but he could not find specimen evidence. Our specimens seem
to extend the known range of this taxon considerably westward.
Grimmia teretinervis Limpr. This taxon was little understood
in North America until Kucyniak (1952) clarified its characteristics
and distribution. This moss is well known in Europe where it
occurs in alpine regions of Austria, Jugoslavia, Hungary and Switzer-
land. In the latter country it seems to be rather common. In
North America the species is reported from Minnesota, Wisconsin
and Quebec by Kucyniak (loc. cit.). Kucyniak also found her-
barium specimens of the species from Kansas and Missouri. The
first of these has questionable data, and therefore the record was
provisional. The Missouri record was also reported as questionable.
The Kansas locality for Grimmia teretinervis was based upon
a specimen in the New York Botanical Garden collected along
Wolf's Creek, near Lucas, Russell county, Kansas on rocks, June 7,
1931, by C. H. Demetrio. Following the report of this collection
by Kucyniak (1952), the senior author visited the Lucas locality
and made collections of Grimmia species. One of these, McGregor
5672, was sent to Kucyniak for his study. He reported that the
plant was certainly G. teretinervis, thus establishing unquestionably
its presence in Kansas. Since then the taxon has been found in
Scott county, as McGregor 2176 and from Norton county, as R. A.
Keen, Sept. 1952.
The range of Grimmia teretinervis does not follow any general
distribution pattern noted for other plants. It appears to be an
alpine, nonarctic species which can occur at low altitudes, even
sea level, where conditions of wind and other climatic factors
somewhat duplicate alpine conditions.
In Kansas the species has been found in two different habitats.
The Russell county station consists of calcareous sandstone escarp-
ments along dry windswept hillslopes. The Scott and Norton
county stations were crevices in calcareous conglomerates called
mortar beds. These beds appear as outcrops along dry windswept
cliffs.
Srupres ON Kansas Mosses II 333
Leptodictyum laxirete (Card. and Ther.) Broth. was found in
a small creek five miles east of Baxter Springs, Cherokee county.
Our specimens, Hartman 140, extend the known range of this
taxon from Missouri to southeast Kansas.
Physcomitrium acuminatum (Schleich.) Bry. Eur. is here re-
ported for the first time from Kansas on the basis of a specimen
collected from Douglas county. This collection, Hartman 334, ex-
tends the known range of the taxon some 150 miles west from the
Nearest known station in Missouri.
Pogonatum pensilvanicum (Hedw.) Paris is added to the Kansas
moss flora on the basis of collections by Elmer Roth, April 4, 1954,
and Hartman 91, from a sandy bank near Sedan, Chautauqua
county. This extends the range nearly 150 miles west from known
Stations in Missouri.
Polytrichum piliferum Hedw. has previously been reported from
Kansas but without specimen evidence. It is now known by a
collection, McGregor 5445, from a sandy hillside in Ellsworth
county where it occurred near the base of Andropogon gerardi
clumps.
BIBLIOGRAPHY
Gren, Lid.
1955. Missouri Bryophytes. Trans. Kansas Acad. Sci., vol. 58, no. 1,
1955, pp. 24-49.
Grout, A. J,
1932. Moss Flora of North America North of Mexico. Publ. by the
Author, Newfane, Vermont, vol. 3, pt. 8, Nov. 1932, pp. 135.
1933. Moss Flora of North America North of Mexico. Publ. by the
Author; Newfane, Vermont, vol. 2, pt. 1, Nov. 1933, pp. 389.
1938. Moss Flora of North America North of Mexico. Publ. by the
Author; Newfane, Vermont, vol. 1, pt. 8, Oct. 1988, pp. 148.
Kucyntax, J,
1952. The Occurrence of Grimmia Teretinervis Limpr. in North America.
The Bryologist, vol. 55, no. 1, Mar. 1952, pp. 35-47, pl. 1.
McGrecor, Leal ie
1950. Studies on Kansas Mosses I, Univ. Kansas Sci. Bull., vol. 33, pt. 2,
no. 7, Mar. 20, 1950, pp. 291-811.
|
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou, XXXVI, Pr. I] December 20, 1956 [No. 4
Notes on the Vegetation of Southeastern
Mount Desert Island, Maine
BY
A. W. Kicuier
Axsstracr: The vegetation of southeastern Mount Desert Island, Maine,
was mapped at 1:25,000 and the physiognomy and floristic composition of
each plant community recorded, For each species the coverage and sociability
are given. These data are presented in tabular form with a brief comment
explaining the methods employed and some observations made in the field.
INTRODUCTION
The field of vegetation mapping is rapidly evolving at the present
time; in the course of this evolution, many problems become ap-
parent and call for their solutions. In an attempt to contribute
to these solutions, the writer investigated some relations between
classifications of vegetation and purposes to which vegetation maps
are applied.* In the summers of 1953 and 1954, the vegetation of
southeastern Mount Desert Island, Maine, was mapped at a large
scale, and colored maps at 1:25,000 have since been published.’ It
was exclusively with reference to the stated problems that this
vegetation was mapped. The purpose of making these maps was
not to make a phytosociological or an ecological study of the vege-
tation. However, it was quite inevitable that much detailed infor-
mation on the vegetation should have been accumulated in the
course of mapping it, even though such data were largely incidental
to the ultimate purpose of the field work. These data are interesting
in their own right, partly because they contribute to an under-
standing of the vegetation of the area, and partly because the vege-
* Sincere appreciation is expressed for the generous support given this project (N6onr
26006) by the Geography Branch of the Office of Naval Research, and for the cordial
Cooperation of the staff of Acadia National Park.
species mentioned in this paper were identified by Dr. George B. Rossbach; the
hames are uniformly based on the 8th edition of Gray’s Manual of Botany by M. L. Fernald.
Doctor Rossbach deserves a special vote of thanks for his untiring enthusiasm and efficiency
Without which it would have been difficult to bring the field work to a successful conclusion.
., 4. A. W. Kiichler: Classification and Purpose in Vegetation Maps. Geogr. Rev., vol.
XLVI, No. 2, 1956.
336 Tue Universiry SCIENCE BULLETIN
tation is in rapid though very uneven evolution. These notes are
therefore documents in the sense that they will be useful in the
future, when the vegetation has passed through new stages of
development. At any time, the study of the evolution of vegetation
san. be so much more meaningful if accurate details of some earlier
period are available. Such details for present or future use are
presented in this paper.
Much of southeastern Mount Desert Island is located in Acadia
National Park. This assures an evolution of the vegetation without
appreciable interference by man, although some indirect effects are
inevitable. For instance, the destruction by man of all large preda-
tors permits the increase of deer to a degree as to affect some
features of the vegetation. But in general, the vegetation evolves
before our eyes unhampered by lumbering or clearing. Not all of
the area lies in Acadia National Park; lumbering is indeed carried
on in restricted areas, and the comparison of the vegetational evo-
lution on protected and on unprotected sites should add further
value to the data embodied in this paper and to future studies of
the vegetation of the island.
The same argument may be applied to the difference between the
burned and the unburned sections of the region. The great Bar
Harbor fire of 1947, which destroyed the vegetation of about twenty-
three percent of the island was, of course, a real tragedy. But now
that disaster has struck, the liability can be turned into an asset by
making careful studies of the vegetation of the burned areas. It
seems a great pity that no major effort was made to compile a de-
tailed record of the regeneration of the vegetation year by year.
The opportunity was singularly favorable but has been missed.
The data here presented can remedy this situation only to a very
minor degree, but their value in this respect is nevertheless beyond
doubt.
This paper consists essentially of three parts; (1) An outline
map of the vegetation of the area; (2) the detailed notes on the
status of the vegetation as given in the “lists”; and (3) some observa-
tions on the vegetation, made during the field work.
The Map
The attached map of southeastern Mount Desert Island shows
the outline of each vegetational area as first ascertained on large
scale aerial photographs and tater checked in the field.|. Each area
Ls A detailed description of the mapping method RO: here has been given earlier:
A. W. Kiichler:; A Comprehensive Method of Mapping Vegetation, Ann. Assoc. Amer,
Geogr., vol. XLV, No. 4. Dec. 1955.
VecEeration or Mount Desert IsuAnp, MAINE 337
on the map has a number which corresponds to the same number
in the “lists”. Whereas the map accurately shows the outline of
each vegetational unit, the detailed description of these units is
given in the “lists”.
The map served as a base map for the construction of the colored
maps, referred to above. It has been printed on transparent paper
and at the same scale (1:25,000) as the colored maps. A correlation
between the material of this paper and the colored maps can be
established by placing the outline map directly on the colored maps.
Such a correlation enhances the value of both.
The Lists
The “lists” is a term which was established in the field as a matter
of convenience; it is brief and clear. The material might have
been referred to as a catalog, a record or even a table. While the
data are, in a sense, tabulated or recorded, it can be said with equal
justification that they are listed. And so the term “lists” was selected
for the large collection of field records.
In these lists, the field data have been arranged in three different
categories: (1) The numbers; (2) the physiognomic formulae; and
(8) the floristic features.
A number precedes the description of every vegetational unit.
The numbers were assigned to the areas only while the field in-
vestigation was in progress, which accounts for their irregular dis-
tribution on the map. Every number in the lists has its counterpart
on the map.
The physiognomic formulae are based on the physiognomic classi-
fication of vegetation presented elsewhere by this author.'\* This
classification is tabulated in an abbreviated form in Table 1. By
arranging the various letter symbols into formulae, the physiognomy
and structure of the vegetation can be described in great detail. An
endless variety of combinations permits this flexible system to de-
scribe any conceivable type of vegetation, The reader is referred
to the sources quoted for a detailed description of this system and
its application, but two examples will here be given to illustrate the
method.
For a first simple example, the reader is invited to turn to No. 96
in the lists; the formula is Em Dlr L. “E” stands for needleleaf ever-
1. A, W. Kichler: A Physiognomic Classification of Vegetation. Ann. Assoc. Amer.
Geogr., vol. XXXIX, No, 8, 1949. pp. 201-210.
2. A. W. Kiichler: Die Physiognomische Kartierung der Vegetation, Pet, Geogr. Mitt.,
vol. 94, No. 1, 1950. pp. 1-6.
12—3378
338 Tue University ScreNcE BULLETIN
green trees, “m” means medium tall (10-25 m); “D” implies broad-
leaf deciduous trees, “I” refers to their low stature (less than 10 m),
and “r” signifies rare but too conspicuous to be ignored. The lower
case letters refer only to the capital nearest to the left, so that the
“P’ and the “r’ refer only to the “D” but not to the “E”. Finally,
there is the “L” which stands for mosses or lichens. Therefore, the
formula Em Dlr L is to be interpreted as follows: the vegetation
consists of a forest of needleleaf evergreen trees of medium height;
low broadleaf deciduous trees are thinly scattered through this
forest and the forest floor has a cover of mosses or lichens.
Taste 1.—A physiognomic classification of vegetation
CAPITAL LETTERS
Woody Vegetation: Herbaceous Vegetation:
B: evergreen broadleaf G: graminoids
D: deciduous broadleaf H: forbs
E: evergreen needleleaf L: lichens and mosses
N: deciduous needleleaf
SMALL LETTERS
Height:
ti tAlle Wumitiitr Melont Ot WeCG so ik ee ce ein as 25 m
minimum height of herbaceous plants................... m
mis edit oleitiot WeeS. sai echo tt sds e ereie te 10- 25 m
Height of Herbaceous planta, 6.5.11 i ee ce th %- 2m
fe lows maxim Heigne of ilees es Ora eee 10 m
maximum height of herbaceous plants................... % m
So BUDD Se In OIUU c,d ee ee lm
a: dwart shrubs) maximum Welght 22). yi ae lm
Density:
continuous growth
interrupted; plants usually do not touch
plants scattered singly, or in groves or patches
rare, yet conspicuous \
barren; vegetation largely or entirely absent
aquatic vegetation
Sound =O
The second example may show a greater complexity, as illustrated
by No. 13: Dsz.lr Hmp Lp bp. “Ds” stands for broadleaf deciduous
shrubs (more than 1 m high) and “Dz” means broadleaf deciduous
dwarf shrubs (less than 1 m). As both are broadleaf deciduous,
they are contracted into “Dsz’. The “I” signifies low trees (less than
10 m high) and “r” stands for rare but conspicuous. The lower case
letters refer to the first capital at their left. As the “r” is to apply
only to the “0”, a period is placed between the “I” and the preceding
letters. “H” means forbs, “m” is medium tall (4-2 m), and “p” im-
plies patchy distribution. “L” stands for mosses or lichens and “b”
VEGETATION or Mount Desert Istanp, MAINE 389
signifies barren. The interpretation of the formula Dsz.lr Hmp Lp
bp is therefore as follows: there is a stand of broadleaf deciduous
shrubs and dwarf shrubs from which emerge occasional low broad-
leaf deciduous trees; patches of medium tall forbs and of mosses or
lichens occur among the bushes as well as an appreciable number
of barren spots.
The physiognomic formulae are followed by a description of the
floristic composition of the vegetation with the major phytosocio-
logical data, coverage and sociability (sensu Braun-Blanquet' ),
given for each species listed. In every vegetational area, all species
were listed if they were in any way prominent in their respective
plant communities. If a species was represented by only one or
very few specimens, its name was not included in the lists. The
name of every species in the lists is followed by two numbers sepa-
rated by a period. The first of these refers to the coverage and
the second number refers to the sociability. The numerical values
of the categories of each of these are given in Table 2.
TABLE 2
Coverage Sociability
under 1% of the area species scattered singly
1 - 5% of the area in small bunches or tufts
5 - 25% of the area in patches
25 - 50% of the area in anton groups
50 - 75% of the area in extensive sheets
over 75% of the area
weeny
TUR hoe
If, therefore, in the lists a species is given thus: Osmunda cinna-
momea 2.8, then this is to be interpreted as follows: Osmunda
cinnamomea covers 5-25% of the ground and grows in patches ir-
regularly distributed throughout the area.
The vegetation as given in the lists has now been described in
detail: (1) It is divided into a large number of distinct vegetation
types; (2) the area (shape and extent) of each type is shown on
the outline map; (3) the physiognomy and structure of each vege-
tation type is described in detail in the physiognomic formula;
(4) the floristic composition (the species) of each type is given;
(5) the coverage and the sociability are given for each listed
species. The map and the lists are therefore an accurate record
of the condition of the vegetation at the time of mapping.
1, J. Braun-Blanquet: Pflanzensoziologie. Springer Verlag, Wien, 1951.
340 Tue Universrry Science BULLETIN
Some Random Observations
The following observations, selected at random from the field
notes, throw additional light on the conditions of the vegetation of
Mount Desert Island. They were made quite incidentally to the
mapping activities. For ecological studies and floras, the reader
is referred to the literature.'**»4 The references given in the foot-
notes usually contain very adequate bibliographies for further re-
search. Two maps of the vegetation of Mount Desert Island have
become available in the past, one (unpublished) was prepared by
the U. S. Forest Service, and one is published in the book by
Moore and Taylor.? Both these maps are on small scales and hence
much more generalized than the maps presented here and in the
Geographical Review.”
The mapped area may well be divided into two parts: one part
burned in the great Bar Harbor fire of 1947, and another part which
escaped that calamity. However, the reader should not conclude
that the vegetation was more or less uniformly destroyed in the
burned area. Indeed, the variations in survival and revival are
striking, ranging from one extreme to the other. There are “burned”
areas where the effect of the fire was slight (191).* A few healing
years and removal of the dead trees leave a vegetation which an
unsuspecting tourist would not recognize as having burned at all.
The other extreme shows tree stumps grotesquely crowning a set
of stilt roots that rise from naked rock. Here the highly humous
soil burned with the vegetation, and what remained of it was soon
eroded and removed. The exposed granite is not conducive to a
rapid recovery and only in small depressions and cracks has enough
humus collected that flowering plants can once again participate
in the seasonal rhythm of organic activity. The red spruce (Picea
rubens ), arbor vitae (Thuja occidentalis ), and the white pine (Pinus
Strobus) were usually destroyed so thoroughly that little or nothing
remained of them. The pitch pine (Pinus rigida) proved more
resistant and is the only conifer on Mount Desert Island able to
produce root sprouts after a fire (127), but it did not do so very
often. Beech, i gus grandifolia) and oak (Quercus rubra, var.
A ee Pea ie “Hill: The Vegetation of the Penobscot Bay Region, Maine, Proc. Portland
Soc. Nat. Hist., vol. 3, , 1923.
9. F. Hyland and ae a Steinmetz: The Woody Plants of Maine. University Press,
ane ere: 1944,
Moore and N. Taylor: Vegetation of Mount Desert Island and Its Environment.
Bioghigss "Bot. Garden Mem. +» 1927
4, E. T. Wherry: Wild Flowers of Mount Desert Island, Maine. Garden Club of
Mount igo Island, 1928.
By Ay . Kiichler; Classification and Purpose of Vegetation Maps, Geogr. Rev., vol.
XLVI, No. ve 1956,
* Numbers in parentheses refer to the same numbers in the lists,
VEGETATION OF Mount Desrerr IsLAND, MAINE 841
borealis) were often able to survive because they could send strong
sprouts from undamaged roots. This frequently resulted in very
dense shrub formations. The grey birch (Betula populifolia) and
the paper birch (Betula papyrifera) were almost invariably killed
by the fire, but the wind scattered the readily available seed over
the burned areas, and much of these is now clothed by a more or
less dense cover of birches. Mixed with these are red maple (Acer
rubrum), both aspens (Populus tremuloides and P. grandidentata),
and the pin cherry (Prunus pensylvanica); the latter, in fact, is
referred to as fire cherry because it colonizes burned areas so read-
ily. Blueberries (Vaccinium angustifolium) and sweet fern (Comp-
tonia peregrina) are also common on many of the burned sections.
Due to circumstances, not investigated, the birch seedlings have
already developed into little trees in some places whereas their
growth is distinctly retarded elsewhere. Presumably, edaphic con-
ditions are responsible for such an irregular development.
Conifers, quite generally, are slow in returning and certainly
much slower than broadleaf deciduous trees. But the long root
sprouts of beech and oak, and the fast growing birches and aspens
often hide little conifers; the conifer seedlings seem scarcer than
they are because they cannot be observed readily. But the fact
remains, that they are also less numerous than other trees. The
regeneration of larches on the burned areas is usually slight.
Many of the burned areas are very steep. The destruction of
plant life and the resulting intensification of soil erosion has had
the effect of making a mottled patchwork of the returning vegeta-
tion. The steeper slopes often remain barren whereas innumerable
little ledges, or simply spots a little less steep and perhaps with
cracks in the rock are once again covered with shrubs and forbs.
Occasionally, the slopes are so steep that they are largely barren in
any case. Trees that become established here escaped the fire be-
cause the latter could not reach them.
Sometimes, distinct plant communities endeavoring to re-establish
themselves, do so without any clearly marked secondary succession,
although some species are unable to return as quickly as others.
For instance, a black spruce bog with the typically associated flora
has returned, except for the black spruce (178). Similar examples
san be given for cedar swamps (3).
The regeneration of the burned sections is now in full progress.
Broadleaf deciduous trees and shrubs are taking over the entire
area and are therefore much more prominent in the landscape than
before the fire. Spruce, fir and pine seedlings appear among their
342 Tur Untiversiry SCIENCE BULLETIN
faster growing companions but usually only in parts where they
had been before the fire. While therefore birch, aspen, red maple
and others have greatly extended their area, spruce, fir and pine
have not. Beech and oak behave like the conifers in that they, too,
do not expand into “new” territory. On the other hand, the great
expansion of birch and aspen is obviously temporary; as the conifer
seedlings mature, the light loving birches and aspens will disappear
again.
Special mention must be made of some water relations. Where
the soil permits ready drainage and seepage, and water may be
assumed to move at an appreciable speed, as along streams in
narrow valleys, hemlocks are a notable part of the vegetation (249);
but they are never dominant. If the water moves more slowly or
even seems to stagnate, a cedar (Thuja occidentalis) bog develops,
or a black spruce (Picea mariana) bog. Larches are common in
bogs, but more so in black spruce bogs than in others; while not
limited to bogs, they are rare on the more mesic sites. Certain
shrubs are commonly found in places where the water table may be
assumed to be high, especially Nemopanthus mucronata, I lex
verticillata, Viburnum cassinoides and Alnus spp. Sometimes a wet
spot accounts for the presence of these species in an otherwise mesic
plant community (151).
On higher and often very rocky slopes, as especially on the upper
western slopes of Mount Cadillac, water seeps along the surface of
the granite. The result is invariably an accumulation of arbor vitae
or white cedar (Thuja occidentalis). This species, incidentally, is
the only one which reflects any zoological components of the biome:
it is usually browsed by deer, sometimes heavily (131). The smaller
plants form very dense bushes, whereas the trees have all their lower
branches eaten off.
In the latitude and climate of Mount Desert Island, it may be
assumed that there exists a sharp contrast between the north and
south facing slopes of the mountains. This contrast is negligible.
Pitch pine (Pinus rigida) occurs usually on well drained slopes of
more or less southern exposure, but there is little else to indicate a
climatic contrast between northern and southern exposures.
A special paragraph must be devoted to the summit type of
vegetation, since it is largely responsible for the island’s name.
Champlain, sailing along the coast of Maine, was impressed with
the absence of forests on the summits and therefore called the island
“Isle des Monts Deserts”; this has been anglicized into Mount Desert
Island, The summit type is rather clearly distinguishable. Although
VEGETATION OF Mount Desert IsLanp, MAINE 343
summits of most mountains are rather bare, they do not reach above
the climatic timberline. This is well shown by the fact that the
highest parts of the highest mountain, Mount Cadillac, have well
developed forests of spruce and fir (46). The red spruce (Picea
rubens) shows little or no effect of altitude, whereas the balsam fir
(Abies balsamea) on Mount Cadillac presents the short and
dense growth that is sometimes associated with altitudinal effects.
Although the summits show much barren ground, they are far
from lifeless. Even very minor depressions contain a considerable
variety of species, nearly all of which occur also at lower altitudes.
The species that may be considered the most characteristic of higher
elevations because they are limited to these sites, are quite insig-
nificant in the general physiognomy of the vegetation. The most
common of these species is Arenaria groenlandica; Hypericum
gentianoides is rarer, and Hudsonia ericoides is found only sporadi-
cally. Pinus Banksiana occurs on Mount Cadillac, forming one very
small grove (303).
The shore lines of an island may be expected to have a vegetation
all their own. In the mapped area of Mount Desert Island, the
coasts are very rocky, often cliff-like and the forest vegetation here
does not seem to differ materially from forests farther inland. In-
deed, it is remarkable how the forest will advance right to the edge
of the salt water without appreciable change in physiognomy or
floristic composition. The white spruce (Picea glauca) is perhaps
the only one that shows a more or less distinct concentration near
the coast, but it is by no means limited to the seashore. The very
rockiness of some of the headlands results in vegetation types
similar to equally rocky sites in the mountains with ground juniper
(Juniperus communis, var. depressa) being representative. Species
that are typical of and limited to the spray region of the sea cliffs
are only few in number, notably Solidago sempervirens and Juni-
perus horizontalis (78). As a general rule, a coastal vegetation is
not clearly developed.
The exception to the tule is a small beach of shell sand and its tiny
hinterland. The beach itself is largely barren, but the grassy area
behind it (21, 22) contains, among other species, Ammophila
breviligulata, Juncus balticus, var. littoralis, and Triglochin maritima
which are not found elsewhere in the mapped area of the island in
appreciable quantity,
The vegetation of the lakes varies considerably in that some lakes
show a varied and extensive development of plant communities
whereas in others vegetation is notably absent. Of all the lakes,
344
Tur Universrry ScrmeNceE BULLETIN
Aunt Betty Pond (171) and the Tarn (42) present quite the most
spectacular sight when the water lilies (Nymphaea alba), the
pickerel weed (Pontederia cordata) or the bladderwort ( Utricularia
purpurea) are in bloom. It may be said parenthetically that these
“flowering lakes” are a real attraction in Acadia National Park, and
quite unappreciated by the annual flood of tourists.
THE LISTS
Em Dlp Lp
Picea rubens 5.1 Acer rubrum 1.1 Betula papyrifera 1.1 Acer pen-
sylvanicum 1.1 Picea glauca +-.1 moss spp. 2.3
Dsi Glp Hmlp bp
Prunus pensylvanica 3.1 Acer rubrum 2.1 Alnus crispa, var. mollis 2.1
Betula populifolia 2.1 Betula papyrifera 1.1 Populus tremuloides 1.1
Populus grandidentata 1.1 Spiraea latifolia 1.1 Diervilla Lonicera 2.1
Rubus idacus, var. strigosus 2.1 Poa compressa 1.3 Solidago canadensis
1.1 Solidago graminifolia 1.1
Gm HImp L
Carex vulpinoidea? 2.2 Carex trisperma 2.2 Scirpus cyperinus, var.
pelius 2.2 Calamagrostis canadensis 2.2 Typha latifolia 1.2 Juncus
effusus, probably var. solutus 1.2 Solidago graminifolia 2.1 Onoclea
sensibilis -+-.1 Dryopteris Thelipteris, var. pubescens +-.2 Viola pallens?
+.1 Eupatorium perfoliatum 1.1 Sphagnum sp. 5.5
Eml Dir Hlr Lp
Picea rubens 4.1 Thuja occidentalis 3.1 Abies balsamea 1.1 Acer
rubrum 1.1 Betula papyrifera 1.1 Trientalis borealis 1.1. moss spp. 2.5
Dis Ely Hm
Acer rubrum 5.1 Betula papyrifera 2.1 Fraxinus americana 1.1 Abies
balsamea 2.1 Aster umbellatus 4.1
Dls.zp Emlp Hlp Bzp
Acer rubrum 5.1 Betula papyrifera 2.1. Fraxinus americana 1.1 Abies
balsamea 2.1 Picea rubens 1.1 Vaccinium angustifolium 1.2 Gay-
lussacia baccata 1.2 Pyrus melanocarpa +.1 Aster macrophyllus 3.4
Clintonia borealis 1.3 Pteridium aquilinum, var. latiusculum +-.1 Aralia
nudicaulis +-.1 Aster acuminatus +-.1 Kalmia angustifolia 2.3
Elp Dzp Lp
Pinus rigida 4.1 Gaylussacia baccata 4.4 Amelanchier laevis a1.
Cladonia spp. 2.3
Dli Eliimp Hip
Acer rubrum 3.1 Betula papyrifera 2.1 Betula populifolia 1.1 Populus
grandidentata 1.1 Acer pensylvanicum 2.1 Picea rubens 2.1 Pinus
Strobus 2.1 Abies balsamea 3.1 Maianthemum canadense 2.3 Aralia
nudicaulis --.1 Aster macrophyllus -+-.2
El Disp Hmlp
Picea rubens 3.1 Abies balsamea 8.1 Pinus Strobus +.1 Betula
papyrifera 2.1 Betula populifolia 1.1 Acer rubrum 2.1 Populus tremu-
loides 1.3. Viburnum cassinoides 1.1 Prunus pensylvanica +-.1 Cornus
10,
19.
VEGETATION OF Mount Desrert ISLAND, MAINE 345
canadensis 2.4 Pteridium aquilinum, var, latiusculum 1.3 Aster um-
bellatus.:1.1. Osmunda Claytoniana +-.4- Osmunda cinnamomea +.4
Sphagnum spp. +.5
Eml Dlr Hlp Lp
- Picea rubens 5.1. Abies: balsamea 2.1 Thuja occidentalis 1.1 Picea
glauca +-.1 Betula papyrifera 1.1. Acer rubrum 1.1 Maianthemum
canadense 1.8 Cornus canadensis 1.3 Trientalis borealis 1.3 moss spp.
44
Dis Emlr Hm rf
Betula papyrifera 4.1 Acer rubrum 2.1 Fraxinus americana 2.1 Bet-
ula populifolia -+-.1 Acer spicatum 2.1 Sambucus pubens 1.1. Acer
pensylvanicum 2.1 Populus tremuloides +-.1 Rubus allegheniensis +-.1
Picea rubens +4-.1 Abies balsamea +-.1 Aster umbellatus 8.1 Maian-
themum, canadense 8.1 . Aralia nudicaulis -+-.1 . Solidago rugosa +.1
_ Cornus canadensis 1.2 Pteridium aquilinum, var. latiusculum -++.1 Aster
macrophyllus +-.3
No symbols
urban and agricultural land
Dszilr Hmp Lp bp
Betula populifolia 3.1. Betula papyrifera 2.1 Populus tremuloides 2.1
Populus. grandidentata. 1.1 Acer rubrum 1.1 Prunus pensylvanica 1.1
Salix discolor -+-.1 Comptonia peregrina 1.2 Rubus allegheniensis 1.
Vaccinium angustifolium 1.3 Aralia hispida +-.1 Solidago bicolor 1.1
Aster macrophyllus --.2 Polytrichum sp. 3.3
Dls. Hp
Acer rubrum 3:1 Betula populifolia 2.1 Fraxinus americana 1.1 Pop-
ulus tremuloides 1.1 Tlex verticillata +-.1 Rubus pubescens 1.1 Os-
munda Claytoniana 1.2 Solidago rugosa 1.1 Aster, umbellatus 2.1
Ds Hlp
Alnus crispa, var. mollis 4.3. Spiraea latifolia 2.4 Solidago rugosa 2.3
Dls Elr Hml
Acer. rubrum 3.1 Betula populifolia 2.1 Fraxinus americana 1.1 Prunus
pensylvanica 2.1 Crataegus sp. 1.1 Thuja occidentalis 1.1 Solidago
rugosa 3.3 Aster umbellatus 2.1
Eml
Picea glauca 4.1 Picea rubens 4.1 Abies balsamea 1.1
El Dsr Hli Glp L
Picea mariana 5.1 Ilex verticillata -+.1 Osmunda cinnamomea 4.4
Smilacina trifolia -+-.2 Carex trisperma 1,3 Carex stricta 1.1 Calama-
grostis canadensis 1.2 Sphagnum sp. 4.5
Dsp Hmlp Glp Lp
Acer rubrum 3.2 Betula populifolia 3.1 Calamagrostis canadensis +-.1
Polytrichum sp. 4.3 in addition there are in: southern section: Rubus
hispidus 8.4 Juncus tenuis +.3 Juncus canadensis +-.2 Scirpus cyper-
inus, var. pelius --.2 Aster umbellatus +-.1 northern section: Nemo-
panthus mucronata 1.3 Viburnum cassinoides +-.1 Tlex verticillata 4-.1
Osmunda cinnamomea 1.3
20.
ai.
22.
23.
24.
25.
26.
27.
28.
29.
80.
Tue Unrversiry SCIENCE BULLETIN
Ds Gmp
Alnus rugosa, var. americana 4.1 Salix discolor 1.1 Calamagrostis can-
adensis 1.8
Gl Hlp Dzr
Ammophila breviligulata 5.5 Lathyrus japonicus, var. glaber 2.2 Lath-
yrus palustris +.4 Myrica pensylvanica +.4
Gl
Calamagrostis canadensis 3.4 Juncus Gerardi 1.4 Scirpus validus, var.
creber 3.5 Juncus balticus, var. littoralis +.4 Hierochloe odorata 1.1
Triglochin maritima 1.4
wp
floating filamentous algae 3.5
Ds Hlp
Acer rubrum 2.1 Quercus rubra, var. borealis 2.1 Betula populifolia
2.1 Betula papyrifera 2.1 Populus tremuloides 2.1 Aster macrophyllus
1.3
Ds.zp Hlp Lp
Betula populifolia 3.1 Betula papyrifera 2.1 Comptonia peregrina 2.2
Acer rubrum 1.1 Populus tremuloides 1.1 Populus grandidentata 2.1
Prunus pensylvanica 1.1 Quercus rubra, var. borealis +-.1 Pteridium
aquilinum, var. latiusculum 1.1 Aralia hispida +.1 Danthonia spicata
+.2 Polytrichum sp. 2.3
Ds Hlp
Quercus rubra, var. borealis 3.1 Acer rubrum 2.1 Betula papyrifera 2.1
Betula populifolia 2.1 Populus grandidentata 2.1 Populus tremuloides
1.1 Prunus pensylvanica -+-.1 Comptonia peregrina 2.2 Acer pensyl-
vanicum +.1 Pteridium aquilinum, var. latiusculum +.1 Aster macro-
phyllus 1.2 Aralia hispida +-.1 Lysimachia quadrifolia +-.1 Danthonia
spicata +.2 Polytrichum spp. +.3
Ds.zp
Quercus rubra, var. borealis 4.1 Betula papyrifera 2.1 Betula populi-
folia 1.1. Acer rubrum +-.1 Acer pensylvanicum -+-.1 Comptonia pere-
grina 2.2
Dm.zp Hip
Quercus rubra, var. borealis 5.1 Betula papyrifera 2.1 Acer rubrum +.1
Comptonia peregrina 1.2 Aster macrophyllus 2.4
Ds.mr
Quercus rubra, var. borealis 8.1 Betula papyrifera 8.1 Fagus grandi-
folia 2.1 Populus tremuloides 1.1 Acer pensylvanicum 1.1 Acer rub-
rum 1.1 Fraxinus americana +.1 Rubus allegheniensis 2.3
Dsz HlIp Lp
Quercus rubra, var. borealis 3.1 Betula papyrifera +.1 Betula populi-
folia 3.1 Acer rubrum 2.1 Populus grandidentata 1.1 Prunus pensyl-
vanica 1.1 Populus tremuloides 1.1 Comptonia peregrina 2.2 Spiraea
latifolia +-.1 Aster macrophyllus 2.2 Aralia hispida +.1 Solidago
canadensis +.1 Solidago Randii +.1 Solidago spp. +.1 Polytrichum
sp. 1.3
Sl,
34.
36.
37.
39.
40.
41,
VEGETATION OF Mount Desert IstAnp, MAINE 347
Gm.wp
Typha latifolia 4.4 Sparganium chlorocarpum 1.3 Calamagrostis cana-
densis 2.4 Eleocharis palustris? +.2 Carex lenticularis +-.2 Nulphar
variegatum 2.3
Ds
Comptonia peregrina 3.3 Spiraea latifolia 3.3
Gl Dzp Elr
Danthonia spicata 4.5 other grass spp. 1.1 Antennaria neodioica +-.1
Vaccinium angustifolium 1.1 Picea glauca 1.1
Dm.sp Elmr Hlp
Acer rubrum 3.1 Betula papyrifera 2.1 Quercus rubra, var. borealis 2.1
Populus tremuloides +-.1 Picea rubens +-.1 Picea glauca +.1 Pinus
Strobus 4-.1 Thuja occidentalis +-.1 Abies balsamea 1.1 Aster macro-
phyllus 2.8 Aster acuminatus +.2 Aralia nudicaulis +-.2 Lysimachia
quadrifolia 1.8 Pteridium aquilinum, var. latiusculum 1.3 Dryopteris
Thelypteris, var. pubescens +-.2
Ds.mlr Emlr Glr Hr
Acer rubrum 2.1 Populus tremuloides 2.1 Betula papyrifera 1.1
Quercus rubra, var. borealis 1.1 Pinus Strobus 1.1 Comptonia pere-
grina 2.1 Salix discolor +-.1 Prunus pensylvanica +-.1 Rhus typhina
+-.1 Diervilla Lonicera +-.2 Festuca rubra -+-.2 Danthonia spicata 1.2
Phleum pratense -+.2 Agrostis alba or A. tenuis? +-.2 Poa sp. +.2
Festuca capillata -+-.2 Aster macrophyllus 1.2
Dls Elp Hir
Acer rubrum 3.2. Populus tremuloides 3.2 Betula populifolia 2.2
Abies balsamea 1.1 Aster macrophyllus 1.8
GI Dlp Hp
Phleum pratense 2.3 Festuca rubra 2.3 Dactylis glomerata 1.2 Poa
pratensis 1.2 Anthoxanthum odoratum +.2 Ranunculus acris -++.2
Pyrus malus (planted )
Elp Dlszpbp Hlp Glr Bzp Lp
Pinus rigida 2.1 Pinus Strobus 1.1 Betula papyrifera 1.1 Acer rubrum
1.1 Quercus rubra, var. borealis +-.1 Prunus pensylvanica +-.1 Vac-
cinium angustifolium 3.3 Diervilla Lonicera -+-.2 Pyrus melanocarpa
+.2 Potentilla tridentata 1.3 Pteridium aquilinum, var. latiusculum 1.1
Deschampsia flexuosa +-.2 Polytrichum sp. 1.8 lichen spp. 2.4 Dan-
thonia spicata +-.2 Kalmia angustifolia 2.8
GI Dszp Hmlp
Festuca capillata 4.5 Agrostis alba? 3.2 Alnus crispa, var, mollis 2.1
Solidago spp. 1.1 Comptonia peregrina 2.2 Vicia Cracca 1.1 Aster
umbellatus 1.1
GI Dszp.|r Hp
Festuca capillata 4.2 Agropyron repens 2.2 Spiraea latifolia 2.1
Prunus pensylvanica 2.1 Comptonia peregrina 2.1 Betula papyrifera 1.1
Betula populifolia 1.1 | Vicia Cracca 1.1
Dml.szp Gmp
Acer rubrum 3.1 Betula papyrifera 2.1 Quercus rubra, var. borealis 2.1
Alnus rugosa, var. americana +-.2 Myrica Gale 8.4 Calamagrostis cana-
densis 1.4
348
42,
43.
44,
45.
46.
AT.
48.
A9.
Tue University SCIENCE BULLETIN
w
Nuphar variegatum 3.4 Najas flexilis 3.5 Nymphaea odorata 2.3
Pontederia cordata 2.8 Utricularia purpurea 1.4 Sparganium fluctuans
+.4
Dmsi Emr Hmlp
Acer rubrum 8.1 Betula papyrifera 2.1 Fraxinus americana 1.1 Betula
populifolia 1.1 Fagus grandifolia 1.1 Betula lutea 1.1 Quercus rubra,
var. borealis 1.1 Populus tremuloides 1.1 Prunus pensylvanica +-.1
Acer pensylvanicum -+-.1 Sambucus canadensis 2.1 Sambucus pubens
2.1 Rubus allegheniensis 3.4 Rubus idaeus, var. strigosus 3.4. Acer
spicatum -++.1 Tsuga canadensis 1.1 Dryopteris noveboracensis +-.3
Osmunda Claytoniana +.2 Solidago rugosa +.1 Aralia nudicaulis 1.1
Aster macrophyllus +-.3
Gm Hmlp Dszp
Calamagrostis canadensis 5.5 Carex spp. 2.3 Impatiens capensis 1.2
Lysimachia terrestris 1.3 Salix discolor and probably S. gracilis 3.2
Populus tremuloides +-.1 Betula populifolia +-.1
Dzp Eszp Glp Bzr Hr Lp bp
Vaccinium angustifolium 2.3 Betula populifolia +-.1 Nemopanthus
mucronata -+-.1 Pyrus melanocarpa +.1 Picea rubens +.1 Juniperus
communis, var. depressa +.2 Abies balsamea -+.1 Deschampsia flexuosa
1.2 Arctostaphylos Uva-ursi, var. coactilis --.1 Kalmia angustifolia 1.1
Potentilla tridentata 4+-.3 Arenaria groenlandica 1.3
El Dsp HlIp
Picea rubens 4.1 Abies balsamea 3.1 Betula papyrifera 2.1 - Viburnum
cassinoides 1.1 Nemopanthus mucronata 1.1 Vaccinium angustifolium
+.1 Kalmia angustifolia -+-.1 Pyrus americana +.1 Cornus canaden-
sis 1.3 Aralia nudicaulis +-.1
Dml.sp Elp.mr Hp
Acer rubrum 3.1 Betula papyrifera 3.1 Fagus grandifolia 1.1 Fraxinus
americana 1.1 Acer pensylvanicum 1.1 Picea rubens 1.1 Pinus Strobus
1.1 Tsuga canadensis +.1 Abies balsamea 2.1 Maianthemum cana-
dense 1.1 Pteridium aquilinum, var. latiusculum 2.3 Aralia nudicaulis
1.1 Aster macrophyllus 1.1 Dennstaedtia punctilobula 1.4
Dls.mp Emlp Hmlp Glp
Betula papyrifera 2.1 Acer rubrum 2.1 Betula populifolia 2.1 Populus
grandidentata 1.1 Viburnum cassinoides 1.1 Prunus pensylvanica 1.1
Acer pensylvanicum 1.1 Spiraea latifolia 1.1 Diervilla Lonicera 1.2
Picea rubens 2.1 Thuja occidentalis 1.1 Pinus Strobus +-.1 Aster
macrophyllus 2.5 Aster umbellatus 1.1 Vaccinium angustifolium -+-.3
Pteridium aquilinum, var. latiusculum +-.3 Danthonia spicata 1.2 Des-
champsia flexuosa 1.2 Cladonia spp. +-.3
Dml.sp Emlp Hmlp Gmp Lp
Acer rubrum 4.1 Betula papyrifera 2.1 Fraxinus americana 1.1 Betula
lutea 1.1 Acer pensylvanicum 2.1 Picea rubens 2.1 Abies balsamea
+.1 Aralia nudicaulis 1.1 Ilex verticillata 1.1 Alnus rugosa, var.
americana 1.1 Nemopanthus mucronata 1.1 Thuja occidentalis 1.1
Osmunda Claytoniana 1.8 Cornus canadensis 1.3 Calamagrostis cana-
densis 1.3 moss spp. 3.3
52.
54,
ye
55.
57,
59.
60,
Gy
62,
VEGETATION OF Mount Desert Istanp, MAINE 349
Dis Elr bp
Acer rubrum 3.1 Betula papyrifera 3.1 Quercus rubra, var. borealis
1.1 Fraxinus americana 1.1 Picea rubens +-.1 Thuja occidentalis +-.1
Em Dlszr Hr Lp
Picea rubens 5.1 Acer rubrum +.1 Betula papyrifera +.1 Acer pen-
sylvanicum 1.1 Vaccinium angustifolium -+-.2 Aralia nudicaulis -+-.1
Maianthemum canadense +.1 Pteridium aquilinum, var. latiuseulum
-+-.2 moss spp. 3.3
b
Eml Dip
Picea rubens 4.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer pen-
sylvanicum 2.1
Dml Hmlp
Betula papyrifera 3.1 Betula lutea 3.1 Acer rubrum 2.1 Acer pen-
sylvanicum 8.1 Acer saccharum 1.1 Dryopteris marginalis +.1 Dry-
opteris spinulosa --.1 Osmunda Claytoniana +-.1 Athyrium Filix-
femina, var. Michauxii? 2.1
Disp Elp bp
Betula papyrifera 8.1 Acer rubrum 2.1 Acer pensylvanicum -+-.1 Acer
spicatum -++.1 Diervilla Lonicera +-.1 Picea rubens 2.1 Thuja oc-
cidentalis 2.1
DI Emp
Betula papyrifera 3.1 Acer rubrum 2.1 Acer saccharum 2.1 Acer pen-
sylvanicum 2.1 Betula lutea 1.1 Picea rubens 2.1 Tsuga canadensis
+.1
Dzi Hlp Glp Lp bp
Betula populifolia 3.3 Populus tremuloides 3.3 Vaccinium angusti-
folium 2.8 Aralia hispida 1.1 Anaphalis margaritacea, var. intercedens
+.2 Solidago graminifolia 4-.1 Danthonia spicata +-.2 Deschampsia
flexuosa 2.2 moss spp. 2.3
Dis Elr
Acer rubrum 3.1 Fraxinus americana 2.1 Betula papyrifera 2.1 Fagus
grandifolia +-.1 Acer pensylvanicum 2.1 Picea rubens 1.1
Dms Hp
Fagus grandifolia 3.1 Acer saccharum 3.1 Betula lutea 2.1 Fraxinus
americana +.1 Betula papyrifera 4-.1 Acer pensylvanicum 2,1 Poly-
stichum acrostichoides +-.1_ Dryopteris spinulosa +-.1
Dz.sp Glp HIp Lp -
Betula populifolia 3.1 Vaccinium angustifolium 3.3 Populus grandi-
dentata 1.1 Prunus pensylvanica 1.1 Acer pensylvanicum +.1 Des-
champsia flexuosa 2.3 Danthonia spicata 2.3. Pteridium aquilinum, var.
latiusculum 1.1 Aralia hispida +-.1_ lichen spp. 1.3 moss spp. 2.3
Emp Dmp
Tsuga canadensis 3.1 Picea rubens 1.1 Acer rubrum 3.1 Acer saccha-
rum 1.1 Betula papyrifera 1.1
Eml Dlszp Hlp L
Picea rubens 4.1 Thuja occidentalis 2.1 Pinus Strobus 1.1 Abies
balsamea 1.1 Betula papyrifera 1.1 Acer rubrum 1.1 Acer pensyl-
63,
64,
65,
66.
67.
69.
70.
Tides
Tue Universiry SCIENCE BULLETIN
vanicum 1.1 Cornus canadensis 1.1 Pteridium aquilinum, var. latius-
culum 2.1 Maianthemum canadense 1.1 Vaccinium angustifolium 2.4
moss spp. 4.3 lichen spp. 3.8
Ds Elp Hm Lp
Betula populifolia 4.4 Vaccinium angustifolium 1.3 Thuja occidentalis
2.1 Aster umbellatus 5.4 Polytrichum sp. 3.3. Sphagnum sp. 1.2 lichen
spp. 1.2
Ds.li Hmlp
Acer rubrum 3.1 Betula papyrifera 2.1 Fagus grandifolia 2.1 Betula
populifolia 2.1 Acer pensylvanicum 1.1 Populus grandidentata 1.1
Rubus idaeus, var. strigosus 2.3 Solidago canadensis +.1 Solidago
rugosa +.1 Aralia hispida +-.3
El Dip Gmlp Hmlp Lp
Thuja occidentalis 5.1 Betula populifolia. 1.1 Calamagrostis canadensis
1.3 Carex crinita +.2 Aster umbellatus 1.1 moss spp. 1.3
Eml Dlzp Hlp Lp
Picea rubens 4.1 Thuja occidentalis 1.1 Betula papyrifera 2.1 Acer
rubrum 1.1 Vaccinium angustifolium 1.3 Cornus canadensis 1.3 Pterid-
ium aquilinum, var. latiusculum 1.3 Maianthemum canadense 1.3 moss
spp. 2.2
Dml.sp Emr
Acer rubrum 8.1 Betula papyrifera 2.1 Fagus grandifolia 2.1 Betula
lutea 1.1 Acer saccharum 1.1 Acer pensylvanicum 3.1 Tsuga canaden-
sis 1.1
Dml Emlp Hli Lp
Betula papyrifera 4.1 Acer rubrum 2.1 Populus grandidentata -++.1
Acer pensylvanicum 2.1 Picea rubens 2.1 Pinus Strobus 1.1 Thuja
occidentalis 1.1 Abies balsamea 2.1 Pteridium aquilinum, var, latius-
culum 2.1 Aralia nudicaulis 2.1 Cornus canadensis 1.1 Trientalis
borealis +-.1 Aster macrophyllus 2.1 Maianthemum canadense 1.1 moss
spp. 2.3
Emli Dmlsi Hmlp
Picea rubens 2.1 Abies balsamea 2.1 Pinus Strobus 2.1 Betula papyri-
fera 4.1 Acer rubrum 1.1 Acer pensylvanicum 1.1 Aralia nudicaulis 1.1
Aster macrophyllus 1.1 Pteridium aquilinum, var. latiusculum 1.1
Dmp Emlp
Betula papyrifera 3.1 Acer rubrum 2.1 Acer pensylvanicum 1.1 Pinus
Strobus 3.1 Thuja occidentalis 1.1
Diszp Elp Glp Lp bp
Betula populifolia 1.1 Betula papyrifera -+.1 Acer rubrum +.1 Gay-
lussacia baccata 1.3 Vaccinium angustifolium 1.2 Pyrus melanocarpa 1.3
Picea rubens +.1 Thuja occidentalis -+.1 Deschampsia flexuosa 1.2
lichen spp. 2.3
Dm Elr Glp
Fagus grandifolia 4.1 Betula lutea 2.1 Betula papyrifera 2.1 Acer sac-
charum 1.1 Acer pensylvanicum 1.1 Picea rubens 1.1 Carex conoidea?
1.2
73.
74,
75.
76.
Ge
78.
79.
80.
81.
82.
VEGETATION OF Mount Desert IsLAND, MAINE 351
Em Dir
Thuja occidentalis 5.1 Picea mariana 1.1 Pinus Strobus 1.1 Acer ru-
brum 1.1
Dzp Elsr Lp Bzr Glr bp
Gaylussacia baccata 1.1 Pyrus melanocarpa +.1 Vaccinium angustifo-
lium -+-.2 Rhododendron canadense 1.1 Nemopanthus mucronata +-.2
Kalmia angustifolia 1.1 Thuja occidentalis +.1 Picea rubens +.1 lichen
spp. 2.3. Deschampsia flexuosa +.2 Danthonia spicata +.2 Arenaria
groenlandica +-.1 Potentilla tridentata +-.1
El Dlszp Lr bp
Picea rubens 5.1 Thuja occidentalis 1.1 Betula papyrifera 1.1 Acer
rubrum 1.1 Vaccinium angustifolium 1.3 Gaylussacia baccata +-.3
' Kalmia angustifolia +.3 lichen spp. 1.2
Emli Dip Hip
Abies balsamea 2.1 Picea rubens 2.1 Pinus Strobus 1.1 Betula papyri-
fera 1.1 Betula populifolia +-.1 Acer rubrum 1.1 Cornus canadensis
+.4 Maianthemum canadense 1.2 Pteridium aquilinum, var. latiuscu-
lum +.2 Aralia nudicaulis +.1 Clintonia borealis +-.1
Dsz.lp Elzr Hmp Glp Lp bp
Viburnum cassinoides 2.1 Prunus pensylvanica 1.1 Betula papyrifera 1.1
Betula populifolia 1.1 Prunus virginiana 1.1 Populus tremuloides +.1
Myrica pensylvanica 2.4 Diervilla Lonicera 1.1 Rubus allegheniensis 2.1
Picea glauca 1.1 Juniperus communis, var. depressa 1.8 Danthonia spi-
cata 2.2 Solidago canadensis 1.1 lichen spp. 2.38 Pteridium aquilinum,
var, latiusculum 1.1 Aster macrophyllus 1.3 Aster umbellatus 1.1
Glp Dzp Ezp.Ir Hlr bp
Festuca rubra 3.3 Festuca capillata 4+-.3 Myrica pensylvanica 1.8 Spi-
raea latifolia 1.1 Juniperus communis, var. depressa 2.3 Juniperus hori-
zontalis 4-.3 Picea glauca 1.1 Aster novi-belgii, probably var. litoreus
+.1 Plantago juncoides, var. decipiens -+-.2 Solidago nemoralis 1.1
Solidago sempervirens -+-.1
Emp Dlp Hlp
Picea rubens 4.1 Picea glauca +-.1 Abies balsamea 1.1 Acer rubrum
8.1 Acer pensylvanicum +-.1 Betula papyrifera 2.1 Aster macrophyllus
1.3 Aralia nudicaulis +-.1 Maianthemum canadense 1.2
Eml Dlsp Lp
Picea rubens 4.1 Abies balsamea +-.1 Thuja occidentalis 1.1 Pinus
Strobus +.1 moss spp. 3.3 Acer rubrum 2.1 Betula papyrifera 1.1
Acer pensylvanicum +-.1
Eml Dir
Picea rubens 5.1 Pinus Strobus 1.1 Abies balsamea +.1 Thuja occi-
dentalis -+-.1 Acer rubrum -+-.1
Dmlp Elp Hmp Glr
Acer saccharum 2.1 Acer rubrum 2.1 Fraxinus americana 2.1 Thuja
occidentalis 3.1 Aster umbellatus 3.1 Osmunda Claytoniana 2.3 Os-
munda regalis, var. spectabilis 2.3 Osmunda cinnamomea 2.8 Carex
gracillima 2.2
83.
84,
85,
86.
87.
88.
89.
90.
QL.
Tue Universrry Science BULLETIN
Di
’ Acer rubrum 4.1 Betula papyrifera 3.1 Acer saccharum -++.1 . Fagus
grandifolia +.1
Elp Dlszp Lp bp
- Picea rubens 4.1 Thuja occidentalis 2.1 Pinus resinosa 1.1 Betula
papyrifera 1.1 Acer rubrum 1.1 Vaccinium angustifolium 1.3. Gay-
lussacia baccata 1.38 Kalmia angustifolia +.3 Pteridium aquilinum, var.
latiusculum -+-.3. lichen spp. 2.3
Emli Dlp Hlp Lp
Picea rubens 4.1 Pinus Strobus +.1 Tsuga canadensis +.1. Betula
papyrifera 2.1 Acer rubrum 2.1 Populus grandidentata +.1 Maian-
themum canadense 1.1 Cornus canadensis -+-.1 Aralia nudicaulis +.1
moss spp. 3.2 .
Elp Dlszp Bzp Glp Lp bp
Picea rubens 2.1 Betula populifolia 2.1 Acer rubrum +.1 Gaylus-
sacia baccata 2.3. Vaccinium angustifolium 1.3 Pyrus melanocarpa 1.3
Kalmia angustifolia 1.2 Danthonia spicata 1.2 Deschampsia flexuosa
1.2 Cladonia spp. 1.8
Dzp Bzp Hlp Glp Lp bp
Pyrus melanocarpa 4.3 Gaylussacia baccata 1.3 Vaccinium angusti-
folia 2.2 Kalmia angustifolia 1.38 Potentilla tridentata +.1 Arenaria
groenlandica +-.2 Deschampsia flexuosa 1.2 Cladonia spp. 2.3
Dm Emr
Fagus grandifolia 4.1 Betula papyrifera 1.1 Acer saccharum 1.1’ Acer
rubrum +.1 Betula lutea +.1 Fraxinus americana +.1] Tsuga cana-
densis +-.1
Dm.lsp Emlr
Betula papyrifera 3.1 Acer saccharum 1.1 Acer rubrum 3.1 Fagus
grandifolia 2.1 Betula lutea +.1 Fraxinus americana +.1 Quercus
rubra, var. borealis --.1 Acer pensylvanicum 1.1 Acer spicatum +.
Tsuga canadensis +-.1 Abies balsamea -+.1 Picea rubens +.1 Pinus
Strobus +-.1
Emp Diszp Hmp Glp
Picea rubens 2.1 Tsuga canadensis +-.1 Pinus Strobus 1.1 Acer rubrum
3.1 Betula papyrifera 2.1 Betula populifolia 2.1 Acer pensylvanicum
1.1 Amelanchier laevis -+-.1 Vaccinium angustifolium 3.3 Aster um-
bellatus| 1.1. Pteridium aquilinum, var. latiusculum 2.2 Danthonia
spicata 2.3 °Deschampsia flexuosa 2.3
Emli Dlszi Glp Lr
Picea rubens 2.1 Pinus Strobus 3.1 Thuja occidentalis 1.1 Betula
papyrifera 3.2 Betula populifolia 2.1° Acer pensylvanicum 2.1 Acer
rubrum 2.1 Vaccinium angustifolium 4.3 Gaylussacia baccata 2.8. Dan-
thonia spicata 1.2 Deschampsia flexuosa 3.3 lichen spp. 1.3
Dli.zr Elr
Acer saccharum 3.1 Betula papyrifera 3.1 Fagus grandifolia 2.1 Betula
lutea’ 2:1 Acer pensylvanicum 1.1 Picea rubens 1.1 Rubus idaeus, var.
strigosus 1.4
93.
94,
96,
97,
98,
99.
100,
LOL,
102.
VEGETATION OF Mount Desert ISLAND, MAINE 853
Dzp Bzp Elp Hlp Glr bp
Pyrus melanocarpa 2.1 ‘Vaccinium angustifolia +.2 Vaccinium Vitis-
Idaea, var. minus +.2 . Kalmia angustifolia 2.2 Picea rubens 1.1
Deschampsia flexuosa +-.2 Potentilla tridentata 1.2 Arenaria groen-
landica +-.2 Rhododendron canadense +.2
Dlisp Elr Hmp Gmp
Acer rubrum 4,4. Betula populifolia 1.1 Viburnum cassinoides 1.1
Nemopanthus mucronata 1.1 Alnus crispa, var. mollis 1.1 Pyrus. amer-
icana +.1. Picea rubens 1.1. Aster umbellatus 3.1 Brachyelytrum
erectum, var. septentrionale syd Calamagrostis canadensis 3.4 Carex
gracillima 2.2
Dml.sp Emlp Hlp Glp
Acer rubrum 4.1 Betula. populifolia 8.1 Acer pensylvanicum 2.1 Vi-
burnum cassinoides 1.1 Picea rubens 3.1 Abies balsamea +-.1 Picea
glauca --.1 Pteridium aquilinum, var. latiusculum -++.1 . Aster umbel-
latus 1,1 Calamagrostis canadensis 2.4 Carex gracillima 2.2 Brachyely-
trum erectum, var, septentrionale 2.4
Em, DireL
» Rieea ‘rubens 5,1. Pinus Strobus. +,1. Thuja occidentalis +-.1 Betula
papyrifera +-.1) Acer rubrum +.1/ moss spp, 4.3
DI El Hlp Lr
Betula papyrifera 3.1 Acer rubrum. 3.1',.Acer pensylvanicum 1.1 Picea
rubens 2.1. Picea: glauca 1.1 Abies balsamea 3.1. Maianthemum cana-
dense 2.4 moss spp. 1,2 lichen spp. 1.8
Eml Dlp.sr Hlp
Picea rubens 5.1 Pinus Strobus 2.1 Abies balsamea 1.1 Betula papy-
rifera 1.1 Acer rubrum 1.1 Acer pensylvanicum 1.1. Maianthemum
canadense 1.2 Aralia nudicaulis +-.1 Trientalis borealis -++.1
Dml Elr
Fagus grandifolia 4.1. Betula papyrifera 2.1 Acer rubrum 1.1 Betula
lutea 1.1 Acer pensylvanicum 1,1 Picea rubens 1.1
Dml Elp Hmlp
Acer rubrum 4.1 Betula papyrifera 2.1 Betula lutea 1.1 Fraxinus
americana 1.1 Picea rubens 2.1 Abies balsamea 1.1 Dryopteris nove-
boracensis +-.3. Aster macrophyllus 1.1 © Osmunda Claytoniana 1.1
Dryopteris Phegopteris --.1
Gl Dzp
Carex lenticularis 2.2 Carex rostrata 2.2 Carex stricta 2.2 Dulichium
arundinaceum 2.2 Typha latifolia 2.2 Myrica Gale 1.2 Alnus rugosa,
var. americana 1.2 Nemopanthus mucronata 1.2. Ilex verticillata 1.2
Dz.lsp Glp Hmp Emzr
Spiraea latifolia 2.2 Diervilla Lonicera 2.2 Vaccinium angustifolium
9.3 Acer rubrum 1.1 Gaylussacia baccata 1.8 Betula papyrifera 2.1
Viburnum cassinoides --.1 Prunus pensylvanica +.1 Betula populifolia
--.1 Festuca rubra 1.2 Danthonia spicata 1.2 Aster umbellatus 1.1
Pteridium aquilinum, var. latiusculum 1.3 | Solidago graminifolia +-.1
Picea rubens 2.1 Juniperus communis, var. depressa +-.2 Picea glauca
lia
112.
118.
1038.
104.
105.
106.
107.
108.
109.
110.
LLG.
Tue Universiry SCIENCE BULLETIN
Em Dlp Nir Hmp
Picea rubens 5.1 Abies balsamea 1.1 Acer rubrum 2.1 Acer pensyl-
vanicum 1.1 Betula papyrifera 1.1 Betula lutea 1.1 Larix laricina +-.1
Kalmia angustifolia +.3 Osmunda Claytonia +-.2 Osmunda cinna-
momea 1,2
Dz.lr Elp Bzp Glp Lp Hmlp bp
Picea rubens 3.1 Pinus Strobus 1.1 Picea glauca 1.1 Vaccinium an-
gustifolium 4.3 Gaylussacia baccata 3.3 Betula papyrifera 2.1 Acer
rubrum 2.1 Kalmia angustifolia 1.2 Danthonia spicata 2.3 Deschampsia
flexuosa 1.3 Pteridium aquilinum, var. latiusculum 3.3 Cladonia spp. 2.3
El HI Glp L
Thuja occidentalis 5.1 Picea rubens +-.1 Osmunda cinnamomea 4.4
Carex trisperma 2.2 Equisetum sylvaticum, forma multiramosum 1.3
Sphagnum spp. 4.4
Em.lp Lp Hlr
Picea rubens 5.1 Maianthemum canadense 1.1 moss spp. 3.3
Eml Hlp Lp Dzp.mlr
Picea rubens 5.1 Abies balsamea 1.1 Cornus canadensis 2.1 Maianthe-
mum canadense 2.1 moss spp. 3.3. Betula papyrifera 1.1 Acer rubrum
1.1 Gaylussacia baccata -+.2 Vaccinium angustifolium +-.2
Gl Ezlr Dzr Hmr
Festuca rubra 4.4 Luzula multiflora 2.3 Festuca capillata 2.8 An-
thoxanthum odoratum 1.8 Juniperus communis, var. depressa +-.1_ Picea
glauca +-.1 Spiraea latifolia 2.8 Alnus crispa, var. mollis +.2 Hyperi-
cum perforatum +.2
Em
Picea rubens 5.1 Pinus Strobus 2.1
Emli Dlzp HJp Glp Lp bp
Picea rubens 3.1 Thuja occidentalis 2.1 Pinus rigida +.1 Betula
papyrifera 1.1 Acer rubrum 2.1 Gaylussacia baccata 2.4 Vaccinium
angustifolium 2.4 Aster umbellatus +.3 Pteridium aquilinum, var.
latiusculum 1.3 Danthonia spicata 1.2 Deschampsia flexuosa 1.2 lichen
spp. 1.2
Dm.sp Emr
Acer rubrum 2.1 Betula papyrifera 2.1 Betula lutea 2.1 Fagus grandi-
folia 2.1 Acer saccharum 1.1 Acer pensylvanicum 3.1 Picea rubens 1.1
Pinus Strobus ++.1
Elp.zr Dzp Bzp Glp Lp Hir bp
Picea rubens 2.1 Pinus rigida 1.1 Juniperus communis, var. depressa
+.2 Gaylussacia baccata 2.2 Betula populifolia -+-.1 Vaccinium angus-
tifolium 1.2 Pyrus melanocarpa 1.2 Rhododendron canadense 1.1 Kal-
mia angustifolia 2.2 Deschampsia flexuosa 1.2 Danthonia spicata 1.2
Cladonia spp. 2.3 Potentilla tridentata +.2 Hypericum gentianoides
+.2 Hudsonia ericoides 1.1
Eli.zr Dzp Glr Bzr bp
Picea rubens 3.1 Pinus rigida 1.1 Juniperus communis, var. depressa
+.2 Pinus resinosa 1.1 Gaylussacia baccata 2.2 Vaccinium angusti-
folium 2.2 Betula populifolia +-.1 Deschampsia flexuosa +-.1 Dan-
thonia spicata +.2 Kalmia angustifolia 1.2
114.
115.
L16.
LT.
118.
119,
120.
VEGETATION OF Mount Desert Istanp, MAINE 355
DI Elp
Acer rubrum 3.1 Betula papyrifera 3.1 Picea rubens 2.1
Not on map.
Dml Elp Hlp
Betula papyrifera 4.1 Acer rubrum 8.1 Acer saccharum 1.1 Acer pen-
sylvanicum 1.1 Betula lutea 1.1 Populus grandidentata 1.1 Fraxinus
americana 1.1 Thuja occidentalis 2.1 Picea rubens 2.1 Dryopteris
noveboracensis --.2 Aralia nudicaulis +.1
Dls Elp.mr Nir
Acer rubrum 3.1. Betula papyrifera 2.1 Betula populifolia 2.1) Frax-
inus americana 2.1 Alnus rugosa, var. americana 2.1 Amelanchier laevis
2.1 Tlex verticillata 2.1 Pinus Strobus 2.1 Abies balsamea +.1 Larix
laricina 1.1
Eml Hip
Picea glauca 5.1 Thuja occidentalis +-.1 Abies balsamea +.1 Maian-
themum canadense 2.4 Aralia nudicaulis 4-.1
Dml Emi
Acer saccharum 2.1 Fagus grandifolia 2.1 Betula lutea 2.1 Betula
papyrifera 1.1 Acer rubrum 1.1 Fraxinus americana 1.1 Picea rubens
3.1 Abies balsamea 2.1 Picea glauca 1.1 Pinus Strobus 1.1
Eml Dlzr Hp
Pinus Strobus 4.1 Picea rubens 2.1 Thuja occidentalis +.1 Abies bal-
samea +.1 Acer rubrum +-.1 Betula papyrifera -+.1 Vaccinium angus-
tifolium 1.3 Maianthemum canadense 1.3 Gaylussacia baccata 1.3
Eml Dip Hp
Picea rubens 4.1 Thuja occidentalis 1.1 Abies balsamea 1.1 Betula
papyrifera 2.1 Acer rubrum 2.1 Betula lutea 1.1 Aster macrophyllus
1.2 Aster umbellatus 1.2 Maianthemum canadense 1.1 Cornus cana-
densis 1.1 Aralia nudicaulis 1.1
Em.lp Dlsr
Picea rubens 5.1. Abies balsamea 1.1 Thuja occidentalis +-.1 Betula
papyrifera 1.1 Acer pensylvanicum 2.1
Dz GI L Hmlp
Myrica Gale 5.1 Vaccinium Oxycoccos 1.1 Chamaedaphne calyculata,
var, angustifolia -+-.1 Rosa nitida --.1 Carex stricta 4.1 Carex sp. 1.1
Carex lenticularis 4.1 Sphagnum spp. 5.5 Aster nemoralis +-.2 Pogonia
ophioglossoides +-.1 Osmunda regalis, var. spectabilis +-.1
Ds Elp Nir HIp Glp
Alnus crispa, var. mollis 3.3 Acer rubrum 1.1 Tlex verticillata 1.1 Ne-
mopanthus mucronata 1,1 Thuja occidentalis 1.1 Larix laricina +.1
Smilacina trifolia 1.1 Carex spp. 1.2
El Dir Hmp L
Thuja occidentalis 5.1 Acer rubrum +4-.1 Osmunda cinnamomea 3.4
Sphagnum spp. 4.4 moss spp. 1.8
Ds.zp Glp Hr
Populus tremuloides 3.1 Betula populifolia 3.1. Prunus pensylvanica 1.1
Acer rubrum -++.1 Quercus rubra, var. borealis 1.1 Comptonia pere-
131.
132.
1338.
134,
135.
127,
128.
129.
130.
Tue UNiversiry ScreNcE BULLETIN
grina 1.2 Carex spp. 2.38 Lysimachia quadrifolia +.8 Aster macro-
phyllus +.3
Dsi.zp Hlp Glr Ezr Lp bp
Betula papyrifera 3.1 Acer rubrum 1.1 Quercus rubra, var. borealis 1.1
Betula populifolia 1.1 Prunus pensylvancia 1.1 Diervilla Lonicera 1.2
Comptonia peregrina 8.2 Vaccinium angustifolium 2.3 Aralia hispida 1.1
Pteridium aquilinum, var. latiusculum 1.1 Danthonia spicata 1.2 Pinus
rigida 1.1 Polytrichum sp. 8.2
Dm.lp Elp.mr
Acer saccharum 3.1 Fagus grandifolia 3.1 Betula papyrifera 1.1
Betula lutea 1.1 Acer pensylvanicum 1.1 Tsuga canadensis 1.1 Picea
tubens 1.1 Abies balsamea 1.1
Eli Dszp Lp
Picea rubens 4.1 Pinus rigida 1.1 Betula populifolia 1.1 Gaylussacia
baccata 2.3 Vaccinium angustifolium 2.3 Lichen spp. 2.3
Dls Elp
Betula papyrifera 3.1 Betula populifolia 3.1 Prunus pensylvanica 1.1
Amelanchier laevis 1.1 Picea rubens 2.1 Abies balsamea 1.1 Thuja
occidentalis +-.1
Dszp Bzp Ezr Glr bp
Pyrus melanocarpa 2.2 Vaccinium angustifolium 2.2 Betula populi-
folia 1.1 Potentilla tridentata 2.2 Kalmia angustifolia 2.2 Juniperus
communis, var. depressa +-.1 Thuja occidentalis --.1 Festuca capillata
-+-.3 Danthonia spicata 1.3 Deschampsia flexuosa +-.3
Dsz Hlp Gmlp Lp
Betula populifolia 4.1 Diervilla Lonicera 2.2 Prunus pensylvanica 1.2
Spiraea latifolia 1.2 Betula papyrifera -+-.1 Aster umbellatus 1.1
Pteridium aquilinum, var. latiusculum +-.1 Lysimachia quadrifolia --.2
Danthonia spicata 1.2 Deschampsia flexuosa 2.2 Calamagrostis cana-
densis 1.2 Polytrichum sp. 2.2
bp Emlp Dlzp Bzr Lp
Picea rubens 3.1 Thuja occidentalis --.1 Betula populifolia 1.1 Betula
papyrifera +-.1 Vaccinium angustifolium 2.3 Kalmia angustifolia 1.3
Potentilla tridentata +.2 Abies balsamea -+-.1 Cladonia spp. 2.3
Emi Dzp.Jsr Glp Hip
Picea rubens 5.1 Abies balsamea 1.1 Betula populifolia +.1 Viburnum
cassinoides +-.1 Vaccinium angustifolium 2.2 Deschampsia flexuosa 1.3
Cornus canadensis 2.4
Dml.szp Elr Hmlp Glr
Betula papyrifera 3.1 Fagus grandifolia 2.1 Betula populifolia 1.1
Acer saccharum 1.1 Acer rubrum 3.1 Fraxinus americana -+-.1 Betula
lutea +-.1 Acer pensylvanicum 2.1 Abies balsamea +-.1 Aster um-
bellatus 2.1 Solidago rugosa 1.2 Aster macrophyllus 1.8 Onoclea
sensibilis 1.2 Osmunda Claytoniana -+-.1 Dryopteris noveboracensis +-.3
Solidago canadensis +-.1 Carex crinita +-.2
(Dslzp Glp HIp Elr: forest openings in No, 135)
Betula populifolia 2.1 Acer pensylvanicum 2.1 Comptonia peregrina 2.2
Vaccinium angustifolium 2.3 Rubus allegheniensis 2.1 Rubus idaecus,
a
136.
137.
138.
139.
140,
141,
142."
1438,
144.
VEGETATION OF Mount Desert IsLAnp, MAINE S57
var. strigosus 2.1 Danthonia spicata 2.3 Deschampsia flexuosa 2.3
Pteridium aquilinum, var. latiusculum 2.1 Aster umbellatus 1.1 Solidago
canadensis 1.1 Picea rubens 1.1
Dlsp.mr Hlp Glp Elr
Betula papyrifera 8.1 Acer rubrum 8.1 Acer saccharum 2.1 Acer
pensylvanicum 2.1 Prunus pensylvanica 2.1 Aster umbellatus 2.1
Spiraea latifolia 1.2 Poa compressa 1.2 Deschampsia flexuosa 1.2 Pinus
Strobus 2.1 Picea rubens 2.1 Thuja occidentalis 1.1 . Festuca capillata
1.2
Dli Elp Hlp Glp Dsp
Betula populifolia 3.1 Betula papyrifera 1.1 Acer rubrum 2.1 Acer
saccharum 1.1 Acer pensylvanicum 1.1 Prunus pensylvanica 1.1 Rubus
allegheniensis 1.1 Thuja occidentalis 2.1 Pinus Strobus 1.1 Picea
rubens 1.1. Aster umbellatus 1.2 Danthonia spicata +.1 Festuca
capillata +-.1 Poa compressa +.1
Eml Dmlzp Hli
Picea rubens 5.1 Pinus Strobus 2.1 Abies balsamea +.1 Betula
papyrifera 1.1 Acer rubrum 1.1 Acer pensylvanicum +-.1 Vaccinium
angustifolium 8.8 Aralia nudicaulis 1.1 Cornus canadensis 2.1 Mai-
QQ
anthemum canadense 1.1 Pteridium aquilinum, var. latiusculum 38.3
Em Dlszp Hlp
Picea rubens 3.1 Pinus Strobus 2.1 Picea glauca 1.1 Abies balsamea
1.1 Acer rubrum 2.1 Betula populifolia 1.1 Betula papyrifera 1.1 Vac-
cinium angustifolium 1.2 Gaylussacia baccata 1.2 Aralia nudicaulis 1.1
Pteridium aquilinum, var. latiusculum 2.4 Cornus canadensis 1.3 Mai-
anthemum canadense 1.1
Eml
Picea rubens 4.1 Picea glauca 1.1 Thuja occidentalis 2.1
Dis Hm Elr Nir
Acer rubrum 2.1 Betula populifolia 2.1. Fraxinus americana 1.1 Ilex
verticillata 1.1 Alnus rugosa, var. americana 3.1 Aster umbellatus 3.2
Onoclea sensibilis 2.2 Lysimachia terrestris 2.2 Osmunda Claytoniana
1.2 Osmunda cinnamomea 1.2 Solidago rugosa 1.1 Thuja occidentalis
1.1 Abies balsamea 1.1 Larix laricina 1.1
Dl.szp Elr Hmlp Glp
Quercus rubra, var. borealis 3.1. Acer rubrum 2.1 Betula populifolia 2.1
Comptonia peregrina 2.3 Vaccinium angustifolium 2.3 Gaylussacia
baccata 2.8 Pinus rigida 1.1 Aralia nudicaulis 1.1 Pteridium aquilinum,
var. latiusculum 2.3 Danthonia spicata 2.2
Em Hlp Dizr
Picea rubens 4.1 Pinus Strobus 3.1 Abies balsamea 1.1 Aralia nudi-
caulis --.2 Pteridium aquilinum, var. latiusculum 1.2 Cornus canadensis
--.2 Maianthemum canadense +.2 Acer rubrum +-.1 Vaccinium
angustifolium +-.2 Gaylussacia baccata +-.2
Dzp Bzp Hlp Glp Lp bp
Gaylussacia baccata 1.8 Vaccinium angustifolium 2.2 Kalmia angusti-
folia 1.8 Arenaria groenlandica +.2 Potentilla tridentata +-.2 Des-
champsia flexuosa 2.2 Cladonia spp. 2.3
148,
149,
150.
151.
152.
Tse University SCIENCE BULLETIN
Dz.sp Glp
Betula populifolia 4.1 Populus grandidentata 1.1 Acer rubrum -+-.1
Vaccinium angustifolium 3.4 Acer pensylvanicum +-.1 Danthonia spicata
-+.1 Deschampsia flexuosa 3.4
Dm! Hmlp Glp
Acer rubrum 5.1 Quercus rubra, var. borealis -+-.1 Fraxinus americana
1.1 Acer saccharum 1.1 Fagus grandifolis +-.1 Betula papyrifera 1.1
Acer pensylvanicum -+-.1 Aster umbellatus 3.3 Pteridium aquilinum,
var. latiusculum -+.1 Brachyelytrum erectum, var. septentrionale 1.2
Dml.sr Elp Hmlp
Acer rubrum 3.1 Quercus rubra, var. borealis +-.1 Betula papyrifera
3.1 Betula populifolia +.1 Amelanchier laevis +-.1 Acer pensylvanicum
+.1 Abies balsamea 3.1 Picea rubens 1.1. Pteridium aquilinum, var.
latiusculum 1.2 Aster umbellatus --.1 Cornus canadensis 1.1 Maian-
themum canadense 1.1
Dsz.lp Elr Glp Hp
Betula populifolia 3.1 Betula papyrifera 3.1 Populus grandidentata 2.1
Fraxinus americana -+-.1 Populus tremuloides --.1 Salix discolor -+.1
Acer rubrum +.1 Vaccinium angustifolium 1.3 Comptonia peregrina
2.2 Pinus Strobus -+-.1 Picea rubens +-.1 Festuca rubra 1.3 Pteridium
aquilinum, var. latiusculum +.3 Solidago graminifolia +..8 Solidago
rugosa 2.3. Aralia hispida --.2
Eml Nlp Hmlp Glp Lp Dar
Thuja occidentalis 4.1 Picea mariana +-.1 Pinus Strobus +-.1 Larix
laricina 1.1 Osmunda regalis, var. spectabilis +-.2 Osmunda cinnamo-
mea +.2 Onoclea sensibilis +.1 Potentilla simplex, var. calvescens +-.1
Solidago graminifolia +.1 Aster umbellatus +.1 Aster radula -}.1
Viola pallens? +.1 Carex spp. 1.2 Carex cephalantha? 2.3 Calama-
grostis canadensis 2.4 Sphagnum spp. 3.3 Rubus idacus, var. strigosus
3
Dsz Hlp Elr
Betula populifolia 3.1 Betula papyrifera 3.1 Populus grandidentata 2.1
Acer rubrum +.1 Populus tremuloides +-.1 Comptonia peregrina 2.2
Vaccinium angustifolium 1.3 Aralia hispida +-.2 Pteridium aquilinum,
var, latiusculum +.3 Pinus Strobus +-.1 Picea rubens +-.1
Gmlp Hmlp Elp Dmlsr
Anthoxanthum odoratum 2.3 Agrostis tenuis? 2.2 Danthonia spicata
2.2 Festuca ‘capillata 1.2 Panicum sp. 1.3 Carex crinita 8.3. Carex
projecta? +-.2 Carex vulpenoidea? +.2 Poa palustris +.1 Glyceria
striata 1.3 Impatiens capensis +.2 Solidago rugosa +.3 Solidago
canadensis 1.2 Hieracium floribundum +-.1 Hieracium aurantiacum
+.1 Hieracium pratense -+.3 Hieracium Pilosella +-.1 Picea rubens
1.1 Picea glauca 4-.1 Thuja occidentalis +-.1 Betula populifolia +-.1
Betula lutea +.1 Fraxinus americana 4-.1 Rubus idaeus, var. strigosus
8
Dmi.sp Elr Gmlp Hmlp
Quercus rubra, var. borealis 2.1 Acer rubrum 2.1 Betula populifolia 2.1
Betula papyrifera 1.1 Fraxinus americana 8.1 Abies balsamea +-.1
Picea rubens +.1 Anthoxanthum odoratum 1.8 Agrostis alba or A.
1538.
154.
155.
156.
157.
158.
159.
160.
161.
VEGETATION OF Mount Desert IsLanp, MAINE 359
tenuis? 1.2 Panicum sp. -+-.3 Danthonia spicata 1.2 Festuca capillata
+.2 Solidago rugosa +.3 Solidago canadensis 1.2 Hieracium pratense
+.3 Hieracium floribundum +.1 Hieracium Pilosella +.1 Hieracium
aurantiacum +.1
Dz.sr Glp Hlp
Betula papyrifera 4.1 Populus grandidentata +.1 Quercus rubra, var.
borealis +.1 Acer rubrum 1,1 Salix discolor -++-.1 Vaccinium angusti-
folium 2.3 Comptonia peregrina 2.2 Danthonia spicata 1.3 Pteridium
aquilinum, var. latiusculum 1,1 Aralia hispida +-.1
Dls.zp Hli Glp
Quercus rubra, var. borealis 5.1 Acer rubrum +.1 Fagus grandifolia
+.1 Betula papyrifera +-.1 Vaccinium angustifolium +.2 Gaylussacia
baccata +.2 Pteridium aquilinum, var. latiusculum 4.1 Danthonia
spicata 1.2
Dizi.sp Elp Glp
Betula populifolia 4.1 Betula papyrifera 2.1 Acer rubrum 2.1 Populus
grandidentata 1.1 Acer pensylvanicum +.1 Vaccinium angustifolium
3.4 Picea rubens 2.1 Deschampsia flexuosa 3.4 Danthonia spicata +-.1
Ds.mli Elr Hlp Glp
Acer rubrum 8.1 Fagus grandifolia 2.1 Betula papyrifera 2.1 Betula
Jutea 1.1 Acer saccharum 1.1 Acer pensylvanicum 2.1 Fraxinus ameri-
cana 1.1 Rubus idaeus, var. strigosus 3.3 Tsuga canadensis 1.1 Dry-
opteris noveboracensis 1.2 Danthonia spicata 2.1
Ds.mp Glp Hlp Elr
Betula papyrifera 2.1 Betula populifolia 1.1 Acer rubrum 1.1 Quercus
rubra, var. borealis +-.1 Populus tremuloides 1.1 Rubus idaeus, var.
strigosus 1.2 Calamagrostis canadensis 1.8 Brachyelytrum erectum, var.
septentrionale 1.8 Pteridium aquilinum, var. latiusculum +.2 Dryop-
teris noveboracensis +-.8 Viola incognita? +.2 Trientalis borealis +-.1
Thuja occidentalis +.1
El Dir Hml L Nir Glp
Thuja occidentalis 5.1 Picea mariana +.1 Acer rubrum +.1 Osmunda
cinnamomea 3.3. Osmunda regalis, var. spectabilis 3.3 Sphagnum spp.
4.4 Trientalis borealis +-.1 Gaultheria hispidula +-.2 Coptis groen-
landica 1.2 Carex trisperma 1.3 Larix laricina 1.1
Eml Dlsp.mzr Lp
Picea rubens 8.1 Abies balsamea 2.1 Thuja occidentalis 2.1 Pinus Stro-
bus 38.1 Fagus grandifolia 2.1 Acer saccharum 1.1 Acer rubrum 2.1
Quercus rubra, var. borealis 1.1 Acer pensylvanicum 2.1 Fraxinum
americana 1.1 Betula papyrifera 1.1 Populus grandidentata +.1 Vac-
cinium angustifolium 1.2 moss spp. 1.3 lichen spp. 1.3
Dmls Emlp
Fagus grandifolia 2.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer
pensylvanicum 1.1 Acer spicatum 1.1 Betula lutea 1.1 Picea rubens 2.1
Abies balsamea 2.1
Eml Dir
Picea rubens 3,1 Pinus Strobus 3.1 Thuja occidentalis 1.1 Pinus resi-
nosa 1.1 Abies balsamea 1.1 Betula papyrifera 1.1
165.
166.
167,
168,
169.
Li,
gtr
dig,
178.
162,
163,
164,
THe Universrry Science BULLETIN
Dlsz Emp
Acer rubrum 3.1 Quercus rubra, var. borealis 1.1... Betula populifolia 3.1
Populus tremuloides..1.1 Comptonia peregrina 1.2 Pinus Strobus 2.1
Picea rubens 1.1
Dl.mzp Elp Hmlp Glp
Quercus rubra, var. borealis 4.1 Acer rubrum 1.1 Betula populifolia 1.1
Vaccinium angustifolium 3.3. Pteridium aquilinum, var. latiusculum 3.1
grass spp. 3.3 Pinus Strobus 2.1 Picea rubens 1.1
DlLzp Elp Hlr
Betula populifolia 4.1. Acer rubrum 1.1 Vaccinium angustifolium 8.3
Gaylussacia baccata 1.3. Abies balsamea 2/1. Picea: rubens 1.1 Pte-
ridium aquilinum, var. latiusculum +-.3
Dls
Fagus grandifolia 4.1 Betula papyrifera 1.1 Acer rubrum 2.1. Quercus
rubra, var. borealis 1.1 Acer pensylvanicum 1.1 Hamamelis virginiana
+.1
Glwp
Sparganium chlorocarpum 1.3
Dz.sp.lr Gmlp Elr
Chamaedaphne calyculata, var. angustifolia 3.3 Myrica Gale 3.8. Cala-
magrostis canadensis 1.3. Carex stricta +.2 Alnus rugosa, var. ameri-
cana -++-.1 Nemopanthus mucronata ---.2 Ilex verticillata 1.2 Viburnum
cassinoides +-.1 Betula populifolia +.3 Acer rubrum --.1 Picea mari-
ana +.] }
Dz.sp Hr
Betula populifolia 4.1 Comptonia peregrina 3.2 Betula papyrifera 1.1
Acer rubrum 1.1 Populus grandidentata 1.1 Prunus pensylvanica --.1
Aralia hispida 1.2
Dsz.lp Emr Hmp
Betula populifolia 3.1 Comptonia peregrina 3.2 Acer rubrum 3.1 Quer-
cus rubra, var. borealis 1.1 Spiraea latifolia +-.1 Salix discolor +.1
Vaccinium angustifolium 3.3 Pinus Strobus 1.1 Pteridium aquilinum,
var. latiusculum 2.1
Dz Glp Lp
Betula populifolia 5.1 Comptonia peregrina 2.8 Vaccinium angustifo-
lium 1,2 Acer rubrum -+.1 Festuca capillata 2.4 Festuca rubra +.1
Agrostis tenuis? 1.3 Danthonia spicata +.2 Polytrichum sp. 2.4
wp Hip Glp
Nymphaea odorata 2.4 Nuphar variegatum +-.2 Eleocharis Smallii? 1.3
Brasenia Schreberi --.3 Pontederia cordata +-.8 Sparganium chlorocar-
pum +.3
Dzp Bzr Gmlp
Chamaedaphne calyculata, var. angustifolia 2.8 Alnus crispa, var. mollis
2.1 Myrica Gale 2.1 Kalmia angustifolia 1.2 Carex lenticularis 3.3
Carex rostrata --.3
Eli Dzp Hr
Picea glauca 2.1 Picea rubens 2.1 Pinus Strobus 2.1 Vaccinium angusti-
folium 2.2 Comptonia peregrina 2.2 Pteridium aquilinum, var. latius-
culum 1.1
ee
174,
175.
176.
Were
Lok
179.
180.
181.
182.
Veceration oF Mount Desert IstANnp, MAINE 361
3ml Dszp Hip Lp
Carex stricta 3.1 Carex rostrata 1.3 Carex spp. 1.1 Calamagrostis
canadensis 2.3 Eriophorum virginicum 2.1 Rhynchospora alba 1.3
Cladium mariscoides 1.3 Myrica Gale 8.1 Chamaedaphne calyculata,
var. angustifolia 1.8 Alnus rugosa, var. americana 1.3 Rubus hispida,
var. obovalis 2.1 Lysimachia terrestris 1.1 Drosera intermedia ++.1
Pogonia ophioglossoides -+-.1 Sphagnum spp. 4.4
Dsz.mlp Bzr Hr
Quercus rubra, var. borealis 3.1 Acer rubrum 2.1 Populus grandidentata
3.1 Betula populifolia 1.3 Comptonia peregrina 2.3 Vaccinium angusti-
folium 1.8 Kalmia angustifolia 1.8 Pteridium aquilinum, var. latius-
culum 1.1
Dsi.zp Elp Hli
Acer rubrum 3.1 Quercus rubra, var. borealis 1.1 Betula papyrifera 3.1
Populus grandidentata 2.1 Vaccinium angustifolium 3.3 | Comptonia
peregrina 2.3 Picea rubens 2.1 Pinus Strobus 1.1 Pteridium aquilinum,
var. latiusculum 4.3. Aralia hispida +-.1
Dzi.sp.mr Hmlp Glp Emlr
Betula populifolia 2.1 Betula papyrifera 2.1 Acer saccharum -+.1
Fagus grandifolia +-.1 Betula lutea --.1 Acer rubrum 2.1 Rubus
idaeus, var. strigosus 1.2 Solidago canadensis 2.1 Solidago graminifolia
--.1° Polygonum cilinode 1.4 Dennstaedtia punctilobula +.3 Aralia
hispida +.3 Agrostis tenuis? 1.2 Danthonia spicata 2.2 Festuca capil-
lata -+-.2 Picea rubens +.1 Abies balsamea +-.1
Dsz Bszp Lp
Chamaedaphne calyculata, var. angustifolia 8.1. Myrica Gale 3.1 Ledum
groenlandicum 1.1 lex verticillata 2.1_ Viburnum cassinoides 2.1 Vac-
cinium corymbosum -+.1 Kalmia angustifolia +-.1 Rhododendron cana-
dense +-.1_ Sphagnum sp. 3.3. Polytrichum sp. -++.2
Em.lp Dszp.lr Hlp Lp
Picea rubens 3.1 Pinus resinosa 3.1 Pinus Strobus 2.1 Abies balsamea
2.1 Viburnum cassinoides 2.1 Gaylussacia baccata 1.2 Vaccinium
angustifolium 3.3 Acer rubrum +-.1 Gaultheria procumbens +-.2 Aralia
nudicaulis 1.1. Cornus canadensis 3.1 Maianthemum canadense -+-.1
Pteridium aquilinum, var. latiusculum 1.1 moss spp. 2.2
Dls Elr Hmi Glr
Fraxinus americana 4.1 Acer rubrum 2.1 Alnus rugosa, var. americana
2.3 Betula lutea +:.1 Acer saccharum 1.1 Ilex verticillata 1.1 Thuja
occidentalis 1.1 Pinus Strobus -+.1 Osmunda regalis, var. spectabilis 3.4
Aster umbellatus 1.1 Solidago canadensis +-.1 Carex spp. 1.2
Eml.zp
Picea rubens 5.1 Pinus Strobus 1.1
Eml H1 Disp.mzr
Picea rubens 4.1 Pinus Strobus 2.1 Abies balsamea 1.1 Pinus resinosa
1.1 Acer rubrum 2.1 Betula papyrifera --.1 Viburnum cassinoides 1.1
Gaultheria procumbens +-.1 Vaccinium angustifolium 1.3 Aralia nudi-
caulis 1.1 Pteridium aquilinum, var. latiusculum 3.3 Maianthemum
canadense 2.3 Mitchella repens -++.1
362
183.
184.
185.
187.
188.
189,
190,
191,
Tue University ScreENCE BULLETIN
Emlp Dlszp Hmlp Glp
Pinus Strobus 2.1 Pinus resinosa +.1 Picea rubens +.1 Picea glauca
+.1 Acer rubrum 1.1 Betula papyrifera 2.1 Betula populifolia 1.1
Populus tremuloides +.1 Populus grandidentata +-.1 Ilex verticillata
1.1 Viburnum cassinoides 1.1 Alnus crispa, var. mollis 1.1 Salix dis-
color +.1 Acer pensylvanicum +.1 Comptonia peregrina 2.2 Vac-
cinium angustifolium 2.3 Aralia hispida 1.1 Pteridium aquilinum, var.
latiusculum 2.38 Aster macrophyllus 2.1 Solidago spp. 1.1 Danthonia
spicata 2.4 Festuca capillata 1.3 Anthoxanthum odoratum 1.8 Agrostis
tenuis? 1.3
Dszi Elsp Glp Hlr L
Populus tremuloides 2.1 Betula papyrifera 2.1 Salix discolor 1.1
Comptonia peregrina 1.3 Betula populifolia 2.1 Picea rubens 1.1
Danthonia spicata 1.3 Aralia hispida +-.1 Polytrichum sp. 5.5
Dmsi.zp Emr Hlp
Quercus rubra, var. borealis 4.1 Betula papyrifera 2.1 Populus grandi-
dentata 1.1 Comptonia peregrina 1.2 Vaccinium angustifolium 1.3
Pinus Strobus 1.1 Danthonia spicata +.2 Pteridium equilinum, var.
latiusculum 3.1
Ds.zp Hlp Glp Lp
Betula populifolia 2.1 Betula papyrifera 1.1 Quercus rubra, var. bo-
realis 1.1 Fagus grandifolia 1.1 Acer rubrum 3.1 Prunus pensylvanica
1.1 Comptonia peregrina 1.2 Populus grandidentata 1.1 Rubus
allegheniensis +-.1 Rubus idaeus, var. strigosus 1.4 Aralia hispida +.3
Pteridium aquilinum, var. latiusculum 1.1 Solidago canadensis +-.1
Festuca capillata 1.4 Danthonia spicata +.4 Deschampsia flexuosa +-.2
Carex pensylvanica?P +.2 Polytrichum sp. 2.4
Em
Picea rubens 5.1 Picea glauca +.1
Dmlz
Quercus rubra, var. borealis 5.1 Acer rubrum 1.1 Betula papyrifera 1.1
Rubus allegheniensis 3.3 Rubus idaeus, var. strigosus 4.3
Dis Elp Hmlp
Betula populifolia 3.1 Fraxinus americana 1.1 Alnus rugosa, var. amer-
icana 3.1 Thuja occidentalis 2.1 Aster umbellatus 2.1 Dryopteris
Thelypteris, var. pubescens 1.3
Gmlp Dsp Hmp
Calamagrostis canadensis 3.3 Scirpus cyperinus, var. pelius +.2 Salix
discolor 1.1 Alnus rugosa, var. americana 3.2 Solidago canadensis 1.1
Dmzp Elp Hmlp Glp
Betula papyrifera 2.1 Betula populifolia 1.1 Acer rubrum 3.1 Quercus
rubra, var. borealis 1.1 Fraxinus americana +.1 Comptonia peregrina
+.2 Picea rubens 1.1 Pinus Strobus 1.1 Tsuga canadensis +.1 Dry-
opteris noveboracensis +.3 Pteridium aquilinum, var. latiusculum 4+-.3
Osmunda Claytoniana +.2 Solidago canadensis +.2 Aster umbellatus
1.1 Aster macrophyllus +.38 Solidago rugosa +.2 Festuca capillata
2.2 Danthonia spicata 1.2 Agrostis alba? 1.2 Calamagrostis canaden-
sis +-.2 Carex spp. 2.3 Scirpus cyperinus, var. pelius 1,2
193.
194.
195.
196.
197.
198.
199.
VEGETATION oF Mount Desert Istanp, MAINE 363
Dz.mlsr Elr Hlp Glp Lp
Betula papyrifera 3.1 Betula populifolia +.1 Comptonia peregrina 2.2
Populus tremuloides +-.1 Prunus pensylvanica 1.1 Salix discolor +-.1
Quercus rubra, var. borealis +.1 Acer rubrum 1.1 Pinus Strobus +.1
Epilobium angustifolium 1.3 Anaphalis margaritacea, var. intercedens
+.2 Erigeron canadensis +-.3 Agrostis scabra --.2 Agrostis alba? or
tenuis? +.2 Polytrichum sp. 2.2
Emlp Nmlp L Dszp Bzp Hmp Glp
Picea mariana 8.1 Thuja occidentalis 1.1 Abies balsamea +.1 Larix
laricina 1.1 Sphagnum sp. 5.5 Gaylussacia baccata +.1 Nemopanthus
mucronata 1.1 Viburnum cassinoides 1.1 Alnus rugosa, var. americana
+.1 Acer rubrum +.1 Ledum groenlandicum -++.1 Rhododendron
canadense +.1 Galmia angustifolia +-.1 Osmunda cinnamomea +.3
Aster radula --.1 Smilacina trifolia +.1 Carex trisperma 2.4
Eml Nmlp Dir Hml L
Picea mariana 2.1 Pinus Strobus 2.1 Thuja occidentalis 4.1 Larix
laricina 3.1 Acer rubrum 1.1 Dryopteris noveboracensis 1.3 Osmunda
regalis, var. spectabilis 2.4 Osmunda cinnamomea 3,4 Impatiens capen-
sis 1.2 Trientalis borealis 1.1 Aster radula 1.1 Sphagnum sp. 5.5
Dzi Bzp Glmp Elp Nir Hlp L wr
Myrica Gale 2.3 Gaylussacia baccata 1.3 Gaylussacia dumosa, var.
Bigeloviana 2.1 Andromeda glaucophylla +.1 Pyrus melanocarpa +.1
Vaccinium Oxycoccos 1.1 Empetrum nigrum +-.4 Chamaedaphne caly-
culata, var. angustifolia 1.4 Ledum groenlandicum +.4 Rhynchospora
alba 2.4 Sarracenia purpurea +.1 Drosera rotundifolia +-.3 Drosera
intermedia +.3 Calopogon pulchellus -++.1 Solidago uliginosa, var.
linoides -+.1 Aster nemoralis +.1 Calamagrostis canadensis 1.4 Eri-
ophorum virginicum -+-.1 Eriophorum spissum -++.1 Carex stricta 1.4
Carex lenticularis 1.4 Carex rostrata +.3 Scirpus cespitosus, var. cal-
losus +.4 Sphagnum spp. 5.5 Picea mariana 1.1 Larix laricina 1.1
Nymphaea odorata -+.5 Nuphar variegatum +-.3
Dm.sp Elp
Betula papyrifera 3.1 Acer rubrum 3.1 Acer saccharum 1.1 Populus
tremuloides 1.1 Betula lutea 1.1 Fraxinus americana 1.1 Acer pensyl-
vanicum 2.1 Abies balsamea 3.1 Picea rubens 2.1
Eli Diszp Hlp Lp
Picea rubens 2.1 Thuja occidentalis 2.1 Abies balsamea 2.1 Acer
rubrum 2.1 Betula papyrifera 1.1 Vaccinium angustifolium 3.3 Vi-
burnum cassinoides +.3 Cornus canadensis 2.3 Cladonia spp. 1.3
Dlp Elp Hlp Lp
Acer rubrum 3.1 Betula papyrifera 2.1 Betula populifolia 1.1 Thuja
occidentalis 8.1 Abies balsamea -+.1 Picea rubens +.1 Cornus cana-
densis 1.3 Maianthemum canadense +.3 Aralia nudicaulis +.1 Trien-
talis borealis --.1 moss spp. 3.4
Dzp Glp
Spiraea latifolia 3.1 Diervilla Lonicera 3.1 Betula populifolia 1.1
Prunus pensylvanica +-.1 Alnus crispa, var. mollis +.1 Solidago Randii
+.3 Festuca capillata 2.3 Calamagrostis canadensis 1.2 Danthonia
spicata +.2
204.
205.
206.
207.
208.
200.
201.
202.
Tue University SCIENCE BULLETIN
Elsp Dip Hml
Picea glauca 3.1 Picea rubens 2.1 Betula papyrifera 3.1 Acer pensyl-
vanicum 1.1 Acer rubrum +.1 Rubus allegheniensis 2.3 Aster macro-
phyllus 4.5 Aster umbellatus 1.1 Pteridium aquilinum, var. latiusculum
2.3 Aralia nudicaulis 1.1
w Gmlp
Nuphar variegatum 3.1 Nymphaea odorata 3.1 Sparganium chloro-
carpum 1.4 Typha latifolia 2.5
Dsi Hmp
Alnus rugosa, var. americana 2.1 Salix gracilis 1.1 Salix discolor 1.1
Prunus pensylvanica 2.1 Betula populifolia 2.1 Betula papyrifera 2.1
Spiraea latifolia 1.1 Solidago canadensis 1.1
Dzi.sp.mlr Bzr Glp Hlp
Quercus rubra, var. borealis 1.1 Betula populifolia 1.1 Vaccinium
angustifolium 4.4 Comptonia peregrina 4.3 Spiraea latifolia 1.2 Kalmia
angustifolia 1.8 Viburnum cassinoides -++.1 Danthonia spicata 3.3
Festuca capillata 1.3 Pteridium aquilinum, var. latiusculum 1.3 Aster
macrophyllus +.1 Aster umbellatus +-.1 Hieracium aurantiacum +-.1
Hieracium pratense +.1 Apocynum androsaemifolium +-.1
Gm Dzp.sr wr
Carex stricta? 5.4 Calamagrostis canadensis -+-.2 Myrica Gale 2.3
Spiraea latifolia +-.1 Alnus rugosa, var. americana +.3 Acer rubrum
-+-.8 Chamaedaphne calyculata, var. angustifolia -+-.2 Nuphar varie-
gatum +.2
Gl Hlp Dlr
Poa pratensis 5.5 Agrostis tenuis 1.1 Festuca capillata +.2 Trifolium
repens 1.38 Ranunculus acris +.2 Ranunculus bulbosus --.1 Cerastium
vulgatum +.2 Rumex acetosella +-.2
Emp Nmp Dzp.lr Lp Hlr Gmr
Picea mariana 3.1 Larix laricina 8.1. Abies balsamea +-.1 Betula populi-
folia +.2 Comptonia peregrina 2.3 Vaccinium angustifolium 3.8
Myrica Gale 3.2 Viburnum cassinoides -+-.1 Chamaedaphne calyculata,
var. angustifolia +.1 Ledum groenlandicum +-.2 Sphagnum spp. 1.3
Osmunda regalis, var. spectabilis +-.2 Osmunda cinnamomea -4-.2
Calamagrostis canadensis +-.3
Dzi.sr Hlp Glp
Comptonia peregrina 4.3 Vaccinium angustifolium 4.4 Betula populi-
folia +.1 Aster umbellatus +-.1 Pteridium aquilinum, var. latiuscu-
lum +.8 Oryzopsis asperifolia --.2 Danthonia spicata 2.2
Emi Nmi Dsp Gmp Hmp L
Picea mariana 3.1 Larix laricina 3.1. Alnus rugosa, var. americana 3.3
Acer rubrum +.1 Viburnum cassinoides --.1 Carex trisperma 8.3 Cala-
magrostis canadensis 2.3 Rubus pubescens -+.1 Cornus canadensis 1.3
Lysimachia terrestris 1.1 Smilacina trifolia 1.1 Coptis groenlandica --.8
Dryopteris Thelypteris, var. pubescens +-.1 Osmunda cinnamomea +-.1
Sphagnum sp. or spp. 5.4
209.
210.
218.
217,
219,
VEGETATION oF Mount Desert IsLaAND, MAINE 865
Dsz Elr Hlp
Comptonia peregrina 4.3 Betula populifolia 2.1 Vaccinium angusti-
folium 1.8 Pinus Strobus 1.1 Pteridium aquilinum, var. latiusculum 2.2
Aralia hispida 1.1
Dszi Gmp Lp
Acer rubrum 1.1 Alnus rugosa, var. americana 2.2 Spiraea latifolia 3.4
Vaccinium macrocarpon -+.3 Typha latifolia 2.4 Carex spp. 1.3 Carex
rostrata, var, utriculata 1.2 Calamagrostis canadensis +-.2 Sphagnum
sp. or spp. 4.4
Dsi.zp.mlr Emlr Hmp Glp
Quercus rubra, var. borealis 3.1 Acer rubrum 8.1 Betula populifolia 1.1
Comptonia peregrina 1.4 Vaccinium angustifolium 2.4 Pinus Strobus 1.1
Pteridium aquilinum, var. latiusculum 2.4 Danthonia spicata 1.2
Dzi.sp Hmp
Quercus rubra, var. borealis 1.1 Acer rubrum +-.1 Betula populifolia 1.2
Comptonia peregrina 3.4. Vaccinium angustifolium 8.4 Pteridium aquili-
num, var. latiusculum 2.2
Dliszp Glp Hlp
Betula populifolia 3.1 Betula papyrifera 1.1 Quercus rubra, var. borealis
2.1 Acer rubrum 2.1 Comptonia peregrina 2.4 Danthonia spicata +.2
Festuca capillata --.2 Poa pratensis +-.2 Pteridium aquilinum, var.
latiusculum 2.4 Aster umbellatus +-.1 Lysimachia quadrifolia +.8
Gm Dszp Lp
Calamagrostis canadensis 5.5 Carex spp. +.2 Typha latifolia +.2
Alnus rugosa, var. americana +.3 Chamaedaphne calyculata, var. an-
gustifolia +-.1 Myrica Gale 1.3 Ledum groenlandicum +-.2 Sphagnum
Sete eke Cheyay tinal
Dsz Glr L
Chamaedaphne calyculata, var. angustifolia 5.5 Myrica Gale +.3 Carex
spp. -+.4 Sphagnum spp. 5.5
Emlp Dzp Bzp L
Picea mariana 3.1 Vaccinium angustifolium 1.1 Chamaedaphne caly-
culata, var. angustifolia 1.2 Rhododendron canadense 8.1 Kalmia an-
gustifolia 3.1 Ledum groenlandicum 1.1 Cladonia spp. 4.3
Dz.sr Glp Hlp
Comptonia peregrina 3.4. Spiraea latifolia 2.3 Alnus rugosa, var. ameri-
cana 2.3 Danthonia spicata 1.1 Festuca capillata 8.2 Agrostis tenuis
1.2 Ranunculus acris +-.1 Chrysanthemum Leucanthemum, var. pinna-
tifidum -+-.2 Ranunculus bulbosus ++.1
El Dsp Nir
Picea mariana 4.1. Acer rubrum -+.1 Alnus rugosa, var. americana 3.4
Larix laricina 1.1
Dsz Hmp Glp
Acer rubrum 2.1 Quercus rubra, var. borealis 1.1 Comptonia peregrina
8.3 Betula populifolia 1.8 Populus grandidentata -+-.1 Gaylussacia
baceata +-.8 Rubus idaeus, var. strigosus 4+-.2 Rubus allegheniensis +-.2
Pteridium aquilinum, var. latiusculum 2.1 Festuca capillata 1.2 Dan-
thonia spicata 1.2
220.
223,
224,
225,
226.
227,
228.
229.
230.
231.
Tue University Science BULLETIN
Dml.zp Elr Hmp
Quercus rubra, var. borealis 5.1 Betula populifolia 2.1 Populus grandi-
dentata +.1 Vaccinium angustifolium +.3 Picea rubens +.1 Pinus
Strobus --.1 Abies balsamea +.1 Pteridium aquilinum, var. latiusculum
21
Not on map.
Eml Dzp.Jsr Nmlp Hmlp Lp
Picea mariana 5.1 Abies balsamea +.1 Acer rubrum -+.1 Alnus ru-
gosa, var. americana +.3 Vaccinium angustifolium 1.3 Larix laricina
1.1 Pteridium aquilinum, var, latiusculum 1.8 Cornus canadensis 1.2
Osmunda cinnamomea +.1 Osmunda Claytoniana +.1 Equisetum syl-
vaticum, forma multiramosum -++.1 Calamagrostis canadensis +-.3 moss
spp. 1.8 Sphagnum sp. or spp. +.3
Eml Dszp Hmlp Glmp L
Picea mariana 5.1 Alnus rugosa, var. americana +-.1 Acer rubrum +-.1
Viburnum cassinoides +.1 Ilex verticillata +-.1 Nemopanthus mucro-
nata +.1 Ledum groenlandicum +.1 Gaultheria hispidula +.1 Os-
munda regalis, var. spectabilis +.1 Osmunda cinnamomea 1.1 Carex
trisperma 4.5 Calamagrostis canadensis +.4 Carex paupercula, var.
irrigua --.1 Sphagnum sp. or spp. 5.5
Eml Nmlp Dlszp Hmlp Glmp L
Picea mariana 5.1 Larix laricina 1.1 Acer rubrum 1.1 Ledum groen-
landicum +.3 Osmunda cinnamomea 3.3. Trientalis borealis +.1 Carex
trisperma 4.5 Calamagrostis canadensis +.3 Sphagnum sp. or spp. 5.5
Not on map.
Gml Dzp Hip
Festuca rubra 4.5 Agrostis tenuis 4.5 Agropyron repens 1.8 Danthonia
spicata --.1 Phleum pratense +-.1 Comptonia peregrina 1.3 Hieracium
pratense +.1 Hieracium floribundum +.1 Hieracium Pilosella +.1
Hieracium aurantiacum +.1 Vicia Cracca +-.3
Eml Dlp Hlp Lp
Abies balsamea 4.1 Picea glauca 2.1 Picea rubens +.1 Betula popu-
lifolia 1.1 Betula papyrifera +.1 Trientalis borealis +.1 Maianthemum
canadense 1.1 Cornus canadensis 1.1 Linnaea borealis, var. americana
+.1 Pteridium aquilinum, var. latiusculum +.1 moss spp. 2.3 Acer
rubrum 2,1
Dli.zp Elp Hp
Quercus rubra, var. borealis 2.1. Acer rubrum 1.1 Betula populifolia 3.1
Vaccinium angustifolium 2.4 Comptonia peregrina 1.3 Pinus Strobus 2.1
Abies balsamea 2.1 Picea rubens 1.1 Pteridium aquilinum, var, latius-
culum 1.1
Not on map.
Not on map.
Gml Hmlp
Dactylis glomerata 2.8 Anthoxanthum odoratum 4.3 Festuca rubra 1.2
Rhinanthus Crista-galli 2.1 Chrysanthemum Leucanthemum, var. pin-
natifidum 1.2 Stellaria graminea 1.1 Ranunculus acris +-.2 Vicia Cracca
1.8 Hieracium pratense 1.3 Hieracium aurantiacum +-.2
232.
233,
234,
236.
238.
239,
240.
VEGETATION OF Mount Desert IsLAND, MAINE 867
Dzp Bzp Hlp Glp Lp bp
Vaccinium angustifolium 2.8 Pyrus melanocarpa 3.8 Alnus crispa, var.
mollis 1.1 Kalmia angustifolia 2.3 Potentilla tridentata +-.1 Arenaria
groenlandica +-.3 Solidago Randii +.1 Hypericum gentianoides +.2
Deschampsia flexuosa 2.3 Cladonia spp. 2.8
Dmls Elr Hml Glp Lp
Acer rubrum 4.1 Fraxinus nigra 1.1 Alnus rugosa, var. americana +-.3
Ilex verticillata +-.1 Abies balsamea 1.1 Thuja occidentalis +-.1 Onoclea
sensibilis 2.1 Aster umbellatus 1.1 Osmunda Claytoniana +-.2 Osmunda
cinnamomea -+.2 Rubus pubescens +-.1 Athyrium Filix-femina, var.
Michauxii? --.2 Arisaema strorubens +.1 Equisetum sylvaticum, forma
multiramosum 1.3 Calamagrostis canadensis 3.3 Carex spp. 1.2 Sphag-
num sp. or spp. 2.3.
Gl Hlp
Festuca rubra 8.2 Anthoxanthum odoratum 2.2 Agrostis tenuis 3.2
Agropyron repens 1.1 Phleum pratense 1.1 Carex spp., incl. especially
Carex scoparia, 1.4 Scirpus spp. -+-.2 Chrysanthemum Leucanthemum,
var, pinnatifidum 1.2 Vicia Cracca 2.3 Rhinanthus Crista-galli 1.1
Hieracium pratense +-.2 Hieracium aurantiacum -++.2 Ranunculus acris
+.1 Stellaria graminea +-.1 Rumex Acetosella +.2 Luzula multiflora
+.
Ds.zp Emlr Hlp Lp
Quercus rubra, var. borealis 3.1 Acer rubrum 3.1 Populus grandidentata
1.1 Populus tremuloides +.1 Betula papyrifera +-.1 Comptonia pere-
grina 2.1 Vaccinium angustifolium 1.2 Diervilla Lonicera 2.4 Pinus
Strobus 1.1 Pinus resinosa 1.1 Pteridium aquilinum, var. latiusculum
1.1 Polytrichum sp. 2.2
Eli.zp Dli
Picea rubens 3.1 Thuja occidentalis 1.1 Juniperus communis, var.
depressa 1.3 Acer rubrum 2.1 Betula populifolia 1.1 Pinus rigida +-.1
Polypodium virginianum +-.2
Dzp Bzr Eszr Hlp Glr Lp bp
Vaccinium angustifolium 2.4 Betula populifolia 1.8 Pyrus melanocarpa
-++.1 Kalmia angustifolia 1.3 Pinus rigida 1.1 Juniperus communis, var.
depressa -+.2 Potentilla tridentata 1.3 Hypericum gentianoides +.3
Arenaria groenlandica +-.2 Deschampsia flexuosa +-.2 lichen spp. 4.4
Dli.sp.zr Eli Hmlp Glr Lp
Betula populifolia 4.1 Alnus rugosa, var. americana 1.1 Vaccinium
angustifolium +-.3 Picea rubens 3.1 Thuja occidentalis 3.1 Picea
glauca 1.1 Abies balsamea 1.1 Aster umbellatus 1.2 Dryopteris nove-
boracensis 1.3 Viola cucullata +-.2 Festuca capillata 1.8 moss spp. 2.3
Gl Dzr Hlr
Festuca capillata 4.2 Agrostis tenuis 3.2 Danthonia spicata 1.2 Vac-
cinium angustifolium +-.3 Vaccinium Vitis-Idaea, var. minus +-.3 Hier-
acium pratense +-.1 Ranunculus acris +-.1 Solidago nemoralis +-.1
Gml Hml
Dactylis glomerata 2.2 Phleum pratense 2.2 Festuca rubra 2.2 Agrostis
tenuis (possibly also A. alba) 2.2 Anthoxanthum odoratum -++-.2 Agro-
246.
247,
248.
241,
243,
244,
245,
THe Universtry Science BuLLeTin
pyron repens +-.2 Rumex Acetosella 3.1 Chrysanthemum Leucanthe-
mum, var. pinnatifidum 2.3 Ambrosia artemisiifolia, var. elatior 1,1
Hieracium pratense 1.1 Hieracium floribundum 1.1 Trifolium agrarium
1.2 Trifolium pratense +.2 Trifolium repens +.2 Trifolium hybridum
-+-.1 Raphanus Raphanistrum --.1 Vicia Cracca +.1 Vicia tetrasperma
+.1 Stellaria graminea +.1 Triticum spp. +.1
Gml Hmlp Dszp Esr
Festuca rubra 3.2 Festuca capillata 2.2 Danthonia spicata 2.2 Agrostis
tenuis 2.2 Agropyron repens 1.2 Luzula multiflora +-.2 Chrysanthe-
mum Leucanthemum, var. pinnatifidum -+-.2 Solidago canadensis -+-.3
Solidago graminifolia --.1 Solidago nemoralis +-.1 Vicia Cracea +.1
Potentilla simplex, var. calvescens +-.1 Comptonia peregrina 2.3 Alnus
rugosa, var. americana 1.4 Pinus Strobus 1.1
Gl Dsp Elr Hmlp
Festuca capillata 3.2 Festuca rubra 2.3 Agrostis tenuis, possibly also
A. alba, 2.3 Agropyron repens 1.3 Phleum pratense --.2 Anthoxanthum
odoratum 2.2 Typha latifolia 1.4 Carex spp., especially C. scoparia,
+.2 Luzula multiflora +.2 Comptonia peregrina 1.4 Alnus rugosa,
var. americana 2.4 Spiraea latifolia +.1 Betula populifolia +-.1 Rosa
carolina? +.1 Pinus rigida 1.1 Potentilla simplex, var. calvescens 1.1
Solidago graminifolia 1.1 Ranunculus acris 1.1 Rhinanthus Crista-galli
1.1 Vicia Cracca +.1 Chrysanthemum Leucanthemum, var. pin-
natifidum +-.1 Aster umbellatus +-.1
Gm Dsp
Calamagrostis canadensis 5.5 Populus tremuloides 3.3. Acer rubrum --.1
Salix discolor and probably S. gracilis +-.1
Ds.lp
Populus tremuloides 4.1 Spiraea latifolia 2.1 Salix discolor and prob-
ably S. gracilis 2.1 Betula populifolia 2.1
Gm DIsp
Calamagrostis canadensis 5.5 Spiraea latifolia 1.1 Salix discolor and
probably S. gracilis 1.1 Populus tremuloides 1.1 Betula populifolia -+-.1
Dls Glr
Populus tremuloides 5.1 Spiraea latifolia 3.1 Comptonia peregrina 1.4
Betula populifolia +-.1 Rosa palustris? (possibly R. carolina) -+.1
Agrostis tenuis, possibly also A. alba, -+.2
Dzp Bzp
Kalmia angustifolia 3.1 Ledum groenlandicum 2.1 Chamaedaphne caly-
culata, var. angustifolia 3.1 Rhododendron canadense 3.1
Emi Disp Hmlp Glp
Pinus resinosa 3.1 Pinus Strobus 1.1. Quercus rubra, var. borealis 2.1
Acer rubrum 8.1 Betula papyrifera 1.1 Populus grandidentata +-.1
Fagus grandifolia --.1 Acer pensylvanicum -+.1 Hamamelis virginiana
+.1 Comptonia peregrina +.3 Pteridium aquilinum, var. latiusculum
+.3 Osmunda Claytoniana +-.3 Lysimachia quadrifolia +-.3 Maian-
themum canadense +.3 Aster macrophyllus --.1 Hieracium pratense
+.1 Solidago bicolor +.1 Poa compressa 1.3 Festuca capillata 1.3
249,
250.
251,
252.
253.
255.
VEGETATION OF Mount Deserr IsLAND, MAINE 869
Ds.mzp Emr Hlp Lp
Acer rubrum 3.1 Quercus rubra, var. borealis 2.1 Betula papyrifera 4.1
Prunus pensylvanica +-.1 Comptonia peregrina 2.3 Vaccinium angusti-
folium 1.3 Pinus Strobus 1.1 Tsuga canadensis 1.1 Pteridium aquili-
num, var. latiusculum 1.1 Aralia hispida +-.1 Gaultheria procumbens
+.1 Polytrichum sp. 1.2
Dz.sp Bzp Hlp Glr Ezr Lp bp
Betula populifolia 8.1 Vaccinium angustifolium 3.3 Prunus pensyl-
vanica +.1 Diervilla Lonicera +.2 Pyrus melanocarpa +-.2 Kalmia
angustifolia 2.3 Pteridium aquilinum, var. latiusculum 1.1 Potentilla
tridentata 1.8 Danthonia spicata -+-.2 Deschampsia flexuosa +-.2 Pinus
rigida +.1 lichen spp. 2.4 Polytrichum sp. 1.3
Dsz.mp Emr HIp
Acer rubrum 8.1 Quercus rubra, var. borealis 2.1 Betula papyrifera 3.1
Populus grandidentata 2.1 Populus tremuloides +-.1 Fraxinus ameri-
cana +-.1 Acer pensylvanicum -+-.1 Comptonia peregrina 8.3 Alnus
rugosa, var. americana +.4 Pinus Strobus 1.1 Pteridium aquilinum,
var. latiusculum 2.1 Aster macrophyllus 2.3 Lysimachia quadrifolia 1.1
Osmunda Claytoniana --.3 Dryopteris noveboracensis +-.2
Dszp Elr Glp bp
Betula papyrifera 4.1 Comptonia peregrina 8.1 Quercus rubra, var.
borealis 2.1 Acer rubrum +.1 Fraxinus americana -+-.1 Prunus pen-
sylvanica +-.1 Vaccinium angustifolium 1.2 Pinus rigida 1.1 Des-
champsia flexuosa 1.2 Danthonia spicata 1.2 Aralia hispida +-.1
Dszi Gmlp Hmlr
Quercus rubra, var. borealis 8.1 Acer rubrum 1.1 Betula papyrifera 3.1
Prunus pensylvanica 1.1 Populus tremuloides +-.1 Diervilla Lonicera
1.1 Comptonia peregrina 1.8 Spiraea latifolia 1.1 Rubus idaeus, var.
strigosus 1.1 Alnus crispa, var. mollis +-.4 Salix discolor -+-.1 Rhus
typhina +-.1 Danthonia spicata 2.2 Deschampsia flexuosa 1.2 ' Festuca
capillata +-.4 Calamagrostis canadensis --.8 Aralia hispida +-.1 Aster
macrophyllus +-.1
GI Hp
Festuca rubra 5.5 Festuca capillata 2.3 Agropyron repens 2.2 Anthox-
anthum odoratum 1.2 Agrostis tenuis 1.2 Phleum pratense 1.2 Poa
compressa 1.2 Dactylis glomerata 1.2 Vicia Cracea 1.1 Hypericum
perforatum +-.3 Chrysanthemum Leucanthemum, var. pinnatifidum -++.3
Cirsium arvense +-.3 Hieracium pratense +-.3 Rosa carolina? +.3
Picea glauca +.1 Alnus rugosa, var. americana 8.3 Betula papyrifera
2.1 Populus tremuloides 2.3 Prunus pensylvanica 2.8 Rubus alleghe-
niensis 1.3 Spiraea latifolia 1.1 Osmunda Claytoniana 1.8
Dszi Gmlp Hmp
Salix discolor 8.1 Salix gracilis? 3.1 Salix rigida +-.1 Spiraea latifolia
3.1 Typha latifolia 1.3 Typha angustifolia --.3 Scirpus cyperinus, var.
pelius +-.3 Agrostis tenuis, perhaps also A. alba, 2.3 Carex scoparia 2.2
Calamagrostis canadensis +-.2 Lythrum Salicarea 2.3 Aster novi-belgii
1.1 Lysimachia terrestris -+.8
18—3378
256.
257.
258.
259.
260.
261.
262.
Tur Universiry SCrENCE BULLETIN
Ds.mlzp Emr Hmlp Glp
Betula papyrifera 3.1 Populus tremuloides 2.1 Prunus pensylvanica 2.1
Fraxinus americana -+.1 Acer pensylvanicum +.1 Acer spicatum +-.1
Acer rubrum +.1 Rubus allegheniensis 2.2 Comptonia peregrina 1.2
Abies balsamea -+-.1 Picea rubens +.1 Aster macrophyllus 1.3 Pterid-
ium aquilinum, var. latiusculum 4-1 Danthonia spicata 1.2
Dz.sp Hmlp Glp Elr
Betula populifolia 2.1 Betula papyrifera 2.1 Populus tremuloides 1.1
Vaccinium angustifolium 3.3 Rubus idaeus, var. strigosus 12) Salix
discolor 1.1. Pteridium aquilinum, var. latiusculum 1.3 Aralia hispida
1.1 Maianthemum canadense 1.4 Cornus canadensis 1.4 Festuca
capillata 2.2 Deschampsia flexuosa 1.2 Picea glauca +-.1 Picea rubens
1.1 Abies balsamea 1.1
Dsz Elr Glp Lp
Quercus rubra, var. borealis 2.1 Prunus pensylvanica 2.1 Betula papy-
rifera 2.1 Betula populifolia 1.1 Diervilla Lonicera 2.3 Spiraea lati-
folia 1.1 Vaccinium angustifolium 1.3 Pinus Strobus 1.1 Deschampsia
flexuosa 2.2 moss spp. 3.3
Dzp Hlp Glp Lp Ezr bp
Vaccinium angustifolium 2.3 Pyrus melanocarpa 1.2 Betula papyrifera
+.1 Betula populifolia +-.1 Prunus pensylvanica +-.1 Gaylussacia bac-
cata --.2 Deschampsia flexuosa 2.2 Danthonia spicata 1.2 Potentilla
tridentata 2.3 Solidago Randii +.1 Arenaria groenlandica +-.2 Aralia
hispida +-.1 Hypericum gentianoides +.3 Polytrichum sp. 1.2 lichen
spp. 2.3 Pinus rigida -+.1 Kalmia angustifolia --.2
Dm.sp Hml Gmp Emlr Lp
Betula lutea 3.1 Fraxinus americana 2.1 Acer rubrum 8.1 Acer saccha-
rum -++.1 Acer pensylvanicum 1.1 Tsuga canadensis --.1 Picea rubens
+.1 Abies balsamea +.1 Thuja occidentalis +.1 Osmunda_cin-
namomea 1.2 Dryopteris noveboracensis 3.4 Viola pallens 1.1 Athyrium
Filix-femina, var. Michauxii? +-.2 Osmunda Claytoniana 1.8 Aster um-
bellatus --.1 Onoclea sensibilis -+-.2 Dryopteris disjuncta +-.2 Aster
acuminatus 1.1 Aralia nudicaulis + .1 Calamagrostis canadensis + .3
Carex crinita 2.2 Brachyelytrum erectum, var. septentrionale 1.2 Carex
lurida --.1 Sphagnum sp. 2.3 Mnium sp, 1.2 moss spp. 1B
Ds.mlp Hmp Gmlr
Populus grandidentata 3.1 Betula papyrifera 3.1 Acer rubrum 2.1 Acer
saccharum 1.1 Fraxinus americana 1.1 Fagus grandifolia 1.1 Betula
lutea 1.1 Rubus idaeus, var. strigosus 2.1 Rubus allegheniensis 4.1
Solidago canadensis 1.1 Solidago rugosa 1.1 Solidago graminifolia 1.1
Aster lateriflorus 1.1 Aster macrophyllus 1.8 Lysimachia quadrifolia 1.4
Carex gracillima 1.2 Carex crinita 1.3
Dms Hmp Gmlr
Populus grandidentata 3.1 Betula papyrifera 3.1 Acer rubrum 2.1 Acer
saccharum 1.1 Fraxinus americana 1.1 Fagus grandifolia 1.1 Betula
lutea 1.1 Quercus rubra, var. borealis +.1 Rubus allegheniensis 4.1
Rubus idaecus, var. strigosus 2.1 Solidago canadensis 1,1 Solidago
rugosa 1.1 Solidago graminifolia 1.1 Aster lateriflorus 1.1 Aster macro-
263,
264,
266.
267.
VEGETATION OF Mount Desert Istanp, MAINE 871
phyllus 1.3 Lysimachia quadrifolia 1.4 Carex gracillima 1.2 Carex
crinita 1.8
Dsz.lp Emr Hmlp Glp
Betula papyrifera 2.1 Betula populifolia 2.1 Populus grandidentata 3.1
Acer rubrum 8.1 Vaccinium angustifolium 3.4 Comptonia peregrina 2.3
Pinus Strobus 1.1 Aster macrophyllus 1.2 Pteridium aquilinum, var.
latiusculum 2.1 Pinus resinosa 1.1 Poa compressa +4-.2 Deschampsia
flexuosa 1.2 Danthonia spicata 1.2
Gml Hmlp Dszp
Festuca capillata 4.5 Agropyron repens 2.4 Agrostis tenuis 2.2 Poa
pratensis 1.2 Festuca rubra 1.2 Phleum pratense +-.2 Aster novi-belgii
1.1 Potentilla tridentata +-.8 Rhinanthus Crista-galli 1.1 Rosa carolina?
3.1 Pteridium aquilinum, var. latiusculum +-.3 Vaccinium angustifolium
2.3 Spiraea latifolia 2.1 Alnus crispa, var. mollis 1.1 Populus alba 1.3
Prunus pensylvanica 1.3 Rhus typhina +-.3
Gml Hml Dsr
Festuca rubra 3.2 Dactylis glomerata 3.2 Poa pratensis 2.2 Phleum
pratense 2.2 Agropyron repens 2.2 Agrostis tenuis 2.2 Aster umbel-
latus 2.1 Vicia Cracca 2.1 Stellaria graminea 1.1 Hieracium aurantia-
cum 1.1 Hieracium pratense 1.1 Aster novi-belgii 1.1 Solidago juncea
+-.1 Solidago rugosa +-.1 Ranunculus acris -+-.1 Hemerocallis fulva
+.3 Rubus idaeus, var. strigosus 2.2 Spiraea latifolia +-.3
Ds Emr Gmlp Hmlp
Alnus crispa, var, mollis 38.1 Alnus rugosa, var. americana 2.1 Spiraea
latifolia 2.1 Rosa carolinaP 2.1 Amelanchier laevis 1.1 Betula papy-
rifera 1.1 Betula populifolia +-.1 Populus tremuloides 1.1 Picea glauca
-+-.1 Agropyron repens, Phleum pratense, Poa pratensis, Festuca rubra,
Agrostis tenuis, Vicia Cracca, Aster umbellatus, Hieracium pratense,
Chrysanthemum Leucanthemum, var. pinnatifidum, Stellaria graminea,
Hieracium aurantiacum
Dls Emlr Hmlp Glr
Betula papyrifera 3.1 Betula populifolia 3.1 Populus tremuloides 1.1
Spiraea latifolia 4.1. Amelanchier laevis +-.1 Picea glauca 1.1 Aster
umbellatus 3.1 Aralia nudicaulis 1.1 Aster macrophyllus +-.2 Maian-
themum canadense -+-.1 Calamagrostis canadensis 1.3
Not on map.
Emli.szp Dlp Hlp Lp
Picea rubens 4,1 Thuja occidentalis 3.1 Pinus Strobus 1.1 Abies bal-
samea +-.1 Juniperus communis, var. depressa +-.1 Acer rubrum 1.1
Populus grandidentata +.1 Betula papyrifera +.1 Betula populifolia
1.1 Vaccinium angustifolium 2.2 Gaylussacia baccata 2.2 Gaultheria
procumbens +-.1 Clintonia borealis +.1 Maianthemum canadense ++.1
Cornus canadensis 4-.1 Aster umbellatus +-.1 Aster macrophyllus -+-.3
Aster acuminatus +-.1 Trientalis borealis +-.1 Pteridium aquilinum,
var. latiusculum +-.3 Solidago squarrosaP +-.1 moss spp. 2.8 Cladonia
and other lichen spp. 1.2 Agrostis scabra --.1
270.
271,
272.
273,
274.
276.
277.
278.
Tur Universrry SCreENCE BULLETIN
Dz.slp Hlp Lp
Betula populifolia 5.1 Acer rubrum 2.1 Populus tremuloides 1.1 Amel-
anchier laevis +.1 Pteridium aquilinum, var. latiusculum 1.2 Solidago
rugosa +.1 Polytrichum sp. 2.3
Dsi.lzp Glp
Betula papyrifera 3.1 Betula populifolia 2.1 Acer pensylvanicum 1.1
Populus grandidentata --.1 Prunus pensylvanica +-.1 Diervilla Loni-
cera 1.3 Vaccinium angustifolium 1.8 Danthonia spicata 1.2 Des-
champsia flexuosa 1.2
Dml Emlp Hmlp
Acer rubrum 4.1 Populus grandidentata 2.1 Betula papyrifera 1.1 Acer
pensylvanicum 2.1 Picea rubens 2.1 Abies balsamea 2.1 Thuja occi-
dentalis 2.1 Maianthemum canadense -++.4 Clintonia borealis 4+-.1 As-
ter macrophyllus +.1 Pteridium aquilinum, var. latiusculum -++-.1 Aster
umbellatus 1.1 Solidago graminifolia 1.1 Solidago rugosa +.1 Dryop-
teris Phegopteris --.1 Osmunda cinnamomea +.1 Osmunda Claytoni-
ana +.1 Equisetum sylvaticum, forma multiramosum +.8
Eml Disp Hmlp L
Thuja occidentalis 5.1 Acer rubrum 1.1 Hex verticillata 2.1 Nemo-
panthus mucronata 2.1 Alnus rugosa, var. americana +.1 Fraxinus
americana -+.1 Osmunda cinnamomea 2.1 Osmunda regalis, var. spec-
tabilis 1.8 Sphagnum spp. 5.5
El Disp Hml Lp
Thuja occidentalis 4.1 Picea rubens 1.1 Abies balsamea 1.1 Acer ru-
brum 3.1 Betula lutea --.1 Betula papyrifera +-.1 lex verticillata 2.1
Nemopanthus mucronata 1.1 Osmunda cinnamomea 2.1 Osmunda
Claytoniana 2.1 Dryopteris noveboracensis 4.5 Sphagnum spp. 3.3
DI Hml Glr
Acer rubrum 4.1 Betula papyrifera 3.1 Populus tremuloides 1.1 Fraxi-
nus americana --.1 Acer pensylvanicum +.1 Aster macrophyllus 4.4
Aster umbellatus 1.1 Lysimachia quadrifolia +-.1 Dryopteris nove-
boracensis 1.3 Osmunda Claytoniana +.3 Pteridium aquilinum, var.
latiusculum -+.1 Poa compressa +-.1
Dls.mzp Hml Glr
Populus grandidentata 8.1 Betula papyrifera 2.1 Populus tremuloides 2.1
Betula populifolia 2.1 Fraxinus americana 1.1 Acer rubrum 1.1 Comp-
tonia peregrina 2.3 Diervilla Lonicera 1.8 Pteridium aquilinum, var.
latiusculum 3.3 Aster macrophyllus 3.8 Lysimachia quadrifolia 1.3
Festuca capillata 1.8
Dsz Hmlp Lp Gmr
Betula populifolia 4.1 Betula papyrifera 2.1 Populus grandidentata 1.1
Populus tremuloides 1.1 Acer rubrum 1.1 Comptonia peregrina 2.2
Rubus idaeus, var. strigosus 2.38 Rubus allegheniensis 1.3 Salix dis-
color +.1 Pteridium aquilinum, var. latiusculum 2.1 Aster acumi-
natus 1.1 Aster umbellatus -+.1 Aralia hispida +.1 Polytrichum spp.
2.8 Scirpus cyperinus, var. pelius +.1
Dms Elr Hmlp Lp
Fraxinus americana 2.1 Acer rubrum 2.1 Acer pensylvanicum 2.1
Populus grandidentata 2.1 Acer saccharum 1.1 Fagus grandifolia 1.1
279.
280.
288.
284,
285.
VEGETATION OF Mount Desert IsLAnp, MAINE 378
Betula lutea 1.1 Sambucus pubens +.1 Tsuga canadensis 1.1 Thuja
occidentalis 1.1 Brachyelytrum erectum, var. septentrionale -+-.1 Rubus
idacus, var. strigosus 8.4 Rubus allegheniensis 1.1 Lysimachia quadri-
folia 1.1 Aster acuminatus 1.1 Aster macrophyllus 1.8 Osmunda
Claytoniana 1.3 Viola pallens 1.1 Athyrium thelypteroides +-.1 moss
spp. 2.2
Dsz Hmlp
Betula papyrifera 3.1 Betula populifolia 3.1 Populus tremuloides 2.1
Populus grandidentata 2.1. Acer rubrum 1.1 Comptonia peregrina 8.2
Pteridium aquilinum, var. latiusculum 2.1 Aster macrophyllus 2.3 Aralia
hispida -+-.1 Solidago rugosa 1.1 Solidago squarrosa --.1
Dli.zp Elp Hmlp Lp
Betula papyrifera 4.1 Fraxinus americana +-.1 Populus tremuloides 1.1
Populus grandidentata 1.1 Acer rubrum 1.1 Acer pensylvanicum 1.1
Comptonia peregrina 2.1 Diervilla Lonicera 1.1 Rubus idaeus, var.
strigosus 1.2 Rubus allegheniensis +-.1 Thuja occidentalis 2.1 Pinus
resinosa +-.1 Pinus Strobus +-.1 Picea rubens 1.1 Aster umbellatus 1.1
Aralia hispida 1.1 Polytrichum sp. 2.2
Eml Dzp.sr Glr
Pinus rigida 3.1 Pinus Strobus 1.1 Thuja occidentalis +-.1 Betula
papyrifera 1.1 Betula populifolia 1.1 Populus grandidentata 1.1 Acer
pensylvanicum 1.1 Vaccinium angustifolium 1.3 Comptonia peregrina
1.1 Spiraea latifolia 1.8 Diervilla Lonicera 1.3 Danthonia spicata 1.2
Deschampsia flexuosa 1.2
Dszp Lp bp
Betula papyrifera 3.1 Betula populifolia 3.1 Populus tremuloides +-.1
Salix discolor +-.1 Prunus pensylvanica +-.1 Polytrichum sp. 28
Dis Hmlp
Acer rubrum 8.1 Betula populifolia 8.1 Alnus rugosa, var. americana 2.1
Salix discolor +-.1 Betula papyrifera 2.1 Spiraea latifolia 2.1 Populus
tremuloides 1.1 Amelanchier laevis -+-.1 Alnus crispa, var. mollis +-.1
Fraxinus americana +-.1 Ilex verticillata +-.1 Rubus hispidus 1.1 Os-
munda Claytoniana 1.1 Osmunda cinnamomea 1.1 Aster umbellatus 3.1
Aster macrophyllus 1.4 Dryopteris noveboracensis +-.4 Solidago rugosa
+.1
Gl Hlp Dzr
Festuca rubra 4.2. Agrostis tenuis 3.2 Anthoxanthum odoratum 2.2 Fes-
tuca capillata 1.2 Phleum pratense 1.2 Dactylis glomerata +-.2 Carex
scoparia +-.2 Luzula multiflora --.2 Potentilla simplex, var. calvescens
2.1 Solidago rugosa 2.1 Solidago juncea 1.1 Solidago graminifolia 1.1
Solidago nemoralis +-.1 Chrysanthemum Leucanthemum, var. pinnati-
fidum 1.1 Stellaria graminea 1.1 Vicia Cracca 1.1 Trifolium repens 1.1
Trifolium hybridum +-.1 Trifolium pratense +-.1 Aster umbellatus +1
Hieracium floribundum +-.1 Spiraea latifolia 1.1
Gml Hlp.mr Dsp.mr
Agrostis tenuis 3.2 Festuca rubra 2.2 Festuca capillata 2.2 Dactylis
glomerata 1.2 Phleum pratense 1.2 Anthoxanthum odoratum 1.2 Poa
pratensis 1.2 Carex conoideaP +-.2 Potentilla simplex, var, calvescens
2.1 Solidago rugosa 2.1 Solidago graminifolia 1.1 Solidago juncea 1.1
874
286.
287.
288.
289,
290.
Tue Universrry SCIENCE BULLETIN .
Rhinanthus Crista-galli 1.1 Aster umbellatus 1.1 Aster novi-belgii +.1
Vicia Cracca 1.1 Stellaria graminea -++.1 Chrysanthemum Leucanthe-
mum, var. pannatifidum +.1 Hypericum perforatum -++.1 Hieracium
pratense +.1 Ranunculus acris +-.1 Alnus crispa, var. mollis 1.1 Rosa
carolina? 1.1 Spiraea latifolia +.1'° Betula papyrifera +.1 Acer rubrum
+.1 Fraxinus americana +.1 Pyrus malus +-.1 Salix alba, var. vitellina
+.1
Dsjzp Hmlp Glmp
Populus tremuloides 3.1 Betula populifolia 3.1 Populus grandidentata
2.1 Betula papyrifera 2.1 Alnus rugosa, var. americana 3.1 Spiraea
latifolia 2.1 Fraxinus americana --.1 Rosa carolina? 1.1 Salix discolor
-+.1 Rubus allegheniensis +.1 Prunus pensylvanica +-.1 Aster umbel-
latus 2.1 Aster macrophyllus 1.3 Solidago rugosa 9.1 Solidago gramini-
folia -+.1 Onoclea sensibilis +-.1 Carex crinita +.1 Festuca rubra 2.2
Agrostis tenuis 1.2 Dactylis glomerata +-.2 Agropyron repens +..2 Fes-
tuca capillata +-.2 Poa pratensis +.2 Chrysanthemum Leucanthemum,
var. pinnatifidum -+.1 Vicia Cracca 1.1. Comptonia peregrina 2.1
Dmli Emlr Hmlp Gmlr
Acer rubrum 3.1 Betula papyrifera 2.1 Alnus rugosa, var. americana 2.1
Fraxinus americana +.1 Populus tremuloides --.1 Picea rubens +.1
Pinus Strobus -+-.1 Abies balsamea +.1 Diervilla Lonicera +-.8 Dryop-
teris noveboracensis 2.3 Pteridium aquilinum, var. Jatiusculum 2.1 Aster
macrophyllus 2.4 Aster umbellatus 2.1 Aster acuminatus +-.3. Osmunda
Claytoniana 1.2 Solidago rugosa -+.1 Solidago graminifolia 1.1 Hiera-
cium pratense +.2 Hieracium floribundum +-.2 Lysimachia quadrifolia
+.8 Carex crinita 1.2 Scirpus cyperinus, var. pelius +-.2 Agrostis
tenuis +.2
Dsz.lp.mr Hmlp Glp Lp
Betula populifolia 4.1 Populus tremuloides 3.1 Betula papyrifera 2.1
Prunus pensylvanica 1.1 Fraxinus americana --.1 Comptonia peregrina
91 Diervilla Lonicera +.1 Aralia hispida 1.1 Pteridium aquilinum,
var. latiusculum 1.1 Hieracium pratense 1.1 Hieracium floribundum 1.1
Solidago juncea +.1 Danthonia spicata 1.2 Carex pensylvanica? 1.2
Agrostis scabra -+.2 Polytrichum sp. 3.2
Gml Hmlp Dsp.lr
Festuca capillata 2.3 Poa pratensis 2.2 Agrostis tenuis 2.2 Agropyron
repens 1.2. Phleum pratense +.1 Solidago rugosa 9.1 Apocynum an-
drosaemifolium 1.5 Vicia Cracca 1.1 Pteridium aquilinum, var. latius-
culum 1.4 Lupinus polyphyllus +.1 Solidago juncea +-.1 Aster novi-
belgii +-.1 Rubus hispidus 2.1 Spiraea latifolia 2.1 Rosa carolina? 2.1
Celastrus scandens +.4 Betula populifolia +-.1 Alnus crispa, var. mollis
+.1 Acer rubrum -+.1 Fraxinus americana -++.1
Ds.mlp Hmlp
Acer rubrum 3.1 Spiraea latifolia 3.1 Betula papyrifera 2.1 Betula
populifolia 2.1 Populus grandidentata -+.1 Populus tremuloides 1.1
Alnus crispa, var. mollis 1.1 Salix discolor -+-.1 Alnus rugosa, var. amer-
icana 2.1 Comptonia peregrina 1.3 Tex verticillata +-.1 Rosa carolina?
--.1 Aster umbellatus 3.1 Pteridium aquilinum, var. latiusculum 1.3
Solidago rugosa 1.1 Agrostis tenuis +.2
291.
292,
293.
294,
296.
297,
298.
VEGETATION OF Mount Desert ISLAND, MAINE 875
Dlsi Ely Hml Glp
Populus grandidentata 8.1 Populus tremuloides 1.1 Acer rubrum 2.1
Fraxinus americana +-.1 Betula papyrifera 1.1 Betula populifolia 1.1
Comptonia peregrina 2.1 Diervilla Lonicera 1.2 Picea rubens 1.1
Aster macrophyllus 3.4 Pteridium aquilinum, var. latiusculum 2.3
Danthonia spicata 2.2 Polytrichum sp. 1.3
Dlsi Elp.mr Hmlp
Acer rubrum 3.1 Alnus crispa, var. mollis 1.3 Betula papyrifera 3.1
Betula populifolia 2.1 Fraxinus americana 2.1 Acer spicatum 2.1
Betula lutea +-.1 Thuja occidentalis 3.1 Abies balsamea 2.1 Picea
rubens 1.1 Tsuga canadensis +-.1 Aster macrophyllus 2.3 Aster acu-
minatus +.1 Solidago rugosa 1.1 Clintonia borealis --.1 Thalictrum
polygamum -+.3 Onoclea sensibilis -+-.3 Dryopteris spinulosa -++.1
Athyrium Filix-femina, var. Michauxii? --.1
Dsz.mlp Emlp Hmlp Glr Lp
Betula populifolia 3.1 Betula papyrifera 2.1 Acer rubrum 2.1 Fraxinus
americana 2.1 Populus tremuloides 2.1 Sambucus pubens +-.1 Salix
discolor +-.1 Rubus allegheniensis 1.2 Rubus idaeus, var. strigosus 1.2
Prunus pensylvanica +.1 Diervilla Lonicera 2.3 Picea rubens 2.1
Pinus Strobus +-.l Thuja occidentalis +-.1 Aster macrophyllus 1.3
Hieracium floribundum 1.2 Agrostis tenuis +-.2 Festuca capillata +-.2
moss spp. 3.3
Emli Dliiszr Hlp Lp
Picea rubens 8.1 Abies balsamea 2.1 Thuja occidentalis 1.1 Pinus
Strobus 1.1 Betula papyrifera 8.1 Acer rubrum 2.1 Acer pensylvani-
cum +.1 Populus grandidentata +-.1 Nemopanthus mucronata +.1
Viburnum cassinoides +-.1 Vaccinium angustifolium 1.3 Aster macro-
phyllus 1.3 Aralia nudicaulis +-.1. Maianthemum canadense 1.1 Pte-
ridium aquilinum, var. latiusculum 1.1 lichen spp. 2.3
Gml Hlp Lr
Poa pratensis 3.4 Festuca rubra 3.5 Agropyron repens 2.2 Festuca
capillata 2.2 Danthonia spicata 1.2 Phleum pratense +.2 Agrostis
tenuis 1.2 Hypericum perforatum 2.1 Stellaria graminea +-.4 Thymus
Serpyllum +-.3 Hieracium Pilosella +.3 Rumex Acetosella +-.1 Lechea
intermedia, var, juniperina +.1 Potentilla simplex, var. calvescens -+.1
moss spp. +.2
Gml Hip
Poa pratensis Festuca rubra Agrostis tenuis Phleum pratense Agro-
pyron repens Rumex Acetosella Hieracium pratense ete.
Gml Hmlp Lp
Festuca rubra Festuca capillata Phleum pratense Agropyron repens
Danthonia spicata Agrostis tenuis Luzula multiflora Hieracium spp.
Rhinanthus Crista-galli Rumex Acetosella Chrysanthemum Leucanthe-
mum, var. pinnatifidum Polytrichum spp.
Gml Hmlp
Agrostis tenuis 4.2 Anthoxanthum odoratum 3.2 Poa pratensis 2.2
Luzula multiflora 2.2 Phleum pratense 1.2 Festuca rubra 1.3 Dactylis
glomerata -+--.1 Carex scoparia +.1 Carex conoideaP +.1 Festuca
876
299.
800.
801.
302.
303.
804,
805.
Tur University ScrencE BULLETIN
capillata +.2 Panicum sp. +.1 Hieracium spp. 1.3 Stellaria graminea
1.1 Hieracium Pilosella -+.1 Solidago canadensis +-.1 Solidago juncea
+.1 Solidago rugosa +.1 Solidago graminifolia +-.1 Ranunculus acris
1.2 Achillea Millefolium +.2 Potentilla simplex, var. calvescens +.1
Dz.sr Hmlp Glp L bp
Betula papyrifera 3.1 Betula populifolia 8.1 Prunus pensylvanica 1.1
Salix discolor 1.1 Alnus crispa, var. mollis +.1 Acer rubrum +.1
Populus tremuloides +.1 Diervilla Lonicera 2.1 Spiraea latifolia -+.1
Aster umbellatus 2.1 Aralia hispida 1.1 Solidago Randii 1.1 Des-
champsia flexuosa 2.2 Polytrichum sp. 4.3
El Diszp Hlp Gmp
Thuja occidentalis 3.1 Picea rubens 2.1 Abies balsamea -++-.1 Betula
papyrifera 3.1 Betula populifolia 2.1 Acer pensylvanicum +1 Acer
rubrum +.1 Spiraea latifolia 1.1 Viburnum cassinoides +.1 Acer
spicatum +.1 Vaccinium angustifolium 2.3 Pyrus melanocarpa +,1
Aster umbellatus 2.1 Polypodium virginianum +-.1 Calamagrostis
canadensis 1.3
Dl.sr Gm Hm
Acer rubrum 5.1 Betula papyrifera 2.1 Betula lutea +.1 Acer pen-
sylvanicum 1.1 Rubus pubescens 1.4 Calamagrostis canadensis 5.5
Brachyelytrum erectum, var. septentrionale 1.3 Aster umbellatus 3.1
Aster acuminatus 2.1 Glyceria canadensis 2.3 Dryopteris novebora-
censis 2.3. Osmunda Claytoniana 2.1
Dl.szp Elr Him Gmp Lp
Acer rubrum 4.1 Betula papyrifera 3.1 Betula lutea 1.1 Acer pen-
sylvanicum 2.1 Populus tremuloides 2.1 Alnus crispa, var. mollis 1.1
Vaccinium angustifolium 1.3 Abies balsamea 1.1 Aster umbellatus 3.1
Pteridium aquilinum, var. latiusculum 8.3 Cornus canadensis 2.3 Dry-
opteris noveboracensis 1.3 Coptis groenlandica +.1 Aster acuminatus
4.1 Osmunda Claytoniana +.3 Calamagrostis canadensis 2.3 Brachy-
elytrum erectum, var. septentrionale +-.3 Polytrichum sp. 1.3
Dsz Elp Glp Bzr bp Hr
Betula populifolia 5.5 Vaccinium angustifolium 2.4 Gaylussacia bac-
cata 1.3 Nemopanthus mucronata +-.1 Pyrus melanocarpa 1.1 Picea
rubens 3.1 Danthonia spicata 2.2 Pinus Banksiana 1.1 Kalmia angus-
tifolia 1.2 Pteridium aquilinum, var. latiusculum 1.1
Dizi.sp Elp Bzp Lp Glr Hr
Betula papyrifera 4.1 Vaccinium angustifolium 8.8 Viburnum cassi-
noides +.1 Pyrus americana +.1 Pyrus melanocarpa -++-.1 Abies bal-
samea 3.1 Picea rubens 1.1 Kalmia angustifolia 8.3 lichen spp. 2.3
Potentilla tridentata 1.2 Danthonia spicata 1.2 Deschampsia flexuosa 1.2
Glp Elr Dszp Hp Bzr bp
Danthonia spicata 3.4 Deschampsia flexuosa 1.4 Festuca capillata 2.4
Agropyron repens 1.3 Agrostis scabra 1.2 Agrostis tenuis 1.2 Festuca
rubra +.2 Abies balsamea 1.1 Picea rubens -+-.1 Thuja occidentalis
--.1 Betula populifolia 2.1 Vaccinium angustifolium 2.3 Pyrus melano-
carpa 2.2 Spiraea latifolia 2.1 Pyrus americana 9.1 Alnus crispa, var.
mollis 2.1 Prunus pensylvanica 1.1 Potentilla tridentata 1.2 Chrysan-
306,
807.
808.
309.
$10,
312.
313.
314,
VEGETATION OF Mount Desrrr IsLAND, MAINE 377
themum Leucanthemum, var. pinnatifidum 2.2 Ambrosia artemisiifolia,
var. elatior 2.1 Potentilla simplex, var. calvescens -+.4 Solidago Randii
+.1 Arenaria groenlandica +.2 Achillea Millefolium +.2 Hypericum
perforatum -+.1 Kalmia angustifolia 2.3
Dszi Elp Glp bp
Betula populifolia 8.1 Vaccinium angustifolium 3.3 Pyrus melanocarpa
2.1 Spiraea latifolia 1.1 Kalmia angustifolia +-.1 Picea rubens 3.1
Thuja occidentalis 2.1. Abies balsamea 1.1 Danthonia spicata 2.2 Des-
champsia flexuosa 1.2 Potentilla tridentata +.1 Cladonia spp. +-.2
Eli Ds.zp Glp Lp bp Hmlr
Thuja occidentalis 3.1 Picea rubens 1.1 Betula populifolia 4.1 Betula
papyrifera +.1 Pyrus americana +.1 Vaccinium angustifolium 2.4
Viburnum cassinoides +-.1 Pyrus melanocarpa -+.1 Deschampsia flexu-
osa 2.2 Danthonia spicata 2.2 Calamagrostis canadensis +.3 Aster
umbellatus 2.1 Potentilla tridentata 1.2 Polytrichum sp. 2.2 Acer
rubrum -++.1
Gl Dszp Lp
Deschampsia flexuosa 3.3. Danthonia spicata 3.3. Betula populifolia 3.1
Spiraea latifolia 2.1. Pyrus melanocarpa 1.2 Diervilla Lonicera 1.1 Vac-
cinium angustifolium 2.4 Potentilla tridentata 4-.2 Cladonia spp. 2.2
Polytrichum sp. 2.2 Prunus pensylvanica +-.1
Dmli.zp Emlr Hmp Glp
Quercus rubra, var. borealis 3.1 Acer rubrum 2.1 Betula papyrifera 3.1
Spiraea latifolia 1.2 Comptonia peregrina 2.4 Vaccinium angustifolium
2.4 Pinus Strobus 1.1 Picea glauca +-.1 Pteridium aquilinum, var.
latiusculum 2.3 Aster umbellatus 1.1 Solidago spp. 1.1 Agrostis tenuis?
3.2
Em.Jp Dmlsp
Pinus Strobus 4.1 Picea rubens 1.1 Quercus rubra, var. borealis 3.1
Fagus grandifolia +-.1 Corylus cornuta +-.1
Dsz.lp Hlp Glr
Betula populifolia 4.1 Populus grandidentata 2.1 Acer rubrum 2.1
Quercus rubra, var. borealis 4-.1 Comptonia peregrina 8.3 Vaccinium
angustifolium 8.3 Acer pensylvanicum ++.1 Pteridium aquilinum, var.
latiusculum 8.4 Danthonia spicata 1.2
Dli.zp Elp Hp
Betula papyrifera 2.1 Betula populifolia 2.1 Acer rubrum 2.1 Quercus
rubra, var, borealis -++.1 Vaccinium angustifolium 1.8 Comptonia
peregrina +.2 Pinus Strobus 2.1 Picea rubens +.1 Pteridium aquili-
num, var, latiusculum 1.2 Aster umbellatus 1.3 Abies balsamea 1.1
Em.lp Dlp Hp
Pinus Strobus 4.1 Picea rubens 3.1 Abies balsamea 3.1 Acer rubrum
1.1 Betula populifolia 1.1 Aster umbellatus 1.1
Dml Elp Hmlp Glr
Quercus rubra, var. borealis 3.1 Acer rubrum 3.1 Fraxinus americana
1.1 Betula papyrifera 1.1 Betula populifolia 1.1 Betula Intea +-.1
Picea rubens 2.1 Pinus Strobus 1.1. Osmunda cinnamomea 2.2 Dry-
opteris spp. 1.1 Aster umbellatus 1.1 Carex crinita +-.3
815,
316.
SLT;
318.
819.
820.
821.
322.
323.
824.
Tue Universiry SCIENCE BULLETIN
Dm.l|si Elr
Quercus rubra, var. borealis 4.1 Acer rubrum 3.1 Acer pensylvanicum
2.1 Fagus grandifolia 1.1 Picea rubens 1.1
Dlimszp Emlp Hml Glp
Acer rubrum 4.1 Quercus rubra, var. borealis 1.1 Vaccinium angusti-
folium 2.8 Picea rubens 2.1 Picea glauca 2.1 Pinus Strobus 2.1
Pinus resinosa 1.1 Maianthemum canadense 3.4 Pteridium aquilinum,
var, latisculum 2.1 Aster umbellatus 8.1 Dryopteris noveboracensis 1.4
Osmunda cinnamomea 1.1 Carex spp. 1.3 Equisetum sylvaticum,
forma multiramosum +-.3 Abies balsamea 1.1
Dis.zp Elr Hlp Glp
Betula populifolia 4.1 Betula papyrifera 2.1 Populus grandidentata 2.1
Acer rubrum 1.1 Fraxinus americana +.1 Gaylussacia baccata 2.3
Spiraea latifolia 1.1 Pinus Strobus -+-.1 Picea rubens +.1 Picea
glauca +.1 Thuja occidentalis 4-.1 Comptonia peregrina 1.3 Aralia
hispida 1.1 Pteridium aquilinum, var. latiusculum -+-.3 Aster macro-
phyllus +.2 Danthonia spicata 1.2
Em.lp Dlzp
Picea rubens 4.1 Pinus Strobus 8.1 Pinus resinosa 2.1 Thuja occi-
dentalis 3.1 Betula papyrifera 1.1 Acer rubrum 1.1 Acer pensyl-
vanicum -+.1 Populus grandidentata +.1 Gaylussacia baccata +4-.3
Elimp Dip L
Picea rubens 5.1 Thuja occidentalis 2.1 Acer rubrum 1.1 Betula
papyrifera 1.1 moss and lichen spp. 4.4
Eli Nlp Dzp.sr Glp Lp
Thuja occidentalis 4.1 Larix laricina 2.1 Acer rubrum 1.1 Alnus crispa,
var. mollis +.1 Rhododendron canadense 1.1 Ilex verticillata +-.1
Osmunda regalis, var. spectabilis +-.2 Scirpus cyperinus, var. pelius 2.3
Carex spp. 2.8 Calamagrostis canadensis --.2 Sphagnum sp. 2.5
Dlsp Elp Hmp ‘
Betula papyrifera 4.1 Acer rubrum 3.1 Betula populifolia 1.1 Comp-
tonia peregrina -+.4 Alnus crispa, var. mollis 2.1 Pinus Strobus 2.1
Abies balsamea 2.1 Aster umbellatus 2.4 Populus grandidentata +-.1
Dml Elp.mr Hlp
Acer rubrum 8.1 Betula papyrifera 3.1 Betula lutea 4+-.1 Acer pensyl-
vanicum +.1 Abies balsamea 8.1 Picea rubens 2.1 Pinus Strobus 1.1
Maianthemum canadense 1.4 Aralia nudicaulis -+.1
DI Elp Hm
Betula papyrifera 4.1 Acer rubrum 8.1 Betula populifolia 1.1 Vi-
burnum cassinoides +-.1 Betula lutea -++-.1 Abies balsamea 8.1 Picea
rubens 1.1 Pinus Strobus -+.1 Aster umbellatus 4.4 Osmunda Clay-
toniana +.3 Equisetum sylvaticum, forma multiramosum +.4
Dms Emlp
Betula papyrifera 8.1 Acer rubrum 3.1 Fagus grandifolia 1.1 Betula
populifolia +.1 Acer saccharum ++-.1 Fraxinus americana -+.1 Acer
pensylvanicum 8.1 Picea rubens 3.1 Tsuga canadensis 1.1
re
825.
326.
327,
828.
329.
830.
332.
333.
334,
VEGETATION OF Mount Desert IsLanp, MAINE 379
Dis Emlp
Acer rubrum 3.1 Betula papyrifera 3.1 Betula populifolia 2.1 Acer
pensylvanicum 1.1 Picea rubens 2.1 Pinus Strobus 2.1 Pinus resinosa
Ll
DI Elp bp
Acer pensylvanicum 3.1 Betula papyrifera 8.1 Acer rubrum 8.1 Picea
rubens 8.1
Elp NIlp Disp Gm ‘Hmp L
Thuja occidentalis 4.1 Larix laricina 8.1. Acer rubrum 2.1 Alnus rugosa,
var. americana 2.1 Ilex verticillata 2.1 Calamagrostis canadensis 5.5
Osmunda cinnamomea 2.2 Sphagnum sp. 5.5
Esi Gmp Hmlp Dmsr
Abies balsamea 8.1 Picea rubens 8.1 Pinus Strobus 1.1 Carex scoparia
1.2 Carex lurida 1.4 Carex crinita 1.2 Carex sp. 1.2 Scirpus cyperinus,
var. pelius 1.2 Danthonia spicata +-.2 Aster umbellatus 2.1 Dryopteris
noveboracensis 2.4 Cornus canadensis 2.4 Rubus hispidus 1.4 Acer
rubrum 1.1
Dls.zp Elp.mr Glp
Betula papyrifera. 4.1 Acer rubrum 3.1 Betula populifolia 1.1 Populus
grandidentata 1.1 Viburnum cassinoides 1.1 Acer pensylvanicum 1.1
Vaccinium angustifolium 2.4 Abies balsamea 2.1 Picea rubens 2.1
Danthonia spicata 1.2 Deschampsia flexuosa +-.2
Dl.sp
Acer rubrum 4.1 Betula papyrifera 3.1 Fraxinus americana 2.1. Betula
populifolia 1.1 Acer pensylvanicum 1.1
Dlszp Elp Glp Hmlp Lp
Betula populifolia 4.1 Acer rubrum 8.1 Betula papyrifera 2.1 Acer
pensylvanicum 1.1 Vaccinium angustifolium 2.4 Gaylussacia baccata
+.3 Viburnum cassinoides +-.1 Picea rubens 2.1 Pinus Strobus +.1
Pinus resinosa +-.1 Thuja occidentalis 1.1 Abies balsamea +-.1 Dan-
thonia spicata 2.3 Deschampsia flexuosa 1.3 Pteridium aquilinum, var.
latiusculum 1.1 Aster umbellatus 1.1 Potentilla tridentata +-.3 Hyperi-
cum gentianoides +-.3 lichen spp. 1.8
Dml Emlp
Acer rubrum 4,1 Quercus rubra, var. borealis 2.1 Betula populifolia 2.1
Populus grandidentata 1.1 Fraxinus americana 1.1 Acer saccharum 1,1
Acer pensylvanicum +-.1 Picea rubens 2.1 Thuja occidentalis 2.1
Dlszp Elp Hlr Bzp Glp Lp
Betula populifolia 8.1 Acer rubrum 3.1 Betula papyrifera 2.1 Viburnum
cassinoides 1.1 Gaylussacia baccata 1.4 Pyrus melanocarpa 1.4 Acer
pensylvanicum +-.1 Prunus pensylvanica 4+-.1 Vaccinium angustifolium
24 Picea rubens 3.1 Thuja occidentalis 1.1 Abies balsamea +.1
Pteridium aquilinum, var, latiusculum 1.8 Kalmia angustifolia 2.4 Des-
champsia flexuosa 2.2 Danthonia spicata 2.2 lichen spp. 2.3
Eli Nmp Hmp L Dir
Thuja occidentalis 4.1 Larix laricina 2.1 Picea mariana 2.1 Picea
rubens +.1 Pinus Strobus +.1 Dryopteris noveboracensis +.3 Os-
380
336.
337.
338.
339.
340.
341.
842.
343.
844.
Tar Universrry Science BULLETIN
munda regalis, var. spectabilis +-.3 Osmunda cinnamomea 1.4. Sphag-
num sp. or spp. 4.5 Acer rubrum +-.1
Dmii.sp GI Hmlp
Acer rubrum 3.1 Quercus rubra, var. borealis 3.1 Betula papyrifera 2.1
Acer pensylvanicum 2.1 Festuca rubra 4.3 Poa compressa 2.2 Dan-
thonia spicata 2.2 Lysimachia quadrifolia +.3 Anaphalis margaritacea,
var. intercedens 2.3 Pteridium aquilinum, var. latiusculum 1.3 Aster
umbellatus 2.1
Dml.sp Glm Elr Hmr
Acer. rubrum 4:1 Betula’ papyrifera’ 3.1: Acer pensylvanicum 2.1
Brachyelytrum erectum, var. septentrionale 5.5 Picea rubens 1.1 Aster
umbellatus 1.1
Eml Dizp Glp
Picea rubens 5.1 Acer rubrum 2.1 Betula papyrifera 2.1 Acer pen-
sylvanicum +.1 Vaccinium angustifolium 3.8 Viburnum cassinoides
+.1 Deschampsia flexuosa 1.2 Danthonia spicata 1.2
Eml Dz.lp Lp Hr
Picea rubens 4.1 Thuja occidentalis 1.1 Pinus resinosa +-.1 Betula
papyrifera 8.1 Acer rubrum 2.1 Vaccinium angustifolium 4,4 Populus
grandidentata +-.1 Gaylussacia baccata -+.3 lichen spp. 1.3. Pteridium
aquilinum, var. latiusculum +-.3
Dmliszp Glp Hip
Quercus rubra, var, borealis 4.1 Acer rubrum 2,1 Betula papyrifera 2.1
Fagus grandifolia +.1 Acer pensylvanicum 2.1 Comptonia peregrina
2.3 Rubus idaeus, var. strigosus 1.3 Danthonia spicata 3.2 Pteridium
aquilinum, var. latiusculum 2.3
Eli Dip
Thuja occidentalis 4.1. Acer rubrum 8.1 Betula papyrifera 2.1
Dsz.lp Hmp
Acer rubrum 3.1 Quercus rubra, var. borealis 1.1 Betula populifolia 3.1
Betula papyrifera 2.1 Prunus pensylvanica 1.1 Comptonia peregrina
9.2. Rubus idaeus, var. strigosus 1.1 Solidago rugosa +.1 Solidago
graminifolia +-.1 Solidago canadensis +.1 Aster umbellatus 1.1 Pteri-
dium aquilinum, var, latiusculum 1.3
Dz Hip Glp
Vaccinium angustifolium 4.3 Comptonia peregrina 3.3 Betula populi-
folia 3.1 Pteridium aquilinum, var. latiusculum 2.1 Danthonia spicata
1.2
Dz.lr Elp Hmlp Lp
Betula populifolia 3.1 Comptonia peregrina 3.3 Pinus Strobus 1.1
Pinus resinosa -—-.1 Pteridium aquilinum, var. latiusculum 2.3 Poly-
trichum sp. 3.3
Eml Nip Ds.lr Gm Hmr L
Thuja occidentalis 4.1 Picea mariana 3.1 Larix laricina 3.1 Tlex
verticillata 3.1 Viburnum cassinoides 2.1 Amelanchier laevis 2.1 Alnus
crispa, var. mollis 1.1 Acer rubrum 1.1 Betula papyrifera +-.1 Cala-
magrostis cariadensis 5.5 Osmunda cinnamomea +.1 Sphagnum sp. 5.5
345,
346.
847,
348.
349,
358.
354,
VEGETATION OF Mount Desert IstANpD, MAINE 881
Dmliszp Emlp Hmp Lp
Quercus rubra, var. borealis 8.1 Acer rubrum 2.1 Betula populifolia 2.1
Betula papyrifera 1.1 Populus grandidentata +-.1 Comptonia_pere-
grina 1.2 Vaccinium angustifolium 2.3 Pinus Strobus 3.1 Abies bal-
samea 2.1 Picea rubens 1.1 Pteridium aquilinum, var. latiusculum 2.3
moss spp, 3.3
GI Dizp Elp
Festuca, capillata 3.3 Agrostis tenuis 8.8 Festuca rubra 3.3. Betula
papyrifera 2.1 Betula populifolia 1.1 Acer rubrum +.1 Spiraea lati-
folia 1.1 Gaylussacia baccata --.2 Comptonia peregrina 2.2 Vaccinium
angustifolium 8.3 Pinus Strobus 2.1
Dsz Hmp Bzp
Populus grandidentata 4.1 Betula populifolia 4.1 Quercus rubra, var.
borealis 3.1 Acer rubrum 1.1 Vaccinium angustifolium 3.3 Comptonia
peregrina 2.2. Gaylussacia baccata +-.1 Pteridium aquilinum, var. latius-
culum 4.1 Kalmia angustifolia 1.1
Dmsz Hmp
Quercus rubra, var. borealis 5.1 Acer rubrum 3.1 Betula populifolia 2.1
Betula papyrifera 1.1 Populus grandidentata +-.1 Gaylussacia bac-
cata 1.3 Comptonia peregrina 1.2 Vaccinium angustifolium 2.3. Pteri-
dium aquilinum, var. latisculum 8.1 Kalmia angustifolia -+-.1
EI Ds.lp Hmp Gmlp L
Thuja occidentalis 4.1 Abies balsamea 2.1 Acer rubrum 2.1 Alnus
crispa, var. mollis 2.1 Ilex verticillata 2.1 Viburnum cassinoides 2.1
Osmunda cinnamomea 4.1 Calamagrostis canadensis 3.3 Carex spp. 2.8
Sphagnum sp. 4.8
Dszilr Glp Hip
Quercus rubra, var. borealis 1.1 Vaccinium angustifolium 8.8 Comp-
tonia peregrina 8.2 Betula populifolia 4.1 Danthonia spicata 2.2
Agrostis tenuis 1.2. Poa sp. 1.2. Pteridium aquilinum, var. latiusculum 2.2
Dlzi.sp.mr Hmp Glr
Quercus rubra, var. borealis 8.1 Acer rubrum 2.1 Betula populifolia 3.1
Betula papyrifera 2.1 Fraxinus americana -+.1 Acer pensylvanicum 1.1
Comptonia peregtina 2.3 Rubus idaeus, var. strigosus 1.1 Aster um-
bellatus +-.3 Pteridium aquilinum, var. latiusculum 2.8 Agrostis tenuis?
+.2 Danthonia spicata +-.2
Dsz.mr Hmp
Betula populifolia 8.1 Betula papyrifera 2.1 Comptonia peregrina 8.1
Populus grandidentata 1.1 © Quercus rubra, var. borealis 1.1 Acer
rubrum -+.1 Rubus idaeus, var. strigosus 1.3 Aralia hispida 1.2 Pter-
idium aquilinum, var. latisculum 2.2
Eml Dip’ Lp
Abies balsamea 4.1 Picea rubens 2.1 Pinus Strobus --.1 Betula papy-
rifera 2.1 moss spp. 2.2
Dmi.lsp Gmp Hmlp Elr
Acer rubrum 4,1 Quercus rubra, var, borealis 1.1 Fraxinus americana
1.1 Acer pensylvanicum 1.1 Rubus idacus, var. strigosus 4+-.3 Betula
papyrifera +.1 Calamagrostis canadensis 2.4 Carex gracillima 2.4
855.
856.
857.
358.
359.
360.
861.
362.
363.
364.
865.
366.
Tur Universrry SCIENCE BULLETIN
Carex crinita 2.4 Equisetum sylvaticum, forma multiramosum +.3 Aster
umbellatus 2.1 Osmunda cinnamomea +.2 Dryopteris noveboracensis
1.3 Picea rubens +.1
Ds
Betula populifolia 4.1 Fagus grandifolia 2.1 Acer rubrum 1.1 Acer
pensylvanicum +.1 Aralia hispida +-.1 Rubus idaeus, var. strigosis +-.1
Elp.mr Nmlp Dmlp
Thuja occidentalis 3.1 Picea rubens 2.1 Pinus Strobus 1.1 Larix lari-
cina 3.1 Acer rubrum 2.1 Fraxinus americana 2.1 Betula papyrifera 1.1
DI El
Acer rubrum 2.1 Quercus rubra, var. borealis 2.1 Betula populifolia 3.1
Populus grandidentata 1.1 Picea rubens 5.1 Abies balsamea 1.1
Dl.sp Elp
Acer rubrum 3.1 Alnus rugosa, var. americana 2.1 Betula papyrifera
2.1 Betula populifolia 2.1 Prunus pensylvanica 1.1 Picea glauca 2.1
Pinus Strobus 2.1
Eml Dl
Picea rubens 3.1 Picea glauca 2.1 Pinus Strobus 2.1 Tsuga canadensis
+.1 Acer rubrum 3.1 Acer pensylvanicum 1.1 Betula papyrifera 2.1
Betula populifolia 1.1 Quercus rubra, var. borealis +-.1
El Dlr Hmp Glp Lp
Thuja occidentalis 5.1 Picea rubens +.1 Pinus Strobus -++-.1 Fraxinus
americana +.1 Osmunda cinnamomea 8.8 Carex spp. 2.2 Sphagnum
sp. 3.3. moss spp. 3.3
Dzi.sp Hmlp
Betula populifolia 4.1 Comptonia peregrina 8.2 Vaccinium angustifolium
2.3 Acer rubrum 1.1 Populus grandidentata 1.1 Aralia hispida +.1
Pteridium aquilinum, var. latiusculum 2.1
Eli
Pinus rigida 3.1 Thuja occidentalis 2.1
Diszp Elp Glp Hmlp
Betula populifolia 2.3 Betula papyrifera 1.1 Acer rubrum 1.1 Acer
pensylvanicum +.1 Vaccinium angustifolium 8.3 Viburnum cassinoides
--.1 Picea rubens 2.1 Deschampsia flexuosa 3.3 Calamagrostis cana-
densis +.3 Aster umbellatus 1.1 Pteridium aquilinum, var. latiusculum
11
Eli Dp bp -
Thuja occidentalis 4.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer
saccharum 1.1 Betula populifolia 1.1
Elp Dlp bp
Thuja occidentalis 3.1 Picea rubens 1.1 Betula populifolia 2.1 Betula
papyrifera 1.1
El Dip bp
Picea rubens 4.1 Thuja occidentalis 2.1 Betula populifolia 2.1 Betula
papyrifera 1.1 '
367,
368.
369.
370.
871.
378.
874,
VEGETATION OF Mount Desert IstANp, MAINE 883
Dm.sp Emlp Hmp Gmp
Fagus grandifolia 38.1 Acer saccharum 2.1 Betula lutea 2.1 Acer
rubrum 2.1 Acer pensylvanicum 2.1 Betula papyrifera 1.1 Fraxinus
americana +-.1 Tsuga canadensis 2.1 Picea rubens 1.1 Osmunda Clay-
toniana 1.8 Aster umbellatus +-.1 Carex crinita +.2 Brachyelytrum
erectum, var. septentrionale 1.8
Em Dzp Hmlp
Pinus Strobus 5.1 Betula populifolia 2.1 Vaccinium angustifolium 8.8
Comptonia peregrina 1.2 Rubus idaeus, var. strigosus 2.3 Pteridium
aquilinum, var. latiusculum 1.1 Cornus canadensis 2.1
Dzp Bzp Hlp Glp Lp Elr bp
Pyrus melanocarpa 3.3 Vaccinium angustifolium 2.3 Alnus crispa, var.
mollis 1.1 Kalmia angustifolia 1.3 Potentilla tridentata +-.1 Arenaria
groenlandica +-.3 Solidago Randii +-.1 Hypericum gentianoides +-.2
Deschampsia flexuosa 2.3 Cladonia spp. 2.3. Picea rubens 1.1
DI] Elr Gm Hmp
Acer rubrum 5.1 Betula lutea 1.1 Acer pensylvanicum 1.1 Picea
rubens 1.1 Abies balsamea 1.1 Calamagrostis canadensis 4.3 Brachy-
elytrum erectum, var. septentrionale 8.8 Aster umbellatus 2.1
Elp Dszp.lr Hlp Glp Lp bp
Picea rubens 3.1 Betula populifolia 3.1 Gaylussacia baccata 2.8 Vi-
burnum cassinoides 1.1 Prunus pensylvanica 1.1 Vaccinium angusti-
folium 2.8 Acer rubrum -+.1 Acer pensylvanicum +-.1 Pyrus melano-
carpa +.2 Aster umbellatus 1.2 Pteridium aquilinum, var. latius-
culum 1.1 Deschampsia flexuosa 2.2 Danthonia spicata -+-.2 moss
spp. 2.2 lichen spp. 2.2
Dmlsp Emlp.si Hlp
Acer rubrum 8.1 Betula papyrifera 8.1 Picea rubens 4.1 Abies bal-
samea 2.1 Pinus Strobus 1.1 Maianthemum canadense 2.1 Cornus
canadensis 1.1
DLmsp Elp.mr Hlp Lp
Betula papyrifera 8.1 Acer rubrum 8.1 Fagus grandifolia 1.1 Acer
pensylvanicum 2.1 Betula lutea +.1 Picea rubens 3.1 Abies bal-
samea -+.1 moss spp. 8.38 Cornus canadensis 1.1 Maianthemum
canadense 1.1 Pteridium aquilinum, var. latiusculum 1.3
Eli Dir.si Hmlp Gir L Nir
Thuja occidentalis 5.1 Larix laricina -+.1 | Nemopanthus mucronata 2.1
Ilex verticillata 2.1 Acer rubrum 1.1 Betula papyrifera +.1 Alnus
rugosa, var. americana 1.1 Osmunda regalis, var. spectabilis 2.8 Os-
munda Claytoniana 1.2 Osmunda cinnamomea 1.2 Cornus cana-
densis 2.1 Sarracenia purpurea --.2 Gaultheria procumbens -+-.1 Carex
spp. 1.2 Sphagnum sp. 4.4
Dszp Glp Hlp Elr Lp bp
Betula papyrifera 2.1 Betula populifolia 2.1 Gaylussacia baccata 2.3
Viburnum cassinoides -+-.1 Vaccinium angustifolium 2.8 Acer rubrum
1.1 Acer pensylvanicum 1.1 Amelanchier laevis +-.1 Pyrus melano-
876.
877.
878.
879.
380.
881.
382.
Tue University SCrENCE BULLETIN,
carpa +.2 Deschampsia flexuosa 2.3 Danthonia spicata 1.2 Hypericum
gentianoides 1.8 Pteridium aquilinum, var. latiusculum 1.1 Arenaria
groenlandica +.2 Potentilla tridentata +.2 Picea rubens 1.1 lichen
spp. 3.3
Em Dlp
Picea rubens 5.1 Acer rubrum 2.1 Betula papyrifera 1.1
Elp Dlszp Hmp Glr Bzp Lp
Picea rubens 3.1 Abies balsamea +.1 Vaccinium Vitis-Idaea, var.
minus +.8 Vaccinium angustifolium 3.3 Pteridium aquilinum, var.
latiusculum 1.1 Danthonia spicata 1.2 Acer rubrum 2.1 Betula papy-
rifera 1.1 Tlex verticillata 1.1 Nemopanthus mucronata 2.1 Rhodo-
dendron canadense +.1 Myrica Gale 2.2 Ledum groenlandicum +-.2
Kalmia angustifolia 1.2 Picea mariana +.1 Osmunda cinnamomea 1.2
Sphagnum sp. 2.3 lichen spp. 3.3
Elp Dlszp Glp Hmlp Lp bp
Picea rubens 1.1 Thuja occidentalis 1.1 Pinus Strobus +.1 Betula
populifolia 2.1 Betula papyrifera -+-.1 Acer rubrum 1.1 Gaylussacia
baccata 2.1 Viburnum cassinoides 1.1 Pyrus melanocarpa 2.3 Vac-
cinium angustifolium 2.8 Fraxinus americana 1.1 Danthonia spicata 2.2
Brachyelytrum erectum, var. septentrionale 2.8 Deschampsia flexuosa 2.2
Hypericum gentianoides 1.8 Aster umbellatus 2.4 Arenaria groen-
landica +-.3 Potentilla tridentata +.2 Cladonia spp. 1.3 lichen spp. 1.5
Emli Dmli.szp Hlp Glr Lp
Pinus Strobus 3.1 Picea rubens 3.1 Abies balsamea 1.1 Thuja occi-
dentalis --.1 Acer rubrum 3.1 Betula papyrifera 2.1 Acer pensy]-
vanicum 1.1 Gaylussacia baccata 2.3 Vaccinium angustifolium 2.2
Gaultheria procumbens +-.2 Maianthemum canadense 2.1 Pteridium
aquilinum, var. latiusculum 2.1 Danthonia spicata 1.2 lichen spp. 2.8
Emli Dmlzp Hlp Lp
Picea rubens 3.1 Abies balsamea 2.1 Thuja occidentalis 1.1 Acer
rubrum 2.1 Betula papyrifera 2.1 Acer pensylvanicum +-.1 Vaccinium
angustifolium 1.3 Cornus canadensis 1.1 Pteridium aquilinum, var.
latisculum 2.3 Maianthemum canadense 1.1 moss & lichen spp. 4.3
Em Dlzp Hlp Lp
Picea rubens 5.1 Pinus Strobus 1.1 Abies balsamea 1.1 Thuja occi-
dentalis +-.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer pensyl-
vanicum 1.1 Vaccinium angustifolium 1.3 Cornus canadensis 2,1
Maianthemum canadense 2.1 Pteridium aquilinum, var. latiusculum 1.1
moss spp. 3.3
Emli Dizp.sr Hlp Glr Bzr
Picea rubens 5.1 Abies balsamea 1.1 Picea glauca +.1 Betula papy-
rifera 2.1 Acer rubrum 2.1 Acer pensylvanicum -++.1 Gaylussacia bac-
cata 1.3 Vaccinium angustifolium 8.8 Vaccinium Vitis-Idaea, var.
minus +.3 Cornus canadensis 3.3 Maianthemum canadense 2.1 Pter-
idium aquilinum, var. latiusculum 1.3 Aster umbellatus +-.2 Danthonia
spicata +.2 Deschampsia flexuosa +.2 Kalmia angustifolia 1.8
383.
884.
885.
387.
388.
889.
890.
891.
VEGETATION OF Mount Desert IsLAND, MAINE 885
Emli Hip Glp
Picea glauca 5.1 Picea rubens 1.1 Pteridium aquilinum, var. latius-
culum 2.1 Aralia nudicaulis 2.1 Maianthemum canadense 1.1 Festuca
sp. 3.3 Pyrus melanocarpa +-.1
El.zp Dz.lsr Hmlp Lp Glr bp Bzp
Picea rubens 4.1 Pinus Strobus 1.1 Juniperus communis, var. de-
pressa 2.8 Thuja occidentalis -+-.1 Gaylussacia baccata 5.4 Vaccinium
angustifolium 8.8 Betula papyrifera 1.1 Acer rubrum 1.1 Viburnum
cassinoides 1.1 Pteridium aquilinum, var. latisculum 1.1 lichen spp.
(mostly Cladonia) 2.8 Danthonia spicata +-.2 Deschampsia flexuosa +-.2
Kalmia angustifolia 2.3
Dmi.lsp.zr Emlp Hlp Lp
Acer rubrum 4.1 Betula papyrifera 2.1 Acer spicatum -++.1 Acer
pensylvanicum +-.1 Vaccinium angustifolium 1.38 Picea rubens 3.1
Abies balsamea 1.1 Thuja occidentalis -+-.1 Cornus canadensis 1.1
Maianthemum canadense 1.1 Pteridium aquilinum, var. latiusculum 1.3
Dryopteris noveboracensis +-.3 moss spp. 2.3
Dsz Elp Nlp Hml Gmp Lp
Alnus rugosa, var. americana 1.1 Acer rubrum 1.1 Betula papyrifera 2.1
Myrica Gale 8.1 Chamaedaphne calyculata, var. angustifolia 2.1 Tex
verticillata 8.1 Thuja occidentalis 2.1 Larix laricina 1.1 Sarracenia
purpurea 2.2 Osmunda regalis, var. spectabilis 3.1 Hypericum vir-
ginicum, var. Fraseri --.1 Aster nemoralis 2.1 Solidago uliginosa +-.1
Carex rostrata 2.4 Carex lenticularis 2.4 Calamagrostis canadensis 1.2
Juncus canadensis -+-.2 Sphagnum sp. 2.3
Elimp Dlp
Picea rubens 4.1 Betula papyrifera 2.1 Acer rubrum 1.1
Dmlsi Elp Hmp
Acer rubrum 8.1 Quercus rubra, var. borealis 1.1 Betula papyrifera 2.1
Betula populifolia 2.1 Populus grandidentata 1.1 Acer pensylvani-
cum 1.1 Fagus grandifolia -++.1 Rubus allegheniensis 8.8 Picea
rubens 2.1 Pinus Strobus 1.1 Pteridium aquilinum, var. latiusculum 1.8
Aralia hispida 1.1 Trientalis borealis +-.1
Em
Picea rubens 5.1
Dz.sp Hmp
Betula papyrifera 2.1 Betula populifolia 2.1 Populus grandidentata 2.1
Prunus pensylvanica 1.1 Acer rubrum 1.1 Comptonia peregrina 2.3
Pteridium aquilinum, var. latiusculum 1.8 Aralia hispida 1.2 Solidago
canadensis -++.2
Dsz Lp
Betula papyrifera 4.1 Betula populifolia 3.1 Acer rubrum 2.1 Populus
grandidentata 1.1 Acer pensylvanicum 1.1 Salix discolor 1.1 Prunus
pensylvanica 1.1 Polytrichum sp. 3.3
Emp Dmzp.si Lp bp
Tsuga canadensis 2.1 Pinus Strobus 2.1 Picea rubens 2.1 Betula
lutea 2.1 Betula papyrifera 1.1 Sambucus pubens 2.1 Prunus pensyl-
394,
396.
397.
398.
399,
893.
395.
Tue Universiry SCIENCE BULLETIN
vanica 1.1 Acer spicatum 1.1 Rubus idaeus, var. strigosus 1.1 moss
spp. 2.8 Rubus allegheniensis --.1 Diervilla Lonicera 1.2
Dz Glp Hlp Lp bp
Betula populifolia 4.1 Betula papyrifera 1.1 Vaccinium angustifolium
2.8 Comptonia peregrina 2.2 Deschampsia flexuosa 1.2 Danthonia
spicata 1.2 Pteridium aquilinum, var. latiusculum 1.2 Potentilla triden-
tata +.1 Solidago sp. +.1 Polytrichum sp. 2.3
Eli Dzp
Pinus rigida 4.1 Betula populifolia 4.1 Vaccinium angustifolium 2.8
Gaylussacia baccata 1.3
Dlszp.mr Emlr Glp Hlr Lp
Betula populifolia 3.1, Betula papyrifera 2.1 Acer rubrum 2.1 Betula
lutea 1.1 Acer saccharum +-.1 Acer pensylvanicum 1.1 Viburnum cas-
sinoides 1.1 Prunus pensylvanica 1.1 Vaccinium angustifolium 2.8
Diervilla Lonicera 1.2 Gaylussacia baccata 1.2 Rubus allegheniensis
1.1 Rubus idaeus, var. strigosus 1.1 Picea rubens 1.1 Deschampsia
flexuosa 2.3 Danthonia spicata 1.2 Calamagrostis canadensis 1.3 Carex
crinita 1.2 Osmunda Claytoniana 1.2 Solidago canadensis 1.2 Poten-
tilla tridentata 1.2 Aster umbellatus --.1 Solidago Randii +.1 Aralia
hispida -++.1 moss spp. 2.2
Dlsp Elr Glp Hlp
Betula populifolia 3.1 Betula papyrifera 2.1 Acer saccharum 1.1 Fagus
grandifolia 2.1 Acer rubrum 1.1 Acer pensylvanicum 1.1 Populus
tremuloides 1.1 Picea rubens 2.1 Osmunda—Dryopteris—Aralia—
Viola—Aster—Hieracium—etc. ete.
Dlszp.mr Emlr Glp Hmlp
Betula papyrifera 2:1 Betula populifolia 2.1 Acer rubrum 2.1 Populus
grandidentata 1.1 Populus tremuloides 1.1 Quercus rubra, var. borealis
1.1 Acer pensylvanicum 1.1 Fraxinus americana 1.1 Prunus pensyl-
vanica 1.1 Salix discolor 1.1 Viburnum cassinoides 1.1 Spiraea latifolia
1.1 Comptonia peregrina 2.2 Diervilla Lonicera 1.2 Rubus allegheni-
ensis 1.1 Vaccinium angustifolium 1.3 Pinus Strobus 1.1 Danthonia
spicata 2.2 Agrostis tenuisP 1.2 Aster macrophyllus 1.3 Aster umbel-
latus 1.1 Solidago canadensis 1.1 Solidago graminifolia 1.1 Pteridium
aquilinum, var. latiusculum 1.2 Hieracium pratense 1.1
Dl.sp Elp Hlp Glp
Betula populifolia 4.1 Betula papyrifera 2.1 Populus grandidentata +-.1
Corylus cornuta 1.1 Picea rubens 2.1 Pinus Strobus 2.1 Abies balsamea
1.1 Thuja occidentalis 1.1 Hieracium pratense 2.1 Aster umbellatus
L.1 Pteridium aquilinum, var. latiusculum 1.1 Aster macrophyllus 1.1
Danthonia spicata 2.2
Dl.sp Elr Himlp Glp Lp
Betula papyrifera 3.1 Betula populifolia 3.1 Acer saccharum 1.1 Fagus
grandifolia 1.1 Acer pensylvanicum 2.1 Picea rubens 2.1 Aster umbel-
Jatus 1.1 Hieracium pratense 1.1 Solidago canadensis 1.1 Solidago
graminifolia 4+-.1 Danthonia spicata 1.2 Agrostis tenuis? 1.2 moss spp.
1.3
400.
401.
402.
408.
404,
405.
406.
407.
408.
409.
VEGETATION OF Mount Desert Istanp, MAINE 387
Emli Dmli.sp
Picea rubens 3.1 Tsuga canadensis 2.1 Thuja occidentalis 2.1 Fagus
grandifolia 2.1 Acer saccharum 2.1 Acer rubrum 2.1 Acer pensylvani-
cum 4.1 :
Dlsi El Hmlp
Betula papyrifera 8.1 Betula populifolia 3.1 Populus grandidentata 1.1
Acer saccharum 1.1 Acer rubrum 1.1 Acer pensylvanicum 2.1 Prunus
pensylvanica 1.1 Picea rubens 1.1 Pteridium aquilinum, var. latiusculum
9.1 Aralia hispida 1.1 Aster macrophyllus +-.1 Solidago Randii +-.1
Eml Dmlp Hip Lp
Abies balsamea 4.1 Picea rubens 2.1 Betula papyrifera 2.1 Betula
populifolia 1.1 Acer rubrum 1.1 Fraxinus americana +.1 Fagus grandi-
folia +.1 Betula lutea 1.1 Acer saccharum +-.1 Maianthemum cana-
dense 2.1 Cornus canadensis 2.1 moss spp. 4.4
Dz.sp.lr Hmlp Gmp Lp
Betula papyrifera 3.1 Betula populifolia 3.1 Populus tremuloides 1.1
Salix discolor 1.1 Acer rubrum 1.1 Acer pensylvanicum 1.1 Rubus
idaeus, var. strigosus 1.1 Aster umbellatus 2.1 Solidago canadensis 1.1
Solidago graminifolia 1.1 Solidago rugosa 1.1 Aralia hispida 1.1
Anaphalis margaritacea, var. intercedens 1.3 Scirpus cyperinus, var.
pelius 2.8 Carex crinita 1.2 Lycopodium sp. 3.3
Dsz.lr Hmp
Betula papyrifera 4.1 Betula populifolia 2.1 Populus grandidentata 1.1
Salix discolor 2.1 Rubus idaeus, var. strigosus 2.1 Rubus alleghenien-
sis 2.1 Solidago canadensis 1.1 Anaphalis margaritacea, var. intercedens
1.3
Dml Elp Gm Hmlp L
Acer rubrum 8.1 Fraxinus americana 8.1 Betula papyrifera 1.1 Betula
lutea 1.1 Picea rubens 2.1 Abies balsamea 2.1 Thuja occidentalis 1.1
Calamagrostis canadensis 4.4 Carex crinita 1.2 Dryopteris novebora-
censis 8.3. Onoclea sensibilis 1.2 Cornus canadensis 1.1 Viola pallens?
1.1 Sphagnum sp. 4.8
Dms Emr
Fagus grandifolia 2.1 Acer saccharum 2.1 Fraxinus americana Ki
Betula papyrifera 2.1 Betula populifolia 1.1 Betula lutea +-.1 Acer
rubrum 2.1 Acer pensylvanicum 3.1 Picea rubens 1.1
Eml Dlr Hlr Lp
Picea rubens 5.1 Betula papyrifera 1.1 Acer rubrum 1.1 Acer pensyl-
vanicum 1.1 Thuja occidentalis 1.1 Maianthemum canadense 1.1
Pteridium aquilinum, var. latiusculum +-.1 Aralia nudicaulis +-.1 moss
spp. 2.2
Dmls Elsp Hlr Lp
Betula lutea 3.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer sac-
charum 1.1 moss spp. 2.2 Fraxinus americana 1.1 Fagus grandifolia 1.1
Acer pensylyanicum 2.1 Picea rubens 2.1 Abies balsamea 1.1 Aralia
nudicaulis 1.1
Eml Dsr Lp
Picea rubens 5.1 Tsuga canadensis 1.1 Thuja occidentalis +-.1 Acer
pensylvanicum 1.1 Acer rubrum -++.1 moss spp. 8.8
412.
413.
414,
415.
416.
ALT,
418.
419,
410.
411.
Tue Universtry SCIENCE BULLETIN
Dzp Bzp Hlp Glp Elr Lp bp
Pyrus melanocarpa 3.3 Vaccinium angustifolium 2.3 Alnus crispa, var.
mollis +-.1 Betula populifolia +-.1 Kalmia angustifolia 2.3. Potentilla
tridentata 1.2 Arenaria groenlandica +-.2 Hypericum gentianoides +.8
Danthonia spicata 2.2 Deschampsia flexuosa 1.2 Picea rubens +.1
Cladonia spp. 3.8
Emli Dlszp Hmlp Glp Lr
Picea rubens 4.1 Betula papyrifera 2.1 Acer rubrum 2.1. Acer pensyl-
vanicum 2.1 Populus grandidentata +-.1 Vaccinium angustifolium 2.8
Pteridium aquilinum, var. latiusculum 3.1 Aralia nudicaulis 1.1 Dan-
thonia spicata 1.2 moss spp. 2.2 lichen spp. +.2
Dlsi Elp Glp bp
Betula papyrifera 3.1 Acer pensylvanicum 1.1 Quercus rubra, var.
borealis 1.1 Betula lutea +.1 Acer spicatum 1.1 Picea rubens 1.1
Deschampsia flexuosa 1.2 Polypodium virginianum +-.1
Dms Emlp Lr
Betula papyrifera 3.1 Acer rubrum 2.1 Acer pensylvanicum 2.1
Betula lutea 1.1 Fagus grandifolia +-.1 Picea rubens 3.1 Abies bal-
samea 2.1 moss spp. 1.2
Emli Disp Hlp Lp
Picea rubens 4.1 Pinus Strobus 2.1 Abies balsamea 1.1 Betula papy-
rifera 2.1 Acer rubrum 2.1 Acer pensylvanicum 2.1 Acer saccharum 1.1
Maianthemum canadense 1.1 Pteridium aquilinum, var. latiusculum -+-.1
moss spp. 1.2
Dml.sp Elp Hmlp
Betula papyrifera 3.1 Acer rubrum 8.1 Acer saccharum 1.1 Acer
pensylvanicum 1.1 Betula lutea 1.1 Alnus rugosa, var. americana 1.1
Abies balsamea 2.1 Picea rubens 1.1 Osmunda cinnamomea 1.2 Os-
munda Claytoniana 1.2 Dryopteris noveboracensis 1.3. Aster um-
bellatus 1.1
Emlp Dlszp Nmr Hmlp Bzr
Picea rubens 3.1 Abies balsamea 1.1 Picea mariana 1.1. Acer rub-
rum 3.1 Betula papyrifera 2.1 Vaccinium angustifolium 1.3 Nemo-
panthus mucronata 2.1 Ilex verticillata 2.1 Viburnum cassinoides 1.1
Spiraea latifolia 2.1) Rhododendron canadense 1.1 Kalmia angusti-
folia 1.2 Larix laricina 1.1 Osmunda cinnamomea 2.1
Dmli Elp.mr Hml Gmr Lp
Betula papyrifera 8.1 Acer rubrum 8.1 Fagus grandifolia 2.1 Betula
lutea 1.1 Picea rubens 2.1. Abies balsamea 1.1. Dryopteris novebora-
censis 2.38. Aster umbellatus 2.1. Cornus canadensis 1.1 Maianthemum
canadense 1.1 Solidago rugosa 1.1 Onoclea sensibilis --.2. Hieracium
pratense +.1 Carex crinita +.2 Agrostis tenuis +-.2 Polytrichum sp. 2.2
Dml.sp Elp
Betula papyrifera 3.1 Fagus grandifolia 2.1. Acer rubrum 2.1 Acer
pensylvanicum 1.1 Betula lutea +-.1 Picea rubens 3.1
Dml.sp Elp Hmlp Lp
Acer rubrum 3.1 Betula papyrifera 3.1 Fraxinus americana 1.1 Vi-
burnum cassinoides 1.1 Alnus rugosa, var. americana 1.1 Picea rubens
Le eee Te eee ee ee tee ee eee ee a eee ee ee a
420.
421.
422,
425,
426.
427,
428,
429.
VEGETATION OF Mount Desert Istanpd, MAINE 389
2.1 Abies balsamea 2.1 Osmunda Claytoniana 2.2 Pteridium aquilinum,
var, latiusculum: 2.4 Aster umbellatus 2.1 Aster macrophyllus 1.1
Maianthemum canadense 2.1 moss spp. 3.3
Dmls Emr
Betula papyrifera 3.1 Betula lutea 2.1. Fraxinus americana 2.1 Acer
pensylvanicum 2.1 Acer saccharum +-.1 Fagus grandifolia +-.1 Populus
grandidentata +-.1 Acer rubrum 3.1 Picea rubens -++.1
Ds Els
Betula populifolia 3.1 Betula papyrifera 2.1 Prunus pensylvanica 1.1
Thuja occidentalis 3.1
Dm.sp Elp
Betula papyrifera 3.1 Acer saccharum 2.1 Fagus grandifolia 2.1 Acer
pensylvanicum 2.1 Picea rubens 3.1
Dml.sp Emlp
Acer rubrum 8.1 Acer pensylvanicum 3.1 Fagus grandifolia 2.1 Betula
papyrifera 2.1 Betula lutea 1.1 Fraxinus americana 1.1 Picea rubens
2.1 Abies balsamea 1.1
Emli Dmlp.si
Picea rubens 4.1 Abies balsamea 1.1 Betula papyrifera 2.1 Fagus
grandifolia 1.1 Acer pensylvanicum 4. 1
Dmls Emlp
Betula papyrifera 3.1 Acer rubrum 8.1 Fraxinus americana 1.1 Acer
pensylvanicum 1.1 Fagus grandifolia +-.1 Picea rubens 2.1
Dli.sp Eli Hmlp
Acer rubrum 4.1 Betula papyrifera 1.1 Ilex verticillata 1.1 Nemopan-
thus mucronata 1.1 Viburnum cassinoides 1.1 Thuja occidentalis 4.1
Abies balsamea 3.1 Picea rubens +-.1 Aster umbellatus 2.1 Osmunda
Claytoniana 1.3 Aster macrophyllus 1.1 Dryopteris noveboracensis +-.3
Pteridium aquilinum, var. latiusculum 1.3
Dl.sp Eli Hmli
Acer rubrum 4.1 Betula papyrifera 1.1 Betula lutea 1.1 Nemopanthus
mucronata 1.1 . Ilex verticillata 1.1 Viburnum cassinoides 1.1 Thuja
occidentalis 4,1 Abies balsamea 3,1. Picea rubens 1.1 Osmunda Clay-
toniana 1.3 Aster umbellatus 2.1 Aster macrophyllus 1.1 Pteridium
aquilinum, var, latiusculum 1.3 Dryopteris noveboracensis +-.3
Eml Dlp Hip
Pinus Strobus 4.1 Picea rubens 2.1 Acer rubrum 1.1 Betula papyrifera
+.1 Cornus, canadensis 2.1 Maianthemum canadense 2.1 Pteridium
aquilinum, var. latiusculum 1.2
Dlszp Elp Hmlp Glp Bzr Lp bp
Betula populifolia 3.1 Betula papyrifera 2.1 Acer rubrum 2.1 Acer
pensylvanicum 2.1 Acer spicatum 1.1 Amelanchier laevis 1.1 Vi-
burnum cassinoides 1.1 Gaylussacia baccata 1.3 Vaccinium angusti-
folium 2.38 Pyrus melanocarpa 1.8 Picea rubens 2.1 Abies balsamea
1.1 Thuja occidentalis 1.1 Aster umbellatus 2.1 Pteridium aquilinum,
var. latiusculum 2.1 Aralia nudicaulis 1.1 Hypericum gentianoides +-.2
Potentilla tridentata +-.2 Danthonia spicata 1.2 Deschampsia flexuosa
1.2 Kalmia angustifolia 1.3 lichen spp. 2.8
430.
432.
433,
435,
436,
487,
438.
439.
Tue Universiry Science BULLETIN
Dl.sp
Fagus grandifolia 4.1 Betula papyrifera 2.1 Acer rubrum 1.1 Fraxinus
americana +.1 Acer pensylvanicum 8.1 Acer saccharum 1.1 Betula
lutea 1.1
Dmli.sp Emlp Hlr Lp
Betula papyrifera 2.1 Fagus grandifolia 2.1. Acer saccharum 1.1 Acer
rubrum 1.1 Acer pensylvanicum 2.1 Fraxinus americana +-.1_ Picea
rubens 3.1 Thuja occidentalis +-.1 Tsuga canadensis +-.1 Aralia nudi-
caulis 1.1 moss spp. 3.2
Dml Elp.mr
Fagus grandifolia 3.1 Betula papyrifera 2.1 Acer saccharum 2.1 Acer
rubrum 2.1 Fraxinus americana 1,1 Betula lutea 1.1 Acer pensylvani-
cum 2.1 Picea rubens 3.1 Abies balsamea 2.1 Thuja occidentalis 1.1
Eml Dlp Hmr Lp
Picea rubens 5.1 Thuja occidentalis 2.1 Betula papyrifera 2.1 Acer
rubrum 1,1 Acer saccharum +.1 Fraxinus americana +-.1 Betula lutea
+.1 Acer pensylvanicum 1.1 Osmunda Claytoniana 1.3 moss spp. 2.3
Dlszp Elp.mr Glp Hmlp Bzr Lp bp
Betula papyrifera 1.1 Betula populifolia 1.1 Acer rubrum 1.1 Vi-
burnum cassinoides 2.1 Spiraea latifolia 2.1 Vaccinium angustifolium
2.3 Diervilla Lonicera 1.2 Gaylussacia baccata 2.3 Picea rubens 2.1
Picea glauca 1.1 Deschampsia flexuosa 2.2 Danthonia spicata 2.2 Aster
umbellatus 2.1 Potentilla tridentata 1.2 Pteridium aquilinum, var.
latiusculum 2.38 Kalmia angustifolia 1.3 lichen spp. 2.3
Dlsi Emlp Hmlp Glp Lr bp
Acer rubrum 3.1 Betula papyrifera 8.1 Acer pensylvanicum 1.1 Vi-
burnum cassinoides 2.1 Prunus pensylvanica 1.1 Spiraea latifolia 1.1
Vaccinium angustifolium 2.4 Picea rubens 2.1 Abies balsamea 2.1
Aster umbellatus 1.1 Pteridium aquilinum, var. latiusculum 2.2 Cornus
canadensis 2.2 Maianthemum canadense 1.1 Aster acuminatus 1.1
Deschampsia flexuosa 1.2 Danthonia spicata 1.2 lichen spp. 1.8
Eml Dmlp Hlp Lp
Picea rubens 5.1 Acer rubrum 2.1 Betula papyrifera 2.1 Acer pensyl-
vanicum 2.1 Cornus canadensis 1.1 Maianthemum canadense 1.1
Clintonia borealis +-.1 Aster macrophyllus +-.1 Aralia nudicaulis +-.1
moss spp. 2.3
Dm.Jp Elimr HI
Acer rubrum 8.1 Betula papyrifera 3.1 Picea rubens 3.1 Abies bal-
samea 2.1 Maianthemum canadense 4.1 Clintonia borealis 2.1 Aralia
nudicaulis 1.1
Eml L
Picea rubens 5.1 Picea glauca 1.1 moss spp. 4.4
Dlsp Eli Hip Lp
Acer rubrum 4.1 Betula papyrifera 3.1 Acer pensylvanicum 1.1 Popu-
lus grandidentata 1.1 Fraxinus americana +-.1 Abies balsamea 4.1
Picea rubens 2.1 Maianthemum canadense 1.1 Pteridium aquilinum,
var. latiusculum 1.1 moss spp. 2.3
440.
441.
442,
Ae
444,
445,
446,
448.
449.
i
a
VEGETATION OF Mount Desert IsLAND, MAINE 391
Hm Gm Li Dsr
Solidago graminifolia 4.1 Onoclea sensibilis 1.1 Osmunda_ cinna-
momea 1.1 Osmunda Claytoniana 1.1 Dryopteris noveboracensis 1.3
Rubus hispida 2.1 Carex gracillima 8.1 Luzula multiflora 1.1 Scirpus
cyperinus, var, pelius 1.1 Carex crinita 1.1 Sphagnum spp. 4.4 Spiraea
tomentosa 1.1 Alnus rugosa, var. americana 1.1
Elmp Dlp Hlp Lp
Picea rubens 5.1 Pinus Strobus 1.1 Abies balsamea 8.1 Betula papy-
rifera 2.1 Acer rubrum 2.1 Maianthemum canadense 1.1 Pteridium
aquilinum, var. latiusculum -++.1 Aralia nudicaulis 1.1 Trientalis
borealis +-.1 moss spp. 2.8
Emli Dlisp Hlp Lp
Picea rubens 4.1 Pinus Strobus 1.1 Abies balsamea 1.1 Acer rubrum 3.1
Betula papyrifera 2.1 Acer pensylvanicum 1.1 Cornus canadensis 1.1
Maianthemum canadense 1.1 Pteridium aquilinum, var. latiusculum 1.1
Aralia nudicaulis 1.1. moss spp. 2.2
Dszp.lr Emlzr Glp Hmp Lp bp
Betula papyrifera 2.1 Acer rubrum 1.1 Gaylussacia baccata 1.3 Pyrus
melanocarpa +-.2 Vaccinium angustifolium 1.8 Picea rubens 1.1 Juni-
perus communis, var. depressa 1.3 Pinus rigida 1.1 lichen spp. 2.8
Deschampsia flexuosa 1.2 Danthonia spicata +-.2 Pteridium aquilinum,
var. latiusculum 1.3 Aralia hispida 1.1 Potentilla tridentata -++.2
Elp.zr Dzp Hlr Lp
Pinus rigida 4.1 Juniperus communis, var. depressa 1.8 Gaylussacia
baceata 1.8 Vaccinium angustifolium 1.8 Pyrus melanocarpa +.2
Aralia hispida 1.1 Potentilla tridentata +-.2 lichen spp. 2.8
Dlsp Eli Hlp Lp
Acer rubrum 4.1 Acer pensylvanicum 2.1 Betula papyrifera 1.1 Abies
balsamea 4.1 Picea rubens 1.1 Thuja occidentalis +-.1 Aster macro-
phyllus 8.3. Aralia nudicaulis 2.1 Cornus canadensis 2.1 Maianthemum
canadense 1.1 Pteridium aquilinum, var. latiusculum 1.1 moss spp. 2.8
Dml.sr El.mr Hlp Li
Acer rubrum 3.1 Betula papyrifera 3.1 Acer pensylvanicum 1.1 Thuja
occidentalis 4.1 Abies balsamea 2.1 Picea rubens 1.1 Trientalis
borealis 2.1 Cornus canadensis 2.1 moss spp. 4.4
Gm Dszp
Calamagrostis canadensis 4.5 Danthonia spicata 1.4 Juncus cana-
densis 1.8 Populus tremuloides 2.1 Betula populifolia 1.38 Spiraea
latifolia 1.8 Salix discolor +-.1
Emlp Nmlp Dlzp
Picea mariana 8.1 Abies balsamea ++-.1 Larix laricina 8.1 Betula
populifolia 1.1 Vaccinium angustifolium 3.8 Comptonia peregrina 2.3
Gl Hmp Dlsp
Poa pratensis 3.2 Phleum pratense 3.2 Agropyron repens 1.2 Agrostis
alba 1.4 Festuca rubra +.2 Hypericum perforatum 2.8 Solidago
canadensis 2.2 Betula populifolia 2.4 Pyrus malus and Pyrus communis
planted
452.
458.
454,
Tue Universiry SCIENCE BULLETIN
Dli Elp
Betula papyrifera 8.1 Acer rubrum 8.1 Picea rubens 1.1
Dz Glp Hlp Lp
Betula populifolia 5.1 Betula papyrifera 1.1 Fagus grandifolia 1.1
Prunus pensylvanica 1.1 Acer rubrum 1.1 Vaccinium angustifolium 1.3
Populus tremuloides ++-.1 Spiraea latifolia 1.1 Rubus idaeus, var.
strigosus 3.3 Rubus allegheniensis 8.8 Comptonia peregrina +.3
Danthonia spicata 1.8 Festuca capillata 1.2 Pteridium aquilinum, var.
latiusculum 1.1 Solidago canadensis 1.1 Solidago rugosa 1.1 Poly-
trichum sp. 3.3
Eml Nip
Thuja occidentalis 4.1 Pinus Strobus +.1 Picea mariana +.1 Larix
laricina 2.1
Dlisp Gm Hmp
Acer rubrum 4.1 Betula populifolia 1.1 Viburnum cassinoides 1.1
Nemopanthus mucronata 1.1 Alnus crispa, var. mollis 1.1 Pyrus ameri-
cana +.1 Aster umbellatus 3.1 Calamagrostis canadensis 3.4 Brachy-
elytrum erectum, var. septentrionale 1.3 Carex gracillima 2.2
El Dzp Bzp Hlp Glr Lp
Picea rubens 5.1 Betula populifolia 1.1 Betula papyrifera 1.1 Gay-
lussacia baccata 1.8 Vaccinium angustifolium 1.2 Kalmia angusti-
folia 8.8 Cornus canadensis 3.3 Danthonia spicata 1.2 Agrostis
scabra +.2 Cladonia spp. 2.8
2 2 344 19 451
229 14 Q 7
Ss 6) 12 170 188 9
=> if
Xe ao aad e\'e , 19 2 192 a0
e “> 168 / a6 346 186 ae 9 x 389 12
216 | (v 213 343 /oa 2
219 \. be 4 7 - 238 S 2
4 a
£18 pts 34) 169 ye 7 "29 ae oh ag 135
ms 42 334 8 “ Se 397 -
©) A 211 5 173 “al nd 338 393 ie am
© é 34 347 9
eases (Leos ‘s ™ “sg e. ae
204 351 449
$0; 61 349 177 24 = 30
208 \ 17) aes 126
Ny 207 153 © 394
e 345 348 2 249 oa 4
- 350 '
209 _— 135 roe] '22
7 “ 106 245 24 35
12 “6, 172 247
©. oe 352 So
> * . 361 154 391 235
: 353 \34 395 “ 246
“a sad ye 133 Wa
A ee ell nw) a
= o 4 43 34 35
ome 456 360 137 [3087 131 202 201
12 163 340 Stl (58 A 357 ais 136 450 127
310 I OR 64 250
355 25! 248
142 162 65 ‘Oo: © 82
339 i ual - 60 as) 2 258\\ , 5
45
139 335 354 % 63 198 299 237
m 312 145 335 <> 135 24
159 1. x) »
is f 2. Al 252
3/3 6 155 61
zg S xO 46 -
314 2 159 | oe 62 304/ 236 La 29
164 6 9, ad 259 447
— 313 : = ha “oe 305 \ 48 28
165 y 45
12 wd ie 3 70 26
©)
13943) 18 165 405 26!
> 399 580 27
12 338 z 453 %2 eal 444 28
¥ 4o 57 406 38 ;
119 » 45 3 -
95 \865 ue 59 200 °
139 Sf as 51 A44 v + .
161 » w 409 ) 404 301
117 337 93 365 39
2372 70
an 333 id
c
- 87 r wi 59 2 bk “~ 25 253
o 30 263 *
— 120 331 $3 42\ 271
4 430 12 £\
% 377 58 9 303
87 . Q a 277 ( 279 ~
25 86 8 $2 255
@) 286
C) ks 110 ae 420 74 asa( \22
% 160
- . ce 270 Pre, 256 ‘98
86 13 2?
376 410 - 291 wy \ 13
325 $9 75 72 87 2
8I iit 375 443 12
/ e 408 122 ey
x I A 422 59X 273 ary 288
es 85 324 432 f
378 407 36 293 290
96 7, 6
N2 oeree - an s2 66 os 7 289
116 6 8 ‘2 ia 7 yt \
!97 3 90 4\\ Fe 423 "1 439 275 12
2 re
8 89 2 7 429 76 1
"3 85 » 121
320 110 433 440 284 294
100 Z le .
318 S) 7\ 283
80 91 MY) 407 s 15
32) 419 67 7 265
3 138 424 | 45 69 266
323 12 264
109 12 43 96 295 ian
os 98 Qe an a4! 8 267
8 6
o 12 i2 as on 99 97 6a P al
106 20 107 427 47 76 .
% S 418
374 1 257,
107, 4ai4 99 426 - 9 441 A
3 373
tt 14 370 98 se Py
id
x 233 - 445
96 435 3
12 - 436
“ 101 234 437 I °
12 454 A.
> a
a
107 ” z= 94 48
|
10
105 So,
384
380
385 103 12 (@
296 77
sas =e 102
12 / »%
12 416 re) 79 VEGETATION OF SOUTHEASTERN
' es (3 ee 102
Sa 751 lo a8 MOUNT DESERT ISLA
“td i 2 L |
i 2.
wl * 383 MAINE
tH i 1:25,000 1953 A.W. KUCHLER
}
e778
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVIII, Pr. 1] December 20, 1956 [No. 5
Biological and Systematic Studies of Two Species of
Cheyletid Mites, with a Description of a New
Species (Acarina, Cheyletidae) *
BY
Rosert E. Beer and Davw T. DatLey
Asstracr: Studies of the developmental stages of two species of cheyletid
mites are reported in this paper. One of these species is new to science. In
addition there is included a discussion of the taxonomy and morphology of
these species as well as detailed remarks on rearing methods used.
Mites of the family Cheyletidae are segregated into nineteen genera, two
of which are represented by species discussed in this work. Cheyletus eruditus
(Schrank) a cosmopolitan species, has on numerous occasions been reported
in the literature especially regarding its predatory habits, yet its complete de-
velopmental history has been unrecorded prior to the publication of this paper.
The new species here described is a member of the genus Cheletophyes and
is the second North American species to be described in this genus.
INTRODUCTION
The objective of this study was twofold. First, an effort was
directed toward establishing the unequivocal identity, in all of its
developmental stages, of a cheyletid mite species that apparently
not only extends its range over a large area of the world, but quite
possibly may be one of the most beneficial species in the family be-
cause of its predatism on many important pests. Second, it was
hoped that some techniques developed in the rearing and observing
of predaceous mite species might be recorded for use by others
working with live cultures of predatory acariens.
The mite family Cheyletidae contains nineteen genera that are
distinguished largely on the basis of characters of the dorsal shields,
and on palpal and pretarsal structure and ornamentation, In the
genus Cheletophyes five species have been described from various
parts of the world, the species here concerned being the second
* Contribution No, 898 from the Department of Entomology, University of Kansas.
(393)
394 Tue Universitry SctENCE BULLETIN
from North America. C. hawaiiensis Baker is recorded from Kailua,
Oahu, Hawaii; C. marshalli Baker was described from specimens
taken at Imboden, Arkansas; C. phillippinensis Baker from Bataan,
Philippine Islands; C. semenovi Kuzin from Uzbekistan, U.S. S. R.;
C. vitzthumi Oudemans from Willomore, Cape Colony, Africa. To
the knowledge of the present authors there are no published ac-
counts of biological observations of any members of the genus.
The genus Cheyletus contains approximately thirty species, the
uncertainty of the exact complement of the genus being due to the
inclusion of some questionable species. Cheyletus eruditus
(Schrank) is apparently a cosmopolitan species, being recorded
from Europe, Asia, South America, Greenland and the United States.
The biology of C. eruditus has been investigated in America by
Ewing (1912) and in England by Newstead e¢ al. (1918). The
species of mite studied by Beck (1866), whose work was repeated
and confirmed by M’Intire (1869), had a life cycle very similar to
that of C. eruditus and may have been this species. Some account
was taken of this mite’s life stages in connection with biological
studies of predaceous mite species, Gause, et al. (1936), and on
studies of interrelations between phytophagous and predaceous
mites by Rodionoy (1940).
C. eruditus has frequently aroused the interest of entomologists
due to its usual role as a predator of arthropods associated with
stored grains and foods. Some tests and observations have been
made regarding its potential value in biological control of such
pests. Ewing (1912), Newstead et al. (1918), Siggaard (1920),
Redikortzev (1924), Rodionov (1987, 1940), Solomon (1946), and
others contributed papers wherein biological observations of this
type were recorded. Rennie and Harvey (1922-1923) state that
the species probably is a valuable predator of Tarsonemus aphis
Rennie, Gamasus sp., and Rhizoglyphus echinopus Fumouze and
Robin, when in situations where these species are destructive in bee
nests. The mite has been found in stocks of bees infested with
Acarapis woodi (Rennie), where, according to Homann (1983), it
was thought to be feeding on the stocks of “wet-mold” mites rather
than on the actual bee parasites.
Cheyletus eruditus (Schrank) has some medical and veterinary
history. Pierce (1921) states that otoacariasis is caused in man by
this species attacking the external auditory meatus and Riley and
Johannson (1938) state that the mite has been taken in a pus
discharge from the ear of man. An outbreak of dermatitis among
American troops in North Wales reported by Hill and Gordon
Srupies or Two Species or CHey.etip MrtEs 395
(1945) was thought to involve C. eruditus. When searching for
the cause, they discovered two species of rodent mites in the troops’
bedding and two additional species in the palliasses used by the
troops. C. eruditus was found only in the stockpile of straw used
for stuffing the palliasses. The authors point out that rodent mites
may have been the primary cause of this dermatitis in humans.
Pillers (1920) lists C. eruditus as an accidental parasite in the ears
of the domesticated rabbit and guinea pig in association with
mites of the families Acaridae and Parasitidae and also mentions
C. eruditus as the causal agent for a skin eruption in horses.
A detailed description of the adult female and developmental
stages of Cheyletus eruditus and of both sexes and the develop-
mental stages of Cheletophyes knowltoni are presented in a por-
tion of this paper that follows. Rearing and observation tech-
niques used in this study are also discussed in detail as an aid
to persons engaged in similar studies,
REARING METHODS
The majority of specimens obtained for this study were col-
lected in conventional berlese traps. Litter of straw and other
detritus on the earth floor of an abandoned barn in Lawrence, Kan-
sas was found to be heavily populated with the two species re-
ported upon in this paper. The trapping jar beneath the funnel
was prepared with a mixture of plaster of Paris and activated char-
coal, utilizing the recipe of Farrell and Wharton (1949), poured
to a depth of one inch. The plaster was kept moist during trap-
ping operations after the early discovery that both species were
extremely sensitive to desiccation. The cheyletids were removed
from the trap jars at frequent intervals and placed in rearing pens.
The stender dish type rearing pens, prepared in the manner
described by Lipovsky (1953a), were used only for mass cultur-
ing of the species being studied and for containing individual
specimens in the deutonymphal or adult stages. For individual
rearing of larvae and protonymphs, stender dishes having an out-
side diameter of 35 mm. and a depth of 18 mm. were used.
Preparation of a substrate within the rearing dish was accom-
plished as follows: The jar was filled with a mixture of plaster of
Paris and charcoal, mixed in the proportion of 500 to 60 respec-
tively by weight. The jar was filled to a depth of one fourth of
an inch, the mixture then wetted and allowed to set. While soft,
some of the mixture was plastered on the sides of the rearing pen
to form a thin, even film about the inside of the pen, terminating
396 Tur Universiry SCIENCE BULLETIN
about one eighth of an inch from the top. Charcoal was added
to the plaster of Paris to make the substrate more porous, so that
when water was added during culture of the mites, it would sink
immediately into the substrate and not leave a drop of moisture
which might drown the mites. As C. eruditus is quite pale in
color in all immature stages, the dark color given to the substrate
by the addition of charcoal made it much easier to find the mites.
The plaster-charcoal substrate provided a roughened surface
which is necessary in order that cheyletids may move freely and
easily after their prey, [noted by Kozulina (1940) ], and it also pro-
vided a method of keeping the relative humidity in the rearing
pen within tolerable limits. However, it was found that humidity
was not an important limiting factor in the adult stage, as C.
eruditus will tolerate a relative humidity of thirty to thirty-five
per cent for four to five days which confirmed observations of
Rodionov (1940). The adults of Cheletophyes knowltoni seemed
to be as resistant to desiccation as the adults of Cheyletus eruditus.
Before the substrate had hardened after being wetted in the
rearing pen, a narrow depression was scratched in the bottom of
the plaster to give the mites a hiding place. C. eruditus would not
feed nor remain quiet unless such a place of concealment was pro-
vided. This practice was early discontinued, however, as it was
possible for mite larvae and protonymphs to hide or molt under
overhanging lips of such depressions and escape detection. As a
substitute, a bit of paper or other foreign material was introduced
into the chamber which seemed to be quite adequate for the pur-
poses mentioned. To complete the rearing pen, a small amount of
vaseline was spread around the top of the rearing pen and then
the lid was pressed firmly into place and rotated until a firm and
air-tight “seat” was effected. The vaseline also served as a repellent
to Cheletophyes knowltoni. The rearing pen remained near labora-
tory temperature (approximately 80-85° F.) which was within the
tolerance limits for C. eruditus. Rodionov (1937) reports the mite
living at a temperature of 95° F. in laboratory culture.
In order to keep large cultures of predaceous mites in the labora-
tory, it was necessary also to rear large numbers of mites as a
source of food. Tyrophagus sp. (Acaridae) was tried and found
to be very useful for this purpose. These acarids have a high re-
productive potential, are easily kept in the mass culture pens men-
tioned above and need to be fed only once a week with a bit of
dried yeast. It was not necessary to add water to this type of cul-
ture.
eta ee
oe
Srupies or Two Species or CHEYLETID MrIrEs 897
As described by M’Intire (1869), acarids may occasionally at-
tack cheyletids and overpower them, resulting in the death of the
latter. This situation in culturing C. eruditus became a problem
only at the time of molting. At that time, if the rearing pen held a
number of semistarved acarids, regardless of the state of develop-
ment, they would attack the molting cheyletid and feed upon it,
usually resulting in the death of the cheyletid. Semistarved acarids
in any appreciable quantity were dangerous to Cheletophyes knowl-
toni, regardless of whether the cheyletid was active or molting and
regardless of the stage of development. Although the cheyletid was
much more agile, the acarids would surround and overpower the
mite, killing it in a short time. In one case a female C. knowltoni
was placed in an individual rearing pen with ten acarids in varying
stages of development and within two hours the cheyletid had been
overpowered and a number of the acarids were feeding upon her.
Within five hours the larger acarids had left the cheyletid and
acarid larvae were devouring what remained of the cheyletid with
the exception of legs and palps. In practice, when Cheletophyes
knowltoni was observed in the premolt condition, all acarids were
removed from the rearing pen.
Cheyletids of both species involved, cultured in individual rear-
ing pens, were fed once a day. A microneedle mounted in a glass
handle made a very serviceable instrument for the transferring of
individual acarids from stock culture to rearing pen.
For the keeping of individual records of the cheyletids reared,
ruled five-by-eight note cards were used. The plain side of the card
was ruled into squares for the daily entry of such information as
feeding and molting data. The ruled side of the record card was
kept for such information as to where the mite was obtained, dates
for stages in the life history, records of activities, etc. The cards
were kept in a convenient file box and were identified with numbers
corresponding to the numbers fastened to the lids of the rearing
pens.
Mites to be mounted were cleared in a commercial solution of
lactophenol, and if stained, a ten-percent solution of acid fuchsin
was used. The specimens were then mounted in polyvinyl alcohol
(PVA) containing lactophenol in the manner described by Lipov-
sky (1958b). Specimens were flattened to some degree in order
to project the legs satisfactorily.
Tue Universiry SCIENCE BULLETIN
DESCRIPTIONS AND BIOLOGICAL NOTES
Cheyletus eruditus (Schrank)
Acarus eruditus Schrank, 1781, Enumeratio insectorum Austriae indigenorum,
p. :
Eutarsus cancriformis Hessling, 1852, Illustr. Med. Zeit., vol. 1, p. 258. Oude-
mans, 1938, Tijdschr. voor Ent., vol. 81, p. Ixxv.
Cheyletus seminivorus Packard, 1878, Guide to the study of insects, (Henry
Holt and Co., New York), p. 665. (new synonymy). Ewing, 1909, Illinois
Univ. Studies, vol. 8, pp. 76-77; 1912, Jour. Econ, Ent., vol. 5, pp. 416-420.
Cheyletus ferox Banks, 1906, Proc. Ent. Soc. Washington, vol. 7, p. 134.
Cheyletus eburneus Hardy, 1933, (in Andre), Ann. des Epiphyt., vol. 19,
pp. 336, 344, 350.
Cheyletus eruditus (Schrank), Oudemans, 1903, Tijdschr. voor Ent., vol. 46
pp. 118-129; 1906, Mem. Soc. Zool. France, vol. 19, pp. 84-88. Newstead
and Duvall, 1918, Reports of the Grain Pests (War) Committee, no. 2,
pp. 14-16. Haarlgv, N., 1942, Medd. Grgnland, no. 1281, p. 171. Hughes,
1948, The mites associated with stored food products, Ministry of Agricul-
ture and Fisheries, London, pp. 102-105. Baker, 1949, Proc. U.S. Natl.
Mus., vol. 99, no. 3238, p. 278.
This is a robust, active mite, light yellow to orange in color as
a young adult, darkening to brown with age. The immature
forms are white to light yellow in color, the deutonymph becom-
ing light brown toward the end of the stadium. The female has
two dorsal plates while the immature stages have only the anterior
dorsal plate present. The dorsum, with the exception of the
gnathosoma, is finely striated where not covered by a plate. The
mite is widest at the region of the shoulder setae, tapering rear-
ward, Structural characteristics of the setae on the legs seem
to vary from specimen to specimen, from finely pilose setae to
simple setae. The degree of ornamentation for a given seta will
be stated only when it is seen to be constant throughout the
specimens examined.
Female: (Plate I, figures 1-4)—Length of body, 360-410,; in-
cluding rostrum, 527-637». Width of body 286-332». Palpus
stout; femur two-thirds as broad as long with convex outer margin
and nearly straight inner margin, the long dorsal seta situated
near mid-segment minutely and sparsely pilose, its length about
equal to length of segment, two small simple setae located ven-
trally on segment, one near inner basal angle, the other near outer
apical angle of segment; genu short, nearly three times as broad as
long, a long minutely pilose seta located dorsally near outer margin
of segment, its length exceeding width of segment; tibia twice as
broad at base as at apex, the inner margin with pronounced con-
cavity extending from basal third of segment to apex, the tarsus
adjoining tibia along posterior half of declivity of this inner tibial
margin, tibia bears a simple dorsal seta one and one-half times
Srupres or Two Species or CHeyLtetip Mires 899
as long as segment situated on inner margin near mid-segment,
and a strong slightly curved blunt claw with two stout protuber-
ances near its inner basal margin, the claw more than one and
one-half times as long as tibia and emanating, with its basal pro-
tuberances, from narrowed apex of tibia; tarsus small, as broad as
long, outer. tarsal comb borne at apex of segment, with thirteen
to fourteen teeth, these decreasing in size toward base, inner tar-
sal comb attached to inner basal margin of segment, about one-
half as long as outer comb, with fourteen to seventeen teeth with
largest teeth in center of comb, segment with two long simple
ventral setae; palpal coxa robust, as broad as long, with one simple
ventral seta near apex; trochanter very short, without setae. Ros-
trum simple, narrow, broadening rearward, with two dorsolateral
setae distally, two short ventrolateral setae subapically; peritreme
segments bent to form letter M, four to five segments lying between
spiracle and longitudinally directed limb. Dorsal propodosomal
shield rounded anteriorly, widening posteriorly, covering most of
propodosoma, with four pairs of narrow lanceolate pilose marginal
setae, three pairs at anterior angles, one pair at posterior angles.
One pair of lanceolate pilose setae on dorsum between propodo-
somal shield and hysterosomal shield. One pair of long lanceolate,
finely pilose, dorsolateral shoulder setae between legs If and IIL.
Hysterosomal shield trapezoidal, with rounded angles, shield slightly
longer than wide, wider anteriorly, smaller than propodosomal
shield, with three pairs of narrow lanceolate pilose marginal
setae, one pair at anterior angles, one pair toward rear, one pair at
posterior angles. One pair of lanceolate, pilose dorsal setae situ-
ated subterminally on opisthosoma. Venter of propodosoma with
two pairs .of submedian setae, first pair located mesad from coxae I,
second pair posteromesad from coxae II. Venter of hysterosoma
with two pairs of submedian setae, first pair mesad from coxae IV,
second pair posteromesad from coxae IV. Venter of opisthosoma
with two pairs of setae on either side of anterior extremity of vul-
vular opening, two pairs of lanceolate setae on folds of vulvular
opening. One pair of lanceolate, finely pilose setae on terminal
margin of opisthosoma; anal opening surrounded by four minute
stout curved bristles. Coxae I, IIT and IV, each with two setae;
coxa II with one seta; trochanters I, IT and IV, each with one seta;
trochanter III with two setae; femora I, II, IIT and IV, each with
two setae; genua I, II, III and IV, each with two finely pilose
setae; genu I with a small rod-shaped seta located dorsally at apex;
tibia I with a small rod-shaped seta and one pilose lanceolate
400 Tue Universiry SCIENCE BULLETIN
seta; tibiae I, II, III and IV, each with four unclothed lanceolate
setae; tarsus I with sensory seta about one-fourth as long as tarsus,
one simple guard seta, one-half as long as sensory seta; tarsi I, II,
III and IV, each with one finely pilose seta in center of segment,
two long simple setae at distal end of segment, terminal on tarsus I,
substerminal on tarsi II, III and IV, two small finely pilose setae
at distal end of segment flanked by two short simple setae; tarsi
I, II, 111 and IV, each terminate with an empodium flanked by two
small, recurved claws. Tarsus I, 107 long; tibia I, 71y long. Leg I,
416y, long; leg IV, 387y long.
Larva: (Plate II, figures 5-7)—The larva has been described and
figured by Newstead and Duvall (1918, pp. 13-14, figure 3). A re-
description follows: Length of body, 156-241; including rostrum,
202-307. Width of body, 94-158y. Palpus slender; femur about
one and one-half times as long as wide, with convex outer margin,
dorsal seta finely and sparsely pilose, about as long as segment;
genu with a dorsal pilose seta; tibia with two ventrolateral setae and
a dorsal seta at distal end of segment, claw slightly bent, no pro-
tuberances at base; tarsus with two subterminal, sicklelike, ventral
setae, outer tarsal comb with twelve teeth of equal size, inner tarsal
comb with ten teeth, outer tarsal comb about two times as long as
inner tarsal comb. Rostrum simple, narrow, broadening rearward,
with two dorsolateral setae distally, two short, ventrolateral setae
distally; peritreme segments bent to form letter M. Propodosomal
shield rounded anteriorly, widening posteriorly, covering most of
propodosoma, with four pairs of narrow, lanceolate, pilose, marginal
setae; one pair of dorsal, pilose setae near posterior angles of shield;
one pair of dorsolateral, lanceolate, shoulder setae. Dorsum of
hysterosoma with three transverse rows of pilose setae, first row
with two pairs, remaining rows with one pair each. Venter of
propodosoma with one pair of setae mesad from coxae I, Venter
of hysterosoma with two pairs of setae, one pair mesad from coxae
III, one pair laterad from anus. Legs subequal in size; coxa I with
one seta; coxae II and III bare; trochanters J, II and III bare;
femora I, II and III, each with two setae; genua I and III, each with
a simple seta; genu II with a pilose seta; tibia I with a short sensory
seta; tibiae I, II and III, each with four simple setae; tarsus I with
one guard seta one-half as long as tarsus, one sensory seta about as
long as guard seta, one finely pilose seta, two lanceolate terminal
setae unequal in length, two short terminal setae; tarsi II and III,
each with two subterminal setae, two short simple terminal setae,
Stupies oF Two Species or CHeyterm MITEs 401
two short terminal finely pilose setae; tarsi I, II and IT, each termi-
nate with an empodium flanked by two small, recurved claws.
Length of tarsus I, 50p; length of tibia I, 23p. Length of leg I, 154y:;
length of leg HI, 145p.
Protonymph: (Plate II, figures 8-10)—The protonymph has been
described and figured by Oudemans (1903, pp. 123-124, figure 84).
A redescription follows: Length of body, 277-298; including ros-
trum, 359-385p. Width of body, 192-214y. Palpus stout; femur
about as broad as long with convex lateral margin, segment with two
ventrolateral setae the outer seta pilose, the inner simple, and one
dorsal seta finely pilose and longer than segment; genual seta on
posterior margin of segment, finely pilose; tibia with two ventro-
lateral setae and one seta at distal end of segment; a strong claw
slightly bent, with two protuberances of equal size at base sur-
mounts segment; tarsus with outer tarsal comb having twelve to
fourteen teeth, teeth decreasing in size toward base, inner tarsal
comb with eight to ten teeth decreasing in size toward base; outer
tarsal comb about one and one-half times as long as inner tarsal
comb; coxa with a simple ventral seta near apex; trochanter very
small and unornamented. Rostrum simple, broadening posteriorly,
with two dorsolateral setae distally, two short, ventrolateral setae
distally, peritreme segments bent to form letter M, four to five seg-
ments lying between spiracle and longitudinally directed limb.
Propodosomal shield rounded anteriorly, widening posteriorly, with
five pairs of lanceolate pilose marginal setae, three pairs toward
anterior angles, two pairs toward posterior angles; one pair of
lanceolate, finely pilose, dorsolateral, shoulder setae between legs
If and III; four transverse rows of lanceolate, finely pilose setae on
dorsum of hysterosoma, each row with two pairs of setae. Venter
of propodosoma with two pairs of lanceolate setae, one pair mesad
from coxae I, second pair posteromesad from coxae II. Venter of
hysterosoma with six pairs of lanceolate setae, one pair mesad from
coxae IV, one pair posteromesad from coxae IV, anus flanked by
four pairs of setae; one pair of terminal pilose setae. Coxae I and
IH, each with two setae; coxa II with one seta; coxa IV bare; tro-
chanters I, II and IV bare, trochanter IIT with two setae; femora I,
I, IM and IV, each with two setae; genua I, II, III and IV, each
with two setae; tibia I with one short rodlike seta, two lanceolate
setae, three finely pilose setae; tibiae II, III and IV, each with four
finely pilose setae; tarsus I with one sensory seta about one-half as
long as segment, one simple, guard seta about one and one-half
14—3378
402, Tse Universtry SCIENCE BULLETIN :
times as long as sensory seta; tarsi I, II, IM and IV, each with one
finely pilose seta in center of segment, two long simple setae distally
on segment, two small pilose setae at distal end of segment flanked
by two short simple setae. Tarsus I, 58p. long; tibia I, 31y long.
Leg I, 222u, long; leg IV, 190y long.
Deutonymph: (Plate II, figures 11-13)—The deutonymph has
been described and figured by Oudemans (1903, pp. 124-125,
figure 88). A redescription follows: Length of body 240-321);
including rostrum, 356-449». Width of body 148-235p. Palpus
robust; coxa with a simple ventral seta near apex; femur longer
than wide, with convex lateral margin, segment with two ventro-
lateral setae, the inner simple, the outer seta finely pilose and
about as long as segment; genu with one ventrolateral, median
seta and one finely pilose seta on posterior margin of segment;
tibia with two ventrolateral setae, a dorsal tibial seta at distal end
of segment and a stout terminal claw slightly bent with two equal
protuberances at base; outer tarsal comb about two times as long
as inner tarsal comb, with eleven to thirteen teeth, these decreasing
in size toward base; inner tarsal comb with twelve to thirteen teeth,
decreasing in size toward base; tarsus with two subterminal ventral
sicklelike setae. Rostrum simple, narrow, broadening rearward,
with two dorsolateral setae distally, two short ventrolateral setae
distally; peritreme segments bent to form letter M, four to five seg-
ments lying between spiracle and longitudinally directed limb.
Propodosomal shield rounded anteriorly, widening posteriorly,
covering most of propodosoma dorsally, with four pairs of finely
pilose marginal setae, three pairs at anterior angles, one pair at
posterior angles, two pairs of submedian finely pilose setae toward
caudal margin of shield. One pair of long lanceolate finely pilose
dorsolateral shoulder setae between legs II and II laterad from
shield. Dorsum of hysterosoma with four transverse rows of finely
pilose setae, four setae in a row, lateral setae longer than median
setae, Venter of propodosoma with two pairs of setae, first pair
mesad from coxae I, second pair mesad from coxae IT. Venter
of hysterosoma with two pairs of submedian setae, first pair mesad
from coxae IV, second pair posteromesad from coxae IV; anal
opening flanked by four pairs of simple setae; one pair of pilose
setae laterad from anus; one pair of subterminal, pilose, ventro-
lateral setae at distal end of hysterosoma. Coxae I, HI and IV,
each with two finely pilose setae; coxa II with one pilose seta;
trochanters I, II and IV, each with two finely pilose setae; trochanter
III with one finely pilose seta; femora I, II and HI each with two
Srupizs or Two Species or CueyLerm MITES 403
finely pilose setae; femur IV with one pilose seta; genua I, I, II
and IV each with two finely pilose setae; genu I with a short rod-
shaped seta distally; tibiae I and II each with four finely pilose
setae; tibiae III and IV each with three finely pilose setae; tibiae I,
III and IV each with a simple, lanceolate seta; tibia I with a short,
rod-shaped seta distally; tarsi I, II, III and IV each with a pilose
seta in center of segment, two long, simple setae distally, two small,
pilose setae flanked distally by two short simple setae; tarsus I
with a sensory seta about one-half as long as segment, guard seta
simple, about one and one-half times as long as sensory seta; tarsi
I, I, III and IV, each terminates with an empodium flanked by two
small recurved claws. Tarsus I, 85y. long; tibia I, 47p long. Leg I,
310p long; leg IV, 280y, long.
Type locality: Austria.
Location of type: Unknown.
Type habitat: Unknown.
Distribution: The species has been recorded from Australia, Hol-
land, India, Portugal, Mexico, Scotland, Jugoslavia, England, Ger-
many, Japan, Colombia, Chile, Brasil, Greenland and the United
States, In the United States it has been taken from house sparrows
at Ithaca, New York, on Sciurus carolinensis at Thomasville, Georgia,
and from California, Minnesota, and Oregon in warehouses, in
grains. The description of the female and immature forms is based
on material taken in stable litter from cattle barns at Lawrence, Kan-
sas,
Cheyletus eruditus was found to be entirely predaceous, feeding
on almost any arthropod small or weak enough to be overpowered.
It has been observed feeding on tineid larvae one fourth of an inch
long, adult psoccids and various collembolans, to mention the larger
prey. The mite has not been observed feeding on spiders or Acarina
of the families Uropodidae, Oribatidae, or Bdellidae, even when
the cheyletid was in a state of semistarvation and these arachnids
abundant.
The mite was cannibalistic by nature. In mass cultures the
larger forms were often seen feeding on smaller active stages of the
same species, and nymphal forms were observed feeding on the
eggs of their own species. Adults kept in the same pen without a
sufficient food supply battled savagely with one another, the victor
feeding on the vanquished. The species would by preference, how-
ever, attack the smallest and weakest prey conveniently available.
This species usually would not feed unless it had a small pit or
404 Tue Universiry ScreNCE BULLETIN
cut on the substrate in which to hide, It would remain at the open-
ing of the depression with palps widespread, ready to seize any
prey which ventured near. Rarely would it dash from the pit to
catch a victim which did not pass close enough to be ‘seized from
ambush. As had been reported by Ewing (1912), the mite, if not
starved, often fed a bit and then released its victim which would
survive if too great a quantity of the victim’s blood had not been
removed, The present authors found that C. eruditus often would
feed a bit on five or six different acarid adults during the course of
an afternoon and all of these acarids usually survived the assault.
However, if the predator was starved, it might completely drain an
adult female acarid in as little as twenty minutes, leaving only an
empty, shriveled skin.
If no pit was provided, the mite would lie in ambush under a
piece of paper or any such concealment that was provided within
the pen. If no method of concealing itself was available, the mite
would usually wander about until it became stuck in the vaseline
at the top of the rearing pen. While wandering about looking for
a place of concealment the mite was quite timid, rushing wildly
from arthropods smaller than itself, no matter how defenseless they
might be.
Once entrenched in a hiding place, the cheyletid seemed able to
cope only with arthropods approaching it from the front. If in some
manner acarid larvae or other small and defenseless prey were in-
troduced to the rear of the pit, the cheyletid would, upon being
touched by the prey, jump forward or sideways. It was assumed
that it was searching for the source of the stimulation. Being un-
able to find the cause of stimulation to the front or sides, the chey-
letid would move from the hiding place and usually move to a new
location. On the rare occasions that it returned to the original spot,
the prey would be devoured if the cheyletid entered the pit head
first. However, if the mite backed into the pit and was again stimu-
lated by the prey which had remained in the pit, the same procedure
of searching forward and sideways followed by leaving the pit, was
repeated,
The mite would, if possible, grasp a leg of its prey with the strong
palps and having inserted its needlelike chelicerae, suck the body
fluids from the victim with powerful pumping movements of the
pharynx. In the case of acarids, the body fluids could be seen
through the transluscent body wall as they flowed down the leg into
the cheyletid. If a leg was not caught at the first thrust, the predator
would usually seize its victim by any part of the body, inserting its
Stupres oF Two Species or CHeyLerm Mrres 405
chelicerae in whatever portion of the anatomy that was readily avail-
able. Observations of great numbers of individual feedings seemed
to indicate that a definite preference was shown by the cheyletid
to grasp the victim by a leg and feed from this appendage.
It is doubted that the mite in any stage of the life history produced
venom or toxic saliva, This assumption rests on the fact that
arthropods attacked by C. eruditus often recovered from the attack
of the predator as discussed above, and also victims usually con-
tinued escape attempts, flexing the legs and body parts while being
fed upon, and if released shortly after the predator started to feed,
seemed to recover fully from the attack.
C. eruditus seemed sensitive to light, making immediate attempts
to escape from the more strongly illuminated parts of the pen. If
the mite was within a hiding spot, it would withdraw as far as pos-
sible into the dark. Mites running loose on the substrate would in-
variably attempt to reach a dark portion of the rearing pen. It is
possible that this was a reaction to the heat accompanying the illumi-
nation, but the same reaction was observed when the pen was illumi-
nated by daylight in a cool, humid atmosphere.
The mite usually molted in the section of the pen where it
normally lived. There was a premolt period which varied in dura-
tion for the different instars. During this premolt period the mite
refused all food and its movements were inclined to be sluggish,
The mite was not observed actually assuming the molting posi-
tion, but once settled, the position of the mite was prone on the sub-
strate, dorsum up, with legs I and II directed forward at full length.
This was a very typical position and was only assumed at the time
of molting.
This species was not observed in copulation. During the course
of this work, no males were taken from berlese funnels nor reared
from eggs. All agamic eggs from laboratory cultures produced
females.
Adult behavior. About forty-eight to sixty hours after emergence
of the female, egg-laying commenced. A maximum of twenty-two
eggs was laid over a period of seven days by one female. The most
frequent procedure was for the female to lay eleven eggs at a time.
The eggs were generally laid in a depression or under a bit of forei en
material and were deposited in a cluster, but not fastened together
nor fastened to the substrate. The eggs had no strands of silk or
other material covering them. The female gradually moved for-
ward as the eggs were deposited, making an oblong pile. Eleven
eggs could be laid in about twelve hours. After laying a quota of
406 Tur Universrry SCIENCE BULLETIN
eggs, the female would stand guard over them for about twenty-
four hours, palps spread in a defensive attitude, and would rush
forward and repeatedly pinch any object presented. No attempt
to feed was made during this period. Occasionally, while the eggs
were being deposited, an acarid would move up behind the female
and feed on the eggs as they were laid. So long as the acarid did
not touch the cheyletid and moved slowly, it was not molested. One
such acarid was prodded with a needle at such a time, the disturb-
ance being enough to greatly excite the cheyletid which rushed from
the pit, turned around and entered it head foremost and rapidly
drove the acarid from the pit with repeated pinchings of legs and
body of the intruder.
About seven days from the time of the first egg-laying, ten to
eleven more eggs usually were deposited as described and guarded
for the same length of time. The female would then live for some
time, actively feeding but on no occasion were more eggs laid,
Egg-laying may be extended over a period of thirty days, two to
three eggs being laid every other day or so until the full comple-
ment was reached, although some females reared have laid as few
as four eggs during their life span. When laid in such a manner, the
eggs may or may not be guarded by the female. Irregular egg
deposition seemed to be caused by the availability of food. If the
female was not fed after emergence, only two to five eggs might be
laid, and if still not fed, the female would die without laying any
additional eggs. However, if given ample food after the initial
starving, a further complement of eggs was usually laid. One female
laid four eggs and then fifty-seven days later deposited five addi-
tional eggs. During this entire period the female had ample food.
The length of the adult stadium varied widely in the laboratory,
running from twenty-five to eighty-three days. However, the last
few days before a normal death, the mite would be very feeble with
poor co-ordination and rarely took food. About a week before
death, drops of some clear exudate became visible, scattered over
the body of the mite. This exudate was quite viscous, debris of all
kinds adhering to it. The exudate seemed to be formed regardless
of the relative humidity of the rearing pen.
The eggs were semitransparent, shiny and oblong-oval in shape
being about two thirds as broad as long when laid. The eggs
averaged 125y, in length and 90p. in width.
Egg:—The eggs laid in small batches, that were usually left un-
protected after laying, were often preyed upon by mites of the
families Acaridae and Tydeidae. In the mass rearing jars used for
Srupies or Two Species or CieyLetip Mires 407
Cheyletus eruditus various immature forms of other predatory
_ mites were occasionally seen to feed on the eggs if there was a
dearth of other food.
At temperatures of 80-85°F., the eggs hatched after two to four
days, averaging three days. The process of hatching was fairly
rapid, taking about thirty minutes. The rostrum was forced through
the egg shell and moved from side to side a bit until an opening,
large enough to allow the palps to protrude, was formed. Legs I
were slowly projected from this opening, one at a time, in a continu-
ous movement. Simultaneously, the palps and rostrum were prying
on the substrate in such a manner as to urge the body forward. At
about the same time, the forepart of the body was forced through
the opening which continued to grow larger. The larvae seemed to
push the forepart of the body from the shell by repeatedly contract-
ing the body, which was followed by an active expansion and push-
ing forward of the body. Legs IJ eventually were released, one at
a time, and fastened to the substrate. At this time legs III could be
seen through the egg shell taking an active part in pushing the body
forward. The mite always paused for a few moments after the
anterior portion of the body was out of the shell and then would
ease the remainder of the body from the shell and move some dis-
tance away.
Occasionally a mite would be found with one or more legs pro-
jecting through small holes that the legs had torn in the egg shell.
Mites in this condition never emerged, although they have been
seen struggling to do so for as long as seventy-two hours before
death terminated the futile struggle to escape from the shell.
Larva:—The larva was able to move speedily about immediately
after leaving the egg but seemed inclined to stay close to the loca-
tion of hatching for five to eight hours; no food was taken during
this period, Later it moved about on the substrate until a suitable
hiding place was found. The larva would back into the hiding
place and remain quiet with the palps projecting. The mite then
would attack passing animals only if they made no threatening ad-
vances. The larva was extremely timid and would attack larvae
much smaller than themselves only after many timid advances and
retreats. The larva must have done much traveling about when
not under observation, because if a number of larvae were confined
in a rearing pen without ample food, within a short while there
would be only one or two individuals remaining, . . . undeni-
able evidence of their cannibalistic habits.
408 Tur Universiry SCIENCE BULLETIN
The larvae were fed on the larvae and first nymphal instars of
Tyrophagus sp., but usually in order to induce them to feed under
observation, it was necessary to place the prey in the pit or depres-
sion with the larval form of C. eruditus and then replace the lid of
the rearing pen for a short period, or else reduce the illumination
of the pen very greatly. When observed a short time later, the
larval predator usually would be feeding.
Six to twelve hours before molting the larva would take no food
and would be very sluggish in its movements. No larvae were
actually observed assuming the molting position although the
typical attitude of forward extension of legs I and II was assumed
before the ecdysis, The larval stadium at 80°-85° F. was from
three to six days with an average of four days.
Protonymph: The protonymph emerged after a twenty-four hour
molting period. The larval skin would split transversely on the
dorsum, the line of suture being between legs II and HI of the
larval skin. During emergence legs IV were not used, but were
held upward and rearward from the body with the genual and
tibial segments appressed to these same segments of the opposite
leg. The tarsi were free and could be seen flexing. The length of
time the rear legs were held in this position is not known, but
twelve hours after completing emergence they were in a normal
position and were functional in locomotion.
The protonymph often refused food for about five to eight hours
after emergence. Some tried to feed soon after emergence, but the
palps did not seem to be hardened enough to hold the prey and,
in one instance where the prey (Tyrophagus sp., larva) remained
quiet, the predator’s chelicerae did not seem to be able to pierce
the body wall of the prey.
Protonymphs were fed on larvae and small nymphal instars of
Tyrophagus sp. It was rare for a protonymph to attack prey its
own size or larger. If several of the same age were confined in
the same pen they were highly cannibalistic, however.
The protonymphal stadium at 80°-85° F, was from four to eight
days, averaging five days, the nymph molting within the depression
where it normally sought shelter. As in the larva, food was refused
for some time (six to twelve hours) before molting actually began.
The protonymph was not observed assuming the molting position,
though the characteristic stance was invariably taken before ecdysis.
Deutonymph: Immediately after emergence, the deutonymph
would usually refuse food. Occasionally, one would attempt to
Srupies or Two Species or CueyLerm Mites 409
feed soon after emergence but, as in other stages, the palps did not
seem strong enough to hold the prey.
Deutonymphs were fed Tyrophagus sp. nymphs and males, but
an attempt to feed them nymphs or females larger than themselves
was usually unsuccessful. Toward the end of the deutonymphal
stadium, the mite would often attack adult acarids or cheyletids, if
these adults were in a weakened condition.
From the remains of prey found scattered about in the rearing
pens, it is assumed that the deutonymphs moved about consider-
ably, when not under observation, in the darkened rearing cham-
bers. However, the deutonymph always seemed to return to the
pit or debris selected for a hiding place, as they would normally be
found in that location.
The deutonymph would molt in that portion of the rearing pen
selected as its retreat after going through a premolt period of six
to twelve hours. During this period it would take no food. The
mite was not seen assuming the molting position. The deuto-
nymphal stadium at 80°-85° F, was from eight to ten days, averaging
nine days.
Cheletophyes knowltoni, sp. nov.
Superficially this species resembles Cheletophyes hawaiiensis
Baker and C. marshalli Baker, but examination of the female will
show distinct differences, Cheletophyes knowltoni is unique in
having but one tooth on the palpal claw. The dorsal palpal
femoral setae are rodlike and spinose. The tarsus of leg I has a
rodlike sensory seta about one-half as long as the segment, with a
finely spinose guard seta twice as long as the sensory seta. There
are five spinose setae and one short peglike sensory seta on tibia I.
Observed under low magnification the dorsolateral body setae
appear as flattened whitish subclavate frondlike erect structures.
The mite is orange to pink in color as an adult, the immature
stages varying from yellow to orange, Legs I of all stages seem to
be used as tactile organs and are not used to support the body.
There is, in the adult stage, a single anterior and a single posterior
dorsal plate. The nymphal stages have one anterior and two small
posterior dorsal plates, while an anterior dorsal plate only is present
in the larval stage. There is a dorsal rostral shield present on all
active stages.
Female: (Plates III and IV, figures 14-16, 20 )—Length of body,
384u; including rostrum, 576; width of body, 287. Palpus robust;
410 Tue University SCIENCE BULLETIN
coxa robust, with one simple ventral seta near apex; trochanter
small, unornamented; femur about as broad as long, strongly con-
vex on outer margin, inner margin slightly concave, with two rodlike
dorsal spinose setae, one ventrolateral lanceolate spinose seta and
two ventrolateral simple setae; genu short and broad, unadorned;
tibia as broad at base as long, inner margin with deep declivity at
basal fourth of segment, width of segment decreasing from declivity
to apex, segment with one dorsolateral simple seta situated near
inner margin at base of segment, one long minutely pilose seta
located near mid-segment at apical third; palpal claw long curved
slender, acuminate at tip, with a single stout blunt tooth on inner
margin at basal third of claw; tarsus small knoblike, as broad as
long, with one apical short peglike seta on tubercle, two apical
whiplike setae; two apical tarsal combs, the outer comb with twenty-
three teeth, teeth decreasing in size toward base, comb stouter and
longer than inner comb, inner comb with forty-two teeth, decreasing
in size toward base. Rostrum with dorsal shield; forepart of shield
split from spiracle forwards, notched distally on inner margins,
longitudinally striated; rear part of shield extending to anterior
margin of propodosoma, longitudinally striated, fine punctations in
striations; peritreme simple, with elongate segments; rostrum with
a pair of ventrolateral simple setae distally and a pair of short
dorsolateral simple setae subapically. Single pair of eyes. Anterior
dorsal shield large, covering most of propodosoma, rounded anteri-
orly, widening posteriorly, with four pairs of long marginal sub-
clavate spinose setae, three pairs at anterior angles near eyes, one
pair at posterior angles; shield coarsely striated. A pair of long
subclavate spinose dorsolateral shoulder setae located slightly
laterad from posterolateral angles of anterior dorsal shield. Posterior
dorsal shield large, covering most of hysterosoma, broadest in front,
rounded posteriorly, about equal in length to anterior shield; shield
with five pairs of long marginal subclavate spinose seta, two
pairs at anterior angles, two pairs at posterior angles, one pair
located submesally at distal end of shield; shield coarsely striated;
a pair of stout subclavate spinose setae located terminally at apex
of opisthosoma behind shield. Dorsum of idiosoma with five pairs
of submedian staghorn setae, two pairs on anterior dorsal plate,
three pairs on posterior dorsal plate. Venter of propodosoma with
two pairs of submedian simple setae, first pair located anteromesad
from coxae I, second pair posteromesad from coxae II. Venter of
hysterosoma with three pairs of submedian simple setae located in
linear arrangement between coxae IV and anterior extremity of
|
|
Stupies or Two Species or CHEyLetip Mites 411
vulva; vulva flanked posteriorly by a pair of small triangular genital
plates each with rounded angles and bearing a pair of small simple
setae, Anal plate small and subtriangular, its broad anterior margin
slightly concave, posterior angle notched distally, with two short
pilose setae and a transverse row of four simple setae distributed
along anterior margin of plate, the medial setae shorter than lateral
setae, Coxa I with two simple ventral setae; trochanter I with one
lanceolate spinose seta; femur I with two spinose setae; genu I
with two long spinose setae; tibia I with three long spinose setae
subbasally, a very long (224y.) spinose seta distally, a long spinose
seta subdistally, and a peglike sensory seta distally; tarsus I as illus-
trated. Coxa II with one simple seta; trochanter IL with one
spinose seta; femur II with two spinose setae; genu II with two
spinose setae; tibia Hl with a simple sensory seta three times as long
as segment and three spinose setae; tarsus If with a short peglike
sensory seta at apical fourth of segment, a spinose seta at midseg-
ment, a pair of simple setae terminally and a pair of finely spinose
setae subterminally; tarsus II terminates in a fringed empodium
flanked by two short recurved claws. Coxa III with two setae, one
simple, one spinose; trochanter III with two spinose setae; femur
III with two spinose setae; genu III like genu JI; tibia III with
three spinose setae and a simple seta; tarsus ITI lacking short peg-
like sensory seta, otherwise like tarsus II. Coxa IV with two simple
setae; trochanter IV with one spinose seta; femur IV with one
spinose seta; genu IV like genu II; tibia IV like tibia III; tarsus IV
like tarsus III; pretarsal elements as on leg II. Length of tarsus I,
156; length of tibia I, 144y. Length of leg I, 700p; length of leg
IV, 560p..
Male: (Plate V, figures 21-23 )—Length of body, 3806p; including
rostrum, 500y; width of body, 230p. Palpus robust; femur broadly
rounded on outer margin, about as broad as long, with two simple
ventral setae, two stout spinose dorsal setae and one stout spinose
ventrolateral seta; genu without ornamentation; tibia with two
simple setae, one dorsal and one ventral, and one ventral spinose
seta; palpal claw short, curved, slender, with a single large blunt
tooth at basal third of claw; tarsus with one apical short peglike
seta on tubercle, two apical simple whiplike setae; two apical tarsal
combs, the outer comb stouter and larger than inner comb, outer
comb with twenty-two teeth decreasing in size toward base, inner
tarsal comb with forty teeth decreasing in size toward base; coxa
with one ventral simple seta near trochanter; trochanter small,
unornamented, Rostrum robust, its dorsal shield extending rear-
412 Tur University SCIENCE BULLETIN
hat
wards to propodosoma, longitudinally striated; anterior portion of
shield lightly punctate; rostrum with a pair of ventrolateral simple
setae distally and a pair of dorsolateral simple setae subapically;
peritreme simple, segments elongate, two to three segments be-
tween spiracle and longitudinally directed member. Single pair
of eyes located laterad from third setae of anterior shield. Dorsum
of idiosoma almost completely covered by the two large dorsal
plates; anterior plate about three-fifths as long as posterior plate,
both plates projecting laterad nearly to margins of body. Anterior
shield rounded anteriorly, posterior extremity slightly wider than
anterior extremity. Posterior shield with hind margin rounded-
truncate and anterior margin broadly and angularly truncate. Six
pairs of large subclavate spinose setae distributed on lateral mar-
gins of plates, three on anterolateral angles of anterior plate, one
on posterior angles of anterior plate, one at mid-length of posterior
plate, one on posterolateral angle of rear plate. Five pairs of large
subclavate spinose setae arranged in two longitudinal rows on
dorsal plates submesally, two on posterior third of anterior plate
the remaining three pairs on posterior plate. Two pairs of stout
subclavate spinose setae located dorsally on idiosoma but not on
dorsal plates, one of these slightly anterior to juncture of anterior
and posterior shields, the other borne subterminally on opisthosoma
in linear alignment with mesal setae of plates. Venter of idiosoma
with two longitudinal rows of small simple setae, the first pair just
mesad to coxae I, the second anteromesad from coxae HI, the
remaining three pairs posteromesad from coxae IV. Genital open-
ing dorsoterminal on opisthosoma with two pairs of small hooked
setae situated laterad from orifice and one pair of minute setae
located dorsally just anterior to orifice; two pairs of small dorsal
setae each borne on a minute plate, located between posterior
margin of the large posterior dorsal plate and the cluster of setae
around genital orifice. Aedeagus lancetlike. Coxa I with two
simple setae; trochanter I with one stout subclavate spinose scta;
femur I with two stout subclavate spinose setae; genu I with two
rodlike spinose setae, unequal in length; tibia I with three sub-
clavate spinose setae located on basal fourth of segment, one
sensory seta subdistally, about one fifth as long as segment, two
subclavate spinose setae subdistally, one very long (227); tarsus I
as illustrated. Coxa II with one simple seta; trochanter If with
one subclavate spinose seta; femur II with two setae, one sub-
clavate spinose, the other lanceolate finely pilose; genu I with
two subclavate spinose setae; tibia If with one long rodlike sensory
Stupies or Two Species or CueyLeTm MIteEs 4138
seta twice as long as segment, three long subclavate spinose
setae, one lanceolate finely pilose seta; tarsus II with a short
spinelike sensory seta and a subclavate spinose supporting seta
distally on segment, two subterminal simple setae, four subterminal
short finely pilose setae; tarsus II terminating in a fringed em-
podium flanked by two small recurved claws, Coxa III with one
subclavate, spinose seta, one lanceolate simple seta; trochanter
III with two pilose setae; femur III with two subclavate spinose
setae; genu IIT like genu II; tibia HI like tibia II; tarsus II
without short spinelike sensory seta, otherwise like tarsus II; pre-
tarsal elements as on leg Il. Coxa IV with two simple setae; tro-
chanter with one subclavate spinose seta; femur IV with one sub-
clavate spinose seta; genu IV like genu III; tibia IV like tibia III,
tarsus IV like tarsus III; pretarsal elements as on leg Il. Length
of tarsus I, 136y; length of tibia I, 146p. Length of leg I, 672p:;
length of leg IV, 508p..
Larva: (Plate IV, figures 17-19)—Length of body, 146; includ-
ing rostrum, 208%; width of body, 1832p. Palpus and rostrum
robust; palpal femur rounded laterally, about as wide as long, with
one dorsal clavate spinose seta; no dorsal genual seta; one dorsal
tibial seta finely pilose; palpal claw long, slender, slightly curved,
with a single tooth at basal fourth; tarsus with two apical whiplike
setae, two ventrolateral setae, the outer finely spinose, the inner
smooth, segment with two combs, outer tarsal comb longer and
stronger than inner comb, with twelve teeth adorning basal two
thirds of comb, teeth decreasing in size toward base, inner tarsal
comb with twenty-seven teeth distributed along entire length of
comb, teeth decreasing in size toward base. Rostrum with dorsal
plate extending posteriorly to anterior margin of propodosoma,
deeply emarginate anteriorly, finely punctate; peritreme simple
with elongate segments, two to three segments between spiracle
and longitudinally directed member; rostrum with one pair of
simple ventrolateral setae distally. Single pair of eyes. Anterior
dorsal plate large, .covering most of propodosoma, rounded an-
teriorly, widened posteriorly, with three pairs of marginal sub-
clavate spinose setae at anterior angles near eyes, one pair of
submedian subclavate spinose setae on posterior margin of plate;
entire plate finely punctate. One pair of dorsal subclavate spinose
setae near but not on posterolateral angles of propodosomal plate
and one pair of dorsolateral lanceolate spinose shoulder setae
situated slightly laterad from these setae. Dorsum of hysterosoma
414 Tue Universrry SCrENCE BULLETIN
with four transverse rows of subclavate spinose setae; first row with
four setae, second, third and fourth rows with two setae each;
dorsal body surface of entire hysterosoma striated. Venter of
propodosoma with one pair of submedian simple setae, situated
mesad from coxae I. Venter of hysterosoma with two pairs of
submedian simple setae, situated mesad from coxae IV and laterad
from anus; anus flanked by three pairs of simple setae. Coxa I
with one simple seta; trochanter I bare; femur I with two spinose
setae, unequal in length; genu I with one spinose seta, tibia I with
three spinose setae and one simple seta; tarsus I as illustrated.
Coxa II bare; trochanter II like trochanter I; femur II with two
setae, one spinose and one simple; genu I like genu I; tibia IT
like tibia I; tarsus II with one small peglike, sensory seta located
distally on segment, two simple subapical setae, four short, finely
pilose setae apically, segment terminated by a fringed empodium
flanked by two small recurved claws. Coxa III like coxa II; tro-
chanter III like trochanter I; femur III with one spinose seta, genu
III like genu I; tibia III like tibia I, tarsus Ill without short peg-
like sensory seta, otherwise like tarsus II. Length of tarsus I, 75p;
length of tibia I, 26y. Length of leg I, 205; length of leg TI, 128p.
Protonymph: (Plate VI, figures 24-26)—Length of body, 251y;
including rostrum, 342; width of body, 189». Palpus and rostrum
robust; palpal femur rounded on outer margin, one simple seta
and two dorsal rodlike spinose setae; genu bare, tibia with two
long and one short setae, the long seta minutely pilose; palpal claw
long, curved, slender, with a single prominent tooth at basal third;
tarsus small knoblike, with two apical whiplike setae, two apical
tarsal combs, the outer comb with fifteen teeth arranged on basal
half of comb, teeth decreasing in size toward base, outer comb
stouter and longer than inner comb, the inner comb with thirty-six
tecth distributed along entire length of comb, teeth decreasing
in size toward base; coxa with one simple, ventral seta. Rostrum
with dorsal shield extending rearward to anterior margin of pro-
podosoma, forepart broadly emarginate; shield finely punctate,
peritreme simple, segments elongate, two to three segments be-
tween spiracle and longitudinally directed limb; rostrum with a
pair of ventrolateral simple setae distally, one pair of shorter, dorso-
lateral simple setae subapically. Single pair of eyes. Anterior
dorsal plate large, covering most of propodosoma, rounded an-
teriorly, broadening posteriorly, finely punctate, with four pairs of
marginal subclavate spinose setae, three pairs at anterior angles
Stupies or Two Species or CaeyLetm Mrres 415
near eyes, one pair at posterior angles; one pair of submedian
subclavate spinose setae near posterior margin of plate. One
pair of long subclavate spinose, dorsolateral, shoulder setae be-
tween legs II and III. Dorsum of hysterosoma with five horizontal
rows of long subclavate spinose setae; anterior row with four
setae, each borne on a minute plate, second row with two setae,
each borne on a small distinct plate, third, fourth and fifth rows
with two setae each; all six of the small dorsal hysterosomal plates
finely punctate. Dorsum of hysterosoma finely striated in regions
not covered by plates. Venter of propodosoma with two pairs
of submedian simple setae, one situated mesad from coxae I,
the other posteromesad from coxae II. Venter of hysterosoma with
a pair of submedian simple setae, situated posteromesad from
coxae IV; terminal anal opening flanked by four pairs of short
simple setae. Coxa I with two simple setae; trochanter I bare;
femur I with one clavate spinose seta and one subclavate spinose
seta; genu I with two subclavate spinose setae; tibia I with four
subclavate spinose setae and one short peglike seta; tarsus I as
illustrated. Coxa II with one simple seta; trochanter II like tro-
chanter I; femur II with one subclavate spinose seta and one
simple seta; genu II like genu I; tibia II with three subclavate
spinose setae and one simple seta; tarsus IT with one finely spinose
seta distally, one short peglike sensory seta distally, two simple
setae and four finely pilose setae subterminally; tarsal segment
terminates in a fringed empodium flanked by two small recurved
claws. Coxa IIT with one lanceolate spinose seta and one simple
seta; trochanter III with one lanceolate, spinose seta; femur III
like femur IT; genu HI like genu I; tibia II like tibia Il; tarsus II
without short spinelike sensory seta, otherwise like tarsus I]. Coxa
IV bare; trochanter IV bare; femur IV with one subclavate spinose
seta; genu IV bare; tibia TV like tibia I]; tarsus TV without finely
spinose seta distally, otherwise like tarsus III. Length of tarsus I,
102p; length of tibia I, 71y. Length of leg 1, 190p; length of leg
IV, 175y.,
Deutonymph: (Plate VI, figures 27-29)—Length of body, 336,
including rostrum, 462; width of body, 238,. Palpus and rostrum
broad; palpal femur broadly rounded on outer margin, with two
dorsal rodlike spinose setae, one ventrolateral finely pilose seta
located near outer apical margin and two simple ventral setae;
ho genual setae; tibia with one long finely pilose seta and two
simple setae, the segment terminated by a long curved simple
416 Tur Universiry ScrENCE BULLETIN
claw having a single tooth at basal fourth; tarsus small, knoblike,
with one apical peglike seta on tubercle, two apical simple whip-
like setae and two apical combs, the outer comb with eighteen
teeth adorning basal two thirds of inner margin, teeth decreasing
in size toward base, this comb stouter and longer than inner
comb, the inner tarsal comb with forty-seven teeth distributed
along its entire inner margin, teeth decreasing in size toward base;
coxa with one ventral simple seta. Rostrum with dorsal shield
broadly emarginate at anterior margin, forepart striate-punctate,
rear part finely punctate; rostrum with two ventrolateral simple
setae distally, two dorsolateral simple setae subapically; peritreme
simple, segments elongate, two to three segments between spiracle
and longitudinally directed member. Single pair of eyes. Anterior
dorsal shield large, covering most of propodosoma, rounded an-
teriorly, widening posteriorly, with three pairs of long marginal
subclavate spinose setae, located at anterior angles of shield near
eyes, one pair of similar setae at posterior angles of shield, two
pairs of spinose, subclavate setae located submesally on posterior
half of shield; entire shield finely punctate. One pair of long
dorsolateral spinose shoulder setae between legs If and III. Dor-
sum of hysterosoma with five horizontal rows of spinose setae;
first row with four setae, each borne on a minute plate; second
row with four setae, two borne on each of the two moderately
large and widely separated dorsal plates; third row with four setae,
each borne on a minute plate; fourth row with four setae; fifth
row consisting of two setae situated terminally on opisthosoma;
all dorsal hysterosomal plates are finely punctate with intervening
integument finely striate. Venter of propodosoma with two pairs
of submedian simple setae, one pair situated mesad from coxae I,
the other pair posteromesad from coxae II. Venter of hystero-
soma with three pairs of submedian simple setae one pair mesad
from coxae IV, the other two pairs posteromesad from coxae IV;
terminal anal opening flanked by four pairs of short stout setae.
Coxa I with two simple setae; trochanter I with one spinose seta;
femur I with two spinose setae; genu I with two spinose setae;
tibia I with three spinose setae basally, one very long, one
short spinose setae subapically, one very short, peglike, sensory
seta situated subapically; tarsus I as illustrated. Coxa II with one
simple seta; trochanter II like trochanter I; femur IT with two
spinose setae; genu II like genu I; tibia If with three spinose
setae, one simple seta; tarsus II with one short peglike sensory
Srupies oF Two Species or CrHrytetm MITES 417
seta, one lanceolate spinose seta, and one short simple seta located
distally on segment; two long simple setae located subterminally,
one short sensory peg and one long, pilose seta located at about
mid-segment; tarsal segment terminates in a fringed empodium
flanked by two small recurved claws. Coxa III with one spinose
seta and one lanceolate simple seta; trochanter II with two spinose
setae; femur III with two spinose setae; genu III like genu I;
tibia III like tibia Ul; tarsus UI without short peglike sensory
seta, otherwise like tarsus II; pretarsal elements as on leg Il. Coxa
IV with two simple setae; trochanter IV with one spinose seta;
femur IV with one spinose seta; genu IV like genu I; tibia IV like
tibia II; tarsus IV like tarsus III; pretarsus as on leg Il. Length
of tarsus I, 110y; length of tibia I, 802. Length of leg I, 4583p;
length of leg IV, 414y.
Holotype: Female, Lawrence, Kansas, August 24, 1951, D. T.
Dailey, from litter in cattle barn.
Allotype: Male, same data as holotype.
Paratypes: Larva (morphotype), Lawrence, Kansas, August 2,
1951, D. T. Dailey, reared in laboratory culture; protonymph
(morphotype), Lawrence, Kansas, August 26, 1951, D. T. Dailey,
reared in laboratory culture; deutonymph (morphotype), same data
as protonymph; fifty female and fourty-six male paratypes, all from
Lawrence, Kansas.
Location of types: Holotype, allotype, morphotypes, forty-two
female paratypes and thirty-eight male paratypes deposited in the
Snow Entomological Museum, University of Kansas. Eight male
and eight female paratypes deposited in the U. S$, National Museum,
Washington, D. C.
Type habitat: Most of the specimens of the type series were col-
lected from straw litter taken on the ground in an abandoned cow
barn in Lawrence, Kansas and removed from the litter by the berlese
trap method or were reared in the laboratory from specimens col-
lected in the field at the locality mentioned. Some specimens in
the paratype series were collected from one year old, rotted hay
taken from a field in Lawrence, Kansas and processed through a
berlese trap. To date, this species is known only from specimens
taken in Lawrence, Kansas.
This species is named to honor Dr. George F. Knowlton, who has
contributed vast numbers of mite specimens to the Snow Entomo-
logical Museum collections and has in many other ways encouraged
acarological studies at the University of Kansas.
418 Tue Universrry SCIENCE BULLETIN
Cheletophyes knowltoni was found to be entirely predaceous,
feeding preponderantly on other species of mites, In only one in-
stance, during the course of this study, was predation on an insect
observed and this involved the feeding of a larva on an immature
psoccid. Within the order Acarina this mite has been seen to feed
on members of the families Parasitidae, Laelaptidae, Cheyletidae,
Bdellidae, Acaridae, and Tydeidae. No attempt was made to feed
upon mites of the family Cunaxidae nor on the members of the
groups Oribatei and Uropodina, even when the cheyletid was in a
state of semistarvation and these mites abundant. They were not
observed to feed on Cheyletus eruditus, although found in close
association with that species in stable litter. However, C. eruditus
was often observed feeding on Cheletophyes knowltoni. Cunaxa
sp. taken in great numbers in berlese funnels along with C. knowl-
toni seemed to prey almost exclusively on this cheyletid. All life
stages of the cunaxid could be found feeding on all stages of the
cheyletid, The cunaxid usually attacked the mite from the rear,
being able to outmaneuver the cheyletid by reason of their greater
agility. Cunaxid larvae have been observed overpowering large
cheyletid nymphs.
The mite was cannibalistic if the opportunity presented itself,
feeding on smaller or weakened members of its own species. This
behavior was usually seen in mass rearing pens when the acarids
used for food became scarce. Adults of this species seemed to
ignore extremely small prey such as acarid larvae and would die,
apparently from starvation, in a rearing pen containing a quantity
of the acarid larvae. If an adult acarid was introduced into this
pen, the Cheletophyes would immediately feed upon it.
The immature stages were not seen feeding upon mite eggs of
any species. However, one male kept in isolation with a female
was observed to feed upon her eggs when there was a dearth of
acarids for food.
This species did not require a pit or hiding place in which to lay
in ambush as did C. eruditus. It would run actively about the rear-
ing pen searching for food or, at times, lie flat on the substrate with
legs and palps widespread until some victim brushed against legs I.
The mite would then spring forward and seize its prey by a leg
with the strong palps. The chelicerae would be inserted into the
struggling victim’s leg and, after about thirty seconds the victim
would cease to offer any resistance, its legs would curl downward
and all body movements would cease. During the period that the
venom or toxic saliva was taking effect, the predator would be
SA
Srupres oF Two Species or CHEYLETID MITES 419
\ tugging and lifting at its prey. When the prey offered no more
resistance, it would, if not too large, be lifted from the substrate and
carried about while its body contents were drained. The venom or
toxic saliva seemed to have little or no effect on other mites of its
own species for, in instances of a cannibalistic attack, the victim
would attempt to escape for some time and if released before too
} great a quantity of its body fluids were removed, seemed to recover
completely.
C. knowltoni actively avoided bright lights and would congregate
in the darker corners of the rearing pen when under illuminated
microscope observation. The mite has one pair of eyes and seems
to be able to distinguish movement, turning actively away from a
dissecting needle moved in front of it.
The long front legs are not used in walking but seem to be pri-
marily tactile appendages, being waved constantly on and about
every object the mite was investigating. In attacking prey, once
the prey was close enough to touch with the palps, the front legs
were held laterally and rearward, apparently an attitude adopted
to protect these appendages. This pose was maintained after the
prey was overcome and was being carried or dragged about.
The mite usually remained on the side rather than on the bottom
of the rearing pen. The molting form was usually found in this sec-
tion of the rearing pen, affixed to the substrate with no attempt at
concealment even when such concealment was provided by pits or
bits of foreign material.
\ There was a period of varying duration before molting in which
the mite would take no food, although its movements were quite
active. The mite was not observed actually assuming the molt-
ing position. In this position the mite was prone on the substrate,
with legs I and II stretched forward at full length. The remaining
legs were extended to the rear at full length. This was a very
typical position and was only assumed at the time of molting.
Behavior of adults. Observations of adult activities were made
repeatedly and on. large numbers of individuals. Typical behavior
patterns are noted in the following. The male was observed
guarding a molting deutonymph twelve hours before its emer-
gence. The male would move rapidly around and around the
molting form, tapping and stroking it with the tactile front legs.
At the approach of other mites, the male would stand over the
quiescent form and stab at the intruders with his palps, eventually
driving them away by making short runs at them pinching vigor-
Tue University SciENCE BULLETIN
420
ously with the palps. He would never move far from the deuto-
nymph but immediately return and commence stroking and tapping
it once more with legs I.
Upon emergence of the female, the male would run rapidly
around her, tapping here and there over her body. The female
would raise her front legs over her body and remain quiet. Even-
tually, the male would approach the female and repeatedly pinch
first one and then the other of her palps. The female would re-
main quiet except for a slight movement of legs I held over her
body. Receiving no rebuff, the male would seize one female palp
in both of his and lead the female forward, sideways and backward
for some time, moving very rapidly in this strange sort of dance.
Then the male would half lift the female and slide beneath her,
turning her in a half circle as he did so, so that she was above him,
both mites facing in the same direction with the female legs placed
indiscriminately on the dorsum of the male and on the substrate.
Legs I of the female would still be held upward and to the side.
Swift and repeated copulation would ensue, the venter of the
female bending downward to make contact with the phallus of the
male. Contact was rapid and frequent. The female would then
move away for some distance (about one centimeter) and remain
quiet. The male would search for her and, upon locating her, the
entire courtship and mating would be repeated. A total of ten
courtships and subsequent matings were observed in a period of
about ten minutes. There was no attempt on the part of the female
to harm the male in any way after the mating activity was com-
pleted. In fact, the two mites shared the same rearing pen for ten
days afterward without harming one another,
About forty-eight hours after copulation, egg deposition began.
The eggs were laid singly, always in a depression or cavity of some
type, if available, and were covered by strands of silk. The egg
itself was not touched by the strands which were affixed at the
mouth of the depression to form a loose network. The silk emerged
from the apex of the rostrum in a continuous strand, The female
walked sideways across the depression, but always turned and
moved backward just before anchoring the strand of webbing.
This was done by pressing the apex of the rostrum against the sub-
strate. The angle formed by two connectively-anchored strands
of silk was variable and did not seem to follow any set pattern.
After a few strands had been deposited, the female would walk
on them while crossing the pit. When a number of strands of silk
Srupires or Two Species or CHEYLETID MITES 421
had been anchored close to and on the edge of the pit, the female
would move away from the edge an increasingly greater distance,
apparently to find fresh locations on which to anchor the silk,
During the deposition of the webbing, legs I were held quiescent,
upward and backward over the body. The palps were not observed
to have any function in depositing the silk. The entire meshwork
of silk was formed in about five minutes. Up to sixteen eggs from
a single female were obtained in the laboratory, the eggs being
laid over a period of nine days. The female usually died in one
to two days after her total number of eggs were laid.
Females kept in isolation during the deutonymphal stage would,
upon emerging as adults, start oviposition within two to twelve
days. A maximum of twelve eggs was recovered from an unferti-
lized female. The eggs were viable, the larvae emerging from
two to four days after the eggs were laid.
During the period of egg-laying the female ate voraciously, feed-
ing on five to six acarid females every twenty-four hours and
draining the bodies of each victim completely.
The male, after mating, was quite active for a period of about
twelve days. It was an active feeder, consuming as many as five
acarid females or nymphs within a twenty-four hour period. In
one instance a male was observed feeding on an egg of its own
species, feeding through the web which covered the eggs.
Egg: The eggs when first laid are ovoid, semitransluscent,
smooth and shiny. The embryo head developed at the slightly
broader end, the appendages being clearly visible before hatching.
In size, the eggs averaged 1837p in length and 103. in width.
The egg hatched in two to three days at 80°-85° F., the average
being two and one-fourth days. The egg shells were very con-
spicuous after hatching had taken place, as they reflected a bril-
liant blue light as has been reported by M’Intire (1869) for an
unidentified species of cheyletid.
Larva: For some time after emergence from the egg, the larva
remained under the strands of silk which had covered the egg. To
get from under this silk, the larva would carefully work two strands
apart and force the body through the opening thus made.
The larva usually refused food the first day of its life, but would
make vigorous attempts to feed on any food presented the follow-
ing day. Female acarids, much larger than the larva, were occa-
sionally attacked, the larva seizing a leg with its palps and attempt-
ing to insert the chelicerae in the acarid’s leg. These attempts
422, Tue Universiry SCIENCE BULLETIN
were usually unsuccessful, the larva being knocked off against the
substrate during the acarid’s efforts to escape. However, larvae
have been observed to overpower acarid nymphs much larger than
themselves. It appeared that if they were able to insert the chelic-
erae and remain attached for a period of time, the toxic saliva
or venom which they seem to produce apparently had a quieting
effect on the acarid, eventually producing a type of stupor during
which the cheyletid could feed without interference. Continued
success in rearing larval C. knowltoni was assured when they were
fed a diet of acarid larvae exclusively.
In two instances, recently hatched larvae which were still under
the protective web were killed by being exposed suddenly to a
strong microscope light. Whether it was the light or the attendant
heat is unknown. The larvae, when struck by the light, would
stiffen and expand all legs and mouthparts. Aside from some
flexing of the palps for a short interval following such exposure,
there was no more movement. The larvae, in each case, were at
once returned to the rearing pen and the relative humidity of the
rearing pen was increased. Such larvae did not recover.
For a period of time before molting, the larva took no food. The
larva was not actually observed assuming the molting position
though this characteristic stance was taken before ecdysis. The
duration of the larval stadium was from two to seven days at 80-85°
F., averaging four days.
Protonymph: The protonymph emerged after a twenty-four to
forty-eight hour molting period, In attempting to escape from the
larval skin, the protonymph would pull backward and upward with
the forepart of the body, stretching the larval skin in the vicinity of
legs I. After some minutes of this stretching, a transverse split
would appear in the dorsum of the larval skin between legs Hl and
Ill. The mite backed from the forepart of the larval skin, freeing
legs I and II along with the gnathosoma. After a pause of a few
seconds duration, the mite stepped out of the rear portion of the
larval skin which often remained attached to the dorsum of the
protonymph for ten minutes or more before dropping off. The
actual molting process lasted about twenty-five minutes.
Legs IV of the protonymph were used for walking as soon as the
mite emerged and were not held from the substrate for a time as
was the case in newly emerged protonymphs of Cheyletus eruditus.
The protonymphs occasionally attempted to feed soon after
emergence, but the palps did not appear strong enough to hold the
Srupirs or Two Species or CHeyLetip MITES 423
prey. The protonymph was not observed feeding successfully until
eight to twelve hours after emergence, when it would attack
mites much larger than itself if they were slow-moving. In the
laboratory the protonymph was fed acarid nymphs.
The protonymphal stadium endured from three to seven days at
80°-85° F., averaging four days, the nymph usually molting on the
side of the rearing pen. As in the larval stage, the protonymph re-
fused food for a short period before molting.
Deutonymph: The deutonymph emerged after approximately a
twenty-four hour molting period and would usually refuse food for
eight to twelve hours. The nymph was fed on acarid adults, having
no trouble in subduing them with its toxic saliva or venom. This
stage would refuse food for a period before molting and would molt
on the side of the rearing pen. The deutonymphal stadium lasted
from four to seven days at 80°-85° F., averaging five days.
BIBLIOGRAPHY
AnpreE, M.
1933. Notes sur les Acariens observés dans les magasins regionaux de
tabacs (Avec reproduction d’un Opuscule inédit de Michel Hardy).
Ann. des Epiphyt., vol. 19, no. 6, pp. 831-356.
BAKER, I, W.
1949. A review of the mites of the family Cheyletidae in the United
States National Museum. Proc. U.S. Natl. Mus., vol. 99, no, 32388,
pp. 267-320.
Banks, N.
1906; Descriptions of some new mites. Proc, Ent. Soc. Washington, vol.
7, pp. 138-142,
Brox, R.
1865. A short description of an Acarus and its agamic reproduction.
Trans. Micro. Soc. London, New Series, vol. 14, pp. 30-34.
Ewine, H. E.
1909. A systematic and biological study of the Acarina of Illinois. The
University Studies, University of Illinois, vol. 8, no. 6, pp. 74-77.
1912. The life history and habits of Cheyletus seminivorous Packard.
Jour, Econ. Ent., vol. 5, no. 5, pp. 416-420.
FARRELL, CHaries E.and Warton, G, E.
1949. A culture medium for chiggers (Trombiculidae). Jour, Parasit.,
vol. 35, no. 8, p. 435.
Gausz, G. F., Smaracpova, N., P., and Wirt, A. A.
1936, Further studies of interaction between predators and prey, Jour.
Anim. Ecol., vol. 5, no. 1, pp. 1-18.
HAARL GY, N.
1942, A morphologic, systematic, ecological investigation of Acarina and
other representatives of the microfauna of the soil around Marke-
fjord, Northeast Greenland, Medd, Grgnland, no, 1281, p, 171.
424 Tse Universrry SCIENCE BULLETIN
Hessiinc, THEODOR VON
1852. Einige notizen tiber den Weichselzopf. TIllust. Med, Zeit., vol. 1,
no. 5, pp. 255-259,
Hitz, M. A., and Gornon, R. M.
1945. An outbreak of dermatitis amongst troops in North Wales caused by
rodent mites, Ann, Trop, Med, Parasit. (Liverpool), vol. 39,
no. 1, pp. 46-52,
Homann, H.
1933. Die milben in gesunden Bienenstécken. Zeitschrift fiir Parasiten-
kunde, (Berlin), vol. 6, no. 3, pp. 350-415,
Hucues, A. M.
1948. The mites associated with stored food products, Ministry of
Agriculture and Fisheries, London: 168 pp.
Kozuuina, O.
1940. Déplacement des différentes espéces de tiques sur les divers sub-
stratums, (In Russian with a summary in French), Uchen, Zap,
mosk. gosud, Univ., Moscow, no. 42, (Zool.), pp. 271-277.
Lreovsxy, Louis J.
1953a. Improved technique for rearing chigger mites. Ent. News, vol, 64,
no. 1, pp. 4-7.
1953b. Polyvinyl alcohol with lacto-phenol, a mounting and clearing
medium for chigger mites. Ent. News, vol. 64, no. 2, pp. 42-44.
M’Intimg, S. J.
1869. Cheyleti. Hardwicke’s Science-Gossip, pp. 5-7.
NewsteEAp, R., and Duvauu, H. M.
1918. Bionomic, morphological and economic report on the acarids of
stored grain and flour, Rep, Grain Pests Comm., Roy, Soc, London,
no, 2, pp. 1-52.
OupEMans, A. C,
1903. Notes on Acari (Eleventh series). Tijdschr. voor Ent., vol. 46,
pp. 98-136.
1906, Révision des Chélétinés. Mem, Soc, Zool, France, vol. 19, pp.
86-144,
1914. Acarologische Aanteckeningen LIV. Ent. Berichten Nederland-
ische, vol. 4, no. 78, pp. 101-102.
1938, Nieuwe vondsten op het Gebied der systematick en der Nomencla-
tuur der Acari Il, Tijdschr. voor Ent., vol. 81, pp, Ixx-Ixxx.
Packarp, A. S.
1878. Guide to the study of insects. Henry Holt and Company, New
York, 715 pp.
Pierce, W. D.
1921. Sanitary Entomology. The Gorhan Press, 518 pp.
Pitters, A, W. N.
1920. Some accidental parasites of the ear of the domesticated rabbit
and guinea pig. Vet. Jour., vol, 76, no, 9, pp. 331-334,
Reprkorrzey, V.
1924, On flour mites, Plant Protection, Leningrad, i, nos, 1-2, pp, 47-48.
Stupies oF Two Species oF CHEYLETID MITES 425
Rennig, J., and Harvey, E. J.
1922. Mites on bee hives and on hives. Proc, Physiol. Soc, Edinburg,
vol, 20, pp. 268-270.
Ritey, W. A., and Jonannsen, O. A.
1938. Medical Entomology (2nd edition). McGraw-Hill Book Company,
Inc.,, New York, N. Y. and London, 438 pp.
Ropionov, Z. S,
1937. Conditions of a mass development of grain mites, (In Russian
with English summary.) Zool, Zh., vol. 16, p. 511,
1940. The qualitative and quantitative damage caused by grain mites.
Uchen. Zap, mosk, gosud, Univ,, Moscow, no, 42, (Zool.), pp.
141-165.
Rovronov, Z. S., and FurMaAN, A. V.
1940. Interrelations between the phytophagous and predaceous mites,
Uchen, Zap, mosk, gosud, Univ,., Moscow, no, 42, (Zool.), p. 197.
Siccaarp, N.
1920. Om Frémidder og deres Bekaempelse. Tidsskr, for Planteavl.,
(Copenhagen ), vol, 27, p. 287.
Scurank, F,
1781. Enumerate insectorum austriae indigenorum. Klett, p, 518.
So_omon, M., E.
1946, Tyroglyphid mites in stored products, Ecological studies. Ann,
Appl. Biol., (London), vol, 33, no. 1, pp. 82-97,
Tue Universiry ScuencE BULLETIN
PLATE |
Cheyletus eruditus (Schrank )
Fic. 1. Dorsal view of female. |
Fic. 2. Ventral view of female. \
Fic. 3. Portion of right palpus of female. |
Portion of leg I of female.
Stupies or Two Species or CHEeyLetrip MIrrs 427
PLATE I
Tue Universiry SCIENCE BULLETIN
PLATE II
Cheyletus eruditus (Schrank )
5. Dorsal view of larva.
6. Portion of right palpus of larva.
Fic. 7. Portion of leg I of larva.
8. Portion of right palpus of protonymph.
Fic. 9. Portion of leg I of protonymph.
Fic. 10. Dorsal view of protonymph.
Fic. 11. Portion of left palpus of deutonymph.
Fic. 12. Dorsal view of deutonymph.
Portion of leg I of deutonymph.
429
StuprEs or Two Species or CHeyLetip Mires
PLATE I]
Tur Universrry ScreNcE BULLETIN
PLATE III
Cheletophyes knowltoni, new species
Fic, 14. Portion of leg I of female.
Fic. 15. Portion of right palpus of female.
Fic. 16. Dorsal view of female. |
Struptes or Two Species or CHeyteri Mrres 481
PLATE III
Tue University SCIENCE BULLETIN
PLATE IV
Cheletophyes knowltoni, new species
Fic. 17. Portion of leg I of larva.
Fic. 18. Dorsal view of larva.
Fic. 19. Portion of right palpus of larva.
Ventral view of female.
Stupies or Two Species or Crryterip Mrres 433
PLATE IV
15—3378
Ture University SCIENCE BULLETIN
PLATE V \
Cheletophyes knowltoni, new species
Fic. 21. Portion of leg I of male.
Frc. 22. Portion of left palpus of male.
Fic. 23. Dorsal view of male.
Stupres or Two Species or Cury.erm Mrres 485
PLATE V
Fic. 24.
Fic. 25.
Fic. 26.
Fic, 27,
Fic. 28.
Fic. 29.
Tue Universrry SCIENCE BULLETIN
PLATE VI
Cheletophyes knowltoni, new species
Portion of right palpus of protonymph.
Portion of leg I of protonymph.
Dorsal view of protonymph.
Dorsal view of deutonymph.
Portion of right palpus of deutonymph.
Portion of leg I of deutonymph.
Stupies or Two Species or CHeytetm Mrres 437
PLATE VI
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVI, Pr. 1] Drcemper 20, 1956 [No.6
Comparative Studies of the Thoracic Musculature of Bees *
BY
ALVARO WILLE
Asstract: This paper consists of a study of the thoracic musculature of
bees, Fifty-seven genera were dissected representing five families, The
musculature of eighty-eight species is described.
Some interpretation of the phylogeny and the functional significance of the
muscles are discussed.
Evolutionary trends in the following five muscles are indicated: lateral
promotor of the middle coxa, pleurotergal muscle of the mesothorax, median
depressor of the abdomen, mesal remotor of the hind coxa, and lateral pro-
motor of the hind coxa.
CONTENTS
PAGE
IMOGMOUGH Ya ak ee i ac ee ee da 439
AChMOWIEMOMENY iiss A ee ee 440
INGtHOG She ee ae et RU ad ee 440
MUSCLES etICled ueieeig. Goth chi deka vcr Gade Se eee 4141
Phylogenetic and functional significance of the muscles................. 443
Trends of specialization of some of the muscles................... 0.0. 444.
IVEROMPUVE MGCOUMIS | 4050s ue \Gh 448
POMUIV OUBHORG cer, crys oan eke 448
PATUY SMOUGUORE Siu te eA ties oc ee 449
PAIV AMCUEINUMOs ois Rue tec i ee 451
PAY WVIOCHOMLUCEE for os i ye 453
PaminApidae Ani er i a ee 456
usiterattive: Gite (piv ee eaieel wets elo ea 469
MUNG URORLONG 3 fe en Fi saiglicn cpt. cree cn crab Ge a 471
INTRODUCTION
This paper is a report of a preliminary study of the thoracic
musculature of bees. The purpose of this work is to shed light on
the relationships of the groups, the possible modification of muscles
in relation to function, and evolutionary trends of the thoracic
musculature,
* Contribution No. 910 from the Department of Entomology of the University of Kansas,
(489)
440 Tue Untiversrry SCiENCE BULLETIN
The scope of the work was limited by the specimens available
for dissection. Nevertheless, it was possible to examine 88 species
representing 57 genera, as follows: Family Colletidae: Colletes (2
species), Ptiloglossa, Hylaeus. Family Halictidae: Halictus (2
species), Agapostemon, Neocorynura, Corynura, Sphecodes, Tem-
nosoma, Lasioglossum (2 species), Coenaugochlora, Augochlora,
Angochlorella, Dufourea, Nomia (3 species). F amily Andrenidae:
Andrena (2 species), Perdita (2 species ), Pseudopanurgus, Calliop-
sis, Protoxaea, Oxaea. Family Megachilidae: M egachile (2 species),
Coelioxys (2 species), Heriades, Ashmeadiella, Anthidium, Dianthi-
dium (2 species), Paranthidium, Lithurge. Family Apidae: Exo-
malopsis (8 species), Ancyloscelis (2 species), Paratetrapedia (8
species), Tetrapedia, Nomada, Neopasites, Triepoelus, Melissodes
(6 species), Florilegus, Thygater, Peponapis, Xenoglossa, Xenoglos-
sodes, Ptilothrix, Diadasia (2 species), Melitoma, Anthophora (2
species), Centris (2 species), Epicharis, Xylocopa (2 species),
Ceratina (2 species), Euglossa, Eulaema, Bombus (5 species),
Melipona, Trigona (3 species ), Lestrimelitta, Apis.
ACKNOWLEDGMENT
The author is especially indebted to Dr. C. D, Michener of the
University of Kansas for the critical advice and suggestions during
this research, for the determination of most of the species, and
for obtaining several specimens used in this work. [ am also
thankful to Wallace E. LaBerge for determining the species of
Melissodes, and to Earle A. Cross for determining the species of
Nomia. Thanks are especially due to Maria Bozzoli for aid with
the drawings.
METHODS
The specimens were preserved in Kahle’s fluid (Dietrich’s fluid).
Usually 24 hours fixation was adequate but in the larger forms,
such as Bombus and Xylocopa, sometimes several days were re-
quired. By soaking dry specimens in either Kahle’s solution or
alcohol for at least 24 hours, it was possible to study the muscles
of a few bees (Lestrimelitta, Lithurge, and Corynura) not available
in fresh condition. This method was not always successful.
Each drawing was made diagramatically from specimens dis-
sected under water. The thorax of each was cut through the
mesal plane, after removing the terga. The left section was used
to study the metathoracic muscles, and most of the drawings were
made from above and slightly toward the right side. The right
STupiEs OF THE THoRAcIC MUSCULATURE OF BEES 44]
section was used to study the mesopleural muscles. ‘The meso-
pleural muscles were drawn from various angles to insure accurate
proportions. It should be mentioned that usually several speci-
mens were dissected for every species studied,
MUSCLES STUDIED
In dissecting the muscles the honey bee was used as the standard
type, because it is the only species well known morphologically
(Morison 1927, Snodgrass 1942). For the musculature the termi-
nology of Snodgrass has been followed, and for the skeletal
structures of Michener (1944),
After a general survey it was found that only a few muscles were
variable to any considerable degree among the bees, It seems
that the variations arise only among those muscles that originate
on wide structures, where they actually would have freedom to
change.
The muscles selected for study were the following:
Musculature of the Mesopleura
These are the direct muscles of the wing, a pleurotergal muscle
and a coxal muscle.
1. Basalar muscle (Fig. 8)—This muscle varies very little. It
is a two-branched muscle; the anterior branch is small and slender,
the posterior one is larger and fan-shaped. Both branches originate
on the anterior part of the episternum, The insertion is on the
basalare by a long tendonlike apodeme. The posterior branch
usually overlaps the inferior part of the second and third pleuro-
axillary muscles. The only important modification is seen in the
honey bee, in which the two branches are united in one broad
muscle (Fig. 56),
2. Pleuroaxillary muscles—There are three pleuroaxillary mus-
cles, the first and anteriormost (Fig. 2a, 3) is a small muscle arising
in a pocket near the summit of the pleuron. This muscle does not
vary much, being confined in the pocket mentioned above. The
other two muscles (Fig. 2b, 2c, and 8) are larger and originate
ventrally on the episternum. They all insert on the third axillary
sclerite.
3. Pleurotergal muscle (Fig, 3)—This is a variable muscle. It
has its origin on the episternum and its insertion on the lateral
margin of the scutum.
4. Lateral promotor of the middle coxa (Fig. 3)—This is also
a highly variable muscle. It usually arises on the under side and
442, Tur Universtry ScrENCE BULLETIN
on the base of the pleural apophysis and on the episternum. It
inserts laterally on the base of the middle coxa.
Musculature of the Metathorax
From this region the following muscles were studied: the coxal
muscles, the thoracic depressor of the trochanter, and two inter-
segmental muscles.
5. Median depressor of the abdomen (Fig. 70)—This muscle
usually originates on the coalesced furcal arms and on the free
distal portion of the metafurcal arm. It inserts medially on the
sternum of the petiole segment (first metasomal segment).
6. Ventral lateromotor of the abdomen (Fig. 21)—This is a
rather small muscle arising on the edge of the vertical place of the
metasternum. It has its origin either at the posterior part of the
edge, or at the anterior part, or along the whole edge. It inserts
laterally on the sternum of the petiole segment.
7. Mesal remotor of the hind coxa (Fig. 24)—This muscle usually
originates on the free basal portion of the metafurcal arm and on
the coalesced furcal arms. In some bees fibers also arise on the
anterior mesal edge of the intercoxal lamella, It inserts on the
posterior basal margin of the hind coxa.
8. Thoracic depressor of the trochanter (Fig. 23)—This muscle
does not show much variation. It arises on the coalesced furcal
arms and on the free distal portion of the metafurcal arm. It in-
serts in the trochanter of the hind leg.
9. Lateral promotor of the hind coxa (Fig. 28)—This muscle
arises on the lateral portion of the intercoxal lamella and on the
propodeum, just behind the third pleural ridge. The insertion is
on the anterior and lateral margin of the hind coxa.
10. Mesal promotor of the hind coxa (Fig, 25)—Showing little
variation, this muscle originates on the vertical plate of the meta-
sternum, where it is thick, and covers much of the plate (except in
the subfamily Megachilinae ) (compare Fig. 25 with Fig. 28). Some
fibers also may arise on the mesal portion of the intercoxal lamella.
Its maximum extension is found in the subfamily Oxaeinae, and the
minimum development in Epicharis (Fig. 65). The insertion is on
the anterior margin of the coxa of the hind leg.
It should be pointed out that a few other muscles, such as the
basalar muscle of the metathorax, and the median levator of the
abdomen, were variable, but these muscles are not considered in
this paper.
SrupiEs oF THE THoRAcIC MUSCULATURE OF BEES 443
PHYLOGENETIC AND FUNCTIONAL SIGNIFICANCE
OF THE MUSCLES
Judging by the specimens dissected three general conclusions can
be drawn. If two bees have almost identical mesopleural and meta-
thoracic muscles then the two forms are related at or below the
subfamily level. The similarity of musculature does not necessarily
indicate whether the relation is at the subfamily, tribe, or generic
level. Only in one instance did the musculature show relationships
above the family. This was the case in the two primitive families,
Collitidae and Halictidae, in which the musculature is almost identi-
cal in all forms studied (except for the genus Nomia). It should
be pointed out that Michener (1953) showed a close relationship
of these two families and the family Andrenidae on the basis of
larval characters. Examples of other bee groups showing almost
identical musculature are the following: subfamily Oxaeinae, tribe
Meliponini, tribe Euglossini, Bombus, Ceratina, Xylocopa, Centris,
Megachile, Coelioxys, Dianthidium with Paranthidium, Perdita.
More species should be dissected to prove or disprove the homo-
geneity of the musculature in each of these eleven groups. In the
genera in which only one species was dissected it is impossible to
predict whether additional species have similar or different muscu-
lature.
The second conclusion to be drawn from the dissections is that
if two bees do not show similarity in either the mesopleura or meta-
thoracic musculature, lack of relationship is not necessarily indi-
cated. This is well illustrated in the honey bee where the worker
cast has a different type of mesopleural musculature from that of
the male (see Fig. 56 and Fig, 58). Other examples are found
among the species of Nomia, Exomalopsis, Paratetrapedia, Melis-
sodes, and Diadasia,
The last logical conclusion to be drawn is that the modifications
of the muscles do not have any obvious relation to function. For
instance, Hylaeus and Petiloglossa are different not only externally
but also in behavior, the first being a small slender, weak flyer, the
second being a large, robust, fast flying form. Yet both have
essentially the same type of musculature, presumably showing their
phyletic relationship to one another. On the other hand, different
species of the genus Nomia differ in the mesopleural muscles al-
though these bees are very similar exteriorly and in behavior (Figs.
6 and 7),
Some of the thoracic muscles in the bees have been modified
444 Tour University ScreENCE BULLETIN
independently in several groups, while the same muscles have re-
tained an unspecialized condition in others. This has resulted in a
great deal of convergence of individual muscles. For example, the
lateral promotor of the middle coxa has the same modification in
some Nomia as in some Melissodes, which are quite unrelated forms
as judged by many other characters, while it has an unspecialized
condition in the family Colletidae, a primitive family, and in the
tribe Meliponini, one of the most specialized groups of Apidae.
Another example is the lateral promotor of the hind coxa which
shows convergence in Anthidium and Thygater; in both forms the
inner lobe has been eliminated, Another specialization of the same
muscle is the elimination of the outer lobe instead of the inner one,
which has been accomplished independently in Neopasites and
Apis.
TRENDS OF SPECIALIZATION OF SOME OF
THE MUSCLES
It was possible to trace the probable trends of specialization for
at least half of the muscles studied. The Colletidae and Halictidae
(except Nomia) are the only families of bees which exhibit almost
identical mesopleural and metathoracic muscles; the other groups
show a great deal of variation. These modifications are easily trace-
able and are derived from the pattern found in the Colletidae and
Halictidae. On the basis of this, a table was made listing five sup-
posed unspecialized types on the left, and their specialized alter-
natives on the right. It should be mentioned that intermediate con-
ditions occur.
Unspecialized Specialized
1. Lateral promotor of the middle 1. Lateral promotor of the middle
coxa fan-shaped. coxa V-shaped or ribbonlike,
2. Pleurotergal muscle of the meso- 2. Pleurotergal muscle of the meso-
thorax twice or almost twice as thorax as long as wide.
long as wide.
3. Median depressor of the abdo- 8. Median depressor of the abdo-
men a triangular muscle. men. ribbonlike.
4, Mesal remotor of the hind coxa 4. Mesal remotor of the hind coxa
not fused anteriorly with the fused anteriorly with the thoracic
thoracic depressor of the hind depressor of the hind trochanter.
trochanter.
5. Lateral promotor of the hind 5. Lateral promotor of the hind coxa
coxa with both outer and inner
lobe.
with only one of the lobes.
STUDIES OF THE THORACIC MusCULATURE OF BEES 445
Lateral Promotor of the Middle Coxa (Fig. 1)
As indicated above, the unspecialized type is a fanshaped muscle
found in all Collitidae and Halictidae (12 genera) except Nomia.*
The main trend of specialization of this muscle is to become V-
shaped by the formation of an inferior arm. In-the family Andren-
idae two genera, Andrena and Pseudopanurgus, six dissections
were found with the fan-shaped type, and in one of these ( Pseudo-
panurgus) the muscle already has some of the inferior fibers ex-
tended anteriorly. In Megachilidae four genera (Heriades, Ash-
meadiella, Coelioxys, Anthidium) eight dissections exhibit the fan-
shaped type, but Coelioxys and Anthidium, like Pseudopanurgus,
have some of the inferior fibers of the muscle extended anteriorly,
without forming a true inferior arm. In the family Apidae only
nine genera of the twenty-eight dissected have retained the un-
specialized type. The genera showing this condition are the fol-
lowing: Neopasites, Thygater, Peponapis, Melitoma, Centris, An-
thophora and the tribe. Meliponini. Of these genera, Neopasites
has some of the inferior fibers well extended anteriorly, almost form-
ing an inferior arm. On the other hand, Thygater, Melitoma, and
Anthophora show partial splitting of the muscle.
It should be mentioned that usually in the V-shaped type the
superior fibers arise on the under side and on the base of the pleural
apophysis, and the inferior arm on the episternum leaving a weak
area in the middle of the muscle. Perhaps because the pull of the
muscle is made by the fibers which are attached to the pleural
apophysis and by the inferior arm, the area in the middle becomes
concave and somewhat overlapped by the inferior arm and superior
fibers. This gives the impression of a muscle having a superior arm,
but this is actually the case only when the area in the middle is
eliminated or partially so, as in many bees,
Two further specializations can be followed from the V-shaped
lateral promotor of the middle coxa. One of these is the enlarge-
ment of the inferior arm as in Melissodes obiqua, Diadasia enevata,
Nomia mortoni, Bombus medius and Nomada. The other line is the
elimination of the inferior arm. This is well illustrated in the honey
bee. In the worker cast the lateral promotor is a V-shaped muscle
with the inferior arm not enlarged, while in the male only the
superior arm is present. Among the thirty males dissected one
specimen was found with an intermediate condition (Fig. 57),
*In_ order to avoid needless repetition only the generic name is used either when one
species has been dissected in the genus or when all the species of a genus have the same
type of muscle under discussion,
446 Ture University SCIENCE BULLETIN
It is surprising that in Vespula the lateral promotor of the middle
coxa does not originate on the mesopleura (Duncan 1939). But
according to Maki (1938) in Vespa“. . . there is a strong pleural
abductor arising on the episternum near the pleural ridge by a very
broad base and attached on the anterolateral basal rim of the coxa
by atendon.” (pp. 234.)
Pleurotergal Muscles of the Mesothorax
The pleurotergal muscle shows some correlations with the lateral
promotor of the middle coxa. The unspecialized type is a dorso-
ventrally elongated muscle twice or almost twice as long as wide,
with a broad base. The inferior end is at the same level or a little
above that of the inferior end of the third pleuroaxillary muscle,
This type is found in Collitidae, Halictidae (except Nomia), An-
drena (one of the two species dissected), Ashmeadiella, Peponapis,
Xenoglossa and Apis (males only). It is correlated with the fan-
shaped type of lateral promotor of the middle coxa, or if the latter
is V-shaped, as in Xenoglossa, the inferior arm is very slender. A
modification from this type is found in Andrena argemonis, Psuedo-
panurgus, Coelioxys, Anthidium, and Neopasites, in which the in-
ferior end of the muscle is pointed, forming a rather leaf-shaped
muscle, This shape is also correlated with the fan-shaped type
of lateral promotor, but the latter has a tendency to have the inferior
fibers extended anteriorly, following the contour of the pleurotergal
muscle.
From these apparently less-specialized types arises a series of
different kinds of pleurotergal muscles. Probably the most special-
ized type of this series is a rounded or quadrangular muscle, as
long as wide and tending to be above the level of the inferior end
of the third pleuroaxillary muscle. This type is usually correlated
with a V-shaped lateral promotor of the middle coxa. It is found
in Nomia (Dieunomia), Calliopsis, Perdita, Oxaea, Protoxaea, Mega-
chile, Exomalopsis, Ancyloscelis, Nomada, Triepeolus, Melissodes,
Thygater, Ptilothrix, Epicharis, Ceratina, Xylocopa, Diadasia, and
Bombus.
Median Depressor of the Abdomen
In all the bees dissected, except those mentioned below, the
median depressor of the abdomen is a large, triangular muscle
arising on the coalesced furcal arms and on the free distal portion
of the metafurcal arm, usually occupying the entire posterior sur-
face of the dorsal process. A modification of this type is a rather
ribbonlike muscle with the origin restricted to the coalesced furcal
STUDIES OF THE THORACIC MUSCULATURE OF BEES 447
arms, This type is found in Perdita, Nomada, Neopasites (with
few fibers on the free distal portion of metafurcal arm), Anthophora,
Ceratina (last two genera with few fibers on mesofurcal bridge )
and Centris (but triangular in shape). In the tribe Meliponini,
including Trigona, Melipona, and Lestrimelitta, the origin is only
on the mesal half of the coalesced furcal arms. Still a further
modification is found in Apis in which the muscle is very slender,
arising on the basal portion of the metafurcal arm.
Mesal Remotor of the Hind Coxa
In most bees this muscle is divided into outer and inner parts.
Intermediate conditions occur between a well-divided muscle (Fig.
26) and one in which no division can be detected (Fig. 24). This
is well exemplified in the subfamily Halictinae. In Halictus the
two parts are well fused. In Augochlora the division is more or
less evident, while in Coenaugochlora the muscle is well divided.
A highly specialized type is that found in Bombus; the muscle
is fused along all the anterior part with the thoracic depressor of
the hind trochanter (Fig. 70). This is also accomplished in smaller
degree in Epicharis, Another modification is the reduction of the
outer part leaving only a ribbonlike muscle as in Ceratina and Apis.
Lateral Promotor of the Hind Coxa (Fig. 2)
This somewhat triangular muscle can be said to be composed of
two lobes, The inner lobe arises on the lateral part of the intercoxal
lamella, just behind the anterior edge. The outer lobe originates,
partly on the intercoxal lamella and partly on the propodeum, just
behind the metapleural ridge. The great majority of bees have
retained this type of arrangement. Nevertheless three lines of
specialization can be followed. In one, the whole muscle extends
over the lateral part of the intercoxal lamella. This trend can
be observed in the family Andrenidae. In Andrena, Calliopsis,
Perdita, and Pseudopanurgus the muscle is already well extended
and fan-shaped, while in the subfamily Oxacinae the lateral pro-
motor of the hind coxa reaches the highest development (Fig. 25).
In the other two lines of specialization either the outer or inner
lobe tends to be eliminated. The suppression of the inner lobe is
well illustrated in certain genera of the family Megachilidae, In
Heriades and Anthidium the inner lobe is already very small, while
in Dianthidium, Paranthidium, and Ashmeadiella the same lobe is
completely eliminated. The same trend is observed in the tribe
Eucerini (family Apidae) in which Peponapis has a small inner
lobe while in Thygater the same lobe is absent, The elimination of
448 Tur Universrry ScrENCE BULLETIN
the outer lobe was observed only in Neopasites and Apis (family
Apidae). In the genera Exomalopsis (2 species of the 3 dis-
sected), Melissodes, Florilegus, Epicharis, Anthophora and Xylo-
copa the lobes of the lateral promotor of the hind coxa are well
separated by a notch,
It should be mentioned that the specialization of the lateral pro-
motor of the hind coxa shows a remarkable parallelism with that of
the lateral promotor of the middle coxa. The specializations of
these two muscles are not correlated, however. The muscle splits,
forming outer and inner lobes (corresponding to superior and in-
ferior arms of the lateral promotor of the middle coxa), The elimi-
nation of the inner lobe (Anthidium) or inferior arm (Apis males )
also occurs in both muscles.
DESCRIPTIVE ACCOUNTS
FAMILY COLLETIDAE
Only three genera were dissected in this family, but all the three
forms presented almost identical musculature,
Subfamily CoLLETINAE
Tre COLLETINI
Colletes sp.
(Female. Carmen, Hidalgo, Mex., July 22, 1954)
Mesopleura divided into anterior and posterior parts by pre-
episternal ridge. Basalar and pleuroaxillary muscles in front of
ridge. Third pleuroaxillary overlapping posterior section of the
second; pleurotergal and lateral promotor of middle coxa on pos-
terior part of pleura. Pleurotergal muscle dorsoventrally elongated;
lateral promotor of middle coxa fan-shaped (Fig.3). In metathorax
median depressor of abdomen a large triangular muscle, originating
on coalesced furcal arms and on free distal portion of metafurcal
arm. Ventral Jateromotor of abdomen small, arising on posterior
ridge of vertical plate of metasternum. Mesal remotor of hind coxa
well divided posteriorly, outer part on mesal section of coalesced
furcal arms and on free basal portion of metafurcal arm, inner part
on base of metafurcal arm. Thoracic depressor of hind coxa on
coalesced furcal arms and on free distal portion of metafurcal arm,
few fibers on lateral portion of intercoxal lamella. Lateral promotor
of hind coxa well lobed. Mesal promotor of hind coxa not covering
small section anteriorly of mesal portion of intercoxal lamella,
Srupies or THE THoracic MuscuLATuRE OF BEES 449
Colletes sp.
(Female. Xilitla, S. L. P., Mex., July 25, 1954)
This species agrees with the description of the previous one.
Trr~ CAUPOLICANINI
Ptiloglossa mexicana (Cresson)
(Female)
This species agrees with Colletes in nearly all details. The fol-
lowing differences exist: Outer part of median remotor of hind
coxa overlapping inner part anteriorly (Fig. 21); thoracic depres-
sor of hind trochanter without fibers arising on intercoxal lamella.
Lateral promotor of hind coxa not so well lobed. Mesal promotor
of hind coxa almost covering mesal portion of intercoxal lamella.
Subfamily HyLarinar
Hylaeus pygmaeus Cresson
(Female )
This species agrees with the description of Colletes.
..Famiry Haricripar
The musculature of the genera dissected (except Nomia) agrees
essentially with that of the family Colletidae (Fig. 4 and 5).
Subfamily Haicrinar
The following species were dissected: Halictus (Halictus) ligatus
Say (female), Halictus (Seladonia) hesperus Smith (worker, queen
and male), Agapostemon radiatus (Say) male and female, Neo-
corynura sp. (female, Xilitla, S.L. P., Mex., July 25, 1954), Cory-
nura rubella (Haliday) female, Sphecodes sp. (female, Filadelfia,
Guanacaste, Costa Rica, March 1954), Temnosoma smaragdinum
Smith (female), Lasioglossum (Paralictus) cephalotes (Dalla Torre),
Lasioglossum (Chloralictus) (worker, Lawrence, Kansas, June,
1954), Coenaugochlora nigromarginata (Spinola) female, Augo-
chlorella aurata (Smith) female, Augochlora pura Say ( female).
Halictus ligatus Say
In comparing this species with that of Colletes the following
points should be mentioned. Ventral lateromotor of abdomen
arises along ridge of vertical plate of metasternum. Méesal re-
motor of hind coxa not divided, originating on coalesced furcal
arms and on free basal portion of metafurcal arm. Thoracic de-
pressor of hind trochanter without fibers arising on intercoxal la-
450 Tue Universiry SCIENCE BULLETIN
mella. Lateral promotor of hind coxa not so well lobed. Mesal
promotor of hind coxa covering the mesal portion of intercoxal
lamella.
All the species of the Halictinae dissected agree with the
description of Halictus ligatus. A few differences, however, were
noted. In Halictus hesperus, Augochloropsis, Augochlorella, Augo-
chlora, and Coenaugochlora the pleurotergal muscle is wider at
the inferior end. In Sphecodes the second pleuroaxillary muscle
is very narrow. In Augochlora and Coenaugochlora the mesal re-
motor of the hind coxa is divided.
Subfamily DurourEInaE
Dufourea marginata (Cresson )
(Female )
The musculature of this species is similar to that of Halictus
ligatus, but differs in having second pleuroaxillary muscle more
extended posteriorly (Fig. 5), and ventral lateromotor of abdo-
men originating on posterior ridge of vertical plate of metasternum,
as is the Colletidae.
Subfamily Nomunar
This group differs from the other two subfamilies in the meso-
pleural musculature.
Nomia (Acunomia) nortoni Cresson
(Male)
Mesopleura not divided by pre-episternal ridge. Lateral pro-
motor of middle coxa with wide inferior arm. Pleurotergal muscle
above level of inferior end of third pleuroaxillary muscle. Third
pleuroaxillary muscle posteriorly overlapping inferior arm of lateral
promotor, and anteriorly second pleuroaxillary muscle (Fig. 6a
Metathoracic musculature (Fig. 22) agreeing with description of
Colletes except as follows: ventral lateromotor of abdomen arises
along ridge of vertical plate of metasternum; outer part of mesal
remotor of hind coxa originating on coalesced furcal arms and on
free basal portion of metafurcal arm.
Nomia (Epinomia) micheneri Cross
(Male)
Mesopleura divided by pre-episternal ridge. Lateral promotor
of middle coxa with slender inferior arm. Third pleuroaxillary
muscle not overlapping either lateral promotor or pleurotergal
SrupiEs OF THE THoRACIC MuscuLATURE OF BEES 451
muscle (Fig. 7). Metathoracic musculature as in N. nortoni, but
thoracic depressor of hind trochanter without fibers arising on
intercoxal lamella. Lateral promotor of hind coxa not as well
lobed, and mesal promotor of hind coxa covering mesal portion
of intercoxal lamella.
Nomia (Dieunomia) sp.
(Female. Western Kansas)
Mesopleura divided by pre-episternal ridge. Lateral promotor
with wide inferior arm but not as wide as in N. nortoni. Pleurotergal
muscle small and rather rounded. Third pleuroaxillary muscle over-
lapping posteriorly inferior arm of lateral promotor and pleurotergal
muscle, but not overlapping anteriorly second pleuroaxillary muscle
(Fig. 8). Metathoracic musculature agrees with that of N. mich-
eneri.
FamiLy ANDRENIDAE
All the six genera dissected agree in having the lateral promotor
of the hind coxa well extended on lateral portion of the intercoxal
lamella. The indication of the outer and inner lobes are suppressed
by the rather fan-shaped muscle. The mesal promotor of the hind
coxa almost covers the mesal portion of the intercoxal lamella.
Subfamily ANDRENINAE
Andrena argemonis Cockerell
(Male)
Third pleuroaxillary muscle of mesopleura posteriorly overlapping
anterior half of pleurotergal muscle. Pleurotergal muscle dorso-
ventrally elongated, but with inferior end pointed, leaf-shaped.
Lateral promotor of middle coxa fan-shaped, as in the Colletidae
(Fig. 9). In metathorax median depressor of abdomen a large tri-
angular muscle, originating on coalesced furcal arms and on free
distal portion of metafurcal arm. Ventral lateromotor of abdomen
on posterior ridge of vertical plate of metasternum. Mesal remotor
of hind coxa well divided posteriorly, outer part triangular in shape,
originating on free basal portion of metafurcal arm, on coalesced
furcal arms, and few fibers on free distal portion of metafurcal arm;
inner part ribbonlike and arising on base of metafurcal arm. Tho-
racic depressor of hind trochanter on coalesced furcal arms and on
free distal portion of metafurcal arm (Fig. 23).
Tue Universrry. ScreNcE BULLETIN
Andrena sp.
(Female, Pachuca, Hidalgo, Mex., July 29, 1954)
This species agrees with the description of the previous one in
nearly all details. The following differences were noted: third
pleuroaxillary muscle of mesopleura posteriorly overlapping less
than half of pleurotergal muscle; pleurotergal muscle not leaf-
shaped; in the metathorax the outer part of mesal remotor of hind
coxa arising only on free basal portion of metafurcal arm.
Subfamily PANURGINAE
The species dissected agree in having the mesal remotor of the
hind coxa either not divided or the division not evident.
Trine PANURGINI
Perdita (Cockerellia) sp.
(Female. Acapulco, Mex., August 6, 1954)
Mesopleura musculature different from that of Andrena. Second
and third pleuroaxillary muscles overlapping anterior part of pleuro-
tergal and lateral promotor of middle coxa. Pleurotergal muscle
rounded and above level of inferior end of third pleuroaxillary.
Lateral promotor of middle coxa V-shaped (Fig. 10). In metathorax
median depressor of abdomen a ribbonlike muscle, arising on
coalesced furcal arms. Ventral lateromotor of abdomen on posterior
half of vertical plate of metasternum. Mesal remotor of hind coxa
divided but not evident, because outer part somewhat overlaps
inner part; outer part on coalesced furcal arm, and on free basal
portion of metafurcal arm, inner part on base of metafurcal arm.
Thoracic depressor of hind trochanter on lateral section of coalesced
fureal arm and on free distal portion of metafurcal arm.
Perdita sp.
(Female. Llera, Tamaulipas, Mex., July 20, 1954)
This species agrees with the description of the previous one.
Pseudopanurgus sp.
(Female. Lagos de Moreno, Jalisco, Mex., August 21, 1954)
Mesopleura musculature somewhat similar to that of Andrena
argemonis (Fig. 11). But metathoracic musculature differs from it
as follows: ventral lateromotor of abdomen arises along ridge of
vertical plate of metasternum; mesal remotor of hind coxa not
divided, arising on free distal portion of metafurcal arm, on coa-
lesced furcal arms, and on free basal portion of metafurcal arm (Fig.
StrupiEs OF THE THoRACIC MusCULATURE OF BEES 453
24); thoracic depressor of hind trochanter a ribbonlike muscle,
originating on coalesced furcal arms, some fibers attach to lateral
promotor of hind coxa.
Calliopsis sp.
(Female, Carmen, Hidalgo, Mex., July 30, 1954)
Mesopleura musculature somewhat similar to that of Perdita
(Fig. 12), but third pleuroaxillary muscle smaller and overlapping
section of second. Methathoracic musculature agrees with descrip-
tion of Pseudopanurgus.
Subfamily Oxarmar
The species in this group are characterized by the great develop-
ment of the lateral and mesal promotors of the hind coxa, both
muscles cover completely the intercoxal lamella (Fig. 25).
Protoxaea nigerrima (Friese )
(Female)
Mesopleura musculature somewhat similar to that of Calliopsis
(Fig. 13), but third pleuroaxillary muscle large. Second pleuro-
axillary ribbonlike. In metathorax median depressor of abdomen as
in Andrena. Ventral lateromotor of abdomen on posterior half of
vertical plate of metasternum, as in Perdita. Mesal remotor of hind
coxa well divided posteriorly, outer part on free distal portion of
metafurcal arm, on coalesced furcal arms, and on free basal portion
of metafurcal arm; inner part ribbonlike and arising on base of
metafurcal arm. Thoracic depressor of hind trochanter on coalesced
furcal arm,
Oxaea sp.
(Female. Independencia, Paraguay, February 20, 1951)
The musculature of this species agrees with the description of
Protoxaea nigerrima.
Famity MEGACHILIDAE
Subfamily MEGACHILINAE
All the species dissected in this group agree in having a thin
and transparent plate which originates from the coalesced furcal
arms and the free basal portion of the metafurcal arm, The furcal
plate (as it is here called) attaches along the mesal portion of the
intercoxal lamella, covering the anterior ridge (Fig. 28), It should
be mentioned that the mesal promotor of the hind coxa originates
at the base of the vertical plate of the metasternum (Fig. 28).
Tue University SCIENCE BULLETIN
Tre MEGACHILINI
Megachile sp.
(Female. Taxco, Guerrero, Mex., August 8, 1954)
Third pleuroaxillary muscle of mesopleura overlapping part of
second pleuroaxillary, and part of pleurotergal and lateral promo-
tor of middle coxa. Pleurotergal muscle rounded; lateral promotor
of middle coxa with inferior arm (Fig. 14). In metathorax median
depressor of abdomen a large triangular muscle, originating on
coalesced furcal arms and on free distal portion of metafurcal arm.
Ventral lateromotor of abdomen on anterior ridge of vertical plate
of metasternum, some fibers on free basal portion of metafurcal
arm. Mesal remotor of hind coxa well divided (Fig. 26), outer
part on furcal lamella, inner part arises along the attachment of
furcal lamella to mesal portion of intercoxal lamella, and on base
of metafurcal arm. Thoracic depressor of hind trochanter on free
distal portion of metafurcal arm, and on lateral half of coalesced
furcal arm. Lateral promotor of hind coxa well extended but
lobed, anterior end of inner lobe under furcal lamella (Fig. 28).
Megachile sp.
(Male, Edinburg, Texas, July 17, 1954)
This species agrees with the description of the previous one.
Coelioxys sp.
(Female, Lawrence, Kan., June 3, 1954)
Mesopleura musculature similar to that of Andrena argemonis
(Fig. 15). Metathoracic musculature like that of Megachile but
parts of mesal remotor of hind coxa not widely separated pos-
teriorly.
Coelioxys sp.
(Female. Xilitla, S. L. P., Mex., July 25, 1954)
The musculature of this species is like the previous one.
Heriades variolosa purpurascens Cockerell
(Male)
Mesapleural musculature somewhat resembles that of Coelioxys
(Fig. 17), but pleurotergal muscle rounded and not overlapped
anteriorly by second pleuroaxillary. In comparing the metathoracic
musculature with that of Megachile the following differences are
noted: ventral lateromotor of abdomen arises along ridge of vertical
plate of metasternum, some fibers on free basal portion of meta-
Srupies OF THE THORACIC MusCULATURE OF BEES 455
furcal arm; parts of mesal remotor of hind coxa not widely sepa-
rated posteriorly, as in Coelioxys; lateral promotor of hind coxa
with small inner lobe.
Ashmeadiella opuntiae (Cockerell)
(Male)
Mesopleural musculature similar to that of Heriades variolosa
(Fig. 16). Metathoracic musculature differs from that of Megachile
as follows: Ventral lateromotor arises along ridge of vertical plate
of metasternum. Outer part of lateral promotor of hind coxa on
mesal half of coalesced furcal arms (or furcal plate), and on free
basal portion of metafurcal arm, both parts not widely separated
posteriorly, as in Coelioxys and Ashmeadiella, lateral promotor of
hind coxa without inner lobe.
Tre ANTHIDIINI
The species dissected in this tribe were found with the mesal re-
motor of the hind coxa not divided.
Anthidium maculosum Cresson
(Female)
Mesopleural musculature similar to that of Coelioxys (Fig. 18),
but second pleuroaxillary muscle smaller. Metathoracic musculature
differs from that of Megachile as follows: mesal remotor of hind
coxa not divided; lateral promotor of hind coxa with small inner
lobe (in one specimen inner lobe absent) Fig. 27.
Dianthidium sayi Cockerell
(Male and Female)
In mesopleura pleurotergal muscle dorsoventrally elongated.
Lateral promotor of middle coxa V-shaped with slender inferior arm
(Fig. 19). Metathoracic musculature like that of Megachile but
mesal remotor of hind coxa not divided, and lateral promotor of
hind coxa without inner lobe.
Dianthidium sp.
(Female, Padilla, Tamaulipas, Mex., August 26, 1954)
The musculature of this species is like the previous one.
Paranthidium sp.
(Female, Taxco, Guerrero, Mex., August 8, 1954)
The musculature of this species agrees with the description of
Dianthidium.
Tue Universrry Science BULLETIN
Subfamily LirHurcinaE
This subfamily contains only the genus Lithurge, and the only
species dissected does not have the furcal plate.
Lithurge sp.
(Female, EI Peumo, Chile. December 28, 1950)
In mesopleura third pleuroaxillary muscle overlapping only
anterior part of pleurotergal and lateral promotor of middle coxa.
Lateral promotor with somewhat slender inferior arm (Fig. 20). In
comparing the metathoracic musculature with that of Megachile
the following difference is noted: mesal remotor of hind coxa not
divided, arising on coalesced furcal arms and on free basal portion
of metafurcal arm.
Famity APIDAE
The bees in this family exhibit considerable diversity in their
musculature.
Subfamily ANTHOPHORINAE
Tre EXOMALOPSINI
Exomalopsis sp.
(Female. E] Puente, Jalisco, Mex., August 20, 1954)
In mesopleura second and third pleuroaxillary muscles overlap-
ping anterior part of pleurotergal and lateral promotor of middle
coxa. Third pleuroaxillary small, with long tendonlike apodeme.
Pleurotergal muscle fan-shaped and above level of inferior end of
third pleuroaxillary. Lateral promotor of middle coxa V-shaped,
with long inferior arm (Fig. 29). In metathorax median depressor
of abdomen a large triangular muscle arising on free distal portion
of metafurcal arm, and on coalesced furcal arms. Ventral latero-
motor of abdomen a small muscle originating on posterior ridge of
vertical plate of metasternum. Mesal remotor divided, outer part
triangular-shaped and arising on free distal portion of metafurcal
arm, on coalesced furcal arms, and on free basal portion of metafur-
cal arm; inner part on basal portion of metafurcal arm, and on
anterior ridge of mesal portion of intercoxal lamella; outer part
overlaps inner part anteriorly. Thoracic depressor of hind tro-
chanter on free distal portion of metafurcal arm and on coalesced
furcal arms. Lateral promotor of hind coxa with outer and inner
lobes separated by a notch. Mesal promotor of hind coxa not cover-
ing whole mesal portion of intercoxal lamella (Fig. 59),
Srupies oF THE THORACIC MusCULATURE OF BEES 457
Exomalopsis: sp.
(Female, San Fernando, Tamaulipas, Mex., August 27, 1954)
Mesopleura musculature differs from above species as shown
in Fig. 30. Metathoracic musculature differs as follows: ventral
lateromotor of abdomen arising along ridge of vertical plate of
metasternum; lateral promotor of hind coxa not divided by a notch.
Exomalopsis snowi Cockerell
(Female )
The description of this species agrees with the above species
except that the lateral promotor of the hind coxa is divided by a
notch,
Ancyloscelis armatus Smith
(Female)
In mesopleura third pleuroaxillary muscle overlapping anterior
part of pleurotergal and lateral promotor of middle coxa. Pleuro-
tergal muscle fanshaped. Lateral promotor of middle coxa V-
shaped (Fig. 31). In comparing the metathoracic musculature
with that of Exomalopsis the following differences were noted:
ventral lateromotor of abdomen arising along edge of vertical plate
of metasternum (which is very short); mesal remotor of hind coxa
apparently without division and only few fibers arising on free
distal portion of metafurcal arm; lateral promotor of hind coxa not
divided by a notch.
Ancyloscelis sp.
(Female, Xilitla, S. L. P., Mex., July 25, 1954)
Mesopleura musculature agrees with description of Exomalopsis
(from E] Puente), except that third pleuroaxillary muscle is larger
than second, and inferior arm of lateral promotor of middle coxa
not so wide. In metathorax ventral lateromotor of abdomen aris-
ing along edge of vertical plate of metasternum (which is very
short), Mesal remotor of hind coxa apparently without division
and only a few fibers arising on free distal portion of metafurcal
arm, Lateral promotor of hind coxa not divided by a notch.
Paratetrapedia sp.
(Male. Xilitla, S. L. P., Mex., July 24, 1954)
In mesopleura third pleuroaxillary muscle overlapping only small
section of pleurotergal and lateral promotor of middle coxa. Pleuro-
tergal muscle leaf-shaped. Lateral promotor of middle coxa V-
shaped, with arched arms (Fig. 32). Metathoracic musculature
458 Tue Universrry Screncek BULLETIN
differs from that of Exomalopsis as follows: Mesal remotor of hind
coxa not divided and with few fibers arising on free portion of meta-
furcal arm, as is in Ancyloscelis. Lateral promotor of hind coxa
not divided by a notch, as in Ancyloscelis.
Paratetrapedia sp.
(Male. Xilitla, S. L. P., Mex., July 24, 1954)
Mesopleura musculature somewhat similar to that of the above
species (Fig. 33), but third pleuroaxillary muscle not as large, and
lateral promotor of middle coxa without arched arms. Metathoracic
musculature similar to that of Exomalopsis, with the following
differences: outer part of mesal remotor of hind coxa arising on
coalesced furcal arms and on free basal portion of metafurcal arm;
lateral promotor of hind coxa not divided by a notch.
Paratetrapedia sp.
(Female. Xilitla, S. L. P., Mex., July 24, 1954)
In mesopleura second and third pleuroaxillary muscle overlapping
anterior section of pleurotergal and lateral promotor of middle
coxa. Pleurotergal muscle elongated dorsoventrally, but above
inferior level of third pleuroaxillary. Lateral promotor of middle
coxa V-shaped (as in Fig. 84). Metathoracic musculature similar
to that of Exomalopsis, but ventral lateromotor of abdomen arising
along ridge of vertical plate of metasternum; division of mesal
remotor of hind coxa not well marked; lateral promotor of hind
coxa not divided by a notch.
Tetrapedia maura Cresson
(Female)
Mesopleura musculature agrees with description of above species
(Fig. 34). Metathoracic musculature similar to that of Exomalopsis,
but mesal remotor of hind coxa apparently without division, and
lateral promotor of hind coxa not divided by a notch.
Treg NOMADINI
Nomada (Micronomada) sp.
(Male. San Luis Potosi, S. L. P., Mex., August 22, 1954)
In mesopleura third pleuroaxillary muscle not overlapping second
one, but overlapping anterior part of pleurotergal and lateral pro-
motor of middle coxa. Pleurotergal muscle quadrangular in shape
and above level of inferior end of third pleuroaxillary muscle.
Lateral promotor V-shaped, with wide inferior arm (Fig. 85). In
|
|
)
Srupies oF THE THoRACIC MusCULATURE OF BEES 459
metathorax median depressor of abdomen a ribbonlike muscle,
arising on coalesced furcal arms, Ventral lateromotor of abdomen
small muscle originating on anterior ridge of vertical plate of
metasternum. Mesal remotor of hind coxa divided, outer part on
mesal half of coalesced furcal arms and on free basal portion of
metafurcal arm, inner part on basal portion of metafurcal arm and
on vertical plate of metasternum, Thoracic depressor of hind tro-
chanter on free distal portion of metafurcal arm and on coalesced
furcal arms. Lateral promotor of hind coxa with small inner lobe.
Mesal promotor of hind coxa covering whole mesal portion of inter-
coxal lamella.
Tre NEOPASITINI
Neopasites sp.
(Female. Lawrence, Kan., June 4, 1954)
In mesopleura third pleuroaxillary muscle overlapping part of sec-
ond one anteriorly and part of pleurotergal posteriorly. Pleurotergal
muscle leaf-shaped. Lateral promotor of middle coxa fan-shaped,
with inferior fibers extended anteriorly (Fig. 86). In metathorax
median depressor of abdomen a rather ribbonlike muscle arising on
mesal part of free distal portion of metafurcal arm, and on coa-
lesced furcal arms. Ventral lateromotor of abdomen originating
along ridge of vertical plate of metasternum. Mesal remotor of
hind coxa divided, outer part on mesal half of coalesced furcal arms
and on free basal portion of metafurcal arm, inner part on basal
portion of metafurcal arm and an anterior ridge of mesal portion
of intercoxal lamella. Thoracic depressor of hind trochanter on
mesal part of free distal portion of metafurcal arm and on lateral
part of coalesced furcal arms. Lateral promotor of hind coxa a fan-
shaped muscle, without the outer lobe. Mesal promotor of hind
coxa not covering whole mesal portion of intercoxal lamella.
Tribe EPEOLINI
Triepeolus sp.
(Female, Mexico City, August 8, 1954)
In mesopleura third pleuroaxillary a large muscle, overlapping
part of second one anteriorly, and part of pleurotergal and lateral
promotor of middle coxa posteriorly. Pleurotergal muscle above
level of inferior end of third pleuroaxillary, Lateral promotor of
middle coxa V-shaped (Fig. 37). In metathorax median depressor
of abdomen a large triangular muscle arising on free distal portion
of metafurcal arm, and on coalesced furcal arms. Ventral latero-
460 THe Untiversiry SCIENCE BULLETIN
motor of abdomen a small muscle originating on anterior ridge of
vertical plate of metasternum. Mesal remotor of hind coxa divided,
outer part on free basal portion of metafurcal arm, inner part on
basal portion of metafurcal arm (Fig. 60). Thoracic depressor of
hind trochanter on free distal portion of metafurcal arm and on
coalesced furcal arms. Lateral promotor of hind coxa lobed. Mesal
promotor of hind coxa covering whole mesal portion of intercoxal
lamella.
Trae EUCERINI
Melissodes (Epimelissodes) obliqua Say
(Female )
In mesopleura third pleuroaxillary muscle overlapping anterior
part of pleurotergal and lateral promotor of middle coxa. Pleuro-
tergal muscle small and rounded, above level of inferior end of
third pleuroaxillary. Lateral promotor of middle coxa V-shaped,
with very wide inferior arm (Fig. 38). In metathorax median
depressor of abdomen a large triangular muscle arising on free
distal portion of metafurcal arm, and on coalesced furcal arms.
Ventral lateromotor of abdomen a rather triangular-shaped muscle
arising along ridge of vertical plate of metasternum. Mesal re-
motor of hind coxa divided, outer part on free distal portion of
metafurcal arm, on coalesced furcal arms, and on free basal por-
tion of metafurcal arm, inner part on basal portion of metafurcal
arm and anterior ridge of mesal portion of intercoxal lamella. Tho-
racic depressor of hind trochanter on free distal portion of meta-
furcal arm and on coalesced furcal arms. Lateral promotor of
hind coxa with outer and inner lobes separated by a notch. Mesal
promotor of hind coxa not covering whole mesal portion of inter-
coxal lamella.
Melissodes (Eumelissodes) agilis Cresson
(Female)
In metathorax third pleurotergal muscle overlapping only anterior
part of pleurotergal and lateral promotor of middle coxa, Pleuro-
tergal muscle rounded but larger than that of M. obliqua. Lateral
promotor of middle coxa V-shaped, but inferior arm not so wide
(as in Fig 40). Metathoracic musculature like that of M. obliqua.
Srupies or THE THoracic MuscuLATURE OF BEES 461
Melissodes (Eumelissodes) coloradensis Cresson.
(Female)
Mesopleura musculature somewhat similar to that of M. obliqua
(Fig. 39), but second pleuroaxillary larger than third one, and
lateral promotor of middle coxa with smaller inferior arm. Meta-
thoracic musculature agrees in detail with that of M. obliqua.
Melissodes (Eumelissodes) sp.
(Female. Lawrence, Kansas, September, 1954)
Musculature agrees. with description of above species
Melissodes (Melissodes) communis Cresson
(Female )
Mesopleura musculature like that of M. agilis (Fig. 40). Meta-
thoracic musculature like that of M. obliqua.
Melissodes (Melissodes) tepaneca
(Female)
Mesopleura musculature agrees in detail with that of M. communis
(as in Fig. 40). Metathoracic musculature like that of M. obliqua.
Florilegus condignus (Cresson )
(Female)
Mesopleura musculature agrees in detail with that of M. agilis
and M. communis (as in Fig. 40). Metathoracic musculature like
that of M. obliqua, but parts of mesal remotor of hind coxa widely
separated posteriorly (Fig. 62).
Thygater near analis Lepeletier and Serville
(Female, Ciudad del Maiz, S. L. P., Mex., August 22, 1954)
In mesopleura third pleuroaxillary muscle overlapping anterior
part of pleurotergal. Pleurotergal muscle rounded. Lateral pro-
motor of middle coxa fan-shaped but with a notch (Fig. 41).
Metathoracic musculature similar to that of M. obliqua, but lateral
promotor of hind coxa without inner lobe.
Peponapis sp.
(Female, Iguala, Guerrero, Mex., August 8, 1954)
In mesopleura third pleuroaxillary muscle overlapping posterior
part of second one and anterior part of pleurotergal. Pleurotergal
muscle elongated dorsoventrally, Lateral promotor of middle coxa
fan-shaped (Fig. 42). Metathoracic musculature similar to that
462 Tue Universiry ScrENCE BULLETIN
of M. obliqua, but outer part of mesal remotor of hind coxa arising
only on coalesced furcal arms, and on free basal portion of meta-
furcal arm.
Xenoglossa near fulva Smith
(Female. Iguala, Guerrero, Mex., August 8, 1954)
Mesopleura musculature differs from above species as shown in
Fig. 43. Lateral promotor of middle coxa with slender inferior
arm, superficial fibers of superior arm continuous with fibers of
pleurotergal muscle. Metathoracic musculature like that of M.
obliqua but outer part of mesal remotor of hind coxa arising only
on coalesced furcal arms, and on free basal portion of metafurcal
arm, as is in Peponapis.
Xenoglossodes spissa (Cresson )
(Female)
Mesopleura musculature agrees in detail with that of M. com-
munis (as in Fig. 40). Metathoracic musculature like that of M.
obliqua.
Tree EMPHORINI
In general, the musculature of the species dissected in this tribe
is very similar to that of the tribe Eucerini. The only constant
feature found in this group and not in the Eucerini, is that the inner
part of the mesal remotor of the hind coxa does not have any fibers
arising on the mesal portion of the intercoxal lamella. The reason
of this is that the mesal promotor of the hind coxa covers entirely
the mesal portion of the intercoxal lamella.
Ptilothrix sumichrastit. (Cresson)
(Female )
In mesopleura third pleuroaxillary muscle overlapping only an-
terior part of pleurotergal and lateral promotor of middle coxa.
Pleurotergal muscle rounded, Lateral promotor of middle coxa
V-shaped, with wide inferior arm (Fig. 44). In comparing the
metathoracic musculature (Fig. 61) with that of Melissodes obliqua
the following differences (besides the ones mentioned above) are
noted: thoracic depressor of hind trochanter on free distal portion
of metafurcal arm and on lateral half of coalesced furcal arms;
lateral promotor of hind coxa not divided by a notch.
Srupres or THE THorAcic MUSCULATURE OF BEES 463
Diadasia near diminuta (Cresson )
(Female. Carmen, Hidalgo, Mex., July 28, 1954)
Mesopleura musculature somewhat similar to that of Ptilothrix
sumichrastii (see Fig. 44 and 45). Metathoracic musculature differs
from that of Melissodes obliqua, besides ones mentioned above, in
that lateral promotor of hind coxa is not divided by a notch.
Diadasia enevata (Cresson)
(Female)
In mesopleura third pleuroaxillary muscle overlapping part of
second one and part of pleurotergal and lateral promotor of middle
coxa. Pleurotergal muscle rounded. Lateral promotor of middle
coxa V-shaped with very wide inferior arm (Fig. 46). The meta-
thoracic musculature differs from the description of Melissodes
obliqua, besides the ones mentioned above, as follows: ventral
lateromotor of abdomen on posterior ridge of verticle plate of meta-
sternum; lateral promotor of hind coxa not divided by a notch.
Melitoma euglossoides Lepeletier and Serville
(Female)
In mesopleura pleuroaxillary muscles like that of Diadasia (near
diminuta) (Fig. 45). Pleurotergal a rather rounded muscle. Lateral
promotor of middle coxa fan-shaped but inferior fibers shorter and
separated by a notch from superior fibers (Fig. 47). In comparing
the metathoracic musculature with that of Melissodes obliqua the
following points (besides the ones mentioned above) are noted:
outer part of mesal remotor of hind coxa arising on small mesal sec-
tion of free distal portion of metafurcal arm, on coalesced furcal
arms, and on free basal portion of metafurcal arm; lateral promotor
of hind coxa not divided by a notch, as all other forms in this tribe.
Trax ANTHOPHORINI
Anthophora californica texana Cresson
(Female)
In mesopleura third pleuroaxillary a large muscle, overlapping
more than half of second one anteriorly, and small section of
pleurotergal posteriorly, Pleurotergal muscle large and dorso-
ventrally elongated. Lateral promotor of middle coxa as in Meli-
toma englossoides, fan-shaped but with inferior fibers shorter than
superior ones (Fig. 48). In metathorax median depressor of ab-
domen a band-shaped muscle arising on coalesced furcal arms and
few fibers on mesofurcal bridge. Ventral lateromotor of abdomen
464 Tue Universtry Scrence BULLETIN
on anterior ridge of vertical plate of metasternum. Mesal remotor
of hind coxa not divided (Fig. 63), originating on coalesced furcal
arms and on free basal portion of metafurcal arm. Thoracic de-
pressor of hind trochanter on free distal portion of metafurcal arm,
occupying entire posterior surface of dorsal process, and on coa-
lesced furcal arms. Lateral promotor of hind coxa with outer and
inner lobes separated by a notch, as is in Melissodes and some
Exomalopsis. Mesal promotor of hind coxa almost covering whole
mesal portion of intercoxal lamella.
Anthopora (near californica) sp.
(Female. Ixmiquilpan, Mex., July 29, 1954)
The description of this species agrees with the previous one.
Tre HEMISINI
Centris near agilis Smith
(Female. Taxco, Guerrero, Mex., August 8, 1954)
In mesopleura third pleuroaxillary muscle overlapping only an-
terior part of pleurotergal. Pleurotergal muscle dorsoventrally
elongated, Lateral promotor of hind coxa fan-shaped (Fig. 49).
In metathorax median depressor of abdomen on coalesced furcal
arms. Ventral lateromotor of abdomen a small muscle arising on
posterior ridge of vertical plate of metasternum. Mesal remotor of
hind coxa divided, (Fig. 64) outer part on coalesced furcal arms
and on free basal portion of metafurcal arm, inner part on base
of metafurcal arm and on anterior ridge of mesal portion of inter-
coxal lamella. Thoracic depressor of hind trochanter on free distal
portion of metafurcal arm, occupying entire posterior surface of
dorsal process, and on coalesced furcal arms (as is in Anthophora
californica), Lateral promotor of hind coxa lobed. Mesal_ pro-
motor of hind coxa not covering whole mesal portion of intercoxal
lamella.
Centris lanosa (Cresson )
(Female)
This species agrees with the description of the above species ex-
cept that some fibers of the outer part of the median remotor of
the hind coxa are fused with the thoracic depressor of the hind
trochanter,
STUDIES OF THE THorACcIC MuscuLATURE OF BEES 465
Epicharis sp.
(Female, Cuernavaca, Morelos, Mex., August 4, 1954)
In mesopleura third pleuroaxillary muscle overlapping anterior
parts of pleurotergal and lateral promotor of middle coxa. Pleuro-
tergal muscle rounded. Lateral promotor of middle coxa V-shaped
(Fig. 50). In metathorax median depressor of abdomen on free
distal portion of metafurcal arm and on coalesced furcal arms.
Ventral lateromotor of abdomen a rather large muscle, arising
along ridge of vertical plate of metasternum, and some fibers on
free basal portion of metafurcal arm. Mesal remotor of hind coxa
not divided, but partially fused anteriorly with thoracic depressor
of hind trochanter (Fig. 66); it originates on coalesced furcal arms,
on free basal portion of metafurcal arm, and some fibers on anterior
tidge of mesal portion of intercoxal lamella. Thoracic depressor
of hind trochanter on free distal portion of metafurcal arm and
on lateral part of coalesced furcal arms. Lateral promotor of hind
coxa a large muscle (Fig. 65), with outer and inner lobes separated
by a notch, inner lobe large. Mesal promotor of hind coxa small
and arising on posterior part of vertical plate of metasternum,
some fibers on mesal portion of intercoxal lamella (Fig. 65).
Subfamily XyLocorinaE
TriBE XYLOCOPINI
Xylocopa fimbriata Fabricius
(Female)
In mesopleura third pleuroaxillary muscle overlapping almost
completely second one. Anterior part of pleurotergal and lateral
promotor of middle coxa overlapped by second and third pleuro-
axillary muscles. Pleurotergal muscle rounded and above level
of inferior end of third pleuroaxillary. Lateral promotor of middle
coxa V-shaped (Fig. 51). In metathorax median depressor of abdo-
men on free distal portion of metafurcal arm and on coalesced
furcal arms. Ventral lateromotor of abdomen originating along
ridge of vertical plate of metasternum. Mesal remotor of hind
coxa not divided and arising on coalesced furcal arms, on free
basal portion of metafurcal arm, and some fibers on anterior ridge
of mesal portion of intercoxal lamella; its insertion covers most
of posterior basal margin of hind coxa (Fig. 67). Thoracic de-
pressor of hind trochanter on free distal portion of metafurcal arm
and on coalesced furcal arms. Lateral promotor of hind coxa with
16—3378
466 Tur Universtry SCrENCE BULLETIN
outer and inner lobes separated by a notch. Mesal promotor of
hind coxa not covering whole mesal portion of intercoxal lamella.
Xylocopa brasilianorum Linnaeus
(Female)
The description of this species agrees with that of the above
species.
Trips CERATININI
Ceratina (Zadontomerus ) ignara Cresson
(Female)
In mesopleura third pleuroaxillary muscle overlapping part of
second one and part of pleurotergal and lateral promotor of middle
coxa. Pleurotergal muscle fan-shaped and above level of inferior
end of third pleuroaxillary. Lateral promotor V-shaped (Fig. 52).
In metathorax median depressor of abdomen a ribbonlike muscle
(fused posteriorly to its counter part (Fig. 69), arising on coalesced
furcal arms, some fibers on mesofurcal bridge. Ventral lateromotor
of abdomen originating along ridge of vertical plate of metasternum,
Mesal remotor of hind coxa a ribbonlike muscle (Fig. 68) arising
only on free basal portion of metafurcal arm. Thoracic depressor
of hind trochanter on free distal portion, occupying entire posterior
surface of dorsal process, and on coalesced furcal arms. Lateral
promotor of hind coxa with small inner lobe. Mesal promotor of
hind coxa not covering whole mesal portion of intercoxal lamella.
Ceratina (Zadontomerus) dupla Say
(Female)
This species agrees with the description of C. ignara.
Subfamily APINAE
Trine EUGLOSSINI
Euglossa (Euglossa) sp.
(Female. Zimapan, Hidalgo, Mex., July 27, 1954)
In mesopleura third pleuroaxillary muscle overlapping more than
half of second one. Second and third pleuroaxillary overlapping
anterior part of pleurotergal and lateral promotor of middle coxa.
Pleurotergal a rather rounded and large muscle. Lateral promotor
of middle coxa V-shaped (Fig. 53). In metathorax median de-
pressor of abdomen on free distal portion of metafurcal arm and
on coalesced furcal arms. Ventral lateromotor of abdomen arising
along ridge of vertical plate of metasternum, Mesal remotor of
Srupies OF THE THORACIC MUSCULATURE OF BEES 467
hind coxa not divided (Fig. 71) and somewhat fused anteriorly
with thoracic depressor of hind trochanter, originating on coa-
lesced furcal arms, on free basal portion of metafurcal arm, and on
anterior ridge of mesal portion of intercoxal lamella. Thoracic de-
pressor of hind trochanter on free distal portion of metafurcal arm
and on coalesced furcal arms. Lateral promotor of hind coxa lobed.
Mesal promotor of hind coxa not covering whole mesal portion of
intercoxal lamella.
Eulaema mussitans (Fabricius)?
(Female )
Musculature like that of Euglossa.
Tre BOMBINI
The species dissected in this group exhibit similar musculature
with that of the tribe Englossini.
Bombus (Fervidobombus) medius Cresson
(Female)
Mesopleura musculature differs from that of Euglossa as shown in
Fig. 54. In comparing the metathoracic musculature with that of
Euglossa the following differences are noted: mesal remotor of
hind coxa with narrower insertion on posterior basal margin of
hind coxa, and entirely fused with anterior part of thoracic de-
pressor of hind trochanter. (Fig. 70).
Bombus (Fraternobombus) fraternus Smith
(Male and female)
Musculature like that of B. medius.
Bombus (Pyrobombus) ephippiatus Say
(Worker and queen)
Musculature as in B. medius but mesal remotor of hind coxa not
so fused with thoracic depressor of hind trochanter.
Bombus sp.
(Female. Zimapan, Hidalgo, Mex., July 27, 1954)
Musculature of this species agrees with that of B. medius, but
mesal remotor of hind coxa not so fused with thoracic depressor
of hind trochanter, as is in B. ephippiatus.
Tur Universiry SCiENCE BULLETIN
Bombus sp.
(Female. Lawrence, Kan., June, 1953)
Musculature like that of B. medius.
Tre MELIPONINI
All the species dissected exhibit very homogeneous type of mus-
culature. The species dissected are as follows: Melipona beechei
Bennett (female), Trigona (Partamona) testacea (Klug) female,
Trigona (Cephalotrigona) capitata Smith female, Trigona (Tetra-
gona) jaty Smith (female), Lestrimelitta (female Bolivia).
Melipona beechei Bennett
In mesopleura second and third pleuroaxillary muscles overlap-
ping anterior part of pleurotergal. Pleurotergal a large muscle
shaped as shown in Fig. 55. Lateral promotor of middle coxa fan-
shaped. In metathorax (Fig. 72) median depressor of abdomen
a ribbonlike muscle originating on mesal half of coalesced furcal
arms. Ventral lateromotor of abdomen small muscle arising on
free basal portion of metafurcal arm. Mesal remotor of hind coxa
on mesal half of coalesced furcal arms and on free basal portion of
metafurcal arm, some fibers are fused anteriorly with thoracic de-
pressor of hind trochanter. Thoracic depressor of hind trochanter
on free distal portion of metafurcal arm and on lateral half of
coalesced furcal arms. Lateral promotor of hind coxa lobed. Mesal
promotor of hind coxa covering whole mesal portion of intercoxal
lamella.
Trine APINI
Apis mellifica Linnaeus
(Male)
In mesopleura third pleuroaxillary overlapping posterior part of
second one and anterior part of pleurotergal. Pleurotergal muscle
dorsoventrally elongated. Lateral promotor of middle coxa a slen-
der muscle, (only superior arm) on pleural apophysis (Fig. 56).
In metathorax median depressor of abdomen ribbonlike, on free
basal portion of metafurcal arm and on mesal section of coalesced
furcal arms. Ventral lateromotor of abdomen on anterior ridge
of vertical plate of metasternum. Mesal remotor of hind coxa on
free basal portion of metafurcal arm, some fibers on coalesced
furcal arms. Thoracic depressor of hind trochanter on lateral half
of coalesced furcal arms, some fibers on free distal portion of meta-
STUDIES OF THE THoracic MuscuLATURE OF BEES 469
furcal arm. Lateral promotor of hind coxa a thin muscle almost
covering mesal portion of intercoxal lamella.
Among the thirty specimens dissected one male was found with
the lateral promotor of middle coxa possessing a small inferior arm
(Fig. 57).
Apis mellifica L.
(Worker )
The worker differs from that of the male in the mesopleura mus-
culature (Fig. 58). Second and third pleuroaxillary muscles over-
lapping anterior part of pleurotergal. Pleurotergal muscle wider.
Lateral promotor of middle coxa V-shaped.
LITERATURE CITED
Duncan, C. D.
1939. A contribution to the biology of North American vespine wasps.
Stanford Univ. Publ., Sci., vol. 8, no. 1, 272 pp., 54 pls.
Maxi, TAKapr,
1938. Studies on the thoracic musculature of insects, Taihoku Imperial
Univ., Formosa, Japan. Memoir vol. 34, no. 1, 343 pp., 17 pls.
MicHEnrr, CHar.es D,
1944. Comparative external morphology, phylogeny, and a classification
of the bees (Hymenoptera). Bull. Amer. Mus, Nat. Hist., vol. 82,
pp. 151-826, 246 figs.
1953. Comparative morphological and systematic studies of bee larvae
with a key to the families of hymenopterous larvae, Kansas Univ.
Sci. Bull., vol. 35, no, 8, pp. 987-1102, 287 figs.
Morison, G, D.
1927. The muscles of the adult honey-bee (Apis mellifera L.). Quart.
Journ. Mier. Sci., vol. 71, pp. 895-463, 12 figs.
Snoperass, R. E.
1942. The skeleto-muscular mechanisms of the honey bee. Smithsonian
Misc. Coll., vol, 103, no. 2, 120 pp., 32 figs.
Tur Untiversrry SCIENCE BULLETIN
Fic. 1, Trends of specialization of the lateral promotor of the middle coxa.
a—as in Colletes sp. (female).
b—as in Pseudopanurgus sp. (female).
c—as in Neopasites sp. (female).
d—as in Dianthidium sayi (female).
e—as in Xenoglossa sp. near fulva (female).
fas in Apis mellifica (worker).
g—as in Apis mellifica (male, see page 469).
h—as in Apis mellifica (male).
i—as in Diadasia near diminuta (female).
j—as in Melissodes obliqua (female). |
Fic. 2. Trends of specialization of the lateral promotor of the hind coxa.
a—as in Megachile sp. (female).
b—as in Heriades variolosa purpurascens (male).
c—as in Ashmeadiella opuntiae (male). '
d—as in Andrena argemonis (male). |
e—as in Protoxaea nigerrima (female).
f—as in Exomalopsis snowt (female).
g—as in Epicharis sp. (female).
h—as in Apis mellifica (worker).
'
StuptEes or THE THoractc MuscCULATURE OF BEES
“’== = pleural apophysis
- = superior arm
outer lobe of =~ |
) -- inner lobe of
pase plete Oe jin eottihmnanecienie
4
é
ih
Tue Universiry SCIENCE BULLETIN
MrsopLeurAL Muscacarure (Right Side)
Fic. 3. Colletes sp. (female).
Fic. 4. Halictus ligatus (female ).
Fic. 5. Dufourea marginata (female).
Fic. 6. Nomia nortoni (male).
Nomia micheneri (male) 1, 2a muscles omitted.
’ STUDIES OF THE THoRACIC MUSCULATURE OF BEES 473
third pleuroaxillary muscle
4
first pleuroaxillary muscle ~-.-----
, -- + == pleurotergal muscle
i second pleuroaxillary muscle ----- - - P §
- = lateral promotor of the middle coxe
basalar muscle ~--- - —
Tur Unriversrry SCIENCE BULLETIN
MrsopLEURAL MuscuLature (Right Side )
Nomia (ieunomia) sp. (female).
Andrena argemonis (male).
Perdita sp. (female).
Pseudopanurgus sp. (female ).
Calliopsis near andreniformis (female).
Protoxaea nigerrima (female).
Tur Universrtry SCIENCE BULLETIN
MrsopLEurAL Muscuature (Right Side )
Megachile sp. (female) 1, 2a muscles omitted.
. Coelioxys sp. (female) 1, 2a muscles omitted.
. Ashmeadiella opuntiae (male) 1, 2a muscles omitted.
. Heriades variolosa purpurascens (male).
. Anthidium maculosum (female).
Dianthidium sayi (female).
Lithurge sp. (female).
STUDIES OF THE THORACIC MUSCULATURE OF BEES 477
We 18
Tue University ScreNCE BULLETIN
Merarnoracic Muscu.ature (Left Side)
Fic. 21. Ptiloglossa mexicana (female). Median depressor of the aldo-
men and lateral remotor of the hind coxa omitted.
Frc. 22. Nomia nortoni (male). Median depressor of the abdomen, ventral
lateromotor of the abdomen, and lateral remotor of the hind coxa omitted.
Fic. 28, Andrena argemonis (male). Median depressor of the abdomen,
ventral lateromotor of the abdomen, and lateral remotor of the hind coxa
omitted.
SruprEs oF THE THorRACIC MuSsCULATURE OF BEES 479
dorsal process of free distal metafurcal arm coalesced furcal arms
; ;
NE net etal . _ free basal portion of metafurcal arm
.inner part of mesal remotor of the hind coxa
-origin of mesal promotor of the hind coxa
~~ vertical plate of metasternum
~ ventral lateromotor of the abdomen
\
I outer part of mesal remotor of the hind coxa
’
yf ~origin of mesal promotor of the hind coxa
lateral promotor of the hind coxa... . —
¥- - ~ verticol plate of metasternum
outer part of mesal remotor of the hind coxa-— ~~ - = ~
Tur Universtry SCIENCE BULLETIN
Meratruoracic Muscutature (Left Side)
Fic. 24. Pseudopanurgus sp. (female). Median depressor of the abdomen,
ventral lateromotor of the abdomen, and lateral remotor of the hind coxa
omitted.
Fic. 25. Protoxaea nigerrima (female). Promotors of the hind coxa.
Fic. 26. Megachile sp. (female). Median depressor of the abdomen, ven-
tral lateromotor of the abdomen, and lateral remotor of the hind coxa omitted. ‘
Fic. 27. Anthidium maculosum (female). Median depressor of the abdo-
men, ventral lateromotor of the abdomen, and lateral remotor of the hind coxa
omitted.
Fic. 28. Megachile sp. (female), Promotors of the hind coxa.
Srupies oF THE THORACIC MUSCULATURE OF BEES 481
jude remotor of the hind coxa mesal promotor of the hind coxa
outer part of mesal remotor of the hind coxa
inner part of mesal remotor of the hind coxa
-
lateral promotor
of the hind coxa
lateral promotor _
of the hind coxa
lateral promotor of the hind coxa---— — - H- - mesal promotor of the hind coxa
intercoxal lamella - - -- —-
University SCIENCE BULLETIN
MEsopLEuRAL MuscuLature (Right Side)
Exomalopsis sp. (female).
Exomalopsis sp. (female).
Ancyloscelis armatus (female ).
Paratetrapedia sp. (male).
Paratetrapedia sp. (male).
Tetrapedia maura (female).
Tse Universiry SCIENCE BULLETIN
MesopLeuraL MuscuLature (Right Side)
Fic. 385. Nomada (Micronomada) sp. (male).
Fic. 86. Neopasites sp. (female).
Fic. 87. Triepeolus sp. (female).
Fic. 88. Melissodes obliqua (female).
Fic. 89. Melissodes coloradensis (female).
. Melissodes communis (female).
36
Ws
Fic.
Fic;
Fie.
Fic.
Fig,
THe Universtry ScieNCE BULLETIN
MrsopLeurAL Muscutature (Right Side)
Thygater near analis (female).
Peponapis sp. (female).
Xenoglossa near fulva (female).
Ptilothrix sumichrastii (female).
Diadasia near diminuta (female).
Diadasia enevata (female ).
SrupiEs OF THE THORACIC MUSCULATURE OF BEES
487
EtG,
Fic.
Fic.
Fic.
Fic,
Tue Universiry SCIENCE BULLETIN
Mrsor.eurat Muscutatune (Right Side )
Melitoma euglossoides (female) 1, 2a, 2b and 2c muscles omitted.
Anthophora californica Texana (female ).
Centris near agilis (female).
Epicharis sp. (female ).
Xylocopa Fimbriata (female).
Ceratina ignara (female).
489
STUDIES OF THE THORACIC MUSCULATURE OF BEES
Fic.
Fic.
Fic,
Fic.
Fic.
THe Universrry Science BULLETIN
MEsopLEuRAL Muscutarure (Right Side)
Euglossa (Euglossa) sp. (female). 1 muscle omitted,
Bombus medius (female),
Melipona beechei (female).
Apis mellifica (male). 2a muscle omitted.
Apis mellifica (male, see page 469). 2a muscle omitted.
Apis mellifica (worker). 2a muscle omitted.
Srupres oF THE THORACIC MusCULATURE OF BEES 49]
Tue University SCIENCE BULLETIN
MeraTrHoracic MuscuLatTure (Left Side)
Fic. 59. Exomalopsis sp. (female). Median depressor of the abdomen
omitted.
Fic. 60. Triepeolus sp. (female). Median depressor of the abdomen
omitted.
Fic. 61. Ptilothrix sumichrastii (female). Median depressor of the abdo-
men and ventral lateromotor of the abdomen omitted.
Fic. 62. Florilegus condignus (female). Median depressor of the abdo-
men and ventral lateromotor of the abdomen omitted.
Fic. 68. Anthophora californica (female). Median depressor of the abdo-
men and lateral remotor of the hind coxa omitted.
Nore: Dotted lines connecting figures 60 and 62 refer to inner part of mesal
remotor of the hind coxae.
Strupirs OF THE THORACIC MUSCULATURE OF BEES 493
thoracic depressor of
the hind trochanter \
lateral remotor
of the hind coxa
ris
N
‘ventral lateromotor of the abdomen
|
|
\
|
|
|
,
}
thoracic depressor of
the hind trochanter”
S
Smesal remotor of the hind coxa
Tue Universiry ScreNCE BULLETIN
Mrrarnoracic MuscuLature (Left Side)
Fic. 64. Centris near agilis (female). Median depressor of the abdomen,
ventral lateromotor of the abdomen, and lateral remotor of the hind coxa
omitted.
Fic. 65. Epicharis sp. (female). Promotors of the hind coxa.
Fic. 66. Epicharis sp. (female). Median depressor of the abdomen and
ventral lateromotor of the abdomen omitted.
Fic. 67. Xylocopa fimbriata (female). Median depressor of the abdomen
and ventral lateromotor of the abdomen omitted.
StupiEs OF THE THORACIC MUSCULATURE OF BEES 495
outer part of mesal remotor of the hind coxa
‘
! inner part of mesal remotor of the hind coxa
/
thoracic depressor of the hind trochanter it 7
*
mesal promotor of the hind coxo
|. thoracic depressor of the hind trochanter
|... — -mesal remotor of the hind coxa
lateral remotor of the hind coxa —=
~ -origin of mesal promotor of the hind coxa
~ =|~ -.mesal remotor of the hind coxa
Tue Universrry ScreENCE BULLETIN
MeEtTaTHORACIC MUSCULATURE
Fic. 68. Ceratina ignara (female). Left side: Promotors of the hind
coxa. Right side: Median depressor of the abdomen omitted.
Fic. 69. Ceratina ignara (female).
STUDIES OF THE THoRACIC MusCULATURE OF BEES 497
mesal remotor of the hind coxa
,
mesal promotor of the hind coxa
17—3378
Tue Universrry SCIENCE BULLETIN
METATHORACIC MUSCULATURE
Fic. 70. Bombus medius (worker).
Fic. 71. Euglossa (Euglossa) sp. Left side: Median depressor of the \
abdomen and ventral lateromotor of the abdomen omitted.
Fic. 72. Melipona beechei. Lateral remotor of the hind coxa omitted,
STUDIES OF THE THORACIC MUSCULATURE OF BEES 499
median depressor of the abdomen
/
- ~~ lateral promotor of the hind coxa
- ~~ —~mesal remotor of the hind coxa
~ — = thoracic depressor of the hind trochanter
Se | lateral remotor of the hind coxa
~* ventral lateromotor of the abdomen
70
lateral remotor of the hind coxo — .. . um frcal remotor of the Wind coxa
thoracic depressor of the hind trochanter! 71
lateral promotor of the hind coxa ~~.
~~ mesal remotor of the hind coxa
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVITI, Pr. 1] = Decemper 20, 1956 [No. 7
Catalogue of the Types in the Snow Entomological
Museum. Part I (Hymenoptera)
BY
Watuace FE. LaBercr *
Axsstracr: This paper is the first part of a catalogue of the type specimens
in the Snow Entomological Museum of the University of Kansas. Types of
746 species of Hymenoptera are listed with references, notes on the condition
of the types, and type numbers as recorded in the catalogue of the types in the
Snow Entomological Museum.
INTRODUCTION
This paper contains a list of the type specimens of Hymenoptera
in the Snow Entomological Museum of the University of Kansas
arranged alphabetically under each family or superfamily. The
types are placed in two lists beneath each family or superfamily,
except in cases in which only one to three species represent the
group. First, the holotypes, allotypes, cotypes and _lectotypes
are listed together with references to the papers in which these
were originally described. Second, the paratypes are listed with-
out references. The names and references cited in both lists have
all been checked for accuracy.
The term cotype is used only in reference to a member of a
syntypical series. Some authors in the past have used the term
cotype as synonymous with paratype. An attempt has been made
to discover all such cases and such specimens are listed as paratypes.
Since it is the hope of the author that all of the types in the
Snow Museum will be catalogued in this and subsequent papers,
the list contains only a minimum of information concerning the
specimens. Mention is made of lost, destroyed or seriously dam-
aged holotypes, allotypes and lectotypes. The sex, the number
@ type specimens, and the Snow Entomoisical a ata vee
* Contribution No. 916 of the Department of stanton: Greve of Kansas, Law-
rence, Kansas,
(501)
502 Tue Universiry SCIENCE BULLETIN
number, as recorded in the catalogue of types in the Snow Ento-
mological Museum, are listed. Secondary types, such as metatypes,
topotypes and homotypes, are omitted from this list and are not
recorded in the catalogue of types in the Snow Entomological
Museum. No notice is taken of synonymy; the genera under which
the species are listed are those under which they were originally
described. Lectotypes have not been designated in this catalogue,
since the author is of the opinion that such designations should
be made only by the author of the species in question or by some
competent authority in the group involved.
ACKNOWLEDGMENTS
The author is grateful to the following persons who have gra-
ciously taken the time to answer questions regarding references to
certain names: R. M. Bohart, University of California at Davis; B. D.
Burks, U.S. N. M.; H. E. Evans, Cornell University; Karl V. Krom-
bein, U.S.N.M. The author also wishes to thank the University
of Kansas for providing aid from its general research fund for
the purpose of cataloguing the types in the Snow Entomological
Museum.
Types of 746 species, of which 373 are represented only by para-
types, are listed in this part of the catalogue.
APOIDEA
Andrena collitoides Viereck and Cockerell, 1914, Proc. U. S. Nat.
Mus., vol. 48, p. 27. @ holotype. 4284.
Andrena kraussi Michener, 1954, Bull. Amer. Mus. Nat. Hist., vol.
104, p.30. ¢ holotype. 4285.
Andrena submariae Viereck and Cockerell, 1914, Proc. U. S. Nat.
Mus., vol. 48, p. 4. ? holotype. 4288.
Anthocopa beameri Michener, 1951, Pan-Pacific Ent., vol. 27, p. 64.
9 holotype, ¢ allotype, 2 ¢ and 15 ¢g paratypes. 5293.
Anthocopa daleae Michener, 1951, Pan-Pacific Ent., vol. 27, p. 62.
@ holotype, g allotype,2 9 and1 g paratypes. 4394,
Anthocopa hurdiana Michener, 1954, Pan-Pacific Ent., vol. 30, p. 47.
2 holotype. 5295.
Anthocopa mirifica Michener, 1954, Pan-Pacific Ent., vol. 80, p. 51.
2 holotype, 10 @ paratypes. 5296.
Anthocopa rubrella macswaini Michener, 1954, Pan-Pac. Ent., vol.
30, p. 45. ¢ holotype, ¢ allotype, 1 9 and 2 ¢ paratypes.
5298.
CATALOGUE OF TypPEs IN SNow MusEUM 503
Anthocopa rubrella rubrior Michener, 1954, Pan-Pac. Ent., vol. 80,
p-46. @ holotype, ¢ allotype. 5299.
Anthocopa segregata Michener, 1954, Pan-Pac. Ent., vol. 80, p. 48.
? holotype. 5300.
Ashmeadiella altadenae Michener, 1936, Pan-Pac. Ent., vol. 12, p. 63.
g holotype. 5271.
Ashmeadiella clypeodentata simplicior Michener, 1951, Pan-Pac.
Ent., vol. 27, p. 70. holotype, 1 9 paratype. 4275.
Ashmeadiella cockerelli Michener, 1936, Pan-Pac. Ent., vol. 12,
p. 62. ¢ holotype. 5272.
Ashmeadiella gillettei rubra Michener, 1951, Pan-Pac. Ent., vol. 27,
p. 67. ¢ holotype,7 g paratypes. 4269.
Ashmeadiella lateralis Michener, 1936, Pan-Pac, Ent., vol. 12, p. 60.
& holotype. 5270.
Ashmeadiella timberlakei Michener, 1936, Pan-Pac, Ent., vol. 12,
p.56. 9 allotype,3 ¢ and8 ¢% paratypes, 4364.
Ashmeadiella truncativentris Michener, 1951, Pan-Pac. Ent., vol. 27,
p. 68. 4366.
Augochlora coloradensis Titus, 1901, Canad. Ent., vol. 33, p. 188.
@ cotype. 2223.
Ceratina glossata Michener, 1954, Bull. Amer. Mus. Nat. Hist., vol.
104, p. 150. 9 holotype, 1 ¢ paratype. 5803.
Ceratina kraussi Michener, 1954, Bull. Amer, Mus. Nat. Hist., vol.
104, p. 153. 9 holotype, 12 ? paratypes. 5304.
Ceratina nigritula Michener, 1954, Bull. Amer, Mus. Nat. Hist.,
vol. 104, p. 153. 9 holotype, ¢ allotype,1 g paratype. 5305.
Ceratina rectangulifera Schwarz and Michener, 1954, Bull. Amer.
Mus. Nat. Hist., vol. 104, p. 154. 9 holotype, 11 ¢ paratypes.
5306,
Chelostoma bernardinum Michener, 1988, Pan-Pac, Ent., vol. 14,
p. 40. ¢ allotype,3 9? paratypes. 4319.
Chelostoma marginatum incisuloides Michener, 1954, Jour. Kansas
Ent. Soc., vol. 27, p.69. ¢ holotype, 3 allotype. 5269.
Colletes aridus Stephen, 1954, Univ. Kansas Sci. Bull., vol. 36, p. 494.
g holotype, ¢ allotype,1 g and 1 ¢ paratypes. 5209.
Colletes beamerorum Stephen, 1954, Univ. Kansas Sci. Bull., vol. 86,
p. 465. g holotype,5 ¢ paratypes. 5208.
Colletes bulbotibialis Stephen, 1954, Univ. Kansas Sci. Bull., vol. 36,
p. 458. ¢ holotype. 5212.
Colletes fulgidus longiplumosus Stephen, 1954, Univ. Kansas Sci.
Bull., vol. 86, p. 248. 5214.
504 Tue Untversrry SCIENCE BULLETIN
Colletes kansensis Stephen, 1954, Univ. Kansas Sci. Bull., vol. 86,
p. 297. g holotype. 5217.
Colletes micheneri Stephen, 1954, Univ. Kansas Sci. Bull., vol. 36,
p. 357. g holotype, ¢ allotype. 5207.
Colletes nigrifrons Titus, 1900, Canad. Ent., vol. 82, p. 804. 9
neotype selected by Stephen, 1954, Univ. Kansas Sci. Bull., vol.
36, p. 429. 9 neotype, ¢ neoallotype. 5219.
Colletes punctipennis maurus Stephen, 1954, Univ. Kansas Sci.
Bull., vol. 36, p. 232. g holotype. 5218.
Colletes saritensis Stephen, 1954, Univ. Kansas Sci. Bull., vol. 36,
p. 341. g holotype, ? allotype,1 ¢ and 20 9 paratypes. 5216.
Colletes swenki Stephen, 1954, Univ. Kansas Sci. Bull., vol. 86, p.
450. g holotype, @ allotype,6 g and 2 ¢ paratypes. 5211.
Colletes timberlakei Stephen, 1954, Univ. Kansas Sci. Bull., vol. 36,
p. 290. ¢g holotype,1 g paratype. 5210.
Diadasia afflicta perafflicta Cockerell, 1905, Amer. Nat., vol. 39,
p. 744. g holotype, 1 g¢ paratype. 1027.
Exomalopsis mourei Michener, 1954, Bull. Amer, Mus, Nat. Hist.,
vol. 104, p. 106. @ holotype, ¢ allotype, 35 9 and 18 ¢g para-
types. 5286.
Exomalopsis snowi Cockerell, 1906, Ann, Mag. Nat. Hist., ser. a
vol. 18, p. 73. g¢ holotype. 1033.
Florilegus isthmicus Michener, 1954, Bull. Amer. Mus. Nat. Hist.,
vol. 104, p. 134. @ allotype,1 ¢ paratype. 4393.
Halictoides altidenae Michener, 1937, Ann. Mag. Nat. Hist., ser. 10,
vol. 19, p. 3895. ¢g allotype,2 9 paratypes. 4337.
Halictoides nemophilae Michener, 1937, Ann. Mag. Nat. Hist., ser.
10, vol. 19, p. 400. ¢@ allotype, 1 ¢ and 8 9 paratypes. 4349.
Hdlictoides rhamni Michener, 1937, Ann. Mag. Nat. Hist., ser. 10,
vol. 19, p. 394. @ allotype,2 9 paratypes. 4348,
Halictus bernardinensis Michener, 1936, Ann. Mag. Nat. Hist., ser.
10, vol. 18, p. 286. @ holotype. 4826.
Heriades micropthalma Michener, 1954, Jour, Kansas Ent. Soc.,
vol. 28, p. 66. ¢ holotype, g allotype,1 ¢ and 10 ¢ para-
types. 5509.
Hesperapis parva Michener, 1937, Ann. Mag. Nat. Hist., ser. 10,
vol. 19, p.327. ¢ allotype,1 g paratype. 4352.
Hoplitis mazourka Michener, 1954, Pan-Pac, Ent., vol. 30, p. 38.
& holotype. 5273.
Lasioglossum kraussi Michener, 1954, Bull. Amer. Mus. Nat. Hist.,
vol. 104, p.50. @ holotype, 1 9 paratype. 5264,
CATALOGUE OF Types IN SNow Museum 505
Lasioglossum strigosigena Michener, 1954, Bull. Amer. Mus, Nat.
Hist., vol. 104, p. 47. 2 holotype, ¢ allotype,3 g and 8 9
paratypes. 5265.
Leptometria pacifica Cockerell, 1914, Jour. New York Ent. Soc., vol.
22,p.319. ¢ holotype, 2 ¢ paratypes. 4482.
Megachile gualanensis Cockerell, 1912, Ann. Mag. Nat. Hist., ser. 8,
vol. 10, p. 27. 2 holotype, 4 @ and 8 ¢ paratypes. 3349.
Megachile pruina nigropinguis Mitchell, 1937, Amer. Ent. Soc,
Trans., vol. 63, p.68. 9 holotype. 1014.
Melissodes aegis LaBerge, 1956, Univ. Kansas Sci, Bull., vol. 37,
p. 959. ¢ holotype, 9 allotype,3 g and4 9 paratypes. 4430.
Melissodes atripes atrimitra LaBerge, 1956, Univ. Kansas Sci. Bull.
vol. 37, p. 949. @ holotype, 9 allotype,8 ¢ and 1 9 paratypes.
4431.
Melissodes bimaculata nulla LaBerge, 1956, Univ. Kansas Sci. Bull.
vol. 37, p. 1109. g holotype, ¢ allotype, 4 g and 5 9 para-
types. 5221.
Melissodes blanda LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 1084. ¢ holotype, @ allotype, 6 ¢ paratypes. 5220.
Melissodes cubensis LaBerge, 1956, Univ. Kansas Sci. Bull. vol. 37,
p. 1078. g holotype, 1 g paratype. 4415.
Melissodes duplocincta Cockerell, 1905, Psyche, vol. 12, p. 103.
Lectotypes designated by LaBerge, 1956, Univ. Kansas Sci. Bull.,
vol. 37, p. 1031. g lectotype, ¢ lectoallotype. 1015.
Melissodes elusa LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 1061. ¢ holotype, ? allotype,1 9 and10 g paratypes. 4440.
Melissodes epicharina Cockerell, 1905, Psyche, vol. 12, D,..o9... 2
holotype. 1016.
Melissodes flexa LaBerge, 1956, Univ. Kansas Sci. Bull. vol. 37,
p. 1064. g holotype, 3 g paratypes. 4455.
Melissodes galvestonensis Cockerell, 1905, Proc. Biol. Soc. Wash-
ington, vol. 18, p. 181. Lectotypes designated by LaBerge, 1956,
Univ. Kansas Sci. Bull., vol 37, p. 1185. 9 lectotype, g lecto-
allotype. 1017. :
Melissodes gilensis crenata LaBerge, 1956, Univ. Kansas Sci. Bull.,
vol. 37, p. 1150. holotype, ¢ allotype,5 ¢ and 9 9 para-
types. 5500.
Melissodes herricki Cockerell, 1905, Psyche, vol. 12, p. 98. 9 co-
type, g¢ cotype. 1018.
Melissodes hexacantha Cockerell, 1905, Psyche, vol. 12, p. 100. é
holotype. 1019.
506 Tue Untiverstry SCrENCE BULLETIN
Melissodes hortivagans Cockerell, 1905. Proc. Biol. Soc. Washing-
ton, vol. 18, p. 180. g holotype. 1020.
Melissodes minima LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 981. g holotype, ¢ allotype, 8 ¢ paratypes. 4274,
Melissodes nigrosignata Cockerell, 1905, Psyche, vol. 12, p. 101.
9 holotype. 1021.
Melissodes nigrosignata pallidisignata Cockerell, 1905, Proc, Biol.
Soc. Washington, vol. 18, p. 180. ¢ holotype. 1022.
Melissodes nitida LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 969. g holotype, allotype, 2 g paratypes. 5502.
Melissodes praelauta Cockerell, 1905, Psyche, vol. 12, p. 102. ¢
holotype. 1023.
Melissodes semitristis Cockerell, 1905, Psyche, vol. 12, p. 102. ¢
holotype. 1024.
Melissodes sila LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37, p.
1002. ¢ holotype, ? allotype, 3 g¢ and 4 9? paratypes. 5504.
Melissodes tenuitarsis Cockerell, 1905, Psyche, vol. 12, p. 99. ¢
holotype. 1025.
Melissodes tepida yumensis LaBerge, 1956, Univ. Kansas Sci. Bull.,
vol. 37, p. 1128. ¢ holotype, ¢ allotype, 2 g¢ and 8 Q para-
types. 5508.
Melissodes tessellata LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 37,
p. 1157. g holotype, ¢ allotype, 86 g and 19 9 paratypes, 5505.
Melissodes thelypodii stulta LaBerge, 1956, Univ. Kansas Sci. Bull.
vol. 37, p. 1073. g¢ holotype, ¢ allotype, 4 g¢ and 3 9 para-
types. 5506.
Neocorynura nuda Michener, 1954, Bull. Amer. Mus. Nat. Hist.,
vol, 104, p. 81. ¢ holotype. 5257,
Neopasites carinatus Linsley, 1943, Trans. Amer. Ent. Soc., vol. 69,
p. 137. ¢ holotype. 3450.
Nomadopsis fracta Rozen, 1952, Jour. Kansas Ent. Soc., vol, 25,
p. 144. g holotype, ? allotype, 24 ¢ and 36 @ paratypes. 5206.
Osmia calcarata White, 1952, Univ. Kansas Sci. Bull., vol. 35, p. 245.
g holotype, ¢ allotype,3 g and3 9 paratypes. 4490.
Osmia crenulaticornis Michener, 1936, Southern California Acad.
Sci, Bull. vol. 35, p. 84. g holotype. 4491].
Osmia elongata Michener, 1936, Canad. Ent., vol. 68, p. 41. 2
holotype. 5279.
Osmia hurdi White, 1952, Univ. Kansas Sci. Bull., vol. 35, p. 273.
? holotype, 14 9 paratypes. 4493.
CATALOGUE oF TypEs IN SNow Museum 507
Osmia integra nigrigena Michener, 1936, Bull. Southern California
Acad, Sci., vol. 35, p. 90. g holotype. 4494.
Osmia obliqua White, 1952, Univ. Kansas Sci. Bull., vol. 85, p. 288.
& holotype, 8 g paratypes. 4489,
Panurginus bilobatus Michener, 1987, Ann. Mag. Nat. Hist., ser. 10,
vol. 19, p. 322. 9 allotype,4 9 and8 ¢ paratypes. 4298.
Perdita bidenticauda Timberlake, 1953, Univ. Kansas Sci, Bull., vol.
35, p. 965. g holotype, ¢@ allotype,38 g and 1 paratypes.
5230.
Perdita coreopsidis kansensis Timberlake, 1953, Univ. Kansas Sci.
Bull., vol. 35, p. 984. holotype, ¢ allotype, 1 ¢ and 1 ¢ para-
types. 5231.
Perdita coreopsidis obscurior Timberlake, 1953, Univ. Kansas Sci.
Bull., vol. 85, p. 984. ¢ holotype, g allotype, 21 @ and 65 ¢
paratypes. 5232,
Perdita fracticincta Timberlake, 1953, Univ. Kansas Sci. Bull., vol.
35, p. 976. 9 holotype. 5234.
Perdita laticauda Timberlake, 1953, Univ. Kansas Sci. Bull., vol. 35,
p. 967. ¢ holotype, ¢ allotype, 10 g and 12 9 paratypes.
5238.
Perdita lepachidis levifrons Timberlake, 1953, Univ. Kansas Sci.
Bull., vol. 35, p. 983. @ holotype, 3 allotype, 8 9° and38 g
paratypes. 5237.
Perdita lobata Timberlake, 1953, Univ. Kansas Sci, Bull., vol. 35,
p. 967. g holotype. 5224,
Perdita peculiaris Timberlake, 1953, Univ. Kansas Sci. Bull., vol. 85,
p.970. ¢ holotype. 5227.
Perdita perpulchra flavidior Timberlake, 1953, Univ. Kansas Sci.
Bull., vol. 35, p.975. 9 holotype. 5238.
Perdita scopata Timberlake, 1953, Univ. Kansas Sci. Bull., vol. 85,
p. 980. 9 holotype, g allotype, 839 9 and 95 g paratypes.
5241.
Perdita tricincta Timberlake, 1953, Univ. Kansas Sci, Bull., vol. 35,
p. 978. 9 holotype, 3 allotype, 28 9 and 40 ¢ paratypes.
5242,
Perdita trifasciata Timberlake, 1958, Univ. Kansas Sci. Bull., vol. 85,
p. 969. ¢ holotype. 5248.
Perdita verbesinae nigrior Cockerell, 1896, Proc. Acad, Sci, Phila-
delphia, vol. 48, p. 102. 9 holotype. 4807.
Proteriades bidenticauda Timberlake and Michener, 1950, Univ.
Kansas Sci. Bull., vol. 33, p. 420. g holotype, ¢ allotype, 6 9
and 16 3 paratypes. 4877.
508 Tue Universrry ScreENCE BULLETIN
Proteriades caudex Timberlake and Michener, 1950, Univ. Kansas
Sci. Bull., vol. 33, p. 418. ¢ holotype, ? allotype,6 ¢ and8 ¢
paratypes. 4378.
Proteriades hamulicornis Timberlake and Michener, 1950, Univ.
Kansas Sci. Bull., vol. 33, p. 425. ¢ holotype, ¢ allotype. 5274.
Proteriades incanescens tota Michener, 1954, Jour. Kansas Ent. Soc.,
vol. 27, p. 72. 2 holotype. 5275.
Proteriades nanula sparsa Timberlake and Michener, 1950, Univ.
Kansas Sci. Bull., vol. 33, p. 410. ¢ holotype, allotype, 2 ¢
and 5 ¢ paratypes. 4382.
Proteriades nigrella attonita Michener, 1954, Jour. Kansas Ent. Soc.,
vol. 27, p.73. ¢@ holotype, ¢ allotype. 5277,
Proteriades truicauda Timberlake and Michener, 1950, Univ. Kansas
Sci. Bull., vol. 83, p. 423. g¢ holotype, ¢ allotype, 1 g andl 9
paratypes. 4386.
Psithyrus consultus Franklin, 1918, Trans. Amer. Ent. Soc., vol. 38,
p. 459. 2 g cotypes. 1035.
Psithyrus tricolor Franklin, 1911, Trans. Amer, Ent, Soc., vol. 37,
p. 167. 1g cotype. 1034.
Spinoliella lawae Michener, 1937, Ann. Mag. Nat. Hist., ser. 10, vol.
19, p. 324. 3 holotype. 4538.
Synhalonia fuscotincta Cockerell, 1905, Proc. Biol. Soc. Washington,
vol. 18, p.178. ¢@ holotype. 1029.
Synhalonia gillettei snoviana Cockerell, 1905, Proc. Biol. Soc. Wash-
ington, vol. 18, p.179. 3 holotype. 1028.
Synhalonia lippiae semilippiae Cockerell, 1905, Proc. Biol. Soc.
Washington, vol. 18, p.179. @ holotype. 1030.
Trigona acapulconis kraussi Schwarz, 1951, Amer. Mus. Nov., no.
1505, p.8. ¢ holotype, 1 ? paratype. 4458.
Trophocleptria schrederi_ Michener, 1954, Bull. Amer. Mus. Nat.
Hist., vol. 104, p. 127. ¢ holotype. 5291.
Xenoglossa angelica Cockerell, 1902, Ent. News, vol. 13, p. 108.
g cotype. © 4435.
Xenoglossa pruinosa limitaris Cockerell, 1906, Ann. Mag. Nat. Hist.,
ser. 7, vol. 18, p. 73. g holotype. 1081.
Xenoglossa strenua kansensis Cockerell, 1905, Psyche, vol. 12, p. 104.
g holotype, 1 g paratype. 1032.
Xenopanurgus readiot Michener, 1952, Jour. Kansas Ent. Soc., vol.
25, p. 25. g holotype. 4532.
CATALOGUE oF TypEs IN SNow Museum 509
APOIDEA—Paratypes
Ancyloscelis hertigi Michener. 8 ¢,8 9. 5282.
Ancyloscelis panamensis Michener. 10 g ,23 9. 5283.
Andrena duboisi Timberlake, 1 ¢, 19. 4855.
Andrena lativentris Timberlake. 19. 4416.
Andrena nigroclypeata Linsley. 2 ¢,29. 5203.
Andrena sieverti opacicauda Cockerell. 19. 4289.
Andrena submaura Linsley. 19. 4287.
Andrena vanduzeei Linsley. 19. 5205.
Andrena vidalesi panamensis Michener. 1 g , 11 ¢. 5204.
Anthocopa alta Michener. 29. 4390.
Anthocopa anthodyta Michener. 6 @. 4391.
Anthocopa anthodyta bequaerti Michener. 1 ¢,19. 4892.
Anthocopa arizonensis Michener. 1 9. 5278.
Anthocopa namatophila Michener. 1 g . 5297.
Anthocopa nigrior Michener. 1 3. 4896.
Anthocopa nitidivitta Michener. 1 3,2 9. 4897.
Anthocopa oregona Michener. 1 ¢,1 9. 4898.
Anthocopa panamintensis Michener. 4 ¢,1 9. 4399.
Anthocopa pycnognatha Michener. 4 ¢. 4400.
Anthocopa pycnognatha solatus Michener. 2 ¢ , 2 9. 4401.
Anthocopa rubrella Michener. 1 ¢,19. 4402.
Anthocopa triodonta usingeri Michener. 8 3,59. 4403.
Anthophora emarginata Timberlake. 14,39. 4448.
Anthophora erythrothorax Michener. 19. 4444.
Anthophora gohrmanae coloradensis Michener. 1 ¢,19. 4445.
Anthophora langi Cockerell. 1 ¢. 4446.
Anthophora rhodothorax Michener. 1 3,19. 4447.
Anthophora simillima californiensis Michener. 1 g. 4448.
Anthophora subspensis Cockerell. 1. 4291.
Ashmeadiella barberi Michener. 1 3. 4276.
Ashmeadiella basalis Michener. 7 2,29. 4280.
Ashmeadiella basalis nigra Michener. 1 9. 4858.
Ashmeadiella breviceps Michener. 89. 4282.
Ashmeadiella cactorum astragali Michener. 10 o,509. 2578.
Ashmeadiella cactorum Cockerell. 1 9. 4277.
Ashmeadiella californica sierraensis Michener. 1 9. 4278.
Ashmeadiella cazieri Michener. 19. 4279.
Ashmeadiella difugita emarginatula Michener. 8 9. 4474,
Ashmeadiella erema Michener. 29. 4281.
Ashmeadiella eurynorhyncha Michener. 2 4,39. 4273.
510 Tre Universiry SCIENCE BULLETIN
Ashmeadiella florissantensis Michener. 13,19. 4272,
Ashmeadiella foxiella Michener. 1g. 4271.
Ashmeadiella gillettei cismontanica Michener. 29. 4334.
Ashmeadiella inyoensis Michener. 3g ,6¢. 4356.
Ashmeadiella maxima Michener. 1¢. 4857.
Ashmeadiella occipitalis Michener. 13,19. 4359.
Ashmeadiella rufitarsis Michener. 13,19. 4860.
Ashmeadiella rufiventris Michener. 1¢. 4861. ,
Ashmeadiella salviae Michener. 3 g,6¢. 4862.
Ashmeadiella submaxima Michener. 1 3. 4863.
Ashmeadiella timberlakei salida Michener. 7 $3 ¢ . 4365.
Ashmeadiella xenomastax Michener. 33. 4867.
Augochlora antonita Michener. 5 3, 29. 5251.
Augochlora chiriquiana Michener. 2g. 5258.
Augochlora clarki Michener., 3 3 , 849. 5254.
Augochlora hallimani Michener. 2 ? . 5255,
Augochlora mulleri mourei Michener. 19. 5256,
Augochlora quiriquensis Cockerell. 1 ¢ . 4315.
Augochlora utahensis Michener. 1 ¢ . 4314.
Augochlorella bidentata Michener. 2 3, 9°. 5252, \
Augochlorella edentata Michener. 19. 5262.
Augochloropsis auriferina Michener. 19. 5247.
Augochloropsis crassiceps Moure. 1¢. 5248.
Augochloropsis fairchildi Michener. 5 2. 5249.
Augochloropsis hebescens fuscocyanea Michener. 19. 5250.
Caenaugochlora macswaini Michener. 2g. 5261.
Calliopsis anomoptera Michener. 13,1¢. 4808.
Calliopsis bernardinensis Michener. 73,89. 4809.
Centris obscurior Michener. 17 3,39. 5801.
Ceratina breviceps Michener. 4 3,29. 5302.
Ceratina nanula rigdenae Michener. 43,59. 4450,
Ceratina sequoiae Michener. 59. 4451.
Ceratina subpunctigena Michener. 5 ? . 4452.
Ceratina tricolor Michener. 2 3,5 9. 5807.
Chelostoma minutum cockerelli Michener. 3¢,29. 4821.
Chelostoma minutum incisulum Michener. 2 3. 4320,
Chelostoma minutum marginatum Michener. 4 3 , 29. 4822.
Chelostoma phacelia Michener. 6 3 , 49. 4830,
Chelostoma tetramerum Michener. 1 3. 4331].
Colletes algarobiae Cockerell. 13. 5499.
Colletes arizonensis Stephen. 1g. 5215.
Colletes brimleyi Mitchell. 4 3. 5198.
CATALOGUE OF TyPEs IN SNow Musrum 511
Colletes bryanti Timberlake. 1 g. 5222.
Colletes carolinus Mitchell. 29. 5199.
Colletes covilleae Timberlake. 1 3. 5192.
Colletes deserticola Timberlake. 2 3,29. 5191.
Colletes hyalinus oregonensis Timberlake. 49. 5196.
Colletes ilicis Mitchell. 2¢,29. 5183.
Colletes laticinctus Timberlake. 19. 5198.
Colletes lutzi interior Timberlake. 1 @. 5184.
Colletes lutzi Timberlake. 14,19. 5185.
Colletes lutzi monticola Timberlake, 2 ¢. 5186.
Colletes pinorum Timberlake. 19. 5187.
Colletes mitchelli Stephen. 23,19. 5213.
Colletes panamensis Michener. 1 ¢,19. 5200.
Colletes paniscus mertensiae Timberlake. 1 ¢, 1 @. 5190.
Colletes rugicollis nigrior Michener. 2¢,19. 5201.
Colletes sphaeralceae Timberlake. 6 g¢ , 6 9. 5188.
Colletes susannae Swenk. 1g. 5197.
Colletes tectiventris Timberlake. 5 ¢ , 9 9. 5194.
Colletes thysanellae Mitchell, 1 By kes 75195;
Diadasia lutzi Cockerell. 14,19. 4484,
Dianthidium subrufulum Timberlake. 14. 5189,
Dufourea cuprea Bohart. 13,19. 4339.
Dufourea dentipes Bohart. 13. 4340.
Dufourea femorata Bohart, 1 3,19. 4842.
Dufourea nudicornis Timberlake. 1 ¢. 5267.
Dufourea trochantera Bohart. 19. 4344.
Dufourea tuolumne Bohart. 1 ¢,19. 4845.
Dufourea vanduzeei Bohart. 1 ¢,19. 4843.
Epeolus ainsliei Crawford. 1 ¢. 4428.
Exomalopsis albata Timberlake. 2¢. 5285.
Exomalopsis compactula completa Cockerell. 19. 4417.
Fiorentinia lanei Moure. 19. 5284.
Halictoides escholtziae Michener. 1 3. 4341.
Halictoides ruficaudus Michener, 79. 4347.
Halictoides sandhouseae Michner. 1 g , 8 9. 4846.
Malictus abanci Crawford. 19. 4328.
Halictus abietum Michener. 1 g,29. 4824.
Halictus albuquerquensis Michener. 29. 4825.
Halictus brimleyi Crawford. 19. 4817.
Halictus brycinci Crawford. 19. 4827.
Halictus euryceps Ellis. 2. 4828.
lalla Tue University Screnck BULLETIN
Halictus marinensis Michener. 1 9 . 4829.
Halictus pasadenae Michener. 2 3 , 209. 4882.
Halictus sequoiae Michener. 29. 4338.
Heriades communis Cockerell. 19. 4298.
Heriades cressoni Michener. 38,69. 4350.
Heriades occidentalis Michener. 38,29. 4335.
Heriades timberlakei Michener. 3%. 4836.
Hesperapis arida Michener. 16 4,19. 4851.
Hesperonomada melanantha Linsley. 13,19. 4418.
Heteranthidium cordaticeps Michener. 19. 4853.
Heteranthidium zebratum mississippi Michener. 13,19. 4854.
Hexepeolus mojavensis Linsley. 28,19. 4421.
Hoplitis bunocephala Michener. 13. 4370,
Hoplitis bullifacies Michener. 19. 4869.
Hoplitis elongaticeps Michener. 19. 4371.
Hoplitis paroselae Michener. 19. 4868.
Hoplitis producta bernardina Michener. 1 3. 4378,
Hoplitis producta panamintana Michener. 1 ¢. 4374.
Hylaeus panamensis Michener. 19. 5202.
Hypomacrotera andradensis Cockerell. 2¢,29. 4810.
Lasioglossum denticeps Michener. 29. 5263.
Megachile monardarum Cockerell. 19. 4415.
Megachile manardarum Cockerell. 1¢ . 4413.
Megachile soledadensis Cockerell. 1g. 4414.
Melipona fasciata paraensis guerreroensis Schwarz. 19. 4454.
Melipona lateralis kangarumensis Cockerell. 19. 4456.
Melipona marginata micheneri Schwarz. 1 ¢ . 4983,
Melissodes confusiformis incondita Cockerell. 19. 4439,
Melissodes maesta LaBerge. 18,19. 5501.
Melissodes sabinensis Cockerell. 1@. 4441.
Melissodes sabinensis nubila LaBerge. 10 ¢,29. 5503.
Melissodes texana eluta LaBerge. 33,29. 5507.
Melissodes toluca Cresson. 29. 5281.
Neocorynura perpectinata Michener. 3 ¢ . 5258.
Neocorynura rufa Michener. 1g,19. 5259.
Neocorynura villosissima Michener. 24,29. 5260.
Neolarra abdominalis Michener. 19. 4428.
Neolarra californica Michener. 13,19. 4424.
Neolarra coloradensis Michener. 49. 4425.
Neolarra vandykei Michener. 49. 4426.
Nomada pallidelutea Swenk. 13. 4419.
CATALOGUE OF TypES IN SNOW MusEUM 518
Nomada panamensis Michener. 1 f. 5287.
Nomada simplicocoxa Swenk. 26,89. 4420.
Nomada trapidoi Michener. 2,19. 5288.
Odyneropsis gertschi Michener. 3 ¢. 5289.
Osmia brevior Michener. 89. 4895.
Osmia claremontensis Michener. 1 ¢. 4407.
Osmia dakotensis Michener. 19. 4405.
Osmia francisconis White. 29. 4492.
Osmia lawae Michener. 14,29. 4872.
Osmia parallela Michener. 1 ¢,29. 4408.
Osmia pingreeana Michener. 19. 5280.
Osmia potentillae Michener. 19. 4409.
Osmia ribifloris biedermannit Michener. 8 ¢,39. 4404.
Osmia sequoiae Michener. 19. 4406.
Osmia subpurpurea jamesi Michener, 3¢@. 4411.
Oreopasites vanduzeei vernalis Linsley. 1 ¢. 4270.
Panurginus atramontensis Crawford. 1 ¢. 4292.
Panurginus gracilis Michener. 19. 4294.
Panurginus nigrihirtus Michener. 1 ¢,19. 4295.
Perdita acapulcona Timberlake, 1 3. 5228.
Perdita ainsliei Crawford. 19. 4800,
Perdita azteca Timberlake. 13. 5229.
Perdita coahuilensis Timberlake. 4¢,39. 5881.
Perdita diversa Timberlake. 14,39. 5882.
Perdita euphorbiae Timberlake. 63,79. 5288.
Perdita giliae Timberlake. 23,29. 5285.
Perdita mellea Timberlake. 43,79. 5225,
Perdita miricornis Cockerell. 1 ¢,19. 4808.
Perdita mitchelli Timberlake. 29. 4804.
Perdita obsuripennis Timberlake. 1 3,59. 5228.
Perdita portalis Timberlake. 19. 5239.
Perdita ruficauda ornata Timberlake. 5 ¢. 5240,
Perdita salicis occidentalis Timberlake. 23,29. 4806.
Perdita signata Timberlake. 5 ¢,79. 5380.
Perdita tristella Timberlake. 19. 5244.
Perdita turgiceps Timberlake. 264, 109. 5245.
Proteriades basingeri Timberlake and Michener. 1 ¢. 4875.
Proteriades boharti Timberlake and Michener. 1g. 4876.
Proteriades cryptanthae Timberlake and Michener. 1 3,22. 4879.
Proteriades deserticola Timberlake and Michener. 4 ¢,7 9. 4880.
Proteriades incanescens nevadensis Timberlake and Michener. 1 @ .
4388.
514 Tue Universtry ScreNcE BULLETIN
Proteriades nanula Timberlake and Michener. 2¢,29. 4881.
Proteriades nigrella Michener. 29. 5276.
Proteriades pygmaea Timberlake and Michener. 2 ¢,39. 4384.
Proteriades reducta Timberlake and Michener. 1 ¢,19. 4885.
Proteriades seminigra Timberlake and Michener. 19. 4887.
Proteriades seminigra yosemitensis Timberlake and Michener. 1 3 ,
1¢o. 4888.
Proteriades tristis Michener. 19 . 4389.
Pseudopanurgus gerardiae Crawford. 14,19. 4296.
Pseudopanurginus nebrascensis timberlakei Michener. 19. 4297.
Sphecodes macswaini Michener. 29. 5266.
Stelis anthracina Timberlake. 1 ¢. 5268.
Stelis arizonensis Swenk. 5 3. 4299.
Stelis ashmeadiellae Timberlake. 19. 4811.
Stelis depressa Timberlake. 3g¢,19. 4313.
Stelis hemirhoda Linsley. 29. 4316.
Stelis micheneri Linsley. 2g. 4818.
Tetralonia lata lautipes Cockerell. 19. 4487.
Townsendiella californica Michener. 69. 4422.
Triepeolus rufithorax Graenicher. 19. 4429.
Trigona katangensis Cockerell. 19. 4457.
Trigona leucogastra chiriquiensis Schwarz. 19. 4433.
Trigona mexicana subobscuripennis Schwarz. 29. 4802.
Trigona pectoralis barrocoloradensis Schwarz. 29. 4312.
Trophocleptria nigra Michener. 23,19. 5292.
Trophocleptria odontothorax Michener. 2 ¢. 5290.
Xylocopa varians rufotegularis Michener. 19. 5308.
BRACONIDAE
Agathis wyomingensis Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 284. holotype. 902.
Aphaereta delosa Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 284. g¢cotype destroyed, ¢ cotype missing abdomen, 1 9
paratype. 885.
Aphaereta subtricarina Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 283. ¢ holotype, 1 ? paratype. 886.
Ascogaster mimeticus Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 277. @ holotype. 901.
Aspilota columbiana Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 282. ¢ holotype. 887.
Atanycolus calophrys Shenefelt, 1948, Washington State Coll. Res.
Studies, vol. 11, p. 72. ¢@ holotype. 3520.
CATALOGUE oF Types IN SNow MusEUM 515
Biosteres indotatus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 278. holotype. 861.
Brachistes nocturnus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 279. gholotype. 895.
Bracon kansensis Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 268. ¢ holotype. 877.
Bracon piceiceps Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 268. g holotype (abdomen missing), 878.
Cardiochiles nigroclypeus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 275. holotype, 2 ¢ paratypes. 858.
Chelonus altitudinis Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 278. 9 holotype. 896.
Chelonus egregicolor Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 278. gholotype. 897.
Chelonus exogyrus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 287. 9@ holotype. 898.
Chelonus nucleolus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 287. 9? holotype. 899.
Chelonus texanoides Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 286. 9 holotype. 900.
Coenocoelius politifrons Viereck, 1905, Trans, Kansas Acad. Sci.,
vol. 19, p. 279. g holotype. 894.
Crassomicrodus nigricaudus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 288. ¢ holotype, 1 ¢ paratype. 903.
Diachasma appalachicola Viereck, 1905, Trans, Kansas Acad. Sci.,
vol. 19, p. 272. 9 holotype. 862.
Diachasma secunda Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 273. 9 holotype. 863.
Dinotrema signifrons Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p.177. holotype. 888.
Doryctes apacheus Viereck, 1907, Trans. Amer, Ent. Soc., vol. 33,
p. 383. holotype. 884.
Doryctes femur-rubrum Viereck, 1907, Trans, Amer. Ent. Soc., vol.
83, p. 384, 9 holotype. 883.
Doryctomorpha shoshonea Viereck, 1907, Trans. Amer, Ent. Soc.,
vol. 33, p. 882. ¢ holotype. 2111.
Hedysomus wichitus Viereck, 1907, Trans. Amer. Ent. Soc., vol. 83,
p. 404. #holotype. 907.
Hormiopterus claripennis Brues, 1907, Bull. Wisconsin Nat. Hist.
Soc., vol. 5, p. 60. ¢ holotype. 879.
Iphiaulax cinnabarinus Viereck, 1905, Trans, Kansas Acad. Sci.,
vol. 19, p. 286. ¢ holotype. 869.
Tur University SCcreENCE BULLETIN
Iphiaulax melanogaster Viereck, 1905, Trans. Kansas Acad, Sci.,
vol. 19, p. 270. ¢@ holotype. 870.
Iphiaulax militaris Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 269. 9 holotype. 871.
Iphiaulax perepicus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 285. @ holotype. 872.
Iphiaulax propinquus Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 286. ¢@ holotype missing abdomen. 873.
Iphiaulax triangulifera Viereck, 1907, Trans. Amer. Ent. Soc., vol. 83,
p. 384. @ holotype. 868.
Ischneutidea (1) proteroptoides Viereck, 1905, Trans. Kansas Acad.
Sci., vol. 19, p. 274. @ holotype missing abdomen. 860.
Lysiphlebus succineus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 282. g holotype, 1 g paratype. 889.
Lytopylus asygos Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 267. holotype. 904.
Melanobracon ulmicola Viereck, 1906, Trans. Amer. Ent. Soc., vol.
32, p.176. g and ¢ cotypes, 2 ? paratypes. 876.
Meteorus campestris Viereck, 1905, Trans, Kansas Acad. Sci., vol. 19,
p. 281. ¢@ holotype. 890.
Meteorus noctivagus Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 281. ¢ holotype. 891.
Meteorus relativus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 280. ¢holotype. 892.
Microdus agathoides Viereck, 1905, Trans. Kansas Acad, Sci., vol. 19,
p. 277. ¢ holotype—lost. 905.
Microdus castaneicinctus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 276. ¢ holotype. 906.
Microdus nigrotrochantericus Viereck, 1905, Trans. Kansas Acad.
Sci., vol. 19, p.275. ¢ holotype. 910.
Microdus pimploides Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 276. holotype. 909.
Microdus wichitaensis Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 276. @ holotype missing head, abdomen and wings. 911.
Microgaster tuckeri Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 274. ¢@ holotype. 859.
Opius aberrans Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 271. ¢ holotype, 1 ¢ paratype. 864.
Opius basiniger Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 270. ¢ holotype missing abdomen. 865,
Opius luteiceps Viereck, 1905, Trans. Kansas Acad, Sci., vol. 19,
p. 271. gholotype. 866.
CATALOGUE OF TypEsS IN SNow MusEUM 517
Opius nigrocastaneus Viereck, 1905, Trans, Kansas Acad. Sci., vol.
19, p. 272. 3g holotype. 867.
Phaenodus caddous Viereck, 1907, Trans. Amer. Ent. Soc., vol. 33,
p. 403. ¢ holotype. 908.
Rhogas melanothorax Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 267. 9° holotype missing head. 882,
Vipio erythrus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 284. ¢ holotype, 1 ¢ paratype. 874.
Vipio piceipectus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 269. 9 holotype. 875.
Zele crassicalcaratus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 279. &holotype—lost. 898.
BRACONIDAE-——Paratypes
Atanycolus crassicruris Shenefelt. 29. 3521.
Atanycolus impressifrons Shenefelt. 19. 3524.
Atanycolus longifemoralis Shenefelt. 19. 3522.
Atanycolus malii Shenefelt. 19. 3525.
Bracon charus Riley. 3 ¢. 3528.
Bracon pascuensis Shenefelt. 29,1 4. 2776.
Heterospilus beameri Rohwer. 59,92. 880.
CHALCIDOIDEA
Leucospis bicincta Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 227. holotype. 918.
CHALCIDOIDEA—Paratypes
Necremnus breviramulus Gahan. 159,38 4. 8282.
Orasema beameri Gahan. 89,3 ¢. 8280.
Orasema cockerelli Gahan. 29. 8281.
Spilochalcis clora Burks. 1 ¢. 8059.
Spilochalcis dema Burks. 19. 8060.
CHRYSIDIDAE
Chrysis equidens: Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32, p.
193. ¢ holotype, 1 ? paratype. 915.
Chrysis kahli Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32, p. 194.
? holotype. 916.
Chrysis kansensis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82, p.
198. 9 holotype. 917.
Chrysis pattonella Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32, p.
194. ¢ holotype. 918.
Tue Universtry Science BULLETIN
518
Chrysis snowi Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82, p. 195.
? holotype. 919.
Notozus connexus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 192. ¢ holotype. 914.
CYNIPOIDEA
Aspicera africana Kinsey, 1919, Psyche, vol. 26, p. 162. 1g and
1 ¢ cotype. 807.
CYNIPOIDEA—Paratypes
Acraspis undulata Gillette. 19. 8451.
Andricus biconicus Weld. 49. 3452.
Andricus foliosus Weld. 29. 8453.
Andricus robustus Weld. 29. 8454.
Callirhytis fructuosa Weld. 49. 8455.
Compsodryoxenus illinoisensis Weld. 19. 3456.
Disholcaspis terrestris Weld. 29. 4488.
Holcaspis monticola Gillette. 19. 8458.
Neuroterus verna Gillette. 29. 8459.
Rhodites arefactus Gillette. 29. 3457.
EVANIIDAE
Hyptia texana Bradley, 1908, Trans. Amer. Ent. Soc., vol. 34, p. 161.
¢ holotype. 788.
FORMICIDAE—Paratypes
Lasius crypticus Wilson. 1 g, 1 worker. 5398.
Lasius sitiens Wilson. 1 worker. 5899.
Lasius fallax Wilson. 8 workers. 5400.
GASTERUPTIIDAE
Tropaulacus torridus Bradley, 1908, Trans. Amer. Ent. Soc., vol. 34,
p. 125. ¢ holotype. 3811.
GASTERUPTILDAE—Paratypes
Gasteruption amputatum Townes. 1g. 38814.
Rhydinofoenus floridanus bradleyi Townes. 19. 8815.
Rhydinofoenus striatus Townes. 2 $. 3818.
Rhydinofoenus tarsatoris solaris Townes. 19. 3812.
ICHNEUMONIDAE
Agathobancus bradleyi Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 308. ¢@ holotype. 843.
Amorphota angusta Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 180. ¢ holotype. 820.
CaraLocuE or Types In SNow MusruM 519
Amorphota confluens Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 217. @ holotype. 821.
Amorphota ferruginosa Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 220. @ holotype. 822.
Amorphota galvestonensis Viereck, 1906, Trans. Amer, Ent. Soc.,
vol. 82, p. 219. ¢ holotype. 828.
Amorphota nocturna Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 308. g holotype. 827.
Amorphota paenexareolata Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 309. g¢ holotype. 829,
Amorphota perrivalis Viereck, 1905, Trans. Kansas Acad, Sci., vol.
19, p. 807. 1 ¢ and 1 ¢ cotypes, 1 ¢ paratype. 880.
Amorphota relativa Viereck, 1902, Canad. Ent., vol. 34, p. 67. 9
holotype. 881.
Angitia autumnalis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p- 181. ¢ holotype (abdomen missing). 838,
Anomalon paeneferrugineum Viereck, 1905, Trans. Kansas Acad.
Sci., vol. 19, p. 310. 9 holotype. 808.
Anomalon fulvescens var. hemimelas Viereck, 1906, Trans. Amer.
Ent. Soc., vol. 32, p. 218. 9 holotype. 826.
Aranetia leucotaenia Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 818. 9 holotype. 808,
Atrometus angitioides Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p.311. ¢ holotype. 804.
Boethus aenigmaticus Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p.314. @ holotype. 796.
Callidiotes kansensis Brues, 1907, Wisconsin Nat. Hist. Soc. Bull.,
vol. 5, p.55. ¢ holotype. 798.
Campoplex photomorphus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p.309. 9 holotype. 805.
Campoplex wyomingensis Viereck, 1906, Trans. Amer. Ent. Soc.,
vol, 32, p. 242. holotype. 806.
Cryptus citrinimaculatus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 820. ¢ holotype. 810.
Cryptus consobrinus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p..225. @ holotype. 811.
Cryptus crassisculptus Pratt, 1945, Amer. Midland Nat., vol. 34,
p.578. 9? holotype. 2407.
Cryptus politicalypterus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 298. g holotype. 812.
Erymotylus felti Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 812. @ holotype. 828.
Tue Universrry ScreENCE BULLETIN
520
Exetastes infumatricus Cushman, 1937, Proc. U. S. Nat. Mus., vol.
84, p. 283. ¢ holotype. 2409.
Exetastes infumatricus Cushman, 1987, Proc. U. S. Nat. Mus., vol.
84, p. 288. 9? holotype. 2887.
Glypta aprilis Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19, p. 316.
? holotype. 853.
Glypta brunneisigna Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 316. @ holotype. 854,
Glypta egregiafovea Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 292. ¢@ holotype. 852.
Glypta succineipennis Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p.3815. 9? holotype. 851.
Harrimaniella paeneimitatrix Viereck, 1905, Trans. Kansas Acad.
Sci., vol. 19, p.817. ¢ holotype. 809.
Hemiteles cushmani Butcher, 1933, Ann. Ent. Soc. Amer., vol. 26,
p. 76. ¢@holotype, allotype, 39 and 4 ¢ paratypes. 2406.
Hemiteles cheboyganensis Butcher, 1933, Ann. Ent. Soc, Amer., vol.
26,p.77. @ holotype. 2408.
Hemiteles lathroscopoides Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 294. g holotype. 815.
Hemiteles manitouensis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 290. g holotype. 816.
Ichneumon arizonensis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 295. ¢ holotype. 789.
Ichneumon citrinifascialis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 297. holotype. 790.
Ichneumon egregiafacialis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p.291. g holotype. 824.
Ichneumon flavofascialis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 291. holotype. 791.
Ichneumon hemimelanarius Viereck, 1906, Trans. Amer. Ent. Soc.,
vol. 32, p. 226. g holotype. 792.
Ichneumon humphreyi Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 296. g holotype. 798.
Ichneumon nigrosignatus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 297. g holotype. 794.
Ichneumon oryxicornis Viereck, 1905, Trans, Kansas Acad. Sci., vol.
19, p. 824. g holotype. 856.
Idecthis biconjunctus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 180. holotype. 835.
Idecthis paenerivalis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 217. 19 and1 3 cotypes. 836.
CATALOGUE OF TYPES IN SNow MusEUM SAE
Ischnoscopus taeniatus Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 305. ¢ holotype. 832.
Limnerium lawrencei Vierick, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 178. 9 holotype. 887.
Limnerium perdistinctum Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 304. 9 holotype, 2 ? paratypes. 838.
Limnerium vigilis Viereck, 1906, Trans. Amer, Ent. Soc., vol. 32,
p. 179. g holotype. 889.
Mesochorus noctivagus Viereck, 1905, Trans. Kansas Acad, Sci.,
vol. 19, p. 303. ¢@ holotype. 842.
Mesostenus discoidaloides Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 819. ¢ holotype. 813.
Metopius grandior Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 814. @ holotype. 795.
Nematopodius exclamans Viereck, 1905, Trans, Kansas Acad. Sci.,
vol. 19, p. 818. ¢ holotype. 3467.
Netelia cushmani Townes, 1939, Lloydia, vol. 1, p. 206. ¢ holotype.
8409.
Netelia glabra Townes, 1939, Lloydia, vol. 1, p. 183. ¢ holotype.
3412.
Olesicampa melanerythrogastra Viereck, 1905, Trans. Kansas Acad.
Sci., vol. 19, p. 806. 19 and 1 gs cotypes. 834.
Ophion idoneum Viereck, 1905, Trans. Kansas Acad, Sci., vol. 19,
p. 3812. g@holotype. 800.
Pezomachus alogus Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 821. holotype. 817.
Pezomachus homalommoides Viereck, 1905, Trans, Kansas Acad.
Sci., vol. 19, p. 294. holotype. 818.
Pezomachus testaceicoxus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol, 19, p. 822, ¢ holotype. 819.
Phygadeuon spinicoxus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 289. ¢ holotype. 857.
Pimpla landerensis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 248. ¢ holotype. 849.
Pimpla parvialba Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32, p.
182. ¢ holotype. 850.
Platylabus omniferrugineus Viereck, 1905, Trans, Kansas Acad. Sci.,
vol. 19, p. 822. ¢ holotype. 855.
Pristomerus appalachianus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 298. 1 (abdomen missing) and 1 g¢ (abdomen miss-
ing) cotypes. 840.
522 Tue Universrry Screnck BULLETIN
Pristomerus appalachianus var. dorsocastaneus Viereck, 1905, Trans.
Kansas Acad. Sci., vol. 19, p. 299. 19 and 1 gcotypes. 841.
Scopiorus monticola Brues, 1907, Wisconsin Nat. Hist. Bull., vol. 5,
p. 56. ¢ holotype. 799.
Sychnoporthus tuckeri Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 815. ¢ holotype. 814.
Syndipius erythrogaster Viereck, 1908, Trans. Amer. Ent. Soc., vol.
33, p. 3885. ¢ holotype (missing abdomen). 797.
Thersilochus egregiacolor Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 302. ¢ holotype, 1 ? paratype. 844.
Thersilochus hamiltonensis Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 301. g holotype. 845.
Thersilochus mimeticus Viereck, 1905, Trans. Kansas Acad. Sci.,
vol. 19, p. 300. ¢ holotype. 846.
Thersilochus quintilis Viereck, 1905, Trans. Kansas Acad. Sci., vol.
19, p. 301. g holotype. 847.
Thersilochus snowi Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 299. 9 holotype. 848.
Thyreodon morio race transitionalis Viereck, 1906, Trans. Amer. Ent.
Soc., vol. 82, p. 225. ¢ holotype. 801.
Thyreodon snowi Viereck, 1905, Trans. Kansas Acad. Sci., vol. 19,
p. 318. holotype (tip of abdomen damaged), 1 ¢ paratype.
802.
ICHNEUMONIDAE—Paratypes.
Conocalama occidentalis var. luteata Hopper. 19. 2427.
Cryptus associatus Pratt. 19. 8299.
Cryptus asymmetricus Pratt. 19,1 ¢. 8297.
Cryptus flagellatus Pratt. 2°. 3300.
Cryptus mutatus Pratt. 19. 3298.
Cryptus temporalis Pratt. 1g. 8296.
Echthronomas ochreofrons lexiphaga Walley. 14. 4222.
Erromenus dolichops Townes and Townes. 1 3. 8605.
Erromenus nasalis Townes and Townes. 1 3. 3604.
Exetastes angustoralis Cushman. 19,1 ¢. 2388.
Exetastes bituminosus Cushman. 19. 2386.
Exetastes geminus Cushman. 1¢,1 3%. 2385.
Hoplismenus teres Swift. 1g. 4221.
Megarhyssa icterosticta Michener. 59,2 ¢. 2579.
Netelia acuminata Townes. 19. 3407.
Netelia magniceps Townes. 19. 8418.
Netelia pallens occidentalis Townes. 1g. 8408.
Pas Snr
CATALOGUE OF TyPEs IN SNOow MusEUM 523
Netelia recta Townes. 1g. 38410.
Netelia ovalis Townes. 19. 9468.
Symboethus communis foraminatus Townes and Townes. 19,13
3606.
Symboethus viator Townes and Townes. 19,14. 3603.
MUTILLIDAE
Dasymutilla arcana Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143, p.
217. & holotype, 3 g paratypes. 1059.
Dasymutilla beameri Mickel, 1936, Ann. Ent. Soc. Amer., vol. 29,
p. 48. ¢ holotype. 2112.
Dasymutilla campanula Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143,
p. 227. @ holotype. 934.
Dasymutilla candida Mickel, 1928, Bull. U. S$. Nat. Mus., vol. 143,
p. 296. ¢holotype. 1038.
Dasymutilla curticeps Mickel, 1928, Bull. U. S. Nat. Mus., vol. 148,
p. 178. ¢ holotype. 2116.
Dasymutilla dionysia Mickel, 1928, Bull. U. S. Nat. Mus., vol. 148,
p. 104. ¢ holotype. 928.
Dasymutilla eminentia Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143,
p. 79. g holotype, 9 g paratypes. 929.
Dasymutilla errabunda Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143,
p. 269. holotype. 1063.
Dasymutilla fulgida Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143,
p. 76. 9 holotype, 8 9 paratypes. 2122.
Dasymutilla furina Mickel, 1928, Bull. U. S. Nat. Mus., vol. 148,
p. 82. @holotype, gallotype. 2121.
Dasymutilla meracula Mickel, 1928, Bull. U.S. Nat. Mus., vol. 148,
p- 201. g holotype, 2 ¢ paratypes. 931.
Dasymutilla myrice Mickel, 1928, Bull. U. S. Nat. Mus., vol. 148,
p. 121. ¢ holotype, 28 ¢ paratypes. 1060.
Dasymutilla nupera Mickel, 1928, Bull. U. S. Nat. Mus., vol. 1438,
p. 108. ¢ holotype. 924.
Dasymutilla saetigera Mickel, 1928, Bull. U. S. Nat. Mus., vol. 148,
p. 211. ¢ holotype. 930.
Dasymutilla satanas Mickel, 1928, Bull. U. S. Nat. Mus., vol. 143,
p. 239. ¢ holotype, 3 ? paratypes. 1070.
Mutilla apachea Viereck, 1908, Trans. Amer. Ent. Soc., vol. 38, p.
386. g holotype. 1065.
Mutilla crepuscula Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 188. g holotype. 1039.
524 Tue Universiry SCIENCE BULLETIN
Mutilla imperialiformis Viereck, 1906, Trans. Amer. Ent. Soc., vol.
32, p. 189. g holotype. 1041.
Mutilla montivagoides Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 185. 9 holotype. 1042.
Mutilla nigricauda Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 187. @ holotype. 1056.
Mutilla phoenix Fox, 1899, Trans. Amer. Ent. Soc., vol. 25, p. 247.
g lectoallotype (designated by Mickel, 1928, Bull. U. S. Nat.
Mus., vol. 148, p. 87). 926.
Mutilla prognoides Viereck 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 186. ¢ holotype. 1057.
Mutilla quintilis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 187. g holotype. 1040.
Pseudomethoca carbonaria Mickel, 1924, Proc. U.S. Nat. Mus., vol.
64, p.43. gf holotype. 1049.
MUTILLIDAE—Paratypes
Dasymutilla atrifulua Mickel. 8g. 1086.
Dasymutilla calorata Mickel. 19. 1071.
Dasymutilla dilucida Mickel. 19. 1064.
Dasymutilla helva Mickel. 39. 1066.
Dasymutilla hirticula Mickel. 1 g. 1054.
Dasymutilla hora Mickel. 1 3. 985.
Dasymutilla intermixta Mickel. 2 4. 1068,
Dasymutilla magnifica Mickel. 29,14. 1069.
Dasymutilla nitidula Mickel. 29. 982.
Dasymutilla nogalensis Mickel. 119. 1067.
Dasymutilla ocydrome Mickel. 2 4. 1037.
Dasymutilla reclusa Mickel. 6 g. 1058.
Dasymutilla reperticia Mickel. 1 3. 1061.
Dasymutilla rubricosa Mickel. 1 3. 986.
Dasymutilla sophrona Mickel. 1g. 1062.
Dasymutilla stevensi Mickel. 29. 925.
Pseudomethoca paludata Mickel. 29. 1053.
PAMPHILIIDAE
Neurotoma crataegi Middlekauff. 2 ¢ paratypes. 3309.
PERGIDAE
Hylotoma spiculata McGillivray. 1 9 paratype. 785.
CATALOGUE OF Types IN SNOow MusEUM 525
POMPILIDAE
Anoplius snowi Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82, p. 202.
& holotype. 940.
Aporinellus taeniatus baboquivari Evans, 1951, Trans. Amer. Ent.
Soc., vol. 77, p. 286. holotype. 4516.
Aporus ferrugineipes Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 208. holotype. 937.
Cryptocheilus paeneparcus Viereck, 1906, Trans. Amer, Ent. Soc.,
vol. 32, p. 202. ¢ holotype. 941.
Pepsis angustimarginata Viereck, 1907, Trans. Amer. Ent. Soc., vol.
33, p. 898. 9? holotype. 939.
Pompilinus splendens Dreisbach, 1949, Ent. Amer. (n. s.), vol. 29,
p. 20. gholotype. 4522.
POMPILIDAE—Paratypes
Anoplius krombeini Evans. 19,18. 4519.
Aporinellus bridwelli Evans. 1 g. 4515.
Aporinellus sinuatus Evans. 39,5 ¢. 4517.
Episyron quinquenotatus Evans. 19,1. 4518.
Pepsis bequaerti Salman. 1 3. 938.
Pompilinus pseudoreductus Dreisbach. 1 g. 4526.
Pompilinus rectangularis Dreisbach. 1 ¢. 4525.
Pompilus parvulus otomi Evans. 19,2 g. 4645.
Pompilus solonus silvivagus Evans. 19. 4520.
SCELIONIDAE
Scelio monticola Brues, 1906, in Viereck, Trans. Amer, Ent. Soc.,
vol. 82, p. 184. 19 holotype. 920.
Scelio striaticollis Brues, 1906, in Viereck, Trans. Amer. Ent. Soc.,
vol. 82, p. 222. ¢ holotype. 921.
Scelio venata Brues, 1906, in Viereck, Trans. Amer. Ent. Soc., vol. 32,
p. 228. ¢ holotype (abdomen missing). 922.
SPHECIDAE
Ancistromma paenerugosa Viereck, 1906, Trans. Amer. Ent, Soc.,
vol. 32, p. 210. gholotype. 942.
Ancistromma tachysphecoides Viereck, 1906, Trans. Amer. Ent. Soc.,
vol. 82, p. 209. g holotype. 947.
Ancistromma vegetoides Viereck, 1906, Trans. Amer, Ent. Soc., vol.
32, p. 208. ¢ holotype. 825.
Ancistromma zerbei Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 208. ¢@ holotype. 946.
526 Tue Universiry Science BULLETIN
Bembix arcuata Parker, 1917, Proc. U.S. Nat. Mus., vol. 52, p. 81.
3 allotype. 963.
Bembix heliathopolis Parker, 1917, Proc. U.S. Nat. Mus., vol. 52,
p- 118. 9 holotype, ¢ allotype, 3 g paratypes. 960.
Bembix latifrons Parker, 1917, Proc. U.S. Nat. Mus., vol. 52, p. 116.
9 holotype. 964. .
Bicyrtes annulata Parker, 1917, Proc. U.S. Nat. Mus., vol. 52, p. 67.
9 holotype, 1 9 paratype. 965.
Bicyrtes gracilis Parker, 1917, Proc. U.S. Nat. Mus., vol. 52, p. 68.
g holotype. 966,
Crabro canonicola Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 403. 9 holotype, 1 ¢ paratype. 974.
Crabro cinctibellus Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 401. 19 and 1 gcotypes, 19 and 2 paratypes. 975.
Crabro clarconis Viereck, 1906, Trans. Amer, Ent, Soc., vol. 32,
p. 213. 9? holotype. 976,
Crabro papagorum Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 401. ¢ holotype. 977.
Crabro viciniformis Viereck, 1907, Trans. Amer. Ent, Soc., vol. 83,
p. 381. ¢ holotype. 978.
Didineis crassicornis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 204. g¢ holotype. 1008.
Diodontus brunneicornis Viereck, 1906, Trans. Amer. Ent. Soc., vol.
82, p.212. gholotype. 5450.
Eucerceris lacunosa Scullen, 1939, Oregon State Monogr., Studies in
Ent., vol. 1,p.19. g holotype. 4210.
Gorytes gulielmi Viereck, 1907, Trans. Amer. Ent. Soc., vol. 38,
p. 408. holotype. 1008.
Gorytes papagorum Viereck, 1907, Trans, Amer. Ent. Soc., vol. 33,
p. 400. 19and 1 4 cotypes, 39 and 1¢ paratypes. 1005.
Harpactus gyponae Williams, 1913, Univ. Kansas Sci. Bull., vol. 8,
p. 223. 9 holotype. 1004.
Larropsis ater Williams, 1913, Univ. Kansas Sci. Bull., vol. 8, p. 146.
& holotype. 945.
Larropsis minor Williams, 1913, Univ. Kansas Sci. Bull., vol. 8,
p. 146. 3 holotype (abdomen missing). 2113.
Niteliopsis foxii Viereck, 1906, Trans. Amer, Ent. Soc., vol. 82, p. 207.
9 holotype. 961.
Niteliopsis kansensis Williams, 1913, Univ. Kansas Sci. Bull., vol. 8,
p. 178. 9 holotype. 962.
Nysson clarconis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 204. g¢ holotype. 1006.
CATALOGUE OF TypEs IN SNow MusruM 527
Nysson intermedius Viereck, 1907, Trans. Amer, Ent. Soc., vol. 33,
p. 409. ¢ holotype. 1007.
Oxybelus exclamans Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 215. 9 holotype (missing from collection, but probably loaned
to some specialist). 1048.
Passaloecus equalis Viereck , 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 212. ¢@ holotype. 971.
Philanthus clarconis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 206. ¢ holotype. 1009.
Philanthus magdalenae Viereck, 1906, Trans. Amer. Ent. Soc., vol.
82, p. 240. ¢ holotype. 1010.
Plenoculus apicalis Williams, 1914, Univ. Kansas Sci. Bull., vol. 8,
p. 175. ¢ holotype. 959.
Podalonia puncta Murray, 1940, Ent. Americana (n.s.), vol. 20,
p- 86. 9 allotype, 4 9 and 1 ¢ paratypes. 5246.
Stictia spinifera Mickel, 1916, Trans. Amer. Ent. Soc., vol, 42, p. 418.
g holotype, 1 ¢ paratype. 970.
Stictiella divergens Parker, 1917, Proc. U.S. Nat. Mus., vol. 52, p. 55.
& holotype, 1 g paratype. 969.
Stictiella melampous Parker, 1917, Proc. U.S. Nat. Mus., vol. 52,
p. 48. holotype, 1 g paratype. 970.
Tachysphex clarconis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 211. ¢ holotype. 948.
Tachysphex consimiloides Williams, 1918, Univ. Kansas Sci. Bull.
vol. 8, p. 164. ¢ holotype. 949.
Tachysphex crassiformis Viereck, 1906, Trans. Amer. Ent. Soc., vol.
82, p. 210. 9? holotype. 950.
Tachysphex crenuloides Williams, 1913, Univ. Kansas Sci. Bull.
vol. 8, p. 168. ¢ holotype. 951.
Tachysphex dentatus Williams, 1918, Univ. Kansas Sci. Bull., vol. 8,
p. 169. ¢ holotype. 952.
Tachysphex glabrior Williams, 1918, Univ. Kansas Sci. Bull., vol. 8,
p.171. gholotype. 953.
Tachysphex intermedius Viereck, 1906, Trans, Amer, Ent. Soc., vol.
82,p. 211. g holotype. 944.
Tachysphex nigrocaudatus Williams, 1918, Univ. Kansas Sci. Bull.,
vol. 8, p. 167. ¢@ holotype. 954.
Tachysphex plenoculiformis Williams, 1918, Univ. Kansas Sci. Bull.,
vol. 8, p. 167. 9 holotype. 955.
Tachysphex robustior Williams, 1913, Univ Kansas Sci. Bull., vol. 8,
p. 164. holotype. 956.
Tue Universiry ScueNce BULLETIN
528
Tachysphex sculptiloides Williams, 1913, Univ. Kansas Sci. Bull.,
vol. 8, p. 166. ¢ holotype. 957.
Tachysphex sepulchralis Williams, 1918, Univ. Kansas Sci. Bull.,
vol. 8, p. 169. ¢ holotype. 958.
Trypoxylon quintilis Viereck, 1906, Trans. Amer, Ent, Soc., vol. 82,
p. 206. ¢ holotype. 972.
Trypoxylon regularis Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 205. ¢ holotype (abdomen missing). 978.
SPHECIDAE—Paratypes
Bembecinus bishoppi Krombein and Willink. 19. 4193.
Bembecinus wheeleri Krombein and Willink, 19. 4194.
Ectemnius scaber rufescens Krombein. 19,14. 5188.
Eucerceris arenaria Scullen, 3 9,7 g. 4209.
Microbembex aurata Parker. 29,14. 967.
Philanthus tarsatus Smith. 19. 2115.
Podalonia clypeata Murray. 49. 2114.
Podalonia melaena Murray. 1g. 1051.
Podalonia mickeli Murray. 49,14. 1052.
Podalonia parallela Murray. 1g. 1050.
Podalonia occidentalis Murray. 39. 1055,
Podalonia pubescens Murray. 39,5 ¢. 8411.
Podalonia sericea Murray. 11 ¢,12 4. 4486.
Sphex ferrugineus Fox. 2 g (used for first ¢ description by Fernald,
1906, Proc. U.S. Nat. Mus., vol. 31, p. 831). 943.
Stictiella callista Parker,1 9. 968.
Trypoxylon sculleni Sandhouse. 1g. 2502.
TENTHREDINIDAE
Dolerus illini Ross. 1 ¢ ,3 g paratypes. 786.
Dolerus simulans Rohwer. 1 $ paratype. 787.
TIPHIDAE
Aglyptacros sulcatus Mickel and Krombein, 1942, Amer. Mid. Nat.,
vol. 28, p. 674. 9? holotype. 8366.
Astigmometopa emarginata Mickel and Krombein, 1942, Amer. Mid.
Nat., vol. 28, p. 668. 9? holotype. 3867.
TIPHITDAE—Paratypes
Brachycistis stictinotus Viereck. 1g. 5449.
Myrmosa nocturna Krombein, 4 3. 2603.
Myrmosa nocturna rufigastra Krombein. 2 4. 2604.
SATALOGUE OF Types IN SNow MusrEuM 529
VESPIDAE
Eumenes bolliformis Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 887. 19 and 1 ¢ cotypes, 1 9 paratype. 1043.
Eumenes cruciferorum Viereck, 1908, Trans. Amer. Ent. Soc., vol. 88,
p. 388. 19 and1 g cotypes. 1044.
Eumenes enigmatus Viereck, 1908, Trans. Amer. Ent. Soc., vol. 38,
p. 889. ¢ holotype, 1 ¢ paratype. 1045.
Eumenes globulosiformis Viereck, 1908, Trans. Amer. Ent. Soc.,
vol, 33, p. 886. 19 and 1 ¢ cotypes. 1046.
Eumenes marginilineatus Viereck, 1907, Trans. Amer, Ent. Soc.,
vol. 33, p. 881. ¢ holotype. 1047,
Odynerus anormiformis Viereck, 1908, Trans. Amer, Ent. Soc.,
vol. 33, p. 407. ° holotype. 980.
Odynerus canaliculatus Viereck, 1908, Trans. Amer. Ent. Soc.,
vol. 33, p. 392. ¢ holotype. 981.
Odynerus cochisensis Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 395. 1¢9and 1 cotypes, 839 and 2¢ paratypes. 982.
Odynerus congressus Viereck, 1908, Trans. Amer. Ent. Soc., vol. 83,
p. 405. ¢@ holotype. 983.
Odynerus delodontus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 196. ¢ holotype. 984,
Odynerus egregius Viereck, 1906, Trans. Amer, Ent. Soc., vol. 82,
p. 197. g holotype. 985.
Odynerus excentralis Viereck, 1906, Trans. Amer, Ent. Soc., vol. 32,
p. 198. 9 holotype. 986.
Odynerus gulielmi Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 408. holotype. 987.
Odynerus hidalgiformis Viereck, 1908, Trans. Amer, Ent. Soc., vol.
33, p. 891. 9 holotype, 1 9 paratype. 988.
Odynerus leionotus Viereck, 1906, Trans. Amer, Ent. Soc., vol. 32,
p. 198. ¢ holotype. 989.
Odynerus maricoporum Viereck, 1908, Trans. Amer, Ent. Soc., vol.
38, p. 897. g holotype. 991.
Odynerus microstictus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 199. 9? holotype. 990.
Odynerus paenevagus Viereck, 1906, Trans. Amer. Ent. Soc., vol. 82,
p. 199. ¢ holotype. 994.
Odynerus papagorum Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 894, 1¢ and 1 ¢ cotypes, 2 ° paratypes. 992.
Odynerus percampanulatus Viereck, 1906, Trans, Amer, Ent. Soc.,
vol. 82, p. 200. ¢ holotype. 996.
18—3378
580 Tue Universiry SCIENCE BULLETIN
Odynerus pimorum Viereck, 1908, Trans, Amer. Ent. Soc., vol. 38,
p. 405. holotype. 993.
Odynerus provisoreus Viereck, 1908, Trans. Amer. Ent. Soc., vol. 38,
p. 396. holotype, 1 g paratype. 995.
Odynerus pulverulenta Viereck, 1908, Trans. Amer. Ent. Soc., vol.
33, p. 406. ¢ holotype. 997.
Odynerus rectangulis Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 393. 19 and 1 g cotype. 998.
Odynerus spectabiliformis Viereck, 1908, Trans. Amer. Bint. .50Ci,
p. 392. gholotype. 1001.
Odynerus trachysomus Bohart, 1940, Pan Pacific Ent., vol. 16, p. 54.
3 holotype, ? allotype. 2974.
Odynerus yumus Viereck, 1908, Trans. Amer. Ent. Soc., vol. 33,
p. 397. gholotype. 1002.
Polistes exclamans arizonensis Snelling, 1954, Jour. Kansas Ent. Soc.,
vol. 27, p. 152. ¢ holotype, ¢ allotype, 5 9 paratypes. 4634.
Polistes exclamans Viereck, 1906, Trans. Amer. Ent. Soc., vol. 32,
p. 201. holotype. 4632.
Pterocheilus arizonicus Bohart, 1940, Ann. Ent. Soc, Amer., vol. 33,
p. 174. gholotype. 2972.
Stenodynerus beameri Bohart, 1948, Florida Ent., vol. 31, p. 75.
& holotype. 1353.
VESPIDAE—Paratypes |
Ancistrocerus neocallosus Bequaert. 19,4 ¢. 8368.
Leptochilus propodealis Bohart, 1 g¢. 3426.
Mischocyttarus flavitarsis var. centralis Bequaert. 89 ,2¢. 2402.
Mischocyttarus flavitarsis var. idahoensis Bequaert. 6 9 ,1 g . 2403.
Mischocyttarus flavitarsis var. navajo Bequaert. 499,22. 2401.
Odynerus boscii albivestis Bohart. 19. 2980.
Odynerus boscii azotopus Bohart. 19,14. 2981.
Odynerus cinnabarinus Bohart. 1g. 2580.
Odynerus erubescens Bohart. 19. 2975.
Odynerus exoglyphus Bohart. 1 3. 2978.
Odynerus exoglyphus albovittatus Bohart. 19. 2979.
Odynerus monotylus Bohart. 19. 2976.
Odynerus tylocephalus Bohart. 1g. 2977.
Polistes hunteri californicus Bohart. 29,1 4. 4205.
CATALOGUE OF Types IN SNOow MusEUM 531
Polistes hunteri clarionensis Bohart. 19. 4206.
Pterocheilus flavobalteatus cyathopus Bohart. 2 ¢. 2971.
Pterocheilus desertorum Bohart. 19,1 4. 4218.
Pterocheilus hirsutipennis Bohart. 1 3. 2970.
Pterocheilus micheneri Bohart. 2 ¢. 2967.
Pterocheilus timberlakei Bohart. 19,14. 2968.
Pterocheilus trachysomus Bohart. 19,1 ¢. 2969.
Pterocheilus trichogaster Bohart. 29,2. 2978.
Stenodynerus acarigaster Bohart. 19,1 ¢. 4637.
Stenodynerus acrophorus Bohart. 19,1 4. 4638.
Stenodynerus apache Bohart. 19,1 g . 4208.
Stenodynerus blandoides Bohart. 19. 3423.
Stenodynerus chiricahuae Bohart. 29,1 g. 4200.
Stenodynerus coronado Bohart. 19,2 ¢. 4204.
Stenodynerus bicornis cushmani Bohart. 1 ¢. 4199.
Stenodynerus fulvipes rufovestis Bohart. 29,1. 4630.
Stenodynerus fundatiformis gonosceles Bohart. 19. 8421.
Stenodynerus lixovestis Bohart. 1g. 4201.
Stenodynerus mcclayi Bohart. 19,1 ¢. 4639.
Stenodynerus minimoferus Bohart. 19,1 ¢. 4196.
Stenodynerus ochrogonius Bohart. 89 ,2 4. 8422.
Stenodynerus parapedestris arenosus Bohart. 19,1 ¢. 4680.
Stenodynerus polingi Bohart. 19,14. 4195.
Stenodynerus pulvivestis Bohart. 29,2 3%. 4198.
Stenodynerus rectangulis fraxieri Bohart. 19. 4641.
Stenodynerus saecularis rufulus Bohart. 19,1 ¢. 4202.
Stenodynerus siccus Bohart. 1 3. 4642.
Stenodynerus valliceps Bohart. 1 ¢. 8424.
Stenodynerus vanduzeei Bohart. 1 ¢. 3425.
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVUI, Pt. 1] DecempBer 20, 1956 [No. 8
A Revision of the Bees of the Genus Melissodes in
North and Central America. Part II
(Hymenoptera, Apidae)"
BY
Watiace E. LABErcE
Asstract: This is the second part of a monographic revision of the bee
genus Melissodes in North and Central America. Ten species belonging to
four subgenera are described. Ten names are relegated to synonymy. One
new species, M. mitchelli, is described.
TABLE OF CONTENTS
PAGE
IGE OCUGUON es pier Web aun P44 At 6 alc y eee 1a nv weale 4 bcos 583
mubgenits, Pachymelssodeg sti. Na cevin els hes cane ees toe 68d
SUDGEniS: APOMOGMSCOUER rumen Oy costs sinuw ves nie mn cutesoli eens 544
Sith emia, PROMONSSOGIOSEH. cu: aux tuden tay sce wll bcc pad lst video ta de an.b cw 558
DO MENUS aM UOMO USSOCOSs hy oc fan wel icone ce hikieuck buh 1046 eee 561
DUCE Cues ahulines ay riuumi hack Goan viol hl asi Banus 4 ue woe dea 557
INTRODUCTION
This is the second part of a revision of the bees of the genus
Melissodes in North and Central America. Ten species belonging
to four subgenera are treated. A total of 2,088 specimens repre-
senting these ten species have been examined.
Explanations of the methods used in describing the species, the
meanings of certain descriptive terminology and methods used in
taking certain measurements are described in a section on descrip-
tive methodology in the first part of this revision (LaBerge, 1956).
The reader is also referred to the first part of this revision for a
complete list of acknowledgments. Certain persons have been
of invaluable aid in comparing specimens sent to them with type
specimens in their care, in lending types and in generously giving
of their time and advice. I am grateful to Dr. I. H. H. Yarrow of
* Contribution No, 914, Department of Entomology, University of Kansas.
(588 )
5384 Tur Universiry SCIENCE BULLETIN
the British Museum (Natural History) for generously giving of
his time to make comparisons with types in his care and for informa-
tion concerning those types. Mr. K. V. Krombein of the U. S.
National Museum is especially thanked for answering questions con-
cerning types in his care and for facilitating the loan of certain
specimens including paratypes and cotypes. Professor P. H. Tim-
berlake of the Citrus Experiment Station, Riverside, California,
Dr. E. S. Ross of the California Academy of Sciences, San Francisco,
California, Dr. Hugo G. Rodeck of the University of Colorado,
Boulder, Colorado, and Dr. T. B. Mitchell of North Carolina State
College, Raleigh, North Carolina, are thanked for the generous loan
of type specimens in their care. I am grateful to Dr. C. D. Michener
of the University of Kansas, Lawrence, Kansas, for his continued
interest and guidance without which this study would have been
much more difficult. Much of the research involved in this work
was done while holding a National Science Foundation Fellowship
at the University of Kansas during the academic years of 1952-
1954 and the author is grateful for this aid.
Subgenus Tachymelissodes LaBerge
Tachymelissodes LaBerge, 1956, Univ. Kansas Sci. Bull, vol.-87, pe L170.
Type species. Melissodes dagosa Cockerell, 1909, by original
designation.
This subgenus includes three species of Melissodes which have
the distal pubescent bands of the metasoma narrow, situated on
the apices of terga 2 to 4, subequal in width to each other and of
about equal width across each tergum in both sexes. In addition,
the males have short antennae which barely reach or do not reach
the first metasomal tergum in repose and have the minimum length
of the first flagellar segment equal to half or more of the maximum
length of the second segment.
Key TO THE SPECIES OF THE SUBGENUS ‘Tl ACHYMELISSODES
Males
J, Apical portion of pygidial plate half or less as broad as basal portion
of plate; labrum black; mandible with large basal yellow spot;
clypeus all or partly black... iy ea, sphaeralceae
Apical portion of pygidial plate broader than half basal width of
plate; labrum often with median yellow spot; mandible usually
with basal yellow spot; clypeus usually yellow except dark spots
at tentorial pits.....+..., ieee Pee: oes
2(3). Basal areas of terga 1 to 3 with ground spaces dulled by dense
transverse shagreening; labrum without yellow spot; clypeus with
MeELIssopEs IN NoRTH AND CENTRAL AMERICA 535
small regular punctures separated by less than one puncture
Widlilivooe £. deeerbene: Walla bale én this’ .. .opuntiella
Basal areas of terga 1 to 8 with ground’spaces shiny, with delicate
transverse shagreening which scarcely dulls surface, or unsha-
greened; labrum usually with small median yellow spot; clypcus
with coarse punctures which are irregular in size and spacing,
dagosa
Females
iy Clypeal punctures small, round, evenly spaced, separated by one
puncture width or less (mostly by, half a puncture width or
less); distal pale band of tergum 2 reaches apex of tergum
medially, not interrupted medially; hairs of inner surfaces of
hind basitarsi yellow or pale red P } 2
Clypeal punctures coarse, some elongate, some deparnts od by: more
than one puncture width at least basally; distal pale band of
tergum 2 separated from apex of tergum medially, usually nar-
rowly interrupted medially hairs of inner surfaces of hind basi-
tarsi often dark brown or black, occasionally yellowish red,
dagosa
2(1). Terga 2 and 3 with interband zone with small shallow punctures
and ground areas dulled by dense transverse shagreening; basal
half of tergum 1 with small shallow punctures, ground areas
dulled by dense shagreening; mesoscutum with no brown hairs,
or brown hair patch extends anteriorly to a transverse line at
about middle of tegulae .... opuntiella
Terga 2 and 8 with interband zone coatiely. and ircegulash punc-
tate, some punctures 2 or 3 times diameter of others, ground
areas shagreened but moderately shiny; basal half of tergum 1
with distinct punctures, ground areas shagreened but moderately
shiny; mesoscutum with patch of brown hairs usually extending
forward to or beyond a transverse line at anterior margins of
ROMUING 4s peels h ceva. Winey Gey age. Antik: | SORBORG Oeae
Melissodes (Tachymelissodes) dagosa Cockerell
Melissodes dagosa Cockerell, 1909, Canad. Ent., vol. 41, p. 128; 1928; Univ.
Colorado Studies, vol. 16, p. 114; Lore Knowlton and Bailey, 1950, Utah
State Agric. Coll., mimeo. ser. no. ‘871,
Melissodes calloleuca Cockerell, 1924, Bin Pacific Ent., vol. 1, p. 55 (new
synonymy ).
This is the most common and widespread species of this subgenus
The females can be recognized by the coarse clypeal punctation,
the usually narrowly interrupted and slightly arched pubescent band
of tergum 2 and the hairs of the inner surfaces of the hind basi-
tarsi often being dark brown to black. The males can be recognized
by the ground areas of the metasoma and clypeus being shiny, the
coarse clypeal punctation and the shape of the pygidial plate as
described below.
Female: Measurements and ratios: N, 20; length, 10-12 mm.;
536 Tur Universiry SCrENCE BULLETIN
width, 3.5-4.5 mm.; wing length, M = 3.54 + 0.169 mm.; hooks in
hamulus, M = 12.10 + 0.049; flagellar segment 1/segment 2, M =
Structure and color: Integument black except as follows: apical
half of mandible usually red with median golden macula; lower
surface of flagellum (except segment 1 and basal half of segment 2)
yellow to dark red; eyes blue or greenish blue; tarsi dark reddish
brown, claws red basally; tibial spurs pale yellow to pale brown;
apices of terga 2-4 (beneath pale pubescent bands) sli ghtly hyaline,
brown; apex of tergum 1 usually narrowly hyaline, yellow to brown;
sterna often red basally and hyaline apically, except last 2 or 3;
wing membranes slightly infumate, veins dark reddish brown to
black; tegulae piceous.
Clypeus coarsely and irregularly punctate, punctures, especially
basally, often separated by one to two puncture widths, several
punctures usually elongate, ground areas shiny, slightly or not at all
shagreened; supraclypeal area with deep round punctures widely
separated medially, shiny; galeae shiny, unshagreened and sparsely
punctate above; maxillary palpal segments in ratio of about
3.0:2.5:2,0:1.5. Mesoscutum with punctures small, round, separated
by 2 to 4 puncture widths posteromedially and crowded anteriorly
and peripherally, ground shiny; scutellum with punctures similar to
those of mesoscutum but separated mostly by about 2 puncture
widths, ground shiny; metanotum with small crowded punctures,
shiny dorsomedially; lateral surfaces of mesepisterna with deep
punctures separated mostly by half of one puncture width or less,
ground areas shiny, unshagreened; propodeum with dorsal surface
irregularly reticulorugose, dulled by fine tessellation. Metasomal
tergum 1 with basal % or slightly more with indistinct round punc-
tures separated by one puncture width or less, ground areas dulled
by fine transverse shagreening, moderately shiny; tergum 2 with
interband zone with small, indistinct punctures separated by 1 to 2
puncture widths medially and by 1 puncture width or less laterally,
ground areas dulled by dense, fine shagreening, moderately shiny,
especially laterally; interband zones of terga 3 and 4 similar to that
of tergum 2 but punctures more crowded.
Hair: On head silvery white except as follows: in darkest speci-
mens hairs of clypeus, labrum, mandibles and vertex between apices
of compound eyes dark brown, in palest specimens labral hairs
ochraceous and with a few short brown hairs on vertex between
apices of compound eyes and lateral ocelli, Mesoscutum often
ME.issopEs In NortH AND CENTRAL AMERICA 587
with posteromedian patch of dark brown hairs extending forwards
to a transverse line at middle of tegulae, occasionally dark meso-
scutal hairs absent, usually posteromedial area with few or no hairs;
scutellum usually with the sparse median hairs brown; mesepisterna
often with hairs of anterior, ventral and lower lateral surfaces dark
brown, occasionally all pale; pale hairs of dorsum of thorax white
to pale yellow, laterally and on propodeum white. Metasomal
tergum 1 with long hairs of basal % white; tergum 2 with distal and
basal pubescent bands silvery white, interband zone with short,
brown, subappressed hairs, distal band usually narrowly inter-
rupted medially and separated from apex of tergum in median
third or less by narrow zone of short, simple, brown hairs (often
rubbed off); tergum 3 and 4 similar to tergum 2 but distal pale
bands not interrupted medially nor separated from apices of terga
by zones of brown hairs and basal bands consist of dark brown
tomentum; terga 5 and 6 with long, appressed, dark brown hairs,
white to ochraceous lateral tufts present in pale specimens; sternal
hairs dark brown with fringes of long white hairs on apices of all
but last two sterna, in darkest specimens apical fringe of hairs pale
only laterally. Legs with white hairs except red or yellow on inner
surfaces of tarsi and dark brown hairs on basitibial plates in palest
specimens; in darkest specimens legs with dark brown hairs except
as follows; a few white to ochraceous hairs on outer surfaces of
fore and middle tibiae, scopal hairs silvery white except just below
basitibial plates and at apices of basitarsi; scopal hairs with rela-
tively few branches (usually 3 to 6 on either side of rachis) and
with strong rachises which extend beyond branched portion of hairs
by about % length of latter.
Male: Measurements and ratios: N, 20; length, 8-11 mm.; width,
2.5-3.5 mm.; wing length, M = 3.25 + 0.258 mm.; hooks in hamulus,
M = 11.10 + 0.280; flagellar segment 2/segment 1, M = 1.69 +
0.029.
Structure and color: Color as in female with the following
differences: clypeus pale yellow except dark spots at tentorial pits;
labrum cream-colored except narrow dark brown apical margin;
mandible usually with large triangular basal yellow spot, often
reduced but rarely absent; flagellum yellow below, except basal
half to two thirds of first segment, dark brown above; eyes green
to blue; terga 1-6 with apical margins (beneath pale pubescence)
hyaline, usually colorless, occasionally yellowish. First flagellar
segment with minimum length equal to half of maximum length of
538 Tur University SCIENCE BULLETIN
second segment or more (Fig, 12); penultimate flagellar segment
approximately two-thirds as broad as long; antennae reaching base
of first metasomal tergum in repose. Integumental sculpturing and
structure as in female with the following differences: clypeal punc-
tures usually smaller; mesoscutum with posteromedian area usually
with slightly more crowded punctures; metasomal terga with
ground areas shiny, with extremely fine and sparse shagreening or
none at all, Pygidial plate with apical portion at least as broad
as half of base of plate and usually much broader.
Gonostylus capitate and slightly pointed inwards; inner surface
of gonocoxite just below gonostylus with abundant short acute
hairs and few blunt hairs admixed; penis valve with abundant short
hairs on lateral surface below dorsal lamella (see LaBerge, 1956,
Figs. 118 to 121). Sternum 7 with median plate with thick short
neck, flat apical margin, and blunt apex directed posterolaterally,
inner margin straight, slanted inwards towards apex, with abundant
short hairs ventrally. Sternum 8 truncate apically with several
relatively long hairs at apex, median ventral carina well separated
from apex, short, blunt, not bilobed apically.
Hair: Much as in female but generally paler; hairs of head, thorax
and legs never dark brown except occasionally with a few brown
hairs on vertex of head, in posteromedian area of mesoscutum and
medially on scutellum; metasomal hairs as in female but distal
band of tergum 2 not interrupted and usually reaching apex medi-
ally, tergum 5 with pale distal band as on tergum 4, and terga 6
and 7 with long pale hairs; legs with yellow to red hairs on inner
surfaces of tarsi, otherwise white. Often all pale hairs are golden
yellow rather than being white; the former bees are considered to
be newly emerged, or at least young adults, and the latter are con-
sidered to be older faded specimens.
Bionomics: Nothing has been published concerning the nesting
habits of this species. Little is known concerning its flower prefer-
ences, but from the available records it appears that M. dagosa
is a polylectic bee and visits a large variety of plants for pollen as
well as for nectar. Females have been taken from plants of the
families Asclepiadaceae, Leguminosae, Compositae and Cruciferae
in that order of abundance. The only records on composites, how-
ever, are from thistles, whereas three genera of crucifers have
yielded females of this species. M. dagosa is probably important
in the pollination of alfalfa at least in Utah where this bee seem-
ingly is most abundant.
ME issopEs IN NortH AND CENTRAL AMERICA 539
Type material: Holotype male of dagosa from Osborn’s Ranch,
Grand Coulee, Washington, is the property of the California Acad-
emy of Sciences but temporarily deposited in the collection of the
Citrus Experiment Station, Riverside, California. The holotype
female of calloleuca from Saltair, Utah, is in the California Academy
of Sciences, San Francisco, California.
Distribution: This species is widespread in the Rocky Mountains
from southern Idaho and Washington in the north to middle eastern
California in the southwest and the western half of Colorado in
the east. It has been collected from May 19 to August 8 but mainly
in June and July. In addition to the type material, 183 females and
184 males have been examined from the localities listed below.
Localities mentioned in the literature and localities of the type
material are included..
Cauirornia: Big Pine, Inyo Co.; Bridgeport, Mono Co.; Cedar-
ville, Modoc Co.; Deep Springs, Inyo Co.; Diaz Lake, Inyo Co.;
Lone Pine, Inyo Co.; Mammoth Lakes, Mono Co.; Topaz, Mono
Co.; Wendel, Lassen Co.; Westgard Pass (7 miles W.), Inyo Co.
Cotorapo: Alamosa; Boulder; Delta; Eckert; Monte Vista. Ipano:
Aberdeen (5 miles N.); Hot Springs, Owyhee Co. Nevapa: Con-
nor’s Pass, White Pine Co.; Dumphy; Fallon; Lovelock; Reno (Sky
Ranch), Orecon: Warner Lake, Lake Co.; Harney Co.; Steen
Mts., Harney Co.; Umapine; Vale. Uran: Callao; Cedar Mts. (S.E,
end of), Tooele Co,; Delta; Dugway Parade Ground (S. E. of Camel
Back Mt.), Tooele Co.; Elberta; Erda; Fillmore; Fish Springs; Fort
Duchesne; Gandy; Grantsville; Hinckley; Jensen (10 miles E.);
Kelton; Layton; Lehi; Little Salt Lake, Iron Co.; Lynndyl; Mc-
Cormick; Milford; Nephi; Parowan; Park Valley; Payson; Penrose;
Promontory; Saltair; South Fork, Ogden River; West Point. Wasn-
incton: Grand Coulee (Osborn’s Ranch); Lind; North Yakima;
Yakima City. Wyominc: Carbon Co.; Green River; New Castle.
Flower records: Asclepias sp., Cirsium sp., Medicago sativa,
Melilotus sp., M. alba, Norta altissima, Sphaeralcea sp., Thelypo-
dium brachycarpa.
Melissodes (Tachymelissodes) opuntiella Cockerell
Melissodes opuntiella Cockerell, 1911, Canadian Ent., vol. 48, p. 131.
Melissodes albocincta Cockerell, 1919, Ann. Mag. Nat. Hist., ser, 9, vol. 2,
p. 119 (new synonymy ).
This is the smallest species of this subgenus. It differs from
M. dagosa in both sexes by the clypeal and pubescence characters
described in the diagnosis of the latter. Females of opuntiella
540 Tur Universiry SCIENCE BULLETIN
can be distinguished from those of sphaeralceae by the punctation
of the metasoma and thorax as described below. Males of opun-
ticlla differ from those of sphaeralceae in having a pygidial plate
similar to that of dagosa and by usually having an all yellow clypeus
(partly black in a few specimens ).
Female: Measurements and ratios: N, 20; length, 8-9 mm.; width,
3-4 mm.; wing length, M = 2.85 + 0.016 mm.; hooks in hamulus,
M — 10.25 -+ 0.138; flagellar segment 1/segment 2, M = 2.05
(0.047.
Structure and color: Integumental color as in M. dagosa except
as follows: apices of terga 2-4 broadly hyaline, colorless to yellow
and apex of tergum 1 narrowly hyaline, colorless, hyaline area of
tergum 1 equals about half of hyaline area of tergum 2 medially;
wing membranes clear; tegulae piceous to clear yellowish brown.
Integumental sculpturing as in M. dagosa except as follows:
clypeus with small round regular punctures usually separated by
less than one puncture width, without markedly elongated punc-
tures; maxillary palpal segments in ratio of about 8.6:2.25:2.1;
mesoscutum with punctures smaller but distributed as in dagosa;
propodeum dulled dorsally by dense fine irregular rugae and dense
fine tessellation; metasomal tergum 1 with basal % to % with minute,
round punctures separated mostly by one puncture width basally
to two to four puncture widths apically, ground area dulled by
exceedingly minute and dense shagreening; terga 2-4 with small
round widely spaced punctures in areas basal to apical pubescent
bands, ground areas opaque, dulled by exceedingly fine and dense
shagreening.
Hair: Color as in M. dagosa with the following differences:
Hairs of head and thorax white except occasionally pale brown in
posteromedian area of mesoscutum; metasomal tergum 2 with apical
pale band not interrupted or separated from apex of tergum medi-
ally; hairs of interband zones of terga 2-4 shorter and usually pale
brown; basal band of tergum 3 white and of tergum 4 pale brown
to white; terga 5 and 6 with lateral tufts of white hairs small or
absent; hairs of legs white except as follows: on hind tibiae pale
yellow, on inner surfaces of hind basitarsi yellow to red, on inner
surfaces of fore and middle tarsi red, on outer surfaces of fore
tarsi brown to red.
0.032.
saul ee
MEtissopEs IN NortH AND CENTRAL AMERICA 541
Structure and color: Integumental color as in M. dagosa except
as follows: labrum entirely black; basal mandibular yellow spot
always large; clypeus occasionally blackened posteriorly. Struc-
ture as in M. dagosa with the following differences: antennae not
surpassing propodeum in repose, penultimate flagellar segment %
to % as broad as long; clypeal punctation and maxillary palpi as in
female; sculpturing of terga as in female but punctures slightly
larger and more abundant.
Genital capsule as in M. dagosa but gonostylus usually less capi-
tate and with fewer hairs on dorsal and outer surfaces, penis valves
with fewer hairs laterally and gonocoxite with more abundant blunt
hairs on inner surface near apex. Sternum 7 with median plate
long-necked, oval in outline and with few minute hairs ventrally;
apodeme not capitate. at apex (Fig. 1). Sternum 8 emarginate at
apex, with a distinct median ventral carina, narrower than in M.
dagosa and distinctly narrower medially than at level of upper
fourth (Fig. 2).
Hair: Much as in M. dagosa but paler; never with brown hairs
on head or thorax; never with hairs golden yellow, but occasionally
pale ochraceous on dorsum of head and thorax.
Bionomics: In keeping with its name this species shows some
preference for pollen of cacti of the genus Opuntia. However, the
few flower records available show that M. opuntiella will take pollen
from other sources such as the legume Lindheimera texana, the
mallow Sphaeralcea pedatifida and the composites Cirsium sp. and
Verbesina encelioides. This variety of flowers bespeaks a rather
polylectic species rather than the oligolecty inferred in its name.
Type material: Holotype male and allotype female of opuntiella
from Brownsville, Texas, October 23, 1908, Jones and Pratt col-
lectors, on Opuntia lindheimeri are in the U. S. National Museum
(U.S.N.M. Type No. 14105). Unfortunately, the allotype female
is not conspecific with the holotype male but belongs to the follow-
ing species, M. sphaeralceae. The holotype female of albocincta
from Mexico is in the U. S. National Museum (U.S.N.M. Type
No. 23218).
Distribution: This species ranges from southern Utah and south-
ern California in the northwest to eastern Texas and northeastern
Mexico in the southeast. It has been collected from March 25 to
October 23 but mainly during April and May. In addition to the
type material, 49 females and 34 males have been examined from
the localities listed below.
542 Tue University ScreENCE BULLETIN
Arizona: Wickenburg. Carirornia: Hopkins Well (1.8 miles
E.), Riverside Co. Texas: Brownsville; Corpus Christi; Cotulla;
Dilley; Hidalgo; Kerrville; Laredo (2 and 18 miles N.); Pharr (10
miles S. W.); Progresso; Rio Grande (6 miles E. and 20 miles S. E.);
San Diego; San Ygnacio; Sonora; Uvalde. Uran: Heber. Co-
anuiLa: San Pedro de Colonias (3700 feet altitude).
Flower records: Cercidium texanum, Cirsium sp., Helenium
microcephalum, Helianthus annuus, Lindheimeri texana, Opuntia
sp., O. lindheimeri, Parkinsonia sp., Prosopis sp., Sphaeralcea sp.,
S. pedatifida, Verbesina encelioides.
Melissodes (Tachymelissodes) sphaeralceae Cockerell
Melissodes sphaeralceae Cockerell, 1896, Entomologist, vol. 29, p. 804; 1897,
Bull. New Mexico Coll. Agric. and Mech. Arts, no. 24, p. 20; 1898, Bull.
Sci. Lab. Denison Univ., vol. 11, p. 66; 1902, Entomologist, vol. 35, p. Wa
1904, Entomologist, vol. 37, p. 8; 1906, Trans. Amer. Ent. Soc., vol. 32, pp.
75, 86; 1906, Trans. Amer. Ent. Soc., vol. 32, p. 309.
This species is closely related to M. opuntiella. The males can
readily be distinguished by the form of the pygidial plate as de-
scribed below. The female can be separated from the female of
dagosa by the finer punctation of the clypeus and from the female
of opunticlla by the large patch of brown hairs on the mesonotum
and by the more coarsely punctate metasomal terga as described
below.
Female: Measurements and ratios: N, 12; length, 9-11 mm.;
width, 3-4 mm.; wing length, M = 3.08 + 0.230 mm.; hooks in
hamulus, M = 11.41 +0.244; flagellar segment 1/segment 2, M =
2.44 + 0.049.
Structure and color: Integumental color as in M. dagosa with
the following differences: eyes gray to grayish blue; tegulae piceous
but clear brown posteromedially; tergum 1 with a broad colorless
hyaline apical margin; terga 2-4 (beneath apical pubescent bands)
hyaline, colorless or yellow; wing membranes clear or somewhat
milky apically.
Clypeus finely and regularly punctate, punctures round, separated
mostly by half of one puncture width or less, ground areas smooth
and shiny; supraclypeal area with round punctures similar to those
on adjacent portion of clypeus, ground areas smooth and shiny;
galeae as in M. dagosa; maxillary palpal segments in ratio of about
9.5:2:2:1, Mesoscutum with large round punctures separated by
one to four puncture widths posteromedially and by half of a punc-
ture width or less anteriorly and laterally, ground areas shiny, often
with exceedingly fine reticulate shagreening; scutellum similarly
Me issopEs IN NortTH AND CENTRAL AMERICA 543
sculptured but punctures distinctly smaller than on mesoscutum;
metanotum and mesepisterna sculptured as in dagosa; propodeum
as in dagosa but dorsal surface with large distinct punctures. Meta-
somal tergum 1 with basal % or slightly more with small round
punctures, in basal half punctures separated mostly by less than
one puncture width, distal to these punctures more widely separated
(especially laterally) except for a single row of close-set punctures
extending across tergum at about % distance from base, ground
areas moderately shiny, dulled by exceedingly fine transverse sha-
greening; interband zones of terga 2 and 3 with round, irregularly
spaced punctures of irregular size, separated from nearest neighbors
by less than half to four puncture widths (sparser medially than
basally or apically), punctures crowded beneath basal pubescent
band and beneath basal portion of apical pubescent band, ground
areas as in tergum 1; tergum 4 with interband zone more densely
punctate than in terga 2 or 8.
Hair: On head and thorax white except long brown hairs on
vertex of head between apices of compound eyes, mesoscutal patch
of brown hairs which extends forwards well beyond a transverse
line at anterior margins of tegulae and scutellum with large median
brown hair patch. Metasomal terga with hairs as in dagosa with
the following differences: hairs of interband zones shorter and
more appressed, basal band of tergum 3 pale, distal pale bands
of terga 2 and 3 consisting of much shorter, appressed pubescence,
that on tergum 2 not interrupted and reaching apex medially.
Hairs of legs white except as follows: ochraceous or pale brown
distally on middle tibiae and basally on hind tibiae, orange to yel-
low on inner surfaces of tarsi and tibiae; scopal hairs with 4 to 10
branches on either side of rachis.
Male: Measurements and ratios: N, 19; length, 9-10 mm.; width,
2.0-3.5 mm.; wing length, M = 3.82 -+ 0.247 mm.; hooks in hamulus,
M = 10.47 + 0.188; flagellar segment 2/segment 1, M —1.79 +
0.010.
Structure and color; Integumental color as in M. dagosa with
the following differences: clypeus partial! or wholly black; man-
dible with large basal yellow spot never reCuced or absent; labrum
black; eyes gray or grayish blue.
Minimum length of first flagellar segment equals more than half
of maximum length of second segment; penultimate flagellar seg-
ment approximately half as broad as long; antennae barely reach-
ing base of first metasomal tergum in repose; galeae and maxillary
palpi as in female. Sculpturing as in female with the following
544 Tur Untversiry SCIENCE BULLETIN
differences: punctation of terga usually somewhat denser, ground
areas of interband zones occasionally dulled by dense shagreening.
Pygidial plate with apical portion less than half as broad as base
of plate and often as narrow as a third basal width.
Gonostylus capitate as in dagosa, otherwise capsule as in opun-
tiella, Sternum 8 as in opuntiella but median ventral carina sharper
and hairs at apex of sternum somewhat more abundant. Sternum
7 as in opuntiella.
Hair: On head and thorax as in female but often without brown
hairs on vertex and mesoscutal brown patch usually smaller and
occasionally absent. Metasomal hairs as in female but tergum 5
with distal pale band as in terga 2-4, tergum 6 usually without
lateral tufts of pale hairs and sternal hairs all white. Legs with
white hairs except orange to yellow on inner surfaces of tarsi. Body
hairs not golden yellow as in dagosa on any specimens now before
me.
Type material: Lectotype male of sphaeralceae, here designated,
from Santa Fe, New Mexico, July, is in the collection of the Univer-
sity of Colorado Natural History Museum, Boulder, Colorado.
Distribution: This species ranges from Durango, Mexico, and
Brownsville, Texas, north to the Sandia Mountains in New Mexico,
and west to Arizona. It has been taken from June 23 to September
4, but mainly in July and August. In addition to the lectotype,
20 males and 12 females have been examined from the localities
listed below. This list includes localities reported in the literature.
Arizona: Navajo Co. New Mexico: Pecos, San Miguel Co.;
Sandia Mountains; Santa Fe. Texas: Brownsville. DuRanco:
Nombre de Dios; Yerbanis (Cuencame District). marco: Ix-
mequilpan. Duistrrrro Frperat: Guadalupe; México (city).
Flower records: Monarda sp., Sphaeralcea sp., S. angustifolia,
S. fendleri.
Subgenus Apomelissodes LaBerge
Apomelissodes LaBerge, 1956, Univ. of Kansas Sci. Bull., vol. 87, p. 1175.
Type species: Melissodes fimbriata Cresson, 1878, by original
designation.
This subgenus, originally intended to include two species (M.
fimbriata Cresson and M. apicata Lovell and Cockerell) which
occur in southeastern United States, now includes four species.
Melissodes baileyi Cockerell is provisionally included at this time
and a new species is described below. M. baileyi is known from a
Me tissopEs 1IN NortH AND CENTRAL AMERICA 545
single female which does not have a protruding clypeus, long galeae
or hooked galeal hairs. The first couplet in the key to the subgenera
of Melissodes (females) published previously (LaBerge, 1956,
p. 922) should accordingly be changed to read as follows:
1. Scopal hairs simple or weakly branched, if weakly branched, then
clypeus in profile protruding beyond eye by % width of eye or
MOTO 404i f5 tl I lv el Aa ine lens Mie Apomelissodes
Scopal hairs branched, usually abundantly so, if weakly branched, then
clypeus in profile not protruding beyond eye by as much as %
WICH MOL MOY Os herr ork Pea rie ein a VA tr Gey ore ws g
Key To THE SPECIES OF THE SUBGENUS APOMELISSODES
Males
1. Galeae (measured from base of first palpal segment) three times
as long as clypeus or longer; pygidial plate notched laterally but
not so deeply as to create the impression of two plates; apical
pubescent bands of terga 2-5 consist of one to three rows of long
silky short-branched hairs arising slightly anterior to one hair length
POM MA UAIS OU OTOA vere. iaive) steady 1 Gan ae hoe fimbriata
Galeae considerably less than three times as long as clypeus; pygidial
plate deeply notched laterally, often appearing like two plates
(a large high anterior plate and a small posterior plate on a lower
level); apical pubescent bands of terga 2-5 consist of several rows
of short, highly plumose hairs arising in a broad zone from bases of
baridsealmost towthirdine-of terfa. 6 3 ee oc 5 enews ee 2
2. Clypeus yellow except black notches posteriorly at tentorial pits,
coarsely punctate with median zone impunctate and moderately
shiny; galeae with sparse short hairs laterally......... _..mitchelli
Clypeus all or partly black, densely punctate and dull; galeae with
abundant short hairs laterally......... cali I Bye TER apicata
Females
Tee Scopele luins” Simiple, MitbreanOnetl: fee) Oe yy i ee eee 2
Scopal airs. weakly.byanchetles sisi, Gs iy ai, See ee ee 8
2. Galeae three times as long as clypeus or longer............ fimbriata
Galeae two times as long as clypeus or less.................. baileyi
8. Galeae with sparse short straight hairs laterally; clypeus with pos-
teromedian area slightly depressed or flattened, sparsely and coarsely
MUMOthle: fee on Ve es iy a levies Gere mitchelli
Galeae with abundant long hooked hairs laterally; clypeus with pos-
teromedian area not depressed or flattened, densely and coarsely
AUN LAL ACh Seema MINIS Far Rroes «Suey NING ete Clara gi i apicata
Melissodes (Apomelissodes) fimbriata Cresson
Melissodes fimbriata Cresson, 1878, Proc. Acad. Nat. Sci., Philadelphia, vol. 30,
p. 203; Fox, 1892, Ent. News, vol. 3, p. 29; Birkman, 1899, Ent. News,
vol. 10, p. 245; Cockerell, 1906, Trans. Amer. Ent. Soc., vol. 82, pp. 77, 87;
Smith, 1910, Ann. Report New Jersey State Museum, 1909, p. 693; Cresson,
1916, Mem. Amer, Ent. Soc., vol. 1, p. 118; Brimley, 1938, Insects of North
Carolina, p. 462.
546 Tue Untversiry ScrENCE BULLETIN
Melissodes perplexa, Cresson, 1878, Proc. Acad. Nat. Sci., Philadelphia, vol. 30,
p. 206 (misidentification of male).
Melissodes fimbriata is easily separated in both sexes from the
other members of this subgenus by use of the characters listed in
the key given above. Several other striking distinctions exist and
are described below.
Female: Measurements and ratios: N, 18; length, 11.0-13.0 mm.;
16), M= 1.95 + 0,002.
Structure and color: Integument black except as follows: lower
surfaces of flagellar segments 2-10, apical half of mandible, legs
(or at least distitarsi) and sterna (at least apically) dark reddish
brown; apical half of mandible usually with median golden macula;
eyes bluish gray to green; tegulae piceous; wing membranes some-
what infumate, yellow or yellowish brown, veins dark brown to
black.
Clypeus with small deep round dense regular-sized punctures
separated by half a puncture width or less, ground areas shiny,
unshagreened except sparsely so laterally and posteriorly; supra-
clypeal area densely punctate laterally, sparsely punctate medially,
ground areas medially dulled by fine tessellation; flattened area of
vertex between compound eye and lateral ocellus densely punctate,
punctures deep, round, separated by one puncture width or less;
galeae from base of first palpal segment to tips equal 3.5 to 4.0
times median length of clypeus, with exceedingly sparse short
straight hairs scattered over surface, ground moderately shiny
above, dulled laterally and below by reticular shagreening; maxil-
lary palpal segment in ratio of about 9.75:3.5:3:1, _Mesoscutum
with small round punctures separated mostly by one puncture width
or less posteromedially and less than half a puncture width ante-
riorly and peripherally, ground areas shiny, unshagreened; scu-
tellum as densely punctate as posterior declivous portion of meso-
scutum; mesepisterna with punctures slightly larger than on anterior
half of mesoscutum, separated mostly by half a puncture width or
less, ground areas shiny, unshagreened; metanotum densely punc-
tate medially, less so laterally, shiny, slightly shagreened laterally;
propodeum with dorsal surface with large coarse punctures except
medially, moderately shiny, surface dulled by irregular tessellation,
with posterior surface densely and coarsely punctate except medi-
ally, dulled by irregular tessellation except dorsomedial area, with
Me.issopEs IN NortH AND CENTRAL AMERICA 547
lateral surface densely and coarsely punctate, ground areas mod-
erately shiny to shiny. Metasomal tergum 1 sparsely punctate in
basal % of dorsal surface, punctures shallow, becoming denser ante-
riorly and laterally, ground dulled by coarse reticulotransverse
shagreening; tergum 2 with interband zone sparsely punctate, punc-
tures shallow and separated mostly by 2 puncture widths or more,
with apical area impunctate except for row of dense punctures near
anterior margin of apical pale band, with basal area densely punc-
tate, ground areas dulled as in tergum 1; terga 3 and 4 as in tergum
2, but interband zones progressively more densely punctate.
Hair: On head white, occasionally a few long dark brown hairs
on vertex behind median ocellus. Thorax with white hairs except
large mesoscutellar patch of dark brown to black hairs which ex-
tends forward beyond.a transverse line at anterior margins of tegu-
lae and scutellar hairs dark brown to black except posterior and
lateral fringes of white hairs; tegulae and humeral angles of wings
with abundant dark brown hairs. White hairs in basal % of dorsal
surface of metasomal tergum | extend to apex laterally, bare apico-
medially; tergum 2 with basal band of white highly plumose pu-
bescence, with apical band of white pubescence composed of long,
silky hairs with short branches, with interband zone with abundant
erect or suberect bristlelike white short hairs which are longer
at extreme sides; terga 3 and 4 similar to tergum 2 but basal pubes-
cent bands dark brown, apical bands denser and bristlelike hairs all
or mostly dark brown and longer; terga 5 and 6 with long ap-
pressed dark-brown hairs, usually with small tufts of white hairs
at extreme sides of tergum 5; sternal hairs reddish brown basally
and medially, white apicolaterally. Legs with hairs white except as
follows: outer surfaces of fore tarsi, outer surfaces of middle and
hind distitarsi, basally on hind tibiae and apically on fore and middle
tibiae brown; inner surfaces of tarsi and hind tibiae yellow to orange-
red; scopal hairs simple, white to ochraceous.
Male: Measurements and ratios: N, 20; length, 9.0 to 18.0 mm.;
width, 3.0-4.5 mm.; wing length, M = 21.34 + 0.869 mm.; hooks in
hamulus, M = 13.55 + 0.271; flagellar segment 1/segment 2, M =
4.20 += 0.256.
Structure and color: Integument black except as follows: clypeus
yellow except black notches posteriorly at tentorial pits, occasion-
ally posterolateral angles (mesad to tentorial pits) darkened and
apical margin dark reddish brown; labrum usually black, rarely with
small posteromedial yellow spot; mandible as in female; lower sur-
548 Tue Untversiry SCrENCE BULLETIN
faces of flagellar segments 2 to 11, distitarsi and apical margins of
sterna dark reddish brown; eyes gray to bluish gray; wing mem-
branes somewhat infumate yellowish brown, veins dark brown to
black.
Galeae long as in female; maxillary palpi as in female; minimum
length of first flagellar segment equals 4 to % of maximum length of
second segment. Sculpturing of head and mesosoma as in female.
Sculpturing of metasoma as in female except as follows: tergum 1
with basal % to % densely punctate; terga 2-5 with interband zones
somewhat more densely punctate, medially on tergum 2 punctures
separated mostly by one to two puncture widths; pygidial plate flat,
notched laterally at about a third of length from apex, apical portion
about 24 width of basal portion or more; sternum 6 shiny, shallowly
grooved medially, slightly emarginate apically, with coarse, round
punctures separated by one puncture width or less on each side
of median groove.
Terminalia described and figured by LaBerge (1956, Figs. li1-
113). Sternum 7 with median plate long-necked. Sternum 8
pointed medioapically.
Hair: Color as in female with the following differences: vertex
of head without dark hairs; long white hairs of metasomal tergum 1
reach apex; terga 2-5 with apical fringes of long white silky
short-branched pubescence arising in one to three rows at about
the length of one hair from apices of terga; terga 3-5 with abundant
erect bristlelike hairs in interband zones, those of tergum 2 white;
terga 6 and 7 with long dark-brown appressed hairs; sternal hairs
white to ochraceous except reddish brown posteromedially on
sterna 2-5 and dark brown on sternum 6. Legs with white hairs
except yellow to orange on inner surfaces of tarsi and hind tibiae.
Bionomics: The evidence available from labels on the specimens
indicates that M. fimbriata is an oligolectic bee dependent upon
plants of the genus Oenothera (Onagraceac) for pollen as well
as nectar. Out of 15 collections with flower data, 13 are from
Oenothera, whereas the other two consist each of one male, the
first from Kneiffia sp. (Onagraceae) and the second from Helian-
thus petiolaris (Compositae ).
Type material: Female lectotype and male lectoallotype of fim-
briata from Texas are in the Academy of Natural Sciences of
Philadelphia, Pennsylvania. The lectoallotype male of perplexa
from Georgia is, unfortunately, not conspecific with the lectotype
female, but is the same as the male lectoallotype of fimbriata.
Distribution: This species ranges from Virginia in the northeast
ME tissopEs In NortH AND CENTRAL AMERICA 549
to Georgia in the southeast, west to Texas and northwest to south-
western Kansas. It has been collected from April 25 to July 27. In
addition to the type material, 18 females and 27 males have been
examined, These records are so widely and sporadically distrib-
uted that it is deemed best to list here the collection data in full.
Grorcra: 1 4, lectoallotype of M. perplexa Cresson. Kansas:
Garden City, 1 ¢ , June 3, 1952, Cheng Liang; Tribune, 1 g, June
16, 1949, C. D. Michener and R. H. Beamer, on Helianthus petio-
laris. Mississippi: Hattiesburg, 1 ¢ , July 22, 1944, C. D. Michener;
Mississippi State College, 1 ¢, April 26, 1949, 19, June 5, 1947,
student collectors. Norra Carourna: Burgaw, 29 ¢?, May 18,
1950, T. B. Mitchell, on Oenothera sp.; Garford, 1 9, June 7, 1927,
C. S. Brimley; Lexington (12 miles E.), 19, June 13, 1950, T. B.
Mitchell, on Oenothera sp.; Merry Oaks, 1 g, July 27, 1926, T. B.
Mitchell, on Oenothera sp.; Raleigh, 1 9 , May 25, 1922, on Oeno-
thera sp.; 2g g, July 18, 1922, on Oenothera sp. 14, July 20,
1927, on Oenothera sp., 19 and 1 3, May 23, 1929, on Oenothera
sp., 29 9, May 25, 1929, on Oenothera sp., 29 9, May 25, 1938,
on Oenothera sp., 1g, July 7, 1933, on Oenothera sp., 1 g¢, May
26, 1935, 1 9 , May 15, 1938, on Oenothera laciniata, 1 9, May 29,
1938, 1 g , May 15, 1939, 1 ¢ , April 25, 1942, 1 g and29 9, May
29, 1950, on Oenothera sp., 4 ¢ g , May 17, 1951, on Oenothera sp.,
T. 'B. Mitchell; Willard, 2g g and 19, May 11, 1951, T. B.
Mitchell. Texas: Kerrville, 1 ¢, May 22, 1908, F. C. Pratt; 13,
“Collection T. Pergande”; 3 ¢ g, “Collection Belfrage”. Vinctnta:
Dawson’s Beach, 1 4, June 6, 1931, on Kneiffia sp., J. C. Brimwell.
A single male in the collection of the Tlinois Natural History
Survey is labeled “Tamarack Bog” with the numeral 4 above.
Melissodes (Apomelissodes) baileyi Cockerell
Melissodes baileyi Cockerell, 1906, Ann. Mag. Nat. Hist., ser, 7, vol. 17, p. 361.
This species, known only from the female holotype, seemingly
is closely related to M. fimbriata. It is similar to the latter in having
simple scopal hairs, but differs in having a relatively flat clypeus
and short galeae. It can be separated from the female of M.
apicata by the lack of abundant hooked galeal hairs. If it were
not for the simple scopal hairs, M. baileyi would probably be
placed in the subgenus Tachymelissodes. However, the simple
scopal hairs, together with the form of the metasomal pubescence,
seems to relate this species to M. fimbriata and it is, accordingly,
placed in the subgenus Apomelissodes. Study of the male ter-
minalia will undoubtedly solve this problem.
Tue Universrry ScreENCE BULLETIN
Female: Measurements and ratios: N, 1; length, 11.0 mm.;
width, 4.0 mm.; wing length, 3.33 mm.; hooks in hamulus, 13;
flagellar segment 1/segment 2, 1.97.
Structure and color: Integument black except as follows: Man-
dible dark reddish brown in apical %, apical half with large
median golden macula; flagellar segments 3 to 10 red beneath;
eyes gray; legs and sterna dark reddish brown; metasomal tergum
1 with apical margin hyaline, colorless; tegulae piceous; wing mem-
branes hyaline, colorless, veins dark brown.
Clypeus with dense round punctures of regular size separated
mostly by half of one puncture width or less, ground areas moder-
ately shiny; supraclypeal area impunctate medially, shiny, coarsely
punctate laterally; flattened area of vertex posteromesad of apex
of compound eye with abundant round punctures of irregular size
separated mostly by one puncture width or less, ground shiny;
galeae about twice as long as median length of clypeus, shiny, un-
shagreened above or only delicately so, with sparse, short, straight
hairs and sparse punctures; maxillary palpi 4-segmented, segments
in ratio of about 2.5:2:2:1. Mesoscutum with small round punc-
tures separated by one or slightly more puncture widths postero-
medially and by half a puncture width in anterior half and _peri-
pherally; scutellum with small punctures of about same size as on
adjacent part of mesoscutum, separated by one to half of one
puncture width medially and less peripherally; ground areas of
mesoscutum, scutellum and mesepisternum shiny, unshagreened;
mesepisternum with large shallow punctures separated by half
of one puncture width or less, many punctures not well defined
posteroventrally; metanotum with abundant small punctures half
size of punctures on posterior part of scutellum or a little larger,
dense medially, sparse laterally, ground shiny medially and dull
laterally; propodeum with dorsal surface reticulorugose except
medially, with a few large punctures posteriorly, surface dulled
by dense fine tessellation, with posterior surface densely punctate
except upper median impunctate triangle, ground areas dulled
by dense tessellation. Metasomal tergum 1 with dorsal surface
punctate in basal half, punctures separated by one to two puncture
widths apically in median half and by half a puncture width
or less basally and laterally to apex, ground areas moderately
shiny, with dense coarse transverse shagreening; tergum 2 with
interband zone with exceedingly small punctures, almost impunc-
tate medially but separated by one puncture width or less laterally,
apical area impunctate, basal area densely punctate, ground areas
ME.issopes in NortH AND CENTRAL AMERICA 551
opaque, dulled by dense fine transverse shagreening; terga 3 and
4 similar to tergum 2 but more densely punctate in interband zones.
Hair: On head white; mesosomal hairs white except as follows:
mesoscutellar patch of dark brown hairs extending forward to a
transverse line at anterior margins of tegulae, scutellar hairs dark
brown except peripherally. Metasomal tergum 1 with long white
hairs in basal half medially and to apex laterally; tergum 2 with
basal band of white pubescence, apical band of long white silky
short-branched pubescence, interband zone of short simple bristle-
like dark-brown hairs about equal in width to apical pale band,
apical and basal pale bands connected laterally by white erect
hairs; terga 3 and 4 similar to tergum 2 but basal bands dark
brown; terga 5 and 6 with long dark-brown hairs, with long white
tufts laterally on tergum 5; sternal hairs reddish brown except
white apicolateral hairs on each sternum but the first and last.
Legs with white hairs except as follows: brown on outer surfaces
of fore tarsi, distally on fore and middle tibiae and basally on
hind tibiae, yellow to orange on middle and hind basitarsi and
hind tibiae; scopal hairs simple, unbranched, white to pale yellow.
Type material: Holotype female of baileyi from Fedor, Lee
County, Texas, April 5, 1904, Rev. G, Birkman coll., is the property
of the California Academy of Sciences, San Francisco, California,
but is temporarily deposited in the collection of the Citrus Experi-
ment Station, Riverside, California.
Melissodes (Apomelissodes) apicata Lovell and Cockerell
Melissodes apicata Lovell and Cockerell, 1906, Psyche, vol. 13, p. 111.
This species is very distinct from the other species in this sub-
genus. The female of apicata can be recognized by the protruding
clypeus, the weakly branched scopal hairs, the short galeae and
the abundant long hooked galeal hairs. The males can be dis-
tinguished by the short galeae which bear abundant short straight
hairs laterally, the peculiar pygidial plate described below and the
all or partially black clypeus. In addition, both sexes often have
three-segmented ‘maxillary palpi, a feature unique in the genus
Melissodes.
Female: Measurements and ratios: N, 20; length, 11.0-14.0 mm.;
width, 4.0-5.5 mm.; wing length, M == 21.56 + 0.252 mm.; hooks in
hamulus, M = 13.85 + 0.379; flagellar segment 1/segment 2, M=
1.98 + 0.075.
Structure and color: Integument black except as follows: mandi-
ble with apical half red with median golden macula; eyes gray to
552, Tse Universiry SCIENCE BULLETIN
bluish gray; flagellar segments 2 to 10 beneath slightly reddish;
distitarsi and often basitarsi, tibiae and femora and occasionally
sterna reddish brown; wing membranes somewhat infumate to
deeply so, brown, veins black; tegulae piceous; metasomal tergum 1
often narrowly hyaline apically.
Clypeus strongly protuberant, with coarse, irregular-sized punc-
tures separated mostly by half a puncture width or less, with irreg-
ular, longitudinal, median carina in apical half or less, ground areas
shiny, unshagreened; supraclypeal area coarsely punctate, ground
area shiny; vertex with flattened areas mesad of apices of compound
eyes coarsely punctate, punctures large, separated mostly by less
than half a puncture width, ground areas shiny, unshagreened,
galeae shiny, unshagreened, with abundant small punctures espe-
cially laterally, length less than three and more than two times
median clypeal length; maxillary palpal segments usually three in
ratio of about 4.0:4.0:2.5, occasionally a minute fourth segment
present. Mesoscutum with abundant round punctures separated by
half to one and a half puncture widths posteromedially and by half
a puncture width or less anteriorly and peripherally, ground areas
shiny, unshagreened; scutellum with punctures about same size as
on adjacent part of mesoscutum, separated by half a puncture
width or less, ground areas shiny, unshagreened; mesepisternum
with large shallow punctures mostly larger than those of postero-
medial area of mesoscutum and separated mostly by less than a
third of one puncture width, ground areas shiny to moderately so,
bottoms of punctures etched by reticulate shagreening; metanotum
with abundant crowded punctures medially, sparser laterally, punc-
tures separated by half a puncture width or less, smaller than those
in middle of scutellum, ground areas shiny medially, shagreened
laterally; propodeum with dorsal surface weakly reticulorugose
except in median sixth, ground areas opaque, tessellate, with pos-
terior surface sparsely punctate except broad, upper, impunctate
triangle, ground dulled by tessellation but impunctate triangle
usually moderately shiny. Metasomal tergum 1 with basal %
weakly punctate, punctures small, round, shallow, separated by
one to two puncture widths apically and by less than one width
basally and laterally, ground areas moderately shiny, with fine
transverse shagreening; tergum 2 with minute punctures in inter-
band and basal zones, punctures separated mostly by two puncture
widths but somewhat more crowded laterally, with apical area
impunctate, ground areas moderately shiny to dull, transversely
Metissopes iy Norta AND CENTRAL AMERICA 553
and densely shagreened; terga 3 and 4 similar to tergum 2 but
punctures more abundant especially in basal areas.
Hair: On head white except abundant long black or dark brown
hairs on vertex between apices of compound eyes and on face
almost to antennae; galeae with abundant long hooked hairs, es-
pecially laterally. Mesosoma with white hairs except large mes-
oscutellar patch of dark brown to black hairs which extends
forward beyond a transverse line at anterior margins of tegulae and
often almost reaches pronotum, posterior lobes of pronotum with
abundant long black hairs mixed with the pale, tegulae and humeral
angles of wings with dark brown hairs, scutellum with hairs dark
brown to black except long white hairs peripherally. Metasomal
tergum 1 with long white hairs in basal half to three fifths and to
apex at extreme sides; tergum 2 with basal and apical bands of
white pubescence connected at extreme sides, apical band with
sinuate anterior margin and almost interrupted medially, interband
zone with abundant black or dark brown, erect or suberect, bristle-
like hairs, apical band consists of several rows of short highly
plumose hairs; terga 3 and 4 with basal bands of dark brown to
black tomentum, with apical bands of white pubescence similar
to that of tergum 2 but not sinuate anteriorly and with abundant
dark brown bristlelike hairs in interband zones; terga 5 and 6 with
long dark brown to black appressed hairs and with lateral tufts
of long white hairs on tergum 5; sternal hairs brown except white
at extreme sides near apices of all but first and last sterna. Legs
with white hairs except as follows: outer surfaces of fore tarsi,
apices of fore and middle tibiae, bases of hind tibiae and basitibial
plates brown; inner surfaces of tarsi yellow (especially distitarsi )
to dark brown, inner surfaces of hind basitarsi usually reddish
brown; scopal hairs weakly plumose, each with one to five branches
on each side of rachis and usually three or four, more highly
plumose on basitarsi than on tibiae, pale ochraceous to white, oc-
‘asionally pale brown on tibiae.
Male: Measurements and ratios: N, 20; 10.0-13.0 mm.; width,
4.0-5.0 mm.; wing length, M = 21.35 =- 0.293 mm.; hooks in hamu-
lus, M = 12.65 -+ 0.215; flagellar segment 2/segment 1, M = 6.11
+ 0,118.
Structure and color; Integument black except as follows: clypeus
yellow or yellowish orange, with black apical margin and black
posterior margin, posterior infuscation often extends forward to
cover half or more of clypeus (one specimen entirely black); man-
dible with apical half red with median golden macula; flagellar
554 Tue Universiry SCIENCE BULLETIN
segments 2-11 yellow to red beneath, dark brown above; eyes dark
violet gray, gray, yellowish green, or bluish gray; distitarsi, often
basitarsi and tibiae, occasionally femora and coxae dark reddish
brown; wing membranes infumate, yellowish brown, veins dark
brown to black, tegulae piceous.
Structural characters as in female with the following differences
and addition: minimum length of first flagellar segment equals
approximately % of maximum length of second segment, often
less; metasomal tergum 1 punctate almost to apex medially; terga
2-5 with ground areas shiny to moderately so, weakly shagreened,
pygidial plate deeply notched at about a third distance from apex
to base, apical margin of apical portion and apicolateral margins
of basal portion upturned, basal portion usually on a slightly higher
level than apical portion so as to give the impression of two pygidial
plates.
xenital capsule essentially as in M. fimbriata but penis valve
with lateral process somewhat more acute (Figs. 3 and 4). Sternum
7 as in fimbriata but median plate larger, with more hairs ventrally
and without a distinct neck but joined to sternum by almost its
entire width, with lateral plates more rounded than in fimbriata
(Fig. 5). Sternum 8 similar to that of fimbriata but with fewer,
shorter hairs apicomedially and with numerous short hairs scattered
over ventral surface in apical half (Fig. 6).
Hair: As in female with the following differences: with few or
no brown hairs on vertex of head; galeal hairs abundant, especially
laterally, but straight, not hooked; mesoscutal and scutellar patches
of dark hairs usually somewhat smaller; metasomal tergum 1 with
pale hairs extending to, or almost to, the apex; tergum 2 with apical
white pubescent band not nearly interrupted medially, not sinuate
anteriorly; terga 2-5 with narrow apical white bands subequal to
each other in width and subequal in width across each tergum,
with dark brown bristlelike hairs more abundant and longer; terga
6 and 7 with long brown hairs; sternal hairs largely white to ochra-
ceous, darker basally on each sternum except the first and last,
apices of sterna with fringes of long hairs white and highly plumose
laterally and ochraceous to reddish brown medially; legs with white
hairs except yellow to brown on inner surfaces of tarsi, occasionally
brown on outer surfaces of hind basitarsi below basitibial plates
and occasionally ochraceous on outer surfaces of tarsi and tibiae
(especially hind tibiae).
Bionomics: This species is apparently oligolectic and restricted
to plants of the genus Pontederia (family Pontederiaceae) for both
Me.issopEs 1N NortH AND CENTRAL AMERICA 555
pollen and nectar. The hooked galeal hairs of the female are
probably an adaptation for removing pollen from the narrow tubu-
lar flowers of this plant. The longer galeae and protruding clypei
to house them are probably adaptations for retrieving nectar from
the deep nectaries of Pontederia.
Type material: Holotype female from Waldoboro, Maine, July
21, 1904, on Pontederia cordata, J. H. Lovell collector, is in the
collection of the North Carolina State College, Raleigh, North Caro-
lina.
Distribution: M. apicata occurs along the Atlantic coast from
Maine to Florida. It has been collected from March 21 to July 21,
but mainly in April and May. In addition to the holotype, 23
females and 45 males have been examined. The collection data for
these are listed below in full, since this is a rare and poorly known
species.
District or Corumpia: Washington, 2 ¢ ¢, July, 1901, J. C.
Bridwell. Fiorma: Alachua Co.,1¢@ and 54 @, April 12, 1955,
on Pontederia sp., 5 ¢ g, April 12, 1955, with no floral data, T. B.
Mitchell, April 13, 1955, 2 g on Melilotus alba and 2g 3 on
Stachys floridana, H. V. Weems Jr., April 12-18, 1955, 73 g on
Melilotus alba, R. A. Morse, April 15, 1955, 2 ¢ @, with no floral
data, R. A. Morse, April 12, 1955, 1 ¢, on Hydrocotyle umbellata,
R. A. Morse; Englewood, 1 ¢ , April 8, 1944, J. G. Needham; Fort
Ogden, 1 @ , April 9, 1952, G. S. Walley; Gainesville, Alachua Co.,
28 g, May 4, 1923, Alexander and Walker; Highlands Hammock
State Park, 5 g g, April 7, 1955, on Pontederia sp., T. B. Mitchell;
[sland Grove, 1 ¢, 1890; Jacksonville, 1 ¢, W. H. Ashmead col-
lection; Lake Iamonia, 1 ¢, May 12, 1916, on Pontederia cordata,
W. D. Pierce; Moon Lake, Pasco Co.; 1¢, April 16, 1952, J. R.
Vockeroth; Orlando, 1 9 , April 4, 1955, taken on sand, K. V. Krom-
bein; Osceola Co., 2 ¢ g, March 27, 1952, F. E. Lutz; Sarasota,
1g, April 2, 1938, F. E. Lutz; Sebring, 1 ¢ , March 25, 1938, F. E.
Lutz. Grorcia: Chauncey, 8¢@ 9? and 14, April 28, 1949, P. W.
Fattig. Maine: Waldoboro. Mary.anp: Mason Spring, 1 ay
July 12, 1931, on Pontederia cordata, P. H. Timberlake. Massacuu-
setrs: Concord, J. Bequaert. New Jersey: Atsion, 13,49 9,
July 9, 1955, on Pontederia sp., A. F. Shinn; Vineland, 1 g , Mourice
River, Norro Carotina: Wilmington, 8? 9 and 1¢, May 20,
1934, on Pontederia sp., T. B. Mitchell. An additional female
labeled “7-14-92” and “Ag. Coll. Mich.” has been identified as of this
species.
556 Tue Universiry ScrENCE BULLETIN
Melissodes ( Apomelissodes) mitchelli, sp. nov.
This species is named in honor of T. B. Mitchell who collected
five of the ten known specimens and who has contributed a great
deal to the knowledge of the species of Melissodes by means of the
specimens collected by him in the eastern states. M. mitchelli is
closely related to M. apicata from which it can be distinguished by
the characters appearing above in the keys. It is so much like
apicata in appearance that a detailed description would largely be
a repetition of the description of that species. The description
which follows, therefore, is largely an innumeration of the charac-
ters in which mitchelli and apicata differ.
Female: Measurements and ratios: N, 6; length, 11.0-12.0 mm.;
hamulus, M = 13.83 + 0.374; flagellar segment 1/segment 2, (N,
5) M=178 + 0.048.
Structure and color: Integumental color as in apicata. Structure
as in apicata with the following differences: clypeus with postero-
median area flattened or slightly depressed, with punctures small,
round, and sparse medially almost to apical margin, ground shiny,
unshagreened, punctures round, regular in size and shape laterally;
galeae shiny, with sparse punctures bearing short straight hairs
laterally; maxillary palpal segments four, in ratio of about 2.5:4:3:1;
propodeum with dorsal face with broader impunctate zone me-
dially; metasomal tergum 1 with basal % or less of dorsal surface
punctate; terga 1-3 with ground areas dulled by fine, dense, trans-
verse shagreening, duller than in apicata; pygidial plate % as
broad at extreme base than median length, distinctly broader than
in apicata.
Hair: Color of hairs as in apicata with the following differences:
vertex with black hairs less abundant; lobes of pronotum without
dark hairs; dark mesoscutellar patch not almost reaching pronotum
but extending forward to a transverse line at or beyond anterior
margins of tegulae; tegulae and humeral angles of wings with pale
hairs; metasomal tergum 2 with apical band separated from apex of
tergum in median third or less and often narrowly interrupted
medially; scopal hairs with one to three branches on either side of
each rachis, never brown on tibiae except immediately below basi-
tibial plates.
Male: Measurements and ratios: N, 4; length, 10.0-12.0 mm.;
width, 3.5-4.0 mm.; wing length, M = 20.90 + 0.643 mm.; hooks in
————————
|
|
|
|
|
MenissopEs iN NortH AND CENTRAL AMERICA 557
hamulus, M = 12.25 + 0.479; flagellar segment 2/segment 1, M =
5.80 + 0.174.
Structure and color: Integumental color as in apicata with the
following differences: clypeus entirely yellow except black notches
posteriorly at tentorial pits; labrum with small median yellow spot
(holotype) or entirely black; mandible with (holotype) or without
small basal pale spot. Structure as in apicata except as follows:
clypeus with coarse round punctures separated from each other by
half to one puncture width laterally and with a narrow longitudinal
impunctate median area, ground areas shiny, unshagreened; mini-
mum length of first flagellar segment equals 4 to % of maximum
length of second segment; maxillary palpi as in female; galeae as in
female; metasomal terga with ground areas dulled by fine dense
transverse shagreening; pygidial plate broader than in apicata and
less deeply notched laterally.
Genital capsule essentially as in M. fimbriata and M. apicata;
penis valve with lateral process short and blunt as in fimbriata.
Sternum 7 as in apicata but with median plates more distinctly
necked, with more abundant hairs ventrally on median plates, with
membranous area between plates much smaller and with apodemes
longer and narrower (Fig. 7). Sternum 8 as in apicata but with
apical margin truncate or slightly concave.
Hair: As in apicata except as follows: tegulae and humeral
angles of wings with pale hairs; posterior lobes of pronotum without
dark hairs; metasomal tergum 2 with apical white band narrowly
interrupted medially; terga 2 and 3 with apical bands not reaching
apical margins of terga in median thirds or less; inner surfaces of
basitarsi and tibiae with yellow hairs.
Type material: Holotype male and allotype female from Levy
County, Florida, April 18, 1955, on Opuntia sp., T. B. Mitchell, is
in the collection of the North Carolina State College, Raleigh, North
Carolina. Four female and three male paratypes are as follows:
Frorma: Alachua Co., 1 9, April 27, 1955, on Opuntia sp., R. A.
Morse; Lake Co.,.1 ¢ , April 20, 1922, T. P. Winter; Gainesville, 1 9 ,
May 1, 1955, on Opuntia sp., R. A. Morse; Orange City, 1 9 , April
22, 1928, C. O. Bare. Grorcia: Wadley, 1 ¢ , April 23, 1938, C. H.
Curran. Norra Caroira: Southern Pines, 29 9, May 20, 1953,
on Opuntia sp., T. B. Mitchell; Wayne Co., 1 ¢ , April 7, 1954, T. B.
Mitchell. The paratypes will be deposited in the collections of the
North Carolina State College, Raleigh, the Snow Entomological
Museum, Lawrence, Kansas, the American Museum of Natural His-
558 Tue Universiry SCIENCE BULLETIN
tory, New York City, the University of Michigan, Ann Arbor, the
State Plant Board of Florida, Gainesville, and in the author's col-
lection.
Subgenus Psilomelissodes LaBerge
Psilomelissodes LaBerge, 1956, Univ. Kansas Sci. Bull., vol. 87, p. 1178.
Type species. Melissodes intorta Cresson, 1872, monotypical and
by original designation.
This subgenus was erected to include a single species which is
distinctive because of its sparse pubescence, the weak scopal hairs
in the female, the short male antennae and the black male clypeus.
A complete subgeneric description is given by LaBerge (1956).
Melissodes (Psilomelissodes) intorta Cresson
Melissodes intorta Cresson, 1872, Trans. Amer. Ent. Soc., vol. 4, p. 278;
Birkmann, 1899, Ent. News, vol. 10, p. 245; Cockerell, 1902, Entomologist,
vol. 85, p. 177; 1906, Trans. Amer. Ent. Soc., vol. 82, p. 75; Cresson,
1916, Mem. Amer. Ent. Soc., vol. 1; p, 121,
Melissodes wickhami Cockerell, 1906, cae Amer. Ent. Soc., vol. 82, p. 112
(new synonymy).
Female: Measurements and ratios: N, 7; length, 11.0-12.0 mm.;
width, 4.0-4.5 mm.; wing length, M = 24.10 + 0.389; hooks in ham-
ulus, M = 18.43 + 0.298; flagellar segment 1/segment 2, M = 2.14
+ 0.082.
Structure and color: Integument black except as follows: mandi-
ble with apical half red or covered by a large median golden-yel-
low macula (in unworn condition); flagellar segments 2-10 dark
reddish brown to brown below; eyes gray; tarsi and often remainder
of legs brown; tegulae piceous, but often clear posteromedially;
wing membranes infumate, brownish, veins brown to black; terga
1-4 with narrow colorless apical margins.
Clypeus with round, regular, small punctures separated mostly
by less than one puncture width, with a short median carina near
apex, ground areas shiny, unshagreened; supraclypeal area with
deep round punctures, ground shiny; galeae sparsely punctate
above, ground shiny, unshagreened; maxillary palpi 4-segmented,
in ratio of about 6:5:3:1; flattened areas of vertex extending mesad
from apices of compound eyes with small round punctures sepa-
rated by one puncture width or less, ground shiny, unshagreened.
Mesoscutum with deep round punctures relatively evenly spaced
but slightly sparser posteromedially than anteriorly and laterally
and becoming distinctly larger and more crowded in posterior
declivity, ground areas somewhat dulled by exceedingly fine tes-
sellation posteromedially, shiny elsewhere; scutellum with abundant
Meuissopes IN NorrH AND CENTRAL AMERICA 559
crowded ‘punctures similar to posterior declivity of mesoscutum,
ground shiny; mesepisterna with large, round, deep punctures
separated by half a puncture width or less, many confluent with
neighbors, ground shiny; metanotum somewhat elevated medially,
punctate in median third, impunctate or indistinctly punctate in
lateral thirds, lateral areas dulled by fine tessellation; propodeum
with dorsal surface irregularly rugose laterally, surface dulled by
tessellation, with posterior surface sparsely punctate laterally,
punctures becoming more abundant at extreme lateral margins,
ground areas dulled by coarse tessellation, with lateral surfaces
coarsely and densely punctate and tessellate. Metasomal tergum
1 with sparse, shallow, indistinct punctures scattered over basal %
of dorsal surface, punctures separated by one puncture width at
extreme base and extreme sides and by two puncture widths or
more elsewhere, ground areas opaque, dulled by dense, coarse,
reticulate shagreening; tergum 2 with interband zone irregularly
and sparsely punctate, punctures of irregular size and separated
by one to several puncture widths from nearest neighbors in any
direction, with dense coarse punctures beneath distal pubescent
band, apical area impunctate, ground areas opaque, dulled as in
tergum 1; terga 3 and 4 similar to tergum 2 but interband zones
with more abundant punctures; sterna coarsely punctate, ground
areas dulled by fine tessellation.
Hair: On head white except abundant brown hairs on vertex
between apices of compound eyes and ochraceous hairs on labrum
and mandibles. Thorax with white hairs except large brown pos-
teromedian mesoscutellar patch which extends forwards beyond
a transverse line at anterior margins of tegulae and scutellar hairs
brown except marginal white hairs. Metasomal tergum 1 with
sparse white hairs basally and laterally on dorsal surface; tergum 2
with sparse basal band of white pubescence, narrow distal band
of white pubescence which becomes thinner and is interrupted
medially, basal and distal bands connected at extreme sides by
white pubescence, apical and interband areas with sparse, white,
short pubescence; tergum 83 similar to tergum 2 but with apical
and interband areas with more abundant pale pubescence and
with short, erect or suberect, spinelike, pale brown hairs present
in interband zone; tergum 4 similar to tergum 8 but pale pubescence
and spinelike hairs of interband zone more abundant and_ the
latter longer and with distal pale band reaching apex of tergum
although often with a bare median patch at apex; terga 5 and 6
with long, dark brown, appressed hairs. Sternal hairs dark brown.
Tue Universiry ScieNCE BULLETIN
560
Legs with hairs as follows: coxae, trochanters and femora white
except brown at tips of femora; fore and middle tibiae with outer
surfaces white basally and pale brown apically; outer surfaces of
middle basitarsi pale brown; inner surfaces of fore tibiae and
middle basitarsi, fore tarsi, middle and hind distitarsi brown; inner
surfaces of hind basitarsi and tibiae reddish yellow to brownish
red; scopal hairs ochraceous medially, brown surrounding and
below basitibial plates and pale brown on apical halves of basi-
tarsi. Scopal hairs weak, sparse, mostly with two or three long
weak usually curved branches on either side of rachises and often
with fewer branches, surface of tibiae is clearly visible through
the weak hairs.
Male: Measurements and ratios: N, 20; length, 11.0-13.0 mm.;
width, 3.0-4.0mm.; wing length, M = 24.18 -+ 0.839 mm.; hooks in
hamulus, M = 12.65 0.335; flagellar segment 2/segment 1, M =
1.12 + 0,001.
Structure and color: Color as in female except as follows: fla-
gellar segments 2-11 yellow to red below; tergum 1 more broadly
hyaline apically; tergum 5 hyaline apically; hyaline marginal zones
of terga 1-5 usually yellow or translucent brown rather than color-
less. Structure and sculpturing as in female with the following
additions: minimum length of first flagellar segment equals maxi-
mum length of second segment or slightly less, usually about %
of second segment (Fig. 13); tergum 1 with basal % or more
punctate, punctures somewhat more abundant than in female,
especially laterally and basally; interband zones of terga 2 and 3
more densely punctate; terga 4 and 5 similar to tergum 3; sternum
6 with prominent short oblique carinae near apex, broadly convex
medially, truncate, with sparse punctures posterior to apical carinae,
ground shiny, with sparse transverse shagreening.
Terminalia described and figured by LaBerge (1956, Figs. 122-
125),
Hair: As in female with the following differences: vertex of
head often without brown hairs; tergum 1 with long white hairs
more abundant; tergum 2 with distal pale band often complete;
interband zones of terga 2-4 with more abundant pale pubescence
and more abundant spinelike hairs; terga 4 and 5 with distal pale
bands as in tergum 3 but with sparse brown tomentum at extreme
bases; terga 6 and 7 with dark brown hairs; sternal hairs pale at
least lateroapically; legs with white hairs except red to yellow on
inner surfaces of tarsi.
ME ttssopes in NortH AND CENTRAL AMERICA 561
Bionomics: It is probable that this is an oligolectic bee dependent
upon malvaceous plants of the genus Callirhoe for its pollen.
However, only eight females are known and among these only
two bear flower labels. These two specimens were collected by
R. H. Beamer visiting flowers of Gaillardia. One of these has
Callirhoe pollen in its scopae, whereas the other bears pollen from
some other plant (it is not certain whether this is Gaillardia pollen
or not). The males have been collected visiting Callirhoe flowers
in several localities in Texas and Kansas.
Type material: The lectotype male of intorta is in the Academy
of Natural Sciences, Philadelphia, Pennsylvania. The holotype
female of wickhami is in the University of Colorado Natural History
Museum, Boulder, Colorado.
Distribution: This species ranges from eastern Texas north to
eastern Kansas. It has been collected from April 14 to June 27.
In addition to the type material, 22 males and 7 females have
been examined from the localities listed below.
Kansas: Baxter Springs (5 miles E. and 5 miles S.); Brookville;
Gove Co.; Manhattan; Riley Co.; Scott City (12 miles N.). Texas:
Childress; Fedor, Lee Co.; Gillespie Co.; Giddings; Goliad (16
miles E.); Lee Co.; Stonewall; Sweet Home; Weser, Goliad Co.
Remarks: A single specimen labeled “Camd. Co., N. J., Collection
W. J. Fox” is evidently mislabeled. I have found several speci-
mens of other species of Melissodes of the W. J. Fox collection,
ostensibly from Camden Co., New Jersey, which belong to western
species and are obviously mislabeled.
Flower records: Asclepias latifolia, Callirhoe sp., C. digitata,
C. leiocarpa, Gaillardia sp., Rorippa sinuata.
Subgenus Heliomelissodes LaBerge
Heliomelissodes LaBerge, 1956, Univ. Kansas Sci. Bull, vol. $7, p, 1178.
Type species. Melissodes desponsa Smith, 1854, by original
designation.
This subgenus consists of two species widely distributed in North
America, The subgenus can be distinguished by the protruding
clypei and long galeae in both sexes and by the male antennae
being of moderate length. A detailed description is given by La-
Berge (1956). The subgenus is restricted to plants of the genus
Cirsium (Compositae ) or related genera for pollen sources.
19—3378
Tue University ScreNcE BULLETIN
Key TO THE SPECIES OF THE SUBGENUS HELIOMELISSODES
Males
1. Metasoma with black pubescence and hairs on terga 2 to 7; meso-
scutum with few or no black hairs posteromedially; legs with black
hairs on outer surfaces of tibiae and tarsi.............. desponsa
Metasoma usually with some pale ochraceous to white pubecent bands
or fasciae medially at least on tergum 2, often on terga 3 and 4 and
occasionally on tergum 5 as well; legs usually with pale hairs on
outer surfaces of tibia and often pale on tarsi as well ...... rivalis
Females
1. Metasoma without pale pubescent bands or fasciae on terga 2 to 4;
mesoscutum without posteromedian patch of dark hairs or this patch
smaller than dark scutellar patch; mesoscutum with punctures pos-
teromedially mostly separated by one puncture width or less ... .
ey lea a a ee eee er er rrr desponsa
Metasoma usually with pale pubescent bands or fasciae at least on
tergum 2 and often on terga 3 and 4 as well, if without pale pu-
bescence, then mesoscutum with posteromedian patch of dark hairs
larger than dark scutellar patch; mesoscutum usually with a postero-
median area of sparse punctures separated mostly by two puncture
WIKENGIOY MOI ae ee a oe ee ... .riwalis
Melissodes (Heliomelissodes) desponsa Smith
Melissodes desponsa Smith, 1854, Cat. Hymen, British Museum, Part IT. Apidae,
p. 310; Cresson, 1879, Trans. Amer. Ent. Soc., vol. 7, p. 925; Provancher,
1882, Le Naturaliste Canadien, vol. 18, p. 174; Robertson, 1894, Trans.
Acad. Sci. St. Louis, vol. 6, pp. 455, 459, 470, 473, 475; 1897, Trans. Acad.
Sci. St. Louis, vol. 7, p. 854; 1901, Canad. Ent., vol. 33, p. 230; Viereck,
et al., 1905, Canad. Ent., vol. 37, p. 820; Lovell and Cockerell, 1906,
Psyche, vol. 13, p. 109; Cockerell, 1906, Trans. Amer. Ent. Soc., vol. 32,
pp. 76, 83; Smith, 1910, Ann. Rept. New Jersey State Museum, 1909, p. 693;
Viereck, 1916, Bull. Connecticut State Geol. and N. H. Surv., vol. 22, p.
732; Criddle, Curran, Viereck and Buckell, 1924, Rept. Ent. Soc. Ontario,
vol. 33, p. 99; Leonard, 1926, Cornell Univ. Agric. Exp. Sta. Mem., No. 101,
p. 1028; Graenicher, 1935, Ann. Ent. Soc. Amer., vol. 28, p. 804.
Pinkie id daponsa (sic) Bridwell, 1899, Trans. Kansas Acad. Sci., vol. 16,
2)
Melissodes nigripes Smith, 1854, Cat. Hymen. British Museum, Part J. Apidae,
p. 811 (a, nec 9); Cresson, 1879, Trans. Amer. Ent. Soc., VO Ti-pr 226:
Provancher, 1888, Addit. Faun. Can., Hymen., vol. 2, p. 300; Smith, 1896,
Rept. Ent. Dept. New Jersey Agric. Coll. Exp. Stat., fig. 21; Robertson,
1900, Trans. Acad. Sci. St. Louis, vol. 10, p. 53; 1901, Canad. Ent., vol. 33,
p. 230; Cockerell, 1906, Trans. Amer. Ent. Soc., vol. 82, p. 102; Smith, 1910,
Ann. Rept. New Jersey State Museum, 1909, p. 693.
Melissodes cnici Robertson, 1901, Canad. Ent., vol. 33, p. 230; Cresson, 1879,
Trans. Amer. Ent. Soc., vol. 7, p. 225; Robertson, 1905, Trans. Amer, Ent.
Soc., vol. 31, p. 369; Cockerell, 1905, Canad. Ent., vol. 87, p. 812; 1906,
Trans. Amer. Ent. Soc., vol. 32, pp. 77, 83, 89; Graenicher, 1911, Bull. Publ.
Mus. Milwaukee, vol. 1, p. 247; Gileson, 1914, Forty-fourth Ann. Rept. Ent.
Soc. Ontario, 1913, p. 125; Robertson, 1914, Ent. News, vol. 25, p. 69;
1926, Ecology, vol. 7, p. 379; 1928, Flowers and Insects, p. 8; Pearson,
1933, Ecol. Monogr., vol. 3, p. 381.
The status of the various names which have been applied to this
bee has been confusing. Cresson in 1879 first synoymized M.
MeE.issopEs in NortH AND CENTRAL AMERICA 563
nigripes Smith with M. desponsa Smith. He was followed in this
by Dalla Torre (1895). Robertson in 1901 named this species M.
cnici, Lutz and Cockerell (1920) recognized that M. cnici Robert-
son was a new name for the male (but not the female) of M. nigripes
Smith and stated that M. cnici might include the female of desponsa.
Dr. I. H. H. Yarrow of the British Museum (Natural History)
has kindly examined specimens of M. cnici sent to him and assures
me in a letter dated August 4, 1954, that the female type of M.
desponsa Smith and the male cotype of M. nigripes Smith are both
conspecific with the specimens of M. cnici. I, therefore, concur
with the opinion expressed by Lutz and Cockerell (1920) that the
names nigripes and cnici should be regarded as synonyms of de-
sponsa. The female cotype of M. nigripes Smith is, unfortunately,
a Tetralonia, according to Dr. Yarrow. In addition to this male and
female of nigripes in the British Museum, Dr. Yarrow has brought
to light two males labeled nigripes Sm. by Smith himself. One of
these is from Mt. Pleasant (R. Forster) and is undoubtedly cnici
Robertson, but the second which is from the United States has a
very swollen clypeus and is probably an Emphor. Dr. Yarrow (in
a letter dated November 23, 1955) agrees that the male of nigripes
which was in the British Museum before Dr. Yarrow found the
additional two males should be designated as the lectotype. This,
in effect, retains the name nigripes Smith in the genus Melissodes
as a synonym of M. desponsa Smith.
This species is distinctive because of the black metasoma and
the pale upper surface of the mesosoma. It converges in this
coloration with a race of the closely related M. rivalis Cresson oc-
curring in the Pacific Northwest. However, it can be distinguished
from these darker specimens of rivalis by the lack of a patch of dark
hairs posteromedially on the mesoscutum, or this patch is small and
no larger than that on the scutellum. Minor differences in puncta-
tion and structure also separate the two species and these are
described below. '
Female: Measurements and ratios: N, 20; length, 11.5-16.0 mm.;
width, 4.0-5.5 mm.; wing length, M = 25.21 ++ 0.157 mm.; hooks in
hamulus, M = 14.00 + 0.229; flagellar segment 1/segment 2, M =
2.08 + 0.020,
Structure and color: Integument black except as follows: fla-
gellar segments 2-10 dark reddish brown below; apical half of
mandible red with broad golden macula apically; apical margin
of clypeus often reddish; eyes grayish blue to green; distitarsi dark
564 Tar Universrry SCIENCE BULLETIN
red; wing membranes infumate, brownish, tegulae piceous, tibial
spurs piceous.
Clypeus with small crowded punctures separated by less than
one puncture width, without a distinct median carina, ground
areas dulled by coarse transverse shagreening, moderately shiny;
supraclypeal area with coarse round punctures, ground areas
dulled by coarse, reticular shagreening; vertex with flattened area
extending posteromesad of apex of compound eye with small, deep,
round punctures separated by less than half to two puncture widths,
ground areas shiny to moderately so, with fine shagreening; maxil-
lary palpal segments in ratio of about 3.5:2.5:2.5:1, the last segment
often much shorter; galea shiny above with small sparse scattered
punctures bearing short straight hairs, shagreened near tip and
laterally. Mesonotum coarsely punctate, punctures separated by
less than half a puncture width anteriorly and peripherally, by
slightly more posteromedially just before posterior declivity, slightly
larger posteromedially than anteriorly, ground areas slightly dulled
by fine reticular shagreening; scutellum coarsely punctate, similar
to mesoscutal sculpturing but shagreening finer or absent; meta-
notum with scattered punctures, ground dulled by tessellation;
propodeum with dorsal surface reticulorugose basally, coarsely
punctate apically, with posterior surface coarsely punctate except
on upper impunctate median triangle, with lateral surfaces coarsely
punctate, with ground areas opaque, dulled by coarse, dense tes-
sellation; mesepisterna with lateral surface with large shallow
punctures separated by less than half a puncture width, some con-
fluent medially, becoming smaller and more distinct dorsally and
posteriorly, ground areas somewhat dulled by irregular shagreening.
Metasomal tergum 1 with a line of close-set punctures just beyond
middle, these separated by a slight gap in median % of tergum
from basal area with abundant small round shallow punctures,
ground areas dulled; tergum 2 with minute, round deep punctures
at extreme base, with interband zone relatively impunctate medially
and with small crowded punctures laterally, apical zone with
minute, sparse punctures laterobasally, ground areas dulled; terga
3 and 4 similar to tergum 2 but with punctures smaller and more
abundant in interband and apical zones.
Hair: Generally shorter and more abundant on mesosoma,
shorter on prosoma and less abundant on metasoma than in M.
mysops. Head with black to dark brown hairs on clypeus, genal
areas, mandibles and often an occiput behind vertex, with hairs
i
Me.issopes IN NortH AND CENTRAL AMERICA 565
of face and vertex all pale ochraceous or ochraceous and brown
mixed, Metasoma with pale ochraceous hairs except as follows:
mesoscutum with posteromedian patch of dark brown to black
hairs often present but never extending forward to a transverse
line at middle of tegulae and scarsely, if at all, any larger than that
on scutellum; scutellum with dark brown to black hairs medially;
lateral surfaces of mesepisterna with dark hairs on lower half or
more; metepisterna and lateral surfaces of propodeum with dark
brown to black hairs. Mesosoma with dark brown to black hairs
except as follows: white to pale ochraceous on basal %% of dorsal
surface of tergum 1 (often mixed with dark hairs), tergum 2 with
basal band often consisting of at least partially pale pubescence.
Legs with dark brown to black hairs except scopal hairs which are
yellow to ochraceous and hairs of inner surfaces of hind tibiae
which are red to yellow.
Male. Measurements and ratios: N, 20; length, 9.0-14.0 mm.;
width, 2.5-4.5 mm.; wing length, M = 23.79 -+- 0.280 mm.; hooks in
hamulus, M = 18.00 + 0.218; flagellar segment 2/segment 1, M =
2,98 == 0.059.
Structure and color: Integument black except as follows: clypeus
pale yellow except reddish brown apical margin and black notches
marking tentorial pits, in specimens from eastern parts of the
range often infuscated in posterior angles from tentorial pits to
posterior margin in middle of clypeus; flagellar segments 2-11
yellow to red below; eyes green to gray; distitarsi, sterna and often
apices of terga dark reddish brown; wing membranes somewhat
infumate, brownish; tegulae and tibial spurs piceous.
Minimum length of first flagellar segment at least as long as one
third of maximum length of second segment and usually longer.
Maxillary galeae, palpae and sculpturing of head and body as in
female with the following differences: clypeal punctures smaller
and denser; lateral areas of vertex often with ground dulled by
shagreening; first metasomal tergum punctate in basal % of dorsal
surface medially and to the apex in lateral thirds, punctures small
and shallow; tergum 2 with interband zone with punctures small,
indistinct but scarsely any sparser than interband zone of terga
3 and 4; last exposed sternum with truncate apical margin, with
strong lateral oblique carinae near apex, with short median de-
pression in apical half between and posterior to apices of oblique
carinae, shiny and unshagreened but with small deep punctures
basally.
566 Tue Universiry SCIENCE BULLETIN
Terminalia as described and figured by LaBerge (1956, Figs.
114-117). Spatha of genital capsule feebly emarginate medially on
apical margin. Sternum 7 with few short hairs apically, narrower
in middle than at anterior third. Sternum 8 with apodemes trun-
cate and slightly capitate.
Hair: Color generally as in the female but usually with more
abundant pale hairs on head and sides of thorax and hairs of hind
legs all dark brown to black except on inner surfaces of hind tarsi.
Bionomics: Nothing has been reported concerning the nesting
sites or nest architecture of M. desponsa. This bee is oligolectic
and prefers flowers of the genus Cirsium as pollen sources, although
it has been collected occasionally on other genera of Composites
and also on flowers of other families. A total of 263 of the specimens
studied by the author bear flower labels. The data from these
specimens are summarized in Table 1. Additional flower records
available in the literature are not included in this table, since no
quantitative data were available. The table clearly demonstrates
the preference of desponsa for flowers of the genus Cirsium.
TABLE 1
Plant data M. desponsa data
Family Number | Number || Number | Number |Number | Total
of of of of of number
genera spp. colls, We oo | of bees
Compositae, other
than Cirsium.... if 9 ak 6 8 14
Compositae,
Oireniamneapp ies ve: 1 8 64. 126 118 244
LIBDINGAG eg ee i} 1 1 | 0 1
Leguminosae....... 1 sl L 0 J 1
Verbenaceae........ 1 2 2 0 3 3
‘TOUIa: (4,0 1 21. 79 133 130 263
Type material: Female holotype of desponsa from Ohio is in the
British Museum (Natural History) (17B855). Male lectotype of
nigripes, here designated, from the United States is in the British
Museum (Natural History) (17B836). Lectotype male and lecto-
allotype female of cnici, here designated, from Carlinville, Illinois,
eee amen aioe
MeE.issopEs IN NorTH AND CENTRAL AMERICA 567
August 18, 1897, on Cnicus lanceolatus, Charles Robertson collec-
tor, are in the collection of the Illinois Natural History Survey,
Urbana, Illinois.
Distribution: This species ranges from Nova Scotia in the north-
east, to North Carolina in the southeast, to Alabama in the south
and to Oklahoma and eastern North Dakota in the west. One
specimen labeled Laramie. Wyoming, was seen by the author.
This label is of doubtful validity. It has been collected from June
26 to October 16, but mainly in August. In addition to the type
material, 474 females and 344 males have been examined from the
localities listed below. This list includes localities reported in the
literature.
AtaBAMA: Kushla. Connecricur: Canaan City; Chappinville;
Colebrook; Greenwich. Ittmors: Attwood; Bath; Beverly Hills;
Bluffs; Carlinville; Champagne; Charleston; Chicago; Downers
Grove; Kankakee; Matteson; McHenry; Olive Branch; Pittsfield;
Urbana; Willow Springs. Inp1ana: Allen Co.; Lafayette; Warren
Co.; White Co. Iowa: Ames; Dickinson Co.; Iowa City; Lake
Okoboji; Ledges State Park; Mt. Pleasant; Nevada; Sioux City.
Kansas: Baldwin; Delavan, Morris Co.; De Soto, Johnson Co.;
Douglas Co.; Iola; Kansas City; Lawrence; Manhattan; McPherson
Co.; Norton, Riley Co.; Olathe; Riley Co.; St. George; Sunflower;
Topeka; Wildcat. Mate: Waldoboro; Winthrop. MARYLAND:
Bethesda; Cabin John; Glen Echo; Millington. MaAssacHusETTs:
Holden; Lexington; Lowell, Montossy; Needham; Wollaston;
Woods Hole. Micrican: Alger Co.; Ann Arbor; Cheboygan; East
Lansing; Grand Rapids; Ingham Co.; Midland; Millington; Otsego
Co.; Pontiac; Tuscola Co. Minnesota: Alex; Dakota Co.; Eitzen;
Erskine; Excelsior; Faribault Co.; Frazer; Freeborn Co.; Grand
Meadow; Hastings; Hayward; Hennepin Co.; Jordan; Lake Park;
Olmsted Co.; Ramsay Co.; Rochester; Rock Co.; Spring Valley;
Stanton; St. Anthony Park; St. Cloud; St. Paul; Warren; Washing-
ton Co.; Zumbra Heights, Carver Co. Mussourt: Columbia; Han-
nibal; Sedalia; Smithton; St. Louis. Nesrasxa: Fullerton; Gordon,
Sheridan Co.; Lincoln; Long Pine; Malcolm; Nebraska City; Neligh;
Omaha; Roca; West Point. New Hampsurre: Alstead; Durham;
Pelham. New Jersey: Arlington; Englewood; Kearny; New Bruns-
wick; Princeton. New York: Brooklyn; Calicoon; Fishers Island;
Flatbush; Fort Wadsworth; Huguenot; Ithaca; Long Island; New
Baltimore; Staten Island; Stockport; Van Cortland Park, New York
City; Westville. Norra Carotima: Black Mts. (valley of); Bryson
568 Tur Universiry SCIENCE BULLETIN
City; Doughton Park. Norra Dakota: Fargo; Henkinson; Oakes.
Ono: Columbia; Ironton; Logan Co. Oxtanoma: Umatilla.
PENNSYLVANIA: Carlisle; Philadelphia; Pike Co. Ruopr IsLanp:
Scituate. Sourm Daxora: Brookings; Vermillion; Yankton. VER-
mont: Chittenden (Rutland); Hartland; Woodstock. Vrrcinta:
Alexander Island; Falls Church; Vienna (2 and 3 miles W.). West
Vircinta: Canaan Valley. Wisconsin: Farmington; Hudson; Mad-
ison; Maiden Rock, Pierce Co.; Milwaukee; Nevers Dam, Polk Co.;
Prescott; Randall, Burnett Co.; Shullsburg, Lafayette Co.; Wash-
ington Co.; Worden Township, Clark Co. Wyomrinec: Laramie
(questionable). New Brunswick: Grand Manan Island. Nova
Scotia: Weymouth. Ontario: Carp (5 miles W.); Marmora,
Ottawa; Toronto; West Hill. Quesec: Cap Rouge; Fort Coulogne;
Lowe; Montreal; Quebec City; Ste. Annes; St. Fargeau.
Flower records: Aster novaeangliae, Brauneria purpurea, Car-
duus crispus, Cassia sp., Centaurea sp., C. jacea, Cirsium sp., C.
altissimum, C. arvense, C. discolor, C. hilii, C. odoratus, C. pumilum,
C. undulatum, C. vulgare, Coreopsis aristosa, Helianthus annuus,
H. grosseserratus, Inula helenium, Monarda fistulosa, Pontederia
cordata, Pycnanethemum sp., Rudbeckia laciniata, Silphium perfo-
liatum, Solidago sp., 8. canadensis, S. puberula, Verbena sp., V.
stricta.
Melissodes (Heliomelissodes) rivalis Cresson
Melissodes rivalis Cresson, 1872, Trans. Amer. Ent. Soc., vol. 4, p. 277; Cock-
erell, 1906, Trans. Amer. Ent. Soc., vol. 32, p. 76; Cresson, 1916, Mem.
Amer. Ent. Soc., vol. 1, p. 129.
Melissodes desponsiformis Cockerell, 1905, in Viereck et al., Canadian Ent.,
vol. 87, p. 820 (new synonymy); 1906, Trans. Amer. Ent. Soc., vol. 32,
p..89.
Melissodes mysops Cockerell, 1905, Entomologist, vol. 38, p. 146 (new synon-
ymy );1905, Canadian Ent., vol. 37, p. 321; 1906, Trans. Amer. Ent. Soc.,
vol, 82, pp. 88, 89, 113; 1907, Entomologist, vol. 40, p. 269; 1910, Ann.
Mag. Nat. Hist., ser. 8, vol. 6, p. 128; 1910, Univ. Colorado Studies, vol. 7,
p. 128; Criddle, Curran, Viereck and Buckell, 1924, Fifty-fourth Ann. Rept.
Ent. Soc. Ontario, p. 99; Scullen, 1928, Pan-Pac. Ent., vol. 4, p. 176; Bohart,
Knowlton and Bailey, 1950, Utah State Agric. Coll., mimeo. series no. 871,
p. 5.
Melissodes hexacantha Cockerell, 1905, Psyche, vol. 12, p. 100 (new synon-
ymy); 1906, Trans. Amer. Ent. Soc., vol. 32, p. 75; Snow, 1906, Kansas
‘Acad. Sci., vol. 20, p. 187; Bohart, Knowlton and Bailey, 1950, Utah State
Agric. Coll., mimeo. series no, 871, p. 5.
Melissodes nigrosignata Cockerell, 1905, Psyche, vol. 12, p. 101 (new synon-
ymy); 1906, Trans. Amer. Ent. Soc., vol. 32, pp. 88, 109, 309; Snow, 1906,
Trans. Kansas Acad. Sci., vol. 20, p. 137.
Melissodes habilis Cockerell, 1925, Ann. Mag. Nat. Hist., series 9, Vol 16,
229 (new synonymy); 1928, Univ. Colorado Studies, vol. 16, p. 114; 1937,
Amer. Mus. Nov., no. 899, p. 5.
This is a highly variable species. No subspecies will be recognized
ME issopEs In NortH AND CENTRAL AMERICA 569
in this paper for reasons expounded below. The males of rivalis
are distinguished by the pale pubescent bands on terga 2 and 3
and often on 4 and 5 as well, and by the posterior edge of the
clypeus usually being darkened. The females of rivalis can usually
be recognized by having pale pubescent bands on terga 3, 2 or 4,
in that order, or at least having lateral fasciae of white pubescence
on one of these terga, or, if without pale pubescence on any meta-
somal tergum, by having a large patch of black hairs on the meso-
scutum which exceeds the size of the dark scutellar patch.
Female: Measurements and ratios: N, 20; length, 12.0-17.0 mm.;
width, 4.5-6.0 mm.; wing length, M = 25.14 + 0.339 mm.; hooks in
hamulus, M = 15.25 + 0.876; flagellar segment 1/segment 2, M =
2.03 = 0.036.
Structure and color: As in M. desponsa with the following differ-
ences: eyes usually grayish blue; wings infumate brown; apical
margin of clypeus black; clypeus with irregular crowded punctures,
with a distinct median longitudinal carina in apical half or less;
mesoscutum usually with posteromedian area with sparse punctures
separated mostly by two puncture widths or more.
Hair: Darkest specimens as in dark specimens of M. desponsa
with the following differences: mesonotum with large patch of dark
brown to black hairs which extends forward to or beyond a trans-
verse line at anterior margins of tegulae; mesepisterna with dark
brown hairs at least on lower two thirds; posterior pronotal lobes
and tegulae with dark hairs; metasomal tergum 1 with long dark
brown hairs basally; inner surfaces of hind basitarsi and tibiae with
dark red to black hairs. Palest specimens with head hairs white
or grayish white except some brown hairs on vertex, with mesepi-
sternal and pronotal hairs white or pale ochraceous, with dark meso-
notal patch extending forward to a transverse line at least at middle
of tegulae; metasomal tergum 1 with long pale hairs basally, tergum
2 with basal band of white pubescence and distal band of pale
ochraceous pubescence (the latter narrowed and often interrupted
medially, separated from apical margin of tergum), tergum 8 with
distal pale pubescent band, tergum 4 with apical pale pubescent
band, tergum 5 with small lateral tufts of white hairs; sterna 2 to 4
with pale hairs apicolaterally, red to brown hairs medially; inner
surfaces of hind basitarsi and tibiae and fore and middle basitarsi
with hairs red to reddish brown. All intergrades occur between
these two extremes of vestiture coloration. The tergal pale hairs
appear in the following order: (1) base of tergum 1 and basally on
570 Tur Universrry SCIENCE BULLETIN
tergum 2, (2) lateral fasciae on tergum 2, (3) lateral fasciae on ter-
gum 8, (4) lateral fasciae on tergum 4, (5) lateral tufts on tergum 5.
The lateral fasciae become progressively wider in paler specimens
forming complete pubescent bands in the same order of appearance.
Male: Measurements and ratios: N, 20; length, 11.0-16.5 mm.;
width, 4.0-5.0 mm.; wing length, M = 23.94 ++ 0.088 mm.; hooks in
hamulus, M = 13.85 + 0.318; flagellar segment 2/segment 1, M =
2.88 + 0.062.
Structure and color: As in M. desponsa with the following differ-
ences: rarely with basal mandibular yellow spot and with small
median labial pale spot (in less than 2 per cent of specimens);
clypeus with posterior margin darkened at least in lateral angles
mesad of tentorial pits, often with posterior third to half black
and rarely more, never entirely black, usually posterior margin of
yellow clypeal area produced posteriorly in median third; sculptur-
ing as in female.
Terminalia essentially as in M. desponsa but with fewer hairs on
gonostyli and sternum 7 with slightly smaller membranous areas
between median and lateral plates and with broader, less capitate
apodemes (Figs. 8-11).
Hair: Darkest specimens as follows: head hairs white to pale
ochraceous except brown on vertex; scutellum with abundant dark
brown hairs medially, mesoscutum with posteromedian patch of
dark brown hairs extending forward to or beyond a transverse line
at middle of tegulae, other mesosomal hairs white to ochraceous;
metasomal tergum 1 with long pale hairs basally, short dark hairs
apically, tergum 2 with complete distal pale pubescent band sepa-
rated from apical margin of tergum and narrowed medially and
with basal pale pubescent band, tergum 3 with distal pale pubescent
band interrupted medially by brown hairs, terga 4-7 without pale
hairs, legs with pale hairs at least on outer surfaces of middle and
hind tibiae, inner surfaces of hind tibiae and fore, middle and hind
basitarsi with hairs red to reddish brown. Palest specimens similar
but with the following differences: without dark hairs on head,
mesoscutum or scutellum, with complete pale pubescent bands on
terga 2, 8, 4 and 5 and lateral tufts of long pale hairs on tergum 6,
with sternal hairs red medially and pale ochraceous laterally on
sterna 2 to 5, brown on sternum 6, legs with white to ochraceous
hairs except pale brown on outer surfaces of fore tarsi and tibiae,
red on inner surfaces of fore middle and hind basitarsi and hind
tibiae. As in the females, all intergrades occur between these two
Menissopes IN NortH AND CENTRAL AMERICA 571
extremes in vestiture coloration. Progressively, from darkest to
palest, pale hairs appear in the following order: (1) lateral fasciae
of pale pubescence on tergum 4 and then on tergum 5 as the inter-
rupted band of tergum 8 closes medially, (2) lateral fasciae of pale
pubescence on tergum 4 closes in medially to form a complete band
and those on tergum 5 close in as the head and thoracic hairs be-
come pale, (3) scutellum is the last area to retain brown hairs.
Bionomics: The nesting of M, rivalis has been described by Scul-
len (1928) (M. mysops). Scullen discovered a group of about
sixty individual nests scattered for a distance of about twenty feet
in a sand bank along the summit of a sea cliff in the Coos Bay
region on the Oregon coast. Both males and females were collected
on nearby thistles (Cirsium sp.). Two nests were opened on July
2, 1926, and were found to contain pollen but no eggs. On July 18
nests were again opened and half-grown larvae and eggs were found.
On July 21 seemingly full-grown larvae were present.
Scullen did not describe the nest architecture in his brief note
and has kindly forwarded his field notes so that the nests can be
more thoroughly described at this time. The following quotation
is taken from a copy of field notes made by H. A. Scullen on
July 138, 1926.
“Nests opened up show two to four branches leading off from
each main gallery. Branches leading in various directions. Leave
main gallery every one to two inches. Branches one to two inches
long, crooked. A mass of pollen apparently moistened with honey
at the bottom with one long egg on top. Pollen looked like thistle
pollen. No cover over pollen. Side gallery not apparently closed
unless by loose soil. Wall of gallery smooth and shiny as though
coated with a secretion but no noticeable layer of special deposit.
“Egg in one cell apparently suspended by a series of web like
threads attached to roof of cell. The egg was in contact with the
honey-pollen mass but was readily lifted by grasping the mass of
webbing. The webbing may have been for preventing the egg
sinking in the semiliquid mass of food material. The web might
have been threads of mould as three other masses of pollen were
found badly moulded.”
This account is interesting for at least two reasons. First, this
is the third species which has definitely been reported in the liter-
ature as having been found nesting in a large group (see M. obliqua
and M. tepida in LaBerge, 1956). However, one of the latter
(M. obliqua) is also known to nest singly. Perhaps species of
Ole Tue Universiry Scoence BULLETIN
Melissodes nest in groups only where certain areas with a favorable
substrate are found to coincide with a large population of bees,
concentrated, perhaps, by the nearby occurrence of favorable or
preferred host plants.
Secondly, the webbing by which the egg in one cell was sus-
pended from the roof of the cell is of interest. The present writer
is inclined to the second opinion expressed by Scullen, that is,
that the webbing was probably threads of mould.
The bees of this species are oligolectic, They, just as M. desponsa,
seemingly are dependent upon composites of the tribe Cynareae
and, particularly, upon plants of the genus Cirsium. Table 2
summarizes the plant-host data taken from labels of 201 specimens
representing at least 74 collections.
TABLE 2
Plant data M. rivalis data.
Family Number |Number || Number | Number |Number | ‘Total
of of of of of number
genera | species colls. 28 oo of bees
Compositae, other
$NA O78 UT ons 6 6 6 8 3 11
Compositae,
CUCU. BD ses acre | 5 58 fl 100 171
LAIAtae es. aa es 2 3 5 4 4 8
Other (4)... a 4 4 5 1 10 Ll
ML OUEIBS sian ano 13 18 74 84 eee 201
Geographical variation: As has been stated above, and as can
be deduced from the description given above, M. rivalis is highly
variable. A number of characters (4 male and 3 female) have been
examined with the viewpoint of describing significant geographical
variation. However, a statistical method for handling these data
has not yet been settled upon. The variation will, therefore, be
described below omitting such variation which is not obviously
significant.
The vestiture of both females and males varies greatly in color.
The darkest specimens occur in the humid coastal area of California
and Oregon. The females from this area can be readily recognized
Me.issopes In NortH AND CENTRAL AMERICA 578
and have been described as a species (M. desponsiformis Cock-
erell). The males, however, are not much darker than specimens
from inland California, Oregon, Washington and British Columbia
and a majority of the males from the coastal area cannot be dis-
tinguished from most of the males from the inland areas. A cline ap-
parently exists from the darkest specimens of the coastal areas to the
paler specimens of the more inland areas and no apparent step
occurs in this cline. Perhaps with many more specimens taken in
series from a number of localities in the critical areas of Oregon
and Washington, a stepped cline might be shown to exist.
The palest specimens occur in Kansas, Texas and New Mexico.
These specimens seem to be quite distinct from the specimens from
the Rocky Mountain regions. However, so few specimens are
known from this large prairie area, that definite conclusions are
unwarranted, Specimens from northern Arizona show an unusual
condition in that the females are as pale as the eastern prairie
specimens, whereas the males are as dark as, and undistinguishable
from, the northern Rocky Mountain specimens. A large gap exists
between the specimens collected in New Mexico, Texas and Kansas
and those collected in Arizona, Utah or Colorado. It is impossible
to see whether smooth or stepped clines occur between the pale
and the darker populations. Specimens from the northern prairie
region of North Dakota, Minnesota, Saskatchewan and Manitoba
are indistinguishable from the Rocky Mountain specimens. Very
few specimens are known from South Dakota and Nebraska and
nothing can be determined about the cline in color that probably
exists between the southern and the northern prairie populations.
In general, specimens from the region from the middle of Oregon
east to Minnesota, north to southern British Columbia and south
to Colorado and Utah are intermediate in color between the Cali-
fornia and Oregon coastal populations and the populations from
the southern prairies. However, each population examined ex-
hibits differences in the average darkness of the vestiture from ad-
jacent populations. The bees from this large region have been
called M. mysops Cockerell in the past.
The coloration of the male clypeus also varies greatly. To study
this character, the median length of the posterior dark area was
measured, It was found that specimens with the greatest amount
of posterior infuscation of the clypeus occur in northern Arizona
and southeastern Utah. The populations with the palest clypei,
on the other hand, were collected in New Mexico, Texas, Kansas
574 Tue Universiry SCIENCE BULLETIN
and eastern North Dakota. Specimens from western North Dakota
and southern Manitoba, eastern Wyoming and northeastern Colo-
rado, central and southern Colorado and northern Utah had the
next palest clypei. Other than the Arizona populations, those with
the darkest clypei were from Oregon and northern California. Two
marked exceptions occurred from this general distribution of the
clypeal coloration of males. First, the northern coastal Californian
specimens and, second, the southern Idaho specimens exhibited very
pale clypei. In general, there appears to be a cline in the amount of
infuscation of the posterior part of the clypei of males from the
darkest in the northwestern to palest in the southeastern and north-
eastern parts of the range of M. rivalis. Three exceptional areas
occurred: an area with dark clypei in Arizona and Utah and the two
above mentioned areas with pale clypei in California and Idaho.
No steps, other than the above mentioned irregularities were ob-
served in this cline.
The specimens also varied greatly in size. ‘Two measurements
were made on both males and females to represent size. The
median length of the clypeus and the wing length were measured
(see section on Descriptive Methodology in LaBerge, 1956, for
method of measuring). These two measurements were highly
correlated in each of the two sexes and, therefore, only one, that of
wing length, is discussed below.
The distribution of wing length means for the two sexes was not
highly correlated so they will be discussed separately. In the males,
specimens with the shortest wings occurred in the coastal areas of
Oregon and northern California. There seems to be an even cline
in increasing male wing length toward the northeast—specimens
with longest wings occurring in north central Washington (state).
The male wings decrease again in length towards the north (British
Columbia), the east (North Dakota and Manitoba) and the south-
west (Utah, Colorado and Arizona), but nowhere average as short
as in the coastal areas of northern California and southern Oregon.
The wings increase in length in New Mexico and Kansas and again
decrease in Texas, although the last decrease is, perhaps, not sig-
nificant. The clines described in this paragraph do not correlate
well with the clines described for color of vestiture or color of male
clypeus.
The female wing length follows the same clines as the male with
the following differences. In the females there appears to be a
sharp step in the cline of increasing wing length from the coastal
|
|
|
|
Me .issopEs IN NorRTH AND CENTRAL AMERICA 575
areas of northern California and southern Oregon towards the north-
east. The shortest winged specimens occur in northwestern Oregon
and southwestern Washington, whereas populations from southern
British Columbia, central Washington, eastern Oregon, eastern
Washington, Idaho and western Montana have mean wing lengths
considerably longer. The long-winged specimens from New Mexico
and Kansas contrast more strongly in the females than in the males
with populations in Texas, Colorado, Arizona and New Mexico.
From the above discussion it can be seen that clines of at least
three characters are not highly concordant. On the other hand,
clines of any two characters examined can be seen to be concordant
in at least some parts of the range of the species. No subspecies are
recognized here and it seems unlikely that any should be recognized
in the future due to the discordant nature of the clines and the
inherent difficulties in recognizing the limits of any of the races,
Perhaps when longer series of specimens from critical areas become
available for a more thorough statistical analysis, subspecies (for
which names already exist) will be recognized.
Type material: Lectotype male of rivalis from Texas is in the
Academy of Sciences of Philadelphia. The female type of despon-
siformis from Corvallis, Oregon, has not been located by the author.
It is presumably lost or destroyed. The lectoallotype male, here
designated, of mysops from Maybell, Colorado, August 1, 1904, is in
the University of Colorado Museum, Boulder, Colorado, and the
lectoallotype female, here designated, with the same data, is the
property of the California Academy of Science, but is on temporary
deposit in the collection of the Citrus Experiment Station, River-
side, California. The holotype male of hexacantha from Arizona,
August, 1902, F. H. Snow, and the holotype female of nigrosignata
from Oak Creek Canyon, Arizona, 6000 feet, August, F. H. Snow,
are in the Snow Entomological Museum at the University of Kansas,
Lawrence. The holotype male of habilis from Colorado Springs,
Colorado, 6000-7000 feet, July 20-26, 1896, H. F. Wickham is in the
American Museum of Natural History, New York City.
Distribution: M. rivalis ranges from northern California north
to southern British Columbia, east to southern Manitoba and north-
western Minnesota, south to south central Texas, central New Mexico
and central Arizona. It has been collected from May 14 to the
beginning of October but most abundantly in July and August. In
addition to the type material, 291 females and 375 males have been
576 Tue University ScreNCcE BULLETIN
examined from the localities listed below. This list includes local-
ities reported in the literature.
Arizona: Flagstaff; Grand Canyon; Houserock Canyon; Kaibab
Forest; Oak Creek Canyon; Phoenix. Catirornia: Camass Swamp,
Glen Blair Road, Mendocino Co.; Carrville; Callahan; Crescent City;
Dyerville; Fairfax; Fort Ross; Guerneville; Kneeland Prairie, Hum-
boldt Co.; Lagunitas; Myers; Point Pinos, Monterey Co.; Salinero,
Shasta Springs; Van Duzen River, Humboldt Co. nano: Burley;
Deary; Gifford; Hazelton (4 mi. N.); Idaho Falls; Lewiston; Mos-
cow; Paris; Shoshone; Twin Falls. Kansas: Dickinson Co.; Ellis
Co.; Meade Go.; Ness Co.; Pratt Co.; Rice Co.; Russell Co. Mrnne-
sora: Marshall Co. Monrana: Como Lake; Flathead Lake; Hot
Springs, Sanders Co.; Missoula; Monterey; Whitehall. Nevapa:
White Pass, White Pine Co. New Mexico: Alto; Nogal; Roswell;
Vaughn. Norra Daxora: Beach; Bismarck; Dickinson; Fargo;
Kenmare; Minot; New Rockford; Oakes; Sentinel Butte, Golden
Valley Co.; Slope Co. (Logging Camp Ranch); Stanley; Williston.
Orecon: Amity; Albany; Baker; Bellfountain; Bend; Canyon City;
Cove; Grand Ronde; Grand Ronde River (N. of Imbler); Gold
Beach; Griffin Creek, Jackson Co.; Heppner Junction, Gilliam Co.;
Hood River, Hood River Co.; Hurricane Creek (Wallowa Canyon),
Wallowa Co.; La Grande, Lake Appelgate, Jackson Co.; Lostine;
Medford; Murphy; Oakridge; Oregon City; Prineville; Redmund;
Richland; Roseburg; Siskiyou Summit, Jackson Co.; Summit Prairie;
The Dalles; Thomas; Three-sisters (Lake Creek Lodge), Deschute
Co.; Umatilla; Union; Wallowa Canyon, Wallowa Co.; Wilderville.
Sourm Daxora: Faulkton; Whitewood. Trxas: Austin; Dallas;
Eastland Co.; Hillsboro; Weser. Uran: Big Cottonwood Canyon
(near Salt Lake City); Brigham; Cache Valley (Beven Dam),
Cache Co.; Clear Creek Canyon, Sevier Co.; Cottonwood Canyon;
Echo; Fillmore; Fort Douglas; Helper; Huntsville; Hyde Park;
Laketown; Logan; Logan Canyon; Manilla; Monticello; Parly’s
Canyon (E. of Salt Lake City); Salt Lake City; South Cove Fort,
Millard Co.; South Willow Canyon, Stansbury Mts., Tooele Co.;
Thistle; Tintic; Uintah Mts., Duchesne Co.; Vivian Park; Wasatch
Mts., Tooele Co.; Wellesville; White Sands. Wasuincron: Asotin
(on Snake River); Blue Mts.; Govan; Liberty; Lind; Packwood;
Pullman; Ritzville; Tucannon Canyon, Garfield Co.; Yakima. Wvyo-
minc: Bridger Basin, Teton Co.; Carbon Co; Cheyenne; Grand
Teton National Park; Jackson; Laramie; Laramie Co.; Lusk; Rock
River, Albany Co.; Shell Canyon Falls, Bighorn Mts.; Uinta Co.;
Me.issopes In NorrtH AND CENTRAL AMERICA 577
Yellowstone National Park. Atserra: Calgary; Lethbridge; Medi-
cine Hat; Scandia. Brrrise: Corumpra: Armstrong; Lillooet (Seton
Lake); Okanagan Falls; Salmon Arm; Spulanchcan; Summerland;
Thompson River; Vernon; Walhackin. Manrrosa: Brandon; Lyle-
ton. SASKATCHEWAN: Swift Current.
Flower records: Apocynum sp., Asclepias sp., Carduus acules-
cens, Centaurea americana, Cirsium sp., C. arvense, C. lanceolatum,
C. pumilum, C. terrae-nigrae, C. undulatum, Grindelia platyphylla,
Helianthus annuus, Penstemon sp., P. cyananthus, Rudbeckia sp.,
Solidago sp., Teucrium sp., Trifolium repens, T. pratense, Verbena
sp.
INDEX
Those names now known to be synonyms or homonyms are marked with
asterisks,
PAGE
MAIIOUIIOLR: nc re cue Ae OA ees vane Cnt ae paar et 539
Shopkel ntl, «cue age Gera ea Gumiere amie Geir ae oy Mee eee te Cr 551
APOMOUSSOUCN 04 tir eh cha a9 Ph EE a Ch Oe CONG ere tte. 544
poulonalge (een ett eee eme ewe tree eee err 549
MGHUGIOUCN ) Ee yc ia homer hes a C0 i te Oe Ec ee 535
MOMIGl es Ch ba YT Od Ad A LOR Oh hs Oe 562
GOUORR erase ba caies vee RENE COs Wad C4 ae CCE ore 535
istchinla)at-|: ed irglnan saneh ade Te eee crer INGEN an err cM re hey Ce ren 562
MCleSPOMSHORENG 2%. verde ch eee Ey esas pred «OAPs 568
Masten atsid:hegy: 2 ou eeee Inrapm ee Cra Meee (ete rye aU ene ce are ae 545
MUSE ie a eet hdc da aks pe ae a es Ce 568
PIONOMONSSNGES or, sie bis Vid a ie AAA Oa ena CE ees 561
ulncyele[ intial pin he ee ee eg ee e ee re Ma eee 568
ATOTUA rs ke ce EV © RE GL OO CR ek eee ge 558
Mee ee es daa Pei ee ee ee ees tad, BOG
MIMIVSODR? 1) cdi fae oie Pls PUL Od eRe ee ee a ee 568
MRIQTINGS 00h eben ea Cee Ga SE eR ere 562
MMIGTOSMQNMLA es reer igs) SA VA pee Ce lee Cee 568
(ohotehoumh Nise gure nem UNO Oa i eerie Geet ey i 539
PSHOMTONSSOOGS 165s vai ee eda CAA Cas G0 Ow ee ee ees ee 558
ivalige ye ee ee As et a gi ek ok 568
SWHAGMNICHAG es ee eis 8 ee ES ie os 542
PO GHVINGUECOMOS @95 9) aa! fi ci Higa lees Ve ta end ia ee re . 584
NHI eg ee Oe i an gd es Vt WE WR Ry A ee ws 558
Tue Universrry ScrENCE BULLETIN
)
Sterna 7 and 8 of M. (Tachymelissodes) dagosa. 3-6, Genital capsule (dorsal
and ventral views) and sterna 7 and 8 of M. (Apomelissodes) apicata, T.
Sternum 7 of M. (Apomelissodes) mitchelli. 8-11. Apex of left gonocoxite
(dorsal view), spatha and sterna 7 and 8 of M, (Heliomelissodes) rivalis, 12-18.
Left antennae (dorsolateral view) of M. (Tachymelissodes) dagosa (12) and
M. (Psilomelissodes) intorta (13).
Ficurns 1-18. Male terminalia and antennae, approximately x45, 1-2.
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVI, Pt. I] Decemper 20, 1956 [No. 9
The Biology and Morphology of Hydrometra
martini Kirkaldy * °
BY
IsABELLE Barrp SPRAGUE
CONTENTS
PAGE
Ubningotshhis nkamhheres eS pheee ce, tReet We SAE Ai esr ey te 3 Sena ee eet! 579
AG OW ECeIMONd ast eis ity Nena by A ee Re ae 580
Nikoruphelel lola) tama Seber Ga ey EN, eer ol meee sot Ni enats Gumateeee perl NE Uy) APES 581
WIGUNOUG sore he ea bas eo eae 582
SEINE ERROR a yrs tig a a ee rn ee 582
GUE DMG erINCIE rant rs bv ee ae ea 582
TRUBYNALI SITIONS V3 aks ou es ve 584
Diunahone of the mouth Parte igs ee 585
IDISCURRION a aici naw x odes oc ee Le 586
BNO: POLE 6 ase hrs to ce wat Cc ee in Se ee 589
TENOTAGIO OXOSKElStON iad. cence ade ic cee ws: Cakes ee 589
PEOUIOIES: Rahat ads di) Flt ieee acaeey CU ie eo 589
WresOUHORhRet nt Genk Gl icra aaa &. Cain les 590
IVUGHAUIORS RE fic yen'% vey naiheyias vacd oda ale rae a Sor 591
WOR i ietace satin cack .) eee Mere so hi eee OLE
MDISCUBBIOIL: yieeee ee oe br eee 0 rei ii oie, 619
Number of generations and longevity ...................5. inc eOL0
WHC CRE eae noi le IAN PA ee 624
ADSORB EAM OO Sci, WN ohana ees vic ev eo Pe os RE ae ee 624
Development 26.0 be ee hs os a es 625
GCOUNG 2 5s eee ee a Oe ae 626
UDINCUIGHLOM oe Ge, soe ern a ac ee Cw ay, ee ed 628
GUNG AVA ce ee el et ee eee 631
PCULV ICR as a de Bee ey re a ae py te ee 631
Growth pita CieremuAuOUe ov ee eee 634
IDISCUSUION 2. sere eed se ot eas ee 642
IDEOTOMBIIE 5 ey es se a ee ae 644
SLAY, OG OMG UMONS 50) gy, ee fe ae 648
pictetonces Cue 68 i er ee, ee ee ee 652
MULCH, POU et ee ie pet ec eh Ge ee 658
Assrracr: Both apterous and the rarer winged forms of Hydrometra
martini occur throughout the northern part of the United States on slow mov-
ing or quiet waters. Overwintering takes place on land. Nymphs, whose
behavior closely follows that of the adults, are found during the spring and
summer, Individuals that will become winged may be distinguished from the
apterous form as early as the third instar.
Morphological development and differential growth of the head, thorax
and abdomen are traced through the less elongate nymphal stadia to the
slender adult forms. The internal organs of the adult animals are similar to
those of other Hemiptera but follow the attenuated body pattern.
INTRODUCTION
The purpose of this investigation is the correlation of form and
function in a single species of insect, Hydrometra martini Kirkaldy.
For this, the internal and external structures of the adults were
examined and the postembryonic development of the nymphs was
followed. The behavior of both nymphs and adults was observed
in the laboratory and in the field throughout the year.
This species is a representative of a small family, the Hydro-
HypROMETRA MARTINI KiIRKALDY 581
metridae. Of its six genera, four are monotypic. Hydrometra, the
only widely distributed genus in the family, now includes 81 species,
47 from the Eastern and 34 from the Western Hemisphere.
The biology and anatomy of the hydrometrids is known almost
entirely from the studies of the European H. stagnorum (Linnaeus )
and the North American H. martini Kirkaldy. Anatomical studies
of Hydrometra consider mainly the head with its mouth parts and
the genitalia.
As in many genera of hemipterans, Hydrometra occurs in both a
long and a short winged form. The micropterous form of H. martini
is notably more abundant in the areas in which observations were
made than is the macropterous form. However, males and females
of both are found throughout the year.
ACKNOWLEDGMENTS
I wish to thank Doctor H. B. Hungerford, under whose direction
this investigation has been carried out, for his patience, his en-
couragement and his helpful criticism. I would also like to ac-
knowledge the critical assistance of Doctors Kathleen Doering and
C. D. Michener of the University of Kansas particularly on the
sections dealing with morphology.
Much of this work has been done at Mount Holyoke College and
[ wish to thank my colleagues there, particularly Doctor Ann Haven
Morgan, for their interest and help. For the opportunity to work
at the University of Michigan Biological Station, I am indebted to
its director, Doctor A. H. Stockard.
MORPHOLOGY
The extremely slender, elongate form of the genus Hydrometra,
its most outstanding characteristic, is especially pronounced in the
adults, Males which average 10 mm. in length are about 0.5 mm.
wide in the region of the metathorax, the widest part of the body.
Females tend to be 1 to 2 mm. longer. The abdomens of those
which are laying eggs are as much as 0.2 mm. wider than the meta-
thorax.
The exoskeleton of adult hydrometrids is both hard and inflexi-
ble. The antennae, mouth parts and legs have motility because
of their membranous connections and muscular attachments. The
head has rather limited movement but the thoracic and pregenital
abdominal segments are completely fused ventrally. The genital
segments, especially those of the male, have considerable flexibility.
Tue Universiry SCIENCE BULLETIN
582
MeErHops
Adults were killed for morphological study in 80% ethy! alcohol
or in hot Dietrich’s fluid, made according to directions in Kingsbury
and Johannsen (1927) and stored in 80% alcohol. For skeletal
structures, the insects were boiled in 10% potassium hydroxide,
washed overnight in tap water and stored in 80% alcohol.
For studies of internal structures, freshly killed or fixed insects
were dissected with fine needles. Muscle and nervous tissue could
be demonstrated best in preserved material which was stained with
eosin during the dissection. Tracheae were visible only when air-
filled and the other internal structures tended to adhere to each
other after storage in alcohol. These were examined in freshly
killed material.
Relationships of internal structures, especially those of the head
region and the nervous system, were checked in serially sectioned
nymphs or recently molted adults. These were fixed in hot Dietrich’s
fluid, dehydrated in dioxane, infiltrated in 53° paraffin and imbedded
in Tissuemat. Sections cut at 7.5% with a Spencer rotary micro-
tome were stained with Delafield’s hematoxylin and eosin and
mounted in Canada balsam.
Tue Heap
External structure
The head of the adult hydrometrid is extremely slender; those
of the nymphs, especially in the early instars, are much less so.
The progressive changes in proportions of the head, antennae and
beak, as well as those of other regions are considered in the sec-
tion on growth and differentiation (see p. 582). Almost as
remarkable as the elongation of the head is its lack of clearly de-
fined sclerites. Except for the eye, the head capsule is a heavily
sclerotized tube for most of its length. The dark red, hemispherical
eyes are on the more posterior part of the head. The antennal
sockets form the widest part of the head. Each is set off by a
suture which is best seen in lateral view (fig. 7). The antennae are
four-segmented (fig. 10). The basal part of the first segment is a
rounded knob which fits into the antennal socket. This segment
is shorter than the second. The third segment is long and slender;
the fourth is subequal to the second.
The most conspicuous sclerite in the anterior part of the head
is the anteclypeus, which in lateral view (fig. 7) projects above
the surrounding areas. The other sclerites associated with the
mouth parts are lateral and anterior to it.
HypROMETRA MARTINI KiRKALDY 583
The broad labrum (figs. 5, 8) is attached to the anterior border
of the anteclypeus. A median ridge of the epipharyngeal surface
of the anteclypeus and labrum is produced distally to form a slender
epipharynx (figs. 8, 9). The labrum forms a dorsal cover for the
anterior part of the labium; the epipharynx lies within the labium,
dorsal to the mandibular and maxillary stylets (fig. 14).
Ventrolateral to the anteclypeus are two pairs of sclerites which
aid in the formation of the cranium. The more dorsal pair are
the so-called lora, lateral extensions of the hypopharnyx (figs. 5, 12).
The ventral pair are the maxillary lobes or plates (figs. 5, 20).
Deep sutures separate the lora from the postclypeus and from the
maxillary plate (figs. 5,7). The outer parts of the latter are lightly
sclerotized and, in younger adults, unpigmented. Ventral pro-
jections of the maxillary plates, the buccalae, partially surround
the base of the beak and prevent its lateral movement (figs. 7, 20).
The membranous proximal part of the labium inserts on the
inner surface of the maxillary plates.
In the resting position, the beak is parallel to the head and
lies close to it (fig. 7). Of the four labial segments, the third is
by far the longest. It is marked, except at either end, with dark
striations. The first and second segments are short and broad.
The fourth, another short segment, terminates in a point. The
labium is slightly flexible but the intersegmental sutures are not
movable,
The edges of the labium fit together tightly (fig. 14). Cross
sections of a fifth instar nymph show variations in the closure at
different levels, as well as differences in width of the labium. At
both the proximal and distal ends, the sides merely meet each other,
whereas in the middle portion they are grooved. The epipharynx,
the most dorsal structure within the labial furrow, extends only
through its proximal segment. The hollow mandibular stylets are
lateral to the maxillary stylets throughout their length (fig. 14).
Both pairs extend to the tip of the labium and protrude from it
when the animal feeds. The tips of the mandibles are spear-shaped
and bear rows of recurved barbs (fig. 11).
A ridge on the right maxillary stylet fits into a groove on the
left one; thus, the maxillae are locked together to form a single
tube. The distal end of each maxilla bears a row of minute setae.
On the left stylet these are spaced evenly (fig. 18) but on the
right, they are arranged in groups of three or four which fasten
together so that they appear to be heavier teeth (fig. 17).
584 Tue Universiry Science BULLETIN
Internal Structure
Two slender apodemes extend posteriorly from the basal segment
of each antenna. A featherlike muscle attaches each of these to the
side of the cranium (fig. 10). The muscle of the outer apodeme
is slightly larger and is lateral to that of the inner apodeme.
Figures 11 and 19 show the mandible and its associated struc-
tures. As the stylets pass between the anteclypeus and the body
of the hypopharynx, they are slightly lateral and dorsal to the
maxillae. Within the head, a membranous fold attaches a vertical
bar of the mandibular stylet of the rectangular mandibular lever.
A second heavy membrane extends from the lower anterior corner
of the lever to the lorum. A slender apodeme continues posteriorly
from the mandibular lever to the occipital region of the head.
The posterior part of this process is surrounded by a retractor mus-
cle which originates on the lateral cranial wall from the posterior
margin of the eye to the occiput. The protractor is a prominent
fan-shaped muscle which attaches by a short, slender ligament-
like apodeme to the lower posterior corner of the lever. This mus-
cle originates on the cranium behind the antennal socket, near
the origin of the muscles of the antenna.
Each maxilla is held in place by a membranous sheath, the
anterior end of which attaches to the posterolateral edge of the
body of the hypopharynx (fig. 12). The sheath extends a half of
the length of the stylet. Both it and the stylet itself are enveloped
by the heavy protractor which has its origin on the maxillary plate.
Since the stylet becomes wider posteriorly, its forward movement is
limited by the sheath. The retractors, which insert on the pos-
terior part of the stylet, originate on the occipital regions of the
head capsule. The protractors insert just anterior to the retractors
and pass under the hypopharynx to attach to the posterior border
of the maxillary plate or lobe.
The hypopharynx consists of a medial bridge which is internal
and a pair of lateral wings, the lora, which form a part of the head
capsule. The maxillary sheaths and salivary syringe (figs. 12, 15)
are derived from the hypopharyngeal bridge. The salivary syringe,
a lightly sclerotized funnel-shaped structure, opens by a duct on the
hypopharyngeal bridge just ventral to the opening of the food tube.
The plunger, fitting closely into the funnel, has a very delicate
apodeme which extends posteriorly near the ventral surface of the
head. A heavy muscle originates on the ventral wall of the head
capsule and inserts on the base of the plunger and along the length
of its apodeme.
HypROMETRA MARTINI KiRKALDY 585
The pharynx (figs. 16, 19) is a lightly sclerotized, slender tube
which extends the length of the head, Anteriorly its floor flattens to
form a narrow lip which rests on the body of the hypopharynx. The
mouth opening is closely associated with the tube formed by the
maxillae as they emerge from their sheaths and unite to make the
sucking tube. A series of straplike muscles which originate on the
cranial wall and insert on the pharynx operate the pharyngeal pump.
Since the origin of these muscles identifies the frons, this area
extends two thirds of the length of the head to the interocular
region,
Function of the Mouth Parts
A description of the food finding process and of the kinds of food
used by Hydrometra martini is given in the section dealing with
behavior. In general, hydrometrids feed on living or recently
killed animals by inserting their mandibular and maxillary stylets
and. sucking out the fluid content.
In preparation for spearing its prey, the animal swings its beak
forward, and thrusts it toward the victim with the whole body.
When this is a dark shiny insect, a minute drop of saliva glistens on
the exoskeleton at the point at which the stylets begin to penetrate.
The saliva, secreted by the paired salivary glands located in the
thorax, enters the salivary syringe through ducts on the ventral side
of the head, and is pumped by the syringe through its medial duct
to the groove of the maxillary stylets.
The spearlike tips of the mandibles pierce the prey and hold it so
securely with their retrorse barbs that small animals can be lifted
and carried considerable distances. Extension of the mandibles is
effected by contraction of their protractors. This action tips the
entire mandibular lever and the attached stylet forward. The
mandibles retain their position while the maxillae protrude between
them in feeding. The contraction of the retractor muscles with-
draws the mandibular stylets.
The slender maxillary tube not only injects saliva into the prey
but also carries the fluid food into the mouth, Quick thrusts of the
maxillae result from alternating contractions of the protractor and
retractor muscles within the head. The tips of the maxillae have
a tonguelike action, lashing and gouging within the soft tissues. The
actual sucking seems to result from the dilation of the food tube
through muscular action of the pharyngeal dilators which originate
in the anteclypeus and frons.
THe University SCreENCE BULLETIN
Discussion
The most thorough study of the structure of the hydrometrid
head is that reported by Ekblom (1926). He worked out the rela-
tionships not only of the mouth parts but also of their musculature,
Severadei (1950) added a description of the salivary syringe.
Spooner (1938) included drawings of three views of the head and
one of the mandibular lever of H. martini in his comparative study
of the head capsules of Hemiptera. Other papers concerning the
homologies of the hemipterous head are those of Snodgrass (1938,
1944, 1947), Butt (1943), DuPorte (1946) and Kramer (1950).
In Hydrometra, as in all Hemiptera, there has been much con-
fusion as to the homologies and hence the names of the various
sclerites. This has been especially true of the two pairs of sclerites
at the anterior end of the head. Ekblom (1926) called the more
dorsal pair the lamina maxillaris and the structures which sur-
round the base of the labium, the lorae. Many taxonomists have
called the dorsal pair the juga in Hemiptera. In drawings of
Oncopeltus, Butt (1943) labeled the external portion of the cor-
responding lobe the juga but he referred to its internal extension
as the lorum. Spooner (1938), considered the more dorsal pair of
plates to be the paraclypeus and the more ventral pair, the maxil-
lary plates.
In a paper published later in the same year, Snodgrass (1938)
discussed the homologies of these sclerites. His conclusion, that
the dorsal sclerites are lateral extensions of the hypopharynx, was
based on the facts that: 1) although the plates adjoin the post-
clypeus, they are continuous with the hypopharynx in the area ven-
tral to the anteclypeus, and 2) the protractor muscles of the mandi-
bles originate on it. The ventral pair, Snodgrass (1944) called
maxillary lobes.
In the only hemipteran which he illustrated, DuPorte (1946) des-
ignated the dorsal sclerites discussed above as the hypopharynx.
Kramer (1950) working with representative auchenorhynchous
Homopetra called it “lorum” and pointed out the relationship with
the hypopharynx.
Both Ekblom (1926) and Severadei (1950) correlated the elonga-
tion of the head and that of the mandibular and maxillary ap-
paratus. Ekblom (1926, 1930) described the attenuated mouth
parts, especially maxillae, in several relatively short-headed semi-
aquatic hemipterans. In these forms, the stylets and their asso-
ciated muscles extend deep into the thorax. The maxillae and
HypROMETRA MARTINI KiRKALDY 587
mandibles of Hemiptera, like those of mandibulate insects, arise
embryologically from paired evaginations of the head (Johannsen
and Butt, 1941). Newcomer (1948), who investigated the develop-
ment of the mouth parts of Oncopeltus found that the maxillary
plates (= maxillary lobes) differentiate from the bases of the
maxillary processes before blastokinesis. This corroborated the
homology established by Snodgrass (1944), who, working with
Magnicicada adults, identified the maxillary lobe as the stipes.
Snodgrass also found that the stylet is homologous with the lacinia,
and a lateral distal process of the maxillary lobe as the galea. The
latter, which is present in the adult homopterans figured by Spooner
(1938), is reduced or absent in his drawings of hemipterans.
In Oncopeltus (Newcomer, 1948), the maxillary and mandibular
evaginations elongate at blastokinesis and their proximal ends sink
deeply into the head. The ectoderm surrounding the base of each
developing stylet invaginates to form a sac or pouch. The levers
and sheaths of the maxilla and the mandibular levers are derived
from the corresponding sacs.
The origin of the protractors of the maxillae on the maxillary lobe
seems to be quite constant. Kramer (1950) reported this relation-
ship in the homopterans which he examined. Ekblom (1926, 1930)
described it in a variety of hemipterans. Butt (1943) and New-
comer (1948) found that in the milk weed bug, Oncopeltus fasci-
atus, the protractors also originate on the maxillary lobe. Griffith
(1945) stated that in Ramphocorixa acuminata the protractor
muscle of the maxillary stylets attach on the margins of the labial
wall and on the attenuated hypopharyngeal processes.
The insertion of the protractor is typically on the maxilla near its
base but below the attachment of the retractor. This is the case
in H. martini and in all of the hemipterans discussed by both Ek-
blom and Butt with the exception of Notonecta (Butt, 1943). Here
the protractor inserts on the end of the maxilla and the retractor
on the side of the stylet, anterior to the attachment of the protractor.
The retractor of the maxilla extends in both homopteran and
hemipteran forms from its insertion on the end of the stylet to the
posterior part of the head capsule. In those forms combining un-
usually long maxillae with head capsules of the normal length
(Gerris, Velia and Mesovelia) a pair of projections of the head
capsule extend into the thorax and serve as places of attachment
for the retractors (Ekblom, 1926, 1930).
A maxillary lever or bar, which is found typically in the Homop-
588
THe Universtry SCruneNCE BULLETIN
tera (Kramer, 1950), is present in Oncopeltus (Newcomer, 1948)
and in some other hemipterans (Ekblom, 1926, 1930). In the
homopterans, this lever is associated with the posterior tentorial
bridge. In some, the cicada, for example, it is retracted by a muscle
which arises on the tentorium and is protracted by one from the
maxillary lobe. According to Ekblom (1926), when a maxillary
lever is present in hemipterans, it lacks independent muscular at-
tachments and seems to serve as a guide.
Another structure closely associated with the maxilla is the maxil-
lary sheath. Earlier workers (Ekblom, 1926, 1930; Hamilton, 1931)
considered this to be the tentorium. Spooner (1938) in his dis-
cussion of the homologies of the head capsule, reported that a true
tentorium appears in the Homoptera together with the sheaths
which serve as guides for the maxillary stylets. Although in the
Hemiptera the tentoria are lacking, the sheaths, which he consid-
ered to be hypopharyngeal evaginations, remain.
The mandibular apparatus of H. martini seems to be typical of
the group of semiaquatic Hemiptera. In all described homopterans
and hemipterans this consists of the paired stylets each of which
is supported by a lever, protractor muscles which insert on the
lever, and a retractor muscle which inserts on an apodeme extending
posteriorly from the stylet.
There is considerable variation in the place of origin of the
protractor. According to Kramer (1950), in the auchenorhynchous
Homoptera it originates on the ventral and anterior margins of the
lorum. In some of the Hemiptera a muscle with these attachments
is also the only mandibular protractor. Butt (1938) reported this
to be the case in Notonecta, Oncopeltus and Benacus, as well as in
the homopterans Tibicina and Cephisus. In the pentatomid Acro-
sternum he found an additional protractor which inserts on the
mandibular lever but originates on the hypopharynx.
Ekblom (1930) described two protractor muscles of the mandi-
ble in only one hemipteran, the corizid Myrmus miriformis Fall.
Both of these muscles insert on the mandibular lever; one on the
lorum and the other on the head capsule. In the other species
which he studied, the only protractor is the fan-shaped muscle aris-
ing on the cranium.
In Hydrometra martini the salivary syringe and its duct are
similar in form, position and function to those found in the homop-
terans and other hemipterans (Butt, 1943; Weber, 1930). The fact
that Hydrometra has only one groove in the maxillary tube has
HyDROMETRA MARTINI KiRKALDY 589
interested various authors since it was first described by Ekblom
(1926). He and Severadei (1946) associated this with the fact
that Hydrometra eats dead or nearly dead insects. Since the sali-
vary glands as well as the salivary syringe are well developed, it
seems likely that the single maxillary channel serves to transport
both saliva and food.
The pharyngeal pump in some predacious Hemiptera (for ex-
ample in Gerris and Reduvius, according to Weber, 1930) is oper-
ated by muscles which form two or three groups. In Hydrometra
these are spread along the very long pharynx but the muscle fibers
bear the usual relationship to it and to the cranium, making a V-
pattern in cross section.
THe THORAX
The thorax of the wingless hydrometrids is more slender and less
flexible than that of the winged forms. Figures 23 through 27 show
the thorax of the wingless and figures 21, 22, 28 and 29 that of the
long winged form. In both, only the metathorax is notably elongate.
In addition to the muscles of the appendages, the thorax contains
the esophagus, the salivary glands, the fused thoracic and abdominal
ganglia, the anterior part of the heart and the thoracic tracheae.
Both the adults and the nymphs lack scent glands. The skeleton
and its musculature will be considered here; the other internal
structures will be discussed later. Comparisons of the nymphs of
macropterous and apterous forms, as well as a consideration of
the progressive changes in proportion and differentiation through
the nymphal instars, are also presented later (pp. 633-34).
Thoracic exoskeleton
PROTHORAX
The anterior margin of the prothorax is a heavy rim formed by
the inflection of its border. The cervical membrane, continuous
with this inflection, is short so that the occiput telescopes slightly
into the prothorax. This arrangement limits the movement of the
head in any direction.
Punctuations surround the anterior margin of the prothorax in
an irregular ring and also mark the dorsal part of the pronotum
(figs. 21, 23). A similar punctation marks each of the supracoxal
lobes, except that of the metathoracic epimeron (figs. 22, 24).
The pronotum is continuous laterally with the pleura and forms
a posterior lobe which covers the mesonotum and the anterior part
of the metanotum. The large size of this posterior lobe in the
590 Tue Universrry ScreNCE BULLETIN
winged individuals corresponds with the general difference in the
size of the two forms. A fissure separates the episternal and epi-
meral supracoxal lobes. The first thoracic (mesothoracic) spiracle
lies just behind the supracoxal lobe of the epimeron. Depressions
of the sternum form the inner walls of the coxal cavities. The small
furca are medial to them.
MESOTHORAX
The mesothorax of the wingless forms (figs. 26, 27) not only is
smaller but also shows less differentiation than that of winged
individuals (figs. 28, 29). The tergal area, which is completely
covered by the posterior lobe of the pronotum, is reduced to a
delicate membrane in wingless hydrometrids. In these animals the
phragmata are not developed. Neither the prealare bridge nor the
prealare membrane is differentiated; apparently they are incorpo-
rated in the tergal membrane. The wing pads, the bases of which
are covered by the pronotal lobe, are small straplike processes near
the midline of the notum.
In winged forms, the tergum is less heavily sclerotized than the
rest of the exoskeleton but it is differentiated into clearly marked
sclerites. The tongue-shaped first phragma projects anteriorly and
somewhat ventrally from the antecostal suture. The acrotergite
(precosta) is narrow. The prescutum is set off from the scutum
both laterally and posteriorly by the parapsidal sutures. The pos-
terior angles of the scutum form the tergal wing process. A wide
membrane connects the scutum and scutellum laterally. The antero-
lateral angles of the scutellum are extended to form the axillary
cords of the first pair of wings. Ventrally the wing bases are sup-
ported by the pleural processes. A pair of short processes of the
scutellum articulates with the postnotum, which is represented by a
pair of lateral sclerites; medially the sclerite is lacking. Neither
a basalare nor a subalare is present.
Although the pleural region is similar in the two forms, it is
larger in winged individuals. The pleural wing processes, which
are more developed in the macropterous hydrometrids, extend dor-
sally and obliquely anterior from the mesothoracic-metathoracic
juncture. A fissure, which is continuous with the lower part of
the pleural suture, separates the anterior supracoxal lobe, formed
by the episternum, from the posterior supracoxal lobe, formed by
the epimeron. Each epimeron bears a second (metathoracic)
spiracle. There is no separation of the mesothoracic sterna from
either the pleura or the metasterna. Deep depressions of the sterna
HypROMETRA MARTINI KiRKALDY 591
form the inner walls of the coxal cavities. The furcae, which lie
just medial to these cavities, are much larger in the long-winged
animals,
METATHORAX
The extremely long metathorax forms an almost inflexible cylinder
which is continuous ventrally with the mesothorax and abdomen.
The large episterna are fused with the sternum (fig. 27). Poste-
riorly these sclerites form the anterior supracoxal lobes which are
separated from the small posterior (epimeral) lobes by fissures
continuous with the pleural suture. The epimera are not differ-
entiated from the abdominal pleura.
The metanotum consists of three areas. For most of its length
it is a narrow internal ridge, compressed by the episterna. The
second phragma, well developed in winged individuals, adheres
along the ridge and expands into a broad, concave vertical plate
anterior to the metapostnotum. ‘The latter, a small triangular
sclerite, lies between the episterna in the middorsal line.
The most conspicuous structure in the anterior part of the meta-
tergum is the heavy ridge on which the posterior pronotal lobe
rests. This forms an arch whose arms are medial to the mesopleural
wing processes. This ridge is especially pronounced in the macrop-
terous individuals. A less heavily sclerotized portion of the meta-
notum extends forward from the ridge and articulates with the
mesopostnotum. Laterally a pair of triangular sclerites, evidently
fragmenta of the notum, extend medially to the wing bases.
The Legs (Figs. 80, 31, 32)
The legs of Hydrometra martini are extremely slender. The
paired claws of each leg are inserted at the apex of the third tarsal
subsegment. The distal end of each tibia and the tarsal subseg-
ments are covered with short stiff setae which are used in grooming.
Each coxa is almost completely enclosed in the cavity formed
by the supracoxal lobes and the sternum. At rest, the prothoracic
and mesothoracic legs extend anteriorly; the metathoracic legs, pos-
teriorly. In each, the very large coxal process articulates with the
supracoxal lobes just dorsal to the fissure between them. This
articulation is lateral in the first and second pairs of legs and nearly
posterior in the third.
Two small sclerites, the basicoxa and the trochantin, are associ-
ated with the articulation of the coxa. Both are present at the base
of each leg and are considerably smaller in the wingless than in the
winged forms. The trochantin is attached to the proximal coxal
592, Tue Universrry Science BULLETIN
margin, anterior to the coxal process. The basicoxal sclerite, which
lies behind this process, is associated with the tendon of the
principal coxal retractor.
The Wings (Figs. 33 and 34)
The base of the mesothoracic wing lies between the scutal wing
process and the lateral arm of the scutellum. The metathoracic
wing attaches just posterior to it. Figure 85 shows the articular
region in side view; the wings are lifted to show the basal sclerites.
The third axillary sclerite of the mesothoracic wing can be iden-
tified positively since the direct flight muscle (M 88), which arises
on the episternum, inserts on this sclerite. Its distal projection
articulates with the posterior margin of the wing. A short thick
sclerite, which extends distally from the pleural wing process, ar-
ticulates with the scutal wing process, the pleural wing process,
the third axillary sclerite and another elongate sclerite. A heavy
sclerite, which articulates also with the scutal wing process, ex-
tends along the anterior margin of the wing. Distally this sclerite
makes contact with the base of a heavy vein which seems to be
the proximal part of the media, since it serves as the pivot of the
wing. A long arm of the first axillary sclerite extends between the
_ second and third axillary sclerites. This sclerite is not closely as-
sociated with any of the wing veins.
There are five longitudinal veins in the mesothoracic wing. ‘Two
of these are short and run into the posterior margin. One vein,
lying in the middle of the wing, extends almost its length. The
other two, near the costal margin, run separately proximally and
fuse with each other distally. Two cross veins connect the
main longitudinal veins in the distal part of the wing.
The base of the metathoracic wing articulates with two small
sclerites. One abuts on the heavy anterior vein and the other ex-
tends from the lateral arm of the notum toward the media. A heavy
vein supports the base of the posterior edge of the wing.
Thoracic Musculature
Larsén (1945a, 1945b) in his comprehensive studies of the he-
mipterous thorax included a winged South American form. The
winged Hydrometra martini resembles it closely, while wingless
individuals lack the flight muscles except M. furcapleuralis (M 89).
The differences, together with specific information as to the origins
and insertions of the muscles, are given here with the names and
numbers used by Larsén. Although they were not studied, the leg
muscles distal to the coxa are listed according to Larsén’s findings.
HypROMETRA MARTINI KiRKALDY 593
Prothoracic Muscles (Figs. 36, 37)
M1. M. pronoti primus. Levator of head. Arises on pronotum
and inserts by apodeme on postoccipital ridge near mid-dorsal line.
M 2. M. pronoti secundus. Rotator and depressor of head.
Arises on pronotum and inserts on ventro-lateral part of postoccipital
ridge.
M 38. M. pronoti tertius. Levator of head. Origin from first
phragma in winged form and from anterior margin of mesonotum
in wingless form. Insertion on dorsal part of postoccipital ridge.
M 5. M. pronoti quintus. Depressor of prothorax. Arises on
lateral margins of first phragma and inserts on lateral part of pro-
notum. Not present in wingless forms.
M 6. M. prosterni primus. Depressor of head. Arises on furca
and inserts on ventral margin of postoccipital ridge.
M 7. M. prosterni secundus. Levator of head. Arises on ster-
num and inserts on lateral part of postoccipital ridge near insertion
of M 2.
M 10. M. proepisterno-postoccipitalis. Rotator and depressor
of the head. Arises on the episternum and inserts on the postoccipi-
tal ridge above insertion of M 6.
M 13. M. noto-trochantinalis, Rotator of coxa and elevator of
leg. Arises on the posterior part of pronotum and inserts by an
apodeme on the trochantin.
M 14. M. noto-coxalis primus. Rotator of the coxa and elevator
of leg. Arises on the notum and inserts with M 16 and M 17 ona
tendon.
M 16. M. noto-coxalis tertius. Rotator of coxa and elevator of
leg. Arises on notum. Insertion on apodeme associated with basi-
coxal sclerite.
M 17. M. pleura-coxalis. Rotator of coxa and elevator of leg.
Arises on episternum and inserts on apodeme with M 14.
M 19. M. furca-coxalis posterior. Retractor of leg. Arises on
furca and inserts.on medial proximal border of coxa.
M 20. M. noto-trochanteralis. Depressor of trochanter. Arises
on notum medial to M 16. Inserts by a long apodeme with M 21
and M 23 on the posterior process of proximal part of trochanter.
M 21. M. pleura-trochanteralis. Depressor of trochanter. Arises
On episternum. Inserts with M 20 and M 23.
M 23. M. coxa-trochanteralis medialis. Depressor of trochanter.
Arises on posterior wall of coxa and inserts with M 20 and M 21.
M 24. M. coxa-trochanteralis lateralis. Levator of trochanter.
20—3378
594 THe Unriversiry SciENCE BULLETIN
Arises on anterior and lateral wall of coxa and inserts on anterior
process of the trochanter.
M 25. M. reductor femoris. Arises on anterior wall of the tro-
chanter and inserts on proximal part of femur.
M 26. M. depressor tibiae. Arises on ventral part of the femur
and inserts by apodeme on the ventral part of tibia.
M 27. M. levator tibiae, Arises on dorsal part of femur and in-
serts by tendon on dorsal part of tibia.
M 28. M. depressor tarsi. Arises on tibia and inserts by apodeme
on ventral part of Ist tarsal segment.
M 29. M. depressor practarsi. Arises on tibia and inserts on the
unguitractor apodeme.
Mesothoracic muscles
M 30. M. mesonoti primus. Indirect flight muscle; depressor
of forewings. From ventral surface of first phragma and prescutum
to anterior surface of second phragma. Lacking in apterous forms.
M 34. M. dorsoventralis primus. Indirect flight muscle; elevator
of the forewings. Attaches to scutum lateral to parapsidal suture.
Lacking in apterous forms.
M 38. M. episterno-alaris. Direct flight muscle; flexor of fore-
wing. Arises on episternum near its upper edge, lateral M 34. Not
present in apterous forms.
M 39. M. furca-pleuralis. Regulates position of the pleural
process. Inserts on this process and arises on furca. Reduced in
apterous forms.
M 40. M. noto-trochantinalis. Rotator of coxa and extensor of
leg. In winged forms, arises on posterior part of scutum; in wing-
less individuals, on mesonotal membrane. In both, inserts by
apodeme on trochantin.
M 41. M. noto-coxalis, Rotator of coxa and elevator of leg.
Arises on scutellum in winged forms and on notal membrane in
wingless forms. Inserts by long apodeme on lateral part of coxal
margin. Insertion associated with small sclerite, free basicoxa.
M 42. M. episterno-coxalis. Rotator of coxa and extensor of
leg. Arises on the episternum and inserts on antero-medial margin
of coxal margin.
M 45. M. furca-coxalis posterior. Extensor of leg. Arises on
furca and inserts on medial margin of coxa.
M 46. M. noto-trochanteralis. Depressor of trochanter. Origin
on scutellum posterior to that of M 40 in winged individuals; in
}
HYDROMETRA MARTINI KirKALDY 595
wingless forms on notal membrane. Inserts on a long apodeme
with M 47 on ventral edge of trochanter.
M 47. M. pleura-trochanteralis. Depressor of trochanter. Arises
on pleura and inserts on apodeme with M 46.
M 48. M. furca-trochanteralis. Depressor of trochanter. Arises
on furca and inserts on trochanter apodeme.
M 49. M. coxa-trochanteralis medialis. Arises on medial coxal
wall and inserts on trochanteral apodeme.
M 50. M. coxa-trochanteralis lateralis. Arises on lateral coxal
wall and inserts on lateral wall of trochanter.
M 51. M. reductor femoris. Like M 26 of prothorax.
M 52. M. depressor tibiae. Like M 27 of prothorax.
M 58. M. levator tibiae. Like M 28 of prothorax,
M 54. M. depressor tarsi. Like M 29 of prothorax.
M 55. M. depressor praetarsi. Like M 80 of prothorax.
Metathoracic Muscles
M 57. M. metanoti secundus. Indirect flight muscle. Extensor
and levator of the hindwing. Arises on third phragma and inserts
on small anterior sclerite of notum, Not present in apterous forms.
M 62. M. furca-pleuralis. No apparent function. Arises on
furca and inserts on episternum. Reduced in wingless individuals.
M 63. M. noto-trochantinalis. Rotator of coxa and promotor of
leg. Arises on anterior part of notum and inserts on trochantin.
M 64. M. noto-coxalis. Rotator of coxa and retractor of leg.
Arises on notal ridge. In winged forms, insertion on medial coxal
margin is associated with free basicoxal sclerite.
M 66. M. episterno-coxalis, Rotator of coxa and extensor of
leg. Arises on intersegmental line and inserts by an extremely long,
slender apodeme which is lateral to other muscles and their tendons
on lateral part of trochantin.
M 69. M. furca-coxalis posterior. Arises on furca and inserts on
median coxal margin. Rotator and adductor of coxa.
M 70. M. noto-trochanteralis. Depressor of trochanter. Arises
on notal ridge posterior to origin of M 64. Inserts with M 71 and
M 72 by a long apodeme on posterior margin of trochanter.
M 71. M. pleura-trochanteralis. Depressor of trochanter. Arises
on pleura and inserts with M 70 and M 73.
M 73. M., coxa-trochanteralis medialis. Depressor of trochanter.
Arises on anterior wall of coxa and inserts with M 70 and M 72.
M 74. M. coxa-trochanteralis. Levator of trochanter, Arises
596 Tue Universiry SciENCE BULLETIN
on posterior and lateral wall of coxa and inserts on posterior (dorsal )
margin of trochanter.
M 75. M. reductor femoris. As M 25 and M 51.
M 76. M. depressor tibiae. As M 26 and M 52.
M 77. M. levator tibiae. As M 27 and M 58.
M 78. M. depressor tarsi. As M 28 and M 54.
M 79. M. depressor praetarsi. As M 29 and M 55.
Discussion of the Thorax
Little was reported on the hemipterous thorax prior to the
thorough studies of Larsén in 1942 and 1945. Both Gulde (1902)
and Brindley (1930) called attention to the fact that Hydrometra
lacks scent glands. In early comparative work on the thoraces of
both Homoptera and Hemiptera, Taylor (1918) considered the ci-
cada in detail. Among the other forms, he included a gerrid.
Differences in the interpretation of the homologies of the various
sclerites and lack of detail in the drawings make this paper and
others, that on the stink bug by Tower (1913) for example, less
valuable than Larsén’s work in present investigations. The struc-
ture of numerous homopterans has been investigated by Kramer
(1950). The thoraces in this group are markedly different from
those of the Hemiptera.
In 1942, Larsén described the metathorax of a number of insects,
a few of which he figured. Gerris was discussed in considerable
detail and Hydrometra was compared with it briefly. In the later,
more comprehensive papers (1945a, 1945b), he discussed the
thoracic structure and its musculature in 26 species from 25 families
of hemipterans, including a macropterous gerrid, an apterous veliid,
and a macropterous hydrometrid. He illustrated the latter with
two figures, one of thoracic musculature and the other of the second
phragma.
The present study compared the winged and wingless forms of
Hydrometra martini with the macropterous South American H ydro-
metra, the macropterous Gerris rufoscutellatus and Velia currens
which were described by Larsén.
The prothorax of Hydrometra martini is apparently very similar
to that of other hemipterans. The length of the pronotal lobe varies
with the size of the mesonotum which it covers; both are markedly
larger in macropterous than in apterous individuals.
Well-developed supracoxal lobes (the acetabular caps of tax-
onomists) are usually found in Hemiptera (Larsén, 1945b). The
leg articulation with the trochantin imbedded in the anterolateral
HyYDROMETRA MARTINI KIRKALDY 597
portion of the coxal membrane and the free basicoxal sclerite asso-
ciated with the tendon of the coxal retractor (M 16) is also typical
(Larsén, 1945a). Both of these sclerites are small in Hydrometra.
The prothoracic muscles in the wingless and winged forms are
the same and like those of Larsén’s species except that the apterous
individuals lack the prothoracic depressor (M5), which inserts
on the base of the first phragma. Although most of the leg move-
ment is in a horizontal plane, some elevation and depression is
possible. Both the prothoracic and mesothoracic legs can swing
from a nearly anterior position to a nearly posterior one.
The mesothorax of both Gerris and Hydrometra closely resembles
that of other Hemiptera, characterized by a single scutal process
for each wing, rather prominent parapsidal sutures, a well-devel-
oped first phragma and a scutellum which articulates with the
postnotum laterally. In many of the forms studied by Larsén,
as in Hydrometra, the postnotum consists of lateral sclerites, the
median section being lacking. Laterally the scutum and scutellum
in Gerris and Hydrometra are separated by a wide, lightly sclero-
tized band, called by Larsén the “tergalspalt.” The flight muscles
in H. martini appear to be identical with those of the species de-
scribed by Larsén.
The mesonotum of the apterous form of Hydrometra martini
shows interesting similarities to that of the wingless Velia currens
studied by Larsén (1945a). In both, the tergal sclerites are re-
duced to an undifferentiated membrane. The phragma is reduced
in Velia; in H. martini it seems to be lacking. In both species there
is a correlated loss of the large indirect flight muscles (M 30 and
M 34) and the direct flight muscle (M 88). Musculus furca-
pleuralis (M 52) which is lacking in Velia is reduced in H. martini.
The leg articulation in the winged individuals differs from that
in the wingless individuals only in having larger basicoxal sclerites
and larger trochantins. The muscles and leg movement are similar
to those of the prothorax.
The metathorax of Hydrometra is markedly different from that
of other hemipterans which have been investigated. In the reduviid,
Emesa, which also has an extremely elongate body, both the meso-
thorax and metathorax are long and both have clearly defined tergal
Sclerites (Taylor, 1918). Hydrometra has not only a metathorax
which is much longer than the mesothorax, but also a reduced ter-
gum. That the narrow middorsal line with its corresponding in-
ternal ridge is the notum is shown by the origin of the leg muscles
(M 63, M 64 and M 70) on it.
598 Tur Universrry Science BULLETIN
The metapostnotum, which according to Larsén (1942) can be
distinguished from the metanotum by the absence of notal leg
muscles, is small. M. metanoti secundus, an indirect flight muscle,
takes its origin on it in winged animals but is lacking in apterous
hydrometrids. This muscle is present in Gerris rufoscutellatus and
absent in wingless specimens of Velia currens.
It is difficult to interpret the homologies of the anterior part of
the metathorax. The heavy ridge which characterizes this area
in apterous as well as macropterous forms of Hydrometra martini
is not mentioned by either Taylor (1918) or Larsén (1945a) in
any of the insects which they described. It seems to be a secondary
development which gives rigidity to the area. No other differentia-
tion is found in the wingless H. martini.
Larsén (1942) pointed out that in Hydrometra as well as in
Gerris, the anterolateral angles of the metanotum are separated
to form small independent sclerites above the wing bases. These
serve as places of insertion for a pair of indirect flight muscles
(M 57) which originate on the third phragma. He found homol-
ogous structures in a number of terrestrial hemipterans which also
have a direct flight muscle (M 61) which inserts on the third axil-
lary sclerite. Neither Gerris nor Hydrometra has this muscle. Three
other landmarks of the typical thoracic region are missing: the
basalare which is never found in Hemiptera as an independent
sclerite, the subalare which is not found in many aquatic or semi-
aquatic bugs and the metapleural wing process which all he-
mipterans lack.
Two sclerites are present in the base of the hindwing. One,
which acts as a fulcrum and extends toward the media, is probably
the second axillary sclerite. The other sclerite articulates with
the anterior margin of the wing and is probably the humeral plate.
If this is so, the first and third axillary sclerites are missing. This
area is not described by Larsén in Hydrometra or in any form which
shows such a marked reduction of sclerites.
The extremely large size of the second phragma is made possible
by the elongation of the metathorax. It has the same relative posi-
tion in Hydrometra that it has in other hemipterans. Another ef-
fect of this elongation is the manner in which the leg muscles insert.
The origins of all of the muscles are anterior to the leg base and
each inserts with a long apodeme. These connections limit the
horizontal movement of the hindlegs. Because of the comparatively
larger size of the metepisternum, the distance between the middle
HypDROMETRA MARTINI KIRKALDY 599
and hind coxae is much greater than that between the fore and mid-
dle coxae.
Wing venation in the Hemiptera has not been the subject of re-
cent investigation. Comstock and Needham (1898) studied a num-
ber of nymphs and figured a nymphal pentatoid wing. They found
that the most conspicuous feature in these wings was the coalescence
of the subcosta and radius. Two papers published in 1926, one
by Hoke and one by Tanaka, described the venation in a large
number of hemipterous nymphs and adults. In general, the
nymphal wing includes the typical veins; the adult wing shows
considerable reduction. Of those described, the wing of Hydro-
metra most closely resembles those of Mesovelia and Merragata,
which were figured by Hoke. However, both of these wings have
a small but clearly developed anal region which Hydrometra lacks.
Unfortunately, neither of these papers included results of a study
of positions of the wing sclerites. A thorough review of the wing
venation with consideration of the basal sclerites seems to be essen-
tial for the establishment of homologies of the veins. Such a study
would be interesting from the point of view of phylogenetic re-
lationships.
THe ABDOMEN
The abdomen of the female hydrometrid consists of seven pre-
genital segments, two genital segments (VIII and IX) (fig. 44)
and a postgenital region, the anal tube or proctiger. In the male,
there are eight pregenital segments the last of which is cylindrical
and partially encloses the ninth (fig. 48). The latter, the genital
segment, forms the capsule which bears the external genitalia. As
in the female, the postgenital area is an anal tube. The female
abdomen is both longer and wider than that of the male (figs. 42,
43). The almost complete sclerotization of the adult abdomen
prevents variation in total body length which is characteristic of the
nymphs. This, and other aspects in the differentiation of the region,
especially of the genital segments, will be considered below (pp.
600-605).
Pregenital Region (Figs. 41, 42, 43)
In both sexes, the first abdominal segment is reduced to a tergal
plate. Its rounded anterior end fits closely behind the metapost-
notum; in winged individuals the small third phragma arises be-
tween them. Ventrally the first segment is indistinguishable from
the metathorax and the rest of the abdomen.
The ventral fusion of the definitive sterna of the pterothorax and
600 Tue Universrtry SCIENCE BULLETIN
abdomen makes flexion of the body as a whole impossible. Varia-
tion in the diameter of the abdomen is possible because of the
arrangement of the dorsal sclerites. In the typical pregenital seg-
ments (II through VII) these consist of a pair of lateral plates
(paratergites or laterotergites and segmentally arranged medial
tergites. Each of the latter is further subdivided into a broad-
ridged median, and a pair of narrow, flat lateral sclerites. The
fused laterotergites of each side form long dorsolateral plates which
taper to a point anteriorly and posteriorly. The junctions of the
sclerites on the dorsum are membranous and allow for variation
in size of the abdomen. In an unexpanded abdomen, the fused
laterotergites cover the lateral part of the medial tergite of each
segment. In a distended abdomen, they are raised (fig. 43).
The ventral part of the abdomen is marked laterally by a pig-
mented band (fig. 41). The spiracles, a pair on each segment from
II through VII, lie between this line and the margin of the fused
laterotergites. Segments I and VIII, which in the nymphs bear
spiracles (see p. 607), lack them in the adult. The spiracles of
the second segment are near the metepisterna and are more dorsal
than the others. The mid-ventral line is very slightly carinate.
The intersegmental lines are indicated on either side by pigmented
lines. In the males, a pair of ridges project from the sternum of
segment VII (fig. 41).
The pregenital region of the mature adult lacks intersegmental
longitudinal muscles. In fifth instar nymphs and in very recently
molted adults, five or six heavy muscle fibers run from one inter-
segmental ridge to another on either side of the middorsal and
mid-ventral lines. In both nymphs and adults, each abdominal
segment has a series of straplike dorsoventral muscles which at-
tach to the laterotergites dorsally and ventrally to the sternum at
the level of the pigmented band.
Female External Genitalia (Figs. 44, 45, 46, 47)
The terminal segments of the female hydrometrid are covered
dorsally by the eighth tergum and ventrally and laterally by the
first valvifers. The first valvifers hinge on the lateral margins of
the triangular eighth tergite (fig. 46) and extend ventrally nearly
to the mid-line (fig. 45). The ventrolateral portions of these
sclerites are partially telescoped into the seventh segment, which
extends back to form a subgenital plate. The proctiger, bearing
the anal opening, and the tips of the second valvulae project
posteriorly between the first valvifers (fig. 46).
HypROMETRA MARTINI KiRKALDY 601
Figure 47 is a diagram of the region with the left anterior valvifer
and the left side of the seventh segment removed to expose the
other structures. The retractor muscles of the valvifer which arise
on the seventh sternum and the membranous lining of this valvifer
are also removed, ‘The first valvula hinges on the inner surface
of the first valvifer. The vagina passes between the first valvulae
which nearly surround the wide vulva. The second valvifers are
dorsal and medial to the first. Together they form an egg guide
rather than a true ovipositor. The second valvulae, or gona-
pophyses, either are lacking or are incorporated in the valvifers.
Two pairs of muscles originate on the seventh sternum. One
pair attaches to the anterior margin of the first valvifer; the other
inserts by a long apodeme on the first valvula. The protractors
of the first valvulae arise on the eighth tergum and attach dorsally.
The two muscles of the second valvifer arise on the lateral margin
of the small ninth tergum. The anal muscle takes its origin on
the seventh tergum and inserts by a slender apodeme which extends
under the ninth tergum to the posterior part of the proctiger.
Male External Genitalia (Figs. 48, 49, 50, 51, 52)
In the male hydrometrid, the membrane which attaches the
eighth abdominal segment is extensible enough to allow flexion
of approximately 60° (fig. 48). Its movement is produced by two
pairs of muscles which arise on the seventh segment, one laterally
and one ventrally. The ninth segment, the genital capsule, is par-
tially telescoped into the eighth. The connecting membrane here
too allows for movement; the capsule can twist laterally about 90°.
Two pairs of intersegmental muscles, originating laterally and ven-
trally on the eighth segment, insert on it.
The outer surface of the entire capsule is rigid but the dorsal
margin, and particularly its lateral angles are extremely heavily
sclerotized. The portions usually enclosed by the eighth segment
are lighter in color and less heavily sclerotized. The ventral wall of
the genital chamber, formed by the concave inner lining of the
capsule, is lightly sclerotized.
The proctiger, or anal tube, projects from the basal part of the
ninth segment and forms a partial cover for the genital chamber
(fig. 50). The large basal plate which articulates with the capsule
posteriorly is nearly concealed (fig. 51). This plate is a sclerotized
ring with a median dorsal projection. The tubular phallosoma or
phallobase extends from it and the genital claspers (parameres,
602 Ture University SCIENCE BULLETIN
harpagones, gonoforceps, genitalhaken ) articulate with it laterally.
The aedeagus is telescoped within the phallobase.
When the genital apparatus is evaginated, the phallobase and
the aedeagus extend perpendicular to the capsule. The lightly
sclerotized valves of the aedeagus are imbedded in its ventral wall.
The tip of the aedeagus is elongate (fig. 52). The claspers are
paddle-shaped and each bears four or five setae on its anterior
margin (fig. 49).
The musculature of the genital capsule is shown in figure 49.
Two small muscles, one of which arises anteriorly and the other
posteriorly on the genital capsule, insert on the base of each clasper.
Protraction and retraction of the phallobase are caused by two
pairs of muscles arising anteriorly and laterally on the genital
capsule and inserting on the basal plate. The protractors are
broad flat muscles which originate on the ventral part of the capsule,
extend under the lining of the genital chamber and attach broadly
to the medial part of the basal plate. The retractors are fan-
shaped muscles which arise on the lateral walls of the capsule and
insert by a tendon on the lateral projections of the plate.
Discussion
Work on the hemipterous abdomen has been concerned especially
with the genitalia. Some studies have included the number of
abdominal segments and others have dealt with patterns of sclero-
tization of this region. The most detailed paper considering all
the aspects of the abdomen and the genitalia is that of Larsén
(1938).
Typically, the first abdominal segment of hemipterans is re-
duced as it is in Hydrometra (Larsén, 1938). In the adult the
first tergum is separated from the second by a faint line. Also,
the presence of eight pregenital segments, the second through the
seventh bearing spiracles and the others lacking them, is typical
of this order (Snodgrass, 1935). In adult hemipterans, the post-
genital region always consists of a small sclerotized tube which is
considered to represent the fused tenth and eleventh abdominal
segments (Snodgrass, 1935; Larsén, 1938). This region has been
referred to as the proctiger (Snodgrass, 1935), Analkonus (Larsén,
1988), anal lid (Ekblom, 1926), Analkegel (Weber, 1930). Eleven
abdominal segments develop embryologically in Naucoris, Nepa
and Notonecta (Heymons, 1899) and in Pyrrhocoris (Seidel, 1924).
The embryological development of Hydrometra is not known but
the nymphs as well as the adults have only ten distinct segments.
HypROMETRA MARTINI KIRKALDY 603
The arrangement of the dorsal and ventral intersegmental mus-
cles of Hydrometra shows clearly the nature of the primary seg-
mentation of the pregenital region. Although the loss of these
muscles is not reported in hemipterans it correlates with the loss
of flexibility, since these muscles, essential for the final molt, have
no function in the adult insect.
The lateral margin of each abdominal segment in many hemip-
terans is differentiated into a laterosternite as well as a latero-
tergite (Snodgrass, 1935; Larsén, 1988). These lateral sclerites
are considered together as the connexivum (Torre-Bueno, 1937),
a structure which is frequently mentioned in taxonomic descrip-
tions (Blatchley, 1926; Hungerford and Evans, 1984). Only the
laterotergites are formed in Hydrometra, and in most species of the
genus, each of these is independent of the others. However, in a
number of species, including H. ambulator Stal, H. albolineata
Scott, H. butleri H. & E. and H. greeni H. & E., which Hungerford
and Evans (1934) figured in dorsal view, as well as in H. martini,
the laterotergites of each side form fused lateral plates.
According to Snodgrass (1935), the spiracles of the insect ab-
domen lie in the tergum. Their location in Hydrometra indicates
the incorporation of a part of the tergum into the definitive sternum,
which Snodgrass illustrated as a variation in the pattern of abdom-
inal sclerotization. The origin of the dorsoventral abdominal
muscles on the wall of the definitive sternum ventral to the spiracles
shows that this region is a part of the sternum. If any pleural area
is present, it must, therefore, be incorporated in the narrow band
between the spiracles and this line of muscle insertion.
By far the most thorough comparative work on hemipterous
genitalia is that by Larsén (1938) mentioned previously, who re-
ported on the reproductive systems of several aquatic species.
The earlier papers of Ekblom (1926, 1980) described the male and
female genitalia as well as head structures in a variety of hemip-
terans, including Hydrometra stagnorum. Comparisons of the
male external genitalia in a large number of species were figured
and discussed by Singh-Pruthi (1925). Snodgrass: (1933) included
descriptions of female genitalia of a coreid, Anasa tristis, and a
saldid, Pentacora ligata.
According to Larsén (1938), the female genitalia of Hemiptera
typically include two pairs of valvifers (gonapophysistrager ), the
anterior bearing the first valvulae (gonapophyses) associated with
the eighth segment and the posterior, associated with the ninth
segment, bearing the medial (second) and lateral (third) valvulae
604 Tue Universtry SCiENCE BULLETIN
(gonapophyses). The lateral gonopophyses do not take part in the
formation of ovipositors in Hemiptera. Snodgrass (1933) and
Michener (1944) homologized the third valvulae with the gono-
styli of primative insects. Larsén (1938) found that in typical
Hemiptera the muscles of the first valvifer include one from the
seventh sternum and two others from the eighth tergum; those
of the second valvifer arise on the ninth tergum. In the corixids,
the only family which he considered in which the female geni-
talia are greatly reduced, the valvulae are small and incorporated
in the valvifers. This may be true also in Hydrometra. The re-
lationships of the sclerites correspond with the description by Snod-
grass (1933) of hemipterans.
Ekblom (1926) found that although the ovipositor in Hydro-
metra was greatly reduced, it resembled those of Gerris and Velia.
There are other minor differences between H. stagnorum as de-
scribed by Ekblom and H. martini. In the former, the ninth tergum
and proctiger are larger and the sternum of the seventh segment
extends much farther posteriorly.
The origins and insertions of the various muscles help to estab-
lish the homologies of the sclerotized parts. These have been
investigated by Snodgrass (1933) and Larsén (1938). The muscles
of the ninth segment are, typically, as they are in Hydrometra, two
pairs arising on the tergum and inserting on the second valvifer.
Usually two ventral muscles which originate on the seventh ster-
pum, insert on the first valvifer. In Hydrometra, only one was
found both in this study and in that of Ekblom (1926). The posi-
tion of the dorsal muscle, arising on the eighth tergum and inserting
on the dorsal part of the first valvula is typical.
The hemipteran male genitalia consistently include a genital
capsule, formed from the ninth segment, which bears the intro-
mittent organ and a pair of articulated appendages (Snodgrass,
1935; Larsén, 1938). The species studied by Ekblom (1926, 1930)
and Larsén: (1935) have intromittent organs consisting of a sclero-
tized proximal portion” (phallobase or phallosoma) and a rather
membranous distal portion (aedeagus). The aedeagus is extremely
variable in shape; in H. stagnorum it is long and tapering (Singh-
Pruthi, 1925).
In Hemiptera, the phallobase attaches to the basal plate. The
retractor and protractor muscles, which arise on the genital capsule,
insert on this plate rather than on the phallobase or the aedeagus.
Both the position and the musculature of the genital claspers in
Hydrometra are of the typical hemipterous pattern.
HyYpROMETRA MARTINI KiRKALDY 605
Although the male external genitalia in Hemiptera are consistent
in their form, the homologies of the various structures are not yet
clearly understood. Snodgrass stated that “concerning the origin
of the insect phallus, we can say only that the facts at present known
about the development of the organs are not sufficient to warrant
any definite statement, but that they appear to favor the view that
the phallus is an independent genital structure” (1941, p. 8). How-
ever, Michener (1944) was able to work out the homologies of
male and female structures by using gynandromorphs and intersexes
of a number of species of Hymenoptera and Orthoptera. Assuming
that these homologies apply to the Hemiptera, the basal plate repre-
sents the fused proximal portions of the gonopophyses and the
genital claspers represent either the gonocoxite alone or a fusion
of the gonocoxite and the gonostylus.
An interesting report by Wygodzinsky (1947) suggested that the
male claspers of the cryptostemmatid, Trichotonannus, are serially
homologous with ventrolateral, spiracle-bearing extensions (which
he termed “parasternites” ) found on abdominal segments seven and
eight. Wygodzinsky proposed the theory that this is a primitive
condition retained by the cryptostemmatids which he believed to
be more closely related than most other families to generalized
Hemiptera.
INTERNAL ANATOMY
The Digestive System (Figs. 53, 54)
The pharynx of Hydrometra martini is readily identified in cross
section by its characteristic U-shape and dilator muscles. It ex-
tends nearly to the posterior margin of the head. Here the di-
gestive tube is surrounded by nervous tissue and changes to the
esophageal pattern, which is round in cross section. In the pro-
thorax, the alimentary canal emerges from the circumesophageal
ring and continues posteriorly, margined on either side by the
principal salivary glands. These are composite structures with
lobules of four types (fig. 53), Those of the most anterior group
are transparent; small and vesicular. The others are translucent
and of variable shapes; one group is elongate, the next, ovate. The
posterior part of the gland is a single large, round acinus. The ac-
cessory gland consists of a median, a lateral and a posterior lobe,
each of which is elongate and thin-walled. A duct connects this
gland with that of the primary gland and a second duct leads an-
teriorly to join that from the opposite side. The common duct
extends anteriorly just ventral to the digestive tube.
606 Tue Universiry SCIENCE BULLETIN
At the level of the mesothorax, the esophagus widens to form the
broad anterior part of the ventriculus, which extends back to the
fourth abdominal segment (fig. 54). The next section of the
ventriculus, a narrow tube, continues posteriorly to the middle of
the fifth segment, loops back to the level of the fourth segment
and then turns posteriorly again. At the level of the sixth abdominal
segment, the ventriculus enters the thick-walled intestine. The
Malpighian tubules, two on each side, enter the alimentary canal
at this level. These twisted, white tubules extend into the fifth
and seventh segments.
The intestine continues to the posterior part of the seventh ab-
dominal segment, where it joins the rectum, a short, thin-walled
tube. A large diverticulum, which is an evagination of the ventral
wall of the rectum, extends under the intestine. The rectum passes
through the proctiger and opens to the exterior at its tip.
The Female Reproductive System (Fig. 55)
The ovaries of Hydrometra martini, consisting of seven ovarioles
or egg tubes, extend from the thorax through the sixth abdominal
segment (fig. 55). Each egg-tube has an anterior terminal fila-
ment, a group of undifferentiated cells and a series of eggs of which
the most posterior is the most nearly mature.
The combined terminal filaments join dorsally between the di-
gestive tube and the metathoracic leg muscles and hold the ovaries
in place. The ovarioles of each ovary open into a short oviduct
which joins with that of the other side to form the common oviduct.
The wall of the common oviduct is thickened anteriorly and leads
posteriorly to the vagina. The seminal receptacle, which also enters
the anterior part of the vagina is made of two parts—a slightly
twisted duct and a flat gland which contains long, coiled and very
slender tubules. Posterior to the base of the seminal receptacle, the
vagina is enlarged to form a muscular bursa copulatrix. On its
dorsal surface is a heavily sclerotized evagination. The vaginal
opening, the vulva, lies between the valvifers of the eighth and
ninth abdominal segments.
Recently molted or overwintering females have inactive ovaries
which are small and compact. The females which are reproduc-
tively active may have tremendously enlarged ovaries with as many
as seven mature or nearly mature eggs. Those females which have
mated may have masses of sperm in both the dorsal evagination of
the bursa and the seminal receptacle.
HyprROMETRA MARTINI KIRKALDY 607
The Male Reproductive System (Fig. 56)
The internal reproductive organs of the male are simple (fig.
56). The elongate testes of the sexually active male extend from
the first through the fifth abdominal segments. In immature and
in over-wintering individuals these may be less than half the size
of the active testes. A slender duct, the vas deferens, leads from
the anterior end of each testis along its median margin to the
seminal vesicle. These vesicles have rather muscular walls and
large lumina. In living males which are dissected, masses of sperm
are moved back and forth in the vesicles by contractions of the
walls. The narrow duct of each seminal vesicle joins that of the
other side to form a common ejaculatory duct which enters the
genital capsule. There are no accessory glands.
Organs of Conservation, Respiration and Distribution
Layers of cells, many of which are fat-ladened, lie between the
exoskeleton and the internal organs. In many individuals, espe-
cially those dissected in the early spring, these cells are bright
green, while in other animals, they are white. The ventral and
lateral walls of the abdomen are lined with a continuous double-
layered sheet of this tissue through which numerous tracheal tubes
spread. A double layer of cells spreads under the tergal area of
the abdomen and both tracheae and a part of the dorsal blood
vessel are imbedded in it.
The positions of the spiracles were described above (see p. 600
and fig. 41). Of the eight pairs, only the first has a closing appa-
ratus. The short atrium lies between the external opening and a
valve consisting of a process on one side which closes against the
opposite wall. The other spiracles open directly into spiracular
trunks. The short spiracular trunk gives off three main tracheae:
one which sends branches ventrad and obliquely caudad, another
which supplies the viscera, and the third which goes to the dorsal
layer of cells, Branches of the dorsal tracheae, which join corre-
sponding tubes of the adjacent body segments, form a dorsolateral
trunk. These large trunks extend anteriorly and posteriorly to the
body segments which lack spiracles. Branches from them supply
the various tissues. No vessels were observed to connect the
tracheae of the left and right sides.
The dorsal blood-vessel extends from the seventh abdominal
segment to the level of the deutocerebrum. In the abdomen, it
lies in the dorsal diaphragm. In the thorax and head it parallels
the anterior portions of the ventriculus and the esophagus. There
608 Tue Universiry SCIENCE BULLETIN
is no obvious variation in diameter in the anterior and posterior
parts of the vessel. The fine structure of the vessel was not studied;
but in stained whole amounts of the dorsal body wall and blood
vessel, two cell layers are evident. The inner layer consists of flat
epithelial cells with flattened nuclei and nongranular cytoplasm.
The cells of the outer epithelium are cuboidal; the nuclei are round
with large nucleoli and the cytoplasm is granular. Many of the
surrounding cells have large fat droplets. Others are smaller with
centrally placed nuclei and granular cytoplasm.
Nervous System (Fig. 13)
The ganglia of the central nervous system are concentrated in
the posterior part of the head capsule and the prothorax. Their
fusion is nearly complete but there are indications of the original
ganglia in both dissected and sectioned material. The optic nerves
extend from the protocerebral ganglia. These and the deutocere-
brum are not contiguous medially since the pharynx with its dilator
muscles lies between them. The dorsal blood-vessel opens between
the bases of the deutocerebral lobes. The large tritocerebral ganglia
are fused and, together with the circumesophageal connectives and
the subesophageal ganglia, make a heavy ring of nervous tissue
through which the esophagus and the dorsal blood vessel pass.
In the thorax the esophagus emerges from this concentration of
nervous tissue and continues posteriorly dorsal to it. The thoracic
concentration consists of four ganglia. Since nerves of the first,
second and third ganglia innervate the legs, these are clearly tho-
racic. The last ganglion gives off two pairs of nerves, one directed
obliquely and the other posteriorly, and represents the fused
abdominal ganglia.
Discussion
In general, the structure of Hydrometra seems to be similar to
that of other hemipterans described in the literature. Among
the studies on the internal anatomy of the group, that of Dufour
(1833) remains the most comprehensive. Weber (1930) and more
recently Poisson (1951) have brought together information on
morphology in their general works on the Hemiptera. Other than
the papers mentioned earlier on the structure of the head, thorax
and external genitalia, only two mention original work on the
anatomy of Hydrometra, One by Gross (1901) concerns the num-
ber of ovarioles in the female H. stagnorum and the other by Mam-
men (1912) deals with the spiracles in the same species. Hydro-
metra martini will be compared here for the most part with Gerris,
which has been studied by numerous morphologists.
HypROMETRA ‘MARTINI KIRKALDY 609
The digestive system of Hydrometra is very similar to that of
Gerris canalium as described by Dufour (1838). In Gerris, the
ventriculus is figured as being constricted anteriorly and having a
prominent pouch just before its entrance into the intestine; in
Hydrometra, it is a simple sac. Both kinds are figured for various
hemipterans by Elson (1937) who also found forms in which the
extremely long ventriculus had four subdivisions and others in
which there were numerous gastric caeca. In this study, in which
he compared the length of the alimentary canal with the total body
length of hemipterans with different feeding habits, Elson found
that the predatory forms, Belostoma and Notonecta, had digestive
tracts three times the body length, the phytosuccivorous forms,
twice the body length and the algophagous forms, one and a half
times the body length. In Gerris, as well as in Hydrometra, the
alimentary canal is about one and a half times the body length
(Dufour, 1833).
Dufour showed three pairs of salivary ducts entering the head
in Gerris. In Hydrometra, as in the coreid, corixid and belostomatid
examined by Elson (1987), a single duct is present on each side.
Dufour mentioned the possibility that the accessory salivary glands
serve as reservoirs, Certainly their thin walls and large lumina
have the appearance of storage rather than of secretory activity.
The posterior part of the digestive tract, in all of the described
Hemiptera consists of a short intestine into which open four Mal-
pighian tubules and a thin-walled rectum with a large ventral
diverticulum (cul-de-sac or sac stercorale) (Elson, 1987). Ac-
cording to Poisson (1951) this structure may serve as a hydrostatic
or a respiratory organ in the aquatic hemipterans. Its function in
other insects is not known.
The internal organs of reproduction of Hydrometra have not
been described except that Gross (1901) reported that each ovary
of H, stagnorum consists of four ovarioles. This is the minimum
number found in hemipterans, whereas seven, the number found
in H. martini, is the maximum (Weber, 1931). Besides the differ-
ences in numbers of ovarioles, variations in the female pattern
seem to involve the vaginal region. Both Dufour (1833) and
Wilke (1908) included figures of the female Gerris but neither
showed this area. Of the aquatic hemipterans considered by
Larsén (1938) only in Micronecta minutissima is the vaginal cuticule
modified as it is in Hydrometra martini to form a chamber in which
sperm can be stored. In both species, sperm are found here as
well as in the seminal receptacle.
610 Tue Untversiry ScreNCE BULLETIN
The male reproductive system of Hydrometra is similar to that
of Gerris lacustris, as described by Wilke (1908) and of Velia
currens as described by Dufour (1833). In Gerris, the testes
are U-shaped; the ducts and the seminal vesicles are broader than
those of Hydrometra. In Velia, the testes are shorter and the
vasa diferentia emerge from nearly the middle rather than at the
anterior end of the gland. In all three species, the ejaculatory
duct is straight and lacks accessory glands.
The tracheae of the abdominal segments of Hydrometra follow
the general pattern described by Snodgrass (1935), The struc-
ture of the spiracles, the first pair having a closing apparatus and
the others lacking it, was found in both Gerris and Hydrometra
stagnorum by Mammen (1912). Dufour (1833) commented on
the sparcity of tracheal tubes in the gerrids which he examined
and correlated this with their low rate of activity and their less
frequent flight. Dufour noted also the relationship of the tracheae
with the tissues under the exoskeleton which serve in fat storage.
The dorsal blood-vessel is usually divided into an abdominal
region which bears ostia and is heavier walled than the anterior
part, or aorta (Snodgrass, 1935). In the stink bug, Nezara, there
are three pairs of ostia in the abdominal section (Malouf, 1933).
These structures were not seen in Hydrometra.
The comparative study of the nervous systems of Hemipterans
published by Brandt in 1878 continues to be the most compre-
hensive work in the field. The nervous system in this group is
characterized by a concentration of ganglia. Various combina-
tions of ganglia occur in the different species. In Gerris, as in
Hydrometra, the subesophageal ganglia are connected broadly with
both the circumesophageal commisures and the combined thoracic
and abdominal ganglia. However, in Gerris the eyes are relatively
close to the central nervous system and the optic nerves are cor-
respondingly short. In other species, the ganglia may be fused in
other patterns; there may be two or three concentrations joined by
relatively long connectives.
BroLocy
Hydrometra martini was collected in and near South Hadley,
Massachusetts from 1948 through the spring of 1955 and in the
vicinity of Douglas Lake, Cheboygan County, Michigan in the
summers of 1948, 1949, 1951 through 1954. In both areas, it was
abundant in pasture ponds, in small Jakes and in other still or slow-
moving waters having plant-lined shores supporting abundant ani-
mal life.
HyDROMETRA MARTINI KiRKALDY 611
Descriptions of Habitats
In the South Hadley area, hydrometrids were found most readily
in and on the shores of spring-fed pasture ponds at Lithia Springs
in South Hadley and at Whately Glen in the town of Whately. They
were also taken at Upper and Lower Lakes on the Mount Holyoke
College campus, Paradise Lake in South Hadley, Aldrich Lake in
Granby and the Old Mill pond in Hadley.
The Whately Glen pond, which is approximately 40 m. long and
30 m. wide, is fed throughout the year by springs at its northeast
end. On the north and west sides of the pond, the banks slope
gradually. The bottom, which is covered here with a thick floc-
culent ooze, becomes sandy near the southern end where the outlet
has been dammed. Cattails (Typha latifolia) are the principal
vegetation of the spring-fed area. Bur-reeds (Sparganium amer-
icanum) emerge from the bottom mud and through the summer
and fall, a thick scum of algae fills the spaces between their bases
and lies over the decaying leaves. Myriophyllum is rooted in the
deeper water.
From late spring to early fall the shore and the matted vegetation
support a varied population of animals. Among them are col-
lembolans, gerrids, lycosid spiders, mesoveliids, hebrids, and Micro-
velia, as well as hydrometrids. Large numbers of entomostracans,
hydrachnids and dipterous larvae swim through the shallow water.
Dragonfly and damselfly naiads, corixids, notonectids, naucorids,
and larval and adult beetles dart through the open water or crawl
over the submerged vegetation. Larger animals, Rana clamitans,
Triturus viridescens and Chrysemys picta also live in the pond.
In the autumn, the leaves of the oak and maple trees and the
needles of the white pines, which surround the pond, drop into the
water. The bur-reeds turn brown and fall, making a floating raft
before they sink to the bottom. In this period the hydrometrids
migrate to land; gerrids and gyrinids disappear from the surface.
Spotted newts become more abundant as the red efts return to the
water and change to their green and yellow colors. On the land,
arachnids, cicadellids, and staphylinid beetles crawl among the
grass hummocks and dead leaves with the hydrometrids. Until
snow falls in the early winter, marsh treaders are found on the moist
earth in protected places, depressions among clumps of grass, and
under dead leaves. In most years, the snow- and ice-covers last
until the spring rains in late March.
The pasture ponds at Lithia, which is also spring fed, is nearly
612 Tue University SCIENCE BULLETIN
round with a maximum diameter of approximately 10 m. It lies in
a hilly pasture surrounded by Tsuga, Chamaecyparis, Juniperus and
Kalmia. Iris and Typha surround the margin of the pond, and
Vallisneria and Myriophyllum cover its floor. In the early spring
as the ice melts, fairy shrimp, Eubranchipus vernalis and larvae of
the limnophilid caddis fly, Platycentropis, appear. Rana sylvatica
and Ambystoma maculatum come in to lay their eggs. Tadpoles of
R. catesbiana and R. clamitans are present through the year as are
Triturus viridescens adults. Many species of water beetles and a
variety of hemipterans (Ranatra, Notonecta, Belostoma and corix-
ids) are to be found in the water, and gerrids and gyrinid beetles,
on its surface. In late summer, if there has been little rain, the pond
shrinks to about a tenth of its maximum size; the animal population
is concentrated. With the fall rains, the water level rises so that
the pond is well filled before the ice forms. This pond, too, is
covered with snow from late December until March.
In the Douglas Lake area, hydrometrids occur in varying numbers
in many habitats, bog lakes, beach pools, slow moving side waters
of creeks and man-made pools. Among these are Nichol’s, Smith’s
and Bryant’s bogs, Bessey Creek, Nigger Creek, Fontinalis Run, the
beach pools at Sedge Point on Douglas Lake and at Black Lake,
Nelson Lake and the gravel pit pools near Pellston. Collections
and observations were most frequently made at the gravel-pit pool
near Pellston and at Nelson Lake because they were more accessible
and hydrometrids were relatively abundant there.
The gravel-pit pools are four miles north of Pellston on the east
side of highway 81. Irregular in shape and in depth, they occupy
an area approximately 25 m. long and 16 m. wide. They were made
by the removal of gravel to a depth of two to three meters at least
15 years ago. The sources of water are springs on the north side
and drainage from the adjacent area; there is no outlet. The water
level fluctuates greatly. In each of the years in which collections
were made, the pools were well filled during the early summer.
In 1951 and 1952, rains were frequent through the summer and the
water level remained rather high in August. Even in these years,
the shallowest place dried completely. In the drier years, 1948 and
1949, the deepest depressions held only about 0.5 m. of water by
mid-August. Emergent vegetation, consisting mainly of Typha, is
limited to these deeper areas in which decomposing material has
formed a deposit. Chara beds cover the bottom and Juncus with
some Carex grows in the shallow water and along the shores. Clumps
HyDROMETRA MARTINI KIRKALDY 618
of Cornus stolonifera, Salix lucida and Salix subsericea overhang the
north banks. A great variety of aquatic and semiaquatic hemip-
terans, notonectids, corixids, Belostoma, Lethocerus, Ranatra, Ger-
ris, Hebrus, saldids as well as Hydrometra, abound here. Other in-
sects include bettle larvae, soldier fly larvae, damsel fly, dragon fly,
and mayfly naiads. Amphibian tadpoles and metamorphosing frogs
and toads are present in numbers.
Nelson Lake lies on either side of a gravel road separating
sections 15 and 22 in Hebron township (Row 8W, Tier 88W),
Cheboygan County. The lake is a low marsh about 8000 m. long
and 3000 m. wide. Earth was thrown up to make a causeway on
which the road runs. The ditches thus made are considerably
deeper than the rest of the area. In dry years, in 1948 and 1949
for example, the water evaporated from the entire area, except
the roadside ditches, causing concentrations of animals here. The
soil is yellowish sand except for the detritus which accumulates in
the deeper water. Spirea and Salix grow along the road and the
willows extend to the edge of the water. The marsh is covered
with a mixture of Juncus and Scirpus with some Carex and Du-
lichium. Potomageton and Polygonum grow in the deeper water.
The animals here are of the same groups represented at Pellston.
However, Spongilla is common and the amphibious snails, Succinea,
are extremely abundant.
Collecting Methods
Adult hydrometrids were caught easily by hand or with a small
tea-strainer and were transferred to an aspirator. They survived
transportation best either in a dry container or in one which was
lined with paper to absorb excess moisture. The nymphs were
collected with a strainer and an aspirator with damp paper in the
bottom. The nymphs were subject to death by desiccation if they
were left too long in dry containers and by drowning if caught in
condensation of moisture on the glass. They survived best when
kept in containers which were lined with moist paper.
Both in the field and in the laboratory, cannibalism was prevalent
among the nymphs. The harder bodied adults seemed invulnerable.
Rearing Methods
In the laboratory, hydrometrids lived well in aquaria with plants
extending to or above the surface or in dishes partially filled with
wet sand. They lived equally well in covered containers lined with
several layers of moist towelling paper. This method has several
advantages; the eggs could be removed and the nymphs and their
Tue Universrry Science BULLETIN
molted skins were easier to see. These containers could be cleaned
by removing the top layer of paper. The eggs were removed and
placed on moist paper in containers. Because of cannibalism, the
mortality rate was high unless the hatching nymphs were isolated
and reared separately.
Hydrometrid nymphs and adults feed on any recently killed in-
sects. Because of the ease with which they can be handled, fruit
flies having vestigal wings were used to a large extent.
BEHAVIOR OF THE ADULTS
Locomotion
Hydrometrids are equally at home on slowly moving water,
on floating or emergent vegetation, or on moist earth. In warm
weather they are most active near aquatic vegetation but sometimes
individuals venture 4.5 or 6.0 m. on the open water. On cool or
cloudy days, hydrometrids most often seek the shore vegetation,
sometimes climbing 25 or 80 cm. on the cattails or sedges.
Their typical gait is the deliberate walk that has given them
both their common names, marsh treader, and water measurer.
They do not glide or hop as do the gerrids, but a disturbed animal
runs rapidly from danger.
As a hydrometrid walks it typically swings its antennae from side
to side. Both in the laboratory and in the field, the adults and
nymphs characteristically elevate and lower their bodies by the
rapid bending of the legs at the femerotibial junction either when
the animal is moving or stationary.
A frequent activity is the grooming of the appendages. The
insects repeatedly draw their antennae or beak between the setose
tarsi of the front legs or rub their legs against each other. Resting
insects often raise one leg while standing on the others. This may
also hasten evaporation of water from the tarsi.
Complete cessation of activity occurs only during cold weather;
the animal takes a flattened position in which the antennae and
legs are outstretched, the front and middle legs anteriorly and the
hind legs posterolaterally.
Under-water activity is limited to accidental incidents since
hydrometrids never seem to submerge themselves deliberately. An
experimentally immersed adult is lighter than water because of the
air film held by body pubescence. It sucks air into its digestive
system either from this film on the ventral side of the body or, if
its body touches the water surface, by extending the beak through
HypROMETRA MARTINI KIRKALDY 615
the film and drawing in air. The insect then swims or sculls along
the surface to emergent vegetation or the shore and walks out. By
intensified grooming, the insect soon dries its appendages.
Macropterous individuals exhibit the same behavior patterns as
the micropterous forms. However they can fly well. Twice ani-
mals kept in the laboratory flew from their containers. The flight,
which was preceded by fluttering of the wings as the animal walked,
was almost vertical for about a meter. One animal flew to the
window; the other disappeared in a few seconds and was not re-
covered, Flight was not observed in the field.
Feeding
When food is available continuously under laboratory conditions,
the adults begin to feed not sooner than three hours but usually
not longer than five hours after the molt.
In the laboratory, hydrometrids appear willing to eat any sort
of freshly killed or inactivated insects. Small living insects, such as
nymphal hydrometrids are devoured readily. They take plankton
animals and aquatic larvae, mosquito wigglers and blood worms,
for example, spearing them under the surface film. In the field
they were observed to eat adult midges, mosquito wigglers, cladoc-
erans, ostracods and collembolans. Usually a feeding insect was
seen only after it had impaled its victim and was carrying it. On
seven occasions, most of them during the summer of 1950, adult
hydrometrids were observed to spear living aquatic forms (3 cla-
docerans, 2. mosquito larvae and 2 ostracods). No field records were
made of hydrometrids feeding on large insects or of several indi-
viduals sharing a victim.
In seeking food, the insect stalks with its lowered antennae
swaying from side to side, as though they were helpful in locating
the prey. As the animal approaches food, it swings its beak forward
and with a movement of the whole body, impales its victim.
The process of securing and holding the prey with the retrorsely
barbed mandibles and probing out the juices with the maxillary
stylets has been described in the section dealing with function of
the mouth parts (p. 585).
The use of saliva in feeding was brought out in the discussion
of the head structure. The size and degree of differentiation of the
salivary glands and the drop of fluid which can be seen occasionally
as the beak of the hydrometrid touches its prey, indicate the activity
of the glands. In addition, the victim is paralyzed quickly; one
which is attacked and almost immediately released soon dies.
616 Tur University SCIENCE BULLETIN
In the laboratory, the imbibing of water by adults was observed
only when the animals were changed from a dry container to one
with water. Those kept in a damp situation probably obtained
enough with their food.
Overwintering
In Massachusetts the hydrometrid population in late September
and October consists of adults, apparently young, and 5th instar
nymphs. The females are sexually immature; the abdomen is
slender and the ovaries are small. When the temperature ap-
proaches 15°, usually in mid-October, the landward migration
starts. In the warm fall of 1951, this movement was delayed until
October 25th. At the beginning of the migration at Lithia, a few
individuals can be seen walking up the gently sloping north bank.
Within ten days of the onset of migration, all of the individuals
are off the water. Nymphs are rarely found; those taken into the
laboratory at this time molt in one to three days.
Through the winter, when the ground is not snow-covered, adult
hydrometrids are found singly or in small groups on the moist
ground near the ponds at Lithia and at Whately. Usually they are
among the decaying vegetation or in depressions among the grass
roots. In the coldest weather, the animals which are completely
inactive assume the resting position described above (see p. 616).
Their inactivity combined with their earthlike coloration renders
them nearly invisible. If these animals are touched, they arouse
themselves and walk away slowly.
Further evidence of their dormancy is the lack of food in the
digestive tubes of the animals taken after the land migration
has begun.
The emergence from hibernation in the spring occurs at about
the time the surface temperature of the water reaches 15°. The
date differs with the year and with the locality. In 1950, a typical
year, early March was very cold and snowy. On March 19th, when
the air temperature was 13°, the snow at Whately Glen was 0.6 m.
deep on the northwest slope of the pond. Only the spring fed
area at the northeastern part of the pond was free of snow; a few
grass hummocks rose above the snow. ‘The water temperature in
the ice free area was 6° A week later the air temperature had risen
to 17° and much of the snow and ice had melted. The water
temperature at the northeastern end of the pond was 10°. About
20 hydrometrids, which were crawling on the grass roots and on the
water-filled depressions between them, were taken. That they had
eater iamaaiaiiaeenntaetaeietiaiaits. * _semenmeeenaineie
HyDROMETRA MARTINI KiRKALDY 617
not yet started to eat was indicated by the completely empty di-
gestive systems of the five that were killed immediately and dis-
sected. On April Ist, the air temperature was 15° and the water,
13°. Hydrometrids were near the shore but none was on the water.
April fourth was a warm day with the air temperature 21° and the
surface water near the shore, 17°. Hydrometrids were both on the
water and on the vegetation near it. The five insects examined had
not eaten, On April 8th, when the air temperature was 19° and
the water temperature 16°, most of the hydrometrids seen were on
the water; of the ten examined, nine had partly filled digestive tracts.
In the same year, open water appeared later at Lithia. On
March 19th the pond here was completely covered with snow and
ice. On the 25th, a sunny day with an air temperature of 14’, the
ice had melted but the water temperature was only 3° and there
were snow banks on the south and west slopes. No hydrometrids
were found. Ten days later, on April 4th the water temperature
was 15°, the air temperature 21° and dozens of hydrometrids were
stalking on the water. Three of the ten examined had eaten re-
cently.
The earliest spring collection of Hydrometra was March 16,
1949, when two wingless females were found on the water at
Whately. This was a rather warm day with an air temperature
of 17° and the water in the place where the insects were taken
was 13°. However, there were large snow patches on the ground
and most of the pond was ice-covered.
In 1951, warm spring weather was late. On April 3rd, the pond
at Whately was almost completely covered with ice and the snow
cover was between 0.5 and 1.0 m, deep. Hydrometrids were not
collected until April 20th.
Reproduction
The mating of Hydrometra can be observed in the laboratory
or in the field in the spring and summer. Typically, as he ap-
proaches the female, the male protracts the eighth abdominal
segment to lower the genital capsule. He may come toward her
from any direction but usually he approaches from the side and,
swinging his body parallel to hers, the male mounts the female
and holds her prothorax with one or both of his forelegs. The
seventh abdominal processes rest on the eighth tergite of the
female; the other pregenital segments do not touch. The eighth
segment of the male projects obliquely ventrad and the ninth
segment (genital capsule) is directly under the ninth segment of
618 Tue Universtry SCIENCE BULLETIN
the female. The phallobase extends to the vulva while the aedeagus
with its process are within the female reproductive tract. The
short parameres do not make contact with the female.
During copulation, the female walks elevating and depressing
her body, carrying the male on her back. Not infrequently she
feeds in the process. The male remains mounted on the female
for as long as 15 minutes but actual mating seems to last not
more than six minutes. After the male releases the female, she may
push him aside with her middle and hind legs.
Males were never observed to copulate with a second female
immediately after a successful mating. However, if the male
failed to mate with the first female he approached, he usually
sought another.
The preoviposition time in Hydrometra martini is variable;
in the first spring generation, this period lasted between nine and
seventeen days for ten animals brought into the laboratory on
June 29, 1949, as fifth instar nymphs. As these animals molted
to the adult stage, each was placed in a 3-inch stender dish with
a sexually mature male. Each dish was examined every other
day; the eggs were counted and removed. In the observation
period, five males died and were replaced. There was no ap-
parent pattern of numbers of eggs laid. The greatest number laid
by any individual in the two-day period was 14 while the average
number laid in two days varied between 3.5 and 6.2. The longest-
lived individual in this series continued to lay eggs until September
5th, and died on September 12th. Between August 15th and
September 9th, she laid 72 more eggs making a total of 161. An
animal which lived until September 2, laid a total of 118 and an-
other laid 121 before she died on September 3rd.
Fifteen females reared from fifth-instar nymphs collected later
in the same summer, matured more rapidly. This group was col-
lected on July 21st and completed the final molt between that
date and July 26th. The average preoviposition time was 8 days,
the minimum 7 days and the maximum 10 days.
In the field, females which undergo their final molts at the end
of the summer or in the early fall seem ordinarily not to lay eggs
until the following spring. However, young females brought into
the laboratory in September and October of 1950 began to lay eggs
after November 20th. The average preoviposition time of six
females collected in September was 69 days with a variation of from
59 to 81 days; that of five females taken in October was 65 days
HypROMETRA MARTINI KiRKALDY 619
with a range of 58 to 78 days. Females collected from hibernacula
in mid-December of the same year laid eggs about four weeks later.
Hydrometrids collected in the early spring mature more rapidly
than those brought into the laboratory in the fall and winter.
A group of five animals taken at Whately, Massachusetts on March
26, laid their first eggs between April 18th and April 18th; the
average interval between collection and egg laying was 19 days.
Ten others taken at Whately two weeks later began laying eggs
within 5 to 13 days with an average of about 10 days. All of the
females brought in after April 25th of that year laid eggs within a
day or two of their collection.
Because observations of the previous year had indicated that
Hydrometra mates before going into hibernation, all of the fe-
males collected for this study in 1950 were isolated. Since the
eggs of all but two females (these were of the group collected on
March 26th) were fertile, copulation must occur normally in the
early fall when the females have small ovaries with no. well-
developed eggs.
The egg laying process is an extremely simple one. In the
laboratory, a female walks about the container slowly, dragging
or touching her abdomen to the floor. Then she stops and raises
the anterior part of her body. Her front legs are almost straight,
her middle legs extend laterad and her hind legs are bent at the
femorotibial junction. After the female has tapped the support-
ing surface with her abdomen for two or three minutes, a small drop
of fluid is expelled from the genital opening.
The base of the egg follows the fluid, and as the egg emerges,
the female lifts her body. In so doing, she draws the egg per-
pendicular to the support. The whole process may be completed
in four to five minutes although it may take as long as 15 minutes.
If the hydrometrid is disturbed in oviposition, she moves to an-
other place and starts the process anew. Some may choose a ver-
tical instead of a horizontal support. In this case, the female with
her head downward, holds her body parallel to this surface.
Discussion
The measured gait of Hydrometra was noted by Arrow (1895 )
and by numerous later authors (Kirkaldy, 1899; Martin, 1900;
Bueno, 1905; Weber, 1930; Jordan, 1931). That the movement of
the legs is principally on a horizontal plane was mentioned by
Weber (1930). This is correlated with the articulation of the
coxae deep within the supracoxal cavities which was mentioned
Tue Universiry ScreENCE BULLETIN
previously (pp. 614-15). The front and middle legs can swing
in a wide arc but the hind legs with its coxal process articulating
with the posterior part of the supracoxal wall, is limited in its for-
ward movement.
The elevation and depression of the body by a curious swinging
movement. which is characteristic of hydrometrids was reported
by Jordan (1931). No function can be ascribed to this behavior
pattern, but the equally characteristic grooming movements, which
Jordan also noted, seem to be important in the animals in keeping
their appendages dry. Both Jordan and Weber believed that waxy
or oily secretions distributed by this action, as well as the fine setae,
help in making the body surface hydrofuge. The distribution of
weight of this slender-bodied insect on its six widely separated legs
and the hydrofuge nature of the exoskeleton (especially of the tarsi)
are sufficient adaptations to enable it to walk on the surface film.
Flight in gerrids in the laboratory has been reported by Wilke
(1908), who found that although recently caught gerrids did not
fly, those which had been kept in tight containers for about ten
hours flew away as soon as they were released. From observations
on gerrids in the field, he concluded that they flew only at night
since pools, in which there had been none in the evening, would
have a large population the next morning. Tevrovsky (1920), also
reporting on gerrids, observed that flight also occurs during the day.
Flight in Hydrometra is not reported in the literature. Beamer
(1949), however, found that Hydrometra as well as many other
water bugs were attracted to lights. A South American species,
Hydrometra mensor, was described by White from a specimen
labeled “Manaos on board at light VIII, 1875” (Hungerford and
Evans, 1934).
An additional record of attraction to light is that of Hungerford
and Spangler (1952) who collected two first-instar and one fourth-
instar nymphs of H. martini in a light trap set at the surface of
Nichol’s Bog, Cheboygan County, Michigan, on the night of August
II, 1952.
Other investigators have mentioned many kinds of insects which
serve as food for Hydrometra. Hydrometra stagnorum has been
reported to feed on springtails (Arrow, 1895) and on dead or
nearly dead insects such as dipterans and mayflies (Ekblom, 1926),
and dead flies (Jordan, 1931). Martin (1900) and Bueno (1905)
emphasized the use of aerial insects caught in the surface film as
food by H. martini, while Hungerford (1917, 1920) observed that
Hydrometra eats a great variety of animals in the field. These in-
HypROMETRA MARTINI KiRKALDY 621
clude: mosquito wigglers and pupae, emerging midges, nymphals
corixids, ostracods and small terrestrial insects. Riley (1918) fed
hydrometrids with entomostracans in the laboratory.
The methods by which Hydrometra obtains these food animals
are typical of the predacious Hemiptera which lack raptorial fore-
legs. The securing and holding of prey with the mandibular stylets
and the probing and sucking by the maxillae were mentioned spe-
cifically by Hungerford (1920) for this genus. The carrying of
small insects on the beak was seen by Arrow (1895) and Ekblom
(1926) as well as by Hungerford (1920). Although more than
one hydrometrid was never seen to feed on a single victim in the
field in the present study, this sharing of an individual food insect
has been cited for both Hydrometra stagnorum (Ekblom, 1926)
and H. martini (Martin, 1900).
Because Hydrometra lacks a separated salivary channel in the
maxillary tube, the use of saliva in the feeding process has been
questioned (Ekblom, 1926; Weber, 1930). However, the evidence
seems overwhelming that saliva is injected by hydrometrids into
their victims and that it is effective in paralyzing them. Bueno
(1905) also thought that living victims were paralyzed by hydro-
metrids, although according to Baptist (1941) the saliva of Hemip-
tera is principally digestive in function and only secondarily toxic.
The hard exoskeletons of the adults seems to render them in-
vulnerable to other insects of the same or larger sizes. This is
probably the advantage of the fusion of body parts and the lack
of thin intersegmental membranes. Cannibalism was never seen
in adults nor were adults observed to be eaten by other animals
in the field although large lycosids and gerrids roam the surface film
with them and Ranatra, Dytiscus and other large carnivorous in-
sects lurk beneath it.
When Hydrometra leaves the water to hibernate, feeding ceases.
Both H. martini and H. stagnorum have long been known to hiber-
nate as adults in the decaying vegetation around the banks of the
waters which they inhabit. This was reported for H. stagnorum
by Wesenburg-Lund (1913), Stichel (1926), Jordan (1931) and
Schumann (1934). Schumann also found a wingless pair in the
woods far from water. Jaczewski (1936) mentioned finding very
few nymphs of H. aegyptia in collections made from September
to March in Egypt and concluded that here, as well as in Central
Europe, most Hydrometra winter as adults. Both Martin (1900)
and Hungerford (1920) saw Hydrometra martini coming out of
622 Tse Untversiry Science BULLETIN
hibernation during the first warm days of spring. Bueno (1905)
found hundreds in May at Staten Island.
Although female hydrometrids are able to lay fertile eggs prior
to mating in the spring, proof that they do so in nature is lacking.
Jordan (1931) observed that H. stagnorum and H. gracilenta mate
at the end of April and early May in Germany; he did not mention
whether egg laying preceded copulation in these species. Martin
(1900) who worked on Hydrometra at Ithaca, New York, collected
females which were laying eggs as early as May Ist, but whether
or not these animals had mated was not reported. Bueno (1905)
bred virgin females with captured males on May 26th, and found
that they had deposited eggs two or three days later.
The first report of mating in Hydrometra is that by Palumba
(1891). According to this account, the female is pursued by as
many as eight males which battle ruthlessly. The victor mates
with the female, but dies a few days later; the female lays her
eggs and then dies. This process of selection, which Palumba be-
lieved to lead to the strengthening of the species, has not been
confirmed.
Both Ekblom (1926) and Jordan (1931) included accounts of
copulation of H, stagnorum. Mating pairs were never seen on the
surface film by Ekblom but Jordan mentioned the depression of
the female by the added weight of the male. They both found that
sometimes, immediately after mating with a female, a male would
mate again with the same or another female. These two authors
agreed in finding the duration of copulation highly variable.
In the northern part of the United States, the egg laying period,
like the mating period, is continuous from April through August.
This is the case in New York (Martin, 1900; Bueno, 1905) and
Kansas (Hungerford, 1920) as well as in Michigan and Massachu-
setts, as shown in the present study. Jordan (1931), working in
Germany, found that the egg-laying periods for H. stagnorum and
H. gracilenta last for at least six weeks in April and May.
The length of the egg-laying period and the number of eggs laid
by a single Hydrometra have been recorded most completely by
Hungerford (1920). One female which had molted to the adult
stage in the laboratory on July 25th, had mated and laid 28 eggs
by August 4th. She had deposited a total of 173 eggs by August
31st, and continued to lay eggs after that date. In part of the period,
she averaged 7.8 eggs per day and on one day, laid 11. This indi-
cates a production of an egg by almost every ovariole each day.
The only reports on numbers of eggs laid by European species are
HypROMETRA MARTINI KirnKALDY 623
those of Jordan (1931) who found that average number of eggs laid
by H. stagnorum was 47 and that a single H. gracilenta deposited
53 eggs.
The simple expulsion of the egg by the female and its attachment
by a sticky disc to a support has been observed by other investi-
gators for several species: H. martini by Martin (1900) and Hun-
gerford (1920), H. stagnorum and H. gracilenta by Jordan (1931)
and H. vittata (probably H. albolineata) by Takahashi (1921).
NUMBER OF GENERATIONS AND LONGEVITY
There seem to be three full generations and a partial fourth
generation of Hydrometra martini in both Michigan and Massachu-
setts, The over-wintering generation begins reproductive activity
in late April or early May in Massachusetts. Some of these indi-
viduals live and lay eggs as late as the second week of July. In
both places, the first nymphs of the spring generation molt to the
adult state about June 20th. In the laboratory and probably in the
field, these females begin to lay eggs in about ten days and live,
laying eggs, until nearly the end of the summer. The first offspring
of this generation, arising from eggs laid about July Ist, are the
summer generation. They complete the nymphal instars and reach
their final molt in three to four weeks. After a preoviposition period
of a week to ten days, the females begin to lay eggs. Eggs of the
summer generation, then appears as early as August Ist and they
are laid continuously until late summer. Some animals arising from
this generation may also reproduce in the same season, and _there-
fore give rise to the fourth generation. Most of them, however,
do not lay eggs until the following spring but they do mate before
going into hibernation.
Because of the long life and extended oviposition period of the
females, members of various generations are sexually active simul-
taneously during the summer. In the laboratory the animals that
lived longest were those of the summer generation, they lived from
late August until the following June or July, having attained an age
of about ten months.
Discussion of Longevity and Number of Generations
The record for the longest life in Hydrometra martini is that re-
ported by Bueno (1905) who found that an individual taken in May
overwintered and lived until the following August. Neither in the
present study nor in others in the literature has a span longer than
from September or October to the following June or July been
reported.
Tue Universiry Science BULLETIN
Bueno (1905) gave an average life cycle of 25 to 85 days and
three to five broods per summer in Staten Island, New York.
Martin (1900) implied that there were several generations in Ithaca,
New York. In Kansas and farther south, the number of generations
may well be more than five. Hungerford (1920) reported an op-
timum life cycle of 15 days from egg to egg although the average in
the summer was about 21 days. With reproductive activity be-
ginning earlier in the year and continuing until fall, there are con-
ceivably as many as eight or nine generations in a season.
Workers in northern Europe (Wesenberg-Lund, 1913; Ekblom,
1926) found only one generation per year of Hydrometra stagnorum
which Ekblom found to reach the adult state at the end of July
but in England according to Arrow (1895) and in Germany accord-
ing to Jordan (1931) there are two broods. Jordan supported his
field observations with laboratory rearings; one generation matured
in late June and the other, the overwintering generation, approxi-
mately seven weeks later.
The number of generations produced in different areas varies
greatly. Temperature probably is the factor responsible for re-
productive activity in Hydrometra.
Tue Eco
Appearance
The spindle-shaped and beautifully sculptured egg of Hydro-
metra martini is comparatively large, about 2.0 mm. long and 0.2
to 0.28 mm. wide. The egg shell or chorion consists of two layers.
The leathery outer exochorion protects the embryo and is respon-
sible for the shape and surface pattern of the egg (fig. 57). The
central portion of the inner layer, or endochorion, is a delicate sheath
which surrounds the embryo proper. From it extends two slender
projections: a spicule leading through the stalk into the basal at-
tachment disk and a delicate tube terminating in the micropyle at
the free end of the egg (fig. 58).
The scuplturing of the exochorion indicates the intricate arrange-
ment of the air spaces within it. Each of the larger divisions, the
basis of which abut on the micropylar tubule, is subdivided into a
myriad of minute alveoli. Their outer surfaces make a diamond
pattern on the exochorion. A cross section of this region shows the
arrangement of these compartments (fig. 60), and the proximal
part of the basal stalk has a similar pattern. Here large air cham-
bers surround the spicule of the endochorion. The distal part of
HypROMETRA MARTINI KIRKALDY 625
the stalk, adjoining the gummy basal disk, consists of simple and
rather transparent air cells which lack internal partitions (fig. 59).
The sculptured pattern of the wider middle area gradually merges
with that of the two ends. In most of the eggs the sculpturing of
that part consists of crests separated by furrows extending from one
end of the central area to the other. In some of the less typical eggs,
transverse ridges connect the longitudinal ones making a more or
less extensive network. The raised surfaces are spongy with a vast
number of minute air spaces. In a few eggs a hexagonal pattern
takes the place of longitudinal grooves.
All of the air-filled spaces in the exochorion make the hydrometrid
egg look opaque and give it excellent flotation. Dry eggs dropped
in water rest on the surface film or float just beneath it. Those
which are submerged, so that some of the air is replaced by water,
soon sink and the outer chorion becomes translucent.
When they are laid, hydrometrid eggs are pearly white. They
darken quickly if dampened either by being attached to wet sur-
faces or by being exposed to moist air. Within five minutes they
turn to a light tan color and fifteen minutes later they are brown.
No further color change occurs during the development of the
embryo. On the other hand, eggs that are completely submerged
in water immediately after they are laid, darken slowly. After 24
hours they are only slightly colored and within the next few days,
they become cream colored.
Eggs of Hydrometra martini vary between 1.8 and 2.2 mm. long
and between 0.20 and 0.22 mm. wide when they are laid. The
length does not change but the width increases notably during
development. During the third day of incubation (at 25°), the
eggs expand to their maximum diameters; some reach 0.28, others
only 0.25 mm. The size of the egg remains constant from this time
until it hatches.
Development
Some features in the development of the embryo can be seen
in living hydrometrid eggs and can be followed only if the eggs
are moist enough to be translucent. Eggs for this study were incu-
bated at 25° in stender dishes lined with moist towelling paper.
The first change that can be seen as the living embryo develops
is the appearance of small red eye spots on the third day after the
egg is laid. There are near the posterior end of the egg where
they remain for four to eight hours (fig. 61). During the next
two hours blastokinesis or inversion of the embryo, occurs and the
21—8378
Tue Universiry SCIENCE BULLETIN
eyes shift to the apical part of the egg (figs. 63 and 64). The eyes
remain in this position, enlarging rather gradually for the next
five days (figs. 65 through 69). The egg burster, a conspicuous
black cross between the eyes, becomes apparent on the eighth day.
On the eighth and ninth days, the margins of the embryonic cuticle
surrounding the labrum begin to darken (figs. 66 and 67). By the
eleventh day, the labrum is clearly outlined and the appendages
of the head and thorax are pigmented enough to be seen (fig. 69).
The antennae and first and second pairs of legs extend posteriorly
and then loop upwards on the embryo’s left side before they con-
tinue backward. The antennae and beak lie ventral to the legs.
The claws of the first and second pairs of legs are close together
posterior to the tips of the antennae. The metathoracic legs extend
caudad almost to the end of the abdomen, and then bend upward
on the opposite side so that the claws are lateral.
Some of the eggs incubated at 25° hatch on the eleventh day,
some on the thirteenth and a few on the fourteenth, but the
highest percentage hatch on the twelfth day of incubation. Of
68 eggs laid on July 22, 1950, by wingless hydrometrids in the lab-
oratory, about 70 per cent hatched on the twelfth day and 17 per
cent on the thirteenth day of incubation. Eleven other eggs laid
on July 22nd failed to hatch. Of these, five did not show any signs
of development and may have been infertile. Three failed to com-
plete blastokinesis, and in each the eyes and egg burster developed
in abnormal positions. Three other embryos, which had appeared
to be normal, turned black without beginning to emerge.
Of the 50 eggs collected in the field on July 28 and July 30, 1950,
only three failed to develop. The others hatched between July 31st
and August 9th.
Hatching
The hatching of a hydrometrid egg follows a fixed pattern. Table
1 is a time schedule of the procedure in a series of five nymphs
which hatched on the morning of August 5, 1949. These eggs of
unwinged adults were laid in the laboratory on July 24th and were
kept on moist towelling paper. The process of eclosion is essen-
tially the same in eggs which are submerged or rest on the surface
film.
The young hydrometrid begins the hatching process by sucking
in the fluid within the egg shell. With the increased pressure from
the enlarging animal, the egg burster moves against the chorion
and splits it. The vertical part of the egg burster is knifelike, hard
HyYDROMETRA MARTINI KIRKALDY 627
Taste 1.—Intervals of time in minutes taken in Eclosion of 5 Hydrometra
martini nymphs on August 5, 1949.
Intervals of time in minutes
Activity after first split of chorion
Thee. DUPSIED OMpOSOCl ee is ey ss hep de le ae
Inner margin of eye exposed............ ee ie ea a
Half of eye exposed. 0. besainior. evn oko ae dR
Eyes entirely exposed ................. 6 6.43 y
WSODEUMIAINOOU Cunt t ree eee 1 ea ee
Thorax entirely exposed .-.............. b18 tk
WORUVING SIU sare acl ce bis os ahi we Ck Ce ee ee a |
Head and thorax out of exuviae.......... 2 See eed
lstileg dreed-or esuvide asain... es ns v 8
Und ang Srd leg Iree; fyi ec ee cee Boe, ees)
Antennae free of exuviae............... Se ea
LiOkS Oh SUDPOM sii austin stk ae ep! oF lpg 7
Abdomen lifted from exuviae ........... O. Syd 2. 8
and slightly raised from the surface of the rest of the embryonic
cuticle. The nymph pushes its head through the slit in the egg
shell until the inner margins of its eyes begin to show. It contin-
ues sucking and swallowing the fluid still surrounding it within the
chorion. Within six minutes of the time the chorion splits, the
eyes are half exposed (fig. 70). Then it is completely quiet for
four to seven minutes. As air replaces the fluid within the chorion,
the continued sucking brings bubbles of air into the digestive sys-
tem. Within the next minute the nymph pushes its head outward,
frees its labrum and thrusts its thorax out of the chorion (figs. 71,
72, 73). It then continues to extend its body from the egg shell by
movements in a dorsoventral plane. As the nymph arches its body,
it lifts its head, holding its antennae and beak rigid (fig. 74).
Straightening the body, it bends its head down (fig. 75), pushes
the thorax out, unfolds and draws out its legs. The tarsal seg-
ments and the tips of the antennae and beak as well as the end
of the abdomen remain in the chorion and support the rest of the
body. With its body perpendicular to the egg shell, the nymph
rests and completes the process of filling its entire abdomen with air.
By muscular contractions that start in the metathorax and pass
forward to the base of the head, the nymph now increases the pres-
sure on the embryonic exuviae and these break along the conspicu-
ous dark sutures of the head (fig. 76). The animal pushes its head
out of the embryonic cuticle rapidly, the egg burster and the mem-
brane attached to it pass backward over the thorax, the bases of
the legs and the abdomen, (figs. 77 and 78) and within three min-
utes frees its legs. During this process, the femora bend at a num-
ber of points as well as at the articulations. After it extricates its
628 Ture Universiry Science BULLETIN
last leg, the nymph pulls the antennae and labium out. The tip of
the abdomen remains for a short time within the molted skin and
within the chorion. This anchor gives support to the nymph while
it struggles to bring its feet into contact with a surface. When the
animal begins to walk, it finally pulls the abdomen free.
The hatching process takes between 20 and 30 minutes. If eclo-
sion is prolonged or if any part of it is out of proper sequence,
the nymph dies. Death may occur at the beginning of hatching
or when the nymph has started to molt its embryonic cuticle. But
the greatest number of fatalities comes as the nymphs free them-
selves from the chorion. The legs are hopelessly caught if they do
not molt before the labium and antennae do.
A hydrometrid hatching from an egg floating on or immediately
under the surface of the water lifts its head above the surface film
so that it swallows air. The chorion, lying parallel to the water,
acts as a platform until the nymph can lower its legs to the surface
film and walk away.
When a nymph emerges from a completely submerged egg,
it sucks in water instead of air to distend the digestive system.
A nymph can live only an hour under water; after that its exo-
skeleton becomes permeable and the tissues become hypotonic.
A nymph which approaches the surface with one side uppermost
cannot lift its body for most of its movement is parallel to the film.
Any nymph which maneuvers in such a way that its dorsum
comes in contact with the surface film quickly raises itself out
of the water. For an instant it lies with its body outstretched,
the first and second pairs of legs extending diagonally forward,
its hind legs stretching parallel to the abdomen. First it lifts its
head and antennae out of the water, next its thorax, then its first
and second pairs of legs and finally its abdomen and hind legs.
Then it dries its appendages by lifting them in the air and by rub-
bing the various parts together.
Discussion of the Egg Stage
The eggs of Hydrometra have been described and illustrated
by a number of workers. The earliest description of a hydrometrid
egg was that published in 1855 by Leuckart whose careful draw-
ing showed the basal shaft with the attachment disc as well as
the micropyle. Some of the information from the literature on the
eggs of three species of Hydrometra is summarized in Table 2.
The eggs of Hydrometra stagnorum are less elongate but other-
wise similar to those of H. martini. Some illustrations in the litera-
HypROMETRA MARTINI KIRKALDY 629
Taste 2.—Summary of information on hydrometrid eggs in the literature
Species Author Wee Width Color
H. stagnorum | Brocher (711)....... ahr Perea grayish, opaque
Jordan (oljia. 4 1.66 0.25 | white when laid;
yellow brown later
SohumanncC@ 3d) a:als ee Cee leeia ss dark brown
H, gracilenta..| Jordan (’81)........ 1.6 0.23 | yellow
H. martini....| Martin (’00)........ BO. le Wi ease bio eel pc
Hungerford (’20)....| 2.07 0.277 | white when laid;
brown later
ture, like that of Leuckart (1885), show a basal shaft only slightly
shorter than that of H. martini, but others (Tevrovsky, 1920; Schu-
mann, 1984) indicate a short base. Brocher (1911) pictured and
described an egg in which the basal air cells are broken and re-
ferred to the egg as being mobile as the clapper of a bell. This
part of his description, as well as his figure showing longitudinal
grooves within an oval outline, were criticized by Jordan (1931).
The illustration of the egg by Jordan shows deep grooves arranged
longitudinally and oval granules at either pole. Schumann (1934)
indicated grooves extending from one end of the egg to the other,
and a little papilla near the micropyle. Tevrovsky (1920) in-
cluded two drawings of the egg of H. stagnorum, one of its ex-
ternal appearance and one as it looked after being cleared in cedar
wood oil. The latter shows the micropylar tube and the basal
shaft, which in this drawing is very short. The egg of H. gracilenta,
which was described by Jordan (1931), was not illustrated.
The drawing of the egg of H. martini by Martin (1900) is well
known because it has been used in textbooks (Comstock, 1936;
Wesenburg-Lund, 1934). As he mentioned in his text, Martin over-
emphasized the hexagonal pattern at the expense of the longitudinal
furrows. Hungerford (1920) showed these clearly in his figure.
His photographs of hydrometrid eggs with sprouting cattail seeds
showed their similarity to one another.
The considerable variation of colors mentioned in the descriptions
of the eggs of each species correlates with the actual differences
in eggs of a large series. Eggs of H. martini, which are grayish
and iridescent when they are dry, become a dark or light brown
when they are wet. That newly laid eggs are white and that they
darken quickly was mentioned by Hungerford (1920) and by
Jordan (1931).
Tre Universiry SciENCE BULLETIN
Poisson (1924) stated that eggs of a variety of water bugs are
white when they are laid and that a secretion of the reproductive
tract is responsible for their darkening. This is certainly true of the
eggs of H. martini. The change in color must be the result of an
interaction of the secreted substance with air and water since all
three are necessary for the color. Eggs dissected from the female
do not darken, those which are submerged as soon as they are laid
darken slowly and those kept dry do not change color at all.
Hydrometra stagnorum, as well as H. martini, lays eggs on any
convenient object. Jordan listed a large number of plants on which
he found eggs and both he and Schumann (1934) described eggs
being laid on the sides of containers in the laboratory. Bueno
(1905) found the aquarium in which he kept two female hydrome-
trids studded with eggs by the end of the summer. Both Martin
(1900) and Hungerford (1919) said that the females placed eggs
on any firm object. Jordan (1931) found the eggs always laid high
above the water surface on dry leaves but he did not discover any
on Nymphaea or on emergent reeds. Brocher (1911) however
collected hydrometrid eggs on floating vegetation and debris; all
of these were covered with water or resting on its surface. Taka-
hashi (1921) working with a Japanese species stated that these
insects deposited their eggs on grass or twigs on the water, but
that the eggs were often found under water.
The length of the developmental period varies considerably.
According to Arrow (1895), eggs of H. stagnorum laid on May 28,
hatched three weeks later. Schumann (1934), working with this
species, found nymphs on June 11th which had hatched from eggs
laid on May 29th. Martin (1900) gave 17 days as the developmental
period for H. martini. According to Hungerford (1920), eggs laid
in May took 23 days to hatch while those laid in July hatched after
only seven days. Bueno (1905) noted that embryonic develop-
ment took nine to ten days in midsummer.
Hungerford (1920) in Kansas in July and Bueno (1905) on
Staten Island, New York in summer, found the incubation period
to be less than 11 days. The temperatures probably were higher
than 25°. The incubation periods of 11 to 13 days were observed
in the present study.
Of the numerous authors who have worked with Hydrometra,
only Hungerford (1920) has discussed the hatching process in
detail. He described the bulging of the head with the eyes and
the egg burster appearing, the pushing of the body at right angles
to the shell followed by the breaking and shedding of the em-
HypROMETRA MARTINI KiRKALDY 631
bryonic cuticle and finally the unfolding of the appendages. He
provided three figureés—two of the emergence of the head and the
other of the exuviae.
Brocher (1911), who referred to the role of the egg burster and
the postnatal molt in H. stagnorum, included a drawing of the
shed cuticular skin caught in the opening of the egg shell. Schumann
(1934) observed a nymph of this species escape from its shell and
mentioned that the process took 5 minutes. His figures are of the
late stages of hatching. In each, the egg is on its side and the
emerging nymph is twisted with its head close to the base of the
egg. With H. martini, this does not occur. Schumann also showed
a torn opening in chorion rather than a simple slit which has been
described by other investigators.
Takahashi (1921) described the hatching of Hydrometra vittata
(probably = H. albolineata) from submerged eggs. This species
seems to be better adapted to life underwater than H. martini for
none of the animals he observed died underwater or while emerging
from it. The nymphs that hatched underwater walked or swam
until they came to rest with their backs against the surface film.
A nymph in this position bent its head and abdomen until it had
lifted its head above the water. Then the thorax, first and second
pairs of legs and finally its hind legs and abdomen were raised.
This procedure is essentially the same as that followed by the
successful nymphs of Hydrometra martini.
Tue ACTIVITIES AND STRUCTURE OF THE NYMPH
Activities of Nymphs
The bright red eyes of the hatching hydrometrid nymph are
conspicuously set off by the paleness and delicate green of its head
and body. Its legs and antennae are transparent. The abdomen
is expanded and through the translucent body wall the anterior
part of the ventriculus can be seen filled with air. At this time the
nymph is 1.5 mm. long. If the nymph is undisturbed, it will remain
close to the egg shell from which it hatched. However, if it is
disturbed, it can easily walk or run. Within 15 minutes after
hatching, the abdomen contracts, forcing the air from the gut of the
abdominal region into that of the second and third thoracic seg-
ments. The contracted abdomen appears a darker green and, by
this time, there is a darkening of the legs and head. The body-
length decreases to about 1.2 mm.
Before the nymph is an hour old, it lowers its rostrum to the moist
substratum to obtain water. The stylets extend beyond the tip of
632 Tue Universitry SctiENCE BULLETIN
the last labial segment and water is pumped up through them by
the muscles of the head. The fluid passes into the pharynx and
through the narrow esophagus into the ventriculus. The first water
that enters the stomach seems to absorb the air there so that after
the animal has been drinking for ten minutes or so, the size of
the air bubble is greatly reduced. Peristaltic contractions of the
wall of the gut, some of which start anteriorly and others, poste-
riorly, churn the contents.
When food is available, the nymph may begin to eat about an
hour and a half after it has hatched. Some nymphs observed in
the laboratory, however, did not feed for as long as 12 hours after
hatching although food was available. When the nymphs begin
to eat, the juices of the prey pour into the ventriculus and mix with
the water there. After 24 hours, the curved posterior ventriculus
which lies in the abdominal segments is distended with dark resid-
ual substances. An animal to which food and water is available
stretches out to about 1.6 mm., nearly its maximum length.
The locomotion and feeding of first instar nymphs, as well as
those of the older nymphs, are essentially those of the adults. They
walk on the surface film, on damp earth or on vegetation, their
antennae swaying from side to side. The nymphs continually
groom their appendages and display the up-and-down movement
characteristic of the adults. The flexibility and longitudinal inter-
segmental muscles of the abdomen make possible its curvature;
nymphs often carry the posterior end of the abdomen arched over
the body.
Submerged nymphs regain the surface rather quickly. Such an
animal swims up so that its back makes contact with the surface
film. By flexion of the head and prothorax, the anterior part of the
body, including the forelegs, breaks through the film. The nymph
then swims until it reaches an emerging leaf or other support, and
crawls up on it. If no suitable substrate is available, the nymph
arches its back, pushes its forelegs close together to raise the thorax
until the middle legs and finally the hind legs and abdomen are
elevated. Since hydrometrid nymphs are usually near land and
hurry to it when they are disturbed, complete immersion probably
is not a frequent occurrence.
In the laboratory, hydrometrids which are not isolated have a
strong tendency toward cannibalism throughout their nymphal sta-
dia. Hatching nymphs may be devoured by others which have
emerged only a short time before. The victims are not necessarily
HypROMETRA MARTINI KiRKALDY 683
the weaker animals; unwary nymphs of the fourth and fifth instars
may be eaten by smaller nymphs. As with the adults, subsurface
Entomostraca and other small animals form the chief food sources,
but any sort of recently killed, or slow-moving, living insect may
be used as food. Nymphs, carrying midges or ostracods with their
beaks, walk on the surface film. In the present study, nymphs in
the field were not seen to feed on organisms too large for them to
carry, but in the laboratory they fed on flies such as Musca and
Drosophila.
The interval between molts is highly variable even in series of
animals kept under similar laboratory conditions. Nymphs hatch-
ing from eggs laid from June 28th to July 8th were isolated and
kept in stender dishes lined with paper towelling. Each animal
was fed one or more fruit flies a day and the exuviae were removed
as they appeared. Under these conditions the intervals between
molts varied between two and five days and the total length of the
developmental period varied between twenty-two and _ thirty-two
days.
As a nymph approaches the time of molting, it reaches nearly
maximum length for its instar. The pigmentation of the cuticle,
which gives a grayish cast to its exoskeleton, is lacking along the
ecdysial line which extends in the mid-line from the metathorax
anteriorly to the middle of the head where the ecdysial line forks
and forms an arm on either side of the frontal region.
In the hour before molting, the nymph imbibes water more
frequently than before. As the gut is distended, the nymph be-
comes a little longer than it had been previously and its entire body
appears turgid. As water is taken in, small droplets of fluid are
emitted through the anus. During the early part of the period, the
animal walks about as much as usual. Then it becomes less active;
if it is undisturbed it moves but little, although it continues to
imbibe water.
The actual molting process takes place rather rapidly. For a
minute or two, the nymph seems to be inactive. Then it resumes
the intake of water and moves backward as it lowers its body.
It pushes its hind legs into the substrate, arranging them so that
they are parallel with the long axis of the abdomen. The middle
legs are planted firmly at right angles with the body and the
front legs extend forward.
When the nymph has placed its legs securely, it lifts its body
and starts to suck air rather than water. As the sucking continues,
634 Tur Universiry SCIENCE BULLETIN
the cuticle of the mesothorax begins to cleave. The head bends
so much that the antennae almost touch the front legs. Air, in
small bubbles, passes through the narrow esophagus into the ven-
triculus. With the distention of the digestive tube, the body length-
ens. Alternate waves of contraction and dilation move up and down
the body.
Since the head is bent ventrally, the prothorax is the most anterior
part of the body at this time. With the posterior end close to the
supporting surface, the long axis of the body forms an angle of
about 45° with the substratum. As the body stretches upward the
proximal parts of the legs emerge from the exuviae and_ begin
outward movements, which help to pull the trunk forward, First
the prothoracic and then the mesothoracic and metathoracic legs
pull away from the exuviae. The legs touch the substrate; the
antennae and finally the beak pull free. The nymph begins to walk
away and the tip of the abdomen slides out from the molted skin.
The color of recently molted hydrometrids resembles that of the
hatching nymphs. The body is pale green and the appendages
are transparent. There is a touch of red on the pleura of the
mesothorax, metathorax and, in the region of the spiracles, on the
abdominal segments. The brownish contents of the posterior ven-
triculus make it distinct. The abdomen of fifth instar nymphs is
marked by an opaque white middorsal stripe and two more trans-
lucent lateral lines. The lora are white and conspicuous.
Within thirty to forty-five minutes a recently molted nymph
again begins to suck up water. When food is available, the nymphs
begin to feed about two hours after they have molted.
Growth and Differentiation
Table 8 presents average measurements of head and body widths,
leg, antennal, head, beak and total body lengths of adult and
nymphal Hydrometra. Because of variations in the length of the
abdomen during the nymphal stadia, total body lengths were meas-
ured at three times. Only the measurements taken at two days after
molting are used in the following discussion of relative growth.
Relative growth rates using these data are shown in figures 1,
2 and 3. When they are plotted semilogarithmically against time
in instars, the average measurements for body and head lengths
fall nearly on straight lines, indicating constant rates of growth
(fig. 1). The points representing measurements of head length,
beak length, antennal length and body width fall on, or nearly
on, straight lines when they are plotted logarithmically against
HypDROMETRA MARTINI KiRKALDY 635
TasBLe 3.—Measurement of nymphal and adults (Hydrometra martini) in milli-
meters (average of 10 animals
Head Leg length
Instar on soft
Length Width 1 2 3
Le ee ewan 0.5 0.17 1.4 1.2 1.3 LS
Be GI ne 0.7 0.18 1.8 ded 1.8 2.4
ccs hier 10 0.2 2.2 2.1 2.4 8.3
TE Pie Sees lve 0.22 2.7 3.2 3.6 4.6
peste eer 2.0 0.3 3.4 3.6 4.5 6.3
PUM ced cage 2.8 0.4 4.0 5.2 5.8 8.4
_ Total body length
ae (interval after hatching) B e ete Bo ay
Immed. 1 hour 2 days
De iepiane sees 1,6 1.2 i6 OF7 0.27
EN ocr prea 1.6 2.3 0.9 0.32
Giviewas or vag 3.0 2.3 3.3 Ll 0.36
ae 4.2 3.1 4.6 1.5 0.41
Weve beves 6.2 5.1 6.5 1.8 0.48
AGUUIYS vein: TOO lye ve eee Ce ns es 2.2 0.50
total body length. Therefore each of these structures grows with
each molt at a rate constant in relation to body length. The slopes
of these lines, however, indicate that only head length increases
at the same rate as body length. The beak grows at 67%, the an-
tennae at 61%, each of the legs at about 87%, and the body width
at 40% of total body length. Through the fourth instar (body length
4.6mm.), the head width increases relatively slowly, that is, at
about 25% of body length and after the third molt the rate ap-
proximates that of body and head lengths.
Data for relative growth of parts of the head are given in table
4 and plotted logarithmically on figure 4. The length of the eye
increases at the slowest rate—58% of total head length. The slopes
of the lines derived from data on preocular and postocular length
636 Tur Universiry ScreNCE BULLETIN
show increase at rates of 108% and 97% of the head length, respec-
tively.
The remarkable variation in the total body length during a
nymphal stadium is the result of a partial telescoping of the ab-
dominal segments and depends mainly upon the degree of dis-
tension of the digestive system (figs. 79 and 80). The minimum
length, which is approximately that of the maximum in the pre-
vious instar, is reached about an hour after the molt is completed.
At this time the air which filled the digestive tube during the
molt has not been replaced by water or food. Later in the stadium,
if the nymph does not maintain a distended gut by feeding fre-
quently, the body length may decrease again. Of the nymphs taken
10.0
° BooY LENGTH
8.0}
6,0}
+OF
3.0-
HEAD LENGTH
LoG OF MEASUREMENT
t a 3 by 5 ADULT
INSTAR
Ficurze 1. Head and body lengths plotted semilogarithmically against time
in instars.
LoG OF Bopy MEASUREMENT
HypROMETRA MARTINI KIRKALDY 6387
5.0r
uot ANTENNAL LENGTH
ig 9 © HEAD LENGTH
BEAK LENGTH
207
1S -
1.0}
Bre
Fr
> 7 Booy Wiort
Al Heap WiptH
ak
are
| i. L AT L
1,0 2.0 4.0 6.0 10.0
LoG oF Bopy LENGTH
Fic, 2, Antennal length, beak length, head length and width, and body width
plotted logarithmically against total body length.
THe Universiry ScieENCE BULLETIN
10.0
LoG OF LEG LENGTH
1.0 n rl rn h n A i
1.0 L5 2.0 3.0 4.0 6.0 3.0 10.0
Loc oF BODY LENGTH
Fic. 3, Leg lengths plotted logarithmically against total body length.
Taste 4.—Comparisons of the sizes of the eye and the preocular and post-
ocular portions of the head of Hydrometra martini nymphs and adults.
Each figure represents an average for 10 animals in millimeters
Instar 1 2 8 4 5 Adult
A utcsapcq hill) die gaan page| 0.25 0.35 0.53 0.71 1.19 1.65
VOT cal iG i a ee 0.08 0.1 0.125 | 0.14 0.17 0.2
Portocular. 43-2... 0.18 0.25 0.35 0.45 0.64 0.95
Total length...... 0.5 0.7 1.0 1.4 2.0 2.8
in the field, most had bodies of nearly maximum length, Food
seems usually to be available to them there,
The nymphal head, like that of the adult, shows little differentia-
tion into sclerites. The dorsal and lateral parts are more heavily
sclerotized and more deeply pigmented than is the ventral part.
HyDROMETRA MARTINI KirnKALDY 639
In the later instars the ventral surface becomes increasingly setose.
The prominent setae, a pair near the posterior end of the head
and a pair lateral to the anteclypeus, remain constant in position.
2.0 fF
PREOCULAR
1.0 F POSTOCULAR
0.6
0.3 +
0.2 Fak EYE LENGTH
Log OF MEASUREMENT
0.04 4. A. A ry A. A. A
04 OS 0.6 as 10 8 2.0 3.0
Log of HEAD LENGTH
Fic. 4. Length of eye, preocular and postocular yi of the head plotted
logarithmically against total head lengths.
640 Tue Universitry Science BULLETIN
In the first and second instars (figs. 82 and 85) the lora and
maxillary plates are difficult to see. During the later stadia, the
maxillary plates are white and conspicuous. Their ventral projec-
tions, the bucculae, appear in the fourth instar nymphs as narrow
rims which are directed anteriorly (fig. 90). In the fifth instar
these lobes are larger (fig. 99) but they do not cover the basal
segments of the rostrum, until the final molt.
The thorax of the hydrometrid nymph shows much less fusion
than that of the adult. In the first instar, the tergum of each
segment is a rather flat broad plate (fig. 80). In the second instar
the posterolateral angles of the mesotergum and metatergum are
produced, indicating the formation of wing pads (fig. 84). The
metatergum is much smaller than the mesotergum in the third
instar nymphs (fig. 88). The wing pads of those individuals which
will become macropterous are much more conspicuously developed
than those of animals which will give rise to apterous adults (cf.
figs. 107 and 108 with figs. 86 and 88). These differences are em-
phasized in the fourth and fifth instar nymphs. In the fourth instar
nymphs, the wing pads reach the second abdominal segment, those
of the metathorax are slightly longer than those of the mesothorax
(figs. 109 and 110). The metathoracic wing pads of the fifth instar
nymph extend to the middle of the third abdominal segment and
the mesothoracic wing pads are slightly shorter (figs. 111 and 112).
The mesothoracic wing anlage of the fourth instar nymphs which
will become apterous adults extend over the bases of those of the
metathorax (fig. 88). In the fifth instar the wing pads of the
metathorax are not developed further, but those of the mesothorax
extend posteriorly as broad lobes (figs. 98 and 99). The posterior
lobe of the pronotum develops gradually; in the fourth instar it
projects over the anterior margin of the mesonotum (fig. 90) and
in the fifth over about a third of it (fig. 99).
The episternum and epimeron are of about the same size in the
prothorax and mesothorax of the first instar (fig. 82). During the
succeeding stadia, the proepimeron and the mesepisternum in-
crease in size more, relatively, than do the proepisternum and the
mesepimeron, so that in the fifth instar there are marked differ-
ences (cf. figs. 82, 86 and 99). In all of the nymphs, the mete-
pimeron is a small posterior lobe, set off by a suture from the
metepisternum. The latter lengthens the most rapidly of any
part of the thorax and by the fifth instar extends toward the mid-
dorsal line. The supracoxal lobes develop gradually and even in
HyYDROMETRA MARTINI KirRKALDY 64]
the fifth instar nymphs are relatively small. The legs of all nymphs
articulate with two processes, one near the plural suture and one
on the sternum (fig. 100).
The thoracic spiracles lie in the membrane between the pro-
thorax and mesothorax and in that between the mesothorax and
metathorax in the first, second and third instars (figs. 82, 85, 86).
In the fourth and fifth the second spiracle becomes associated with
the posterior part of the mesepimeron and the first, with that of
the proepimeron (figs. 90 and 99).
The abdomen of the first and second instar nymphs is marked by
intersegmental lines which are especially noticeable if the abdomen
is contracted. The spiracles of the first segment are more dorsal
than those of the second through the eighth segments (figs. 82 and
85). The tenth segment is separated from the ninth by a ventral
groove (figs. 81 and 83). The dorsal surfaces of the seventh, eighth
and the terminal segments bear more heavily sclerotized setose
plates. In the third and succeeding instars, the dorsal interseg-
mental boundaries are marked by dark lines (fig. 88). Well-de-
veloped intersegmental muscles run between these dark lines on
the dorsal body wall and between the ventral intersegmental lines,
indicating that these lines mark the primary segmentation of the
abdomen.
Differentiation of the external genitalia begins to become ap-
parent in the fourth instar. In the female, a small dorsal projection
of the eighth tergum extends over the reduced ninth segment and
the protiger (figs. 95 and 96). A pair of lobular sclerites represent
the sternum of the ninth segment (fig. 96). In the female fifth instar
nymph the dorsal projection of the eighth tergum is larger (figs.
101, 103); ventrally the sclerotized plates which are the anlage of
the first valvifers of the adult female, are joined by a membrane.
The ventral sclerites of the ninth segment are partially covered by
the eighth segment (fig. 102).
In the male fourth instar nymphs, the eighth abdominal tergum
also has a dorsal projection (figs. 92 and 93). Caudad to this
the terminal segments are differentiated, forming a dorsal scleroti-
zation which will become the proctiger and a ventral scleroti-
zation, the anlage of the genital capsule (fig. 93). In the fifth
instar nymph, tergal projection at the eighth segment is relatively
longer but not as long as the proctiger. The genital capsule is
larger (fig. 105) and the ninth tergum appears as a dorsolateral
lobe (figs. 104, 106).
642 Tue Universiry SCIENCE BULLETIN
Discussion of the Nymph
Although several papers consider the nymphal instars of Hydro-
metra stagnorum, comparatively little work has been done on the
postembryonic development of H. martini. The nymphs of only
two other species are mentioned in the literature, those of H. vittata
(=H. albolineata) by Takahashi (1921) and those of H. gracilenta
by Jordan (1981).
The first description of the complete life history of a hydrometrid
was that of Hungerford (1920). Working in Kansas, he found
that the nymphal stadia lasted on the average two days, with some
taking three to four days. Jordan (1931) found that the nymphal
stadia of H. stagnorum lasted from four to six days. The average
length of the instars of 11 H. martini nymphs reared in Michigan
in the summer is 3.2 days, with variations of two to five days. In
these animals, the first and last stadia were, on the average, longer
than the others. The rapid postembryonic development of H. mar-
tini reported by Hungerford probably is correlated with the warm
Kansas temperatures. There is great variation, however, not only
in the length of time of development (from egg to adult) of the
various individuals, but also in the period that each nymph spends
in the different instars.
In an earlier paper Bollweg (1915) described only four nymphal
instars in Hydrometra stagnorum, but Tevrovsky (1920) and Lund-
blad (1921a, 1921b) as well as Jordan (1931) found five nymphal
instars.
By far the most detailed description and drawings of the hydrome-
trid nymph are those of Lundblad (1921a, 1921b), who was par-
ticularly interested in the proportional growth. He found that the
lengthening of the head, which had also been mentioned by Arrow
(1895), was especially the result of growth of the “praokularen
parallelseitigen Partie.” In H. martini, also, this region grows most
rapidly (see Table 4, fig. 4). Jordan (1931) found, however, that
the preocular region is less elongate in nymphs of H. gracilenta than
in those of H. stagnorum.
The observations made by Lundblad (1921b) on the develop-
ment of the wing pads in the apterous form were also confirmed
in the present work. In both species, these are apparent first in
the second and third instar nymphs as slightly produced corners
of the terga. In the fourth instar these tergal angles are differenti-
ated as wing pads and in the fifth continue to elongate. Lundblad
did not include descriptions of the nymphs of the winged form,
nor has any subsequent paper.
HypROMETRA MARTINI KiRKALDY 6438
Lundblad was concerned also with the segmentation of the abdo-
men. He identified nine segments including the first which Bollweg
(1915) seems not to have found in the late instars because of the
overgrowth of the metathorax and small size of the first segment.
Lundblad mentioned the sclerotization of the dorsal plates of the
terminal abdominal segments but he did not consider the differenti-
ation of the external genitalia.
One of the most interesting features in the development of the
hydrometrid nymph is the extreme variation in the total body
length. This was recognized by Hungerford (1920) who used the
length of the animal from the tip of the head rather than the length
of the body as a criterion for the recognition of the various instars.
The lengths of the appendages, which were tabulated by Lundblad
for H. stagnorum and by Hungerford for H. martini, are constant
throughout each stadium. Both Jordan and Lundblad, found that
the body length varied during each nymphal stadium. Bollweg
(1915) included measurements for four instars; he omitted the
second,
None of these investigators correlated the differences in size in
a given stadium with the expansion of the abdomen which results
from distention of the gut with air or food and water. Both Tev-
rovsky (1920) and Lundblad (1921a) figured first instar nymphs
with extremely contracted abdomens. In Hydrometra martini, the
abdomen is of minimum size approximately a half hour after eclo-
sion and later in the stadium if the nymph does not eat or becomes
dry. The larger size of adult females was mentioned by Jordan
(1981) but he did not correlate variation of size of the nymphs
with sex.
Allometric growth, that is, “growth of a part at a different rate
from that of the body as a whole or of a standard” (Huxley and
Tessier, 1986, p. 380), is a pattern frequently found in insects.
Clark and Hersch published a very comprehensive study of relative
growth of Notonecta undulata in 1940. By measuring anesthetized
animals, they were able to determine growth gradients in individual
insects as well as to calculate averages for males and females. On
the average, each of the legs increased in length at about the rate of
total body length in Notonecta. In Hydrometra leg length averaged
87% of body length. Body width increased an average of 76% in
the Notonecta males in contrast to the 40% increase found in Hydro-
metra.
The similarities in behavior, locomotion and feeding habits of the
nymphs to those of the adults which are reported in the present
Tue Universiry ScieENCE BULLETIN
644
study in general confirm the observations of other investigators.
Arrow (1895) stated that nymphs reared in the laboratory fed only
on springtails until the last instar when they took aphids. Hunger-
ford (1920) observed that the nymphs as well as the adults eat
entomostracans and a variety of insects, including weaker hydro-
metrids. Both Jordan (1931) and Schumann (1934) also men-
tioned cannibalism.
Investigators working with both Hydrometra stagnorum (Arrow,
1895; Jordan, 1931; Schumann, 1934) and H. martini (Martin, 1900)
have found that the nymphs stay closer to land and are more sus-
ceptible to drowning than are the adults. On the other hand,
Takahashi (1921) described the nymphs of H. vittata (= H. albo-
lineata) as swimming under the water and crawling from it to the
surface film. Certainly, under laboratory conditions, H. martini
nymphs are capable of this type of locomotion.
Two curious habits of the nymphs, the elevation-depression move-
ments which were observed also in the adults and the carrying of
the abdomen curved over the back, have been noted by Schumann
(1934) and Jordan (1931).
DIMORPHISM
Observations
Dimorphism in Hydrometra martini is evidenced by two patterns
of wing development; an apterous form which has vestigial wing
pads projecting just beyond the pronotum and a macropterous form
in which the two pairs of wings extend beyond the middle of the
sixth abdominal segment. In the field, the adults of these cate-
gories are differentiated easily and the wing pattern of the fourth
and fifth instar nymphs can be identified with good light. Third
instar nymphs have been separated only with the aid of a binocular
microscope.
Hydrometrids with long wings are scarce and those without
wings are abundant in both Massachusetts and Michigan. In the
four summers in Michigan, an average of about six adults or
nymphs of the macropterous form were taken whereas hundreds
of apterous animals were seen. The ratio was perhaps one winged
to 100 or more wingless hydrometrids. In Massachusetts a higher
percentage of winged forms was collected; it was estimated that
one in 45 was macropterous. In the autumn of 1950, collections at
the Lithia pond yielded a ratio of about one winged individual to
ten of the wingless type. At that time, the pond had little water
HyYDROMETRA MARTINI KirKALDY 645
and the animals including the hydrometrids, were concentrated.
Twenty-seven winged forms were taken.
Apterous and macropterous males and females have been col-
lected through the year. At least four mating pairs of which one
individual was winged and the other wingless have been collected
in the field. In the laboratory they mate readily and produce
fertile eggs.
Studies in the laboratory have been directed toward gaining an
understanding of the cause of development of long wings in some
individuals and of only vestiges in others. In the summer of 1949,
a series of nymphs, 26 of which lived to or beyond the fourth instar,
were reared from eggs of apterous females crossed with apterous
males, under constant light with a temperature of about 30°. All
of the 26 individuals which lived to or beyond the fourth instar
were apterous. In the summer of 1950, a group of similar nymphs
were reared at 30° in a constant temperature oven. The glass door
of this incubator allowed normal daylight to enter. The 31 ani-
mals of this series were also wingless. Of the hundreds of nymphs
reared from eggs of apterous individuals under normal laboratory
conditions during all reasons of the year, none were macropterous.
Offspring of other crosses (winged 9 x winged ¢, wingless 9°
x winged g, winged 9 x wingless ¢ ) have been reared in the
laboratory. Since female hydrometrids store sperm for long periods
of time, only those which emerged from isolated fifth-instar nymphs
or teneral individuals were used. Winged females produced fewer
eggs and were shorter lived in the laboratory than wingless indi-
viduals. Because of the small numbers of animals, the results of
my studies are not conclusive but they suggest that some form of
inheritance governs wing form. ‘These results are as follows:
winged 2 x wingless g gave 16 wingless offspring; wingless 9 x
winged g gave 9 wingless offspring; winged 9 x winged g gave
21 offspring of which 19 were wingless and 2 winged.
Some information on the distribution of winged and wingless
forms of Hydrometra martini has been assembled from the series
in the Snow Entomological Museum at the University of Kansas.
The collection includes 140 specimens taken from three localities
in Kansas in the autumns of 1924, 1925 and 1926, of which 3 or
about 2 per cent are macropterous. A series of 104 hydrometrids
taken from Sedge Point Pool, Douglas Lake, Michigan in June and
July of 1923 is made up of 101 apterous and 3 or approximately
8 per cent of macropterous individuals.
646 Tue Universiry Scrence BULLETIN
Discussion
The presence or absence of wings, as well as variations in their
length, has been of interest to investigators of many groups of
Pterogota. In relatively few species are the causes of these varia-
tions well understood. Poisson (1946) summarized the problem of
apterism in the Hemiptera. Ford (1940) reviewed polymorphism
and taxonomy of insects.
In an earlier review article, Larsén (1930) discussed the loss of
flight by reduction of wing musculature and also variation in types
of wings found in the aquatic and semiaquatic Hemiptera. Of the
insects in these groups, the mechanism of wing development is
best understood in Aphelocheirus aestivalis, a truly aquatic bug.
Larsén (1931) found in the laboratory that individuals reared in
aquaria with running water were winged whereas wing develop-
ment was suppressed in animals kept during their last nymphal
instar in aquaria equipped with air pumps.
As many as six distinct wing types are known in some European
species of Gerris (Ekblom, 1927; Jordan, 1947). All of these may
occur in one locality. Some species, however, are known only
from macropterous and others only from apterous or micropterous
specimens. Observations in Finland of distribution of polymorph-
ism in Gerris najas and G. lacustris led Sahlberg (1868) to believe
that brachypterous forms were significantly more abundant in the
north, and macropterous forms in the south. This was substanti-
ated by Lindberg (1929) who found that brachypterous Gerris
lacustris was the common form in the coastal areas of the outer
archipelago as well as in northern Lapland. Lindberg also corre-
lated the collection of a larger ratio of winged to wingless forms of
G. najas in Finland with the warm temperature in 1917-1925. Jordan
(1943) has found that Gerris odontogaster is also found most often
in the brachypterous form in the north and the macropterous form
in the south of Finland. Ekblom (1950) collected only long
winged G. odontogaster at Skellefteé in Sweden.
Another worker, von Mitis (1937), correlated temperature with
wing form. Working with Gerris in Germany, he found that the
spring generation was brachypterous but the generation which
developed during the warm summer weather was macropterous.
Similarly, Ekblom (1927) collected fewer winged animals at Pited
than at colder Stockholm. Larsén (1950) observed G. lacustris
in southern Sweden and found more rapid postembryonic develop-
ment in individuals with reduced wings than in those with long
wings.
HypROMETRA MARTINI KiRKALDY 647
The only studies reported in the literature of crossing the winged
and wingless forms of gerrids are those of Ekblom (1927 and 1950)
and those of Poisson (1924). Poisson found that in some crosses
of G. lacustris macropterous x macropterous gave about 91 per cent
macropterous and 9 per cent brachypterous; in others this cross
yielded only macropterous individuals. In the same species, bra-
chypterous x brachypterous gave above 80 per cent brachypterous
and 20 per cent macropterous. Poisson also varied environmental
conditions in some of the rearings but.he found that the variations
of food and temperature which he used did not influence the ex-
pression of wings. In 1927, Ekblom working with Gerris asper
crossed apterous x apterous and micropterous x macropterous forms.
Both yielded only macropterous offspring. Four macropterous
females which had resulted from the micropterous x macropterous
cross were mated with an apterous male. Most of the offspring
of these crosses were macropterous but a few were micropterous
and a few, apterous,
Ekblom reported in 1950 on crosses made in the laboratory with
Gerris odontogaster, all of the adults at SkellefteA, Sweden were
macropterous. Of the 91 offspring reared in the laboratory 90
were macropterous and one brachypterous.
Poisson (1924) has also described sexual dimorphism which is
correlated with wing dimorphism in Gerris odontogaster. Microp-
terous males of this species have lateral teeth on the 6th abdominal
segment, which never occur in macropterous individuals.
Comparatively little work has been done on variations of wing
form in Hydrometra. In contrast to the dimorphic Hydrometra
martini, both of the European species, H. stagnorum and H. grac-
ilenta, are polymorphic. Poisson (1924) and Walton (1938) have
found that the apterous form of Hydrometra stagnorum is the
most common in France and England, respectively, but that macrop-
terous and brachypterous forms are also found. Polymorphic
forms like those of H. stagnorum have been described in H. graci-
lenta by Walton (1948) from animals taken in England. Both there
and in Germany H. gracilenta is less frequently collected than is
H. stagnorum, That the brachypterous and macropterous forms
are rare in Germany is obvious from the report of Schumann (1934)
who collected only one winged H. stagnorum and from that of
Jordan (1931) who collected only two winged forms, one of each
species.
Poisson (1924) reported that the offspring of wingless individuals
were always wingless in his rearings and that only winged offspring
648 Tue Universiry SCIENCE BULLETIN
resulted from two pairs of macropterous individuals, which he |
crossed in the laboratory. Although he attempted to cross winged ,
x wingless ¢ and wingless 9 x winged g, these animals did not |
mate. This condition, which he termed “amixie,” he believed im- |
portant in the origin of new species.
However, Jordan (1931) reported collecting mating pairs in
the field in which one individual was winged and the other wingless.
Such a pair of H. gracilenta continued to mate after they were
bought into the laboratory. Jordan, realizing that the female might
have sperm stored from a previous mating, destroyed the eggs laid
during the first ten days. Those deposited after that time were
fertile.
No study of the length of time which females may retain viable
sperm has been reported for Hydrometra stagnorum. If, like H.
martini, this species can retain sperm for a period of months, the
eggs observed by Jordan possibly were fertilized by a previous
impregnation.
The occurrence of winged and wingless individuals in a rela-
tively constant ratio indicates that this is a balanced population.
Balanced populations are defined by Ford (1940, p. 493) as being
those in which “two or more forms of the same species are main-
tained in optimum proportions, a departure from which, in either
direction, constitutes a disadvantage.” In Hydrometra, the most
apparent advantage of the winged condition is motility; these indi-
viduals can escape an unfavorable environment by flying to different
areas. Laboratory studies indicate lower reproductive and survival
rates which, if they occur in nature, would be disadvantageous.
The possibility exists that the winged condition is inherited with a
lethal factor in some parts of its range. The inheritance of a re-
cessive lethal factor with a factor for color occurs in dimorphic
females of some species of the pierid genus Colias (Ford, 1987).
The lower reproductive capacity of winged hydrometrids might
result from: the greater demands for nourishment by the thoracic
muscles and the subsequent deficiency for optimum ovarian growth.
SUMMARY AND CONCLUSIONS
Hydrometra martini is the only species of the family Hydromet-
ridae in the northern part of the United States. Water measurer
and marsh treader are vernacular names applied to insects of this
family and indicate their characteristically deliberate gait. Adults
of H. martini are slender; they measure approximately 10 mm. in
length and 0.5mm. in width. The exoskeleton is heavy and, be-
HypROMETRA MARTINI KIRKALDY 649
cause of the fusion of the sternal area, rigid. Most individuals are
apterous, the wings being represented by small straplike processes;
a few have well-developed wings which extend to the sixth abdomi-
nal segment. The nymphs, especially those of the early instars,
are proportionately less elongate and their exoskeletons are lightly
sclerotized,
These insects are found throughout the spring and summer
on still or slowly moving waters and on the banks. They walk on
the surface film and crawl over the floating and emergent vegeta-
tion, preying upon recently dead or living insects. Reproductive
activity is continuous from early spring until late summer, There
are three generations and a partial fourth generation per year in
both Michigan and Massachusetts. In the fall, when the water
temperature approaches 15°C., the adults of the overwintering
generation crawl on land and seek shelter in depressions of the
earth or under decaying plants. Here they remain until spring
when the migration to water takes place. Since females collected
in the winter and early spring lay fertile eggs in the laboratory,
viable sperm must be stored from copulation in the fall.
The elaborate sculpturing of the egg of Hydrometra martini re-
sults from the subdivision of the exochorion into minute air cham-
bers. The length of time of development of the embryo depends
upon the temperature; at 25° this period is 11 to 14 days. The hatch-
ing process, which is rapid and uniform in nature includes the
molting of an embryonic cuticle.
From the time the nymph walks away from the egg shell, its
activities closely resemble those of the adult. This is especially true
of its locomotion, its grooming and its feeding.
The morphological changes are more marked and involve both
proportional growth and differentiation. With each molt, the
head increases in length to a much greater extent than in width.
This growth is especially marked in the preocular region, The head
capsules of the nymphs are lightly sclerotized, but in them, as in
the adults, there are differentiated sclerites associated with the
mouth parts.
The mouth parts, like those of all hemipterans, consist of a labium
which forms a sheath enclosing the mandibular and maxillary
stylets, The labrum partly covers the base of this sheath and the
elongate epipharynx extends into it. The maxillae form a single
channel through which saliva passes to the prey and food materials
are pumped back to the pharynx. The complex musculature of the
mouth parts is contained entirely within the elongate head capsule.
650 Tue Universrry SciENCE BULLETIN
The well-developed salivary glands lie in the anterior part of the
thorax.
Changes in the developing thorax involve the fusion of sclerites
and the differentiation of the tergum associated with wing forma-
tion. The fusion in the thorax includes the union of the pleura
with the sterna which occurs only in the adult and also the reduction
of the membranes joining the regions of the thorax in the nymphs to
the sutures found in the adults. The development of the supracoxal
lobes of the episterna and epimera begins in the fourth instar
nymphs. The articulation of the legs of both the nymphs and adults
is such that ‘movement of the coxae is on a horizontal plane. The
legs are slender and especially in the adults, elongate. The setae
of the tarsi and the distal parts of the tibiae are used in the almost
continuous grooming motions which help in keeping these surfaces
hydrofuge.
Differentiation of the tergum is especially associated with wing
formation. The pronotum, which completely covers the mesotergum
in the adults, is not markedly developed even in the fifth instar
nymphs. The mesoterga and metaterga of the first instar nymphs
are rectangular plates; the posterior angles of these sclerites indicate
the development of wing pads in the second and succeeding instars.
In the third instar, the wing pads of individuals which will become
macropterous are somewhat larger and in the fourth and fifth instar
nymphs these are conspicuously larger than the wing pads of poten-
tially apterous individuals. The mesonota of macropterous nymphs
are not differentiated into sclerites as are those of adults. The
metanotum is progressively compressed by the upward growth of
the metaepisterna. In the fifth instar nymphs, the metanotum is a
rather wide membrane but in the adult, it is represented by an ex-
ternal line and an internal fold on which the metanotal leg muscles
take their origin.
The mechanisms regulating wing growth in Hydrometra are not
known. About three per cent of the specimens of H. martini which
were collected in Michigan and Kansas are macropterous. Genetic
control is suggested by crosses made in the laboratory. The off-
spring of two apterous individuals are always apterous. In the
small numbers of crosses involving macropterous individuals, only
a few of the offspring of two macropterous parents and none of
those having one apterous and one macropterous parent, were
macropterous.
The first pair of thoracic spiracles lies in the posterior border of
the proepimera and the second, in that of the mesoepimera in the
HypROMETRA MARTINI KiRKALDY 651
nymphs. Each pair shifts anteriorly to lie just behind the supra-
coxal lobe of the adult epimera. In the nymph each of the abdominal
segments I through VIII bears a pair of spiracles. Those of seg-
ments I and VIII are lost in the adult. The tracheal system consists
of two independent dorsolateral trunks which extend into the head,
tracheae to the viscera, and tracheae to the ventral body-wall.
The nymphal abdomen consists of ten segments. In the first three
instars, each of these, except the reduced tenth segment, is a
simple cylinder separated from adjacent segments by secondary
segmentation. In the third and succeeding instars the dorsolateral
parts of the intersegmental boundaries are marked by heavy dark
lines, and in the fourth and fifth instar nymphs the terminal seg-
ments are modified with the differentiation of the external genitalia.
In the adult, the first abdominal segment is greatly reduced and
all of the pregenital segments are fused ventrally and laterally;
segmentation is indicated only by the terga and by the positions
of the abdominal spiracles.
The external genitalia of the female consist of broad plates which
serve as a guide in the deposition of the egg; they show considerable
reduction in comparison with the oviposition apparatus of more
nearly typical hemipterans. The male external genitalia closely
resemble those of related insects; a genital capsule formed from the
ninth abdominal segment bears a telescoped aedeagus and a pair
of claspers or parameres,
The internal organs follow the pattern of other hemipterans
although these organs are attenuated, following the body form.
The female reproductive system includes a pair of ovaries each of
which consists of seven ovarioles. Under optimum conditions, each
ovariole can produce an egg per day. The ducts from the ovaries
are simple; the seminal receptacle associated with them may store
sperm for a period of months. The male reproductive system con-
sists of a pair of elongated testes, each of which is connected by a
vas deferens with a seminal vesicle, and a common ejaculatory duct.
The digestive system is a simple tube; the ventriculus is especially
long. The salivary glands lie in the thorax and open at the junction
of the maxillae by a single duct. Their secretion has a toxic effect
on the prey. The compact central nervous system is composed of
the fused ganglia of the head, thorax and abdomen and is found in
the posterior part of the head and the anterior part of the thorax.
The dorsal blood vessel extends from the seventh abdominal seg-
ment to the posterior part of the head.
652 Tue Universiry ScieNcE BULLETIN
REFERENCES CITED
Arrow, G. J.
1895. Habits of Hydrometra stagnorum. Science Gossip, 11:201-208.
Baptist, G. A.
1941. The morphology and physiology of the salivary glands of Hemip-
tera-Heteroptera. Quart. Rev. Micr. Sci., 83:91-139.
Beamer, R. H.
1949. Personal communication.
Buatcuiery, W. S.
1926. Heteroptera of Eastern North America. xii-+ 1116 pp. Nature
Publishing Co., Indianapolis, Indiana.
Botitwec, W.
1915. Beitrag zur Faunistik und Okologie der in der umbegung Bonns
vorkommenen aquatilen Rhynchoten, mit besonderer Beriicksichti-
gung ihrer Larvenverhiltnisse. Werhandle. des Naturhistor. Ver-
eins der preufs. Rheinlande und Westfalens, 71:127-187.
Branpt, E.
1878. Vergleichend-anatomische Untersuchungen iiber das Nervensystem
der Hemipteren. Horae Soc. Ent. Rossicae, 14:496-505.
Brinpuey, M. D. H.
1930. On the metasternal scent glands of certain Heteroptera. Trans.
Ent. Soc. London, 78:199-207.
Brocuer, F.
1911. La ponte Yoeuf et la larva der Limnobates. Ann. Biologie La-
custre, 4:367-379.
Bueno, J. R. pe LA Torre
1905. Notes on Hydometra martiv Sirk. (= lineata Say), Canadian
Ent,, $7: 12-15,
1926. The family Hydometridae o1 the Western Hemisphere. Ent. Amer-
icana (n. s.), 7:83-128.
1937. A Glossary of Entomology. Brook. Ent. Soc., Brooklyn, New York.
Burr, bia,
1943. Comparative study of the mouthparts of representative Hemiptera-
Homoptera. Mem. Cornell Univ. Agric. Exp. Sta., 254:1-20.
Crank, B. and A. H. Herscu
1939. A study of relative growth in Notonecta undulata. Growth,
8:847-872.
Comstock, J. H.
1936. An Introduction to Entomology. xix +- 1044, Ithaca, N. Y.
Comstock, J. H. and J. G. NrrpHam
1898. The wings of insects. American Nat., 32:249-252.
Durour, L.
1833. Recherches anatomique et physiologiques sur les Hemiptera, ac-
compagnées de considerations relatives 4 Vhistoire naturelle et a la
classification de ces insectes. Mem. Acad. Sci. Inst. France, Sav.
Etr., 4:131-461.
HypDROMETRA MARTINI KIRKALDY 653
DuPortg, M.
1946. Observations on the face in insects. J. Morph., 79:371-418.
Expsiom, T.
1926. Morphological and biological studies of the Swedish families of
Hemiptera-Heteroptera. Part I. Zool. Bidrag Fran Uppsala, 10:31-
179.
1927. Vererbungsbiologische Studien tiber Hemiptera-Heteroptera, Her-
editas, 10:3833-359.
1930. Morphological and biological studies of the Swedish families of
Hemiptera-Heteroptera. Part II. Zool. Bidrag Fran Uppsala,
12:118-150.
1950. Uber die Fliigelpolymorphismus bei Gerris odontogaster Zett. Notu-
lae Entomologicae, 30:41-49,
Exson, J. A.
1987. A comparative study of Hemiptera. Ann. Ent. Soc. America,
80:579-597,.
Forp, E. B.
1987. Problems of heredity in the Lepidoptera. Biol. Rev., 12:461-508.
1940. Polymorphism and taxonomy, in The New Systematics. viii 4- 583,
J. Huxley, ed. Oxford, London.
Grirriru, M. E.
1945. The environment, life-history and structure of the water boatman,
Ramphocorixa acuminata, Univ. Kansas Sci. Bull., 30:1-241.
Gross, J.
1901. Untersuchungen iiber das Ovarium der Hemiptera, zugleich ein
Beitrag zur Amitosenfrage. Zeitshrif. wiss. Zool., 69:139-201.
Guvpr, J.
1902. Der Dorsaldriise der Larven Hemiptera-Heteroptera, Sencken-
bergische naturforschende gesellschaft, Frankfurt, Bericht., 1902:
85-136.
Wamiuron, M. A.
1931. The morphology of the water scorpion, Nepa cinerea L, (Rhyn-
chota-Heteroptera). Proc. Zool. Soc. London, 1931:1067-1136.
Hrymons, R.
1899. Beitriige zur Morphologie und Entwicklungs-geschichte der Rhyn-
choten, Nova Acta, Abhandl, d Kaiserl, Leop.-Carol, deut, Akad.
d. Naturf., 74:351-456,
Hoxg, S.
1926. A preliminary paper on the wing-venation of the Hemiptera
(Heteroptera). Ann, Ent, Soc, America, 19:18-34,
Huncerrorp, [. B.
1917. Notes concerning the food supply of some water bugs, Science
(N. S.), 45:336-337,.
1920. The biology and ecology of aquatic and semiaquatic Hemiptera,
Kansas Univ, Sci. Bull., 11:3-265.
654 Tue Universiry Scrence BULLETIN
Huncerrorp, H. B. and N, E. Evans
1934. The Hydrometridae of the Hungarian National Museum and other
studies on the family (Hemiptera), Ann, Mus, nat. Hungarici
Budapest, 28:31-112.
Huncrrrorp, H, B. and P. J, SPANGLER
1952. Personal communication.
Huxxey, J. S. and G. Tressrer
1936, Terminology of relative growth,
Nature, 137:380-381.
Jaczewsxi, T.
1936. Contributions to the knowledge of aquatic Heteroptera of Egypt.
Ann, Mus, Zool, Polonaes, 11:171-211,
Jouannsen, O. A, and F. H. Burr
1941, Embryology of insects and Myriapods, xi + 462. McGraw-Hill,
New York.
Jorpan, K. H. C.
1931, Uber die Entwicklung und Lebenweise von Hydrometra stagnorum
L. and H. gracilenta Horv. Isis Budissina, Bantzen, 12:19-27.
1948, Zum Flugelpolymorphismus von Gerris odontogaster Zett. (Hemip.-
Heter,). Arb, morph, Taxon, Ent, Berlin-Dahlem, 10:157-163.
1947, Brachypterni forma Gerris gibbifer Schum. flaviventris Put, (Hem.
Het. Gerridae). Casopis Csl, spolecnosti Entomologicke, 44:47-50.
Kirxatpy, G, W.
1899. Guide to the study of British waterbugs. Entomologist, 32:108-
115.
1900. Recent notes on Hyrometra martini = lineata Say, ibid, 33:175-
176,
Kuncssury, B, F. and O. A. JonannsEn
1927. Histological Technique. vii +- 142. J. Wiley & Sons, New York.
KraMe_r, S.
1950. The morphology and phylogeny of Auchenorhynchus Homoptera
(Insects), Ilinois Biol. Mong., 20(4):1-78.
Larsin, O,
1931. Beitrag zur Kenntnis des Pterygopolymorphismus bei den Wasser-
hemipteren, Kungl. Fysiografiska Siillskapets, Lund Forhand.,
1:8-11.
1938. Untersuchengen iiber den Geschlectsapparat der aquatilen Wanzen.
Opusc. Entom., Suppl., 1:1-388.
1942, Bisher unbeachtete wichtige Ziige im Bau des Metathorax bei den
Heteropteren. Kungl. Fysiografiska Sillskapets, Lund Forhand.,
12:149-1638.
1945a. Der thorax der Heteropteren, Skellet und Muskulature. Lunds
Univ. Arsskraft N. F., 41(3):1-96.
1945b. Das thorakale Skelettmuskelsystem der Heteropteren. Ibid.,
41(11):1-83.
1950a, Der Veranderungen im Bau der Heteropteren bei der Reduktion
des Flugapparates. Opusc, Ent., 15:17-51.
HypROMETRA MARTINI KiRKALDY 655
1950b. Einiger Beobachtungen an Gerris lacustris L. (Hem, Het.) Opus.
Ent., 15:131-138.
Leucxart, R.
1855. Uber die Mikropyle und den feinenren Bau der Schalenhaut bei den
Insekteneieren. Arch. Anat. Physiol. u. wiss. Medicin. 1855:90-
264,
Linpperec, H.
1929. Vingdimorfismen hos Hemiptera-Heteroptera. Berentning om det
18 skand. Naturforskermg¢de, Copenhagen, 1929:405-410.
Lunpsiap, O.
1921a. Zur Kenntnis der jungen Larven einiger in Wasser lebenden Rhyn-
choten. Sirtryck ur Entomologists Tidskrift, 41:53-59.
1921b, Zur Kenntnis der jungen Larven einiger in Wasser Iebenden Rhyn-
choten. II-VI. Ibid., 41:93-106.
Mauour, N. S. R.
1933. Studies on the internal anatomy of the stink bug. Bull. Soc. Roy.
Entom. D’Egypte, 1933:96-119,
Mammen, H.
1912. Uber die Morphologie der Heteropteren-und Homopterenstigmen.
Zool. Jahrb., Anat., 34:121-178.
Martin, J. O.
1900. A study of Hydrometra lineata. Canadian Ent., 32:70-76.
Micuener, C. D.
1944, A comparacive study of the appendages of the eighth and ninth
abdominal segments in insects. Ann. Ent. Soc. Amer., 37:336-351.
Miris, H. von
1987. Oekologie und Larvenentwicklung der mitteleuropeischen Gerris-
arten (Heteroptera). Zool. Jahrb., 69:3387-371.
Newcomer, W. S.
1948, Embryological development of the mouth parts and related struc-
ture of the milkweed bug, Oncopeltus fasciates (Dallas). J. Morph.,
82:365-411,
PaLtumBa, G.
1891. Gli amori del ’'Hydrometra stagnorum. Revista Italiana Sci. Nat.
11:1-4.
Poisson, R.
1924. Contribution a T’etude des Hemipteres aquatiques. Bull. Biol.
France Belgique, 58:49-805.
1946, L’apterisme chez les insectes. Anneé de la Revue Scientifique,
84:605-617.
1951. Ordre des Heteropteres in Traite de Zoologie, Tome X, Fasc. II pp.
972-1948. Edited by P. Gresse, Masson et Cie., Paris.
Rirry, C. F, C.
1918. Food of aquatic Hemiptera. Science (N.S.), 48:545-547.
Sauxzere, J.
1868. Bidrag till kannedom om Finlands Dimorph Insekt-arten. Notiser
ur Sallsk. pro Fauna et Flora Fennica forh., 9:199-220.
Tue Universiry Screnck BULLETIN
656
ScHUMANN, W.
1934. Beitriige zur Biologie einiger Heteropteren. Mitt. ent. Ges. Halle,
13:54-60.
SEwEL, F.
1924, Die Geschlechtsorgane in der embryonal Entwicklung von Pyr-
rhocoris apterus. Zeitschr. f. Morph, u. Oekol. d. Tiere, 1:429-506.
SERVADEI, A.
1946. Reparti sulla costituzione dell’ apparato bocalle della specie ap-
partenenti alla famiglie: Gerridae, Veliidae, e Hydrometridae
(Hem, Heteropt.). Bull. Int. Ent. Bologna, 15:217-228.
Sincu-Prutui, H.
1925. The morphology of the male genitalia in Rhynchota. Trans, Ent.
Soc. London, 1925:127-267.
Snopcrass, R. E,
1933. Morphology of the insect abdomen. Part II. The genital ducts
and oviposition. Smithsonian Misc. Coll., 89(8):1-148.
1935. The Principles of Insect Morphology. xi-+- 667, McGraw-Hill,
New York,
1938. The loral plates and the hypopharynx of Hemiptera. Proc. Ent.
Soc. Washington, 40:228-236,
1941. The male genitalia of Hymenoptera. Smithsonian Misc. Coll.
99(14):1-86.
1944. The feeding apparatus of biting and sucking insects affecting man
and animals. Smithsonian Misc. Coll., 104(7):1-113.
1947. The insect cranium and the “epicranial suture.” Smithsonian
Mise. Coll., 107(7):1-52.
Spooner, C. S.
1938. The phylogeny of the Hemiptera based on a study of the head
capsule. Illinois Biol. Mong., 16:1-102.
SticuEL, W.
1926. Die Fauna der Phaueninsel. Abh, u. Ber. der Pommerschen
Naturf. Gesellschaft, 7:103-169.
Tanaka, T.
1926. Homologies of the wing veins of the Hemiptera. Annot. Zool.,
Tokyo, Japan, 11:83-53.
Taytor, L. H.
1918. The thoracic sclerites of Homoptera and Hemiptera. Ann. Ent.
Soc. Amer., 11:225-249.
Taxanasut, R.
1921. Observations on Hydrometridae (2). Insect World (Konchu sckai),
25:8-12.
Trvrovsky, V.
1920. Ceske Vodomerky (Gerroideae). Sonderabdruck aus Entomol-
ogické prirucky, Prague, 9:1-51.
Tower, D. G.
1918. The external anatomy of the squash bug Anasta tristis Deg. Ann.
Ent. Soc. Amer., 6:427-442.
HyYpDROMETRA MARTINI KIRKALDY 657
Watton, G. A.
1938. A water bug new to Great Britain—Hydrometra gracilenta Horvath,
Ent. Mon. Mag., 74:278-274.
1943. The water bugs (Rhynchota-Hemiptera) of North Somerset. Trans.
Soc. British Ent., 8:281-290.
Wenner, H.
1926. Problem der Gleiderung des Insektenthorax I-III. Zool, Anz.,
66:9-31.
1980. Biologie der Hemipteren. vii + 544. Springer, Berlin.
Wikre, G.
1908. Einiges iiber die Biologie und Anatomie der Wasserlaufer (Hydro-
metridae). Naturwissenchaftliche wochenschrift, 23:209-218.
WiccLeswortn, V. B.
1939. The Principles of Insect Physiology. viii + 434. Dutton & Co.,
New York.
WeEsENBERG-LuUND, C,
1918. Fortpflanzungsverhiltnisse, Paarung und Ejiablage der Siisswasser-
insekten, Fortschritte der naturw. Forschung, 8:161-286.
1948. Biologie der Siisswasserinsekten. vii + 682. Springer, Berlin.
Wycopzinski, P.
1947, Sur le Trichotonannus setulosus Reuter, avec un theorie sur lorigine
des harpagones des Heteropteres males (Hemiptera-Heteroptera,
Crytostemmatidae). Revue francaise d’entomologie, 14:118-125.
22-—3378
Tae Universiry ScrENCE BULLETIN
PLATE I
Fic. 5. Dorsal view of the head.
Fic. 6. Ventral view of the head.
Fic. 7. Lateral view of the head.
Fic. 8. Anteclypeus, labrum and epipharynx in lateral view.
3. 9. Anteclypeus, labrum and epipharynx in ventral view.
HypROMETRA MARTINI KIRKALDY 659
PLATE I
LABRUM
MAXILLARY PLATE ANTECLYPEUS
LORUM POSTCLYPEUS
10 MM (MAXILLARY PLATE
LABRUM ANTECLYPEUS.
BUCCALA. EPIPHARYNX ——
ANTENNAL SUTURE
LABIUI
LABUM ANTECLYPEUS
LABRUM
EPIPHARYNX 8
EYE
ANTECLYPEUS
LABRUM
EPIPHARYNX 9
Tue Universiry SCIENCE BULLETIN
PLATE II
Fic. 10. Right antenna in medial view.
Fic. 11. Mandibular apparatus. Right mandible in median view.
Fic. 12. Maxillary apparatus. Dorsal view with protractor and retractor
muscles of the right maxilla removed to show the maxillary sheath.
HYpROMETRA MARTINI KIRKALDY 661
PLATE II
MAXILLAE
HYPOPHARYNX LORUM
ANTENNAL MUSCLES
MAXILLARY PROTRACTOR -
SHEATH
MANDIBULAR LEVER
MANDIBULAR PROTRACTOR
MAXILLARY RETRACTOR l2
MANDIBULAR RETRACTOR
MANDIBLE
II
Tur University Science BULLETIN
PLATE WI
Fic. 18. Ganglia of the central nervous system.
Fic. 14.. Cross section of the labium through the third segment.
Fic. 15. Salivary syringe.
Fic. 16. Cross section of head at the level of the mandibular lever.
Fic. 17. Tip of right maxilla. ‘ |
Fic. 18. Tip of left maxilla.
Fic. 19. Anterior part of the head dissected to show the mandibular ap-
paratus and the pharynx. |
Fic. 20. Frontal view of the head.
HypROMETRA MARTINI KiRKALDY
PLATE III
LABIUM
OPTIC NERVE
PHARYNX
PROTOCEREBRAL GANGLION
HEART
DEUTOCE REBRAL GANGLION
TRITOCEREBRAL GANGLION bUCT
SUBESOPHAGEAL GANGLION
ESOPHAGUS SALIVARY SYRINGE
THORACIC GANGLIA
ABDOMINAL GANGLIA
16 PHARYNX
MANDIBULAR LEVER MANDIBLE
MAXILLA
SALIVARY APPARATUS 7
19 MANDIBLE
HYPOPHARYNX
| MANDIBULAR LEVER
> PHARYNX
MANDIBULAR PROTRACTOR
LORUM
20
LABRUM
MANDIBULAR RETRACTOR
BUCCALA
~ EPIPHARYNX
ANTECLYPEUS.
MAXILLARY PLATE
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Tur Untiversiry SciENCE BULLETIN
PLATE IV
Pronotum of winged adult. Dorsal view.
Pronotum of winged adult. Lateral view.
Pronotum of wingless adult. Dorsal view.
Pronotum of wingless adult. Lateral view.
Pterothorax of wingless adult. Lateral view.
Pterothorax of wingless adult. Dorsal view.
Thorax of wingless adult. Ventral view.
HypDROMETRA MARTINI KirRKALDY
PLATE IV
SPIRACLE
10 MM
—-EPISTERNUM Hiceiaiaky 7
——- EPIMERON
— WING PAD
N SPIRACLE
fp EPISTERNUM
ME TANOTUM
EPIMERON
25 26 rag
24
665
Tue Universrry ScteNCE BULLETIN
PLATE V
Fic. 28. Pterothorax of winged adult in lateral view.
Fic. 29. Pterothorax of winged adult in dorsal view.
HyYpDROMETRA MARTINI KinKALDY
PLATE V
merrermnacee une eR INAIEN
ACROTERGITE ——_
“= PREALARE BRIDGE
PREALARE MEMBRANE}
————= PRESCUTUM \
PARAPSIDAL SUTURE
SCUTUM
}—— MESOEPIST ERNUM——+}-—
SCUTAL WING PROCESS.
~PLEURAL WING PROCESS
“mee SCUTELLUM
MESOPOSTNOTUM ~~
Ny
METATHORACIC WING
METANOTUM
METATHORACIC RIDGE ~~
MESOEPIMERON=——————
SECOND SPIRACLE
anne ME TAEPISTERNUM ‘
ae SECOND PHRAGMA*
/ zener THIRD PHRAGMA
ME TAEPIMERON
AXILLARY CORD- a 2 .
MESOTHORACIC WING - ; &
29
667
Fic.
Fic.
Fic.
Fic.
Tue University SCIENCE BULLETIN
PLATE VI
Right prothoracic leg in median view.
Right mesothoracic leg in median view.
Right metathoracic leg in median view.
Right mesothoracic wing.
Right metathoracic wing.
HypROMETRA MARTINI KirnKALDY 669
PLATE VI
L—~ TIBIA
Tue University SCIENCE BULLETIN
PLATE VII
Fic. 35. Wing bases of the mesothoracic and metathoracic wings, Both
wings are lifted to show the basal sclerites. Lateral view.
Fic. 36. Prothoracic muscles of the head. Base of the head and muscles
of the right side in lateral view.
M 1. M. pronoti primus,
M 2. M. pronoti secundus.
M 38. M. pronoti tertius,
M 6. M. prosterni primus,
M 7. M. prosterni secundus,
M 10. M. proepisterno-postoccipitalis,
Fic. 87. Lateral view of the coxa and trochanter of the right prothoracic
leg showing notal and pleural muscles,
18. M. noto-trochantinalis,
M 14. M. noto-coxalis primus.
M 16. M. noto-coxalis tertius,
M 17. M. pleura-coxalis.
M 20. M. noto-trochanteralis,
M 21. M. pleura-trochanteralis,
Fic, 38. Lateral view of the coxa and trochanter of the right mesothoracic
leg showing notal and pleural muscles,
M 40. M. noto-trochantinalis,
M 41. M. noto-coxalis,
M 42. M. episterno-coxalis.
M 46. M. noto-trochanteralis,
M 47. M. pleura-trochanteralis,
Fic, 39, Lateral view of the coxa and trochanter of the right metathoracic
leg showing notal, pleural and trochanteral muscles,
M 63. M. noto-trochantinalis,
M 64, M. noto-coxalis,
M 66. M. episterno-coxalis.
M 70. M. noto-trochanteralis,
M 71. M. pleura-trochanteralis,
M 73. M. coxa-trochanteralis medialis,
M 74, M. coxa-trochanteralis lateralis,
Fic, 40, Pterothorax dissected to show the principal flight muscles of the
right side, Lateral view. The leg muscles are removed,
M 30. M. mesonoti-primus,
M 34. M. dorso-ventralis primus.
M 38. M. episterno-alaris,
M 39. M. furca-pleuralis (mesothoracic),
M
M 57, M. metanoti-secundus,
M 62. M
. furca-pleuralis (metathoracic),
HyYDROMETRA MARTINI KirnKALDY
PLATE VII
RIDGE
THIRD AXILLARY
SCLERITE.
PLEURAL WING PROCESS
LOMM
nies
THIRD PHRAGMA
as uM
M62
i
SECOND PHRAGMA
671
Tue Universiry SCIENCE BULLETIN
PLATE VIII
Fic. 41, Abdomen of adult male, Lateral view of left side,
Fic, 42. Abdomen of adult male. Dorsal view.
Fic. 43. Abdomen of adult female, Dorsal view.
HyYpROMETRA MARTINI KIRKALDY 6738
PLATE VIII
OS MM,
\e
9 soft SPIRACLE
N
° —_—
4~— LATEROTERGITE ++
r $5 |4+ LATEROTERGITE
ee
t—— GENITAL CAPSULE"
TERGUM Wt -——
4| 42
43
Tue Universiry SCIENCE BULLETIN
PLATE IX
TERMINAL SEGMENTS OF THE ADULT FEMALE
Fic. 44. Dorsal view.
Fic. 45. Ventral view.
Fic. 46. Lateral view.
Fic. 47. Lateral view. Left half of exoskeleton of the seventh segment and
the left parasternite of the eighth segment have been removed to expose the
genitalia and the musculature of the region.
(Diagrammatic. )
HyYpDROMETRA MARTINI KirKALDY 675
PLATE IX
TERGUM MI cIRST
jERGUM Sat CL ae
FIRST VALVIFER
44
TERGUM MN
45
LATEROTERGITE
TERGUM Xa
-PROCTIGER
SECOND VALVIFER
FIRST VALVIFER
46
SUBGENITAL PLATE
MUSCLES OF SECOND VALVIFER
PROTRACTOR OF FIRST VALVULA\
TERGUM MIL
~PROCTIGER
———-ANAL OPENING
— SECOND VALVIFER
HIND GUT—————
RETRACTORS OF
ra oer
FIRST VALVULA Seema reed FIRST VALVIFER
VULVA
AT FIRST VALVULA
Tue Universiry SCIENCE BULLETIN
PLATE X
TERMINAL SEGMENTS OF THE ADULT MALE
Fic. 48. Lateral view of the left side. The eighth segment is deflected ven-
trally.
Fic. 49. Genital capsule in lateral view. The left side of the capsule is
removed to demonstrate the muscles of the basal plate and the parameres.
The proctiger and the phallobase are partially protracted.
Ftc. 50. Dorsal view of the ninth segment.
Fic. 51. Dorsal view of the ninth segment, the proctiger removed.
Fic. 52. Lateral view of the ninth segment, with the intromittent apparatus
everted,
HYDROMETRA MARTINI KIRKALDY CIT
} PLATE X
LATEROTERGITE,
TERGUM ‘VC TERGUM UI
LASERS
NC —.
z % iF
= e hy
a { { i |
TE
STERNUM ‘Ut 48
PROCTI
GENITAL CAPSULE: ares
_PHALLOBASE
RETRACTOR OF BASAL PLATE wo CLASFER
—=CLASPER MUSCLES
PROTRACTOR OF BASAL PLATE
BASAL PLATE
10MM
—— TERGUM I
GENITAL CHAMB ER
PHALLOBASE
PROCTIGER————
CLASPER
BASAL PLATE
GENITAL CAPSULE
SO
10MM
joan
PROCTIGER
GENTAL CAPSULE
Tue Universrry Scrence BULLETIN
PLATE XI -
Fic. 53. Salivary glands of the left side in dorsal view.
Fic. 54. Digestive system in dorsal view.
Fic. 55. Female reproductive system in dorsal view.
Male reproductive system in dorsal view.
HyYpDROMETRA MARTINI KrRKALDY 679
PLATE XI
TERMINAL, FILAMENT as
DUCT ene
PRINCIPAL, SALIVARY
GLAND.
a
ACCESSORY SALIVARY
GLAND
33 VENTRICULUS
MATURING EGGS
TESTIS
“MATURE EGG
VAS DEFERENS
MALPIGHIAN ye
COMMON OVIDUCT UBE
I GLAND OF SE
INTESTINE
MINAL.
RECEPTACLE
SEMINAL VESICLE
RECTAL DIVERTICULUM
» VAGINA
SEMINAL, RECEPTACLE
RECTUM
BURSA COPULATRIX
54
EJACULATORY DUCT
bs fe
56
Tur Universiry SCIENCE BULLETIN
PLATE XII
Fic. 57. External view of an egg of Hydrometra martini, showing the
sculpturing of the exochorion.
Fic. 58. Cleared egg, showing the endochorion, micropylar tube and basal
spicule.
Fic. B9. Cross section through the basal region of the egg, above the basal
disc.
Fic. 60. Cross section through the apical region of the egg, below the
micropyle.
Series or Diacrams SHowinc THE DrvELOPpMENT oF Livinc Ecos
From tue Sixra THroucu tHe E.evenrn Day or INCUBATION AT 25°
Fic. 61. Egg showing the position of the eyes at 9:30 and 10:30 AM
on the sixth day of incubation.
Fic. 62. Egg showing the position of the eyes at 11:30 AM on the sixth
day of incubation.
Fic. 63, Egg showing the position of the eyes at 12:00 noon on the sixth
day of incubation.
Fic. 64, Egg showing the position of the eyes at 12:15 and 12:45 on the
sixth day of incubation.
Fic. 65. Egg showing the development on the seventh day of incubation.
Fic. 66. Egg showing the development on the eighth day of incubation.
Fic. 67. Egg showing the development on the ninth day of incubation.
Fic. 68. Egg showing the development on the tenth day of incubation,
Fic. 69. Egg showing the development on the eleventh day of incubation.
HypROMETRA MARTINI KIRKALDY 681
PLATE XII
MICROPYLE
I \ MICROPYLAR TUBE
XN
EMBRYONIC REGION
RY a SPICULE
o
ey,
59
Sy
SPICULE \
MICROPYLAR TUBE 60 62
57 lartacument vis 58 6!
ii
SERIES
Fic.
Fic.
Pic.
Fic.
Fic.
Fie,
Fic.
Fic.
Fic.
or SkercuEes to Suow THE Hatcuinc Process 1s Hydrometra martini
78.
of abdomen still within exuviae.
THe Universiry Science BULLETIN
PLATE XIII
Head pushing through split in chorion; half of eyes exposed.
Head exposed; eyes and labrum free of the chorion.
Thorax nearly exposed.
Thorax exposed. \
Nymph arching back, pulling legs, antennae and beak out.
Nymph straightening back and flexing head.
Exuviae split; beginning of “prenatal molt.”
“Prenatal molt” nearly complete.
Nymph approaching supporting surface; antennae, beak and tip
HypROMETRA MARTINI KIRKALDY 683
PLATE XIII
BiG;
abdomen.
EiG,
Fic.
Fic.
Fic.
Fic.
Fic.
19.
80.
81.
82.
83.
84.
85.
Tue Unrverstry Science BuLLetTiIn
PLATE XIV
Drawincs oF Fimst AND Seconp Instar NymMpus
Dorsal view of first instar nymph with an extremely contracted
Dorsal view of first instar nymph.
Ventral view of first instar nymph.
Lateral view of first instar nymph,
Ventral view of second instar nymph.
Dorsal view of second instar nymph,
Lateral view of second instar nymph.
685
HypROMETRA MARTINI KiRKALDY
PLATE XIV
THe Universiry ScieNCE BULLETIN
PLATE XV
Drawincs oF Turrp Instar NyMpus
Fic. 86. Lateral view.
Fic. 87. Ventral view.
Fic. 88. Dorsal view.
687
HypDROMETRA MARTINI KiRKALDY
PLATE XV
Fic.
Fic.
Fic.
Fic.
Fic,
Fic.
Fic.
Fic.
Fic,
89.
90,
Ol.
92,
93.
94,
95.
96,
97.
THe Untiversrry Screnck BULLETIN
PLATE XVI
Drawincs or Fourts Instar Nympus
Ventral view of head and thorax of apterous form.
Lateral view of head and thorax of apterous form.
Dorsal view of head and thorax of apterous form.
Dorsal view of terminal segments of the male.
Lateral view of terminal segments of the male.
Ventral view of terminal segments of the male.
Dorsal view of terminal segments of the female.
Lateral view of terminal segments of the female.
Ventral view of terminal segments of the female.
HYDROMETRA MARTINI KIRKALDY
PLATE XVI
9|
89
O.SMM
PROCTIGER PROCTIGER
ue 92 i
95
wa
Mar
ee O ee PROCTIGER
xk 93 x 96
Var wa
: Le PROCTIGER oe
* 94 x 97
23—3378
689
690 THe University ‘SCIENCE’ BULLETIN
PLATE XVII
Drawines or Firri Instan Nympus
Dorsal view of head and thorax of apterous form.
Lateral view of head and thorax of apterous form.
Ventral view of head and thorax of apterous form.
Lateral view of terminal segments of the female.
Ventral view of terminal segments of the female.
Dorsal view of terminal segments of the female,
Lateral view of terminal segments of the male.
Ventral view of terminal segments of the male,
Dorsal view of terminal segments of the male,
HYDROMETRA MARTINI KiRKALDY
PLATE XVII
e
99 rele)
PROCTIGER
PROCTIGER
98
ER. sages a “5
Drawincs oF THE THoRACES OF Nympus or THE MacroprTerous Form
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
. Dorsal view of the third instar nymph.
. Lateral view of the third instar nymph,
. Lateral view of the fourth instar nymph,
. Dorsal view of the fourth instar nymph,
. Lateral view of the fifth instar nymph,
. Dorsal view of the fifth instar nymph,
Tue Universrry ScreENCE BULLETIN
PLATE XVIII
HypROMETRA MARTINI KirRKALDY
PLATE XVIII
es:
693
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVIII, Pr. 1] DrcempBer 20, 1956 [No. 10
A Taxonomic Study of the Genus Rhagovelia (Hemiptera,
Veliidae) of the Western Hemisphere *
1D
Joun A. Bacon
A. B., University of Kansas, 1941
Apstractr: This paper includes descriptions of seventy-cight species of the
| genus Rhagovelia (Hemiptera; Veliidae) from North America, Central America,
South America and the Antilles. In addition to the descriptions, a bibliography
and the synonymy is given for each species, and detailed data on distribution
is presented, Keys to the species groups and to the species within the groups
are included,
Twelve new species are named and described: Rhagovelia acuminata from
Panamd, Rhagovelia formosa from Guatemala, Honduras and Mexico, Rhago-
velia gracilis from Mexico, Rhagovelia horrida from Mexico and Guatemala,
Rhagovelia impensa from Pert, Rhagovelia merga from Panamé, Rhagovelia
modesta from Brazil, Rhagovelia palea from Pert and Bolivia, Rhagovelia scabra
from Costa Rica and Panama, Rhagovelia scitula from Brazil, Rhagovelia solida
from Costa Rica, Rhagovelia viriosa from Pert.
The following specific names become synonyms: Rhagovelia arctoa Bueno
(= Rhagovelia obesa Uhler), Rhagovelia flavicincta Bueno (= Rhagovelia
obesa Uhler), Rhagovelia gregalis Drake and Harris (= Rhagovelia tenuipes
| Champion), and Rhagovelia regalis Drake and Harris (= Rhagovelia tenutpes
|
Champion), The following varietal names become synonyms: Rhagovelia
distincta arizonensis Gould (= Rhagovelia distincta Champion), Rhagovelia
distincta cadyi Gould (== Rhagovelia distincta Champion), Rhagovelia distincta
harmonia Gould (= Rhagovelia distincta Champion), Rhagovelia distincta
modesta Gould (= Rhagovelia distincta Champion), Rhagovelia distincta
Proxima Gould (=:Rhagovelia distincta Champion), Rhagovelia distincta val-
entina Gould (= Rhagovelia distincta Champion), Rhagovelia sinuata calcaris
Drake and Harris (= Rhagovelia robusta Gould), and Rhagovelia collaris
pulchra Gould (== Rhagovelia collaris (Burmeister) ). One subspecies Rhago-
velia collaris planipes Gould has been raised to species level as Rhagovelia
f * 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 Doctor
Philosophy. Contribution No, 927 of the Department of Entomology, University of Kansas.
(695)
:
696 Tue University SciENCE BULLETIN
planipes Gould. One species Rhagovelia williamsi Gould, has been re-estab-
lished as a valid species, removing it from the synonomy of Rhagovelia amazon-
ensis Gould.
Brief notes are presented on the biology of Rhagovelia rivale Bueno. The
history of the family is presented and generic concepts are discussed,
Nine species groups are established and their phylogenetic arrangement
within the genus is discussed. The probable phylogenetic arrangement of these
groups of species begins with the angustipes group. Two paths of development
are followed from this beginning. The first includes the abrupta group, the
elegans group, and the crassipes group. The second includes the collaris group,
the obesa group, the ainsliei group, the spinigera group, and the hirtipes group.
CONTENTS
PAGE
UWGOQUIOUION: Sin) es Pe ye a en 697
CKMOWIGCOINEICS 7, vu Gs ieee cdt ak (oak ys ee 698
Biology ov the Gonus Wi tpoVela wis yc ah i tn he 698
PLDI eek a a Es eek 699
AGHV Iti sir in sak 17 eS DR, a ane GRIN Bs 6 TA Ee 699
Hood Frise) sc0 nly ted. on lana, prebiaenl hairs 700
VERTIS a. ete eid th eae al, 1 ty aA ee ew ae Al 701
OVIDOSIHOM. Sick Bane a ee Rs i i a ean ET 702
ihmmidtiine Stages is ust sh Ua, eats FV. OOS eh eaves leony 702
HINDCPURHONM 5 ieee Ge ee ak oe 702
Collecting: Teoliniqued: ican vt des. dei AS Aosnere ee 702
‘laxoromysot the GenussRhagoveliseisens.). casas mea oo ww dl ces 703
Taxonomic History and Generic Concepts...............00.s0e eee 703
Biylogeny aie). tev. ohne, Hv ss IO aor) aes Gales 704
Disttibtitiontas alk Liege, a anwralsauaule kash aan ens 705
Idemtificatior: Techulquei. dena. aaa inna) on al au). 706
Generic: Characteristics 4... 4277 ie an ener ees oh 709
Réy.to.the Species Groupsiivis. ecwivas pees He bawia aor deacon 709
Angus pes Groupeinwiaa.. wield, alll pad Oa 710
Abripia Group aad ale ahiente Alaa o fanless 754
legans.@toup: jell eee eed! etiam isasonh ls. lana 768
Orissiper Group sossd vata didi piiwiiel aa Bd eed 778
COLANS GTOUD. +. eisaiiln, elated) el. Bless aeceterins Ge 816
Obes Grapy a viicieed. wis alana eh and hee 837
Alnehen God... aidan.) wisely mie A OL ts Noes 857
SpinigerasGtoup saisiadi! owutits. Gradiyedan ts Pina ae 863
Hn ee Groupes 1 ina} aia al ev ene aac ine) 8 814
Bibliography, seaneenl. .) vad) pend, vinieaets 15 Glued os 889
TDQGSs a sane baa: | Bike ae ey: ean eh alas Peon 895
BIGtSE® seloerion in: cisGir id ikrutve etait hcp Pibtnvaps wid ene eras Pea Ges
Stupy oF THE GENUS RHAGOVELIA 697
INTRODUCTION
The genus Rhagovelia Mayr (Hemiptera, Veliidae) was sug-
gested by Professor Herbert B. Hungerford as worthy of taxonomic
study. No correlation on a generic level had been made between
Species which were described after the last generic revision by
Doctor George E. Gould in 1931, and those which were described
previously. A large number of specimens had been added to the
Francis Huntington Snow Entomological Collections in the last
two decades giving much additional material for study. At the
time of the revision of the genus by Gould in 1981, the entire col-
lection of Rhagovelia in the Francis Huntington Snow Entomolog-
ical Collections, University of Kansas, numbered approximately
5000 specimens, whereas at this writing the author has had approxi-
mately twenty thousand specimens available. Most of the addi-
tional material was undetermined when the study was begun.
Available keys to the species of the genus Rhagovelia have proven
unserviceable to the average entomologist. Doctors Carl J. Drake
and Halbert M. Harris published descriptions of twenty-eight
Species and one subspecies subsequent to Gould’s 1931 paper. The
claspers of the male genitalia, which are of great taxonomic value,
were not figured for any of these species, nor was a key incorporating
these new species presented. Descriptions by earlier workers were
based mostly on color.
The present account provides unified descriptions of all the pre-
viously known species represented in the Francis Huntington Snow
Entomological Collections, University of Kansas, as well as of the
new species which have been found in the course of this work.
Workable keys, based on clear-cut structural characteristics wher-
€ver possible, and divided into groups of related species of the genus
Rhagovelia of the Western Hemisphere, are included.
The work of O. Lundblad published in 1933 in Stockholm,
Sweden, entitled “Die altweltlechen Arten der Veliidengattungen
Rhagovelia und Tetraripis” will serve as an aid for determination
of specimens from the Eastern Hemisphere.
Two species, Rhagovelia trailii (White) and Rhagovelia bakeri
Bergroth, are not adequately represented by type material; the
Original descriptions are not detailed enough to permit their being
included in the groups of species or in the keys. Original descrip-
tions of these two species are included at the end of the section
on taxonomy.
698 Tue Universiry SctiENCE BULLETIN
An addenda follows the section on taxonomy. This addenda in-
cludes the recent species published by Dr. C. J. Drake and Dr. R.
F. Hussey after 1953. These species were published after the body
of the paper was written and thus, in most cases, could not be
worked into the body of the paper.
ACKNOWLEDGMENTS
The writer is grateful to éach of the following persons: Professor
H. B. Hungerford suggested and directed this study and gave
valuable advice and criticism. Professor Raymond H. Beamer, in
charge of the Francis Huntington Snow Entomological Collections,
has given valuable advice and furnished a majority of the speci-
mens studied; many of these specimens were collected by biological
survey trips under his direction. Professor E. Raymond Hall has
given valuable advice on the preparation of the manuscript. Pro-
fessor Kathleen C. Doering has given advice on morphological prob-
lems. Professor Charles D. Michener has given valuable suggestions
regarding intraspecific variation, phylogeny, and general procedure.
Dr. Edward ‘A. Chapin of the United States National Museum
has permitted the study of the Rhagovelia in the collection under
his charge. The collection of the late Mr. J. R. de la Torre-Bueno,
which contains many specimens valuable in the determination of the
previously named ‘species, as well as many type specimens, has
been made available and is now in the Francis Huntington Snow
Entomological Collections. Dr. W. E. China of the British Museum
compared specimens with the types of the species described by
Champion. Professor H. M. Harris of Iowa State College has
loaned specimens from the collection under his charge.
BIOLOGY OF THE RHAGOVELIA
Individuals of the genus Rhagovelia are both small and timid,
making it difficult for accurate observations to be carried on under
natural conditions. Attempts to observe them in nature rarely are
satisfactory. Since Rhagovelia is small, an observer must approach
closely to see what is going on, and when an attempt is made to get
within several inches of individuals of the genus they dart away
rapidly just out of range, where they again take up their stations,
darting and drifting with the current.
When confined in an aquarium these bugs at once begin frantic
efforts to escape, pressing their heads against the glass sides of the
aquarium and rowing frantically with their long, strong middle
legs. When these efforts to escape fail they take to diving, and
Srupy oF THE GENUS RHAGOVELIA 699
swim about under the surface of. the water. They swim readily
under water and carry down with them a supply of air trapped in
the waterproof pile which covers the body. However, they soon
tire, and die without trying to regain their position on the surface
of the water.
Various attempts, all in vain, have been made by the author to
rear Rhagovelia. Tanks with running water, and tanks with cur-
rents set up by jets of compressed air, did not meet the needs. At-
tempts to rear specimens on damp sphagnum or on clear water
have repeatedly failed; two days was the longest time of survival
obtained with either method. With damp filter paper in the bottom
of finger-bowls, both adults and nymphs have been kept alive for
four and five days. This filter paper must be kept damp or the
bugs will perish in a few hours.
Torre-Bueno (1907) has published the most thorough account of
the biology of the Rhagovelia based on R. obesa Uhler. Bueno’s
observations, amplified by those of the writer on R. rivale Bueno,
may be summarized as follows:
Habitat
Rhagovelia inhabits principally the swifter streams and rivers.
Two species, Rhagovelia plumbea Uhler and Rhagovelia salina
(Champion) are salt- and brackish-water inhabitants, living in the
bays and coves along the shores, Most of the species live on fresh
water, and prefer swift running streams, one common name of
these bugs being “riffle bugs.” Doctor Raymond H. Beamer, how-
ever, captured a pair of Rhagovelia distincta Champion in Arizona
on the surface of the water in a horse-tank forty miles from the
nearest running water (oral information), In general Rhagovelia
congregates in schools on the surface of the water near riffles and
rapids,
Activity
Riffle bugs are strong striders on the surface and can maintain
their position for some time against a rapid current. Appearing to
tire of striving against the current they drift to quieter water where
they row slowly near the bank. In nature I have never seen one on
the bank or shore, nor swimming under water. The nymphs con-
gregate close to the banks in sheltered areas where they dart about
rapidly. In slow flowing water they are most often seen gliding
slowly against the current a few inches from the bank. In riffles
and rapids they are most often found in the middle of the stream
rowing strongly against the current and resting for only a few sec-
Tue Universiry Science BULLETIN
700
onds before again gaining their positions. These insects seem to be
gregarious as most often many are congregated in a few pools or
rifles while long stretches of the stream will be uninhabited.
Rhagovelia glides by means of a vigorous rowing action of the
middle pair of legs while the body is supported by the two other
pairs of legs. Observations have been made on the swimming
motions of Rhagovelia by Champion (1901), Torre-Bueno (1907),
and Coker, Millsap and Rice (1936). Rhagovelia is clumsy on
land, often attempting to row with the middle pair of legs. In-
dividuals are able to walk on a dry surface if not frightened, but
are slow and hesitant in comparison with their behavior on the
water. On the surface of the water they are vigorous and adept,
darting so rapidly that the eye can hardly follow them. Their skill
in “swimming” is due to the peculiar construction of the last tarsal
segment of the intermediate legs. That segment is cleft for three
fourths of its length, and there is a series of long ciliated hairs aris-
ing from a common stem in this cleft. This series of hairs, the
swimming tuft, can be folded within the cleft of the tarsus or ex-
tended fanwise from the cleft. When in use, the entire length of
the tarsus is in contact with the water and the slit is vertical to the
surface. When in this position the spread tuft of hairs projects
beneath into the water and is a powerful auxiliary in swimming.
When the insect is swimming under water the tuft of hair is ex-
panded and is of great assistance. The bug glides on the tarsal
segments of its anterior and posterior legs which have hairs to aid
in supporting the insect when it is resting on the surface film. In
captivity the bug often glides with its venter resting on the surface
of the water, but in this position it does not move swiftly.
Much time is spent in cleaning the legs and special care is used
in the cleaning of the intermediate legs. The intermediate leg is
vigorously scrubbed between the anterior femur and the anterior
tibia. Another cleaning action, seen frequently, also involves the
middle leg. In this action the middle leg is scraped against the
ventral surface of the posterior femur. The posterior femur in most
species of Rhagovelia is set with long, sharply pointed spines, and
the intermediate leg is drawn repeatedly back and forth over these
spines, which act as a comb to clean and straighten the swimming
plume of the intermediate tarsus.
Food
It is difficult to determine exactly the food used by Rhagovelia.
In common with other members of the family Veliidae, riffle bugs
are predaceous, living on various microcrustaceans and small in-
Srupy oF THE GENUS RHAGOVELIA 701
sects trapped in the surface film. Numerous attacks on small
lepidopterous larvae, which were being carried by the current,
have been observed but in no instance did the Rhagovelia remain
with the larvae for more than a few seconds. Nymphs of Rhagovelia
probably feed on microcrustaceans which are picked up from the
surface film, and on other types of animal matter. Second and
third instar nymphs have been observed to feed on mosquito larvae
which were stranded but still alive, as well as on abdomens which
had been removed from grasshoppers. When feeding on insects
as hard-bodied as grasshoppers the bugs probe with their beaks
until unsclerotized areas are located and then settle down to feed
for ten to fifteen minutes. As feeding progresses the abdomen of
the feeding bug becomes distended with the material taken in.
One observation of a third instar nymph of R. rivale feeding on
a live stranded fourth-instar larva of a Culex mosquito showed the
feeding process to last approximately ten minutes. The nymphal
bug was confined in a finger-bowl with damp filter paper on the
bottom. A fourth instar mosquito larva was placed on this damp
filter paper. In the course of its restless wanderings the Rhagovelia
nymph soon walked over the mosquito larva. Seemingly recog-
nizing its danger, the mosquito larva began thrashing about vigor-
ously; nevertheless the nymphal bug inserted its beak between
the third and fourth abdominal segments and proceeded to suck
the contents from the body of the mosquito larva.
Mating
Torre-Bueno has described the mating behavior in some detail
for R. obesa; the author has observed copulation for R. rivale, and
the details are the same for the two species. In copulation the
male is above. The male sets his hind femora at right angles to
his body, bending the tibiae under, and by means of them holds
the female’s second and third pairs of legs straight and close to
her body. Once he is firmly on her, he releases this hold, but
maintains his position by the anterior legs, which clasp the fe-
male over the. prothorax. He is not connected with the female
continuously while on her back. The actual connection is only
for a few seconds’ duration; each time that it occurs the genital
segments of the male are bent downward and under to effect con-
nection, After the act the male lies quiescent upon the back of
the female. As long as the male is on her the female does all
the skating.
702 THe Universiry Science BULLETIN
Oviposition
Little is known about the oviposition of Rhagovelia. Eggs are
probably glued at the waterline on stones in the streams, or to
the upper surface of floating leaves of water plants. Since there
is great difficulty in keeping Rhagovelia alive in aquaria, it has
not been possible to determine the preference of locality for ovi-
position. Live adults, which were frequently copulating, were
kept under close observation, but no oviposition was observed.
The adults were confined two days before the last female died.
Hight days after the death of the last adult female first instar
nymphs were observed in the aquarium. The period of incuba-
tion would thus fall somewhere between eight and ten days.
Immature Stages
The nymphs of Rhagovelia are flattened and oval, with the
pronotum sutured off from the mesonotum in all stadia. There
are five nymphal stadia, during which progressive development
takes place. The wing pads of those individuals which will de-
velope into winged adults first appear in the fourth instar and are
well developed in the fifth instar. The length of time required
for each instar has not been reported, since no means has been
found for keeping them alive in captivity for more than five days.
Several nymphs were observed in the process of molting, First
the old nymphal skin splits along the mid-line of the back from
between the eyes to approximately the middle of the abdomen.
Next the thorax is freed and it is followed by the head, antennae,
front legs, hind legs and finally the long intermediate legs. Sev-
eral nymphs have been kept in captivity until time to shed the
skin, but each was unable to survive this ordeal, either drown-
ing if kept on water, or dying if kept on damp filter paper. How
the skin is shed in nature is not known,
Hibernation
Hibernation probably is in the adult stage. The collections
made earliest in the spring included only adult males and females.
Later in the season only nymphal forms are to be found, and by
midsummer mixed adult-nymph collections can be made, Where
hibernation takes place is unknown; the litter at the edge of the
stream probably is the place.
Collecting Techniques
Rhagovelia is comparatively easy to collect, but requires a quick
hand and proper equipment. Equipment needed consists of a
water net, or, more easily handled, a six-inch tea strainer which
Stupy OF THE GENUS RHAGOVELIA 708
will offer less resistance to the current than does a net. The strainer
can be tied on the end of a stick or pole that can be picked up
at the scene; also several jars or wide-mouthed bottles one-fourth
full of damp sphagnum moss or damp leaves are a necessity if the
specimens are to be brought to the laboratory alive. If the speci-
mens are to be killed a wide-mouthed cyanide killing bottle may
be used, or a little tobacco smoke can be blown into the jars con-
taining the damp sphagnum, which are then sealed. In a short
time all the specimens will be dead and can be transferred to vials
of seventy percent alcohol, or into pillboxes lined with several
layers of cellucotton, Often Rhagovelia congregates in such num-
bers that a single dip with the net will capture a dozen individuals.
Usually, however, only one individual can be captured at a time.
To bring specimens into the laboratory alive it is only necessary
to cover the top of the jars containing damp sphagnum with cheese-
cloth or to punch several holes in the metal screw top cover with
which such jars are usually equipped. Treated in such a manner,
specimens will stay alive for at least one day. If placed in bottles
containing water the specimens will drown before they can be
brought into the laboratory.
The sexes skate together in the riffles and along the banks and
both sexes are generally represented in equal proportions in any
random collection.
TAXONOMIC HISTORY AND GENERIC CONCEPTS
The genus Rhagovelia was established in 1865 by G, L. Mayr
to receive three species which had been described in the genus
Velia by H. C. C. Burmeister in 1832. Mayr’s generic concept
included three-segmented tarsi throughout and the third tarsal
segment of the intermediate leg cleft for three fourths of its length.
In 1871 P. R. Uhler described the species Rhagovelia obesa from
the United States, and, in 1872, Rhagovelia plumbea from the
British West Indies. The minute basal tarsal segments of the
anterior tarsus on some species were difficult to observe. This
led to the establishment of two new genera based on the reduced
number of segments in the anterior tarsus. In 1879 F. B. White
erected the genus Neovelia to receive his trailii, and 1898 G. H.
Carpenter erected the genus Trochopus to receive the species
marinus. In 1898 G. C. Champion described the species salinus
and placed it in the genus Trochopus thus making two species
in that genus. Also in 1898 G. Breddin described a second species,
whitei, in the genus Neovelia.
Tue Universiry ScreENCE BULLETIN
704
In 1901-02, G. W. Kirkaldy published a series of papers in
which he stated that he had examined material from the type
locality of Trochopus marinus Carpenter and after careful study
of the anterior tarsus had reached the conclusion that Carpenter's
Trochopus marinus was identical with Rhagovelia plumbea Uhler,
and that the names Rhagovelia and Trochopus applied to the
same genus. In the same series of papers he also placed the
genus Neovelia White as a synonym of the genus Rhagovelia.
Champion’s description of nine new Rhagovelia in the “Biologia
Centrali-Americana,’ more than doubled the number of known
species in the genus. Kirkaldy described two new species in his
1901 paper, and published a check list of the nineteen species
known from the Western Hemisphere. Since then several writers
have added to our knowledge of the genus. J. R. de la Torre-
Bueno, C. J. Drake, H. M. Harris, and George E. Gould, among
others, have described many new species since the turn of the
century. Gould, in 1931, published keys and descriptions of the
then known species, which totaled forty for the Western Hemi-
sphere. After 1931 Drake and Harris continued their work on
the genus Rhagovelia, adding eighteen new species and one variety
between the years 1938 and 1940,
PHYLOGENY
The genus Rhagovelia is one of the two most highly specialized
genera in the family Veliidae. Both Rhagovelia and Veloidea have
the peculiar construction of the third tarsal segment of the inter-
mediate leg. The genus Veloidea has the posterior as well as the
intermediate tarsi split and set with a swimming plume. The pos-
session of these modified tarsal segments is a factor which enables
the members of these two genera to adapt themselves to their
specialized environment.
Of the groups of species within the genus as set up in this paper,
the angustipes group is considered to be the most primitive because
of the sutured off pronotum (which agrees with the condition
found in all Rhagovelia nymphs) and the broad, evenly tapering
dorsum of the abdomen of the apterous female such as is found
in the other members of the family Veliidae. Two distinct lines of
development can be traced from this beginning. In the first line
the abdomen of the apterous female remains much as in the primi-
tive condition. This line includes such groups of species as the
abrupta group, the elegans group, and the crassipes group. The
second line is characterized by the apterous female having the
Srupy oF THE GENUS RHAGOVELIA 705
dorsum of the abdomen narrow after the first three segments. This
second line of development includes the collaris group, the spinigera
group, the obesa group, the ainsliei group, and the hirtipes group.
The spinelike termination of the last genital segment which
occurs on several species was found to have arisen independently in
at least three groups. The shape of the claspers of the male gen-
italia generally agree as to basic type within any one group. With
this fact in mind, an examination of the species which have the
spinelike termination of the last genital segment shows, at once,
that several different types of claspers are involved. Rhagovelia
uncinata Champion is closely related to the other members of the
elegans group, yet possesses the mucronate genital segment. R.
acuminata sp. nov. is clearly in the collaris group, yet R. acuminata
also possesses the mucronate genital segment. R. ainsliei Drake and
Harris, R. becki Drake and Harris, and R. gracilis sp. nov. all agree
in having a mucronate genital segment, but are closely related to
the obesa group in the form of the dorsum of the abdomen of the
apterous female. In this one case it was thought convenient to
include these three species in a group, known as the ainsliei group,
as they are so well set off from the obesa group by the form of the
male clasper, as well as by the mucronate genital segment.
DISTRIBUTION
Each description in the taxonomic part of this paper is followed
by a section entitled “Data on Distribution.” Previous published
records for the species and its synonyms are indicated in the first
paragraph of the section; the remainder of the section is devoted
to listing the specimens examined by the writer in the course of
preparation of this paper. It must be borne in mind when inspect-
ing the published records of distribution that the records have not
been verified as pertaining to correctly identified species.
The known geographic ranges of most species is extended by the
record-stations of occurrence mentioned in this paper; all such rec-
ords are indicated by an asterisk immediately before the name of
the country or state concerned. The countries are listed alpha-
betically. The states of Mexico and the United States are listed
alphabetically under the country to which they pertain, and the
locality, date, c@llector and number of specimens of brachypterous
and macropterous forms of each sex are listed. Unless otherwise
indicated specimens are in the Francis Huntington Snow Entomo-
logical Collection. A parenthetical entry indicates the source of the
other specimens from each locality. The type specimens in the
706
Tue Universiry ScreENCE BULLETIN
Francis Huntington Snow Entomological Collections, University of
Kansas which are mentioned in the “Data on Types” are not re-
listed in the “Data on Distribution” section.
As far as can be determined, there is no species with a circum-
polar distribution in the genus Rhagovelia. Several species of the
Eastern Hemisphere fit well into groups established in this paper.
Rhagovelia is mainly tropical and semitropical. _Mexico, Central
America, the islands of the Caribbean, and northern South Americ:
contain the major number of species within the genus. One group,
the obesa group, occurs throughout the United States. The species
R. obesa Uhler has the most northerly distribution of any Rhago-
velia, having been taken as far northward as Ontario, Canada.
A study of the Rhagovelia collection of the Francis Huntington
Snow Entomological Collections, University of Kansas, indicates
that winged forms are more frequently taken on the small, tem-
porary brooks and streams, while apterous individuals are mostly
found on the larger, more permanent bodies of water.. The winged
form is unknown in the two salt-water species, R. salina (Cham-
pion) and R, plumbea Uhler. This would indicate that the presence
or absence of wings is controlled by the environment. The individ-
uals which live on water which will continue to support them have
no need of wings, whereas the individuals living on water which is
only temporarily flowing, or that is apt to evaporate, need wings
to seek out other water’ on which to continue their existence.
It is the opinion of the author that the distribution of the species
of the genus Rhagovelia as recorded in this paper represents merely
the’ areas of collection rather than the range of the species. It is
felt that many more species of this genus will be discovered and
the present known range of many existing species will be increased
when a more complete coverage of many areas in Central and
South America is undertaken.
IDENTIFICATION TECHNIQUES
Early workers depended almost entirely on color and color pat-
terns for the differentiation of species in the genus Rhagovelia.
Structural characteristics were ignored or unobserved while a slight
variation in color was considered sufficient basis for establishing
a new species. This led to much confusion sincep in most cases,
the color descriptions applied in full only to the specimen from
which they were written. In this paper a description of the color
is given for each species. This color description is intended to
Srupy OF THE GENUS RHAGOVELIA 707
serve only as guide to the most common color pattern and must be
flexibly interpreted.
Descriptions and keys in the present paper are based on the
apterous forms as this seems to be the normal condition for the
genus; more than one third of the species are known only from the
apterous forms. However, most winged forms can be determined
by noting group characteristics which apply to the winged forms
and by the shape of the male clasper which will, in most species,
clearly show group relationships if not specific identity. The apter-
ous male usually shows the best specific characteristics, while the
apterous female shows the group relationships most clearly.
The principal structural characteristics which have proven valu-
able are the proportions of the segments of the antennae (princi-
pally in the angustipes group ), and the comparative length to width
of the pronotum of the apterous forms. The armature of the an-
terior and posterior trochanter, the pattern of armature of the
posterior femur, the clasper of the male genitalia, and the form
of the venter of the last abdominal segment of the male provide
other useful characters. A character which has not been used
previously, and which is apparently constant for a given species,
is the presence or absence of certain minute conical black setae
or spicules (Pl. I, fig. 4) on parts of the venter of the body. These
minute conical setae are arranged in the same manner in apterous
and winged forms of both male and female specimens except that
those of the females tend to be slightly reduced in many species.
Characteristics which have been found to be variable in many
species include the incrassate condition of the posterior femur
of the male, the proportions of the last two tarsal segments of the
intermediate leg, and in some species the armature of the posterior
femur, All such variation noted is mentioned in the description
of the species involved.
The measurements given for each species were made with a
combination of 9X oculars with a squared reticle, and 6.8X_ ob-
jective lenses on the binocular microscope. At this magnifica-
tion each ten units of the reticle equals .15mm. Conversion to
millimeters has been made only for length and width of the entire
insect; ‘all other measurements are in units of the reticle. In the
measurement of the segments of the antennae the minute “node”
at the base of the third segment is considered to be a part of that
segment. In measuring the length and width of the pronotum as
well as segments of the legs care must be taken to have the body
708 Tue Universrry SctieNCE BULLETIN
of the insect or the part being measured as nearly horizontal as
possible. Limited variations in the proportions given in the descrip-
tions of the species may be expected. Whenever proportions
are used as key characteristics it is intended that the couplets of
the keys be strictly interpreted, since they are phrased in such a
manner as to be mutually exclusive after considering the maximum
variation to be expected. The width of the insect is the greatest
width of the dorsum; the length of the pronotum, mesonotum
and metanotum is measured on the mid-line. In measuring the
projecting spine on the pronotum of the winged forms the projec-
tion beneath the spine, which marks the apex of the pronotum,
is used as a base. Some variation must be expected from that
given for the armature of the posterior femur. In general the
descriptions are stated in such a way as to allow for known varia-
tion.
In order to examine the male genitalia it is necessary to remove
the genital capsule. After relaxing the male specimen with a
drop of five percent alcohol it is easy to remove the genital cap-
sule with a pair of fine-pointed forceps. The capsule is boiled
in ten percent potassium hydroxide for approximately five minutes
and is then neutralized in a one percent solution of hydrochloric
acid. The capsule is then placed under glycerin on a well-slide. -
Dissecting needles made from “minutennadeln” can be used to
tease the claspers from their attachment within the capsule. The
clasper is examined under the dissecting microscope with the
blade pointed toward the observer's right. Extreme care must be
taken to obtain a flat surface view of the blade of the clasper.
This is best accomplished by submerging a fragment of a broken
glass cover-slip beneath the glycerin and propping the free end
of the clasper on this support. After observation of the clasper,
it, along with the genital capsule, may be stored in a tiny glass vial
containing a small amount of glycerin. The vial may be pinned
through. its cork beneath the specimen from which the genital
capsule was removed. Permanent slides of the claspers have
been found to be highly unsatisfactory as the angle of view can-
not be accurately controlled. The “penis appendages” of Gould
were not used in the determination of specimens because these
appendages are entirely too fragile and too little chitinized to re-
tain their shape after boiling in potassium hydroxide. The aedeagus
of the males and the ovipositors of the females are unsatisfactory
from the standpoint of specific determination.
Strupy or THE GENUS RHAGOVELIA 709
Genus Ruacoverra Mayr
Logotype nigricans Burmeister
1865. Rhagovelia Mayr, Vehr. Zool. -bot, Ges, Wien, Bd. 15, p. 445.
1865. Baecula Stal, Hemiptera Africana, vol. 3, p. 157.
1879. Neovelia White, Jour. Linn. Soc. London, Zool., vol. 14, p. 487.
1898. Trochopus Carpenter, Ent. Mo. Mag., vol. 34, p. 78.
Generic characteristics: Minute to medium-sized bugs, oval to
fusiform in general shape. Head marked with a median impressed
line and several lateral dots. Rostrum extending to mesosternum;
three segmented. Antennae long, four segmented, first segment
curved and longest; first three segments bearing long, scattered,
erect setae, Wings without clavis, corium or embolium. Veins
well-developed on basal two thirds of wings, becoming obsolete
toward apex. Apterous forms common and with no trace of wings
or wing pads. Anterior legs not fitted for grasping. Intermediate
legs modified as swimming legs. Third tarsal segment of inter-
mediate leg split for three fourths its length and this split furnished
with retractable, feathered hairs arising from a common base.
All tarsi three-segmented, the first segment usually much reduced.
Third tarsal segment of all legs provided with a pair of claws set
in before the apex. Posterior femur generally spinose beneath,
especially in the male. Abdominal segments and genitalia bi-
laterally symmetrical.
Kry To THE Spectres Grours OF THE GENUS RHAGOVELIA
CE alt)
1, Pronotum of apterous forms shorter than length of eye, posterior
margin straight or slightly concave.......... angustipes group
Pronotum of apterous form longer than length of eye, posterior
Mar olCONVEN ic. i A ee Cer nh sd 2
2.(1) Pronotum of apterous forms much less than three times as long
as exposed portion of mesonotum............-. abrupta group
Pronotum of apterous forms more than, or approximately three
times as long as exposed portion of mesonotum, or pronotum
Covering mesonolUm 666. hye ee 8
8.(2) Posterior tibia armed at apex with a sickle-shaped spur,
; elegans group
Posterior tibia with or without a distinct terminal spur which may
be slightly bent but never with a sickle-shaped spur at apex.... 4
4.(8) Dorsum of abdomen of apterous female definitely narrowed
abruptly after first three segments... 6... k ee. 6
Dorsum of abdomen of apterous female tapering more or less
Oven) AG: Apekicacs (las bape Gre TA 5
710 Tue Universiry Sctrence BULLETIN
5.(4) Intermediate leg of apterous female with tarsal segment II subequal
to tarsal’ segment Us). A ees S hirtipes group
Intermediate leg of apterous female with tarsal segment II definitely
shorter than tarsal segment II]. ...0. 0... crassipes group
6.(4) Intermediate femur of female transversely constricted at middle,
spinigera group
Intermediate femur of female not thus constricted..............
7.(6) Intermediate femur of female flattened beneath on basal one fourth;
male usually with anterior tibia greatly dilate and excavate be-
Cl laee Ue eer ene ee wee ee peeved cE Srna collaris group
Intermediate femur of female dorso-ventrally flattened from basal
one fourth to apical one fourth; male with anterior tibia never
greatly dilate and excavate beneath... 0)... ie,
8.(7) Last genital segment terminating in an elongate spinelike process,
ainsliet. group
Last genital segment not as above.................. obesa group
ANGUSTIPES GROUP
Group characteristics: The angustipes group can be character-
ized as consisting of those species of the genus Rhagovelia in which
the pronotum of the apterous form is shorter than the length of
the eye, with the posterior margin straight or concave. The dor-
sum of the abdomen of the apterous female tapers rather evenly
to the apex. Winged forms are rare; the wings extend well beyond
the apex of the genital segments.
The following species comprise this group:
1. R. angustipes Uhler 13. R. modesta sp. nov.
2. R. bisignata Bacon 14. R. novana Drake *
8. R. callida Drake and Harris 15. R. paulana Drake
4, R. calopa Drake and Harris 16. R. plana Drake and Harris
5. R. deminuta Bacon 17. R. plumbea Uhler
6. R. evidis Bacon 18. R. rioana Drake *
7. R. festae Kirkaldy 19. R. salina (Champion )
8. R. fontanalis Bacon 20. R. spinosa Gould
9, R. hambletoni Drake and 21. R. tantilla Drake and Harris
Harris 22. R. tenuipes Champion
10. R. imitatrix Bacon 23. R. velocis Drake and Harris
ll. R. janeira Drake 94. R. versuta Drake and Harris
12. R. longipes Gould 25. R. viriosa sp. nov.
Kry to Sprcms or THE ANGusTiPES GRouP
li Posterior femur of both sexes unarmed.................00005
Posterior femur armed.at least in male...0)...c 0.00.04,
2, (1) Posterior tibia mommealin length, (ie ted dn a festae
Posterior tibia at least one third longer than posterior femur... .
3. (2) Antennal segment II distinctly shorter than III (at least in
apterous: female iz ce) une | en i longipes
Antennal segment II subequal to III......0.......4.. imitatrix
w
* See addenda at end of section on taxonomy for the description of this species which
was published after the body of this paper was written,
Srupy OF THE GENUS RHAGOVELIA Kala
4, (1) Venter of last abdominal segment of male bearing at base a
stout, slightly, curved Spine 5. ysis teh et ei ce spinosa
Base of last abdominal segment of male without such a spine... 5
5. (4) Dorsum of all abdominal segments of apterous forms with median
shining black areas (that of the first segment may be very
SA i eae eA ah lies Corbet AP Gian ais: 8 6
Dorsum of abdewven of apterous forms with no, or at most four
to six segments showing median shining black areas......... 7
6. (5) Posterior femur not projecting beyond apex of pais segments,
frontanalis
Posterior. femur projecting beyond apex of genital segments in
both sexes ... tenuipes
7. (5) Posterior femora of most male s enormously jncrassabes posterior
trochanter armed with several teeth........,.......calopa
Posterior femur of male never. enormously incrassate; posterior
Krochamber unarmed: 9a 6.8 eV Che we are es 8
8. (7) Venter of abdomen of male with a prominent carina with abdo-
men depressed to form a trough on each side of the carina on
last three or more segments...... heise 9
Venter of abdomen of male with no, or only. slightly produces ed
varina, or carina produced on last segment only. ..... on 11
9, (8) Posterior femur of apterous male extending only to apex of gen-
ital segments, Dorsum of abdomen of female concave for
last five segments. (00 ie oe gies an a viriosa
Posterior femur of apterous male,¢ ex stending well beyond apex of
genital segments. Dorsum of abdomen of female convex or
flat forslast Ave Segments. us 40 te hee) eer 10
10. (9) Posterior femur with no spine longer than diameter of posterior
tibia; posterior femur of male sharply r reduced a little before
apex -. = toda seed a7 ae Teaeren en callida
Posterior femur with at least one spine clearly longer than
diameter of posterior tibia. Posterior femur of male tapering
fO.ADEX saaie: co dition « Meuers onion: by coaeae te tenuipes
Ll. (8) Connexiva margined in brown-orange; posterior femur with
yellow to brown-yellow areas on both basal and apical por-
tions with brown to black central area........... deminuta
Jonnexival margins not orange; posterior femur with no, or only
basal light colored area ee ee ny id eae eas 12
12.(11) Antennal segment II shorter. than Tl a eee ee ee oe eee
Antennal segment Il longer than or subequal to TIT. ..........- 22
13,(12) Entire pronotum behind vertex of head yellow to orange..... 14
Only anterior half of pronotum with yellow to orange band be-
ore 15
hind. vertex af dieadinc, vey cash Bcen vapiign sone ee
14,(13) Posterior margin of pronotum sinuate; connexiva of female re-
flexed; anterior trochanter of male unarmed....... plumbea
Posterior margin of pronotum straight; connexiva of female not
reflexed; anterior trochanter of male armed with a long spur,
salina
15.(13)
16,(15)
17.(16)
18,(17)
19,(18)
20.(19)
21.(18)
22.(12)
23, (22)
24, (23)
Tue UNiversiry SCIENCE BULLETIN
Posterior femur of female unarmed, connexiva reflexed and con-
tiguous for last segment (if only apterous males are at hand
poe fiext OOUplet) iy. 6d eee en a5 janeira
Posterior femur of female armed, connexiva may be reflexed but
if 80 marging are fot COUCWING, 7 ).0 0. ri ceive a ns
Intermediate trochanters yellow to white; venter of male with a
small, blue-gray triangle on anterior margin of last abdominal
Segitient ea eet er ere Gere ett ys evidis
Intermediate trochanters black or brown; venter of male without
lich murkings as described above, ii pokey ce
Posterior tibia of both male and female armed with subequal
teeth along entire length (teeth of tibia of female may be
weak, “bitvare present)... <4. ci heels yes tenuipes
Posterior tibia of female unarmed along inner surface, posterior
tibia of male may be armed with subequal teeth on basal one
third or tiniermed, cy 1 a PE rece
Tarsal segment II of intermediate leg subequal to II (seg. II
more than four titths'as long as seg. Wl). o.0.7.....,...
Tarsal segment II of intermediate leg shorter than III (seg. II
approximately or less than four fifths as long as seg. III)....
Pro- and mesonotum flattened; venter gray-black........ plana
Pro- and mesonotum arched above; venter blue-gray..........
Coxae of intermediate legs yellow; female without armature on
POSIGMOL LOM es aes ce ain Pre er janeira
Coxae of intermediate legs dark brown to black; female with
armed” postenlor- femur. eh, a hambletoni
Posterior tibia unarmed at apex, apterous female with shining
black spots on dorsum of last two abdominal segments,
tantilla
Posterior tibia armed with a spur at apex (at least in apterous
male), female with shining black spots on dorsum of last four
Or tive abdominal segmiente
Size: Length Width
4,25 mm. apterous male 1.47 mm, apterous male
4.50 mm. apterous female 1.59 mm. apterous female
4.98 mm. winged male 1.73 mm. winged male
4.85 mm. winged female 1.79 mm. winged female
Color: General color dark brown, covered with golden pubes-
cence. Head with frons yellow and vertex brown; on some speci-
mens there is small, median, yellow spot on posterior border of
vertex. Anterior portion of pronotum with broad uninterrupted
yellow band; disk of pronotum very dark brown; apex of pronotum
yellow. Anterior basal cell of wing pruinose. Wings lighter brown
than thorax. Connexival margins broadly yellow. Dorsum of
abdomen pruinose at each side of brown, median area. Genital
Segments and venter yellow except for small area between inter-
mediate and posterior coxae. Base of antennae, all acetabulae,
all coxae, and all trochanters yellow. Anterior femur yellow for
basal four fifths; base of anterior tibia yellow beneath. Inter-
mediate femur yellow beneath for almost its entire length. Pos-
terior femur yellow with exception of brown stripe on dorsal surface.
Posterior tibia yellow beneath on basal half.
Tue Universiry Science BULLETIN
788
Structural characteristics: Pronotum of apterous. forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Apter-
ous male: anterior trochanter unarmed. Anterior tibia not dilate,
only slightly excavate on apical one third. Pronotum wider than
long (L. 77, W. 87); rounded behind. Metanotum short on mid-
line (L. 5, W. 90). Proportions of antennae: Seg. I: II: Iff: IV::
74: 44: 39: 39; of intermediate legs: Fem.: Tib.: Tars, I: Tars. III::
140: 100: 40: 64; of posterior legs: 145: 120: 12: 28. Venter with
prominent median carina extending to last segment. Venter dis-
tinctly impressed on each side of median carina. Minute, black
conical setae on base of jugum of head, proepisternum, meso-
sternum and along posterior margin of last abdominal segment at
sides. Posterior femur greatly incrassate (L. 145, W. 50); armed
after basal one third with four irregular rows of spines with three
anterior rows all small and approximately same size, posterior row
larger with three widely-spaced spines before apical third and one
closely-set row on apical one third with one or two spines larger.
Posterior tibia curved inward to middle and curved outwards from
there to apex; slightly laterally flattened; armed with small teeth
within with last two or three teeth enlarged; armed with stout
spur at apex. Apterous female: pronotum broadly rounded be-
hind (L. 79, W. 96); metanotum as in male. Proportions of an-
tennae: 68: 40: 35: 35; of intermediate legs: 183: 95: 48: 60; of
posterior legs: 125: 112: 11: 28. Venter with faint median carina
becoming evanescent on next to last segment. Minute, conical,
black setae as in apterous male. Posterior trochanter unarmed.
Posterior femur not as incrassate as in male (L. 125, W. 35); armed
before middle with one long spine, followed by row of approxi-
mately seven smaller, subequal spines to apex, also armed with an
anterior, quite irregular row of approximately five small spines on
apical one third. Posterior tibia slightly arcuate; armed within
with small, subequal teeth with apical one somewhat enlarged;
also armed at apex with straight spur. Winged forms: proportions
and armature similar to apterous forms. Winged male: pronotum
sharply pointed at apex (L. 125, W. 118); not produced into spini-
form process. Winged female: pronotum formed same as in apter-
ous male,
Comparative notes: This species may resemble Rhagovelia panda
Drake and Harris of which there are no specimens in the University
of Kansas collections. , Unfortunately the type material is in the
personal collection of Dr. C. J. Drake and not available for study.
Srupy oF THE GENUS RHAGOVELIA 789
The antennal proportions of R. horrida sp. nov. differ from those of
R. panda Drake and Harris as given in the original description, as
well as the unusual color variations on the vertex of the head and
the apex of the pronotum. The original description of R. panda
Drake and Harris is not sufficiently detailed to determine its re-
lationship with R. horrida sp. nov. This species is also close to R.
varipes Champion, but can be separated from R. varipes Champion
by the differences in the armature of the posterior femur. The
males of R. varipes Champion and R. horrida sp. nov. may be sepa-
rated by the two larger teeth at the apex of the posterior tibia, and
by the ventral carina extending to the last abdominal segment in
R. horrida sp. nov. The clasper of the male also sets the two apart
readily,
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype winged male, allomorphotype, winged fe-
male; paratypes, 1 apterous male, 1 apterous female; paramorpho-
types, 2 winged males, 6 winged females. The apterous males were
captured at Hda La Libertad, Chiapas, Mexico, Sept. 1, 1937, by H.
D. Thomas. The apterous females were captured at Guadalupe,
Chiapas, Mexico, 1-15-38, 800 m.a.s.l., Octavio Utrilla L. The
morphotype series was captured at Rincén, Guerrero, Mexico, kil.
360 s. Mex. City, 10-31-36, H. D. Thomas. All type specimens are
in the Francis Huntington Snow Entomological Collections, Univer-
sity of Kansas.
Data on distribution: In addition to the three localities mentioned
in the discussion under “Data on types”, material from the follow-
ing localities has been examined:
Mexico: Chiapas: Comitan, 1-18-36, 1800 m.a.s.l., Octavio
Utrilla L., 1 winged male.
GuaTEMALA: Sta. Lucia, 31-Jan.-05 (J. R. de la Torre-Bueno
Collection), 1 apterous male; Agua Caliente, 9-Feb.-05 (J. R. de la
Torre-Bueno Collection), 2 apterous females.
Rhagovelia jubata Bacon
P (Pl. V, fig. 6)
1948, Rhagovelia jubata Bacon, Jour. Kansas Ent. Soc., vol. 21, p. 78.
Size: Length Width
4.55 mm. apterous male 1.49 mm. apterous male
4.53 mm. apterous female 1.52 mm. apterous female
4.72 mm. winged female 1.66 mm. winged female
Color: General color red-brown, clothed with golden pubescence.
Pronotum with yellow band on anterior portion becoming pruinose
Tue Universiry SCIENCE BULLETIN
790
behind eyes. Pronotum slightly lighter in shade than metanotum.
Connexivum broadly yellow. Venter dark gray; venter of last ab-
dominal segment and genital segments of male brown with median
yellow area. Basal portion of first antennal segment, margins of all
acetabulae, all coxae, bases of anterior and posterior trochanters,
bases of anterior and posterior femora yellow. In addition anterior
femur marked with yellow line running full length of ventral sur-
face, also posterior femur conspicuously striped with anterior and
posterior-ventral surfaces yellow for their full length. The dorsum
of fifth abdominal segment of female darker than other abdominal
segments.
Structural characteristics: Pronotum of apterous forms covering
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex.
Apterous male: anterior trochanter unarmed. Anterior tibia flat-
tened within for apical one third; not dilate. Pronotum wider than
long (L. 68, W. 85); metanotum much wider than long (L. 10, W.
95). Proportions of antennae: Seg. I: II: III: [V:: 84: 47: 44: 42;
of intermediate legs: Fem.: Tib.: Tars. II: Tars. III:: 175: 80: 48:
64; of posterior legs: 152: 136: 8: 26. Abdomen tapers evenly to
apex. Connexiva semivertical. Dorsum with scattered long hairs.
Venter with median carina thickly-set with long hairs; ventral carina
extending to last abdominal segment. Minute, black, conical setae
on base of jugum of head, proepisternum, and in patch at side of
last abdominal segment. Venter of last abdominal segment some-
what excavate on posterior one half. Posterior trochanter armed
with one to several very short, and one to three longer teeth. Pos-
terior femur greatly incrassate (L. 150, W. 50); armed on basal
one third with row of very small teeth ending in one long spine,
followed by two irregular rows of about seven smaller spines to
apex. Posterior tibia slightly arcuate; armed within with subequal
teeth except for apical tooth which is elongate and in apposition
to longer teeth on trochanter; armed at apex with long spur. Apter-
ous female: pronotal proportions same as in male. Abdomen gradu-
ally tapers posteriorly. Connexiva flat. Proportions of antennae:
75: 45: 43: 39; of intermediate legs: 155: 111: 46: 64; of posterior
legs: 185: 117: 8: 26. Minute, black, conical setae as in male.
Posterior trochanter unarmed; posterior femur moderately incras-
sate (L. 135, W. 35) and armed with from none to four very small
teeth and one small spine on basal one third, then at middle
with one large spine and two irregular rows of approximately seven
Srupy oF THE GENUS RHAGOVELIA 791
smaller spines to apex. Posterior tibia very slightly arcuate and
armed with equal teeth with elongate spine at apex. Winged fe-
male: proportions and armature similar to apterous female. Apex
of pronotum set with long dark hairs; not produced into a spiniform
process. Winged male: unknown.
Comparative notes: This species resembles R. palea sp. nov. R.
jubata Bacon can be separated from R. palea sp. nov. by the color of
the legs which are dark brown to black in R. jubata Bacon with
conspicuous stripes of yellow on the anterior and posterior femora
while the legs of R. palea sp. nov. are yellow-brown. The venter of
R. jubata Bacon is brown-black, while the venter of R. palea sp. nov.
is light yellow-brown. The posterior femora of all forms are also
quite different in armature.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; allomorphotype, winged female; paratypes, 40 apterous males,
24 apterous females. Described from specimens from Aguaitia
Dept. Loreto, Pert, collected by F. Woytkowski, Oct. 1, 1946. All
type specimens are in the Francis Huntington Snow Entomological
Collections, University of Kansas.
Data on distribution: In addition to the type series, specimens
from the following localities have been examined:
Peru: Loreto Dept., Aguaitia, [X- 19-46, F. Woytkowski, 17 apter-
ous males, 13 apterous females, 2 nymphs; Loreto Dept., Roqueron
del Padre, Abad Cordillern, VIII-3-46, F. Woytkowski, 1 apterous
male; Loreto Dept., Santa Elena, Roqueron Padre, Abad, VIII-8-46,
F. Woytkowski, 2 apterous males, 1 apterous female.
Rhagovelia nitida Bacon
(Pl. V, fig. 7)
1948, Rhagovelia nitida Bacon, Jour, Kansas Ent, Soc., vol. 21, p. 79.
Size: Length Width
4,89 mm. apterous male 1.49 mm. apterous male
4.39 mm. apterous female 1.49 mm. apterous female
4,52 mm. winged male 1.82 mm. winged male
4.62 mm. winged female 1.80 mm. winged female
Color; General color brown, with prominent yellow markings,
clothed with short golden pubescence. Pronotum tan with unin-
terrupted yellow band on anterior one fourth; slightly lighter line
Tunning longitudinally down middle of pronotum. Metanotum and
dorsum of first abdominal segment darker than pronotum. Re-
mainder of dorsum of abdomen tan to yellow with narrow darker
areas at juncture of each of segments. Venter, antennae, and geni-
Tue Universrry Science BULLETIN
792
tal segments practically unicolorous; legs slightly darker. Posterior
femur slightly lighter at base and for three fourths its length ven-
trally. Wings red-brown; veins darker, not prominent.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed. Anterior tibia not dilate;
slightly excavate on apical third. Pronotum wider than long (L. 75,
W. 90); metanotum much wider than long (L. 10, W. 94). Pro-
portions of antennae: Seg. I: II: III: IV:: 77: 49: 52: 44; of inter-
mediate legs: Fem.: Tib.: Tars. II: Tars. III:: 150: 117: 45: 60;
of posterior legs: 150: 163: 15: 30. Angle of taper of connexiva in-
creased for last three segments. Ventral carina sharply elevated
between coxae of posterior legs with depression on each side, be-
coming evanescent on fourth segment. Genital segments usually
retracted partially into last abdominal segment on at least some
specimens. Posterior trochanter thickly set with long hairs; un-
armed. Posterior femur greatly incrassate (L. 150, W. 57); ventral
surface flattened and beset on basal one third with long hairs; armed
on basal half with three irregular rows of small, dark teeth and one
long stout isolated spine on posterior margin of ventral surface
of leg slightly beyond basal third, armed on apical half of femur
with approximately six small, stout spines set in two irregular rows.
Posterior tibia slightly arcuate; armed within with two rows of black
teeth, teeth at apical three fourths tending to be slightly enlarged;
armed at apex with brown spur. Apterous female: anterior tibia
formed as in apterous male. Pronotum wider than long (L. 70, W.
90); metanotum much wider than long (L. 7, W. 92). Proportions
of antennae: 70: 44: 45: 48; of intermediate legs: 130: 108: 42:
58; of posterior legs: 112: 147: 11: 26. Dorsum of abdomen taper-
ing evenly to apex. Connexiva semivertical. Genital segments
small. Posterior trochanter unarmed. Posterior femur only mod-
erately incrassate (LL. 112, W. 24) and armed at apical one third
with one small spine followed by three decreasing spines to apex.
Posterior tibia seemingly unarmed. Winged forms: proportions
and armature similar to apterous forms. Pronotum not produced
into spiniform process.
Comparative notes: This species resembles R. sinuata Gould.
R. nitida Bacon can be separated from R, sinuata Gould by the
unarmed posterior trochanter of the male and the unarmed pos-
terior tibia of the female. The male clasper.is quite distinctive and
Srupy oF THE GENUS RHAGOVELIA 793
easily sets R. nitida Bacon apart from all other closely related
Species.
Data on types: Holotype, apterous male; allotype apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 4 apterous males, 3 apterous females; paramorpho-
types, 3 winged females. Described from Jamaica, B.W.I. All
type specimens are in the University of Kansas Collections.
Data on distribution: In addition to the type series, specimens
from the following localities have been examined:
Brrriso West Inpres: Jamaica: St. Andres, Shooters Hill, e. 1800
ft., 8-xii-46, G. B. Thompson, 1 winged female; St. Thomas, XI-14-
46, G. B. Thompson, 2 apterous males, 1 apterous female, 4 nymphs;
St. Andreas, IV-17-47, G. B. Thompson, 7 apterous males, 8 ap-
terous females, 15 nymphs.
Rhagovelia ornata Bacon
(Pl. V, fig. 8)
1948. Rhagovelia ornata Bacon, Jour. Kansas Ent. Soc., vol. 21, p. 80.
Size: Length Width
5.98 mm. apterous male 1.63 mm. apterous male
6.00 mm. apterous female 1.63 mm. apterous female
Color: General color red-brown; practically unicolorous. Last
two segments of antennae and last tarsal segments of intermediate
legs darker brown. Pronotum without lighter band at anterior
Margin,
Structural characteristics: Pronotum of apterous forms covers
Mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed. Anterior tibia not dilate,
slightly flattened on apical one fifth. Pronotum wider than long
(L. 90, W. 103); metanotum much wider than long (L. 11, W. 115).
Proportions of antennae: Seg. I: II: III: I1V:: 107: 55: 56: 55; of
intermediate legs: Fem.: Tib.: Tars. II: Tars. IlI:: 200: 150: 62:
75; of posterior legs: 192: 178: 41: 20. Margins of connexiva taper
slightly to last two segments which have an increased angle of
taper. Ventral carina pronounced between posterior coxae where
segment is depressed on each side, becoming evanescent on next
to last segment. Posterior trochanter armed with approximately
twelve small, knoblike teeth. Posterior femur greatly incrassate
(L. 192, W. 62) on most specimens; armed on basal half with one
row of approximately eighteen equally spaced spines which are
794 Tue Universiry ScreENCE BULLETIN
graduated in ‘size from small teeth on basal area to small spines at
middle. The apical half of femur is armed with one isolated larger
spine on posterior margin at apical third in those specimens on
which posterior femur is greatly incrassate. This isolated spine is
not conspicuous in specimens with only moderately incrassate pos-
terior femora. Also with two smaller rows of spines, the posterior
row consisting of seven spines and the anterior row of six spines
decreasing in size to apex. None of spines of posterior femur
greatly enlarged. Posterior tibia curved inward at first and out-
ward beyond apical one third; armed with two rows of small teeth,
with one stout, greatly enlarged tooth at apical one third followed by
two slightly enlarged and several smaller teeth; armed with stout
spur at apex. Apterous female: anterior tibia not dilate, slightly
flattened on apical one fourth. Proportions of antennae: 100: 51:
55: 50; of intermediate legs: 185: 140: 57: 73; of posterior legs:
160: 176: 18: 36. Abdomen with only slight taper from base to
apex. Ventral carina distinct on basal segments, becoming evanes-
cent on last three abdominal segments. Posterior trochanter armed
with approximately two small dark teeth. Posterior femur only
slightly incrassate (L. 160, W. 30) and armed at apical one third
with one small spine followed by four or five smaller, rapidly de-
creasing spines to apex. Posterior tibia straight; armed on basal
half with several small teeth; armed with spur at apex. Winged
forms: unknown.
Comparative notes: This species resembles R. robusta Gould.
The males of R. ornata sp. nov. can be separated from those of R. ro-
busta Gould by the location of the enlarged spine of the posterior
tibia which is at the apical one third in R. ornata sp. nov. while the
enlarged spine of the posterior tibia of R. robusta Gould is at the
apical one fifth. The females can be separated by the armature of
the posterior femur which has the long spine after the apical one
third in R. ornata sp. nov. while the long spine of the posterior femur
of R. robusta Gould is at the middle. The absence of minute, coni-
cal, black setae on the lower margin of the jugum will set R. ornata
sp. nov. apart from the other species which it resembles, R. crassipes
Champion and R. varipes Champion.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 32 apterous males, 59 apterous females. Described
from series collected in Bolivia, Miguelito, May, 1938, A. M. Olalla.
All type specimens are in the Francis Huntington Snow Entomo-
logical Collections, University of Kansas.
Data on distribution: Known only from the type series.
Srupy oF THE GENUS RHAGOVELIA 795
Rhagovelia palea sp. nov.
(Pl. V, fig. 9)
Size: Length Width
4.05 mm. apterous male 1.21 mm. apterous male
4.26 mm. apterous female 1.42 mm. apterous female
4.31 mm. winged male 1.60 mm. winged male
4.66 mm. winged female 1.74 mm. winged female
Color: General color yellow-brown. The anterior margin of pro-
notum yellow behind vertex of head becoming pruinose behind eyes.
Head and pronotum tan in apterous forms; posterior portion of
pronotum tan in winged forms. Venter yellow-brown. Base of
anterior femora, base of first antennal segments, all coxae and tro-
chanters yellow-white. Metanotum and dorsum of abdominal seg-
ments red-brown. Wings brown, veins prominent, anterior basal
cell of wing pruinose.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Apter-
ous male: anterior trochanter unarmed; anterior tibia slightly dilate,
flattened on apical one third. Pronotum wider than long (L. 57,
W. 76); metanotum much wider than long (L. 12, W. 80), Propor-
tions of antennae: Seg. I: II: III: IV:: 70: 39: 40: 38; of intermedi-
ate legs: Fem.: Tib.: Tars. II: Tars. II:: 145: 105: 40: 53; of pos-
terior legs: 117: 104: 9: 24. Connexiva tapers to apex, angle of taper
increasing for last three segments. Inconspicuous, hairy ventral
carina extending to last abdominal segment. Minute conical, black
setae on proepisternum and in patch on side of last abdominal
Segment. Posterior trochanter armed with from two to six very
small teeth and one larger tooth. Posterior femur moderately incras-
sate (L. 117, W. 35); armed on basal two fifths with one row of small,
black, knoblike teeth, from basal two fifths to apex with two rows of
spines. The anterior row of spines begins just before middle with
one long spine followed by eight to ten smaller, subequal spines;
the posterior row begins after middle with one long spine displaced
toward posterior margin, then with approximately six gradually de-
creasing spines to apex. Posterior tibia slightly arcuate; armed
with two rows of teeth with apical tooth usually noticeably enlarged;
armed at apex with prominent spur. Apterous female: pronotum
and metanotum formed much as in male. Proportions of antennae:
70: 89: 87: 87; of intermediate legs: 135: 104: 40: 55; of posterior
legs: 117: 100: 8: 24. Posterior trochanter unarmed, or may be
armed with one tooth in some specimens. Posterior femur moder-
ately incrassate (L. 117, W. 30) and armed with one long spine at
796 Tue Universiry Science BULLETIN
middle followed by five or six decreasing spines to apex. Posterior
tibia very slightly arcuate; armed with two rows of equal teeth and
stout spur at the apex. Dorsum of first abdominal segment thickly
beset with dark hairs on posterior portion. Winged forms: propor-
tions and armature same as for apterous forms. Tip of pronotum
in both sexes. thickly beset with long dark hairs. Pronotum not
produced into a spiniform process. Wings just covering apex of
genital segments.
Comparative notes: This species resembles R. jubata Bacon. R.
palea sp. nov. can be separated from R. jubata Bacon by the light
yellow-brown venter and yellow-brown legs of all forms. The arma-
ture of the posterior femur of all forms is also quite distinct for
each of the two species.
Data on types: Holotype, apterous male; allotype, apterous
female; holomorphotype, winged male; allomorphotype, winged
female; paratypes, 12 apterous males, 11 apterous females; para-
morphotypes, 2 winged females. Described from series labeled:
“Peri, Dept. Ayacucho, Prov. La Mar, Sivia, Jungle 790 m. a. s. L.,
Bks. Apurimac riv., June 15-28-1941, F. Woytkowski, No, 4212.”
All type specimens are in the Francis Huntington Snow Entomologi-
cal Collections, University of Kansas.
Data on distribution: In addition to the type series, specimens
from the following localities have been examined:
Botivra: Road between Tedos Santos and Palmer, R. Chapare,
March, 1938, A. M. Olalla, 10 apterous males, 12 apterous females;
R. Beni Cachuela, Esperanza, 9-37, A. M. Olalla, 2 apterous males,
1 winged male, 1 winged female.
Peru: Dept. Ayacucho, Prov. La Mar, Sivia, jungle, 790 m.a.s.L.,
stagnant pools, June 24-30, 1941, F. Woytkowski, No. 426, 3 apterous
males, 1 apterous female; Dept. Ayacucho, Prov. La Mar, Sivia,
jungle, 790 m.a.s.l., stagnant boggy pools, June 18-19, 1941, F.
Woytkowski, No. 428, 1 apterous male, 2 apterous females; Dept.
Ayacucho, Prov. La Mar, Sivia, jungle brooks, 790 m.a.s.l., June
24-30, 1941, F’. Woytkowski, No. 425, 3 apterous males; Vic. San
Pedro, 90 m.a.s.l., pools and ponds, May 15-19, 1935, F. Woyt-
kowski, 2 apterous males, 38 apterous females; Vic. San Pedro, 900
m.a.s.l., pools in jungle, May 15-29, 1935, F. Woytkowski, 3 ap-
terous males, 1 winged male, 4 apterous females; Vic. Sani Beni,
840 m.a.s.l., tiny brook in open land, Oct. 31, 1935, F, Woytkowski,
2, apterous males, 2 apterous females; Vic. Sani Beni, 840 m.a.s.L,
brooks and pools of Sani Beni, Aug. 31, 1935, F. Woytkowski, Field’
Srupy oF THE GENUS RHAGOVELIA 797
Note 3560, 1 apterous male, 2 apterous females; Vic. Sani Beni, 840
m.a.s.l., brook on open cultivated land, Oct. 12, 1935, F. Woyt-
kowski, Field Note 3566, 3 apterous males, 3 apterous females; Vic.
Rio Negro, 790 m.a.s.l., River R. Negro, Nov. 4, 1935, F. Woyt-
kowski, Field Note 3550, 8 apterous males, 1 winged male, 5 apter-
ous females; Vic. Rio Negro, 790 m.a.s.l., shady jungle brook, bed
of clay, Oct. 29, 1935, F. Woytkowski, Field Note 3564, 2 apterous
males, 5 apterous females; Vic. Rio Negro, 790 m.a.s.l., in R.
Negro, Nov., 1985, F. Woytkowski, Field Note 8558e, 5 apterous
males, 1 apterous female; Dept. Hudnuca, Vic. of Afilador, jungle,
800 m.a.s.l., June 3-7, 1937, F. Woytkowski, No, 3765, 7 apterous
males, 7 apterous females; Aguaitia, Dept. Loreto, Sept. 1, 1946,
F. Woytkowski, 9 apterous males, 5 apterous females; Aguaitia
Dept. Loreto, Sept. 19, 1946, F. Woytkowski, 3 apterous males, 1
winged male, 5 apterous females; Satipo, Nov., 1942, Pedro Papr-
zycki, 83 winged males, 2 apterous females, 2 winged females; Satipo,
Dec., 1942, Pedro Paprzycki, 86 apterous males, 18 winged males,
27 apterous females, 13 winged females.
Rhagovelia panda Drake and Harris
1935, Rhagovelia panda Drake and Harris, Proc. Biol. Soc. Washington, vol.
48, p. 193.
This species is not represented in the material in the Francis
Huntington Snow Entomological Collections, University of Kansas.
The original description is quoted below:
“Moderately large, blackish, thickly clothed with golden pubes-
cence. Antennal formula, 54: 34: 30: 26; first and second segments
with a few bristly hairs, the first with basal half yellowish. Ros-
trum extending on basal portion of mesosternum.
“Apterous form.—Pronotum long, indistinctly carinate, broadly
rounded behind; in front broadly yellowish.
“Winged form.—Pronotum broadly yellowish in front as in ap-
terous form, indistinctly carinate, triangularly produced behind, the
humeri prominent.
“Male.—Hind femora strongly developed, in some individuals
enormously enlarged, armed beneath with numerous irregularly
placed, black teeth of various sizes. Coxae and trochanters, the
femora beneath and base and sides within yellowish white, Inter-
mediate tarsi with segment three longer than two (35: 47). Hind
tibia nearly straight or strongly curved, denticulate beneath, armed
at apex with straight spur and before apex usually with three large,
stout teeth. Connexiva yellowish white, the narrow margin and
798 Tue Universtry SCIENCE BULLETIN
the broad basal stripe above blackish. Venter with a very promi-
nent median carina extending to last segment, distinctly impressed
on each side of carina; last segment yellowish, scarcely excavated
behind. Clasper long, narrow, slightly curved, faintly narrowed
towards apex.
“Female.—Color markings about as in male. Hind femora mod-
erately swollen, with a large curved tooth just before middle, thence
to apex with two rows of shorter black teeth; hind tibia nearly
straight, finely denticulated.
“Length, 4.90-6.00 mm.; width, 2.00 mm.
“Holotype, apterous male; allotype, winged female, and paratypes,
several apterous and macropterous males and females, Chiquimula,
Guatemala, June, 1930; authors’ collection.
“The authors desire to express their appreciation to Mr. W. E.
China of the British Museum for comparing this species with the
type of R. femoralis Champion. The clasper is longer and narrower
at the base than in femoralis. The last segment of the intermediate
tarsi is slightly longer than the second. The venter, in some speci-
mens, is mostly yellowish, also the coxae and trochanters. The
hind trochanters are armed with two or three short teeth in the
male.”
Comparative notes: This species resembles R. horrida sp. nov. As
there are no specimens of R. panda Drake and Harris available for
study it has been necessary to attempt to determine this species by
the original description. There are differences in color pattern and
significant proportional differences in the antennae, which in R. hor-
rida sp. nov. are formed with the last two segments equal in length
while in R. panda Drake and Harris segment IV is definitely shorter
than III. It has been thought best to consider these two separate
species. A fresh evaluation of these two species can be made when
Dr. Drake’s type material which is in his personal collection be-
comes available for study.
Rhagovelia perfidiosa Bacon
(Pl. V, fig. 10)
1948. Rhagovelia perfidiosa Bacon, Jour, Kansas Ent. Soc., vol. 21, p, 81.
Size: Length Width
2.98 mm. apterous male 1.16 mm, apterous male
8,32 mm. apterous female 1.88 mm. apterous female
8.52 mm. winged male 1.46 mm. winged male
8.52 mm. winged female 1.50 mm. winged female
Color: General color brown, clothed with golden pubescence.
Pronotum with yellow band on anterior one fifth, becoming pruinose
Srupy OF THE Genus RHAGOVELIA 799
behind eyes; posterior lateral margins and median longitudinal line
orange. Dorsum of first abdominal segment blue-gray except for
median dark brown area; fourth and fifth abdominal segments blue-
gray with smaller median dark brown areas; sixth abdominal seg-
ment brown with yellow median area; seventh abdominal segment
and genital segments yellow. Connexiva broadly yellow. Venter
blue-gray. Venter of last abdominal segment and genital segments
yellow. Base of antennae, margins of all acetabulae, all coxae and
trochanters, base of anterior and posterior femora, and total length
of dorsal and ventral surfaces of intermediate and posterior femora
yellow. Basal half of wings lighter than apical half. Anterior basal
cell of wing pruinose.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex, Posterior tibia without sickle-shaped spur to apex. Ap-
terous male: anterior trochanter unarmed. Anterior tibia slightly
dilate and flattened within on apical one half. Pronotum wider than
long (L. 50, W. 67); metanotum much wider than long (L. 7, W.
57). Proportions of antennae: Seg. I: I: TIT: IV:: 50: 29: 29: 30;
of intermediate legs: Fem.: Tib.: Tars. II: Tars. III:: 110: 80: 29:
47; of posterior legs: 88: 78: 4: 18. Abdomen tapers more sharply
to apex on last three segments. Venter without median carina.
Minute, conical, black setae on base of jugum of head, and on
proepisternum. Posterior trochanter armed with from one to nine
very small knoblike teeth with one larger tooth near apex. Pos-
terior femur greatly incrassate (L. 88, W. 30). Armed on basal
One third with a row of small teeth; after basal one third with
two irregular rows of spines. The anterior row of spines begins
at basal one third with one long spine followed by three smaller
Spines followed by another long spine offset toward anterior margin
and three smaller spines to apex. The posterior row begins approxi-
mately at middle with one large spine followed by four decreasing
Spines to apex. Posterior tibia slightly arcuate and armed with two
rows of equal teeth throughout; armed at apex with a long stout
Spur, Apterous female: proportions of antennae: 55: 28: 80: 30; of
intermediate legs: 110: 83: 34: 51; of posterior legs: 92: 80: 5: 22.
Abdomen tapers evenly to apex. No median ventral carina. Mi-
nute, black, conical setae as in male. Posterior trochanter unarmed.
Posterior femur greatly incrassate (L. 92, W. 80) and armed just
before middle with one long spine followed by approximately five
smaller spines to apex; also armed with row of four small, equal
800 Tur Universiry ScrENCE BULLETIN
spines on anterior margin of ventral surface beginning at apical
one third and continuing to apex. Posterior tibia slightly arcuate
and armed with two rows of teeth that decrease in size to apex;
apex armed with long, stout spur. Winged forms: Proportions and
armature same as for apterous forms. Apex of pronotum thickly
beset with long, dark hairs; not continued into a spiniform process.
Comparative notes: This species resembles R. amazonensis Gould
and R. williamsi Gould. R. perfidiosa Bacon can be separated from
either of these species by the presence of the minute, conical, black
setae on the inner portion of the proepisternum of all forms. The
armature of the posterior femur is distinct from either of the above
mentioned species, as is the clasper of the male genitalia.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 81 apterous males, 85 apterous females; paramor-
photypes, 5 winged males, 14 winged females. Described from a
series collected in Brazil, A. M. Olalla, No. 379. All type specimens
are in the Francis Huntington Snow Entomologist Collections, Uni-
versity of Kansas.
Data on distribution: In addition to the type series, specimens
from the following localities have been examined:
Brazit: Rio Negro, Mandos Region, Oct., 1935, A. M. Olalla, 3
apterous males, 2 winged males, 7 apterous females; Rio Purts,
Castanha Region, A. M. Olalla, Sept., 1935, 1 winged female; Vic.
Jofio Pesson (So Phelipe), River Jurud, 7-10-, 9-20-36, A. M. Olalla,
No. 875, 21 apterous males, 1 winged male, 24 apterous females, 2
winged females; State of Para, Lago Grande, Feb, ’39, A. M. Olalla,
19 apterous males, 27 apterous females.
Rhagovelia relicta Gould
(Pl. V, fig. 11)
1931. Rhagovelia relicta Gould, Kansas Univ. Sci. Bull. vol. 20, p. 39.
1935. Rhagovelia relicta Drake and Harris, Proc, Biol, Soc. Washington, vol.
48, p. 36 (record color variations),
Size: Length Width
3.80 mm. apterous male 1.16 mm. apterous male
8.85 mm. apterous female 1.3838 mm. apterous female
Color: General color black, clothed with golden pubescence. An-
terior band of pronotum broadly yellow becoming somewhat pru-
inose behind eyes. Dorsum of abdomen with median, black, shin-
ing spots on last two segments. Margins of connexiva yellow.
Venter blue-black to brown-black. Venter of last abdominal seg-
Srupy oF THE GENUS RHAGOVELIA 801
ment brown beneath. Base of antennae, margins of all acetabulae,
all coxae, anterior and posterior trochanters and prominent stripes
on anterior and posterior femora yellow.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed. Anterior tibia only
slightly dilate and flattened within toward apex. Pronotum rather
sharply rounded behind (L. 67, W. 75). Metanotum short on mid-
line (L. 4, W. 78). Proportions of antennae: Seg. I: I: III: [V::
60: 30: —: —; of intermediate legs: Fem.: Tib.: Tars. If: Tars. HI::
121: 90: 383: 55; of posterior legs: 105: 93: 7: 22. Abdomen tapers
rather evenly to apex. Connexiva semivertical. Venter without
median carina. Venter of last abdominal segment roundly excavate
beneath on posterior one half. Minute, black, conical setae on base
of jugum of head, and on inner surface of proepisternum. Posterior
trochanter armed with from one to three small, subequal teeth.
Posterior femur moderately incrassate (L. 105, W. 33); armed on
basal one third with one row of small, knoblike teeth; apical two
thirds armed with two rows of teeth, the posterior row with one
long spine after basal one third followed by two short spines, one
long spine somewhat offset to posterior margin, followed by four
or five small spines to apex, the anterior row with six to eight small
subequal spines in an irregular row beginning after basal one third
and continuing to apex. Posterior tibia almost straight; armed
within with two rows of subequal teeth with tooth at apical one fifth
enlarged; armed at apex with stout spur. Apterous female: anterior
leg as in male. Pronotum broadly rounded at apex (L. 68, W. 80).
Metanotum short on midline (L. 7, W. 84). Proportions of anten-
nae: 60: 82: —: —; of intermediate legs: 115: 85: 35: 56; of pos-
terior legs: 97: 90: 5: 26. Dorsum of abdomen tapers rather evenly
to apex. Connexiva vertical. Minute, black, conical setae as in
male. Posterior trochanter unarmed. Posterior femur moderately
incrassate (L. 97, W. 24); armed beyond middle with one long
spine followed by two or three small spines. Posterior tibia straight;
armed within with small teeth with those of basal one half stouter;
armed at apex with spur. Winged forms: unknown.
Comparative notes: This species resembles R. femoralis Cham-
pion. R. relicta Gould can be separated from R. femoralis Cham-
pion by the nature of the minute, black setae of the proepisternum
which are conical in R. relicta Gould while being bristlelike and
802 Tue Universiry Science BULLETIN
more elongate in R. femoralis Champion. R. relicta Gould is also’
related to R. robusta Gould. R. relicta Gould can be separated from
R. robusta Gould by the nature of the armature of the posterior
fermmur and the color of the intermediate trochanter which is black in
R. relicta Gould, yellow in R. robusta Gould. Drake and Harris
(1935 a; p. 86) record color variations and indicate that R. relicta
Gould may be identical with R. robusta Gould. It has been thought
best to consider these two species as distinct until additional speci-
mens will permit a study of the limits of variation.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; both collected in Campinas, Estado de Sao Paulo, Brazil,
3-10-24, by F. X. Williams. These types are in the Francis Hunting-
ton Snow Entomological Collections, University of Kansas.
Data on distribution: Recorded from Brazil. Represented in the
Francis Huntington Snow Entomological Collections only by the’
type series.
Rhagovelia robusta Gould
(Pl. V, fig. 12)
1931. Rhagovelia robusta Gould, Kansas Univ. Sci. Bull., vol. 20, p. 41.
1933. Rhagovelia robusta Gould, Ann. Ent. Soc. America, vol. 26, p. 469.
1935. Rhagovelia sinuata calcaris Drake and Harris, Proc, Biol, Soc, Washing- -
ton, vol. 48, p. 86 (describes from Pert),
Size: Length Width
4.45 mm. apterous male 1.40 mm. apterous male
4.75 mm. apterous female 1.50 mm. apterous female
Color: General color red-brown varying in shade from light to
dark. On dark red-brown specimens anterior band of pronotum
yellow as is indefinite median line on disc of pronotum; median
spot on dorsum of last abdominal segment brown; margins of con-
nexiva yellow-brown; also base of posterior femur yellow. On
lighter red-brown specimens anterior band of pronotum not set off
by definite markings; median spot on dorsum of last abdominal
segment yellow; margins of connexiva yellow; base of antennae,
margins of all acetabulae, all coxae, anterior and posterior tro-
chanters, and base of anterior and posterior femur yellow to white.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Apter-
ous male: anterior trochanter unarmed. Anterior tibia slightly
dilate and flattened within. Pronotum rounded behind (L. 73, W.
83). Metanotum short on midline (L. 7, W. 86). Proportions of
Srupy OF THE GENUS RHAGOVELIA 803
antennae: Seg. I: II: II: IV:: 75: 42: 43: 37; of intermediate legs:
Fem.: Tib.: Tars. II: Tars. II:: 150: 107: 41: 60; of posterior legs:
185: 120: 9: 27. Abdomen tapers slightly at first with angle of taper
increasing for last three segments. Connexiva semivertical. Venter
of abdomen with more or less distinct median carina extending to
last segment. Venter of last abdominal segment roundly excavate
on posterior one half beneath. Black, minute, conical setae on base
of jugum of head and on inner surface of proepisternum. Posterior
trochanter armed with two or three subequal, small teeth. Posterior
femur greatly incrassate (L. 135, W. 55); armed on basal one fourth
with one row of five or six small teeth; apical three fourths armed
with three to five irregular rows of subequal spines; armed near
posterior margin of ventral surface just beyond middle with one
long spine with three to five smaller spines in rough semicircle on
posterior margin of ventral surface of femur. Posterior tibia arcuate,
armed within with two rows of teeth with one tooth at approximately
apical one seventh greatly enlarged; armed at apex with stout spur.
Apterous female: anterior leg as in male. Pronotum rounded be-
hind (L. 75, W. 87). Metanotum as in male (L. 6, W. 89). Pro-
portions of antennae: 82: 46: 47: 38; of intermediate legs: 118: 95:
87: 57; of posterior legs: 102: 100: 7: 238. Dorsum of abdomen
tapers evenly to apex. Connexiva semivertical. Minute, black,
conical setae as in male. Posterior trochanter unarmed, Posterior
femur not as incrassate as in male (L. 102, W. 30); armed at middle
with one long spine followed by eight or ten smaller, rapidly de-
creasing spines to apex. Posterior tibia straight; armed within with
two rows of small spines with those on basal one half stouter; armed
at apex with slender spur. Winged forms: unknown.
Comparative notes: This species resembles R. relicta Gould. R.
robusta Gould can be separated from R. relicta Gould by the differ-
ence in armature of the posterior femur, which in R. robusta Gould
is more strongly developed and of a different arrangement in both
the males and females of the two species. The intermediate tro-
chanter of R. relicta Gould is black to dark brown on the specimens
at hand while that of R. robusta Gould is yellow to orange. There
is some variation in incrassateness of the posterior femur, and on
those specimens which has a less incrassate posterior femur the
armature is reduced but of the same general arrangement. Rhago-
velia sinuata calcaris Drake and Harris as represented by paratypes
in the Francis Huntington Snow Entomological Collections is in-
distinguishable from R. robusta Gould, and the name R. sinuata cal-
caris Drake and Harris becomes a synonym of R. robusta Gould.
Tue University SCIENCE BULLETIN
804.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 1 apterous male, 4 apterous females. The above
type series is in the Francis Huntington Snow Collections, Univer-
sity of Kansas. Collected in Paraguay.
Data on distribution: In addition to the type series, specimens
have been examined from the following localities:
Peru: Vic. Pampa Hermosa, 1600 m.a.s.l., stream pools, May
1-5, 1935, F. Woytkowski, 25 apterous males, 15 apterous females;
Rio Paucartambo, Quiroz, Dec. 1933 (paratypes of R. sinuata cal-
caris Drake and Harris), 1 apterous male, 1 apterous female; Dept.
Huanuco, Vic. Leonpampa, jungle, 800 m.a.s.l., Dec. 11-30, 1987,
No. 3811, F. Woytkowski, 8 apterous males, 4 apterous females;
bought from Juan D. Rivas, 3 apterous males, 8 apterous females;
Vic. San Pedro, 900 m.a.s.l., muddy ponds, May 15-19, 1935, F.
Woytkowski, 1 apterous male, 3 apterous females; Vic. Pampa Her-
mosa, 1600 m.a.s.l., slow muddy brook, May 1-5, 1935, F. Woytkow-
ski, 13 apterous males, 7 apterous females.
Rhagovelia scitula sp. nov.
(Pl. V, fig. 18)
Size: Length Width
3.83 mm. apterous male 1.20 mm. apterous male
4.16 mm. apterous female 1.36 mm. apterous female
Color: General color dark brown, covered with golden pubes-
cence. Anterior one fifth of pronotum yellow. Margins of connex-
iva and spot on dorsum of last three (male) or four (female) ab-
dominal segments yellow-brown. Venter with mesosternum tan,
metasternum and first three or four abdominal segments brown-
black, remaining segments progressively lighter brown, first genital
segment yellow, except at apex, second genital segment brown.
Base of antennae, all coxae and trochanters, and bases of anterior
and posterior femora yellow when viewed from above; all femora
yellow in ventral view.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed, anterior tibia not dilate,
very slightly excavate for anterior half. Pronotum wider than long
(L. 65, W. 82) and covering mesonotum. Metanotum (L. 3, W.80)
almost hidden in middle by apex of pronotum. Proportions of an-
tennae: Seg. I: II: III: 1V:: 64: 34: —: —; of intermediate legs:
Srupy oF THE GENUS RHAGOVELIA 805
Fem.: Tib.: Tars. II: Tars. III:: 127: 938: 38: 54; of posterior legs:
108: 97: 8: 26. Abdomen tapering evenly to apex. Ventral carina
present, becoming evanescent on last two or three abdominal seg-
ments. Minute, conical, black setae on jugum of head, proepister-
num, mesosternum and venter of abdomen with exception of last
segment on which minute setae are concentrated into one row along
each side of posterior edge of segment. Posterior trochanter armed
with one long spine and two or three very small, knoblike teeth.
Posterior femur greatly incrassate (L. 108, W. 37) and flattened
beneath. Armed on basal one half with median row of very small,
knoblike teeth, apical one half armed with two lateral rows of
spines, the anterior row starting just before middle with approxi-
mately four small, knoblike teeth followed by one bent, moderate-
sized spine just before apex, the posterior row starting at approxi-
mately middle with one moderate-sized spine followed by three or
four smaller spines increasing in size toward the apex. Posterior
tibia arcuate and denticulate within, apical three or four teeth being
larger; armed with curved spur at apex. Apterous female: pro-
notum as in male (L. 68, W. 90) covering mesonotum and most of
metanotum. Proportions of antennae: 67: 85: 87: 35; of inter-
mediate legs: 125: 94: 87: 55; of posterior legs: 108: 100: 9: 25.
Abdomen tapers evenly to apex as in male. Minute, conical, black
setae as in male except not as apparent on last abdominal segment
at sides. Posterior trochanter unarmed. Posterior femur not as in-
crassate as in male (L. 108, W. 26); armed on basal one half with
one or two small teeth, apical one half armed with two rows of
spines, the anterior row begins at apical one fourth and consists of
three or four small spines decreasing in size to apex, the posterior
row starts just beyond middle with one long, bent spine followed by
four or five smaller, decreasing spines to apex. Posterior tibia al-
most straight and denticulate within, teeth more closely set on basal
two thirds; armed at apex with straight spur. Winged forms: un-
known.
Comparative notes: This species resembles R. whitei (Breddin)
from which it can be separated by the yellow brown spots on the
dorsum of the last three (male) or four (female) abdominal seg-
ments in R. scitula sp. nov. Also the armature of the posterior femur
of both male and female is somewhat different. The posterior tibia
of the apterous male of R. whitei (Breddin) is flattened and twice
curved while that of R. scitula sp. nov. is cylindrical and curved in-
ward,
Tue Universiry Science BULLETIN
806
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 2 apterous males, 4 apterous females. Described
from specimens collected in Sao Paulo, Capital Brazil, 9/XI/40,
Kurt. All type specimens are in the Francis Huntington Snow En-
tomological Collections, University of Kansas.
Data on distribution: In addition to the type series specimens
from the following localities have been examined:
Peru: Loreto Dept., Aguaitia, [X-19-46, F. Woytkowski, 17 apter-
ous males, 13 apterous females, 2 nymphs; Loreto Dept., Roqueron
del Padre, abad Cordillern, VIII-3-46, F. Woytkowski, 1 apterous
male; Loreto Dept., Santa Elena, Roqueron Padre, Abad, VIII-8-46,
F. Woytkowski, 2 apterous males, 1 apterous female.
Rhagovelia sinuata Gould
(Pl. V, fig. 14)
1931. Rhagovelia sinuata Gould, Kansas Univ. Sci. Bull., vol. 20, p. 42.
Size: Length Width
4.35 mm. apterous male 1.40 mm. apterous male
4,65 mm. apterous female 1.60 mm. apterous female
5.05 mm. winged male 1.76 mm. winged male
5.40 mm. winged female 1.76 mm. winged female
Color: General color red-brown to brown, clothed with golden.
pubescence, Anterior band of pronotum not well defined, slightly
lighter in shade. Venter concolorous with dorsum. Base of an-
tennae, margins of all acetabulae, and all coxae and trochanters
yellow-brown. Wings dark brown; anterior, basal cell of wing
pruinose.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Apter-
ous male: anterior trochanter unarmed. Anterior tibia not dilate;
flattened within on apical one half. Pronotum rounded behind
(L. 73,.W. 85). Metanotum short on midline (L. 9, W. 89), Pro-
portions of antennae: Seg. I: Il: HI: IV:: 78: 48: 47: 42; of inter-
mediate legs: Fem.: Tib.: Tars. Il: Tars. III:: 140: 120: 40: 56;
of posterior legs: 182: 117: 9: 28. Abdomen tapers slightly at first,
angle of taper increasing on last two segments. Connexiva semi-
vertical. Venter without definite median carina. Minute, black,
conical setae on base of jugum of head, on proepisternum, and on
venter of all abdominal segments except last. Venter of last abdomi-
nal segment depressed on posterior one half on each side of median
line beneath. Posterior trochanter armed with two to four small,
Srupy or THE GENUS RHAGOVELIA 807
subequal teeth. . Posterior femur usually strongly incrassate (L. 182,
W. 45); armed on basal one third with one row of small knoblike
teeth; armed after basal one third with two rows of spines, the
posterior row consists of a long spine, followed by four short spines,
another long spine and series of shorter spines to apex, the anterior
row consists of short subequal widely-spaced spines from basal one
third to apex. Posterior tibia arcuate; armed within with two irregu-
lar rows of subequal teeth and curved spur at apex. Apterous fe-
male: anterior leg as in male. Pronotum rounded behind (L. 76,
W. 93). Metanotum as in male (L. 10, W. 100). Proportions of
antennae: 69: 41: 46: 40; of intermediate legs: 137: 118: 38: 54; of
posterior legs: 118: 119: 10: 29. Dorsum of abdomen tapers evenly
to apex. Minute, conical, black setae as in male. Posterior tro-
chanter unarmed. Posterior femur less incrassate than in male (L.
118, W. 80); armed beyond middle with one long spine followed
by six or eight smaller, decreasing spines to apex; also armed on
apical one half with anterior row of small subequal spines. Posterior
tibia straight; armed within with two rows of small, subequal teeth;
armed at apex with spur. Winged forms: proportions and armature
similar to apterous forms. Wings extend beyond apex of genital
segments. Pronotum formed same in both sexes; not continued
into spine at apex.
Comparative notes: This species resembles R. nitida Bacon. R.
sinuata Gould can be separated from R. nitida Bacon by the armed
posterior trochanter of the male, and the armed posterior tibia
of the female. In specimens with a less incrassate posterior femur
in the male the armature is reduced but of the same general pattern.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; morphotypes, winged male and winged female. Gould states
the morphotype is a winged female. In examining the type series
only one winged form is present and it is a winged male bearing
the label: “Morphotype, Rhagovelia sinuata Gould.” This winged
male is taken to be the holomorphotype and a winged female is
established as the allomorphotype. The original type series was
collected in Tunguragu Vale, Banos, Ecuador, Jan, 1923, by F. X.
Williams. The allomorphotype was collected in Rio Blanca, Ecua-
dor, 1800 m.a.s.l., May, 1936, by Clarke-MaclIntyre. All type
specimens are in the Francis Huntington Snow Entomological Col-
lections, University of Kansas.
Data on distribution: Recorded only from Ecuador. In addition
to the type series, specimens from the following localities have been
808 Tue UNiversiry ScreENCE BULLETIN
examined (new distribution record for major political areas is indi-
cated with an asterisk) :
Ecuapor: Rio Blanco, swift river, April, 1936, Clarke-MaclIntyre,
No. 3622, 35 apterous males, 27 apterous females; Rio Blanco, run-
ning stream, May, 1936, Clarke-MacIntyre, No, 3686, 18 apterous
males, 1 winged male, 12 apterous females; Juangolo Creek, Banos,
March 15, 1986, Clarke-MacIntyre, No. 8618, 7 apterous males, 12
apterous females; Rio Blanco, 1800 m.a.s.l., May, 1936, Clarke-
MacIntyre, No. 3683, 27 apterous males, 8 winged males, 26 apter-
ous females, 2 winged females; Mera, 1000 m.a.s.l., Nov. 5, 1936,
Clarke-MaclIntyre, No. 3684, 8 apterous males, 12 apterous females.
Peru: Vic. Rioja, Dept. San Martin, jungle, 900 m.a.s.I., Sept.
9, 1936, F. Woytkowski, No. 3682, 10 apterous males, 14 apterous
females, 1 winged female.
Rhagovelia varipes Champion
(Pl. V, fig. 15)
1898. Rhagovelia varipes Champion, Biol. Centr, Amer., Het., vol. 2, p. 138.
1901. Rhagovelia varipes, Kirkaldy, Ent., vol. 34, p, 308.
1931, Rhagovelia beameri Gould, Kansas Univ. Sci. Bull., vol. 20, p. 18 (de-
scribes from Arizona).
1931, Rhagovelia varipes, Gould, Kansas Univ, Sci. Bull., vol. 20, p, 46.
1936. Rhagovelia varipes, Drake and Harris, Proc. Biol, Soc, Washington, vol. .
49, p. 106 (places R. beameri Gould as a synonym of R. varipes Cham-
pion).
Size: Length Width
5.70 mm. apterous male 1.66 mm. apterous male
6.45 mm. apterous female 1.92 mm. apterous female
6.52 mm. winged male 2.25 mm. winged male
6.40 mm. winged female 2.30 mm. winged female
Color: General color black, clothed with golden pubescence. An-
terior band of pronotum orange behind vertex of head, becoming
black behind eyes. Margins of connexiva orange. Dorsum of last
one or two abdominal segments with median shining brown areas.
Venter blue-gray at base becoming red-brown to brown for last four
or five segments. Base of antennae, margins of anterior and pos-
terior acetabulae, all coxae, all trochanters (intermediate pair only
in part), and anterior and posterior femora beneath orange to red-
brown.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed. Anterior tibia flattened
within; not dilate. Pronotum rounded behind (L. 91, W. 110).
Srupy oF THE GENUS RHAGOVELIA 809
Metanotum short on midline (L. 6, W. 105). Proportions of an-
tennae: Seg. I: II: III: IV:: 95: 48: 60: 52; of intermediate legs:
Fem.: Tib.; Tars. II: Tars. III:: 170: 140: 58: 68; of posterior legs:
163: 176: 20: 35. Abdomen tapers only slightly to apex. Connex-
iva semivertical, Venter with median carina on basal three or four
segments. Minute black conical setae on base of jugum of head,
proepisternum, and along venter of abdomen except for last ab-
dominal segment where conical setae are thinly scattered along pos-
terior margin of venter of last abdominal segment at sides. Posterior
trochanter armed with five to nine small subequal teeth. Posterior
femur strongly incrassate (L. 163, W. 54); armed with two rows of
spines, the posterior row begins near base with small, gradually in-
creasing spines with three longer spines near middle followed by
smaller increasing spines with another long spine at apical one third
followed by two or three much smaller spines to apex, the anterior
row composed of irregularly placed, small, subequal spines begin-
ning near base and continuing to apex. Posterior tibia slightly arcu-
ate; armed within with two rows of teeth with those on apical one
third longer and stronger; armed at apex with small, blunt spur.
Apterous female: anterior legs as in male. Pronotum broadly
rounded behind (L. 104, W. 116). Metanotum short on midline
(L. 4, W. 116). Proportions of antennae: 93: 52: 64: 52; of inter-
mediate legs: 173: 144: 62: 74; of posterior legs: 160: 186: 21: 38.
Dorsum of abdomen tapers evenly to apex. Connexiva semivertical.
Minute, black, conical setae as in male, except not apparent on pos-
terior margin of venter of last abdominal segment. Posterior tro-
chanter unarmed. Posterior femur not as incrassate as in male
(L. 160, W. 33); armed on apical one half with one row of spines
consisting of an increasing series of four small spines then one
somewhat larger spine at apical one third followed by four or five
small, rapidly decreasing spines to apex; on some specimens with
reduced armature apparently only armed on apical one third. Pos-
terior tibia straight and apparently unarmed. Winged forms: pro-
portions and armature similar to apterous forms. Pronotum not
produced into spiniform process at apex in either sex.
Comparative notes: This species resembles R. crassipes Champion,
R. varipes Champion can be separated from R. crassipes Champion
by the lack of the abruptly larger tooth at the apical one third of the
posterior tibia of the male; by having only two or three rows of
spines on the posterior femur of the male; and by the posterior
femur of the female being armed at the apical one third with the
810
Tue Universiry Science BULLETIN
longest spine, while the longest spine of the posterior femur of the
female of R. crassipes Champion is placed near the middle.
Data on types: Champion’s type is an apterous male collected in
Bilimek, Mexico. The type specimen is in the Vienna Museum.
Data on distribution: Recorded from Arizona, New Mexico, and
Mexico. Specimens have been examined from the following local-
ities:
Mexico: México: Real de Arriba, Dist. of Temascaltepec, Alt.
1960 meters, May-June, 1933, H. E. Hinton, 18 winged males, 15
winged females.
Hidalgo: San Antonio, near E] Salto, 5000 ft. a.s.l., semitropical,
June 10, 1937, Meldon Embury, 20 apterous males, 2 apterous fe-
males, 17 winged males.
Unrtep States: Arizona: Chiricahua Mts., 7-8-32, R. H. Beamer,
9 apterous males; Cochise Co., 7-29-27, R. H. Beamer (type series
of R. beameri Gould), 17 apterous males, 1 winged male, 50 apter-
ous females, 2 winged females; Ramsay Co., VII-30-41, B. Hodgden,
12 apterous males, 17 apterous females; Huachuca, 5-30-37, W.
Benedict, 9 apterous males, 11 apterous females; Huachuca Mts.,
7-8-32, R. H. Beamer, 6 apterous males, 2 winged males, 13 apterous
females, 2 winged females.
Rhagovelia whitei (Breddin)
(Pl. V, fig. 16)
1898. Neovelia whitei Breddin, Jahrb. Nat. Ver. zu Magee, p. 14.
1901, Pi te whitei, Kirkaldy, Ent., vol. 34, p. 308 (places in genus Rha-
govelia).
1931. Rhagovelia whitei, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 47 (quotes
original description ).
1931. Rhagovelia whitei, Drake and Harris, Pan-Pacific Ent., vol. 8, p. 34
(record from Chapada, Brazil).
Size: Length Width
4,00 mm. apterous male 1.33 mm. apterous male
4.11 mm. apterous female 1.40 mm. apterous female
Color: General color brown, covered with golden pubescence.
Pronotum brown-black with orange-brown band on anterior fifth.
Connexiva broadly margined with orange-brown. Venter with
mesosternum orange-yellow, metasternum and first three or four
abdominal segments brown, remaining segments progressively
lighter brown, first genital segment yellow except at apex, second
genital segment brown. Base of antennae, all coxae and trochanters,
bases of anterior and posterior femora yellow when viewed from
above. All femora yellow in ventral view.
Srupy or THE GENUS RBAGOVELIA 811
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers evenly
to apex. Posterior tibia without sickle-shaped spur at apex. Apter-
ous male: anterior trochanter unarmed, anterior tibia only slightly
dilate and slightly excavate on anterior fifth. Pronotum wider than
long (L. 65, W. 83) and covering mesonotum. Metanotum (Li 5,
W. 83) nearly hidden by apex of pronotum in middle. Proportions
of antennae: Seg. I: II: III: IV:: 68: 85: —: —; of intermediate
legs: Fem.: Tib.: Tars. Il: Tars. I[I:: 185: 98: 85: 57; of posterior
legs: 120: 107: 7: 25. Abdomen tapers evenly to apex. Ventral
carina present, becoming evanescent on last two abdominal seg-
ments. Minute, black, conical setae on jugum of head, proepister-
num, mesosternum and venter of abdomen with exception of last
segment on which minute conical or tooth-shaped setae are concen-
trated into one row along each side of posterior edge of segment.
Posterior trochanter armed with from no to three very small knob-
like teeth and one long, straight spine. Posterior femur greatly
incrassate (L. 120, W. 50) and flattened beneath. Armed on basal
half with median row of very small knoblike teeth; apical half
armed with two lateral rows of spines, the anterior row starts just
before middle and consists of about five knoblike teeth, then a short
spine followed by a stout, curved spine beneath apex, the posterior
row starts at approximately middle with one small spine followed
by three very small, knoblike teeth then two increasing spines to
apex. Posterior tibia curved inward to apical one fourth and curved
slightly outward from there to apex; noticeably laterally flattened;
denticulate within with last three or four teeth increasingly enlarged;
armed with spur at apex. Apterous female: pronotum formed as in
male (L. 68, W. 87). Proportions of antennae: 68: 87: 87: 32; of in-
termediate legs: 125: 92: 34: 56; of posterior legs: 110: 97: 9: 25.
Abdomen tapers evenly to apex. Minute, conical, black setae ar-
ranged as in male except not apparent on last abdominal segment.
Posterior trochanter unarmed, Posterior femur not as incrassate
as in male (L. 110, W. $2); armed on basal half with one median
knoblike tooth followed by one moderate spine just before middle,
apical half of femur armed with two rows of spines, the anterior
row starts at apical one fourth and consists of about three small
spines decreasing in size to apex, the posterior row of spines starts
after middle with one long bent spine followed by approximately
four, smaller, decreasing spines to apex. Posterior tibia almost
straight and denticulate within, teeth more closely set on basal two
thirds; armed at apex with straight spur. Winged forms: unknown.
812
Tue Universiry ScrENCE BULLETIN
Comparative notes: This species resembles R. scitula sp. nov. The
posterior tibia of the male of R. whitei (Breddin) is definitely
laterally flattened on the basal two thirds and curved outward for
the apical one third while that of R. scitula sp. nov. is cylindrical and
arcuate. The posterior femur of the female of R. whitei (Breddin)
is armed with a moderate spine before the long spine at the middle
while the moderate spine is missing in those females of R. scitula
sp. nov. at hand. The color of the spots on the dorsum of the abdo-
men is constant on all the specimens available for study, being dark,
shining brown in R. whitei (Breddin) and yellow to yellow-brown
in R. scitula sp. nov.
Data on types: Breddin’s types are supposed to be in the Berlin
Museum, but Dr. Hungerford, in his notes on his 1928 European
trip, does not mention them as being there. The type locality is
given as “Lagoa Santa” Brazil.
Data on distribution:
Braz: Chapada, June (U.S. N.M.), 1 apterous male; Chapada,
Aug. (Jowa State College Collection) 1 apterous male, 1 apterous
female.
Rhagovelia williamsi Gould
(Pl. V, fig. 17)
1981. Rhagovelia williamsi Gould, Kansas Univ. Sci. Bull., vol. 20, p. 47.
1935. Rhagovelia williamsi, Drake and Harris, Proc. Biol. Soc, Washington
vol. 48, p. 83 (incorrectly place R. williamsi Gould as a synonym of
R. amazonensis Gould).
Size: Length Width
3.56 mm. apterous male 1.17 mm, apterous male
3.69 mm. apterous female 1.35 mm. apterous female
Color: General color brown-black, clothed with golden pubes-
cence. Anterior one fifth of pronotum yellow, becoming pruinose
at sides. Dorsum of abdomen with first and fourth segments prui-
nose, Margins of connexiva orange to brown. Venter brown-black
to gray-black. Venter of last abdominal segment brown beneath.
Base of antennae, margins of all acetabulae, all coxae, anterior and
posterior trochanters, and bases of anterior and posterior femora
yellow to orange.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen tapers evenly to apex in apterous
female, Posterior tibia without sickle-shaped spur at apex. Ap-
terous male: anterior trochanter unarmed, Anterior tibia only
slightly flattened within and not dilate. Pronotum broadly rounded
behind (L. 52, W. 71). Metanotum short on midline (L. 7, W. 72).
Srupy oF THE GENUS RHAGOVELIA 818
Proportions of antennae: Seg. I: II: II: TV:: 54; 31: 82: 32; of in-
termediate legs: Fem.: Tib.: Tars. Il: Tars. II:: 116: 87: 85: 50;
of posterior legs: 98: 78: 4: 19, Abdomen tapers rather evenly to
apex with slight increase in angle of taper on last three segments.
Connexiva semivertical. Venter without median carina. No black,
minute conical setae on base of jugum of head, proepisternum or on
posterior edge of venter of last abdominal segment at sides. Pos-
terior trochanter armed with three or four short teeth and one longer
tooth. Posterior femur moderately incrassate (L. 98, W. 80); armed
on basal one half with one row of knoblike teeth ending in one
moderate spine before middle; armed at middle with one long
spine followed by seven or eight rapidly decreasing spines to apex,
and an anterior row of five or six decreasing spines to apical one
third. Posterior tibia armed within with two rows of subequal
teeth with those of basal one third slightly stouter; armed at apex
with stout spur. Apterous female: anterior leg formed as in male.
Pronotum broadly rounded behind (L. 60, W. 80). Metanotum
short on midline (L. 6, W. 82). Proportions of antennae: 60: 34:
34: 33; of intermediate legs: 123: 91: 34: 54; of posterior legs: 100:
80: 5: 22. Dorsum of abdomen tapering rather evenly to apex.
Connexiva vertical. Minute, conical setae absent as in male. Pos-
terior trochanter unarmed. Posterior femur moderately incrassate
(L. 100, W. 25); armed on basal one half with one moderate spine
which may be reduced or absent; armed at middle with one long
spine followed by six or seven rapidly decreasing spines to apex;
usually armed with an anterior row of inconspicuous spines on
apical one third. Posterior tibia slightly arcuate; armed within
with two rows of subequal teeth with those of basal one third some-
what stouter; armed at apex with stout spur. Winged forms: un-
known.
Comparative notes: This species resembles R. amazonensis
Gould. R. williamsi Gould can be separated from R. amazonensis
Gould by the armature of the posterior femur of the male which
has a moderate sized spine preceding the long spine at the middle,
and by the longer than wide (L. 29, W. 93) dorsum of the first
genital segment of the female. The dorsum of the second abdominal
segment of the female does not have the longer hairs and golden
pubescence such as is on the dorsum of the first and third ab-
dominal segments in R. williamsi Gould, while in R. amazonensis
Gould the second segment is clothed the same as the first and third.
This species was said by Drake and Harris to be inseparable from
R. amazonensis Gould but, in view of the characters which can be
814 Tue Universtry Science BULLETIN
used to separate the two species, it is the opinion of the author that
R. williamsi Gould is closely related to, but distinct from R. ama-
zonensis Gould and should be regarded as a valid species.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 9 apterous males, 11 apterous females. The type
series was collected in Tena, Ecuador, Feb. 28, 1923, by Bix.
Williams. The above type series is in the Francis Huntington Snow
Entomological Collections, University of Kansas.
Data on distribution: In addition to the type specimens, speci-
mens from the following locality have been examined:
Ecuapor: Oriente, Jatun Yacu, 700 m.a.s.l, E. Rio Napo Water-
shed, Mar., 1937, Clarke-MacIntyre, 8 apterous males, 4 apterous
females.
Hirtrees Group
Group characteristics: The hirtipes group can be characterized
as consisting of those species of the genus Rhagovelia in which the
dorsum of the abdomen of the apterous female is carinate on the
midline after the first two segments, and tapers more or less evenly
to the apex. The intermediate femur of the female is cylindrical
throughout. The last two tarsal segments of the intermediate leg
of the apterous female are equal in length. (The distinctive pro-
portionate measurements and the peculiar shape of the male
clasper have made it desirable that this single species be placed
in a group by itself pending the discovery of related forms. Ob-
viously, group characteristics and specific characteristics can be
easily confused in this case. Further clarification of these matters
must await additional species in this group.)
Rhagovelia hirtipes Drake and Harris
(Pl. VI, fig. 9)
1927. Rhagovelia hirtipes Drake and Harris, Proc. Biol. Soc. Washington, vol.
40, p. 186 (describe from he female from Honduras),
1931. Rhagovelia hirtipes, Drake and Harris, Pan Pacific Ent., vol. 8, p. 35
(ada descriptions of winged male and apterous forms; record from
Guatemala).
1931. Rhagovelia hirtipes, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 34.
Size: Length Width
4.40 mm. apterous male 1,33 mm. apterous male
4.45 mm. apterous female 1.50 mm. apterous female
4.33 mm. winged male* 1.60 mm. winged male
4.28 mm. winged female* 1.66 mm. winged female
Color: General color pruinose black, clothed with golden pubes-
cence. Pronotum with anterior band yellow behind vertex of head,
bs Wings broken away at apex, The measurement given is to apex of genital segments.
Srupy OF THE GENUS RHAGOVELIA 815
becoming pruinose black behind eyes. Margins of connexiva brown.
Dorsum of last abdominal segment with brown to yellow shining
median spot. Venter blue-gray. Venter of last abdominal segment
brown beneath. Base of antennae, margins of all acetabulae, all
coxae and trochanters, and bases of anterior and posterior fe-
mora beneath yellow.
Structural characteristics: Pronotum of apterous forms covers
mesonotum. Dorsum of abdomen of apterous female tapers rather
evenly to apex. Posterior tibia without sickle-shaped spur at apex.
Apterous male: anterior trochanter unarmed. Anterior tibia flat-
tened within; not dilate. Pronotum with width subequal to length
(L. 80, W. 81). Metanotum short on midline (L. 6, W. 81). Pro-
portions of antennae: Seg. I: II: II: IV:: 69: 40: 44: 48; of inter-
mediate legs: Fem.: Tib.: Tars. II: Tars. III:: 148: 115: 59: 56; of
posterior legs: 125: 135: 20: 25. Abdomen tapers evenly to apex at
first; angle of taper increasing on last three segments. Connexiva
semivertical. Venter of abdomen without median carina. Minute,
conical setae absent on base of jugum of head, on proepisternum,
and on posterior margin of venter of last abdominal segment at
sides. Posterior one half of venter of last abdominal segment be-
neath depressed on each side of median line. First genital segment
strongly curved downward on posterior one half. Posterior tro-
chanter armed with from four to twelve small, knoblike teeth. Pos-
terior femur strongly incrassate (L. 125, W. 43); armed on basal
one third with a row of knoblike spines, armed at basal one third
with one long spine followed by eight to twelve much smaller, grad-
ually decreasing spines to apex; also armed with anterior row of
small, subequal spines from before middle to apex. Posterior tro-
chanter straight; armed within with irregular row of stout, subequal
teeth; armed at apex with short spur. Apterous female: anterior
leg as in male. Pronotum broadly rounded behind (L. 82, W. 90).
Metanotum short on midline (L. 11, W. 96). Proportions of an-
tennae: 70: 40: 43: 42; of intermediate legs: 140: 115: 59: 59; of
posterior legs: 110: 132: 20: 27. Dorsum of abdomen tapering
more or less evenly to apex. Margins of connexiva semivertical to
flat at first, abruptly becoming vertical over dorsum of third ab-
dominal segment, usually becoming semivertical again for last two
segments. Dorsum of abdomen usually noticeably carinate on mid-
line of third, fourth, and fifth segments with segments somewhat
depressed at the sides. Posterior trochanter unarmed. Posterior
femur only slightly incrassate (L. 110, W. 20); armed at apical one
Tue Universiry SciENCE BULLETIN
third with one moderate spine followed by four or five smaller,
decreasing spines to apex. Posterior tibia straight and apparently
unarmed. Winged forms: proportions and armature similar to ap-
terous forms. Pronotum (L. 111, W. 102) clothed with brown hairs
at apex; not produced into spiniform process in either sex.
Comparative notes: This species resembles R. nitida Bacon. R.
hirtipes Drake and Harris can be separated from R. nitida Bacon by
the lack of a ventral carina and by the armed posterior trochanter
in the male; the females can be separated by the nature of the con-
nexiva and by the dorsal, median carina on the abdomen of R. hir-
tipes Drake and Harris. The intermediate tarsal segments are
longer than the tibia, and the second tarsal segment of the posterior
leg is over three fourths as long as the third tarsal segment in R.
hirtipes Drake and Harris.
Data on types: Holotype, winged female, collected in San Pablo,
Honduras. Allotype, winged male, morphotypes, apterous male
and female, all collected in Chiquimula, Guatemala. The type
material is in the personal collection of Dr. C. J. Drake.
Data on distribution: Recorded from Honduras and Guatemala.
Specimens have been examined from the following localities (new
distribution for major political areas are indicated with an asterisk):
GuatEeMata: Chiquimula, June, 1930, (metatypes), 1 apterous
male, 2 apterous females.
* Mexico: * Distrito Federal: Xochimilco, June 21, 1934, H. E.
Hinton, 2 apterous males, 2 apterous females.
* Guerrero: Malimaltepec, 4-30-30, Prof. L. Schultze, 8 apterous
males, 8 apterous females; Puente de Ixtla, 7-12-37, H. D. Thomas,
1 winged male, 1 winged female.
* Morelos: Cuernavaca, 7-8-36, H. D. Thomas, 9 apterous females;
Cuernavaca, 10-5-36, H. D. Thomas, 4 apterous males, 12 apterous
females; 7-14-86, H. D. Thomas, 1 apterous male.
Co.LLaARris Group
Group characteristics: The collaris group can be characterized as
consisting of those species of the genus Rhagovelia in which the
dorsum of the abdomen of the apterous female is narrow after the
first three segments, the intermediate femur of the female is flattened
beneath at the base. The anterior tibia of the male is usually
greatly dilate and excavate within. Winged forms are common:
the wings just cover the apex of the genital segments,
Srupy OF THE GENUS RHAGOVELIA
The following species comprise this group:
SRS bom
—7
R. planipes Gould
. acuminata sp. nov. 6.
7. R. scabra sp. nov.
8.
9.
. armata (Burmeister )
. collaris (Burmeister )
. cuspidis Drake and Harris
. impensa sp. nov.
R. solida sp. nov.
R. tayloriella Kirkaldy
BRAS
Key To Species oF THE CoLLARnis Group
Terminal genital segment prolonged into a spinelike process.......
Terminal genital segment not prolonged into a spinelike process. . . .
Connexivum terminating in a spine only in female..... acuminata
Connexivum terminating in a spine in both male and female,
cuspidis
Minute, conical, black setae thickly scattered on base of jugum of
head and on posterior edge of last abdominal segment at the
SUGGS es es ie Rg rr Oe OU et OR by ea ee
WAtHOUIEeSTIGH ESOte 5a OTR Tee a es ee os
Minute, conical, black setae thickly scattered on proepisternum,
mesosternum and venter of abdomen. «.s..... 0.05615. eae ee
Without such setae on proepisternum or venter of abdomen, or very
BPAMsCly SCHUIGIOW Carly te rsa er ec s ee ce
Anterior tibia greatly dilate and excavate in male, venter of abdo-
men of female with minute, black, conical setae on all segments
Hout the: last chy /2328. tes eeasee Oeiateey, eee, Sore scabra
Anterior tibia not greatly dilate and excavate in male, venter of
female without conical setae on last 3 or 4 abdominal segments,
solida
Whole body practically the same red-brown color........ planipes
Dorsum of body brown-black, venter red-brown in part........
Venter of male depressed between the posterior trochanters and
coxae; abdomen of female horizontal................. armata
Venter of male not depressed on basal segments; abdomen of fe-
male turned upward at an angle of 15°, genital segments bent
IOWNNVHIC: MOS er el ee ed impensa
Venter of last abdominal segment of male roundly excavate on pos-
terior 2/3 with a median furrow on anterior 1/3; intermediate
trochanter brown to black beneath on all forms... ... tayloriella
Venter of last abdominal segment of male flattened beneath but not
as above; intermediate trochanter of all forms yellow beneath,
collaris
Rhagovelia acuminata sp. nov.
(Pl. VI, fig. 5)
Size: Length Width
5.60 mm. apterous male 1.42 mm. apterous male
6.01 mm. apterous female 1.56 mm. apterous female
5.78 mm. winged male 1.79 mm. winged male
6.25 mm. winged female 1.93 mm. winged female
817
on
Ps
Color; General color red-brown, covered with golden pubes-
cence. Anterior one sixth of pronotum with band of yellow-brown;
27—8378
‘818 Tue Universiry SCIENCE BULLETIN
pronotum also with slightly lighter longitudinal median line becom-
ing evanescent on posterior portion. Margins of connexiva yellow
brown. Venter slightly lighter in color than dorsum. All legs
slightly darker than venter. Wings brown, veins prominent and
darker brown.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrowed after first three segments, connexiva vertical; anterior
tibia moderately dilate and excavate in male. Apterous male:
anterior trochanter unarmed; anterior tibia moderately dilate (W.
15) and slightly excavate for apical half. Pronotum with length
subequal to width (L. 93, W. 91); metanotum much wider than
long (L. 5, W. 94). Proportions of antennae: Seg. I: II: II: IV::
85: 50: 45: 41; of intermediate legs: Fem.: Tib.: Tars. I: Tars. IIT::
195: 135: 60: 67; of posterior legs: 167: 140: 14: 31. Abdomen
tapers evenly to apex. Terminal genital segment mucronate. Ven-
tral carina most strongly produced between posterior coxae where
venter is depressed on each side, gradually becoming evanescent
just before last segment of abdomen. Minute, black, conical setae
scattered on base of jugum of head, metasternum, venter of ab-
domen, and along posterior margin of last abdominal segment at
sides. Venter of last abdominal segment flattened for its entire
length, bordered on each side with slight callus. Posterior tro-
chanter armed with approximately three small teeth. Posterior
femur moderately incrassate (L. 167, W. 35) and armed with small
spines on basal two fifths, followed by one long spine just before
middle and approximately nine decreasing spines to apex; also with
an anterior row of approximately eight small spines beginning
just beyond middle and continuing to apex. Posterior tibia straight;
closely set with two rows of subequal teeth with last tooth slightly
enlarged; posterior tibia also armed with stout, slightly bent spur
on apical margin. Apterous female: anterior tibia flattened for
apical half, scarcely dilate. Pronotum slightly wider than long
(L. 95, W. 103); metanotum slightly longer than in male (L. 9,
W. 95). Proportions of antennae: 85: 49: 44: 40; of intermediate
legs: 190: 185: 55: 65; of posterior legs: 165: 145: 13: 30. Con-
nexiva vertical; narrowed in at first, then widened slightly, nar-
rowed at apex and reflexed so as to have points nearly touching;
each connexivum ends in a mucronate process. Dorsum of abdo-
men narrow after first three segments. Dorsum of first abdominal
segment swollen, shelflike and partially covering second from
above, thickly beset with long, dark hairs. Terminal genital seg-
ment mucronate. No ventral carina, and abdomen not depressed
Srupy oF THE GENUS RHAGOVELIA 819
between posterior coxae. Minute, black, conical setae as in male.
Posterior trochanter unarmed. Posterior femur slightly incrassate
(L. 165, W. 28); armed at middle with one long, dark-tipped,
amber spine followed by approximately six much smaller, subequal,
dark spines; also with an anterior row of approximately four very
small, widely-spaced spines on apical one third. Posterior tibia
straight; armed for its entire length with small teeth, those of the
basal half slightly larger; armed at apex with stout spur. Winged
male: proportions and armature same as for apterous male. Mu-
cronate genital segment projects from under wings so as to be
visible from above. Connexiva not mucronate at apex. Pronotum
drawn out into short spine (L. 15). Winged female: proportions
and armature same as for apterous female. Connexiva mucronate
at apex, not reflexed, tips visible from above, projecting from under
wings. Genital segment mucronate and projects from under the
wings so as to be visible from above. Pronotum drawn out into
long, slightly-bent process which tapers to point at its apex (L.
68).
Comparative notes: This species resembles R. cuspidis Drake
and Harris. R. acuminata sp. nov. can be distinguished from R.
cuspidis Drake and Harris by the lack of a terminal spine on the
connexivum of the male, by the red-brown color, and by the
larger size.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 8 apterous males, 1 apterous female; paramorpho-
types, 81 winged males, 90 winged females. Described from a
Series labeled: “Panama, Feb. 4, 1935, Col. by J. W. McSwain for
Robert Wind.” All type specimens are in the Francis Huntington
Snow Entomological Collections, University of Kansas.
Data on distribution: In addition to the type material, specimens
have been examined from the following localities:
Panama: Potrerillos, 2-19-34, D. V. Brown, 7 apterous males,
22 winged males, 1 apterous female, 17 winged females; Potrerillos,
2-15-35, D. V. Brown, 8 winged males, 1 winged female; Potrerillos,
2-18-35, D. V. Brown, 15 winged males, 16 winged females; Po-
trerillos, 2-20-35, D. V. Brown, 2 winged males, 2 winged females;
Potrerillos, 2-22-35, D. V. Brown, 6 winged males, 7 winged fe-
males; Potrerillos, 2-25-35, D. V. Brown, 7 winged males, 5 winged
females; Potrerillos, 3-9-35, D. V. Brown, 8 winged males, 3 winged
females,
Tue University Scrence BULLETIN
Rhagovelia armata (Burmeister )
(PL. VI, fig. 6)
1835. Velia armata Burmeister, Handb. der Ent., Bd. 2, p. 212.
1865. Rhagovelia armata, Mayr, Verh. Zool.-bot. Ges. Wien, Bd. 15, p. 445
rece in the genus seach tg
1896. Rhagovelia armata, Lethierry and Severin, Cat. Gén. des Hémip. 8, p. 55.
1898. Rhagovelia armata, Champion, Biol. Centr. America, Het., vol. 2, p. 186
(redescribes from Burmeister’s type).
1901. Rhagovelia armata, Kirkaldy, Ent., vol. 34, p. 308 (mentions in key).
1910. Rhagovelia armata, Banks, Cat. of Nearctic Hemip.-Heter., p, 27.
1916. Rhagovelia armata, Van Duzee, Check List of Hemip.
1917. Rhagovelia armata, Van Duzee, Cat. Hemip. America N. of Mexico,
p. '
1919. Rhagovelia armata, Hungerford, Kansas Univ. Sci. Bull., vol. 11, p. 130
(quotes Champion’s redescription, records from Texas).
1931. Ricsovelis armata, Gould, Kansas Univ. Sci. Bull. vol. 20, p. 17 (rede-
scribes, gives distribution as Mexico and Texas).
1981. Riagooela armata, Drake and Harris, Pan Pacific Ent., vol. 8, p. 34
(record from Guatemala).
1935. Rhagovelia armata, Drake and Harris, Proc. Biol. Soc. Washington, vol.
48, p. 33 (record from West Indies, describe variations in armature,
etc. ).
Size: Length Width
4,80 mm. apterous male 1.33 mm. apterous male
5.15 mm. apterous female 1.46 mm. apterous female
5.40 mm. winged male 1.73 mm. winged male
5.45 mm. winged female 1.82 mm. winged female
Color: General color brown-black; clothed with golden pubes-
cence. Anterior band of pronotum yellow behind vertex of head,
becoming pruinose behind eyes. Dorsum of last abdominal segment
shining black. Margins of connexiva yellow. Mesosternum more
or less yellow as are venters of the last three or four abdominal seg-
ments; remainder of venter blue-gray to gray-black. Base of an-
tennae, margins of all acetabulae, all coxae, trochanters, base of an-
terior femur, base of posterior femur above and all of posterior
femur beneath yellow. Wings dark brown; veins black.
Structural characteristics: Dorsum of abdomen of apterous female
narrow after first three segments, connexiva vertical; anterior tibia
moderately dilate and excavate beneath in male. Apterous male:
anterior trochanter unarmed; anterior tibia moderately dilate and
excavate on apical one half, Pronotum rounded behind (L. 75, W.
85), and covering mesonotum. Metanotum short on midline (L. 7,
W. 85). Proportions of antennae: Seg, I: II: III: IV:: 70: 40; 42:
40; of intermediate legs: Fem.: Tib.: Tars, II: Tars. II:: 103; 106:
47: 60; of posterior legs: 130: 111: 13: 27. Abdomen tapers evenly
to apex. Venter of abdomen depressed for first three segments;
ventral carina present on basal segments of abdomen, becoming
evanescent on next to last segment. Minute, black, conical setae on
base of jugum of head, sparsely on venter of abdomen, and along
Stupy OF THE GENUS RHAGOVELIA 821
venter of last abdominal segment at sides. Venter of last abdominal
segment shallowly excavate on posterior one half, anterior one half
with two hairy prominences with shallow trough between. Pos-
terior trochanter armed with several small teeth. Posterior femur
greatly incrassate (L. 180, W. 44); armed on basal two fifths with
one row of short, subequal spines; armed on apical three fifths with
a larger posterior row which begins at basal two fifths with one
long spine followed by eight or ten smaller, gradually decreasing
spines, and a smaller, anterior row of subequal spines which begins
at basal two fifths and continues to apex. Posterior tibia straight;
armed within with small teeth with those at apical one third tend-
ing to be stouter; armed at apex with stout spur. Apterous female:
anterior tibia shallowly excavate and not as dilate as in male. Pro-
notum rounded behind (L. 77, W. 101); metanotum as in male
(L. 6, W. 100). Proportions of antennae: 71: 38: 40: 86; of inter-
mediate legs: 146: 105: 50: 59; of posterior legs: 130: 120: 14: 30.
Dorsum of abdomen narrow after first three segments. Connexiva
Vertical; narrowing after first three segments, then diverging slightly,
harrowing again over the last segment and diverging at apex. Ven-
ter not depressed as in male; no ventral carina. Minute, black,
conical setae as in male, except much more reduced at sides of last
abdominal segment. Intermediate femur somewhat flattened be-
neath at base. Posterior trochanter unarmed. Posterior femur not
as incrassate as in male (L. 130, W. 30); armed with one long spine
after middle followed by approximately six, smaller, gradually
decreasing spines to apex. Posterior tibia straight; armed within
with dark teeth and spur at apex. Winged forms: proportions and
armature similar to apterous forms. Wings barely covering apex
of genital segments. Winged male: venter formed as in apterous
male, Pronotum continued into short, pointed process (L. 18).
Winged female: pronotum continued into long, curved, sharply
Pointed process (L. 50).
Comparative notes: This species has been misidentified frequently
in the past. Much of this is due to Champion’s figuring an apterous
female (1901-pl. VII) which belongs to the crassipes group as the
female of R. armata (Burmeister). R. armata (Burmeister) is
closely related to R. planipes Gould, but can be separated from that
Species by the formation of the venter of the male, which is de-
pressed for the basal three segments, and excavate on the posterior
One half of the apical segment. The females of R. armata may be
Separated from those of R. planipes by the darker colored venter of
the abdomen which is orange in R. planipes. R. armata (Burmeis-
822, Tue Universtry SCIENCE BULLETIN
ter) is also closely related to R. scabra sp. nov. but can be separated
from that species at once by the lack of the thickly scattered minute,
conical, black setae on the venter of the thorax.
Data on types: Burmeister’s types are in the Berlin Museum. The
type specimens were collected in Mexico. The female figured by
Champion was not from the type series but was from the Vienna
Museum and had been identified by Mayr.
Data on distribution: Recorded from Mexico, Guatemala, West
Indies, and Texas. How many of these records are based on mis-
identified specimens, it is impossible to state. Specimens from the
following localities have been examined (new records for major
political areas are indicated with an asterisk) :
* Costa Rica: Rio Virilla, 12-26-31, Heinrich Schmidt, 2 apterous
males, 2 winged males, 2 apterous females, 2 winged females; San
Isidro del Gen., 2000 ft., Feb., 1939, Dean L. Rounds, 2 apterous
males; San José, purchased 1932, Heinrich Schmidt, 1 apterous male,
1 winged male, 8 apterous females, 1 winged female.
GuateMata:., Amatitlan, Feb. 8, 1905 (J. R. de la Torre-Bueno
Collection) 14 apterous males, 20 apterous females; Mazatenango,
Feb. 3, 1905 (J. R. de la Torre-Bueno Collection) 3 winged males,
1 winged female.
* Honpuras: Copan, 2-18-1937, Chester Roys, 1 apterous male,
2 apterous females.
Rhagovelia collaris (Burmeister )
(Pl. VI, fig. 7)
1835, Velia collaris Burmeister, Handb. der Ent., Bd. 2, p. 212.
1857. Velia fiebrii Guérrin, in Sagra’s Nat. Hist. Cuba, vol. 2, 7, p. 174.
1865. Rhagovelia collaris, Mayr, Verh. Zool-bot. Ges. Wien, Bd. 15, p. 445.
1867. Rhagovelia collaris, Mayr, Reise d Oster Freg. Noy., Zool. Bd. 2, p. 180.
1872, Rhagovelia collaris, Uhler, Proc. Boston Soc. Nat. Hist., vol. 19, 4, p. 434.
1892. Rhagovelia collaris, Johnston and Fox, Ent. News, vol. 3, p. 60 (record
from Jamaica).
1896. Rhagovelia collaris, Lethierry and Severin, Cat. Gén, des Hémip., vol. 3,
p. 55).
1898. _Rhagovelia collaris, Champion, Biol. Centr. Amer., Het., vol. 2, p. 184.
1900: Rhagovelia collaris, Kirkaldy, Ento., vol. 33, p. 71.
1901... Rhagovelia collaris, Kirkaldy, Ento., vol. 34, p. 308.
1910. Rhagovelia, collaris, Banks, Cat. of Nearctic Hemip.-Hetr., p. 27.
1931. Rhagovelia collaris, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 21 (re-
describes, gives distribution as San Domingo, Jamaica and Cuba).
1931, Rhagovelia collaris pulchra Gould, Kansas Uniy. Sci. Bull., vol. 20, p. 22
'- (describes from B. W. I.).
1931, Rhagovelia collaris, Drake and Harris, Pan Pacific Ent., vol. 8, p. 34
(record variations and give distribution of Guatemala and Arizona).
1933. -Rhagovelia collaris pulchra, Gould, Ann. Ent. Soc. America, vol, 26, p.
466 (records from Santo Domingo).
1933. Rhagovelia collaris, Gould, Ann. Ent. Soc. America, vol. 26, p. 466
(records from Cuba and Jamaica).
Srupy OF THE GENUS RHAGOVELIA 823
Size: Length Width
4,60 mm. apterous male 1.40 mm. apterous male
5.05 mm. apterous female 1.53 mm. apterous female
4.58 mm. winged male 1.78 mm. winged male
4.95 mm. winged female 1.80 mm. winged. female
Color: General color brown-black, clothed with golden pubes-
ence. Pronotum with anterior band yellow behind vertex of head,
becoming pruinose behind eyes. Dorsum of abdomen pruinose
toward sides. Margins of connexiva yellow. Venter orange be-
neath except metasternum and first two abdominal segments. Base
of antennae, margins of all acetabulae, all coxae and trochanters,
base of anterior femur, and base of posterior femur above, all of
posterior femur beneath yellow.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexivum vertical; ante-
rior tibia greatly dilate and excavate in the male. Apterous male:
anterior trochanter unarmed; anterior tibia greatly dilate (L. 87,
W. 23) and excavate beneath after basal one third. Pronotum
broadly rounded behind (L. 78, W. 88); mesonotum hidden; meta-
notum short on midline (L. 11, W. 85). Proportions of antennae:
Seg. I; Il: III: IV: : 74: 42: 44: 38; of intermediate legs: Fem.:
Tib.: Tars. Il: Tars. III: : 150: 112: 47: 58; of posterior legs:
126; 124: 138: 27. Abdomen tapers evenly to apex. Venter with
fairly distinct median carina on basal segments, becoming evanes-
cent on next to last segment. Venter of last abdominal segment
slightly excavate on posterior one half or can be flattened beneath.
No minute, conical, black setae on base of jugum of head, on venter,
or on last abdominal segment at sides. Posterior trochanter armed
With several small teeth. Posterior femur greatly incrassate (L.
126, W. 38); armed on basal one third with one row of short, sub-
equal spines; armed on apical three fifths with one larger posterior
tow which begins at basal two fifths with one long spine followed
by ten or twelve gradually decreasing spines to apex, and a smaller,
anterior row of subequal, small spines which begins at basal two
fifths and continues to apex. Posterior tibia straight; armed within
with small teeth with those on basal two thirds tending to be
stouter; armed at apex with slender spur. Apterous female: anterior
tibia slightly dilate and flattened within. Pronotum as in apterous
male (L. 80, W. 92); metanotum as in male (L, 10, W. 93). Pro-
Portions of antennae: 75: 40: 44: 38; of intermediate legs: 148:
115: 48: 61; of posterior legs: 128: 182: 18: 27, Dorsum of
abdomen narrow after first three segments. Connexiva semiver-
tical to vertical; narrow after first three segments and continuing
824 Tue University ScreNcE BULLETIN
subparallel. Minute, conical, black setae absent. Intermediate
femur slightly flattened beneath at base. Posterior trochanter un-
armed. Posterior femur not as incrassate as in male (L. 128, W.
27); armed at apical two fifths with one long spine followed by
four or five smaller, decreasing spines to apex. Posterior tibia
straight; armed with inconspicuous teeth and a spur at apex.
Winged forms: proportions and armature much as in apterous
forms. Wings barely covering apex of genital segments. Winged
male: venter formed as in apterous male. Pronotum produced into
a short process at apex (L. 10). Winged female: pronotum pro-
duced into a long, bent, sharply-pointed process at apex (L. 45).
Comparative notes: This species resembles R. tayloriella Kirk-
aldy, but can be separated from that species by the intermediate
trochanter of all forms being yellow beneath in R. collaris (Bur-
meister). Also the venter of the last abdominal segment of the
male is differently formed in the two species. There is some varia-
tion in the incrassateness of the posterior femur in different speci-
mens. This has been noted before by Drake and Harris (1931;
p. 84). The specimens from Puerto Rica are somewhat foreshort-
ened; however, all other characters are within logical range of
variation so it was thought best not to describe these Puerto Rica
specimens as a new subspecies. R. collaris pulchra Gould was
found to be indistinguishable from R. collaris (Burmeister) and
is made a synonym of it.
Data on types: Burmeister’s type specimens are located in the
Berlin Museum. The type series was collected in Port au Prince,
Haiti.
Data on distribution: Recorded from Haiti, San Domingo, Ja-
maica, Cuba, Guatemala, and Arizona (?). Specimens from the
following localities have been examined (new records for major
political areas are indicated with an asterisk) :
Cupa: Loc. Arroyo, Santa Clara Prov., June 22, 1932, P. J. Ber-
mudez, 26 apterous males, 46 apterous females; Matanzas, Yumuri
Valley, 9-19-33, P. J. Bermudez, 18 apterous males, 12 apterous fe-
males; Havana, F. Z. Cervera (J. R. de la Torre-Bueno Collection )
1 apterous male, 3 apterous females; Trinidad Mts., Mina Carlota,
22 March, 1925, 2 apterous males, 1 winged female; Matanzas,
Yumuri Valley, 12-9-33, P. J. Bermudez, 2 apterous males, 4 apterous
females; arroyo in Mena, West Matanzas, Yumuri Valley, 5-16-32,
P. J. Bermudez, 6 apterous males, 1 winged male, 7 apterous fe-
males, 2 winged females.
Stupy oF THE GENUS RHAGOVELIA 825
Dominican Repusuic: Roseau, 1-29-26, P. G. Howes (type series
of R. collaris pulchra Gould) 4 apterous males, 1 winged male,
3 apterous females; Cristébal, 7-10-30, R. M. Bond, 1 apterous male,
2 apterous females; Laudet, June 10, 1911 (U. S. National Museum
Collection) 2 apterous males, 1 apterous female.
Hartt: Port au Prince, Mar. 1927, 5 apterous males, 3 apterous
females; Mirebalais, 12-2-28, 15 apterous males, 6 apterous females.
* Puerto Rica: Naranjito, July 6, 1915 (U.S. National Museum
Collection), 1 winged male; Aibonito, July 14-17, 1914 (U. S. Na-
tional Museum Collection), 1 apterous male; Adjuntas, June 8-13,
1915 (U. S. National Museum Collection), 1 apterous female;
Barranguita, 2-20-27, W. A. Hoffman, 1 apterous male, 1 winged
male, 2 apterous females; Anasco, 2-1-36, H. Lamoutte, 1 apterous
male, 1 winged male, 1 apterous female; Luquillo Mts. (swim
pool), No. 10-1, Feb. 18, 1935, Needham and Diaz, 3 apterous
males, 1 winged male, 8 apterous females; Coquitas River, No.
829-11, March 4, 1935, Needham and Diaz, 1 winged male, 1 winged
female; Rio Blanco, No. 373-12, Mar. 6, 1935, Needham and Diaz,
2 apterous males, 2 apterous females; Micaco River, No. 428-14,
March 7, 1935, Needham and Diaz, 1 winged female; Prieto River,
No. 448-15, March 7, 1935, Needham and Diaz, 3 apterous males,
3 apterous females; Sabana River, No. 465-16, March 7, 1935,
Needham and Diaz, 4 apterous males, 9 apterous females; Que-
brada, La Joba, No. 414-2, March 8, 1985, Needham and Diaz, 11
apterous males, 2 winged males, 16 apterous females, 1 winged
female; Carreras River, No. 504-14, March 9, 1935, Needham and
Diaz, 1 apterous male, 1 winged male, 2 apterous females, 1 winged
female; Arecibe Utuado, No. 545-22, March 13, 1935, Needham and
Diaz, 1 apterous male, 1 winged male, 1 apterous female; Iuebrada,
Tomey, No. 96-8, Feb. 21, 1985, Needham and Diaz, 2 apterous
males; Lares, Guajataca River, No. 747-26, March 22, 1935, Need-
ham and Diaz, 1 apterous male, 3 apterous females; Adjuntas
Juyaya Rd., No. 868-30, March 24, 1935, Needham and Diaz, 38
apterous females, 1 winged female; Luquillo Mt. Rd. Km. 11.7,
No. 1117-41, June 8, 1935, J. G. Diaz, 6 apterous males, 5 winged
males, 6 apterous females, 1 winged female; El Yunque Trail, No.
1173-43, June 10, 1935, J. G. Diaz, 2 apterous males, 2 winged males,
2 apterous females; Luquillo Mts. (swim pool), No. 11-1, Nov. 18,
1935, Needham and Diaz, 1 apterous male, 10 apterous females;
Luquillo Mts. (swim pool), No. 1773-39, Nov. 17, 1935, Needham
and Diaz, 5 winged females; Luquillo Mts. (picnic grounds), No.
826 Tue Universiry SCIENCE - BULLETIN
23-1, Nov. 12, 1985, Needham and Diaz, 3 apterous males, 1 winged
male, 1 apterous female.
Rhagovelia cuspidis Drake and Harris
1933. Rhagovelia cuspidis Drake and Harris, Proc. Biol, Soc. , Washington,
vol. 46, p. 51.
This species is not represented in the material in the Francis
Huntington Snow Entomological Collections, University of Kansas.
The original description is quoted below:
“Black, thickly clothed with fine golden pubescence, the front
margin of pronotum, connexival margins, base of first antennae,
prosternum, all coxae, anterior and posterior trochanters, base of
anterior femora and part of hind femora testaceous to brownish
testaceous. Head with the usual impressed lines. E'yes large,
coarsely faceted. Antennal formula, 45: 25: 27: ?, the second seg-
ment with long hairs on dorsal surface, the third slightly ‘expanded.
Pronotum extending over mesonotum, broadly rounded behind,
with an indistinct median ridge, with rather numerous indistinct,
but deep punctures. Abdomen tapering slightly posteriorly, the
margins nearly straight, terminating behind in prominent somewhat
laterally projecting spine like processes; last segment a little longer
than preceding, with a patch above, shiny, the apex truncate. Last
genital segment ending in a long sharp process. Legs moderately
hairy, the anterior tibia strongly compressed, somewhat expanded
and shallowly excavated beneath. Intermediate legs long; formula,
102: 76: 35: 38.
“Apterous male.—Venter bluish-black, clothed with longer hairs,
last segment shiny black, strongly depressed behind, the apex trun-
cate. Genital segments plump, the first segment carinate along
median line at base. Clasper long, very broad at base, the terminal
portion sub-cylindrical, of about equal width throughout, slightly
curved inwardly, blunt at apex and about three times as long as
basal portion. Hind femora rather strongly incrassate, reaching
to genital segments, armed with nine or ten progressively shortened,
black-tipped testaceous spines; the first of these is much longer than
the others and situated slightly before the middle; also armed along
the basal half, before the long spine, with a distinct row of closely
set, black teeth and along the distal half, beneath the spines with
an irregular row of short black teeth. Posterior trochanter with
numerous short black teeth. Hind tibia nearly straight, armed
within with numerous short black teeth, the apex with a long, stout,
slightly-bent black spur.
Srupy oF THE GENUS RHAGOVELIA 827
“Apterous female.—Posterior femora slightly incrassate, with a
row of black-tipped, testaceous spines in male, Connexivum
broader than in male, the outer margins nearly straight, the apical
spine as in male.
“Length, 4.42 mm.; width, 1.15 mm.
“Holotype, apterous male, and allotype, apterous female, Punta
Gorda, Br. Honduras; authors’ collection. Paratypes, three apter-
ous males, taken with types.
“The connexival spines, the mucronate terminal abdominal seg-
ment, and the character of the armature of the hind femora will
serve to separate this species from allied forms.”
Comparative notes: This species may resemble R. acuminata
sp. nov. The type material is all in the personal collection of Dr. C. J.
Drake and has been unavailable for study. R. cuspidis Drake and
Harris may be separated from R. acuminata sp. nov. by the spinelike
termination of the connexivum of the male, by its black color, and
by its smaller size.
Rhagovelia impensa sp. nov.
(Pl. VI, fig. 8)
Size: Length Width
4,36 mm. apterous male 1.20 mm, apterous male
4.77 mm. apterous female 1.40 mm. apterous female
4.90 mm: winged male 1.62 mm. winged male
5,80 mm. winged female 1.92 mm. winged female
Color: General color brown-black, clothed with golden pubes-
cence. Pronotal band yellow behind vertex of head, becoming
pruinose behind eyes. Dorsum of abdomen pruinose at sides. Mar-
gins of connexiva yellow. Venter yellow-brown to red-brown;
darkest between intermediate coxae becoming almost brown-black
on some specimens. Base of antennae, margins of all acetabulae,
all coxae and trochanters, basal one half of anterior femur, and all
posterior femur beneath yellow. Wings dark brown; veins black.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva vertical to slightly
reflexed, anterior tibia moderately dilate and excavate in male. Ap-
terous male: anterior trochanter unarmed. Anterior tibia moder-
ately dilate (L. 80, W. 20) and excavate on apical half. Pronotum
rounded behind (L. 72, W. 80); metanotum short on midline (L. 9,
W. 80). Proportions of antennae: Seg. I: I: II: IV:: 67: 39: 40:
86; of intermediate legs: Fem.: Tib.: Tars. II: Tars. III:: 186: 105:
44: 58; of posterior legs: 120: 110: 15: 29. Abdomen tapers evenly
828 Tue Universiry ScieNCE BULLETIN
to apex. Connexiva semivertical. Ventral carina only evident on
basal two’ segments becoming evanescent between posterior tro-
chanters. Venter of last abdominal segment flattened to slightly
excavate beneath. Minute, conical, black setae on base of jugum
of head and along sides of venter of last abdominal segment. Pos-
terior trochanter armed with four to five small teeth. Posterior
femur moderately incrassate (L. 120, W. 35); armed on basal one
third with row of short teeth and beyond basal one third with one
long spine followed by approximately twelve smaller, gradually de-
creasing spines to apex; also armed on apical one half with an ante-
rior row of small, subequal spines. Posterior tibia straight; armed
within with closely set teeth on basal half, with those at approxi-
mately middle stouter; teeth of apical one half smaller and more
widely spaced; armed at apex with stout spur. Apterous female:
anterior tibia much as in male. Pronotum broadly rounded behind
(L. 80, W. 95); metanotum as in male (L. 9, W. 95). Proportions
of antennae: 70: 42: 44: 39; of intermediate legs: 142: 107: 47: 63;
of posterior legs: 118: 120: 18: 80. Dorsum of abdomen narrow
after first three segments. Connexiva vertical to semireflexed tent-
like over dorsum of abdomen. Brown hairs thickly set on dorsum
of last abdominal segment. Venter with no trace of median carina.
Minute, black, conical setae as in male, except sparsely set on sides
of venter of last abdominal segment. Intermediate femur flattened
beneath for basal one third. Posterior trochanter unarmed. Pos-
terior femur not as incrassate as in male (L. 120, W. 23); armed
beyond middle with one long spine followed by four or five smaller,
rapidly decreasing spines to apex; there may or may not be one or
two small spines anterior to main row after apical one third. Pos-
terior tibia straight; seemingly unarmed, or armed on basal one
half with inconspicuous teeth; armed with spur at apex. Winged
forms: proportions and armature similar to apterous forms. Wings
just cover apex of genital segments. Winged male: pronotum pro-
longed at apex into a short, pointed process (L. 10). Winged
female: pronotum prolonged at apex into long, narrow, cylindrical
process (L. 50), which arises almost vertically from pronotum at
first, then curves so that apex is directed slightly downward.
Comparative notes: This species resembles R. armata (Burmeis-
ter), but can: be separated from that species by the normal basal
segments of the venter of the abdomen in the male. The male
clasper is also distinct, being more abruptly curved and with the
blade widened: near the apex, while the clasper of R. armata (Bur-
Stupy oF THE GENUS RHAGOVELIA 829
meister) is gradually curved and the blade tapers more evenly to
the apex. The females of R. impensa sp. nov. can be separated from
those of R. armata (Burmeister) by the abdomen being bent up-
ward at an angle of fifteen degrees with the genital segments bent
downward at an angle of forty-five degrees, while the abdomen
and genital segments of the female of R. armata (Burmeister) are
practically horizontal.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 7 apterous males, 6 apterous females. The apterous
specimens were collected in Lagunas Villa, Peru, on June 8 to
July 1, 1934, by F. Woytkowski. The winged specimens were
collected in the vicinity of Balsas, Peru River Marafon, 1179
m.a.s.l., June 26-29, 19386, by F. Woytkowski. All type specimens
are in the Francis Huntington Snow Entomological Collections,
University of Kansas.
Data on distribution: In addition to the type specimens, speci-
mens from the following localities have been examined:
Peru: Dept. Cajamarco, 1179 m.a.s.l., River Marafion, Vic. of
Balsas, June 26-29, 1936, F. Woytkowski, No. 3634, 4 apterous
males, 1 apterous female; Santa Clara, River Rimac, 17 km. E. of
Lima, Sept. 22-25, 1934, F. Woytkowski, 7 apterous males, 8 apterous
females; Dept. Cajamarco, Andes, River Lallanga, Vic. Llangua,
June 20, 1936, F. Woytkowski, No. 3633, 5 apterous males, 1 winged
male, 6 apterous females; Dept. Cajamarco, Andes, 1600 m.a.s.L,
Vic. Jerez, Rivulet Mineral, July 12, 1936, F. Woytkowski, No.
3635, 9 apterous males, 5 apterous females; Dept. Lima, Luria, 45
km. S. of Lima, pools near sea, Nov. 3-5, 1934, F. Woytkowski, 17
apterous males, 11 apterous females; Rio Rimac, 17-44 km. E. of
Lima, Nov. 27-29, 1934, F. Woytkowski, 17 apterous males, 20
apterous females; mouth of Rio Rimac, close to Pacific, June, 1934,
F, Woytkowski, 17 apterous males, 41 apterous females.
Rhagovelia planipes Gould
(Pl. VI, fig. 9)
1931. Rhagovelia collaris planipes Gould, Kansas Univ. Sci. Bull., vol. 20, p. 22.
Size: Length Width
5.75 mm. apterous male 1.66 mm, apterous male
6.15 mm. apterous female 2.06 mm. apterous female
Color: General color red-brown, clothed with golden pubes-
cence, Pronotum with uninterrupted tan band on anterior one
fifth, becoming slightly pruinose behind eyes. Three apical seg-
830 Tue Universiry SCrmeNCE BULLETIN
ments of antennae, anterior tibia and tarsus, apical half of. inter-
mediate tibia and all of tarsus, apical third of posterior tibia and
all of tarsus brown. Venter entirely red-brown.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva vertical; anterior
tibia moderately dilate and excavate beneath in male. Apterous
male: anterior trochanter unarmed. Anterior tibia moderately
dilate (L. 110, W. 20) and excavate on apical half. Pronotum
slightly wider than long (L. 95, W. 106); metanotum emarginate at
apex (L. 8, W. 110). Proportions of antennae: Seg. I: II: ILI: IV::
90: 55: 47: 44; of intermediate legs: Fem.: Tib.: Tars. II: Tars. III::
185: 135: 62: 72; of posterior legs: 183: 147: 20: 35. Abdomen
tapers evenly to apex. Minute, conical, black setae on base of
jugum and at sides of posterior edge of last abdominal segment.
Metasternum slightly raised in middle so as to appear as continua-
tion of ventral carina. Ventral carina strong, with venter depressed
on each side, extending to next to last segment; on some specimens
only well developed on basal three segments. Last segment flat-
tened. Posterior trochanter armed with approximately five small,
dark teeth. Posterior femur greatly incrassate (L. 183, W. 65);
armed on basal third with row of small closely-set dark spines
followed by two rows of spines to apex, the posterior row begins
at basal one third and consists of one long spine followed by series
of seven rapidly decreasing spines, then another spine at apical
one fourth fully as large as first spine at apical third, followed by
two decreasing spines to apex; the anterior row consists of approxi-
mately fifteen subequal, small, closely-set spines which occur only
in region between the two larger spines of femur. Posterior tibia
curved inward for basal one half, outwards for apical one half,
inner surface swollen from just before apical one third; armed with
two rows of small, knoblike teeth on basal two thirds, with larger
stouter teeth on apical one third; armed at apex with stout spur.
Apterous female: anterior tibia moderately dilate (L. 107, W. 18),
inner surface excavate for apical third. Pronotum wider than long
(L. 98, W. 115); metanotum short on midline (L. 12, W. 122).
Proportions of antennae: 85: 50: 42: 41; of intermediate legs:
168: 118: 54: 70; of posterior legs: 150: 146: 18: 33. Connexiva
vertical, converging at first, then continuing subparallel. _Dorsum
of last abdominal segment thickly set with long, dark hairs. Minute
setae on jugum and posterior edge of last abdominal segment as
in male. No indication of median ventral carina. Posterior tro-
Stupy or THE GENUS RHAGOVELIA 831
chanter unarmed. Posterior femur moderately incrassate (L. 150,
W. 385) and armed at middle with one long spine, followed by
approximately six smaller, decreasing ‘spines to apex; also armed
with an anterior row of approximately four widely-spaced, small,
knoblike spines. beginning just before apical third. Posterior tibia
straight; armed with subequal teeth with those of basal half more
closely-spaced; armed with stout spur at apex. Winged forms:
unknown.
Comparative notes: This species resembles R. armata (Bur-
meister), but can be separated from that species by the almost
unicolorous venter and the characteristic armature of the posterior
femur of R. planipes Gould. This species was originally named
as a subspecies of R. collaris (Burmeister) however, the relation-
ship of R. planipes to R. collaris is comparatively distant. In order
to remain consistent, the variety planipes is elevated to species level.
Data on types: Holotype, apterous male; allotype, apterous fe-
male, Both specimens collected at Mt. Cacaquatique, Dept. St.
Miguel, Salvador, 12-14-25, by R. A. Stirton.
Data on distribution: Recorded only from Salvador. In addition
to the type. specimens, specimens have been examined from the
following locality (new records for major political areas are indi-
cated with an asterisk):
* Mexico: * Chiapas: Sierra Madre, btw. Finca Vergel and F.
Victoria, 900 m.a.s.l., brk.-dp. shade, June 3, 1935, A. Dampf, 1
apterous male, 1 apterous female.
Rhagovelia scabra n. sp.
(Pl. VI, fig. 10)
Size: Length Width
5.20 mm, apterous male 1.53 mm. apterous male
5.65 mm. apterous female 1.738 mm. apterous female
6.38 mm. winged male 2.02 mm. winged male
Color: General color red-brown, clothed with golden pubescence.
Apical segments of antennae and legs brown. Anterior band of
pronotum tan, not interrupted at middle. Disc of pronotum with
more or less distinct median, longitudinal, tan line. Venter tan.
Base of antennae, base of anterior and posterior femora, all coxae,
and trochanters tan. Winged forms darker brown than apterous
forms with veins of wings black.
Structural characteristics: Dorsum of abdomen of apterous female
narrow after first three segments, connexiva vertical to slightly re-
flexed; anterior tibia of male greatly dilate and excavate. Apterous
832 Tue. Universrry SCIENCE BULLETIN
male: anterior trochanter unarmed: anterior tibia greatly dilate (L.
100, W. 25) and excavate on apical half. Pronotum wider than long
(L.91, W. 100); mesonotum hidden by pronotum,; metanotum slightly
emarginate on median line (L. 3, W. 100). Proportions of antennae:
Seg. I: II: III: IV:: 70: 44: 47: 41, of intermediate legs: Fem.: Tib.:
Tars. II: Tars. II:; 163: 122: 56: 63; of posterior legs: 135: 128:
18: 30. Abdomen tapers evenly to apex, connexiva semivertical.
Venter, between posterior coxae, depressed on each side of median
carina which becomes evanescent on next to last abdominal segment,
last abdominal segment depressed on apical half. Minute black con-
ical setae thickly scattered on jugum of head, venter of thorax and ab-
domen with exception of last abdominal segment on which minute,
conical setae are slightly bent inward and arranged in one row along
each side on posterior edge of segment. Posterior trochanter armed
with several small, knoblike teeth. Posterior femur moderately incras-
sate (L. 185, W. 35) and armed along basal half with row of small,
black, knoblike teeth; armed at middle with one large, curved spine
followed by approximately seven smaller, decreasing spines to apex,
also with an anterior row of subequal, small spines from middle
to apex. Posterior tibia straight and armed within with small black
teeth, also armed at apex with broad spur. Apterous female: an-
terior tibia only slightly dilate (L. 104, W. 20), slightly excavate on
inner surface of apical one fourth. Pronotal proportions same as
‘in male. Proportions of antennae: 73: 45: 47: 41, of intermediate
legs: 160: 124: 57: 67; of posterior legs: 185: 143: 17: 30, Con-
nexiva converging abruptly over dorsum of third abdominal seg-
ment then continuing subparallel to last segment where they diverge
slightly. Dorsum of last abdominal segment between diverging
connexiva thickly set with dark hairs. Venter with same minute,
black, conical setae as in male, with those of last segment being
concentrated in two or three rows along each side on posterior
edge of segment. Intermediate femur flattened beneath on basal
one third. Posterior trochanter unarmed; posterior femur only,
slightly incrassate (L. 135, W. 27) and armed at apical two fifths
with one long, bent spine followed by five rapidly decreasing spines
ending before apex; also armed with two short spines anterior to
first and fourth spines of main row of spines. Posterior tibia straight
and armed on basal half with small scattered teeth, apparently un-
armed at apex. Winged male: anterior tibiae as in apterous male.
Pronotum (L. 162, W. 133) prolonged into short spiniform process
at apex (L. 10). Proportions of antennae and legs as in apterous
male. Wings extend beyond apex of genital segments. Venter
Srupy oF THE GENUS RHAGOVELIA 833
without carina well defined for first two segments, which are de-
pressed, but otherwise as in apterous male. Minute, conical setae
on venter as in apterous male. Form and armature of posterior
legs same as in apterous male.
Comparative notes: This species resembles R. armata (Burmeis-
ter) and R. planipes Gould, but can be separated from either of
those species by the extent of the minute, black, conical setae
which extend down on the proepisternum, mesosternum and venter
of abdomen in R. scabra sp. nov. R. scabra sp. nov. also resembles
R. solida sp. nov. R. scabra sp. nov. can be separated from R. solida
sp. nov. by the greatly dilate and excavate anterior tibia in the male,
and by the presence of the conical setae beneath the venter on all
but the last segment of the female.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype, winged male. Paratypes, 8 apterous males,
6 apterous females; paramorphotypes, 4 winged males. Described
from following specimens: Apterous forms collected in Costa Rica
at an altitude of 2000 mtrs. Rio Sarapiqui, by Heinrich Schmidt.
Winged males, collected in Panama, C. A. Potrerillos, 2-18-1935, by
D. V. Brown. All type specimens are in the Francis Huntington
Snow Entomological Collections, University of Kansas.
Data on distribution: In addition to the type specimens, speci-
mens from the following localities have been examined:
Costa Rica: San José, 6 & 7, 1931, Heinrich Schmidt, 4 apterous
males, 2 apterous females.
PanaMa: Potrerillos, 2-22-1935, D. V. Brown, 1 winged male;
Coll. by J. W. McSwain for Robert Wind, Feb. 4, 1935, 2 winged
males.
Rhagovelia solida sp. nov.
(Pl. VI, fig. 11)
Size: Length Width
5.11 mm. apterous male 1.53 mm. apterous male
5.48 mm. apterous female 1.96 mm. apterous female
Color: General color red-brown. Anterior portion of pronotum
Slightly lighter in color than remainder of pronotum but not set
off as distinct band. Venter yellow-brown with last three segments
of antennae, tibiae and tarsi of all legs darker brown.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva vertical; anterior
tibia moderately dilate and excavate in male. Apterous male: an-
terior trochanter unarmed; anterior tibia only moderately dilate
834 Tue University Science BULLETIN
(L. 100, ‘W. 20) and very slightly excavate on apical third. Pro-
notum broader than long (L. 84, W. 100) and evenly rounded be-
hind. Mesonotum hidden by pronotum. Metanotum much wider
than long (L. 7, W. 95). Proportions of antennae: Seg. I: TH: IIT:
IV:: 80: 45: 49: 43; of intermediate legs: Fem.: Tib.: Tars. I:
Tars. III:: 160: 120: 55: 65; of posterior legs: 140: 125: 17: 32.
Abdomen tapers evenly to apex. Connexiva semivertical. Venter
with pronounced median carina between posterior coxae becoming
evanescent. between posterior trochanters. Posterior trochanter,
seemingly unarmed. Posterior femur greatly incrassate (L. 140, Ww.
47) and armed on basal one third with row of uniform, closely set,
small teeth, followed just after basal one third with one large curved
spine then approximately seven smaller spines to apex. Also armed,
with uniform row of small spines anterior to main row of spines.
Posterior tibiae straight; armed with one row of subequal teeth,
with those on basal half being larger and much more closely-set
than those of apical half; armed at apex with slightly curved spur.
Apterous female: pronotum formed much as in male (L. 90, W.
108), metanotum also as in male (L. 7, W. 100). Proportions of an-
tennae: 84: 43: 46: 40; of intermediate legs: 153: 115: 53: 68;
of posterior legs: 130: 182: 16: 33. Dorsum of abdomen narrow
after first three segments. Connexiva vertical converging abruptly
after first three abdominal segments, then continuing almost par-
allel. Thick patch of long brown hairs at apex of abdomen. Venter
without median carina. Intermediate femur flattened beneath on
basal one third. Posterior trochanter unarmed. Posterior femur not
as incrassate as in male (L. 180, W. 32) and armed at middle with
one long spine followed by seven rapidly decreasing spines to apex.
Posterior tibia straight, armed on basal half with close set black
teeth, seemingly unarmed on apical half except for spur at apex.
Winged forms: unknown.
Comparative notes: This species resembles R. scabra sp. nov. but
can be separated by the absence of the greatly dilate and excavate
anterior tibia and the unarmed posterior trochanters in the male.
The absence of the minute, black, conical setae beneath the venter
of the female on the second and third from the last abdominal seg-
ments of R. solida sp. nov. separates the females. R. solida sp. nov.
resembles R. armata (Burmeister) and R. planipes Gould, but can
be separated from either of those species by the extent of the
minute, black, conical setae which extend down on the proepister-
num, mesosternum, metasternum and at least the basal two ab-
dominal segments in R. solida sp. nov.
Srupy oF THE GENUS RHAGOVELIA 835
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 10 apterous. males, 10 apterous females. De-
scribed from specimens labeled: “Costa Rica, Alt. 2000 mtr., Rio
Sarapiqui, Heinrich Schmidt.” All type specimens are in the Francis
Huntington Snow Entomological Collections, University of Kansas.
Data on distribution: Known only from type series.
Rhagovelia tayloriella Kirkaldy
(PI. VI, fig. 12)
1900. Rhagovelia tayloriella Kirkaldy, Ent., vol. 33, p. 72.
1901. Rhagovelia tayloriella, Kirkaldy, Ent., vol. 84, p. 808 (mentions in key).
1931. Rhagovelia tayloriella, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 44
(redescribes; gives distribution as Jamaica).
Size: Length Width
4.35 mm. apterous male 1.33 mm. apterous male
4.50 mm. apterous female 1.50 mm. apterous female
4.55 mm. winged male 1.70 mm. winged male
4.85 mm. winged female 1.80 mm. winged female
Color: General color brown-black, clothed with golden pubes-
cence. Anterior band of pronotum yellow behind vertex of head,
becoming pruinose behind eyes. Dorsum of last five abdominal
segments of female blue-gray to gray-black, not clothed with
golden pubescence or longer hairs. Dorsum of abdomen of male
pruinose at sides, thickly clothed with golden pubescence and
scattered long, brown hairs. Margins of connexiva yellow. Venter
blue-gray with last two (male) to five (female) segments yellow
beneath. Base of antennae, margins of all acetabulae, all coxae,
anterior and posterior trochanters, and base of anterior femora
yellow. Wings brown; veins black.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva vertical; anterior
tibia greatly dilate and excavate in male. Apterous male: anterior
trochanter unarmed. Anterior tibia greatly dilate (L. 85, W. 22)
and excavate on apical one half. Pronotum rounded behind (Li
81, W. 85); metanotum short on midline (L. 6, W. 87). Proportions
of antennae: Seg. I: Il: III: IV:: 70: 40: 45: 36; of intermediate
legs: Fem.: Tib.; Tars. I: Tars. II:: 143: 110: 50: 60; of posterior
legs: 118: 120: 13: 28. Abdomen tapers evenly to apex. Con-
nexiva semivertical. Venter with median carina becoming evanes-
cent on third or fourth from last segment. No minute, conical,
black setae on base of jugum of head ‘or along sides of venter of
last abdominal segment. Venter of last abdominal segment beneath
roundly and deeply excavate on posterior two thirds with median
836 Tue Universiry Science BULLETIN
furrow on anterior one third. Posterior trochanter armed with
four or five small teeth. Posterior femur moderately incrassate
(L. 118, W. 80). Armed on basal one third with inconspicuous
row of small teeth; armed just beyond basal one third with long
spine followed by approximately ten smaller, gradually decreasing
spines to apex; also armed on apical one half with an anterior row
of small, inconspicuous, subequal spines. Posterior tibia straight;
armed within with subequal teeth, and spur at apex. Apterous
female: anterior tibia only slightly dilate (L. 87, W. 15) and
flattened within. Pronotum rounded behind (L. 75, W. 92); meta-
notum as in male (L. 7, W. 95). Proportions of antennae: 74: 40:
42: 36; of intermediate legs: 145: 108: 45: 62; of posterior legs:
118: 122: 14: 26. Dorsum of abdomen narrow after first three
segments. Margins of connexiva vertical and subparallel after
first three segments. Dorsum of last five abdominal segments with-
out golden pubescence and long brown hairs except at apex of last
segment where there is a clump of long brown hairs. Venter
without median carina. Minute, black, conical setae absent as in
male. Intermediate femur flattened within on basal one fourth.
Posterior trochanter unarmed. Posterior femur not as incrassate
as in male (L. 118, W. 23); armed beyond middle with one slender
spine followed by four or five much smaller, rapidly decreasing
spines to apex. Posterior tibia straight and apparently unarmed
except for slender spur at apex. Winged forms: proportions and
armature similar to apterous forms. Wings just covering apex of
genital segments. Winged male: venter formed as in apterous
male. Pronotum continued into short (L. 14) process at apex.
Winged female: pronotum continued at apex into long, curved
process which arises at angle of forty-five degrees and curves
around near apex and is directed slightly downward (L. 40).
Comparative notes: This species resembles R. collaris ( Burmeis-
ter). R. tayloriella Kirkaldy can be separated from R. collaris
(Burmeister) by the formation of the venter of the last abdominal
segment in the male, and by the dorsum of the abdomen of the fe-
male being without short or long hairs for the last five segments.
The armature of the posterior femur is also somewhat reduced in all
forms of R. tayloriella Kirkaldy. Kirkaldy gives as one of his char-
acters the incrassate condition of the third antennal segment of the
male. This is an extremely variable condition and occurs to some
extent in all the species of the collaris group.
Data on types: Kirkaldy states that the type is the winged male.
In the J. R. de la Torre-Bueno Collection (now at the University of
Stupy OF THE GENUS RHAGOVELIA 837
Kansas) is a winged male bearing two hand-written labels: “Jam.
Black R. Elysium Estate, Portland (2), C. B. T., 4-IV-99.” and
“Rhagovelia tayloriella Type Male.” This winged male is the holo-
type of this species. With this specimen in the Bueno Collection
were a winged female and 8 apterous males. These specimens bear
labels in the same hand-writing as the type and are doubtlessly co-
type material. These specimens are all in the Francis Huntington
Snow Entomological Collections, University of Kansas.
Data on distribution: Recorded only from Jamaica. In addition
to the type specimens, specimens have been examined from the
following localities:
Jamaica: Buff Bay R., 3-20-37, Chester Roys, 18 apterous males,
20 apterous females, 1 winged female; Claremont, Baron Hill,
Trelawney, 3-4-28, L. G. Perkins, 4 apterous males, 1 winged male,
1 winged female; St. Ann, 8-1929, Lilly Perkins, 3 apterous males,
12 apterous females; Bath, St. Thomas, 8-29-37, Chester Roys, 8
apterous males, 11 apterous females; Spanish Town, April 10, 1937,
Chester Roys, 52 apterous males, 4 winged males, 20 apterous fe-
males; Pt. Antonio, April, 1906, Van Duzee (J. R. de la Torre-Bueno
Collection), 1 apterous male, 1 apterous female.
Oxsesa GROUP
Group characteristics: The obesa group can be characterized as
consisting of those species of the genus Rhagovelia in which the
dorsum of the abdomen of the apterous female is narrow after the
first three segments, with the connexiva reflexed for the last four
segments; all females have the intermediate femur dorsoventrally
flattened; the terminal genital segments of both sexes are rounded
or triangular but not mucronate. The winged forms vary in fre-
quency of occurrence in the different species; the wings just cover
the apex of the genital segments.
The following species comprise this group:
1. R. distincta Champion 4, R. oriander Parshley
2. R. knighti Drake and Harris 5. R. rivale Bueno
8. R. obesa Uhler
Kry to Species oF THE OsresA Group
di Apterous female with apex of pronotum continued into a long, ele-
vated process; apterous male with pronotum triangular with apex
extending onto metanotum, mesonotum exposed at sides,
oriander
» Apterous female with pronotum not produced at apex; pronotum of
apterous male not extending back over metanotum......... 2
8.(2)
4,(3)
Tue Universiry Scruence BULLETIN
Connexiva of apterous female diverging over apex of last abdominal
segment, apex of connexiva rounded as seen from the side; venter
of last segment of male flattened beneath with prominent hairy
SidGes lo CoG ee, i a Gee distincta
Connexiva of apterous female close together or meeting at apex;
apex of connexiva at least a 90° angle as seen from the side;
male with last.segment notas above i...) ou elias 8
Dorsum of abdomen of apterous male with only traces of median
shining areas on a few segments; posterior trochanter of male
unarmed, Mesonotum of apterous female tumid ...... rivale
Dorsum of abdomen of apterous male and dorsum of metanotum
with broad shining areas; posterior trochanter armed with several
small teeth in male. Mesonotum of apterous female tumid only
AG SIGS dh At Albee ciadin 1 PUN, Ape ws a ce iie ays 4
Margins of connexiva of apterous female with first two segments
curved; exposed portion of mesonotum longer than exposed
portion; Of metanotims Gnd. batviad. 0) ls eal. obesa
Margins of connexiva of apterous female with first two segments
straight; exposed portion of mesonotum shorter than exposed
POTION OL MevAMOUI sp ys eta eee 7 ig knighti
Rhagovelia distincta Champion
(Pl. VII, fig. 4)
Rhagovelia mexicana Signoret, Bull. Soc. Ent. Fr. (5); 7: p. 53 (no
description ).
Rhagovelia distincta Champion, Biol. Centr. Amer. Het., vol. 2, p. 135.
Rhagovelia distincta, Kirkaldy, Ento., vol. 83, p. 72.
Rhagovelia distincta, Kirkaldy, Ento., vol. 34, p. 808.
Rhagovelia distincta, Banks, Cat. Nearctic Hemip.-Heter., p. 27.
Rhagovelia distincta, Van Duzee, Check List of Hemip.
Rhagovelia distincta, Van Duzee, Cat. of Hemip. of America N. of
Mexico, p. 435.
Rhagovolte distincta, Hungerford, Kansas Univ. Sci. Bull., vol. 11, p. 129.
Rhagovelia distincta, Torre-Bueno, Ent. News, vol. 32, p. 274. ?
Rhagovelia distincta, Drake and Harris, Proc. Biol. Soc. Washington, vol.
AO, p. 134 (report on variations in posterior legs).
Rhagovelia excellentis Drake and Harris, Proc. Biol. Soc. Washington,
vol. 40, p. 131 (describe from Colorado).
Rhagovelia distincta, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 25 (re-
describes, gives distribution as South-western United States and Mexico;
places R. excellentis Drake and Harris as a synonym).
Rhagovelia distincta arizonensis Gould, Kansas Univ. Sci. Bull., vol. 20,
p. 26 (describes from Arizona). ;
Rhagovelia distincta cadyi Gould, Kansas Univ. Sci. Bull., vol. 20, p. 27
(describes from Wyoming).
Rhagovelia distincta harmonia Gould, Kansas Univ. Sci. Bull., vol. 20,
p. 28 (describes from Arizona).
Rhagovelia distincta modesta Gould, Kansas Univ. Sci. Bull., vol. 20,
p. 28 (describes from Mexico).
Rhagovelia distincta proxima Gould, Kansas Univ. Sci. Bull., vol. 20,
p. 29 (describes from Colorado).
Rhagovelia distincta valentina Gould, Kansas Univ. Sci. Bull., vol. 20,
p. 29 (describes from Texas and Arizona).
Rhagovelia distincta, Gould, Ann. Ent. Soc. America, vol. 26, p. 466.
Srupy oF THE GENUS RHAGOVELIA 839
Size: Length Width
4.50 nim, apterous male 1.33 mm. apterous male
4.30 mm. apterous female 1.40 mm, apterous female
4.92 mm. winged male 1.66 mm. winged male
5.20 mm. winged female 1.72 mm. winged female
Color: General color black, clothed with fine, brown pubescence.
Anterior band of pronotum interrupted at middle; orange behind
vertex of head becoming pruinose behind eyes. Median, black,
shining areas on all segments of dorsum of abdomen of male, and
on, more or less, basal three segments of dorsum of abdomen of
female. Margins of connexiva orange. Venter blue-gray to gray-
black; venter of last abdominal segment black beneath. Genital
segments orange to brown beneath. Base of antennae, margins of
all acetabulae, anterior and posterior trochanters, all coxae, and
base of anterior femora orange. Wings brown; veins darker at base
becoming inconspicuous on apical one half of wing.
‘Structural characteristics: Dorsum of apterous female narrow
after first three segments; connexiva of apterous female reflexed for
last four segments. Apterous male: anterior trochanter unarmed,
or with agglutinated brown hairs sometimes resembling feeble spine.
Anterior tibia only slightly dilate and somewhat excavate on apical
one half. Pronotum slightly carinate down middle; broadly rounded
behind (L. 72, W. 84); occasionally exposing apex of mesonotum.
Metanotum short on midline (L. 8, W. 87). Proportions of an-
tennae: Seg. I: I: IH: IV:: 73: 45: 44: 85; of intermediate legs:
Fem.: Tib.: Tars. II: Tars. IfI:: 160: 115: 54: 65; of posterior legs:
130: 115: 15: 28. Abdomen tapers evenly to apex. Connexiva
semivertical. Venter without median carina. Sparsely scattered,
minute, conical, black setae on base of jugum of head, inner margin
of proepisternum, and in row along posterior margin at sides of
venter of last abdominal segment. Venter of last abdominal seg-
ment flattened beneath, somewhat excavate on posterior one half;
flattened median portion bounded at sides by more or less distinct
ridges which usually bear a hairy prominence at approximately
middle of segment. Posterior trochanter armed with five or six
small, brown teeth. Posterior femur moderately incrassate (L. 180,
W. 30); armed on basal one third with a row of short, closely-spaced
spines, and just before middle with one long spine followed by nine
or ten shorter, gradually decreasing spines to apex; also armed with
an anterior row of inconspicuous spines on apical one half. Pos-
terior tibia straight; armed within with one row of teeth with those
at apical one fourth sometimes stouter; also armed with slender
spur at apex. Apterous female: anterior tibia flattened within on
840 Tue Universiry ScieNcE BULLETIN
apical one third, and not dilate. Pronotum broadly rounded at
apex; depressed in center of posterior margin (L. 70, W. 88); meso-
notum emarginate on midline, protruding slightly (L. 8) from under
pronotum on each side of midline. Metanotum short on midline
(L. 10, W. 89). Proportions of antennae: 70: 40: 38: 34; of inter-
mediate legs: 143: 100: 50: 62; of posterior legs: 120: 118: 15: 27.
Dorsum of abdomen narrow after first three segments. Connexiva
with group of long brown hairs at apex of second segment; reflexed
over dorsum of abdomen for last four segments; apices of connexiva
diverging over last abdominal segment. Intermediate femur dorso-
ventrally flattened. Posterior trochanter unarmed. Posterior femur
only slightly incrassate (L. 120, W. 18); armed just before apical
one third with one long spine followed by four or five much smaller,
rapidly decreasing spines to apex. Posterior tibia straight; armed
within with inconspicuous teeth with those on basal one half stouter,
also armed with slender spur at apex. Winged forms: proportions
and armature similar to apterous forms. Winged male: Venter of
last segment of abdomen formed as in apterous male. Pronotum
produced into short (L. 10) process at apex. Winged female: pro-
notum with variable apex. Three types can be found: (1) as in
winged male with only short process, (2) with long, bent process
swollen at apex (L. 87), or (8) with long bent process with tip ex-
panded and emarginate on midline (L. 40).
Comparative notes: This species is closely related to R. obesa
Uhler. The males of R. distincta Champion can be separated from
the males of R. obesa Uhler by the differently formed venter of the
last abdominal segment which is slightly raised on the midline with a
depression each side in R. obesa Uhler, while in R. distincta Cham-
pion that segment is flattened beneath with carinate ridges at the
sides. The apterous females can be separated by the apices of the
connexiva which are diverging and rounded in R. distincta Cham-
pion while contiguous and produced into short knoblike processes in
R. obesa Uhler. So far as is known the winged females of R. obesa
Uhler always have the apex of the pronotum produced into a long
process swollen at the apex and deeply emarginate, while the pro-
notum can be formed in any of three ways in R. distincta Champion.
Type 2 (see under structural characteristics; winged female) is by
far the most common type in R. distincta Champion. The types of
apices to the pronotum are found mixed in many long series. Type
1 and type 2 occur in the same series, and type 2 and type 3 occur
together. In no case has there been found characters on the apter-
ous forms or on the winged males which would allow separation
Srupy OF THE GENUS RHAGOVELIA 841
into groups to correspond to the different types of winged females.
The varieties of R. distincta arizonensis Gould, R. distincta cadyi
Gould, R. distincta harmonia Gould, R. distincta modesta Gould,
R. distincta proxima Gould, and R. distincta valentina Gould, are
indistinguishable from R. distincta Champion.
Data on types: This species was described from specimens from
Mexico, Champion’s type material is located in the British Mu-
seum.
Data on distribution: Recorded from Southwestern United States
and Mexico. Specimens have been examined from the following
localities (new distribution for major political areas are indicated
with an asterisk):
* GuaTEMALA: Trib. de las Vacas, Guatemala City, [V-4-47, M.
Holloway (U. S. National Museum Collection ), 12 apterous males,
2 apterous females; 1 km. S. Palin, IV-13-47, R. R. Miller (U. S.
National Museum Collection), 1 apterous male, 1 winged male, 1
apterous female; Rio Momostenango, 2 mi. E. Momostenango, Il-
31-47 (U. S, National Museum Collections), 1 apterous male, 1
winged male, 1 apterous female, 2 winged females; Agua Caliente,
28-1-05 (J. R. de la Torre-Bueno Collection), 1 apterous female,
1 winged female.
* Honpuras: Tegucigalpa, VI-18-18, F. J. Dyar Coll. (U.3.
National Museum Collections), 1 apterous male, 1 apterous female.
Mexico: Chihuahua: San Pedro Rio, between Chihuahua and
Naica, 6-22-34, Smith and Dunkle, 7 apterous males, 20 winged
males, 2 apterous females, 19 winged females; San Antonio, 7-15-27,
R. H. Beamer (Type series of R. distincta modesta Gould), 19
apterous males, 1 winged male, 9 apterous females, 1 winged female.
Distrito Federal: Xochimilco, June 21, 19384, H. E. Hinton, 40
apterous males, 16 winged males, 16 apterous females, 19 winged
females.
Guerrero: Puente de Ixtla, 7-12-37, H. D. Thomas, 17 apterous
males, 15 winged males, 14 apterous females, 7 winged females.
Hidalgo: Durango, July 28, 1938, H. D. Thomas, 19 apterous
males, 1 winged male, 17 apterous females, 4 winged females;
Durango, Sept. 24, 1938, H. D. Thomas, 1] apterous males, 1 winged
male, 17 apterous females, 2 winged females; San Antonio, near
El Salto, 5000 ft. a.s.l., semi-tropical, June 10, 1937, M. Embury,
11 winged males, 5 winged females.
842
Tue Universiry Science BULLETIN
Jalisco: Mazatepec, IV-29-44, J. G. Shaw, 23 apterous males, 4
winged males, 44 apterous females, 5 winged females.
México: Real de Arriba, Alt. 1960 mtrs. May-June, 1933, H. E.
Hinton, 23 apterous males, 74 winged males, 20 apterous females,
§2 winged females; Progreso Industrial, 2300 m.a.s.l., in brook,
Feb, 12, 1938, A. Dampf, 2 apterous males, 2 winged males, 5 apter-
ous females, 2 winged females.
Michoacdn: Morelian, 9-4-88, H. D, Thomas, 11 apterous males,
2 winged males, 11 apterous females; 20 mi. E. Zitacuaro, [X-19-38,
H. D. Thomas, 8 apterous males, 6 winged males; El Sabino,
Uruapan, 7-10-36, H. D. Thomas, 91 apterous males, 101 apterous
females; Tancitar, Alt. 6586 ft., Perm. Pool, VIII-8-40, F. Schacht,
63 winged males, 48 winged females.
Morelos: 7-14-36, H. D. Thomas, 8 apterous males, 2 winged
males, 4 apterous females, 2 winged females; Cuernavaca, 7-8-36,
H. D. Thomas, 2 apterous males, 2 apterous females, 3 winged
females; Cuernavaca, April 15, 1946, J. & D. Pallister (U.S. National
Museum Collection), 7 apterous males, 4 apterous females; Cuer-
navaca, 10-5-36, H. D. Thomas, 18 apterous males, 3 winged males,
10 apterous females,
Nuevo Leén: Ville Santiago, Alt. 2000 ft., Aqueduct pool, VI-18-
40, F'. Schacht, 4 winged males, 3 apterous females, 8 winged fe-
males; Galeana, VII-30-39, H. Heogstraal, 16 apterous males, 8
winged males, 5 apterous females, 1 winged female; Galeana, VII-
8-39, H. Heogstraal, 8 apterous males, 14 winged males, 2 apterous
females, 3 winged females; Ville de Santiago, VIJI-8-39, H. Heog-
straal, 8 apterous males, 6 winged males, 3 apterous females, 5
winged females.
Oaxaca: Oaxaca, 5000 ft., semi-desert, Irrig. Id., semi-tropical,
Aug. 20, 1937, M. Embury, 13 apterous males, 2 winged males, 2
apterous females; Oaxaca, Aug. 25, 1937, H. D. Thomas, 22 ap-
terous males, 3 winged males, 8 apterous females, 1 winged female.
Puebla: Tehuacin, July 19-25, 1987, H. D. Thomas, 29 apterous
males, 2 winged males, 8 apterous females, 4 winged females.
Vera Cruz: Orizaba, July 30, 1987, H. D. Thomas, 14 apterous
males, 2 winged males, 10 apterous females, 2 winged females.
Unrrep States: Arizona: Shinono Creek, 1000 ft. above mouth
Grand Canyon, 9-83-23, R. C. Moore (type series R. distincta arizon-
ensis Gould), 37 apterous males, 12 winged males, 42 apterous fe-
males, 8 winged females; Oak Creek Cn., 8-9-32, R. H. Beamer, 19
Srupy or THE GENUS RHAGOVELIA 843
apterous males, 1 winged male, 15 apterous females; Huachuca
Mts., 7-8-32, R. H. Beamer, 16 winged males, 27 winged females;
Huachuca, 5-27-37, W. Benedict, 6 winged males, 10 winged fe-
males; Granite Dells, 7-12-47, L. D. Beamer, 4 apterous males, 8
apterous females; Baboquivari Mts., 7-18-32, R. H. Beamer, 8
winged males, 8 winged females; Cottonwood River, 7-12-47, A.C;
Michener, 7 apterous males; Oak Creek Canyon, TAlB-ATS. La Di
Beamer, 8 apterous males, 7 apterous females; Grand Canyon, 8-
11-27, P. A. Readio (type series of R. distincta harmonia Gould),
2 apterous males, 4 apterous females; Janez Springs, VII-1-41, R. H.
Beamer, 2 apterous males; Santa Rita Mts., 7-17-82, R. H. Beamer,
2 winged males, 1 winged female; Sabino Canyon, XII-20-41, H. B.
Hungerford, 3 apterous males, 4 apterous females; Chiricahua Mts.,
7-8-32, R. H. Beamer, 3 winged females.
California: San Diego Co. 7-7-29, L. D. Anderson, 5 apterous
males, 1 winged male, 32 apterous females, 1 winged female. San
Diego Co:, 7-4-29, P. W. Oman, 31 apterous males, 5 winged males,
39 apterous females, 1 winged female; San Diego Co., 7-28-29, P. W.
Oman, 9 apterous males, 3 winged males, 24 apterous females; San
Diego Co., 7-4-29, L. D. Anderson, 6 apterous females; San Diego
Co,, 7-4-29, R. H, Beamer, 1 apterous male, 4 apterous females;
Dulzura, 8-9-35, R. H. Beamer, 45 apterous males, 54 apterous fe-
males; Dulzura; 8-9-35, Jack Beamer, 20 apterous males, 83 ap-
terous females; India, 7-24-29, L.'D. Anderson, 39 apterous males,
41 apterous females, 1 winged female; Lemon Cove, 7-24-29, R. H.
Beamer, 15 apterous males, 15 apterous females; Lemon Cove, 7-
96-29, L. D. Anderson, 12 apterous males, 1 winged male, 19 ap-
terous females; Campo, VII-18-40, L. C. Kuitert, 34 apterous males,
33 apterous females; Lene, 8-19-38, R. I. Sailer, 12 apterous males,
15 apterous females; Nipomo, 7-28-35, R. H. Beamer, 19 apterous
males, 8 apterous females; Arroyo, Seca R. 8-8-38, R. I. Sailer, 5
apterous males, 4 apterous females; Palm City, 8-7-35, R. H. Beamer,
2 apterous males; Fresno, June 20, 1926, C. J. Drake, 1 apterous
male, 1 apterous female; Jacumba, VII-17-40, L. C. Kuitert, 1 ap-
terous male, 8 winged males, 2 apterous females, 1 winged female;
Pine Valley, 7-27-36, D. W. Craik, 12 apterous males, 2 winged
males, 25 apterous females; Pine Valley, 7-27-36, R. I. Sailer, 4 ap-
terous males, 2 winged males, 8 apterous females; Winters, Aug. 6,
1928, R. H. Beamer, 4 apterous males, 1 winged male, 13 apterous
females, 1 winged female; Pacific, VII-9-40, L. C. Kuitert, 8 apter-
ous males, 13 apterous females.
844 Tue Universiry ScreNce BULLETIN
Colorado: Grand Junction, 8-15-36, M. B. Jackson, 14 apterous
males, 33 apterous females; N. W. Colorado, Aug. 25, 1920, R. C.
Moore, 12 apterous males, 27 apterous females; Dolores, Aug. 15,
1925, C. J. Drake (paratypes of R. excellentis Drake and Harris),
4 apterous males, 1 apterous female; Boulder Creek, 8-16-25, Beamer
& Lawson (type series of R. distincta proxima Gould), 11 apterous
males, 5 winged males, 9 apterous females, 3 winged females; Craig,
VIII-18-40, L. C. Kuitert, 30 apterous males, 9 apterous females.
New Mexico: Las Cruces, 1925, C. B. Lebert, 1 apterous male,
1 apterous female.
Texas: Ft. Davis, 6-20-47, L. D. Beamer, 6 apterous males, 2 ap-
terous females; Valentine, July 18, 1927, R. H. Beamer (type series
of R. distincta valentina Gould), 43 apterous males, 34 apterous fe-
males; Jeff Davis, 7-19-38, 7 apterous males, 1 winged male, 2 ap-
terous females; Austin, 4-01 (J. R. de la Torre-Bueno Collection),
8 apterous males, 1 apterous female.
Utah: Zion Canyon, 4600 ft., Oct. 1, 1934, Bryant, 1 apterous
male, 5 apterous females; Vernal, 8-2-47, R. E. Elbel, 7 apterous
males, 8 apterous females.
Wyoming: North Platte, Sept. 2, 1926, G. Cady (type series of
R. distincta cadyi Gould), 59 apterous males, 60 apterous females.
Rhagovelia knighti Drake and Harris
(Pl. VIL, fig. 5)
1927, arewreie knighti Drake and Harris, Proc. Biol. Soc. Washington, vol.
» p. 138.
1931, Rhagovelia knighti, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 35 (re-
describes ).
1933. Rhagovelia knighti, Gould, Ann. Ent. Soc. America, vol. 26, p. 468 (rec-
ords from Arkansas).
Size: Length Width
8.65 mm. apterous male 1.16 mm. apterous male
3.73 mm. apterous female 1,33 mm. apterous female
Color: General color black, clothed with golden pubescence.
Anterior band of pronotum interrupted at middle; orange behind
vertex of head, becoming pruinose behind eyes. Dorsum of all
abdominal segments and metanotum with median, black, shining
spots in male; female with last three segments of dorsum of ab-
domen occasionally so marked. Margins of connexiva orange. Ven-
ter blue-gray. Last segment of venter of abdomen orange to brown.
Base of antennae, margins of all acetabulae, anterior and posterior
coxae, all trochanters, and base of anterior femur yellow to orange.
Srupy OF THE GENUS RHAGOVELIA 845
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments; connexiva of apterous fe-
male reflexed for last four segments. Apterous male: anterior tro-
chanter armed only with group of agglutinated brown hairs which
may resemble a spine. Anterior tibia flattened within; not dilate.
Pronotum indistinctly carinate; rounded or somewhat triangular be-
hind (L. 59, W. 67). Mesonotum usually somewhat exposed.
Metanotum short on midline (L. 7, W. 70). Proportions of an-
tennae: Seg. I: I: Il: IV:: 56: 33: 36: 33; of intermediate legs:
Fem.: Tib.: Tars. II: Tars. III:: 132: 90: 40: 53; of posterior legs:
100: 85: 10: 22. Abdomen tapers evenly to apex. Connexiva semi-
vertical. Venter without median carina. Venter of last segment of
abdomen depressed on each side of a median, somewhat triangular,
anterior area. Sparsely scattered, minute, black, conical setae on
proepisternum, or seemingly absent. Posterior trochanter armed
with eight to ten small teeth. Posterior femur moderately incras-
sate (L. 100, W. 27); armed with row of small teeth on basal two
fifths, and with one long spine at basal two fifths followed by eight
or nine smaller, gradually decreasing spines to apex; also armed with
inconspicuous row of small spines anterior to main armature. Pos-
terior tibia straight; armed within with equal teeth and slender spur
at apex. Apterous female: pronotum broadly rounded behind;
slightly depressed on posterior one fourth (L. 64, W. 75). Posterior
margin of mesonotum broadly emarginate behind. Metanotum
longer than in male (L. 10, W. 85). Proportions of antennae: 57:
84: 87: 35; of intermedite legs: 122: 84: 44: 56; of posterior legs:
97: 95: 13: 25. Dorsum of abdomen narrowed after first three seg-
ments, Connexiva reflexed over abdomen on last four segments,
nearly touching or touching at apex; produced at apex into short,
knoblike processes which are elevated less than the length of tarsal
segment II of intermediate leg above dorsum of first genital seg-
ment. Minute, conical, black setae as in male. Intermediate femur
dorso-ventrally flattened from basal one fourth to apical one fourth.
Posterior trochanter unarmed. Posterior femur not as incrassate as
in male (L. 97, W. 16); armed at apical two fifths with one long
spine followed by three or four much shorter, inconspicuous spines
to apex. Posterior tibia unarmed except for inconspicuous spur at
apex. Winged forms: unknown.
Comparative notes: This species resembles R. obesa Uhler. The
apterous female of R. knighti Drake and Harris can be separated
from R. obesa Uhler by the lack of the tumid abdominal segments,
846 Tue Universiry SCIENCE BULLETIN
and by the extent of the apical processes of the connexiva which are
much shorter and closer to the dorsum of the first genital segment.in
R. knighti Drake and Harris; the males can be separated by the
less exposed mesonotum which is only partly exposed in R. knighti
Drake and Harris whereas the mesonotum of the male of R. obesa
Uhler is generally exposed and approximately equal in length with
the metanotum.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, several apterous males and females; all taken at
Hollister, Missouri, Sept. 5-10, 1925, H. H. Knight, collector. The
holotype, allotype, and several paratypes are in the personal col-
lection of Dr. C. J. Drake. Paratypes are also in the collection of
H. H. Knight, Iowa State College, and the Francis Huntington Snow
Entomological Collections, University of Kansas.
Data on distribution: Recorded only from Missouri and Arkansas.
In addition to the paratypes, specimens have been examined from
the following localities (new distribution records for major political
areas are indicated with an asterisk) :
Unrrep States: Arkansas: Scott Co., 8-23-28, R. H. Beamer, 2
apterous males, 3 apterous females; Carroll Co., 9-17-32, G. E.
Gould, 2 apterous males, 2 apterous females; Sharpe Co., 9-14-82,
G. E. Gould, 1 apterous male, 1 apterous female; Polk Co., 8-21-28,
R. H. Beamer, 2 apterous males, 3 apterous females.
Missouri: Barton Co., 9-17-32, G. E. Gould, 1 apterous male,
2 apterous females; Jasper Co., 9-17-32, G. E, Gould, 1 apterous
male.
* Oklahoma: Ottawa Co., Dev. Promenade, 9-21-32, L. D. Tuthill,
10 apterous males, 7 apterous females,
Rhagovelia obesa Uhler
(Pl. VII, fig. 6)
1871. Rhagovelia obesa Uhler, Proc, Boston Soc. Nat, Hist., vol. 14, p. 107.
1878. Rhagovelia obesa, Uhler, Proc, Boston Soc, Nat, Hist., sell 19. p. 434
(amplifies original description).
1884, Rhagovelia obesa, Uhler, in Kingsley’s Natural History, vol. 2, p. 260.
1886. Rhagovelia obesa, Uhler, Check List Hemip., p, 18.
1894. Rhagovelia obesa, Uhler, Proc, Zool, Soc, London, p. 215.
1894, ree end obesa, Uhler, California Acad, Sci., ser, 2, vol. 4, p, 258
(probably should refer to R. distincta).
1896, Biazovelt ia obesa, Lethierry and Severin, Cat, Gén, des Hémip,, vol. 3,
0. 00,
1898. Ei hgobette obesa, Champion, Biol, Centr, Amer, Het., vol. 2, p. 185
(gives as closely related to R. distincta).
1901. Rhagovelia obesa, Kirkaldy, Ent., vol. 34, p. 308.
1907. Rhagovelia obesa, Torre-Bueno, * dauddins Ent., vol, 39, —p. 61 (good
on biology).
Srupy oF THE GENUS RHAGOVELIA 847
1910. Rhagovelia obesa, Banks, Cat. of Nearctic Hemip.-Heter,, p. 28.
1911. Rhagovelia obesa, Van Duzee, Check List Hemip. i
1917. Rhagovelia obesa, Van Duzee, Cat. of Hemip, of America N. of Mexico,
1919, Rhagovelia obesa, Hungerford, Kansas Univ. Sci. Bull., vol. 11, p. 128
(quotes Uhler’s 1878 description ).
1922. Rhagovelia obesa, Parshley, S. Dakota St. Col. Tech, Bull., vol. 2, p. 20
(compares to R. oriander).
1923. Rhagovelia obesa, Torre-Bueno in Connecticut Geo. and Nat. Hist.
Survey Bull., vol. 34, p. 418 (gives distribution as N. E, United States).
1924, Rhagovelia flavicincta Torre-Bueno, ‘Trans, American Ent. Soc., vol. 50,
p. 249 (describes from North Carolina).
1924. Rhagovelia arctoa Torre-Bueno, Trans, American Ent. Soc,, vol. 50, p.
250 (describes from Minnesota ).
1925, Rhagovelia obesa, Torre-Bueno, Trans, American Ent. Soc., vol. 50, p.
250 (redescribes ).
1925. Rhagovelia obesa, Hussey, Jour. New York Ent. Soc., vol, 83, p. 68
(compares with R. choreutes ).
1926, Rhagovelia flavicincta, Blatchley, Heter. of East. N. America, p. 998
(redescribes ).
1926. Rhagovelia obesa, Blatchley, Heter, of Eastern N. America, p. 996
(redescribes ).
1927, Rhagovelia obesa, Drake and Harris, Proc. Biol, Soc, Washington, vol.
40, p. 181 (suggest synonomy of R. arctoa and R. flavicincta with
R. obesa).
1928, Rhagovelia obesa, Torre-Bueno, Cornell Univ, Memoir 101,
1931, Rhagovelia obesa, Gould, Kansas Univ, Sci. Bull., vol, 20, p. 86 (re-
describes ).
1931, mihi flavicincta, Gould, Kansas Univ. Sci, Bull., vol. 20, p. 31
(redescribes, records from Virginia).
1931. Rhagovelia arctoa, Gould, Kansas Univ, Sci, Bull, vol. 20, p. 16 (re-
describes, records from Michigan, and Ontario, Canada).
1938. Rhagovelia obesa, Gould, Ann, Ent. Soc, America, vol. 26, p. 469
(records from Indiana).
1936. Rhagovelia flavicincta, Coker, Millsap and Rice, Bull. Brooklyn Ent. Soc.,
vol, 31, p. 84 (excellent discussion of the function of the swimming
plume of the intermediate leg).
Size: Length Width
8.75 mm. apterous male 1.13 mm. apterous male
8.73 mm. apterous female 1.33 mm. apterous female
4.00 mm, winged male 1.53 mm. winged male
4.55 mm. winged female 1.62 mm. winged female
Color: General color black, clothed with golden pubescence.
Anterior band of pronotum interrupted at middle; orange behind
vertex of head becoming pruinose behind eyes. Dorsum of male
with black shining spots on all abdominal segments, on metanotum
and usually a spot on each side of midline of mesonotum; dorsum
of female pruinose black with occasionally indefinite black shining
areas on some of basal abdominal segments. Margins of connexiva
Orange. Venter blue-gray. Venter of last abdominal segment black
beneath. Base of antennae, all acetabulae, anterior and posterior
coxae, all trochanters, and basal one fifth to one half of anterior
femur yellow to orange.
Structural characteristics: Dorsum of abdomen of apterous. fe-
male narrow after first three segments, connexiva reflexed for apical
848 THe Universiry ScreNcE BULLETIN
four segments. Apterous male: anterior trochanter armed only with
group of agglutinated hairs which may resemble a spine, but which
can be easily rubbed off. Anterior tibia flattened within; not dilate.
Pronotum inconspicuously carinate on midline; rounded behind (L.
53, W. 62). Mesonotum exposed; usually somewhat truncate at
apex (L. 12, W. 64); metanotum short on midline (L. 7, W. 72).
Proportions of antennae: Seg. I: II: III: IV:: 56: 82: 34: 32; of
intermediate legs: Fem.: Tib.: Tars. II: Tars. III:: 180: 93: 48: 52;
of posterior legs: 100: 83: 12: 20. Abdomen tapers evenly to apex.
Connexiva semivertical. Venter without median carina. Venter
of last segment of abdomen depressed on each side of midline be-
neath. Minute, black, conical setae not conspicuous if present.
Posterior trochanter armed with several dark teeth. Posterior femur
somewhat incrassate (L. 100, W. 21); armed on basal one half with
row of small, inconspicuous spines; armed just before middle with
one long spine followed by eight or ten smaller, gradually decreas-
ing spines to apex; also armed on apical one half with inconspicuous
row of smaller spines anterior to main row of spines. Posterior
tibia straight and armed within with small, regular teeth and spur
at apex. Apterous female: anterior trochanter without agglutinated
hairs resembling a spine. Pronotum almost straight to slightly
emarginate on midline at apex; depressed on midline on posterior
one third (L. 60, W. 80). Mesonotum somewhat flattened in center,
becoming tumid toward sides; broadly emarginate on midline at
apex (L. 8, W. 81). Metanotum (L. 18, W. 84) and first two ab-
dominal segments noticeably swollen. Proportions of antennae: 56:
31: 33: 32; of intermediate legs: 125: 90: 45: 57; of posterior legs:
98: 95: 12: 22. Dorsum of abdomen narrowed after first three
segments. Connexiva usually with group of brown hairs at apex of
second segment; reflexed for last four segments; usually contiguous
at apex; prolonged into spinelike points at apex, these points stand-
ing well above dorsum of first genital segment (height, 16). Ab-
domen usually bent upward at an angle of 25 degrees. Intermediate
femur dorso-ventrally flattened from basal one fourth to apical one
fourth. Posterior trochanter unarmed. Posterior femur only slightly
incrassate (L. 98, W. 16); armed at apical one third with one small
spine followed by five or six much smaller, gradually decreasing
spines to apex. Posterior tibia apparently unarmed except for spur
at apex. Winged forms: proportions and armature similar to ap-
terous forms. Wings just cover apex of genital segments. Winged
male: venter formed as in apterous male. Pronotum triangular be-
Srupy OF THE Genus RHAGOVELIA 849
hind (L. 110, W. 103), not produced into spinelike process. Winged
female: pronotum produced at apex into long process which is
abruptly expanded and deeply emarginate at apex (25).
Comparative notes: This species resembles R. knighti Drake and
Harris. The females of R. obesa Uhler can be separated from those
of R. knighti Drake and Harris by the curved first two segments of
the connexiva, which in R. knighti Drake and Harris are straight,
also the swollen condition of the dorsum of the first two abdominal
segments in R. obesa Uhler will serve to separate the females. The
males of the two species can be separated by the reduced condi-
tion of the spines on the basal one third of the posterior femur
which in R. obesa Uhler number six to eight, and are small, widely-
spaced spines; the spines on the base of the femur of R. knighti
Drake and Harris number from 12 to 16 closely-spaced spines which
are variable in size but always in a closely-packed row.
R. arctoa Bueno and R. flavicincta Bueno are indistinguishable
from R. obesa Uhler. The spine on the anterior trochanter of the
male is formed from agglutinated hairs which easily become broken,
and the color variations are no more than would be expected from
a widely distributed species. Therefore, the names R. arctoa Bueno
and R. flavicincta Bueno become synonyms of the name R. obesa
Uhler.
Data on types: This species was described from specimens from
Massachusetts. Ubhler’s type material is in the United States Na-
tional Museum.
Data on distribution: Recorded from Ontario, Canada and the
following states of the United States: California (R. distincta [?]),
Colorado (R. distincta [?]), District of Columbia, Florida (R.
choreutes [?]), Illinois, Indiana, Maine, Maryland, Massachusetts,
Michigan, Minnesota, New Jersey, New York, North Carolina,
Pennsylvania, South Carolina, Tennessee, Utah (R. distincta [?]),
Vermont, and Virginia. Specimens from the following localities
have been examined (new distribution records for major political
units are indicated with an asterisk ):
Unrrep Sratrs: * Alabama: Burnsville, 7-20-30, R. H. Beamer,
6 apterous males, 17 apterous females, 2 winged females.
* Georgia: Perry, 8-12-39, J. D. Beamer, 10 apterous males, 7 ap-
terous females; Macon, 7-25-30, Paul W. Oman, 27 apterous males,
14 apterous females.
28—3378
850 THe Untiverstry ScteENCE BULLETIN
Maryland: Lalseland, 9-12-31, P. W. Oman, 7 apterous males, 1
winged male, 2 apterous females, 1 winged female; Beltsville, 9-25-
32, P. W. Oman, 30 apterous males, 50 apterous females, 1 winged
female.
Michigan: Lake Gogebic, 8-18-37, R. H. Beamer, 28 apterous
males, 9 apterous females; Iosco Co., Aug. 22, 03, E. H. Frothing-
ham (J. R. de la Torre-Bueno Collection), 1 apterous male, 1 ap-
terous female; Cheboygan Co., 8-2-37, H. B. Hungerford, 4 apter-
ous males, 6 apterous females; Cheboygan Co., 1935, M. Sanderson,
2 apterous males, 2 apterous females; Cheboygan Co., 7-16-39, H. B.
Hungerford, 3 apterous males, 1 apterous female; Cheboygan Co.,
7-15-42, E. L. Todd, 3 apterous males, 2 apterous females; Douglas
Lake, 7-23-27, H. B. Hungerford, 1 apterous male, 4 apterous fe-
males; Douglas Lake, 7-24-27, H. B. Hungreford, 3 apterous males,
9 apterous females.
Minnesota: Cook Co., L. Devil Track Cr., Aug. 11, 1922, H. B.
Hungerford, 52 apterous males, 60 apterous females; Cook Co., L.
Devil Track Cr., Aug. 11, 1922, H. B. Hungerford (type series of
R. arctoa Bueno, in part), 11 apterous males, 2 apterous females;
H. B. Hungerford, 9 apterous males, 6 apterous females; Cook Co.,
Little Devil Track Cr., Aug. 12, 1922, Wm. E. Hoffman (paratypes
of R. arctoa Bueno, in part), 3 apterous females; Bengal, Aug. 18,
1922, Wm. E. Hoffman (paratypes of R. arctoa Bueno, in part), 3
apterous males, 1 apterous female; Bengal, Aug. 18, 1922, Wm. E.
Hoffman (J. R. de la Torre-Bueno Collection), 9 apterous males,
7 apterous females.
North Carolina: Northampton Co., 15 Sept. 1929, G. E. Gould,
6 apterous males, 4 apterous females; Raleigh, Oct. 20, 1920, C. 5.
Brimsley (J. R. de la Torre-Bueno Collection, paratypes of R. flavi-
cincta Bueno, in part), 1 apterous male, 3 apterous females; Swan-
nanoa, Oct. 5, 1915, R. W. Leiby (J. R. de la Torre-Bueno Collec-
tion, paratypes of R. flavicincta Bueno in part), 4 apterous females;
Raleigh, Oct. 18, 1915, R. W. Leiby, 3 apterous males.
* New Hampshire: Glen, 8-20-34, P. M. McKinstry, 108 apterous
males, 70 apterous females.
New Jersey: Rahway, R., Cranford, 6-8-04, (J. R. de la Torre-
Bueno Collection), 9 apterous males, 1 winged male, 8 apterous
females, 1 winged female; Lakehurst, 5-9-03, (J. R. de la Torre-
Bueno Collection), 8 apterous females; Glen Ridge, 23-4-06, (J. 8:
de la Torre-Bueno Collection), 2 apterous males, 2 apterous females.
Srupy oF THE GENUS RHAGOVELIA 851
New York: Staten Id. 9-7-05, (J. R. de la Torre-Bueno Collection ),
14 apterous males, 7 apterous females; Ithaca, 1917, H. B. Hunger-
ford, 4 apterous males, 4 apterous females; Niagara; 8-9-05, Chick.
de la Torre-Bueno Collection), 2 apterous males, 2 apterous females;
White Plains, 5-9-20, Bueno (J. R. de la Torre-Bueno Collection )
1 apterous female; Nepperham, 30-9-05 (J. R. de la Torre-Bueno
Collection), 2 apterous females,
Ohio: Columbus, 10-9-15, C. J. Drake (J. R. de la Torre-Bueno
Collection), 1 winged male.
Pennsylvania: Dingmans Fry, 8-20-46, L. D. Beamer, 8 apterous
males, 8 apterous females.
South Carolina: Batesburg, 8-24-30, P. W. Oman, 20 apterous
males, 27 apterous females; Tigerville, 8-26-30, P. W. Oman, 26
apterous males, 9 winged males, 17 apterous females, 7 winged
females; Batesburg, 8-24-30, P. W. Oman, 74 apterous males, 70
apterous females, 1 winged female.
Tennessee: Coal Creek, 8-27-30, P. W. Oman, 8 apterous males,
10 apterous females; Coal Creek, 8-27-30, J. Nottingham, 16 apter-
ous males, 17 apterous females; Clarksville, July 15, 1989, E. G.
egenek, 8 apterous males, 8 apterous females; Clarksville, July
12, 1989, J. D. Beamer, 7 apterous males, 1 apterous female; Elk
Valley, 8-27-30, P. W. Oman, 2 apterous males.
Vermont: Springfield, 8-22-84, P. McKinstry and M. Griffith, 48
apterous males, 86 apterous females.
Virginia: Brunswick Co., Sept. 15, 1929, George E. Gould and
Thelma Gould, 4 apterous females; Warrenton, Aug. 21, 1947,
Shirley Bacon, 3 apterous males, 2 apterous females.
Rhagovelia oriander Parshley
(Pl. VIL, fig. 7)
1922, Rhagovelia oriander Parshley, S$. Dakota State Ent, Tech, Bull, vol. 2,
p. 19
1923, Rhagovelia oriander, Torre-Bueno, in Connecticut Geo, & Nat, Hist.
Survey Bull., vol. 34, p. 418 (mentions possible occurrence in Con-
necticut ).
1925, Rhagovelia oriander, Torre-Bueno, Trans, American Ent, Soc., vol, 50,
19 p. 250 (redescribes ).
25. Rhagovelia oriander, Hussey, Jour. New York Ent, Soc., vol, 33, p. 69
19 (records some variations from Bueno’s redescription ).
26. Rhagovelia oriander, Blatchley, Het. of East. N, America, p. 999 (re-
19 describes, records from Indiana and Illinois). :
27. Rhagovelia oriander, Drake and Harris, Proc, Biol. Soc, Washington,
vol, 40, p, 131 (describes winged forms; record from Ohio, Iowa, South
193) Dakota and Minnesota). :
1. Rhagovelia oriander, Gould, Kansas Univ, Sci. Bull., vol. 20, p. 88 ( gives
new record from Kansas, redescribes ).
852 Tue Universiry Science BULLETIN
1933. Rhagovelia oriander, Gould, Ann, Ent, Soc. America, vol, 26, p. 469
(gives new record from Michigan).
Size: Length Width
3.59 mm. apterous male 1,13 mm. apterous male
8.65 mm. apterous female 1.83 mm, apterous female
3.44 mm. winged male 1.48 mm. winged male
8.89 mm. winged female 1.63 mm. winged female
Color: General color brown-black, clothed with golden pubes-
cence. Pronotum with interrupted orange band behind vertex of
head becoming pruinose behind eyes. Median area of dorsum of all
abdominal segments of male shining brown-black only last one or
two of female shining brown-black. Margins of connexiva orange.
Venter blue-gray. Venter of last abdominal segment black beneath.
Base of antennae, margins of all acetabulae, anterior and posterior
coxae, all trochanters and base of anterior femora yellow. Wings
dark brown-black.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva reflexed for apical
four segments. Apterous male: anterior trochanter armed with
brown spine. Anterior tibia flattened within for apical two thirds,
and not dilate. Pronotum with posterior margins almost straight,
angulate at apex (L. 67, W. 71); mesonotum covered by pronotum
on median line, but exposed at sides; metanotum partially (4) cov-
ered by apex of pronotum. Proportions of antennae: Seg. I: I: III:
IV:: 54: 82: 37: 33; of intermediate legs: Fem.: Tib.: Tars. II:
Tars. III:: 124: 90: 45: 58; of posterior legs: 100: 85: 11: 24. Ab-
domen tapers evenly at first, angle of taper increasing on last three
segments. Venter without median carina. Posterior trochanter
armed with approximately eight small teeth. Posterior femur mod-
erately incrassate (L. 100, W. 23); armed on basal half with row of
small spines, armed at middle with one long, slender spine followed
by five or ten smaller, gradually decreasing spines to apex; also
armed with an anterior row of very small spines on apical one half.
Posterior tibia straight; armed within with one row of closely set
teeth which become slightly stouter toward apex; armed at apex
with small spur. Apterous female: anterior trochanter unarmed;
anterior tibia as in male. Pronotum with apex produced into 4
long (L. 24), knobbed process elevated at an angle of approximately
thirty degrees; longer than wide (L.—to tooth under apical process
—93, W. 82); mesonotum exposed at sides of pronotum. Metanotum
and first abdominal segment covered by apex of pronotum. Pro-
portions of antennae: 57: 32: 37: 33; of intermediate legs: 123: 85:
45: 55; of posterior legs: 100: 95: 18: 24. Abdomen broad at first,
Srupy OF THE GENUS RHAGOVELIA 853
narrowed on last three segments. Connexiva reflexed tightly against
abdomen for last three segments; gradually narrowing and touch-
ing at apex or may continue subparallel and not touch at apex.
Intermediate femur dorso-ventrally flattened from basal one fourth
to just before apex. Posterior trochanter unarmed. Posterior femur
less incrassate than that of male, (L. 100, W. 15); armed at apical
one third with sharply-bent, long spine followed by approximately
seven smaller, decreasing spines to apex. Posterior tibia straight
and apparently unarmed except for small spur at apex. Winged
forms: Wings reaching slightly beyond apex of genital segments.
Winged male: pronotum acutely triangularly produced at apex,
with an inconspicuous median carina. Winged female: pronotum
produced into a semierect, somewhat cylindrical process.
Comparative notes: The characters of the pronotum serve to
separate this species from all known Rhagovelia. The angulate
pronotum of the male exposing the mesonotum only at the sides,
and the pronotum of the apterous female with a knobbed elevated
Process at the apex serve to separate R. oriander Parshley from R.
knighti Drake and Harris, which it mostly closely resembles.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; numerous male and female paratypes. The. type series was
collected at Brookings, South Dakota, 7-8-21, by H. C. Severin.
The holotype, allotype and many paratypes are in Parshley’s per-
sonal collection. Paratypes are also in the collections of the South
Dakota State Agricultural Experiment Station and Torre-Bueno’s
collection (now in the Francis Huntington Snow Entomological
Collections, University of Kansas). Morphotypes and paramorpho-
types are males and females taken at Rockbridge, Hocking Co.,
Ohio, July 7, 1916, and are in the personal collection of Dr Ged:
Drake,
Data on distribution: Recorded only from the United States.
States havin g records of occurrence of this species are South Dakota,
Thdiana, Illinois, Ohio, Iowa, Minnesota, Kansas and Michigan.
In addition to the two paratypes the following specimens have been
examined:
Unrrep States: Illinois: Chicago, New Lenox, 28-VII-27, H. M.
Parshley, 1 apterous male, 1 apterous female; Piatt Co., 4-25-32,
Geo. E. Gould, 2 apterous males.
Indiana: Kosciusko Co., Aug. 11, 1982, Geo. E. Gould, 1 apter-
ous male, 1 apterous female; DeKalb Co., Aug. 11, 1982, G. E.
Gould, 1 apterous male; Lafayette, 1-Oct.-1931, 1 apterous male,
Tue Universiry SCIENCE BULLETIN
854
1 apterous female; Warren Co., Aug. 28, 1932, G. E. Gould, 1 ap-
terous female; Parke Co., Aug. 28, 1932, G. E. Gould, 1 apterous
male; Lafayette, 26-Sept.-31, 1 apterous male, 1 apterous female;
White Co., July 25, 1932, G. E. Gould, 1 apterous male; Montgomery
Co., Aug. 1982, G. E. Gould, 1 apterous female; Howard Co., Oct.
28, 19382, G. E. Gould, 1 apterous male.
Kansas: Woodbine, Aug. 26, 1925, Beamer and Lawson, 35 ap-
terous males, 42 apterous females.
Michigan: St. Joseph Co., Aug. 7, 1932, G. E. Gould, 1 apterous
male.
South Dakota: Brookings, Sept. 27, 1922, H. C. Severin Coll., 1
apterous male, 1 apterous female (labeled “Topotype, R. oriander
Parsh. by Parshley.”).
Rhagovelia rivale Bueno
CPI, Vil, 8g. 8)
1924, Begoves rivale Torre-Bueno, Trans. American Ent. Soc., vol. 50,
p. 247.
1927. Rhagovelia rivale, Drake and Harris, Proc. Biol. Soc, Washington, vol.
AO, p. 133 (record from Colorado).
1931. Rhagovelia rivale, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 40 (re-
describes ).
1933. Rhagovelia rivale, Gould, Ann. Ent. Soc. America, vol. 26, p. 469 (rec-
ords from Missouri).
Size: Length Width
3.50 mm. apterous male 1.16 mm. apterous male
4.32 mm. apterous female 1.33 mm. apterous female
3.83 mm. winged male 1,53 mm. winged male
4.75 mm. winged female 1.78 mm. winged female
Color: General color brown to pruinose black, clothed with
golden pubescence. Anterior band of pronotum partially inter-
rupted at middle in some cases; orange behind vertex of head be-
coming pruinose behind eyes. Lateral and posterior margins of
pronotum usually orange. Dorsum of abdomen of male with last
abdominal segment with rectangular, shining, brown to black
area; three or four other abdominal segments may show smaller,
irregular, median, shining areas. Dorsum of abdomen of female
without median shining areas. Margins of connexiva orange tO
yellow. Venter blue-gray. Venter of last abdominal segment
yellow to orange. Base of antennae, margins of all acetabulae, all
coxae and trochanters, and base of anterior femora yellow.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrow after first three segments, connexiva reflexed for apical
four segments. Apterous male: anterior trochanter armed only
with agglutinated hairs which may resemble a spine. Anterior tibia
Srupy oF THE GENUS RHAGOVELIA 855
arcuate; not dilate nor excavate. Pronotum sharply rounded at
apex: indistinctly carinate on midline (L. 55, W. 64). Mesonotum
broadly exposed; truncate at apex (L.10, W.66). Metanotum
short on midline (L. 7, W. 72). Proportions of antennae: Seg. I:
II: III: IV:: 57: 34: 34: 32; of intermediate legs: Fem.: Tib.: Tars.
Il: Tars, II[:: 125: 90; 47: 51; of posterior legs: 93: 92: 10: 22.
Abdomen tapering rather evenly to apex, angle of taper increasing
for last three segments, Connexiva semivertical. Minute, black,
conical setae absent, Venter without median carina. Venter of
last abdominal segment shallowly depressed on each side of rather
broad, central area on posterior one half. Posterior trochanter un-
armed, Posterior femur slightly incrassate (L. 98, W. 15); armed
on basal one half with one or two small spines or may be unarmed
on basal one half; armed at middle with one slender spine followed
by approximately four much smaller, subequal spines to apex.
Posterior tibia straight; armed within with subequal teeth and
stout spur at apex. Apterous female: anterior femur slightly
arcuate; anterior tibia straight. Pronotum usually wider than long
(L. 73, W. 84) but may vary to subsequal or even longer than wide.
Mesonotum tumid; truncate to slightly emarginate at apex (L. 14,
W. 84). Metanotum slightly tumid in center and at sides (L. 10, W.
85). Proportions of antennae: 67: 40: 40: 38; of intermediate legs:
155: 108: 56: 57; of posterior legs: 112: 111: 15: 25. Dorsum of
abdomen narrow after first three segments. Connexiva reflexed for
last four segments; occasionally contiguous over dorsum of abdo-
Men for a few segments and then separating, or may be parallel
and not contiguous. Intermediate femur dorso-ventrally flattened
from basal one fourth to apical one fourth. Posterior trochanter
unarmed, Posterior femur scarcely incrassate (L. 112, W. 16);
armed just before apical one third with one long spine followed by
three much smaller, subequal spines toward apex. Posterior tibia
straight and seemingly unarmed. Winged forms: Proportions and
armature similar to apterous forms. Wings just cover apex of
8enital segments. Winged male: pronotum continued into short
Process at apex (L. 10). Venter formed as in apterous male.
inged female: pronotum continued into long, cylindrical process
Which is rounded at apex (LL. 85).
Comparative notes: This species resembles R. obesa Uhler and
- knighti Drake and Harris. R. rivale Bueno can be separated
from both of those species by the unarmed posterior trochanter in
the male, by the tumid mesonotum in the female, and by the lateral
856 THe University SCtENCE BULLETIN
and posterior margins of the pronotum being more or less orange.
In addition the dorsum of the apterous male has some segments
without or at most inconspicuously developed median shining areas.
Data on types: The holotype is an apterous female. Bueno states
that the morphotype is a winged female; however the specimen
bearing the label “Morphotype, female” is a winged male, Paratypes
are 5 apterous males, 1 apterous female, 1 winged male, 1 winged
female and 1 nymph. The specimens were all collected at Dyke-
man’s Bridge, Kansas, July 1922, by H. B. Hungerford. The type
series is in the Francis Huntington Snow Entomological Collections,
University of Kansas.
Data on distribution: Recorded from Kansas, Missouri, and Colo-
rado. In addition to the type series, specimens from the following
localities have been examined (new distribution for major political
areas is indicated with an asterisk):
Unrrep Starrs: Colorado: Wray, Aug. 4, 1925, C. J. Drake, 1
apterous male.
Kansas: Kiowa Co., July 5, 1923, R. H. Beamer, 2 apterous males,
1 apterous female; Wilson Co., 866 ft., 1916, R. H. Reamer, 2 apter-
ous males, 2 apterous females; Neosho Co., 9-20-31, G. E. Gould, 2
apterous males, 1 apterous female; Osborne Co., 1557 ft., Aug. 3;
1912, F. X. Williams, 1 apterous male, 1 apterous female; Marshall
Co., 9-25-32, G. E. Gould, 1 apterous male, 1 apterous female; Cloud
Co., 9-24-82, G. E. Gould, 2 apterous males, 1 apterous female; Phil-
lips Co., 8-8-25, H. J. Grady, 1 apterous male, 1 apterous female;
Riley Co., June 17, Popenoe, 1 apterous male, 1 apterous female;
Scott Co., 8-10-25, H. J. Grady, 1 apterous male, 1 apterous female;
Republic Co., 7-11-25, R. H. Beamer, 17 apterous males, 26 apterous
females; Dykeman’s Bridge, July, 1922, H. B. Hungerford, 1 apterous
male, 2 apterous females, 1 winged female, 14 nymphs; Dickinson
Co., 6-25, C. H. Martin, 5 apterous males, 5 apterous females; Mar-
shall Co., July 13, 1925, R. H. Beamer, 18 apterous males, 13 aptet-
ous females.
Missouri: Macon Co., 9-25-32, G. E. Gould, 2 apterous males, 2
apterous females; Linn Co., 9-25-82, G. E. Gould, 1 apterous male,
2 apterous females.
* Oklahoma: Ottawa Co., Dev. Promenade, 9-21-32, L, D, Tut-
hill, 27 apterous males, 2 winged males, 25 apterous females, 2
winged females.
* Texas: Kerr Co., 7-20-28, R. H. Beamer, 2 apterous females.
Srupy oF THE GENUS RHAGOVELIA 857
AINSLIEL GROUP
Group characteristics: The ainsliei group can be characterized as
consisting of those species of the genus Rhagovelia having the dor-
sum of the abdomen of the apterous female narrow after the first
three segments, the intermediate femur of the female dorso-ventrally
flattened, and the terminal genital segment prolonged into a sharply-
pointed process in both sexes. The winged forms are rather rare;
the wings just cover the apex of the genital segments.
The following species comprise this group:
I. R. ainslieci Drake and Harris 8. R. gracilis sp, nov.
2. R. becki Drake and Harris
Kry To Species oF THE AINSLIEL Group
1. Body more than or approximately 4 times as long as wide. . . . becki
Body less than or approximately 3.5 times as long as wide... ..
2.(1) Terminal spines on connexiva of female large and extending almost
back to last genital segment; metasternum of male carinate with
ies)
the segment depressed on each side................. gracilis
No terminal spines on apex of connexiva of female; metasternum
Ol MBI NOW aS BOVE cs es oye me ee ey ainsliei
Rhagovelia ainsliei Drake and Harris
(Pl. VII, fig. 1)
1933, Rhagovelia ainslici Drake and Harris, Proc. Biol. Soc. Washington, vol.
46, p. 50
Size: Length Width
4.00 mm. apterous male 1.20 mm. apterous male
3.65 mm, apterous female 1.32 mm. apterous female
4.20 mm. winged male 1.52 mm. winged male
4.15 mm, winged female 1.50 mm. winged female
Color; General color black; clothed with brown pubescence.
P ronotum with interrupted yellow band on anterior margin becom-
mg pruinose behind eyes. Margins of connexiva yellow. Venter
blue-gray. Venter of last abdominal segment brown beneath. Base
of antennae, margins of all acetabulae, anterior and posterior coxae,
all trochanters (intermediate trochanter varies from yellow to:
brown), and base of anterior femur yellow. Wings brown with
black veins.
Structural characteristics: Terminal genital segment mucronate
at apex. Apterous male: anterior trochanter armed with prominent
Prown spine. Anterior tibia straight, not dilate, and flattened on
Mner surface for apical one half. Pronotum with length subequal
to width (L. 74, W. 73), broadly rounded behind, coarsely punc-
tate, and with fairly distinct median, longitudinal carina; metanotum
858 Tue Universtry ScreNcE BULLETIN
short on midline (L. 70, W. 80). Proportions of antennae: Seg. I:
Il: UL: IV:: 67: 44: 30: 30; of intermediate legs: Fem.: Tib.: Tars.
Il: Tars. III:; 146: 95: 56: 56; of posterior legs: 110: 112: 14: 26.
Abdomen tapers evenly to apex. Venter with median carina well
defined on basal two abdominal segments; last ventral abdominal
segment strongly impressed on each side of median, somewhat tri-
angular portion. First genital segment conspicuously clothed with
long, pale hairs beneath; apex of last genital segment mucronate.
Minute, black, conical setae on base of jugum of head, on pro-
episternum, and on venter of abdomen. Posterior trochanter armed
with four or five small, black teeth. Posterior femur moderately to
strongly incrassate (L. 110, W. 32). Armed on basal two fifths
with row of closely-set, short spines, and at basal two fifths with one
long spine followed by approximately seven shorter, subequal
spines, gradually decreasing to apex; also armed on apical one half
with an anterior row of small, subequal spines. Posterior tibia
straight, or on specimens with greatly incrassate hind femur can be
curved outward on apical one third; armed within with one row of
stout teeth with those at apical one third often larger than others;
armed at apex with stout spur. Apterous female: anterior trochan-
ter unarmed. Anterior tibia as in male. Pronotum formed as in
male (L. 78, W. 77); metanotum short on midline (L. 7, W. 82).
Proportions of antennae: 60: 35: 30: 28; of intermediate legs: 182:
87: 50: 58; of posterior legs: 107: 117: 15: 27. Connexivum strongly
reflexed against sides of abdomen; margins not contiguous. Abdo-
men bent upwards at angle of forty-five degrees for last three seg-
ments. Venter without median carina. Apex of last genital seg-
ment mucronate. Minute, black, conical setae as in male, Inter-
mediate femur dorso-ventrally flattened and excavate beneath from
basal one fourth to apical one fourth. Posterior trochanter unarmed.
Posterior femur not incrassate (L. 107, W. 15). Armed at apical
two fifths with one long spine followed by four or five much smaller,
gradually decreasing spines to apex. Posterior tibia straight, seem-
ingly unarmed except for slender spur at apex. Winged forms:
proportions and armature similar to apterous forms. Apex of geni-
tal segments mucronate. Pronotum of male triangular behind; apex
raised and slightly produced into a short process (L. 7). Pronotum
of female produced into long (L. 55), bulbous process curved
downward near apex, with its apical portion densely clothed with
long, brown hairs.
’
Stupy OF THE GENUS RHAGOVELIA 859
Comparative notes: This species resembles R. gracilis sp. nov. R.
dinslici Drake and Harris can be separated from R. gracilis sp. nov. by
the lack of a spinelike process on the apex of the connexiva of the
female, and by the lack of a median carina with the segment de-
pressed on each side on the metasternum of the male.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; morphotype, winged male; paratypes, several winged and ap-
terous males and females. These specimens were collected in
Guatemala City, Guatemala by C. N. Ainslie. The holotype, allo-
type and morphotype, as well as paratypes, are in the personal col-
lection of Dr. C. J. Drake. A pair of apterous paratypes are in the
Francis Huntington Snow Entomological Collections, University of
Kansas.
Data on distribution: Recorded only from Guatemala. In addi-
tion to the paratype material specimens have been examined from
the following localities (new records for major political areas are
indicated with an asterisk ) :
GuateMaLa: Guatemala City, Ainslie, 1 apterous female (this
Specimen bears a paratype label of R. hambletoni Drake and Har-
tis); Gualan, Jan. 23, 05 (J. R. de la Torre-Bueno Collection), 21
apterous males, 2 winged males, 21 apterous females, 5 winged
females.
* Honpuras: Negrito, Mar. 27, 1923, 1412, T. H. Hubbell, 1 ap-
terous male, 1 winged male.
Rhagovelia becki Drake and Harris
(Pl. VII, fig. 2)
1936. Rhagovelia becki Drake and Harris, Proc. Biol. Soc. Washington, vol. 49,
p. 106.
Size: Length Width
5.44 mm. apterous male 1.29 mm. apterous male
5.23 mm. apterous female 1.24 mm. apterous female
Color: General color black; clothed with light brown pubescence.
Pronotum with anterior band interrupted at middle; orange behind
vertex of head becoming pruinose behind eyes. Pronotum with
darker, median, longitudinal line becoming evanescent on posterior
One fifth, Margins of connexiva orange. Venter blue-gray. Venter
of last abdominal segment black beneath. Base of antennae, mar-
Sins of all acetabulae, anterior and posterior coxae, anterior and
Posterior trochanters, basal one half of intermediate trochanter, and
asal one third of anterior femur orange to yellow.
860 Tue Universiry ScreENCE BULLETIN
Structural characteristics: Terminal genital segment mucronate
at apex. Apterous male: anterior trochanter armed with prominent
brown spine. Anterior tibia straight, flattened on inner surface, and
not dilate. Pronotum coarsely punctate, each puncture surrounded
by a shallow, pruinose depression; with faint, median carina; rather
sharply rounded behind; longer than wide (L. 93, W. 80). Meta-
notum short on midline (L. 6, W. 82). Proportions of antennae:
Seg. I: If: II: IV:: 85: 52: 35: 32; of intermediate legs: Fem.: Tib.:
Tars. II: Tars. III:: 190: 130: 68: 66; of posterior legs: 150: 136: 19:
31. Abdomen elongate, tapers evenly to apex. Terminal genital
segment mucronate. Venter with well defined median carina with
segments depressed on each side on basal two abdominal segments.
Minute, black, conical setae thinly scattered on base of jugum of
head, proepisternum and mesosternum. Venter of last abdominal
segment broadly flattened behind, with ridge along each side of
flattened portion. First genital segment clothed with long yellow
hairs. Posterior trochanter armed with four to six brown teeth.
Posterior femur moderately incrassate (L. 150, W. 35). Armed with
one long spine before middle followed by ten or eleven much
shorter, subequal spines to apex; also armed on basal one half with
row of closely-spaced, small, subequal spines, and from middle to
apex with anterior row of short subequal spines. Posterior tibia
straight; armed within with stout teeth, with those at apical one
third being larger; armed at apex with stout spur. Apterous female:
Anterior trochanter unarmed. Pronotum as in male (L. 91, W. 80);
metanotum as in male (L. 6, W. 78). Proportions of antennae:
77: 46: —: —; of intermediate legs: 165: 110: 61: 66; of posterior
legs: 128: 137: 19: 30. Abdomen above narrow on last four seg-
ments, the posterior margin with two irregular tufts of easily-broken
hairs. Connexiva vertical; last two segments clothed with brown
hair meeting above abdomen; each connexivum ending in mucro-
nate, semierect process thickly clothed with long, dark-brown hairs.
Terminal genital segment mucronate. Venter without median ca-
rina. Minute, black, conical setae as in male. Intermediate femur
flattened beneath, not excavate. Posterior trochanter unarmed. Pos-
terior femur not incrassate (L. 128, W. 18); armed after middle with
one long spine followed by three or four smaller, rapidly decreas-
ing spines to apex. Posterior tibia straight; armed within with sev-
eral inconspicuous, widely-spaced teeth; armed at apex with small
spur. Winged forms: unknown.
Stupy OF THE GENUS RHAGOVELIA 861
Comparative notes: This species resembles R. ainsliei Drake and
Harris. R. becki Drake and Harris can be separated from R. ainsliei
Drake and Harris by the elongate body, the longer than wide pro-
notum, and the mucronate processes at the apex of the connexiva of
the female. R. becki Drake and Harris is also close to R. gracilis
n. sp. but can be separated from that form by the more elongate
body, by the lack of a carina on the mesothorax of the male, and
by the last three segments of the abdomen not being bent upward
in the female.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 27 apterous males and females. The holotype, allo-
type, and 3 male paratypes were collected in the state of Nuevo
Leén, Mexico, June 17, 1934, by D. E. Beck. The remaining para-
types were collected at Reagan Wells, Texas, March 6, 1936, by
M. J. James. All type specimens are in the personal collection of
Dr. C. J. Drake.
Data on distribution: Recorded from Mexico and Texas. Speci-
mens have been examined from the following localities (new dis-
tribution records for major political areas are indicated with an
asterisk ):
Mexico: * Tamaulipas: 3% mi. west Forlon, Aug. 8, 1934, Smith &
Dunkle, 1 apterous male.
Unitep States: Texas: Cocan, 7-6-36, R. H. Beamer, 2 apterous
males, 2 apterous females.
Rhagovelia gracilis sp. nov.
(Pl. VIL, fig. 3)
Size: Length Width
4,15 mm, apterous male 1.26 mm, apterous male
4.33 mm. apterous female 1.36 mm. apterous female
4,25 mm. winged male 1.46 mm, winged male
4,70 mm. winged female 1.53 mm. winged female
Color: General color gray-black. Anterior band of pronotum
interrupted at middle; orange behind vertex of head, becoming
pruinose behind eyes. Pronotum with darker, median, longitudinal
line. Margins of connexiva yellow to orange. Venter gray to blue-
gray. Venter of last abdominal segment brown to black beneath,
Base of antennae, margins of all acetabulae, anterior and posterior
coxae, all trochanters, and base of anterior femora yellow to orange.
Wings brown becoming lighter toward apex, veins black.
Structural characteristics: Terminal genital segment mucronate at
apex. Apterous male: anterior trochanter armed with slender
862
Tue Universiry Scrence BULLETIN
brown spine. Anterior tibia not dilate, slightly flattened within on
apical one third. Pronotum coarsely punctate, sharply rounded
behind, and longer than wide (L. 82, W. 73); metanotum short on
midline (L. 5, W. 75). Proportions of antennae: Seg. I: II: TI:
IV:: 65: 42: 28: 30; of intermediate legs: Fem.: Tib.: Tars. II: Tars.
III:: 148: 94: 55: 55; of posterior legs: 115: 109: 15; 26. Venter
with mesonotum depressed triangularly in center; metanotum de-
pressed on each side of median carina; abdomen with median carina
on basal two segments with venter depressed on each side. Minute,
black, conical setae on base of jugum of head and on proepisternum.
Venter of last abdominal segment excavate beneath with hairy
ridges on each side of excavate area. Terminal genital segment
mucronate. Posterior trochanter armed with six to eight small
teeth. Posterior femur moderately incrassate (L. 115, W. 30).
Armed on basal one half with row of short, closely-set spines, fol-
lowed just beyond middle with one long, curved spine, and six or
seven smaller, subequal spines to apex; also armed with anterior
row of small teeth on apical one half, Posterior tibia straight;
armed with one row of teeth, with those of basal one half being
much more closely set than those on apical half; teeth at apical
one third slightly larger than rest. Apterous female: anterior tro-
chanter unarmed. Anterior tibia not dilate, slightly flattened on
apical one third. Pronotum coarsely punctate, rather broadly
rounded, and slightly longer than wide (L. 87, W. 84); mesonotum
short on midline (L. 6, W. 85). Proportions of antennae: 75: 47:
32: 29; of intermediate legs: 157: 100: 62: 63; of posterior legs:
125: 130: 19: 28. Abdomen narrow above for last four segments;
bent upward at an angle of forty-five degrees for last three segments.
Connexiva closely reflexed over abdomen, converging to third from
last segment, then spreading to just before apex where they con-
verge again. Connexiva terminate in mucronate processes with
their apices almost touching and which are reflexed over dorsum
of first genital segment and extend length of first genital segment.
Stiff groups of agglutinated hairs may be present at posterior angles
of first genital segments or may have been broken away. Last genital
segment mucronate. Venter of abdomen without ventral carina.
Minute, black, conical setae on base of jugum of head and on pro-
episternum. Intermediate femur dorso-ventrally flattened, slightly
excavate beneath from basal one fourth to apical one fourth. Pos-
terior trochanter unarmed. Posterior femur not incrassate (L. 125,
W. 16); armed beyond middle with one long spine followed by five
or six very short, blunt spines to apex. Posterior tibia straight and
)
Srupy oF THE GENUS RHAGOVELIA 863
armed with several feeble teeth on basal one third; armed at apex
with slender spur. Winged forms: proportions and armature similar
to apterous forms. Winged male: pronotum longer than wide (L.
125, W. 95), and drawn out at apex into short-process (L. 11).
Wings just cover mucronate genital segment. Venter formed as in
apterous male. Winged female: pronotum drawn out at apex into
long, bent process (L. 50), which is subequal in diameter through-
out, and thickly beset with short hairs. Connexiva not reflected over
abdomen. Each connexivum terminating in mucronate process
which projects from beneath wing. Terminal genital segment
mucronate and projects from beneath wing.
Comparative notes: This species resembles R. ainsliei Drake and
Harris. R. gracilis sp. nov. can be separated from R. ainsliei Drake
and Harris by the carina of the metasternum of the male, and the
mucronate apices of the connexiva of the female. R. gracilis sp. nov.
is also closely related to R. becki Drake and Harris and can be dis-
tinguished from that species by the carinate metasternum of the
male; by the abdomen of the female which is bent upward at an
angle of forty-five degrees in R. gracilis sp. nov. while remaining
horizontal in R. becki Drake and Harris. Also the general body shape
of R. gracilis sp. nov. is not as elongate as that of R. becki Drake and
Harris.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 6 apterous females. The type series was collected
in the state of Chiapas, Mexico, Gutierrez, Aug. 27, 1937, by H. D.
Thomas. All type specimens are in the Francis Huntington Snow
Entomological Collections, University of Kansas,
Data on distribution: In addition to the type material specimens
have been examined from the following localities:
Mexico: Guerrero: La Sabana, kil. 226 S. of Mex. City, 10-20-36,
H. D. Thomas, 1 apterous male, 9 winged males, 21 winged females;
Rio Agua, kil. 437, S. Mex. City, 10-31-36, H. D. Thomas, 3 apter-
ous males.
Sinaloa: Mazatlan, May 1934, H. Hinton, 2 apterous males.
SPINIGERA Group
Group characteristics: The spinigera group can be characterized
as those Rhagovelia in which the intermediate femur of the female
is transversely constricted at the middle. The dorsum of the abdo-
men of the apterous female is narrow after the first three segments;
864 Tue Universrry ScreNCE BULLETIN
the connexiva is reflexed for the last four segments. Winged forms
are common, the wings just cover the apex of the genital segments.
The following species comprise this group:
1, R. choreutes Hussey 8. R. ignota Drake and Harris
2. R. formosa sp. nov. 4, R. spinigera Champion
Key To THE SPECIES OF THE SPINIGERA GROUP
lL Posterior portion of first genital segment of male very swollen be-
neath; venter of last abdominal segment of male deeply and
roundly excavate behind: ie... jie ni et ..ignota
Posterior portion of first genital segment of male normal beneath;
venter of last abdominal segment of male not formed as above, 2
2.(1) Intermediate femur of male armed with short black spines for
Passel Gx ah. tee i Vis SOL eerste eet ee spinigera
Intermediate femur of male unarmed.............. 0.00.0 e eee 8
3.(2) Anterior trochanter of male armed with a prominent brown spine;
basal one fourth of posterior femur of male without a row of
small teeth; pronotum of apterous forms definitely wider than
Orne MSs od. Var ty ea there ts ‘caeinsy Fits formosa
Anterior trochanter of male unarmed; basal one fourth of posterior
femur of male with a row of small teeth; pronotum of apterous
forms with width subequal to length............. choreutes
Rhagovelia choreutes Hussey
CELE Vil Heh)!
1925. Rhagovelia choreutes Hussey, Jour. New York Ent. Soc., vol. 33, pp.
31-69
1926. Rhagovelia choreutes, Blatchley, Het. of East. N. America, p. 997 (gives
redescription ).
1927. Rhagovelia choreutes, Drake and Harris, Proc. Biol. Soc. Washington,
vol, 40, p. 134 (record additional specimens from Gainsville, Florida,
and new record from Texas).
1931, Rhagovelia choreutes, Gould, Kansas Univ, Sci, Bull., vol. 20, p. 19
(records from Florida, Oklahoma, and New Mexico; gives redescription;
describes winged forms).
Size: Length Width
3.72 mm. apterous male 1.19 mm. apterous male
4,00 mm. apterous female 1.83 mm, apterous female
4,22 mm. winged male 1.70 mm, winged male
4.58 mm. winged female 1.86 mm. winged female
Color: General color black, covered with golden pubescence.
Pronotum with anterior band interrupted at middle, orange behind
vertex of head, becoming pruinose behind eyes. Margins of con-
nexiva orange. Venter blue-gray, Venter of last abdominal seg-
ment brown to black beneath. Base of antennae, margins of all
acetabulae, all coxae and trochanters, and basal one half of anterior
femora yellow. Posterior femora light brown on basal one sixth.
Apical segments of legs brown. Wings brown, veins black.
Srupy OF THE GENUS RHAGOVELIA 865
Structural characteristics: Intermediate femur of female trans-
versely constricted at middle. Apterous male: anterior trochanter
unarmed; anterior tibia not dilate, flattened within, and slightly
excavate on apical one third, Pronotum distinctly but sparsely
punctate, rounded behind, and almost as long as wide (L. 72, W.
73); metanotum short on midline (L. 6, W. 75). Proportions of
antennae: Seg. I: II: II: IV:: 62: 36: 35: 34; of intermediate legs:
Fem.: Tib.: Tars. II: Tars. III:: 117: 92: 44: 60; of posterior legs:
94: 100: 11: 24. Abdomen tapers rather evenly to apex, angle of
taper increases on last three segments. Venter without median
carina, Venter of last two abdominal segments slightly flattened
beneath, lateral margins of flattened area being slightly raised into
hairy calli. Minute, black, conical setae on base of jugum of head
and scattered thinly on proespisternum and venter. Posterior tro-
chanter armed with several small dark teeth. Posterior femur
moderately incrassate (L. 94, W. 24); armed just before middle
with one long spine followed by approximately five smaller,
gradually decreasing spines to apex; also armed with row of small,
knoblike teeth on basal one third, and row of small, subequal teeth
anterior to main row of spines. Posterior tibia straight; armed on
inner surface with small, subequal teeth, with those of basal two
thirds more closely-set and somewhat larger than those of apical
one third; armed at apex with bent spur. Apterous female: Pro-
notum truncate at apex (L. 75, W. 80); metanotum as in male (L.
6, W. 83). Proportions of antennae: 63: 38: 35: 34; of intermediate
legs: 133: 95: 45: 67; of posterior legs: 100: 105: 12: 27, Inter-
mediate femur transversely contricted at middle. Connexiva far
apart at base of abdomen, converging abruptly on fourth segment
from apex and contiguous for last one to three segments; diverging
slightly on posterior half of last segment. Minute, conical, black
setae as in male. Posterior trochanter unarmed. Posterior femur
slightly incrassate (L. 100, W. 22); increasing gradually in diameter
to first spine; armed at middle with one long spine followed by six
or seven shorter, gradually decreasing spines to apex. Posterior
tibia straight, inconspicuously armed on inner surface for basal one
third; armed at apex with bent spur. Winged forms: proportions
and armature similar to apterous forms. Pronotum continued into
almost horizontal process at apex; for male this process is short (L.
10), for female long (L. 43) and thickly set with long hairs. Apex
of wings extend slightly over apex of genital segments.
866 THe UNiversiry SCIENCE BULLETIN
Comparative notes: This species is clearly in the spinigera group,
as the females possess the transversely constricted intermediate
femur, although that fact has not been noted previously. It most
closely resembles R. formosa sp. nov. R. choreutes Hussey can be
separated from R. formosa sp. nov. by the unarmed anterior tro-
chanter in the male, by the armed basal one third of the posterior
femur of the male, and by the length of the pronotum, in the apterous
forms, which in R. choreutes Hussey is approximately as long as
wide. The posterior femur of the female is also formed differently
in the two species, being less abruptly expanded before the middle
in R. choreutes.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; paratypes, 170 apterous males and females. These type
specimens were collected at Gainsville, Florida, on October 13, 1928,
and April 9, 1925, by T. H. Hubbell. The holotype, allotype, and
numerous paratypes are in R, F, Hussey’s collection. Paratypes
are also in the collection of H, G. Barber, J. R. de la Torre-Bueno
(now in Kansas University Collection), C. J. Drake, H. B. Hunger-
ford (in Kansas University Collection), H. M. Parshley, the
museums of the University of Michigan and the University of
Florida, the United States National Museum, and the American
Museum of Natural History. Morphotype specimens are hereby
designated as a winged male as holomorphotype, and a winged fe-
male as allomorphotype. These specimens were collected in Eddy
Co., New Mexico, 7-9-27, by P. A. Readio and are in the Francis
Huntington Snow Entomological Collections, University of Kansas.
Data on distribution: Recorded from Florida, New Mexico, Okla-
homa and Texas. In addition to the type specimens mentioned
above as being in the collection of the University of Kansas, speci-
mens from the following localities have been examined (new rec-
ords on distribution for major political areas are indicated with an
asterisk) :
* Mexico: * Chiapas: La Libertad, 500 m.a.s.l., 1-1-38, Octavio
Utrilla, 20 apterous males, 8 apterous females, 30 nymphs,
* Nuevo Leon: Sabinas Hidalgo, Alt. 1000 ft., pool in river June
14, 1940, H. Heogstraal, 92 apterous males, 63 apterous females.
Unrtep States: Florida: Branford, 7-31-30, Paul W. Oman, 115
apterous males, 124 apterous females; Branford, 7-16-34, P. Mc-
Kinstry, 10 apterous males, 2 apterous females; Branford, 7-16-34,
R. H. Beamer, 2 apterous males, 7 apterous females.
Srupy or THE GENUS RHAGOVELIA 867
* Mississippi: Ireland, 7-7-34, R. H. Beamer, Jr., 3 winged males,
4 winged females.
New Mexico: Eddy Co., 7-9-27, P. A. Readio, 22 apterous males,
14 apterous females.
Oklahoma: Arbuckle Mts., Davis, July, 1925, R. H. Beamer, 22
apterous males, 29 apterous females,
Texas: Brazoria Co., 8-12-28, L. D. Beamer, 1 apterous male;
Hidalgo Co., 7-30-28, R. H. Beamer, 1 apterous male; Kendall Co.,
7-22-28, R. H. Beamer, 9 apterous males, 29 apterous females; Ken-
dall Co., 7-22-28, A. M. James, 4 apterous males, 4 apterous females;
Kerr Co., 7-20-28, R. H. Beamer, 7 apterous males, 40 apterous fe-
males; Menard Co., 7-19-28, R. H. Beamer, 9 apterous males, 12
apterous females; Pinto, 7-7-38; R. I. Sailer, 5 apterous males, 2
winged males, 6 apterous females; Pinto, 7-7-88, D. W. Craik, 7
apterous males, 9 apterous females; San Antonio, 6-25-38, D. W.
Craik, 8 apterous males, 7 apterous females; Sequin, 6-26-38, R. I.
Sailer, 4 apterous females; Starr Co., 7-30-28, R. H. Beamer, 11
apterous males, 4 apterous females.
Rhagovelia formosa sp. nov.
(Pl. VI, fig. 2)
Size: Length Width
8.45 mm. apterous male 1.138 mm. apterous male
8.63 mm. apterous female 1.33 mm, apterous female
8.82 mm. winged male 1.46 mm. winged male
8.92 mm. winged female 1.53 mm. winged female
Color: General color very dark brown, clothed with golden pu-
bescence, Anterior band of pronotum silver-yellow, interrupted
at middle; posterior margins of pronotum slightly tinged with blue-
gray. Connexiva margined with light brown. Venter blue-gray;
venter of last abdominal segment and genital segments yellow to
light brown. Base of antennae, all coxae, all trochanter, and bases
of anterior and posterior femora yellow to light brown. Dorsum
with blue-gray pilosity at juncture with connexivum. Wings lighter
brown than pronotum; veins black.
Structural characteristics: Intermediate femur of female trans-
versely constricted at middle. Apterous male: anterior trochanter
armed with conspicuous brown spine, Anterior tibiae slightly
dilate, slightly flattened on apical half. Pronotum most thickly
punctate along posterior margin and wider than long (1,62, W.
72); metanotum much wider than long (L. 6, W. 70). Proportions
of antennae: Seg. I: I: II: TVs: 55: 28: 28: 29; of intermediate
THe Universiry ScieNCE BULLETIN
868
legs: Fem.: Tib.: Tars. I: Tars. III:: 118: 75: 34: 53; of posterior
legs: 100: 83: 10: 20, Connexiva taper evenly to apex. Venter
without median carina, except occasionally on basal one or two seg-
ments. Minute, black, conical setae on base of jugum of head,
dorsal one half of proepisternum, and at sides of posterior
margin of venter of last abdominal segment. Venter of last
three abdominal segments with median depression forming
longitudinal trough; depression of last abdominal segment bordered
at sides with rather strong, hairy calli, Occasional specimens
may not show this longitudinal depression on venter to any
great extent. Genital segments hairy beneath, but otherwise
normally formed. Posterior trochanter armed only with two or
three feeble brown teeth. Posterior femur greatly incrassate (L.
100, W. 86) and armed before middle with one long, brown-tipped
spine followed by eight smaller, decreasing spines to apex; also
armed with anterior row of eight small, subequal spines which
begins at approximately middle and continues to apex. Posterior
tibiae straight and armed with two rows of stout teeth, with two
or three teeth at middle and apical tooth somewhat enlarged; also
armed with stout spur at apex. Apterous female: pronotum broadly
depressed at middle on posterior margin and coarsely punctate
along posterior border (L. 65, W. 88); metanotum broadly truncate
and wider than long (L. 7, W. 85). Proportions of antennae: 60:
80: 80: 80; of intermediate legs: 113: 73: 33: 56; of posterior legs:
95: 88: 10: 23. Abdomen without ventral carina. Minute, conical,
black setae on jugum of head and proepisternum. Abdomen narrow
beyond first three segments with connexiva reflexed and_ their
margins close together over last three abdominal segments. Apices
of connexiva diverging slightly over apical half of last abdominal
segment, Occasional specimens show dorsum of last one or two
abdominal segments swollen and projecting above connexiva. First
genital segment sloping downward at angle of forty-five degrees.
Intermediate femora transversely constricted at approximately
middle. Posterior trochanter unarmed. Posterior femur with basal
one third subcylindrical (W. 15) and unarmed, abruptly expanding
after basal third (W. 24) and armed just before middle with one
long spine followed by six or seven smaller, decreasing spines to
apex. Also armed after middle with anterior row of approximately
three, small, black, subequal teeth running to apex. Posterior tibia
straight, armed on basal half with fairly strong teeth which are much
reduced on apical half, armed at apex with rather small spur.
Srupy or THE GENUS RHAGOVELIA 869
Winged forms: last segment of antennae of winged male slightly
longer than in apterous male, otherwise proportions and armature
as in apterous forms. Connexiva not reflexed. Venter of abdomen
of male as in apterous male. Formations of intermediate and
posterior femora as in apterous forms. Posterior tips of wings not,
or just barely covering genital segments. Pronotum of male pro-
duced at apex, into short process, while that of female is produced
into long, pointed process standing from dorsum at an angle of
approximately forty-five degrees.
Comparative notes: This species is in the group with R. spinigera
Champion because of the constricted intermediate leg of the fe-
male. It is most closely related to R. ignota Drake and Harris from
which the males may be separated by the formation of the venter
of the abdomen and the normal ventral surface of the first genital
segment, which is greatly enlarged in R. ignota Drake and Harris.
The females may be separated by the form and armature of the
posterior femur. The male claspers are of the same general type
as in R. ignota Drake and Harris, but are broader at the apex and
smaller than those of R. ignota Drake and Harris.
Data on types: Holotype, apterous male, allotype, apterous fe-
male; holomorphotype, winged male; allomorphotype, winged fe-
male; paratypes, 8 apterous males, 3 apterous females; paramorpho-
types, 23 winged males, 12 winged females. Described from a
series labeled: “Acapulco, Guerrero, Kil. 442 S. of Mex. City, 11-
1-36, H. D. Thomas.” All type specimens are in the Francis Hunt-
ington Snow Entomological Collections, University of Kansas.
Data on distribution: In addition to the type. series, specimens
have been examined from the following localities:
GuatemMata: Amatitlan, 8-11-05 (J. R. de la Torre-Bueno Col-
lection), 50 apterous males, 106 apterous females; Gualan, 23-1-’05
(J. R. de la Torre-Bueno Collection), 7 apterous males, 5 apterous
females, 4 winged females.
Honpuras: Copan, 2-7-37, Chester Roys, 1 apterous male, 1 ap-
terous female, 1 winged female; Copan, 2-18-37, Chester Roys, 2
apterous males, 1 winged male, 1 apterous female, 2 winged females.
Mexico: Chiapas: Hda. La Libertad, Sept. 1, 1987, H. D. Thomas,
1 apterous male.
Guerrero: Rio Agua, Kil. 487 S. Mex. City, 10-80-86, H. D.
Thomas, 7 apterous males, 6 apterous females, 7 winged males,
5 winged females; Garrapatas, Kil. 369 S. Mex. City, 10-30-36, 1
870 Tue Universiry Science BULLETIN
apterous male, 3 apterous females, 17 winged males, 22 winged
females,
Michoacén: El Sabino, Uruapan, 7-24-36, H. D. Thomas, 21 ap-
terous males, 31 apterous females, 5 winged males, 4 winged fe-
males,
Morelos: Acatlipa, el. 4000 ft., 1-1946, Coll. J. C. Shaw, 5 ap-
terous males, 1 apterous female; Cuautla, 10-12-36, H. D. Thomas,
30 apterous males, 42 apterous females; Cuntlan del Rio, IV-26-44,
J. G. Shaw, 3 apterous males, 3 apterous females.
Puebla: Tehuacan, July 18-25, 1937, H. D. Thomas, 54 apterous
males, 12 apterous females.
Tamaulipas: 5 mi. N. of Ciudad, Victoria, 11-5-36, H. D. Thomas,
1] apterous males, 4 winged males, 15 apterous females, 5 winged
females.
Rhagovelia ignota Drake and Harris
(Pl. VI, fig. 3)
1933. Rhagovelia ignota Drake and Harris, Proc. Biol. Soc, Washington, vol.
46, p. 51.
Size: Length Width
, 3.72 mm. apterous male 1.19 mm. apterous male
4.05 mm. apterous female 1.42 mm. apterous female
4.20 mm. winged male 1.52 mm. winged male
4.38 mm. winged female 1.58 mm. winged female
Color: General color brown-black, clothed with golden pubes-
cence. Anterior band of pronotum yellow, interrupted at middle, be-
coming pruinose behind eyes. Mesonotum with pruinose areas sur-
rounding each puncture. Shining black spots on dorsum of last one
to several abdominal segments. Margins of connexiva orange to
yellow. Venter blue-gray. Venter of last abdominal segment
yellow beneath. Base of antennae, margins of all acetabulae, all
coxae, all trochanters, and bases of anterior and posterior femora
yellow. Wings brown-black or pruinose brown on basal one half,
brown on apical one half,
Structural characteristics: Intermediate femur of female trans-
versely constricted at middle. Apterous male: anterior trochanter
unarmed, Anterior tibia dilate (L. 70, W. 15) and flattened on
inner surface. Pronotum rounded at apex (L. 70, W. 77); meta-
notum short on midline (L. 7, W. 78). Proportions of antennae:
Seg. I: II: II: IV:: 57: 32: 80: 82; of intermediate legs: Fem.:
Tib.: Tars. I: Tars, III:: 121: 83: 40: 54; of posterior legs: 104:
94: 13: 24, Abdomen tapers evenly to apex. Venter with slight
median carina on first two segments of abdomen. Minute, conical,
Srupy oF THE GEeNus RHAGOVELIA 871
black setae on jugum of head, proepisternum, scattered irregularly
on venter, and on posterior margin of last abdominal segment at
sides. Venter depressed in middle on posterior three segments; last
segment deeply and roundly excavate behind with prominent callus
on each side, crest of each callus being somewhat in front of middle
of segment. First genital segment with posterior portion beneath
greatly swollen and hairy. Posterior trochanter armed with several
knoblike teeth. Posterior femur greatly incrassate (L. 104, W. 36)
and armed with long, black-tipped, yellow spine at basal one third
followed by nine or ten smaller, gradually decreasing spines to
apex. Posterior femur also armed with row of very small, knoblike,
inconspicuous teeth on basal one third, and row of small subequal
spines anterior to primary armature. Posterior tibia straight; armed
with two rows of small teeth with those of basal one half stronger
and more closely-set; armed also with spur at apex. Apterous fe-
male: pronotum formed as in apterous male (L. 70, W. 83), meta-
notum as in male (L. 5, W. 83). Proportions of antennae: 57: 29:
26: 28; of intermediate legs: 115: 78: 35: 55; of posterior legs: 95:
93: 10: 24. Intermediate femora transversely constricted at middle.
Abdomen tapers at first, practically parallel for last two segments.
Connexiva vertical, not contiguous. Venter without median carina.
Minute, conical, black setae on base of jugum of head, proepister-
num, and scattered irregularly on venter, Posterior trochanter un-
armed, Posterior femur moderately incrassate (L. 95, W. 23);
gradually increasing in diameter for basal one half, increasing
abruptly in diameter at middle, then tapering to apex. Posterior
femur armed at middle with one long, yellow spine followed by
seven smaller, gradually decreasing spines to apex; also armed after
middle with anterior row of five or six small, subequal teeth, Pos-
terior tibia straight; armed on basal one half with closely-set teeth,
apical one half armed with two or three scattered small teeth and
inconspicuous spur at apex. Winged forms: Proportions and arma-
ture similar to apterous forms. Pronotum carinate on midline; pro-
longed into short spine at apex in male (L. 15), and a long, slender
spine in female (L. 45). Genital segments of male formed as in
apterous forms. Intermediate and posterior legs of female formed
as in apterous female, Pronotum coarsely and thickly punctate
posterior to anterior band. Wings barely cover apex of genital
segments.
Comparative notes: This species resembles R. spinigera Cham-
pion, R. ignota Drake and Harris can be separated from R. spini-
Tue Universtry ScreNcE BULLETIN
872
gera Champion by the first genital segment of the male which is
greatly swollen beneath in R. ignota, and the formation of the pos-
terior femur of the female.
Data on types: The holotype is a winged male from Guatemala,
and is in the personal collection of Dr. C. J. Drake. The allotype
is a winged female, and paratypes are two males and four females
taken with the allotype from Honduras. These specimens are in
the U. S. National Museum.
Data on distribution: Recorded from Guatemala and Honduras.
Specimens from the following localities have been examined (new
records for major political areas are indicated with an asterisk ) :
Guatemata: Gualan, 23-I-05 (J. R. de la Torre-Bueno Collec-
tion), 2 apterous males, 3 apterous females; Agua Caliente 9-II-’05
(J. R. de la Torre-Bueno Collection), 2 apterous males; Mazate-
nango, Feb. 3, 1905 (J. R. de la Torre-Bueno Collection ), 2 apterous
males, 1 apterous female.
Honpuras: Tegucigalpa, VII-18-18, F. J. Dyer Coll. No. 43870
(U.S. N. M. Collection), 3 apterous males; Dept Morazan, Esc. agr.
Pan., Zomorano, 2600 ft. (creek bank) Aug. 2, 1948, T. H. Hubbell
181, 3 apterous males, 3 winged males, 1 apterous female,
*Mexico: * Chiapas: Tuxla Gutierrez, Aug. 27, 1937, H. D.
Thomas, 17 winged males, 12 winged females; Guadalupe, 800
m.a.s.l., 1-15-38, Octavio Utrilla L., 1 apterous male, 6 winged males,
1 apterous female; Hda. La Libertad, Sept. 1, 1937, H. D, Thomas,
5 winged males, 1 winged female.
* Chihuahua: Carimechi, Rio Mayo, 1-7-35, H. S. Gentry, 3 apter-
ous males, 1 winged male, 7 apterous females, 1 winged female.
* Guerrero: Chilapa, Oct. 29, 1930, Prof. L. Schultze, 1 winged
female; Iguala, 10-7-36, H. D. Thomas, 25 winged males, 3 apterous
females, 23 winged females; La Sabana, 226 kil. S. of Mex. City,
10-20-36, H. D. Thomas, 34 winged males, 40 winged females; Gar-
rapatas, 369 kil. S. of Mex. City, 10-31-36, H. D, Thomas, 1 winged
male,
* Jalisco: Tecotlotlan, 9-17-38, H. D. Thomas, 1 winged male, 3
winged females.
* México: Zamora, Sept. 9, 1938, H. D. Thomas, 2 winged males,
5 apterous females, 3 winged females; E] Sabino, Uruapan, 7-24-36,
H, D. Thomas, 1 winged male.
* Morelos: Acatlipa, el. 4000 ft., 1-1946, J. G. Shaw, 1 apterous
male, 4 apterous females; Acatlipa, May 5, 44, J. G. Shaw, 1 apterous
Stupy oF THE GENUS RHAGOVELIA 873
male, 1 winged female; Acatlipa, 88 km. on Hgw. Mex. City to
Acapulco, May 6, 45, J. G. Shaw, 1 apterous male, 1 apterous fe-
male; Cuernavaca, 10-5-36, H. D. Thomas, 1 winged male; Cuautla,
10-12-36, H. D. Thomas, 1 winged male, 1 winged female; Cuntlan
del Rio, IV-26-44, J. G. Shaw, 1 apterous male, 1 winged male, 2
apterous females; Hda. Cocoyotia, IV-14-44, J. G. Shaw, 1 apterous
male, 3 winged males, 1 apterous female, 5 winged females.
* Nuevo Leén: Galeana, VII-3-39, H. Hoogstraal, 1 winged male.
* Sonora: San Bernardo, Rio Mayo, 10-14-34, H. S, Gentry, 1
winged male, 21 winged females; Arroyo de los Mescales, Rio Mayo,
9-16-35, H. S. Gentry, 14 winged males, 25 winged females.
* Tamaulipas: Victoria, 5 mi. N. of Ciudad, 11-5-86, H. D.
Thomas, 2 apterous males, 3 winged males, 3 winged females;
Victoria, VIII-1-41, Henry Thomas, 2 apterous males, 1 winged
male, 2 apterous females, 1 winged female.
Rhagovelia spinigera Champion
(Pl. VI, fig. 4)
1898. Rhagovelia spinigera Champion, Biol. Centr, Amer., Het., VOL, By Di leds
1901. Rhagovelia spinigera, Kirkaldy, Ento, vol. 34, p. 808 (mentions in key).
1931. Rhagovelia spinigera, Gould, Kansas Univ. Sci. Bull., vol. 20, p. 48 (re-
describes, records from Guatemala and Costa Rica).
1931. Rhagovelia spinigera, Drake and Harris, Pan Pacific Ent., vol. 8, p. 35
(record from Mexico as well as Costa Rica and Guatemala ).
1933. Rhagovelia spinigera, Gould, Ann. Ent. Soc. America, vol. 26, p. 469
(records from Mexico).
19385. Rhagovelia spinigera, Drake and Harris, Proc, Biol. Soc. Washington,
vol. 48, p. 36 (describe apterous form).
Size: Length Width
4.05 mm. apterous male 1.33 mm. apterous male
4.45 mm. apterous female 1.56 mm. apterous female
4.50 mm. winged male 1.56 mm. winged male
5.05 mm. winged female 1.78 mm. winged female
Color: General color brown-black, clothed with brown pubes-
cence, Pronotum with silver-gray to pruinose orange band on an-
terior margin interrupted at middle by median, dark brown, longi-
tudinal line which is more or less distinct for length of pronotum.
Punctures of pronotum surrounded by narrow gray margin. Mar-
gins of connexiva orange. Venter blue-gray. Venter of last ab-
dominal segment and genital segments yellow to light brown. Base
of antennae, margins of all acetabulae, all coxae and trochanters, and
basal one half of anterior femora yellow. Wings brown, lighter at
apex; veins black.
Structural characteristics: Intermediate femur of female trans-
versely constricted at middle. Apterous male: anterior trochanter
874
Tue Universitry ScrENCE BULLETIN
unarmed; anterior tibia only slightly dilate, flattened within for
anterior one half. Pronotum coarsely and thickly punctate, rounded
at apex (L. 72, W. 84); mesonotum hidden; metanotum short on
midline (L. 7, W. 85). Proportions of antennae: Seg. I: Il: III:
IV:: 56: 88: 80: 32; of intermediate legs: Fem.: Tib.: Tars. II:
Tars. III:: 128: 85: 88: 55; of posterior legs: 102: 95: 12: 26,
Slight ventral carina between posterior coxae. Minute, black,
conical setae on base of jugum of head, proepisternum, and scat-
tered irregularly on venter. Venter of last two segments of abdo-
men flattened beneath with slight, hairy calli on each side of me-
dian area. Intermediate trochanter armed with from four to ten
small teeth. Intermediate femur armed on ventral surface with
single row of approximately twelve small teeth on basal one third.
Posterior trochanter armed with approximately ten very small,
faint teeth. Posterior femur moderately incrassate (L. 102, W. 28)
and armed on basal one third with row of small teeth, armed just
before middle with one long spine followed by nine or ten smaller,
gradually decreasing spines to apex. Posterior tibia straight; armed
within with row of black teeth, with those of basal half stronger
than those on apical half; armed at apex with inconspicuous spur.
Apterous female: anterior tibia formed as in male. Pronotum punc-
tate as in male (L. 88, W. 100); mesonotum hidden, metanotum
flattened in middle, turgid to each side (L. 6, W. 104). Propor-
tions of antennae: 66: 86: 34: 33; of intermediate legs: 180: 95:
40: 60; of posterior legs: 105: 115: 15: 27. Intermediate femur
unarmed; with transverse constriction at middle; venter without
median carina. Minute, conical, black setae as in male. Posterior
trochanter unarmed. Posterior femur not as incrassate as that of
male; subcylindrical for basal two fifths (W. 15), expanded abruptly
at basal two fifths (W. 24); armed just beyond middle with one
long, bent spine followed by seven or eight shorter, decreasing
spines to apex. Posterior tibia straight; armed on basal one half
with subequal, black teeth; seemingly unarmed at apex. Winged
forms: proportions and armature similar to apterous forms. Pro-
notum of male produced into short process (L. 10); of female into
long spiniform process projecting backwards at angle of forty-five
degrees (L. 36). Trochanter and femur of intermediate leg of
male armed as in apterous male. Femora of intermediate and pos-
terior legs of female formed as in apterous female. Wings cover-
ing apex of genital segments.
Comparative notes: This species resembles R. ignota Drake and
Harris. R. spinigera Champion can be separated from R. ignota
Srupy OF THE GENUS RHAGOVELIA 875
Drake and Harris by the armed intermediate trochanter and femur
of the male, and by the formation of the posterior femur of the fe-
male, which is abruptly expanded before the insertion of the long
spine, rather than being expanded at the point of insertion of the
long spine as is the posterior femur of the female of R. ignota Drake
and Harris. The males also lack the swollen genital segment which
characterizes R. ignota Drake and Harris. Champion described this
species from only the winged female or he would doubtlessly have
noticed the armed intermediate femur of the male. R. spinigera
Champion is the only species of Rhagovelia known to have such an
armature.
Data on types: Champion's type material is in the British Mu-
seum. He gives the type locality as: “Guatemala, San Gerénimo in
Vera Paz.” Specimens in the Francis Huntington Snow Ento-
mological Collections have been compared with the type material
by Dr. H. B. Hungerford in 1928 and determined by him to be
conspecific.
Data on distribution: Recorded from Guatemala, Costa Rica, and
Mexico. Specimens from the following localities have been ex-
amined (new records for distribution in major political areas are
indicated with an asterisk):
Costa Rica: San José, Alt. 1160 m.a.s.l., Acequias, agua corriente,
VIII, 1905, P. Biolley (J. R. de la Torre-Bueno Collection), 22
winged males, 2 apterous females, 12 winged females; R. Maria
Aguilar, alt. 1160 m.a.s.l., Jan. 1905, P. Biolley (J. R. de la Torre-
Bueno Collection), 5 apterous males, 8 winged males, 8 apterous
females, 9 winged females; San José, Purchased 6-31, H. Schmidt,
- 10 apterous males, 6 apterous females, 3 winged females; San José,
Heinrich Schmidt, purchased 1932, 2 apterous males, 1 winged
male, 3 apterous females, 4 winged females; San José, 6 & 7, 1931,
Heinrich Schmidt, bought from Schmidt, 11 apterous males, 40
winged males, 18 apterous females, 25 winged females; Rio Torres,
2-10-32, Heinrich Schmidt, bought from collector, 3 apterous males,
2 winged males, 4 apterous females, 9 winged females; Rio Tiribi,
1100 m.a.s.L., IV-06, P. Biolley (J. R. de la Torre-Bueno Collection ),
1 winged male, 3 winged females; Rio Virilla, 12-26-31, Heinrich
Schmidt, purchased from collector, 38 apterous males, 63 winged
males, 25 apterous females.
GuatemMata: Mazatenango, 3 Feb., 05 (J. R. de la Torre-Bueno
Collection), 6 apterous males, 3 apterous females; Agua Caliente,
28-1-05 (J. R. de la Torre-Bueno Collection), 6 apterous males, 5
876 Tue Universtry SCIENCE BULLETIN
winged males, 6 apterous females, 3 winged females; Agua Caliente,
9-II-05 (J. R. de la Torre-Bueno Collection), 18 apterous males, 1
winged male, 3 apterous females, 8 winged females.
* Honpuras: Copan, 2-7-37, Chester Roys, 2 winged males, 1
winged female; Tegucigalpa, VIJJ-18-18, F’. J. Dyer Coll. No. 84870
(U. S. National Museum Collection), 1 apterous male; Dept. Mora-
zn, Esc. Agr. Pan., Zomorano, 2600, creek bank, Aug. 2, 1948; T. H.
Hubbell, 181, 3 apterous males, 2 apterous females.
Mexico: Chiapas: Hda La Libertad, Sept. 1, 1937, H. D. Thomas,
15 winged males, 4 winged females; Guadalupe, 1-15-38, 800
m.a.s.l, Octavio Utrilla L., 7 apterous males, 17 winged males, 14
apterous females, 11 winged females, 10 nymphs; Tuxtla Gutierrez,
Aug. 27, 1937; H. D. Thomas, 6 winged males, 6 winged females.
Guerrero; Garrapatas, kil. 360 S. of Mex. City, 10-20-36, H. D.
Thomas, 7 apterous males, 20 winged males, 18 apterous females,
11 winged females; La Sabana, kil. 226 S. of Mex. City, 10-20-36,
H. D. Thomas, 6 winged males, 8 winged females.
Jalisco; Tecolotlan, 9-17-38, H. D. Thomas, 2 winged males.
México: Tejupilco, Dist. of Temascaltepec, alt. 1840 m.a.s.1., June-
July, 1933, H. E. Hinton, 21 winged males, 1 apterous female, 8
winged females. ;
Michoacén: Zamora, Sept. 9, 1938, H. D. Thomas, 1 winged
male, 1 winged female; El Sabino, Uruapan, 7-24-36, H. D. Thomas,
5 apterous males, 5 winged males, 3 apterous females, 2 winged
females.
Morelos: Hda Cocoyotla, IV-14-44, J. G. Shaw, 2 apterous males,
2 winged males, 2 apterous females.
UNKNOWN SPECIES
The following two species have not been included in any group
or key as the descriptions are not sufficiently detailed nor specimens
available,
Rhagovelia bakeri Bergroth
1914. Rhagovelia bakeri Bergroth, Psyche, vol, 21, p. 74.
1931. Rhagovelia bakeri, Gould, Kansas Univ. Sci. wBull, vol, 20, p. 17 (re-
describes ).
This species is not represented in the material in the Francis
Huntington Snow Entomological Collections, University of Kansas.
The original description is quoted below:
“Above black, beneath grayish black, legs greenish black, some-
what aenescent, pronotum with an apical whitish fascia interrupted
Stupy OF THE GENUS RHAGOVELIA 877
in the middle, abdomen at the sides immediately within the margin
narrowly and obscurely streaked with yellowish brown, basal part
of the first antennal joint, anterior acetabula, all coxae and trochan-
ters, and basal part of fore femora yellow, hind femora at the base
above and beneath also tinged with yellow. Head with an im-
pressed median line in front, rostrum passing the middle of the
mesosternum, antennae with the first two joints pubescent, with
scattered longer hairs, outer margin of the last three joints straight,
inner margin slightly convex, first joint one half longer than second,
third a little shorter than second, fourth hardly shorter than third,
pointed at tip. Mesosternum with the ridges between the anterior
and the middle coxae well marked, pubescent, curved inwardly,
strongly divergent posteriorly. Abdomen gradually tapering to the
tip, not carinated beneath. Legs pubescent, with scattered longer
hairs, middle femora thickened toward the base, middle tarsi a little
longer than the tibiae, the second joint a little shorter than the
third, posterior tibiae with a short straight spur at apex.
“Apterous male: Pronotum a little broader than long, rounded
behind. Last dorsal segment of abdomen truncate at apex, last
ventral segment arcuately emarginate at apex. First genital seg-
ment ferruginous beneath at the base. Posterior, femora very
strongly incrassate, not reaching the tip of the apical genital seg-
ment, spined beneath from the apex to a little beyond the middle,
the first spine (near the middle) the longest, the following gradu-
ally diminishing in length toward the apex. Posterior tibia straight,
finely denticulate beneath down their whole length, without large
teeth, the teeth of the middle third slightly longer, the apical third
slightly narrower than the rest. Length, 4 mm.
“Locality: Nicaragua (Granada).
“Allied to Rh. femoralis Champ., but the antennae are differently
constructed; the venter is not ridged in the middle; the posterior
tibiae have no large teeth, etc.”
Comparative notes: From the description quoted above it would
appear that R. bakeri Bergroth belongs in the crassipes group. Speci-
mens will have to be examined before a more complete relationship
can be established. The incomplete description will not allow this
species to be placed in a key. Bergroth’s type material is apparently
lost, making it impossible to determine the relationships of this
species,
Tue Universiry Science BULLETIN
Rhagovelia trailii: (White )
1879. Neovelia trailii White, Jour. Linn. Soc. London, Zool., vol. 14, p. 487.
1901. Rhagovelia trailii, Kirkaldy, Ent., vol. 34, p, 308 (places in genus
Rhagovelia).
1931. Rhagovelia trailiit, Gould, Kansas Univ. Sci. Bull., vol, 20, p. 45.
This species was originaly described from two specimens, both
winged females. The description is mostly based on color. The
relationships of this species cannot be determined with sufficient
accuracy to permit its being placed in any of the groups of species
or included in the keys. The original description is quoted below.
“Nigro-Brunea, pubescentia confera concolori vestita, capite, an-
tennis, pedibus, pronoto postico abdominisque lateribus parce nigro-
setulosis; pronoto antice, prostethio, connexivo, ventre ad medium,
antennarum atriculo primo ad basin, coxis, trochanteribus, femorum
anticorum macula et vitta subtus, femorum posticorum basi et spinis
ad basin flavido-fulvis; pronoti carina centrali sub-elevata plus
minus, praecique antrosum, rufo-fulva. Male long. 4, lat. 1% millim.
“Hab. Brasiliam borealem. (Mandos, August, 1874, ‘at light,’
JW. Eb, Urails) 7
The type and the one cotype.are both winged females and are in
the British Museum.
Data on distribution: Known only from the type series.
ADDENDA
The following pages contain description of seven new species of
Rhagovelia, These were described by Dr. C. J. Drake after the
main body of this paper was completed. Specimens of R. citata
Drake and R. itatiaiana Drake were available to the author and
redescribed by him; other descriptions are the original descriptions
by Dr. Drake. Dr. Drake’s comparative measurements are to the
same scale, at the same magnification, and such that 80 units equal
one millimeter. The types are in the collection of Dr. Drake.
Due to the fact that specimens of most of these species were
unavailable to the author no attempt has been made to include these
species in either groups or keys. The species included in the ad-
denda are:
1. R. acapulcana Drake 5. R. rioana Drake
2. R. citata Drake 6. R. triangula Drake
3. R. itatiaiana Drake 7. R. torreyana Drake and Hussey
4, R. novana Drake 8. R. zeteki Drake
Strupy oF THE GENUS RHAGOVELIA §79
Rhagovelia acapulcana Drake
1953. Rhagovelia acapulcana Drake, Proc. Biol. Soc. Washington, vol. 66, p.
145.
“Apterous female: Large, fusiform, black with a short interrupted
band near the front margin of pronotum orange-brown; connexiva
broadly margined with yellowish brown, the large terminal spines
black; basal part of antennae, most of all acetabulae, middle and
hind coxae, all trochanters and basal part of fore femora pale testa-
ceous. Body beneath, head in front and fore part of pronotum
bluish. Length, 4.10 mm.; width, 1.36 mm.
“Head with usual impressed lines and marks. Antennae with a
few scattered stiff hairs on first two segments, measurements—seg-
ment I, 76; II, 45; III, 80; IV, 35. Pronotum large, covering most
of mesonotum, punctate, broadly rounded behind, with median
carina, beset with long erect dark hairs along entire hind margin,
the width and length subequal (100; 98). Abdomen sharply nar-
rowed on basal part, then slowly tapering for the last three seg-
ments; connexiva moderately wide, strongly reflexed with the outer
margins not touching and projecting above the tergites, each con-
nexivum termining behind in a very long, stout, black, hairy spine,
which is directed obliquely inwardly so that the tips of the two
spines contact each other. Apex of genital segment broad, sharply
truncate. Anterior trochanters unarmed; tibiae slightly dilate and
slightly excavate on apical half. Measurement of middle femora,
164; tibiae, 100; tarsi IT, 16 and III, 28. Male and winged form
unknown.
“Holotype (female), Acapulco, Mex., July 23, 1950 and 1 para-
type, same locality, Aug. 3, 1951, both by C. J. Drake.
“The extremely large black spine at the posterior end of each
connexivum separates this insect at once from other American
species of the genus.”
Rhagovelia citata Drake
(PL, VIII, fig. 1)
1953. Rhagovelia citata Drake, Proc. Biol. Soc, Washington, vol, 66, p. 146.
Size: Length Width
5.80 mm. apterous male 1.50 mm. apterous male
5.80 mm, apterous female 1.70 mm, apterous female
5.50 mm. winged female 2.00 mm. winged female
Color: General color red-brown, clothed with a golden pubes-
cence, Anterior one sixth of pronotum with band of yellow brown;
pronotum also with median longitudinal line and thin posterior
880 Tue Universitry SCIENCE BULLETIN
margin slightly lighter. Disc of pronotum lighter in color than
dorsum of abdomen. Connexiva broadly margined with yellow-
brown. Venter lighter in color than dorsum. All legs yellow-
brown. Wings brown, veins prominent and black.
Structural characteristics: Dorsum of abdomen of apterous fe-
male narrowed after first three segments, connexiva vertical; an-
terior tibia moderately dilate and excavate within in male. Apter-
ous male: anterior trochanter unarmed; anterior tibia moderately
dilate (W. 17) and slightly excavate for apical half. Pronotum
wider than long (L. 76, W. 88); metanotum much wider than long
(L. 2, W. 90). Proportions of antennae: Seg. I: II: III: IV:: 80:
48: 46: 43; of intermediate legs: Fem.: Tib.: Tars. I: Tars, III::
170; 129: 61: 70; of posterior legs: 136: 138: 18: 83. Abdomen tapers
evenly to apex. Ventral carina most strongly produced between
posterior coxae where venter is depressed on each side; carina
only slightly produced between posterior trochanters. Minute black
conical setae sparsely scattered on base of jugum of head and one or
two along posterior margin of last abdominal segment at sides. Ven-
ter of last abdominal segment depressed for posterior two thirds, bor-
dered with a slight callus, raised longitudinal carina faintly indicated
on posterior one half. Posterior trochanter armed with many small,
light colored teeth. Posterior femur moderately incrassate (L. 136,
W. 34) and armed with a row of closely set teeth on basal two fifths,
armed at basal two fifths with a long, black-tipped spine followed by
a row of twelve to thirteen decreasing spines to apex; also with an an-
terior row of subequal teeth from the middle continuing to the apex.
Posterior tibia straight; armed beneath with a row of closely set teeth
and a partial second row with a few scattered teeth; posterior tibia
also armed with a stout slightly bent spur at apex. Apterous female:
anterior tibia flattened within for apical half, slightly dilate. Pro-
notum wider than long (L. 98, W. 114); metanotum as in male (L. 5,
W. 110). Proportions of antennae: 94: 53: 49: 45; of intermediate
legs: 179: 136: 67: 74; of posterior legs: 142: 154: 20: 82, Con-
nexiva meeting after first three segments, spreading slightly for
last segment and terminating in a thick clump of long dark hairs.
Dorsum of abdomen narrow after first three segments, No ventral
carina. Minute, black, conical setae as in male. Posterior trochanter
unarmed. Posterior femur only slightly swollen (L. 142, W. 23);
armed at apical third with one medium sized spine followed by a
rapidly decreasing series of four or five smaller spines to apex.
Posterior tibia unarmed except for spur at apex.
Stupy oF THE GENUS RHAGOVELIA 881
Winged female: proportions armature, and arrangement of black,
minute, conical setae much as in apterous female. Pronotum pro-
duced into a long, bent process (L. 62) which tapers to point at its
apex. Winged male: unknown.
Comparative notes: This species is in the collaris group and ap-
pears to be a close relative of R. armata (Burmeister) but can be
separated at once from that form by the pattern of the minute,
black, conical, setae which are only thinly scattered on the base
of jugum of head and occasionally along the posterior margin of
the last abdominal segment at the sides.
Data on types: Holotype, apterous male; allotype, apterous fe-
male; allomorphotype, winged female. Described from a series
labeled: “Canal Zone, Panama, Feb. 12, 1939, C. J. Drake.” Para-
types, 48 specimens taken with types and several collections from
Venezuela. The type material is in the personal collection of Dr.
C. J. Drake. Three apterous males, 1 apterous female, and one
winged female paratypes are in the Francis Huntington Snow Ento-
mological Collections of the University of Kansas.
Data on distribution: Known only from the type series Panama
and Venezuela.
Rhagovelia itatiaiana Drake
1953. Rhagovelia itatiaiana Drake, Proc. Biol. Soc. Washington, vol. 66, p. 147.
Size: Length Width
8.15 mm. apterous male 1.14 mm. apterous male
8.40 mm. apterous female 1.19 mm. apterous female
Color: General color brown-black with prominent yellow mark-
ings, clothed with a golden pubescence. Head with yellow stripe
in front, extending onto vertex in some specimens. Pronotum with
anterior one third yellow; mesonotum yellow except for faintly
darker margins. Metanotum with yellow areas at humeral angles
on some specimens. Connexiva, on both sides, broadly yellow
except for thin brown outer margin. Yellow beneath except for
venter of abdomen which is blue-gray; last two or three segments
of venter of abdomen with yellow spots at sides. All coxae, all
trochanters, basal one half of anterior femora and basal one half
of posterior femora of female and whole of posterior femora of
male beneath yellow.
Structural characteristics: Pronotum in apterous forms abbrevi-
ated, rounded behind, and exposing much of mesonotum. Apter-
ous male: anterior trochanter unarmed. Anterior tibia not dilate,
29—3378
Tue Universiry ScteENCE BULLETIN
882
flattened within. Pronotum short (L. 31, W. 61); mesonotum
truncate at apex (L. 20, W. 65); metanotum short (L. 3, W. 70).
Antennal segments and intermediate legs broken off on specimens
at hand.* Proportions of posterior legs: Fem.: Tib.: Tars. II: Tars.
III:: 92: 68: 4: 22. Abdomen tapers rather evenly to apex, angle
of taper increasing slightly for last three segments. Venter with
slight median carina, becoming more pronounced on the anterior
one half of the last abdominal segment. Posterior trochanter un-
armed. Posterior femur strongly incrassate (L. 92, W. 29). Armed
on basal one third with a single row of small dark teeth; armed
after basal one third with a closely set row of thirteen to fifteen
gradually decreasing spines to apex. Also armed with a few short,
dark teeth in an anterior row on apical one fifth of the femur.
Posterior tibia slightly arcuate; armed within with a row of closely
set teeth; armed with a stout spur at apex. Apterous female: pro-
notum rounded behind (L. 28, W. 62); mesonotum truncate at apex
(L. 22, W. 68); metanotum short on midline (L. 5, W. 77). An-
tennae broken.f Proportions of intermediate legs: 100: 79: 32: 45;
of posterior legs: 79: 79: 19: 7. Abdomen tapers evenly to apex.
Venter without median carina. Posterior trochanter unarmed. Pos-
terior femur moderately incrassate (L. 79, W. 18) and armed after
the middle with a dark spine followed by seven or eight regularly
spaced, decreasing spines to apex. Posterior tibia straight, un-
armed. Winged forms: unknown.
Comparative notes: This species is in the abrupta group and is
very similar in appearance to R. mira Drake and Harris. R. itatiaiana
Drake is much smaller than that form and the strongly produced
ventral carina on the anterior half of the last abdominal segment
of the male will set it apart at once.
Data on types: Holotype, apterous male, allotype, apterous fe-
male and 22 apterous paratypes in the personal collection of Dr.
C. J. Drake. One male and one female paratype are in the Francis
Huntington Snow Entomological Collections of the University of
Kansas. The type series is labeled “Fas Pendo, Italiaia, Guapimir
M. and Gavea, Rio Janeiro, Brazil.”
Data on distribution: Known only from the type series.
* Dr. Drake gives these measurements in his original description as follows: for an-
tennae: “I, 65; II, 40; III, 45; IV, 38”; for intermediate leg: “femora, 120; tibiae, 95; tarsi
UJ, 45 and Ill, 56.” Dr. Drake used a comparative scale in which 80 units equalled 1 mm.
for these measurements,
+ Dr. Drake gives these measurements in his original description as follows: “I, 60; II,
85; III, 40; IV, 35.”
Strupy oF THE GENUS RHAGOVELIA 883
Rhagovelia novana Drake
1953, Rhagovelia novana Drake, Proc. Biol. Soc. Washington, vol. 66, p. 150.
“Small, blackish with a short, median, interrupted, yellowish
stripe on pronotum; pubescence dark brown; body beneath dark
bluish, the pronotum also a little bluish. Antennae brownish black
with basal part of first segment testaceous. Legs brownish black,
the anterior coxae testaceous. Length (male) 2.30 mm., (female)
2.50 mm.; width (male), 1.10 mm.; (female), 1.25 mm. Winged
form unknown.
“Male: Head with usual impressed lines and scattered long hairs.
Antennae shortly pilose, with the usual bristly hairs on first two
segments: measurements—I, 65; II, 45; III, 88; IV, 35 (same in
both sexes). Pronotum very short, not produced posteriorly over
mesonotum, much wider than long (68: 18). Mesonotum large,
convex above, covering most of metanotum, wider than long (82:
62). Abdomen strongly tapering posteriorly; connexiva wide, grad-
ually tapering to the end of the last tergite, reflexed obliquely up-
ward. Legs long, slender, all femora unarmed in both sexes; an-
terior trochanters unarmed; hind tibiae without distinct spur at
apex, Anterior tibiae slightly dilate apically; measurements of
middle legs—femora, 150; tibiae, 100; tarsal segment IT, 32 and HI,
28. Hind legs—femora 110; tibiae, 116; tarsal segment IT, 4 and
IL, 10. First genital setment (sic) rounded above, feebly roundly
excavated behind.
“Female: Broader than male; pronotum very short, not produced
behind over mesonotum. Antennal measurements as in male.
Mesonotum large, wider than long (92: 80). Abdomen strongly
narrowed posteriorly, unusually strongly constricted at the base
of last segment, there narrow and nearly flat laterally, then abruptly
widened behind, the apical part enlarged and about three times as
wide as at constriction; connexiva strongly reflexed, not quite rest-
ing on tergites on basal half, from the constricted part posteriorly
resting on tergites with their outer margins in contact with each
other; abdominal tergite at base of constriction with a median semi-
ovate plate raised vertically and projecting obliquely posteriorly
between the outer edges of the reflexed connexiva, the outer edge
of the plate clothed with rather long hairs; genital segments also
with long brownish hairs. Legs unarmed, about the same thick-
ness as in male. Measurements—middle femora, 142; tibiae, 108;
tarsi II, 64 and III, 56. Hind femora, 100; tibiae, 144; tarsi I, 11
and III, 23.
Tue Universiry Scrence BULLETIN
884.
“The extremely constricted abdomen a little before apex with
enlarged short apical part and the singular vertical plate-like struc-
ture before apex of abdomen separate this peculiar insect at once
from its congeners. The hind femora of the male is unarmed as in
R. longipes Gould, but the entire insect is much smaller.”
Rhagovelia rioana Drake
1958. Rhagovelia rioana Drake, Proc. Biol. Soc. Washington, vol. 66, p. 152.
“Apterous form: Small, black with the short pronotal band inter-
rupted at the middle; pubescence brownish; entire body beneath
bluish gray, sometimes also the fore part of pronotum. Antennae
black with proximal part of first segment pale testaceous. Legs en-
tirely black, sometimes the fore coxae partly flavous. Length
(male), 2.75 mm. and (female), 3.00 mm.; width (male), 1.30 mm.
and (female), 1.42 mm.
“Male: Head with the usual impressed median line and scattered
long hairs. Antennae shortly pilose, with a few scattered bristly
hairs on first two segments; measurements—I, 92; II, 48; III, 45;
IV, 50. Pronotum very short, not produced over mesonotum, much
wider than long (85: 25). Mesonotum large, covering most of
metanotum, roundly narrowed on the sides posteriorly, blunt and
rounded at the apex, wider than long (100: 70). Metanotum largely
concealed; with hind margin and sides visible. Abdomen strongly
tapering posteriorly; connexiva slightly reflexed, with outer margin
feebly rounded. First genital segment beneath almost cylindrical.
Legs long and slender. Anterior trochanters unarmed; tibia not
dilate. Length of middle femora, 180; tibiae, 120; tarsi IJ, 68; III,
64, Hind femora not incrassate, unarmed, the tibiae also unarmed
and without apical spur: measurements—femora, 120; tibiae, 130;
tarsi II, 10 and III, 35.
“Female: Broader than male, with appendages about the same
size. Pronotum short and sutured-off as in male. Mesonotum
large, covering most of metanotum, broader than long (104: 76),
with apex broad and subtruncate. Abdomen strongly tapering
posteriorly, slightly constricted a little in front of genital segment,
connexiva completely reflexed behind, with outer margins barely
in contact on penultimate tergite with outer margins raised and
flaring anteriorly, obliquely narrowed on last tergite leaving the
median part of tergite uncovered, last tergite raised medially api-
cally so as to form a narrow plate with the rounded apex projecting
Srupy OF THE GENUS RHAGOVELIA 885
a little over narrow basal part of genital segment. Last ventrite
nearly as long as the two preceding segments.
“Alate form: Pronotum moderately convex, with a fairly distinct
median carina, nearly triangular behind with apex rounded, very
little wider than long (124: 110). Hemelytra dark fuscous, wings
fuscous, both longer than abdomen; tips of both wings broken-off
in front of the genital segments in both male and female.
“Holotype (male) and allotype (female), both aperous (sic),
Guapimir M., Rio de Janeiro, Braz. Paratypes, 14 specimens, same
data as type.
“The small size, measurements of appendages, unarmed and not
swollen hind femora and the connexiva and last tergite in female
distinguish R. rioana from R. longipes and allied forms and allied
forms with slender and unarmed legs.”
Rhagovelia torreyana Drake and Hussey
(Pl. VIII, fig. 4)
1955. Rhagovelia torreyana Drake and Hussey.
Size: Length Width
3.75 mm. apterous male 1.26 mm. apterous male
3.80 mm. apterous female 1,28 mm. apterous female
Color: General color black, clothed with golden pubescence.
Anterior one fifth of pronotum with orange band interrupted at
middle. Dorsum of first two abdominal segments pruinose at sides
in female. Margins of connexiva orange. Venter gray-black. Ven-
ter of last abdominal segment orange beneath. Base of antennae,
all acetabulae, all coxae, all trochanters, and femora of all legs
beneath yellow to orange.
Structural characteristics: Pronotum of apterous form covers
mesonotum. Dorsum of abdomen of apterous female slightly con-
stricted after first three segments. Posterior tibia without sickle-
shaped spur at apex. Apterous male: anterior trochanter unarmed.
Anterior tibia only slightly dilate and flattened within. Pronotum
broadly rounded behind, slightly depressed along posterior margin
(L. 62, W. 70) and covering mesonotum. Metanotum short on
midline (L. 6, W. 73). Proportions of antennae: Seg. Te Bh is
IV:: 59: 32: 40: 39; of intermediate legs: Fem.: Tib.: Tars. IT:
Tars. III:: 113: 100: 40: 53; of posterior legs: 104: 95: 9: 22, Ab-
domen tapers evenly to apex. Connexiva semi-vertical. Venter
with no trace of median carina. No minute, conical, black setae
on proepisternum. Posterior trochanter armed with from four to
six short teeth in a single row. Posterior femur moderately in-
886 Tue University SCLENCE BULLETIN
crassate (L. 104, W. 87); armed on basal one third with a median
row of short, knoblike teeth, armed before the middle with a row of
eight to ten equidistant, gradually decreasing spines to apex. Also
armed with an anterior row of short subequal spines beginning at
basal one third and continuing to apex. Posterior tibia almost
straight and armed within with a single row of short, black teeth
and a slender spur at apex. Apterous female: anterior tibia as in
male. Pronotum formed as in male (L. 55, W. 77). Metanotum
as in male (L. 6, W. 80). Proportions of antennae: 59: 28: 36: 36;
of intermediate legs: 122: 93: 36: 55; of posterior legs: 92: 100: 9:
25. Dorsum of abdomen tapering inward for first four segments
then subparallel to apex. Connexiva almost vertical. Posterior
trochanter unarmed. Posterior femur only slightly incrassate (L.
92, W. 22); armed after the middle with one long black tipped
spine followed by four to five much smaller, rapidly decreasing
spines to apex. Posterior tibia straight and armed with a few,
widely scattered black teeth or seemingly unarmed except for a
slender spur at apex. Winged forms: unknown.
Comparative notes: This species is in the crassipes group and
closely related to R. amazonensis Gould. R. torreyana Drake and
Hussey can be separated from R. amazonensis Gould by the arma-
ture of the posterior femur in both males and females.
Data on types: The type series was collected in Liberty Co.,
Florida, Torreya St. Park, May 22, 1954, by R. F. Hussey. The
holotype and allotype are in the personal collection of Dr. C. J.
Drake, numerous paratypes are in the collection of R. F. Hussey.
One male and one female paratype are also in the Francis Hunting-
ton Snow Entomological Collections, University of Kansas.
Data on distribution: Known only from the type locality.
Rhagovelia triangula Drake
1958. Rhagovelia triangula Drake, Proc. Biol. Soc. Washington, vol. 66, p. 148.
“Apterous male: Small, fusiform, with a short, transverse inter-
rupted orange band near front pronotol margin; pubescence brown-
ish. Base of first antennal segment, all coxae, all trochanters (save
sometimes median), and basal third of anterior femora testaceous.
Rustrum (sic) dark ferrugineous. Body beneath, front of head and
fore part of pronotum bluish. Length, 2.85 mm.; width, 1.25 mm.
“Pronotum produced posteriorly, covering nearly one-half of
mesonotum, broadly rounded behind, wider than long (78: 44).
Mesonotum not as long as pronotum, wider than long. Antennal
Srupy oF THE GENUS RHAGOVELIA 887
measurements—I, 58; II, 32; III, 40; IV, 40. Anterior trochanter
unarmed; tibiae slightly dilate apically. Measurements of middle
femora, 120; tibiae, 95; tarsi II, 52, and III, 56. Hind trochanters
unarmed; femora moderately incrassate, armed near the basal third
with a long dark spine, thence to apex with seven or eight smaller
and rapidly decreasing spines; tibiae nearly straight; armed beneath
with numerous, closely-set, short, stout teeth, and at the apex with
a sharp spur; measurements—femora, 84; tibiae, 90; tarsi II, 12 and
Til, 25.
“Winged female: Head and antennae as in male. Pronotum
moderately convex, extremely long, with hind part longly produced,
nearly flat, coarsely punctate, rounded at apex, distinctly longer
than wide (160; 120), Hemelytra dark brown, much longer than
abdomen, the veins fairly distinct. Venter with the hairy median
carina not present on last segment; last ventrite longer than the
preceding two segments, strongly narrowed ventrally and laterally
to hind margin. Measurements of middle femora, 100; tibiae, 92;
tarsi II, 45, and III, 55. Hind femora a little incrassate, not as large
as in male, armed at the apical third with a rather short spine, then
followed with three or four shorter ones. Wingless female unknown.
Length, 4.00 mm.; width, 1.25 mm.
“Macropterous male: Similar to female in general aspect. Pro-
notum much shorter and equal in width and length (110: 110), the
posterior triangular part not unusually extended behind.
“Holotype (apterous male), and allotype (alate female), Guapi-
mir M. Rio de Janeiro, Braz. Paratypes, 4 specimens, taken with
the type.
“Separated from its congeners (pronotum concealing about one-
half of mesonotum in apterous form) by the measurements of
appendages, male parameres and the unusually longly produced
hind part of the pronotum in the winged female. The pronotum in
the alate female does not bear a horn or other modifications, just
the regular hind triangular part longly extended.”
Rhagovelia zeteki Drake
1958. Rhagovelia zeteki Drake, Proc. Biol. Soc, Washington, vol, 66, p. 145.
“Apterous form: Small, fusiform, dark brown with a solid orange-
yellow band near anterior margin of pronotum, clothed with short
golden pubescence and fine erect brownish hairs (hairs a little
more numerous in male). Basal part of antennae, fore and hind
coxae and the basal part of fore femora (nearly half beneath) pale
888 Tue Universiry Science BULLETIN
testaceous. Legs and antennae black-fuscous. Length, 310 mm.,
width, 1.12 mm.
“Male: Antennae with the first three segments beset in front with
numerous long brown hairs; measurements—I, 55; II, 35; III, 31; |
IV, 31. Legs with numerous long dark brown hairs; anterior tro-
chanters unarmed, the tibiae a little dilate and flattened apically.
Measurement of middle femora, 180; tibiae, 73; tarsi II, 40 and III,
58. Hind femora considerably swollen, armed near the basal third |
with a large bent black-tipped spine, thence to apex with a double
row (8-10 spines in row) of smaller-decreasing spines; tibiae a little
bent, with a spur at apex, armed beneath with a double row of
numerous short blunt teeth; measurements—femora, 102; tibiae,
84; tarsi II, 9 and III, 21. First genital segment beneath a little
impressed basally, especially on the sides. Pronotum large, wider |
than long (85: 72), with median carina, broadly rounded behind,
covering most of metanatum (sic).
“Female: Antennal hairs and measurements almost identical with
male. Pronotum large, covering most of mesonotum, with feeble
median carina, wider than long (92: 70). Abdomen more sharply
narrowed on basal part, narrow and nearly parallel on last three
segments; connexiva moderately wide, reflexed almost vertically,
with upper edge subparallel on last three segments. Anterior tro-
chanters unarmed. Measurements of middle femora, 120; tibiae
102; tarsi II, 45 and III, 56. Hind femora incrassate but not as thick
as in male, with similar armature; tibiae as in male; tarsi IJ, 6 and
IIT, 20.
“Macropters form: Pronotum with distinct median carina;
humeral angles a little elevated; hind triangular part with apex
rounded; very little wider than long (110: 100). Hemelytra dark
fuscous with veins darker, longer than abdomen. Male not quite
as broad as female. Length, 3.23 mm.; width, 1.38 mm.
“Holotype (male) and allotype (female), both apteous, (sic)
taken in a small stream, Barro Colorado, Canal Zone, Panama, Feb. |
6-8, 1939, C. J. Drake. Paratypes: 1 apterous and 1 alate male, 2
alate and 6 apterous females taken with type.
“The color, markings and armature of hind legs separate this
species from R. femoralis, Champion.”
Strupy OF THE GreNus RHAGOVELIA 889
BIBLIOGRAPHY
Amyor, C. J. B., and Srrviie, J. G. A.
1843. Histoire Naturelle des Insects. Hémiptéres. Lib. Enc. de Roret,
Paris Ixxvi ++ 675 pp., 12 pls.
Bacon, J. A.
1948. Some New Species of Rhagovelia (Hemiptera, Veliidae). Jour.
Kansas Ent. Soc., vol. XXI, no. 8, pp. 71-87.
Banks, N.
1910. Catalog of the nearctic Hemiptera-Heteroptera, American Ent. Soc.,
Philadelphia. viii + 103 pp.
Barper, H. G.
1914. Insects of Florida II. Hemiptera. Bull. American Mus. Nat. Hist.,
vol. XXXII, pp. 495-535.
Brercroru, E.
1914, Four New American Hemiptera. Psyche, vol. XXI, pp. 73-75.
Biatcuiey, W. S.
1926. Heteroptera of Eastern North America. Nat. Pub. Co., Indian-
apolis. pp. 1-1116.
Breppin, G.
1898. Studia hemipterorlogic. Jahr. Nat. Ver. Magdeburg. p. 14.
Burmeister, H. C. C.
1835. Handbuch der Entomologie, Bd. II, Berlin. ii 4- 400 pp., 2 pls.
Burier, A. G.
1876. Preliminary Notice of New Species of Orthoptera and Hemiptera
Collected in the Island of Rodriquez by Naturalists Accompanying
the Transit of Venus Expedition. Ann. Mag. Nat. Hist., Fourth
Ser., vol. XVII, London. p. 411.
Carpenter, G. H.
1898. Trochopus and Rhagovelia. Ent. Month. Mag., Ser, 2, vol. IX,
pp. 109-112.
Cuamprion, G. C.
1901. Rhynchota Heteroptera Il in Biologia Centrali-Americana, London.
xvi + 416 pp., 22 pls.
Coxer, R. E., Mrusar, V., and Ricr, R.
1936. Swimming Plume and Claws of Broad Shouldered Water Strider
Rhagovelia flavicincta Bueno. Bull. Brooklyn Ent. Soc., vol. XXXI,
pp. 81-84. .
Distant, WM. L.
1901. Rhynchotal Notes, XI. Heteroptera. Ann. Mag. Nat. Hist., vol.
VIII, London, pp. 464-486,
1903, Report on Rhynchota, Pt. I, Heteroptera, Fascicula Malay Zoo...
vol. I, London, pp. 219-272, pl. 15.
Dover, C.
1928. Notes on a Collection Acquatic Rhynchota from the Buitenzorg
Museum. ‘Treubia, vol. X, Buitenzorg. pp. 65-72.
890 Tue Unriversiry SCIENCE BULLETIN
Drake, C. J.
1948. Notes on Philippine Rhagovelia, Proc. Ent. Soc. Washington,
vol. L, no. 8, pp. 61-62.
1949. Concerning some Tropical Rhagovelia (Hemiptera: Veliidae). Bol.
Ent. Venezolana, vol. VII, pp. 141-144.
1953. New Neogaean Rhagovelia (Hemiptera: Veliidae). Proc. Biol.
Soc. Washington, vol. XLVI, pp. 145-152.
Drake, C. J., and Harris, H. M.
1927. Notes on the Genus Rhagovelia with Descriptions of Six New
Species. Proc. Biol. Soc. Washington, vol. XL, pp. 131-138.
1931. Further Notes on the Genus Rhagovelia. Pan Pacific Ent., vol.
VIII, pp. 38-85.
1933. New American Veliidae. Proc. Biol. Soc. Washington, vol. XLVI,
pp. 45-54.
1935a. Concerning Neotropical Species of Rhagovelia, Proc. Biol. Soc.
Washington, vol. XLVIII, pp. 33-38.
1935b. New Veliidae from Central America. Proc. Biol. Soc. Washington,
vol. XLVIII, pp. 191-194.
1936. Notes on American Water Striders. Proc. Biol. Soc. Washington,
vol. XLIX, pp. 105-108.
1938. A New Rhagovelia from Cuba. Pan Pacific Ent., vol. XIV, p. 152.
Exsiom, T.
1926. Morphological and Biological Studies of the Swedish Families of
Hemiptera-Heteroptera. Zool. Bid. Fran Uppsala, vol. X, 1, pp.
29-279, 267 figs.
Goutp, G. E.
1928. A New Species of Rhagovelia. Trans. American Ent. Soc., vol.
XXI, pp. 417-418.
1931. The Rhagovelia of Western Hemisphere with Notes on World
Distribution. Kansas Univ. Sci. Bull., vol. XX, pp. 5-61.
1933. Some Notes on Genus Rhagovelia with Descriptions of a New
Species. Ann. Ent. Soc. America, vol. XXVI, pp. 465-470.
1934a. A Change in Name in the Genus Rhagovelia. Bull. Brooklyn Ent.
Soc., vol. XXIX, p. 56.
1934b. A New Genus of Veliidae. Jour. Kansas Ent. Soc., vol. VII, pp.
56-61.
Gutrin-MiGnevILye, F. E.
1857. Hémiptéres in Historia Fisica Politica y Natural de la Isla de Cuba
(by D. Ramon de la Sagra). vol. VI, Paris. pp. 1-868, 20 pls.
Hacuunp, C, J. E.
1895. Verzeichniss der von Yngve Sjéstedt im Nordwestlichen Kamerunge-
biete eingesammelten Hemipteren. Ofversigt K. Vet. Akad. Ford-
handl., Stockholm. p. 476.
Hate, H. M.
1926. Studies in Australian Aquatic Hemiptera. VII. Rec. S. Aus-
tralian Mus., vol. III, Adelaide. no. 2, p. 203.
Hopervanpt, L.
1941. Sbornik entom. odd. Zemskeho Musea v. Praze., pp. 72-73, figs.
go
STUDY OF THE GENUS RHAGOVELIA 891
Huncrrrorp, H. B.
1920a. The Biology and Ecology of Aquatic and Semiaquatic Hemiptera.
Kansas Univ. Sci. Bull., vol. XI, pp. 1-328, 80 pls.
1920b. The Male Genitalia as Characters of Specific Value in Certain
Cryptocerata (Hemi-Het.). Kansas Univ. Sci. Bull., vol. XI, pp.
329-335, 2 pls.
1944, Some Venezuelan Aquatic Hemiptera. Zoologica, vol. XXIX, 3,
p. 129,
Hussry, R. F.
1925, Some New or Little Known Hemiptera from Florida and Georgia.
Jour. New York Ent., Soc., vol. XXXIII, pp. 61-69.
Jounsron, C. W., and Fox, W. J.
1892. A List of Hemiptera Collected in Jamaica, West Indies. Ent.
News, vol. III, pp. 59-60.
Kirpy, W. F.
1890. Insecta, excepting Coleoptra, in H. N. Ridley’s “Notes on the
Zoology of Fernando Noronha.” Jour. Linn. Soc., vol. XX, pp.
530-548.
Kirxatpy, G. W.
1899a. Sur quelques Hémiptéres aquatiques noveaux ou peu connus. Rev.
Ent. franc., vol. XVIII, pp. 85-96.
1899b. Viaggio del Dr. Enrico Festa nell’ Ecuadore e regioni vicine., XIX.
Aquatic Rhynchota. Boll. Mus. Torino, vol. XIV, no, 350, p. 9.
1899c, Aquatic Rhynchota. Boll. Mus. Torino, vol. XIV, no. 350, p. 4.
1900, Notes on Jamaican Rhynchota No. 2. Ent., vol. XXXII, London.
pp. 70-78.
1901, On some Rhynchota, Principally from New Guinea (Amphibi-
corisae and Notonectidae). Ann. Mus. Civ. de Stor. Nat. di
Genova, vol. XX, ser. 2, pp. 804, 810.
1901-1902. Notes on the division Veliiaria (Rhynchota subfam. Velidae).
Ento., vol. XXXIV, London. pp. 285-286, 308-310; vol. XXXV,
1902, pp. 63-64,
1908. Memoir on a few Heteropterous Hemiptera from Eastern Australia.
Proc. Linn. Soc. New South Wales, vol. XXXII, Sidney. pp. 768-
788.
Kmarpy, G. W., and Torre-Bueno, J. R. pe LA
1909, Catalogue of American Aquatic and Semiaquatic Hemiptera. Proc.
Ent. Soc. Washington, vol. X, pp. 173-215.
Lrernimerry, L., and Severin, G.
1896. Catalogue Général des Hémiptéres. vol. III, R. Friedlander, Berlin.
pp. 1-275,
Lunpstap, O.
1933. Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von
Sumatra, Java und Bali, Arch, Hydrobiol. Supp., Bd. XII, Stutt-
gart. pp. 1-195, 263-489.
1936. Die altweltlechen Arten der Veliidan-gattungen Rhagovelia und
Tetraripis. Ark. Zool., vol. XXVITIA, Stockholm. no. 21, pp. 1-63.
1937, Eenige neve oder wenig bekannte ostaseatesche Rhagovelia. Arten.
Ent. Tidskr., vol. LVIII, pp. 1-9,
892
Tue Universrry ScreNcE BULLETIN
MatsumaRa, S.
1913. Thousand Insects Japan Addit., vol. I, p. 98.
Mayr, G. L.
1865. Diagnosen neuer Hemipteren. II in Verh. Zool-Botan. Ges in
Wien, vol. XV, Wien. pp. 429-446.
1867. Reise der Osterreichischen Fregatte Novara. Zool. Thiel, Bd. Il,
Hemiptera, p. 180.
Parsuiey, H. M.
1922. Report on a Collection of Hemiptera-Heteroptera from South Da-
kota. South Dakota State Ent. Tech. Bull, Brookings, S. D. no. 2,
pp. 1-22.
PeyrourEau, A.
1895. Contribution a etude de la Morphologie de larmure genital des
Insects. Bordeaux. pp. 1-248.
Porsson, R.
1926. Hémiptéres aquatiques noveaux ou peu connus de l’Afrique du nord.
Bull. Soc, Hist. Nat. Afrique N., vol. 17, Alger.
1945. Bull. Soc. Ento. France, July, Paris. pp. 91-92, figs. 4 & 5.
Reuter, O. M.
1882. Ad cognitionem Heteropterorum Africae occidentalis. Ofversigt
finska Vet. Soc. Férhandl., vol. XXV, Helsingfors. pp. 1-48.
SicnoretT, V.
1877.
Espéces composant le genre Rhagovelia. Bull. Soc. Ent. France
(5), vol. VII, Paris. pp. 53-55.
Sincu-Prirut, Hem.
1925. The Morphology of the Male Genitalia in Rhynchota. Trans. Ent.
Soc. London, Aug., pp. 127-267.
Smiru, J. B.
1909. A Report on the Insects of New Jersey—Hemiptera. Ann, Report
New Jersey Mus., pp. 131-170.
Sraz, C,
1865. Hemiptera Africana. Officina Norstedtiana, vol. III, Stockholm.
pp. 1-200.
Torre-Burno, J. R. DE LA
1907.
1916.
1921.
1923.
1924,
1928,
On Rhagovelia obesa Uhler. Canadian Ento., vol, XXXIX, pp-
61-64, 3 figs.
The Veliinae of the Atlantic States. Bull. Brooklyn Ent. Soc.,
vol. XI, pp. 52-61.
New Records of Aquatic Hemiptera for the United States. Ent.
News, vol. XXXII, pp. 273-276.
in Hemiptera or Sucking Insects of Connecticut. State Geo, Nat.
Hist. Surv. Bull., vol. XXXIV, pp. 1-807, 20 pls.
The Neartic Rhagoveliae. Trans. American Ent., vol. L, pp. 243-
252.
Rhagovelia in a List of the Insects of New York with a List of the
Spiders and Certain other Allied Groups. Cornell Agri. Expt. Sta.
Memoir 101, pp. 1-1121, 1 map.
Srupy OF THE GENUS RHAGOVELIA 893
Unter, P. R.
1871. Notices of some Heteroptera in the Collection of Dr. T. W. Harris.
Proc. Boston Soc. Nat. Hist., vol. XIV, pp. 93-109.
1878. Notices of the Hemiptera-Heteroptera in the Collection of the Late
T. W. Harris, M. D. Proc. Boston. Soc. Nat. Hist., vol. XIX, pp.
865-446.
1884, Hemiptera in Kingsley’s Standard Natural History, vol. I, S. E.
Cassino and Co., Boston. pp. 204-296, 1 pl.
1886. Check List of Hemiptera-Heteroptera of North America. Brooklyn
Ent. Soc., Brooklyn, pp. 1-34.
1893, A List of the Hemiptera-Heteroptera Collected in the Island of
St. Vincent by Mr. Herbert H. Smith, with Descriptions of New
Genera and Species. Proc. Zool. Soc. London, pp. 705-719.
1894a. On the Hemiptera-Heteroptera of the Island of Grenada, West
Indies. Proc. Zool. Soc. London, pp. 167-224.
1894b. Observations Upon the Heteropterous Hemiptera of Lower Cali-
fornia, with Descriptions of New Species. Proc. California Acad.
Sci., ser. 2, vol. IV, pp. 223-295,
Van Duzer, E. P.
1916, Check List of Hemiptera (excepting Aphididae, Aleurodidae, and
Coccidae) of America, North of Mexico. New York Ent. Soc.,
New York. xi-+ 111 pp.
1917. Catalogue of Hemiptera of America North of Mexico excepting
Aphididae, Coccidae, and Aleurodidae. Univ. California Press,
Berkeley. xiv + 902 pp.
Wuiter, F. B.
1879. Description of New Hemiptera (I). Journ. Linn. Soc, London.
Zoology, vol. XIV, pp. 482-489,
Srupy OF THE GENUS RHAGOVELIA 895
INDEX
PAGE
Dina (OU e luis vada Gee har nds Coro Lanes ee 766
PADEUMUA TOUR se ty y iy cche tein tins ac leserQtcniies rit G 754
USlpbaRvauday inanaythene) CO NEANC pra Geter ee aur ne iy ee aS 754
GOW PULOONAAIDERRE ei hiv (eS CES EEE ORG, CO EI et re 879
PROMMOWIGULOINENTS i578 Cel oe cee CPE a cad eet 698
ORGUVINU See Ae Re es ek ta CPLA OE 1S Ear Oa Oe ee 699
GOUT GU: BON TIOVG ve Cee ek CUCL TN Eta Oe eat et er i i 817
GUS) TOEB RO ONG MIOITIS bibs cece eek O Sie 0 8 en Os 857
PRINCE COU at ey reed COL CLINE ECAR CREST TCC UCL EEN Na 857
EXITS EL ALOUD ROY, UOe crc 41a 8 bins ae ct 5 cs 8 id eNO hs SN 857
WTRRONE SISA SOUL 6 7 eee Cu 0F Hiv V6 e tie eons oy & 5/4 Ries iy alee, Gee ee 780
RUST GC. WHIGK hp 0 Wea y cove cr ta ee be bs cd aad ens ved Ve ey 713
RUSTIC RIOUD «+c vinrke hve eta UC 4 tha ROCI4 bay wy vee es SIN 710
ATOUSUeS BrOUD, KOY (Orecryrr ss reece ecererreus etry iii CENA Sie RK 710
GinOrd: (GBUDIMOISUOL) ste ones cee cer br py CRE EV CC nt yy bese ever ene 820
Buner BEVetoth, (tia cs50 0h Chis) bavared bia etd cee oye ee 876
Deaki lWihie ANG TIGItSs vac nieve ee ya oy vce evr sieves eee 859
Uitollo}a/ose oer a Ue a eae tae ee ee re 698
MASEHOLO BAC OWc «it cc on el Cs 5 ys ee ee 715
GUUAHO: OTRO: At GRLAtIS «oak as Cie 4 cee 4k a ee ge Tee
GuiOny IorAke ANC Tents: cv ca eaves 1 ie OC ce ee 718
iclelrged al ela tel @ oh bila esr eRe ey! URW ee en eee) eer era Oe ree ten 782
Ger eUyas LSCOU cc ree ue ws Cv 1 WERE 1 a Pes ee 864
CGT DUR G ote Reiss ets 0 was NW 8 dn i ce 879
UHOre GISULINGIStRD) (ea cts Fi aid <3 vet 0 0d ae hay eet es eae 822
MOUIRIIG TOU aie or ee es Chek pe ae 816
MOUS MEG Uo hey NOs 10s Cine 0 eis aN eu ae ee 817
NSOIOGtING TEGHMEUCS 15 si its cs is Git delat 5 ross ene este pe 702
MOSUL IDTUCO! Cay cn et Vis PUA ey eA ee 768
GrassvpRs GHAMPION «yoshi eS tect Ave bette ter VA VE esas ba es 783
Steosies STOUP 4 eds cere ssi eee sos steak ee ee. 778
MeTesa Mes GlOUb) REY tOs chat Cures i Weeden hie ise 778
Ousmane Irae ent) FIAT. c. 66h ich ine dh eee aOhl Oi eek eet 826
ee AGO cn 0 he er cs es ee a ee 721
Gienicte GUBTOTION 6, ri ules Clie ee ee 838
IPN isinalloihialosal ae te arpraetee uth) eee eer ty tre fe eC ey 705
lee Mat vets oe a se A ee ee 770
MHIGWHASGETOUD. 6.6 he ee tc CO Oe RE, ee Le Grek GN ener Gs 768
MUGUanMN TOLD) MOY W011 veces oer k se yc Ce i ayes og gg eee 768
UVES UCOH ane fv ccn ice y cde ures 150 WS Ak ee de er 722
Herroraie: GHBMNENOT fia oa rick ee vs rs he 786
UCAS hd cai hid is ee eye erg reer ce 723
MOMEOROUe BAOOR ayy eae ty lr es se cas 724
(ojopal Ae eee oh eoee s)he ee oe Cee ay Pe eer erg 700
OTOSN SD, HOV. fics sie 11 ve Carey sd Creuse PO UEN COs UCC UVES CREE OE 867
896 Tue University SCIENCE BULLETIN
Getietic COnCepts icp mrs PMG omer cer Pi rs Cents
Genie RRGGOCEHA ied cof ger eh eter OR Te HCCI TES EUs
Genus, key to species ZrOUPS.... 66. cece cece teen eect eee e ete c ts
Bro GD. WON cw care vn es eee e ee cn GaP Cares eh Os Oks AES he
HAA gs re ah ae Gy
hambleiont Drake and Harvis......... 6.0 ete recess ees geen! anne
TeraAlOn ives ck a bs ce ee Og tenga iets 4)
hirtipes Drake and Harris.......:-.ce cece certs ented eters ae ce se nes
Hittipes LOUD «ihc ch ce res tea eee Orc gee tal mt
Hirtipes group, key to..s. ccc cee eee e eens eee tee ena nein spent eet as
Void BD) DOV eres ret Ere CCC ree ts Ces big ten
Identification techniques ...... cc cc cee eee eee e ete e een n ewe meetin
ignota Drake and Harris......... cesses sere cere eee e tne enen eens
Anibatiit BACON Gc. ccc ee Oe art OVC EEE NOEs ORO
Trimature StACCS cece tee reece cece ncn reteset ee renee ee Ee
WAPCNEG BD. MOV. eo. cei ee era ee ea een! Bes
treularis CHAMOION. cc ee oh Ve ee eg
HigHalan WUERS a. vec heh ei ei ed Cra eae 1 clic 8 ety aan tog hm
faneita, DAK ic ce cee eek me ROT re are
piihtd BACON Vi igi Cee eae cn a) Ly ca ee
knighti Drake and Harris. ....... 0000s seven eens rece een n nee een e ans
lonaipes Gould icc cc rhe en cree nha, ce A NTE T
Nidldla Gould 66h te ie COO era ERT EVs DAE ish
MAB i ee le rs a et aoe cus COE i
Mieiad EDV TIOV he bia REL Cer te teen ays
mira Drake and Haris. ccc i ice cei lie iene ieee ies
WOU OME SDL TOV) FFU eh EIT nee ces)
WIC ACGN he va ek oe ee VOY La ee evi k ge byl a io OT ts
Wound DOUAKe Sickie iene er ON ees er er ee
phesa Ube iii ii lev issues tise el nie’ (line han elas D
Opes OUD eG cena Navt Fidel Coho TSS a cee
@hesa group, Key tO... ee Coe yan
Odin BACON ee i eed 1s eed le BE ONE Ae te Ay Us
ONHUSr CAPSHIBV fii Fes OO i) 1s cow god fit
Ovlpostton: 5s ir er oe a Hy Ch Ger te re gs
POled SPL HOV: (bebe cue eh er es Fi haeaa eT
panda Drake and Harris......... 06. . secs see e cee tree een eee eens
POUNONG TRO Foe cee et rE Te OL i Os yaa ees
MEPACOGG BACON. ives cece ge ete tie cure ene eee ee MAS
i MedlelCsrn/ oe 1 te, Oe iw ieee 2 ea ee
plana Drake and Harts: s. 1.6. ieee nem ic:
Pisnipee GOW as eke eer rte Unie ts Cen cis
Wlaiben UM pias ek Gh el rs at eee en eva a
Wollota GOW ie ce ps AV oe er rd Gun esd
#OUNG DLAC i oi ed hd WEA ed Ie OA re een
pple CGO ys oe cs Cee Le a ee
FODUSA GOW 7 i ea en ete er i ere wa
Srupy oF THE GENUS RHAGOVELIA 897
PAGE
galing (Champion) ........ces cece cece seen eect e ence nee e een ees 736
SCGDTA SP. NOV, os. sce e cece seer e ener e eee een seen nese cee ewen een ees 831
SCHULTE SP. NOV, bic r cesses ve ered esac Seceeeneueeunvasareerseusssees 804
ily rleti sa Ota] rike (ater Pompe rarg ere eer ee eee 806
GOlG BP. DOVe os ide cv cere eevee seer es even teer steed eset e sant eueys 833
Species groups, key tO... ....ccecec ence eee eee e tenn nee e ee ee eee ees 709
spinigera Champion ........ cece cence eee e nee nee e ene eennnes 873
Spinigera group ...... cscs screenees ences ena ee ence sees asaeensenes 863
Spinigera group, key to.... 0.60. s cece eee e seen eee e nn en eee eeenes 864
Roh COU i ir tes cs eens Cnt Cea deen e res ee cent 737
tantilla Drake and Harris... 0... 0.0 cece cee cece eee eee e eee en eens 739
toylonielia Kirkaldy oi cv cue c ce gee eee cree encase cenee yes 835
Taxonomic histOry 1... css w eee ces ener eneeseneseeeeeeeseeeerens 703
tenuines Champion .......ccsesccs ere en eee nee neseneeeeseeuwanenes TAL
torquata Bacon os .ciccyeveceeccsceveeeteeuneeeeceeeeeeereuss tens 760
torreyana Drake and Hussey...........cssceeeseeeee eet en een en eens 885
PATE OVER ei ie ci ok 0 Ay © ESN USS Cu 8 ONES CS 878
Hepida BACON iin ee ee ie eed le Ek ees es se rect eens 761
Uonpula Intake 2g ks ok be RMA hehe COL Che ies sae es 886
WTUTOIG IOTAMA: 6 oy ee i AGO gs CELA FORO ee eS 775
Crista: Gold wre ice EE SR hd a ro 763
Uncintta Champlon oe 0 Oh cee ee cs vee ha eo eae ess Certs 776
Daripds CHOTODION os cc et cere e cee yee e te et Taper as 808
welocis Drake and Harris... ... 0. ccc cece cee e cena secu ences ernsess 745
wersuta Drake and Hartis........ ccc cece cee eee ee een eee ene eee ees 747
IORI Bp. TOV is cok ee are a ed on Seg AS es EE 751
OUST BACON ik be bole Fy ONE 4 8 ERE AE OC EES 765
whitel (Breddin) ii. .ci suse s cece cece ces eee une cet e ee seaeuress 810
Willlamst Gould i. cic ves ce vee ere en 0 oa yess abet eter ete tees 812
Botert Wiebe oc es ea ee as yy a Oe 5 ee 887
Tue: Universiry ScriENCE BULLETIN
PLATE I
Fic. 1. Sutured off pronotum of angustipes group, dorsal view.
Fic. 2, Abbreviated pronotum of abrupta group, dorsal view.
Fic. 3, Pronotum covering mesonotum, dorsal view.
Fic. 4. Head, pro-, and mesothorax, showing minute, conical, black setae
on base of jugum of head and proepisternum, ventral view.
Tue Universiry Science BULLETIN
PLATE II
Fic. 1, Dorsum of abdomen of apterous female tapered evenly.
Fic. 2. Dorsum of abdomen of apterous female narrowed after first three
segments,
Fic. 3. Front tibia excavate and expanded, lateral view.
Fic. 4, Intermediate femur of female constricted near middle, lateral view.
Fic. 5. Intermediate femur of female flattened from basal one fourth to
apical one fourth, lateral view.
Fic. 6. Normal femur of female, lateral view.
Fic. 7. Posterior femur armed at apex with hook; elegans group.
Fic. 8. Mucronate last genital segment of female, dorsal view.
Fic. 9. Mucronate last genital segment of male, dorsal view.
Fic.
THE
University ScreNcE BULLETIN
PLATE III
Rhagovelia angustipes Uhler; right clasper, lateral view.
Rhagovelia bisignata Bacon; right clasper, lateral view.
Rhagovelia callida Drake and Harris; right clasper, lateral view.
Rhagovelia calopa Drake and Harris; right clasper, lateral view.
Rhagovelia deminuta Bacon; right clasper, lateral view.
Rhagovelia evidis Bacon; right clasper, lateral view.
Rhagovelia fontanalis Bacon; right clasper, lateral view.
Rhagovelia hambletoni Drake and Harris; right clasper, lateral view.
Rhagovelia imitatrix Bacon; right clasper, lateral view.
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia janeira Drake; right clasper, lateral view.
longipes Gould; right clasper, lateral view.
modesta Bacon; right clasper, lateral view.
plana Drake and Harris; right clasper, lateral view.
plumbea Uhler; right clasper, lateral view.
salina (Champion); right clasper, lateral view.
spinosa Gould; right clasper, lateral view.
tantilla Drake and Harris; right clasper, lateral view.
tenuipes Champion; right clasper, lateral view.
velocis Drake and Harris; right clasper, lateral view.
versuta Drake and Harris; right clasper, lateral view.
paulana Drake; right clasper, lateral view.
viriosa Bacon; right clasper, lateral view.
STuDY OF THE Genus RHAGOVELIA
PLATE Ill
ANGUSTIPES GROUP
Qt oe
R. ANGUSTIPES R. BISIGNATA R. CALLIDA R.CALOPA
i 2 3
4
Oy ONa owe CO,
R.DEMINUTA R.EVIDIS R.FONTANALIS RHAMBLETONI — R. IMITATRIX
R. LONGIPES R.MODESTA R.PLANA R.PLUMBEA R. SALINA
i " i iB \4
R. VELOCIS
R. SPINOSA R. TANTILLA R. TENUIPES i‘
Is 17
R. VERSUTA RPAULANA R. VIRIOSA R JANEIRA
19 20 2\ 22
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
FIG,
Fic.
Fic.
Tue Universiry Science BULLETIN
PLATE IV
1. Rhagovelia abrupta Gould; right clasper, lateral view.
2. Rhagovelia lucida Gould; right clasper, lateral view.
8. Rhagovelia torquata Bacon; right clasper, lateral view.
4, Rhagovelia trepida Bacon; right clasper, lateral view.
5. Rhagovelia trista Gould; right clasper, lateral view.
6A. Rhagovelia vivata Bacon; right clasper, lateral view.
6B. Rhagovelia vivata Bacon; right clasper, dorsal view.
7, Rhagovelia elegans Uhler; right clasper, lateral view.
8. Rhagovelia insularis Champion; right clasper, lateral view.
9. Rhagovelia merga Bacon; right clasper, lateral view. I
10. Rhagovelia uncinata Champion; right clasper, lateral view.
Srupy or THE GENUS RHAGOVELIA 905
PLATE IV
ABRUPTA GROUP
po) as
R. ABRUPTA R. LUCIDA
2
oo ee ee ee
R TORQUATA R. TREPIDA R.TRISTA
3 4 5
on} fees
R.VIVATA
6
ELEGANS GROUP
ian 2
R. ELEGANS R. INSULARIS
7 8
R. UNCINATA
R. MERGA (0
9
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic,
Fic.
Fic,
Fic.
Fic.
Fic.
Fic.
iG,
THE
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
Rhagovelia
UNIVERSITY SCIENCE BULLETIN
PLATE V
amazonensis Gould; right clasper, lateral view.
castanea Gould; right clasper, lateral view.
crassipes Champion; right clasper, lateral view.
femoralis Champion; right clasper, lateral view.
horrida Bacon; right clasper, lateral view.
jubata Bacon; right clasper, lateral view.
nitida Bacon; right clasper, lateral view.
ornata Bacon; right clasper, lateral view.
palea Bacon; right clasper, lateral view.
perfidiosa Bacon; right clasper, lateral view.
relicta Gould; right clasper, lateral view.
robusta Gould; right clasper, lateral view.
scitula Bacon; right clasper, lateral view.
sinuata Gould; right clasper, lateral view.
varipes Champion; right clasper, lateral view.
whitei (Breddin); right clasper, lateral view.
williamsi Gould; right clasper, lateral view.
Srupy oF THE GENUS RHAGOVELIA
PLATE V
CRASSIPES GROUP
see
| R. AMAZONENSIS 2: R.CASTANEA 3 RCRASSIPES
}
4 R. FEMORALIS « RLHORRIOA
| 6 RJUBATA
Se ntyneeee g R. ORNATA
| 7 RNITIDA
9 R PALEA IO R PERFIDIOSA Il R. RELICTA
2 RROBUSTA 13 R.SCITULA 14 RSINUATA
| bn OL
IS RVARIPES 16 R. WHITE! 17 R. WILLIAMS!
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
EKG,
Fic.
Fic.
Fic.
Fic.
. 12. Rhagovelia tayloriella Kirkaldy; right clasper, lateral view.
Tue Untversiry Science BULLETIN
PLATE VI
Rhagovelia choreutes Hussey; right clasper, lateral view.
Rhagovelia formosa Bacon; right clasper, lateral view.
Rhagovelia ignota Drake and Harris; right clasper, lateral view.
Rhagovelia spinigera Champion; right clasper, lateral view.
Rhagovelia acuminata Bacon; right clasper, lateral view.
Rhagovelia armata (Burmeister); right clasper, lateral view.
Rhagovelia collaris (Burmeister); right clasper, lateral view.
Rhagovelia impensa Bacon; right clasper, lateral view.
Rhagovelia planipes Gould; right clasper, lateral view.
10. Rhagovelia scabra Bacon; right clasper, lateral view.
11. Rhagovelia solida Bacon; right clasper, lateral view.
Sera Atkonr
(=
Oo
Srupy oF THE GENUS RHAGOVELIA 9
PLATE VI
SPINIGERA GROUP
R. CHOREUTES R. FORMOSA
y 2
R.IGNOTA R. SPINIGERA
3 4
COLLARIS GROUP
.
R. ACUMINATA
R. ARMATA
6
5
R. COLLARIS R. IMPENSA
ig
9
8
R.PLANIPES R. SCABRA
10
R SOLIDA R. TAYLORIELLA
i
12
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Fic.
Tue Universtry SCIENCE BULLETIN
PLATE VII
Rhagovelia ainslici Drake and Harris; right clasper, lateral view.
. Rhagovelia becki Drake and Harris; right clasper, lateral view.
Rhagovelia gracilis Bacon; right clasper, lateral view.
Rhagovelia distincta Champion; right clasper, lateral view.
Rhagovelia knighti Drake and Harris; right clasper, lateral view.
Rhagovelia obesa Uhler; right clasper, lateral view.
Rhagovelia oriander Parshley; right clasper, lateral view.
. Rhagovelia rivale Bueno; right clasper, lateral view.
. Rhagovelia hirtipes Drake and Harris; right clasper, lateral view.
Srupy oF THE GENUS RHAGOVELIA 911
PLATE VII
AINSLIE| GROUP
a) Gx
R. AINSLIE! R. BECKI
| 2
R. GRACILIS
3
OBESA GROUP
R. KNIGHT
R. DISTINCTA .
4
R. OBESA R ORIANDER R RIVALE
6 7 8
HIRTIPES GROUP
R_ HIRTIPES
Tue University ScreENCE BULLETIN
PLATE VIII
Fic. 1. Rhagovelia citata Drake; right clasper, lateral view.
Fic. 2. Rhagovelia costalimai Drake; right clasper, lateral view.
Fic. 3. Rhagovelia mira Drake and Harris; right clasper, lateral view.
. 4, Rhagovelia torreyana Drake and Hussey; right clasper, lateral view.
Srupy OF THE GEeNuS RHAGOVELIA
PLATE VIII
ADDENDA
fm 2
R.CITATA
\
R. COSTALIMAI
2
we I
R MIRA
80—3378
R.TORREYANA
4
918
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVIII, Pr. I] = Drcemper 20, 1956 [No. 11
A Supplementary Taxonomic Study of the Genus Rhago-
velia (Hemiptera, Veliidae) of the Western
Hemisphere. A Deductive Method’
BY
Ryvicut Matsupa
Department of Entomology, University of Kansas
Assrract: This paper is intended to supplement Bacon's “A taxonomic
study of the genus Rhagovelia (Veliidae, Hemiptera) of the Western Hemi-
sphere.”
Various taxonomic characters especially in winged forms are described and
discussed. It is found that the genus Rhagovelia should be subdivided into
three subgenera on the basis of three correlated characteristics, namely the wing
venation, the pronotum in wingless forms, and the longitudinal carinae in dorsal
abdominal segments in winged forms.
Descriptions of two subgenera, Rhagovelia s, str. and Neorhagovelia, are
given. Also a few additional characters which this study has revealed are dis-
cussed from the viewpoint of phylogeny.
INTRODUCTION
As a supplement to Bacon’s paper, “A taxonomic study of the
genus Rhagovelia (Hemiptera, Veliidae) of the Western Hemi-
sphere,” 2 the present study was undertaken to expand Bacon's work
referring especially to winged forms of this genus, which have
hitherto been more or less neglected by workers with this group of
insects.
The problem with which the writer is primarily concerned in
this study is, therefore, to determine the taxonomic value of the
winged forms in this genus. This involves the study of a peculiar
structure that occurs on the basal dorsal abdominal segments be-
neath the hemelytra in many families of Hemiptera-Heteroptera.
The taxonomic significance of this structure was suggested by
1. Contribution No. 921 from the Department of Entomology of the University of
Kansas.
2. Bacon, J. (1956), Univ. Kansas Sci. Bull., vol. 38, pp. 695-913.
( 915 )
916 Tse Universiry SCIENCE BULLETIN
the writer ® who indicated that each particular pattern of fusion or
modification in the first and second abdominal tergites and the
resulting structures occur (1) at the subgeneric or generic level,
or (2) in more than one genus within a tribe or subfamily, (3) in
more or less constant form within a tribe or subfamily.
The writer’s first step was to investigate the structure in question.
It soon became evident in the course of the study that there occur
three distinct forms of the structure which have been shown to be
represented by paired carinae in this genus. This indicates that
there should exist three distinct groups within the genus if the first
category of the hypothesis introduced above holds true. The next
step taken was to correlate these three distinct types of carinae with
other structures in other parts of the body in series of species in both
winged and wingless forms for the test of the first category of the
hypothesis. Forewing venation, pronotum in winged form, pro-
portional length between dorsal abdominal segments in both winged
and wingless forms, etc., were observed and described for each
species for this purpose. The relative lengths of femur, tibia, and
tarsal segments in intermediate and hind legs given by Bacon in
his descriptions were also freely analyzed to find significant correla-
tions.
Forty-five out of seventy-three species, i. ¢., over sixty percent of
species treated by Bacon, were available for the study of winged
forms. This number of species will probably be large enough so
that a significant conclusion can be drawn with a fair degree of
certainty.
ACKNOWLEDGMENTS
The writer is grateful to Professor H. B. Hungerford for his kind
guidance, to Professor R. E. Beer and Professor A. R. Barr for their
constructive criticism.
MORPHOLOGY AND DESCRIPTIVE TECHNIQUE
The morphology of the family Veliidae was studied by Spooner
(1938) on the head of Rhagovelia obesa, by Larsén (1945) on the
thorax of Velia currens, and by Singh-Pruthi (1924) on the genital
segments of Velia currens.
In the following report a rather simple morphological treatment of
winged forms of the genus Rhagovelia is given together with the
descriptive technique associated with the present study.
8. Matsuda, R. (1955), The morphological and taxonomic significance of the basal
dorsal abdominal segments in Hemiptera-Heteroptera. Pan-Pac, Ent. 31, 2 pp. 73-74.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 917
Tue Heap (Pl. 1)
The compound eyes are large, usually as wide as the vertex at the
posterior margin. The cleavage line begins at the posterior mar-
gin of the vertex and extends down to the anteclypeus without
forming a typical Y-shaped divergence anteriorly. The labrum is
a small plate located anterior to the anteclypeus and with a slender
median projection, apically more or less enclosed by the bucculae
Qleavage \ine /Nertex
~~ / 4 Oa
Antenna aS Z Se
~ = TA A Ped
Babe N é vy
Anteclypeus— one P araclypeus
~- Maxillary
Plate
~SRostrum
~~Metasternum
Transverse ~ se
suture ~ “Coxa
N
N
«
Trochanter
/
/
7
2nd Abd. Sea.”
L RAR.) insularis
Puare 1. Rhagovelia (Rhagovelia) insularis Champion. Ventral view.
918 Tue University Science BULLETIN
(maxillary plate). The paraclypeus of Spooner (1938) lies laterad
to the anteclypeus. The gena is the area between the lateral mar-
gin of the paraclypeus and the eyes. The rostrum is four-segmented
with the second segment the longest. The antenna is four-seg-
mented with a conspicuous protuberance at base between the
lateral margin of the paraclypeal plate and the compound eyes.
The ocellus is absent in this genus.
Tue THORAX
The prothorax (Pl. 1): The pronotum is well developed poste-
riorly, and usually extends back to the first or second abdominal seg-
ment. In females of some species groups of this genus the pronotum
develops into a long and narrow process. The pronotum, the pro-
Preacutum , 7 Prescutum,
a
_Seutum 2
3 ; = ¥
Ay K SS _-P ostnotum
‘ o 2
Scutellum = \
3 ‘SN
ca Antero-lateral
impression
Longifudinal
. te
carina~ Antero-mesol\
impression
\
Postero-latera|
impression
Intevseqmental
qroove
i! ey ‘sd ss 8th Abd. Seg.
Paratergite-~
2 RAR insularis
Puate 2. Rhagovelia (Rhagovelia) insularis Champion. Dorsal view.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 919
pleurum and the presternum are indistinguishably fused. The ster-
nal region is depressed longitudinally to accommodate the rostrum.
The coxal cleft runs anteriorly.
The mesothorax (Pls. 1,2): The mesonotum is completely hidden
beneath the pronotum. The prescutum is the median elongate sub-
pentagonal region defined posteriorly and laterally by the scutum
on which the forewings are articulated. The scutellum is located
posterior to the scutum as a transverse sclerite; it does not develop
posteriorly to be exposed between the wings as in most families of
Heteroptera. The postnotum is a very narrow sclerite running
along the posterolateral margin of the scutum. The pleurum and
the sternum are indistinguishably fused. The sternal region is
slightly depressed. The coxal cleft goes anteriorly beyond the
middle of the ventral surface of the mesothorax. A paired oblique
row of long and white pubescence is disposed from near the middle
of the posterior margin of the prosternum to the inner posterior
angle of the acetabula. This oblique row of pubescence is lacking
in marine forms.
The metathorax (Pls. 1,2): The usual four component parts of
the notum in the metathorax are clearly recognizable. The prescu-
tum is a transverse plate directly behind the posterior margin of the
mesothoracic scutellum. The scutum is an oblique, narrow plate
along the mesothoracic postnotum and the hind wings are artic-
ulated on this plate. The scutellum is clearly defined from the
scutum by an oblique suture and occupies a lateral, rather large
subtriangular area of the metanotum. The postnotum is here in-
terpreted as being represented by the plate behind the scutellum
defined anteriorly by a sinuate suture. The usual scent gland-
opening is sometimes clearly visible behind the posterior margin of
the mesopleurum. The pleurum and the sternum are indistinguish-
ably fused. Another characteristic scent gland in the Veliidae on
the lateral region of the acetabula is provided with a tuft of erected
hairs. Each scent gland opening is connected by a transverse su-
ture across the sternum. This suture is here termed “the ventral
transverse suture” for descriptive purpose. There is no coxal cleft
in the metathorax. According to Taylor (1918) the pleural suture
is frequently absent in Heteroptera. How far the coxal cleft is
associated with the pleural suture in this genus was not investigated,
but the indication is that the coxal clefts in the pro- and mesothorax
do not seem to contain a part of the pleural suture judging from the
course of the cleft.
920 Tue Universitry SCIENCE BULLETIN
Tue Winc (PI. 3)
The wing base’ was not studied.
The forewing: The forewing is not well differentiated into dis-
tinct corium, clavus, and membrane as in other families of Heterop-
tera. The uppermost vein is apparently the subcosta. The next
vein is a union of the radial, median and cubital veins basally.
The lower vein is anal. In one major group of the genus Rhagovelia
two closed apical cells are formed in the distal half. The apical
cells in the other major group of the genus are more or less obliter-
ated or small when present.
The hind wing: The more generalized wing venation is seen in
the hind wing. ‘The second vein is apparently R -+- M basally in-
stead of R-++-M + Cu in the forewing. Cu is basally fused with
anal veins along the anal fold. There is no considerable difference
in hind wing venation between the two major groups of the genus
Rhagovelia.
Tue ABDOMEN
The first dorsal abdominal segment is reduced but still retained
as a distinct plate. This segment is always with a median elevated
area, The second and third dorsal abdominal segments and also
occasionally the fourth segment are provided with paired longitudi-
nal carinae arising from the lateral part of the median elevated
area of the first segment. The second and third segments are always
provided with three more or less distinct laevigate impressions on
the outer sides of the longitudinal carinae; the one immediately on
outer sides of the carinae is called “the anteromesal impression”;
the other two impressions along the inner margin of the connexivum
are called “the anterolateral and posterorlateral impression” respec-
tively in description. The posterior margin of each second and
third segment between the posterior ends of longitudinal carinae
is more or less lengthened and deeply defined obscuring its original
posterior margin and the anterior margin of the succeeding segment.
This region is, therefore, called “the intersegmental groove” in de-
scription, and the numerical value of each segment refers to the
length from the middle of anterior margin of the intersegmental
groove to the same of the intersegmental groove of the succeeding
segments. On the eighth segment there is a laterally partitioned
area; this is especially clearly seen in the winged female and this
plate is supposedly a fragmented part of the tergum, and called
the paratergum in taxonomy. The first ventral abdominal segment
is completely lost. The second segment contributes towards the
SUPPLEMENTARY STUDY OF THE GENUS, RHAGOVELIA 921
{formation of the ventral wall of the metathoracic coxal cavity. The
third to seventh segments are occasionally armed with a median
longitudinal elevation. The description of the genital segments is
left for a study planned in the near future by the writer.
Group CHARACTERISTICS OF THE ANGUSTIPES GROUP
The winged forms of following twelve species out of twenty-three
species treated by Bacon were available for study:
Rh. angustipes Uhler Rh. janeira Drake
Rh. bisignata Drake and Harris Rh, paulana Drake
Rh. calopa Drake and Harris Rh. spinosa Gould
Rh. fontanalis Bacon Bh. tantilla Drake and Harris
Rh. imitatrix Bacon Rh. tenuipes Champion
Rh. longipes Gould Rh. versuta Bacon
In this species group the longitudinal carinae on the basal dorsal
abdominal segments occur always on the second segment usually
as distinct and well-developed carinae reaching to the posterior
margin. The carinae also often occur on the third segment as less
distinct ones which are subject to individual variation as seen in
Bh, janeira, Rh. fontanalis, Rh. spinosa, Rh. versuta, Rh. tenuipes,
Rh. longipes, Rh. calopa, and Rh. bisignata.
Forewing venation is subject to specific variation within the
group which fits more or less three types shown in the accom-
panying text figures (Pl. 3). Small apical cells range from one
to two in number, and these cells are often obliterated in some
individuals within the species. The constant feature of the fore-
wing venation in this species group is that the subcosta never de-
velops further than the middle of anterior margin of the wing and
the apical cell or cells are always formed within the proximal half
of the wing.
The pronotum in winged forms is always bluntly rounded apically
and without sexual difference.
Proportional Jength between sixth and seventh dorsal abdominal
segments in winged males ranges from 32:60 in Rh, janeira to 40:58.
in Rh. fontanalis, that in wingless males ranges from 22:42 in Rh.
spinosa to 38:48 in Rh. tantilla. The proportional length of seventh
and eighth dorsal abdominal segments in wingless females ranges
from 46:35 in Rh. paulana to 43:51 in Rh. tantilla, that in wingless
females ranges from 58:37 in Rh. modesta to 41:48 in Rh. longipes.
In the hind leg the femur is shorter than the tibia in the majority
of species within the group (twenty out of twenty-two species
measured). Only in Rh. callida and Rh, tenuipes the femur is defi-
nitely longer than the tibia.
922
Tue University Science BULLETIN
In the intermediate leg the tibia is less than twice as long as the
second tarsal segment (sixteen out of twenty-two species). The
proportional length of the second and third segments seems to give
group characters. In six species the second tarsal segment is longer
than the third tarsal segment (Rh. bisignata, Rh. callida, Rh. imita-
trix, Rh. longipes, Rh. modesta, Rh. plumbea); in three species (Rh.
plana, Rh. salina, Rh. tenuipes) the second tarsal segment is longer
than the third segment in males, equal or slightly shorter in females;
in the rest of species in this group the second segment is shorter
than the third segment which is true of all species in this genus
except for four species that will be noted later.
In wingless forms the pronotum is shorter than the length of eye,
with the posterior margin straight, sinuate or concave. The dorsum
of the abdomen of the wingless female tapers rather evenly to the
apex.
In spite of slight deviation in some characters in some species
this group seems to be a well-defined natural group.
Rhagovelia angustipes Uhler
(Pl. 8, fig. 6; pl. 4, figs. 8a, b)
1894, Rhagovelia angustipes Uhler, Proc. Zool. Soc. London, p. 125.
Winged female: Pronotum reaching the basal region of second
dorsal abdominal segment. Forewing fuscous, with two well-defined
small apical cells. Proportional length of dorsal abdominal segments
(second to eighth):: 33: 88: 88: 89: 42: 47: 42. Second dorsal
abdominal segment with longitudinal carinae rather narrow, reach-
ing to posterior margin of the same segment; usual three laevigate
impressions shallow, clearly separated from each other; interseg-
mental groove rather shallow. Third dorsal abdominal segment
without longitudinal carina, anteromesal laevigate impression trans-
verse, anterolateral and posterolateral impressions are contiguous.
Connexivum rather strongly reflexed. Visible first connexival seg-
ment short, visible third and fourth connexival segments transverse.
Seventh dorsal abdominal segment transverse, posterior margin
feebly sinuate, seventh connexival segment with lateral margin
straight. Eighth segment with posterolateral margin feebly rounded,
posterior margin substraight.
Wingless female: Posterior margin of pronotum concave, feebly
produced posteriorly at middle, posterior margin of mesonotum
obliterated, Posterior margin of metanotum broadly sinuate. Pro-
portional length of dorsal abdominal segments (first to eighth) ::
\
}
|
|
|
|
|
t
t
SUPPLEMENTARY STUDY OF THE GENUS REIAGOVELIA 923
31: 26: 27: 80: 85: 40: 48: 52. Posterior and anterior margins of
first dorsal abdominal segment broadly rounded. Metasternum
behind transverse suture shorter at middle.
Rhagovelia bisignata Bacon —
(Pl. 8, fig. 5; pl. 4, figs. 9a, b)
1948. Rhagovelia bisignata Bacon, Jour. Kansas Ent, Soc., vol. 21, no. 8, p. 71.
Winged female: Pronotum reaching the base of first dorsal ab-
dominal segment. Forewings yellowish brown, veins dark fuscous,
with one lower apical cell. Proportional length of dorsal abdominal
segments (second to eighth):: 39: 37: 40: 39: 41: 45: 42. First
dorsal abdominal segment with posterior margin obliterated in the
middle. Second segment with longitudinal carinae roundly pro-
duced laterally in the middle, usual three laevigate impressions
shallow and small, clearly separated from each other; intersegmen-
tal groove between second and third segments is represented by
a transverse row of shallow obscure impressions. Third dorsal
abdominal segment with longitudinal carinae obliterated, sometimes
invisible; anteromesal impression obliterated, anterolateral and
posterolateral impressions continuous and well marked; interseg-
mental groove between third and fourth segments is represented by
a transverse row of shallow impressions. Seventh dorsal abdominal
segment transverse, posterior margin rounded; seventh connexival
segment with lateral margin slightly sinuate. Eighth segment with
posterior margin substraight, rounded laterally.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 22: 35: 38: 40: 88: 41: 52: 50, Con-
nexivum more strongly reflexed than in male, apical end almost
reaching to the middle of eighth segment.
Wingless male: Proportional length of dorsal abdominal segments
(first to eighth):: 18: 22: 22: 28: 23: 28: 50: 40 (holotype). Pos-
terior margin of pronotum shallowly concave. Mesonotum long,
posterior margin almost reaching to anterior margin of first dorsal
abdominal segment. Metasternum behind transverse suture rela-
tively long, shorter at middle.
Rhagovelia callida Drake and Harris
1935. Rhagovelia callida Drake and Harris, Proc. Biol. Soc. Washington, vol.
48, p. 84
Wingless male: Posterior margin of pronotum roundly concave,
feebly produced posteriorly in the middle. Posterior margin of
mesonotum obscurely bisinuate. Proportional length of dorsal ab-
924 Tue University ScreNcE BULLETIN
dominal segments (first to seventh):: 23: 26: 26: 23: 24: 33: 60.
Anterior margin of first dorsal abdominal segment feebly sinuate.
Eighth segment dilated apically. Metasternum behind transverse
suture shorter at middle, inclined posteriorly.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 30: 83: 35: 88: 38: 46: 58: 50, Connexivum
subvertically reflexed.
Rhagovelia calopa Drake and Harris
(PL. 4, figs. 10a, b)
1927. ei owes calopa Drake and Harris, Proc. Biol. Soc, Washington, vol. 40,
p. 135.
Winged male: Pronotum reaching the basal region of second
dorsal abdominal segment. Forewings fuscous, with two closed
apical cells. Proportional length of dorsal abdominal segments
(second to eighth) :: $2: 25: 30: 32: 36: 60: 50, First dorsal abdomi-
nal segment with median elevated area straight on its posterior
margin. Second segment with longitudinal carinae slightly di-
vergent posteriorly; three laevigate impressions are connected by
well-sculptured depressions along posterior margin of first segment
and inner margin of second connexival segment; intersegmental
groove between second and third segments finely longitudinally
rugose. Third segment with longitudinal carinae finer than those
on the second segment, subject to individual variation in length,
obliterated behind middle in one male, almost reaching to posterior
margin of third segment in one female examined; three impressions
on the segment are connected in the same way as in second segment;
intersegmental groove between third and fourth segments obscurely
longitudinally rugulose.
Wingless male: Pronotum with posterior margin substraight. Mes-
onotum with posterior margin well defined and straight. Propor-
tional length of dorsal abdominal segments (first to seventh):: 33:
33: 81: 31: 30: 35: 55. Posterior margin of first dorsal abdominal
segment bisinuate, anterior margin of the same segment feebly pro-
duced. Metasternum behind transverse suture shortened medially
and inclined posteriorly.
Winged female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 20: 87: 33: 86: 40: 50: 52: 42. Connexivum
horizontally spread laterally. Seventh segment a little wider than
long, lateral margin subparallel; seventh connexival segment with
lateral margin straight. Eighth segment a little shorter than seventh
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 925
segment, posterolateral margin feebly sinuate, posterior margin sub-
straight.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 80: 80: 31: 34: 84: 38: 48: 33. Metaster-
num behind transverse suture as in male.
Rhagovelia deminuta Bacon
1948, Rhagovelia deminuta Bacon, Jour. Kansas Ent. Soc., vol. 21, no. 8, p. 72.
Wingless male: Pronotum with posterior margin bisinuate on
either side of the middle. Mesonotum with posterolateral margin
feebly sinuate. Posterior margin of metanotum shallowly concave.
Proportional length of dorsal abdominal segments (first to eighth) ::
14: 22: 20: 19: 22: 23: 39: 89 (holotype). Metasternum behind
transverse suture relatively long, slightly shortened medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 16: 23: 24: 23: 28: 82: 42: 40 (allotype).
Connexivum more strongly reflexed than in male; the last three seg-
ments reflexed at a little more than ninety degrees. Metasternum
behind transverse suture slightly shortened medially.
Rhagovelia evidis Bacon
1948. Rhagovelia evidis Bacon, Jour. Kansas Ent. Soc., vol. 21, no. 3, p. 73.
Wingless male: Posterior margin of pronotum broadly sinuate,
posterolateral margin feebly sinuate, posterior margin well defined
and feebly rounded. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 23: 19: 19: 20: 20: 25; 88: 28 (holotype).
Posterior margin of first dorsal abdominal segment broadly rounded.
Metasternum behind transverse suture feebly shortened medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 31: 21: 23: 25: 28: 35: 42: 35.
Remarks: In the male of this species and in both sexes of Rh.
deminuta the femur is equal in length to the tibia in the hind leg.
Rhagovelia fontanalis Bacon
(Pl. 4, figs. Lla, b)
1948. Rhagovelia fontanalis Bacon, Jour. Kansas Ent. Soc., vol. 21, no. 3, p. 74.
Winged male: Pronotum reaching to the basal region of second
abdominal segment. Forewing fuscous, veins black, with two dis-
tinct apical cells. Proportional length of dorsal abdominal segments
(first to seventh) :: 9: 87: 33: 83: 36: 40: 58. First dorsal abdominal
segment with posterior margin well-defined and roundly emargin-
926 Tue Universiry Scrence BULLETIN
ated in the middle. Second segment with longitudinal carinae
slightly roundly divergent posteriorly, reaching to posterior margin;
anteromesal impression rather obliterated, anterolateral and postero-
lateral impressions clearly separated; intersegmental groove be-
tween second and third segments rather shallow. Third dorsal
abdominal segment with longitudinal carinae subject to individual
variation ranging from basal one third to almost reaching posterior
margin of third segment; anteromesal impression obliterated, antero-
lateral and posterolateral impressions narrow and contiguous; inter-
segmental groove between third and fourth segments extends almost
to inner posterior angle of third connexival segment. Metasternum
behind transverse suture lengthened laterally.
Wingless male: Posterior margin of pronotum broadly concave.
Posterior margin of mesonotum obliterated. Proportional length of
dorsal abdominal segments (first to eighth) ::; 22: 30: 33: 33: 38: 43:
65: 45. Posterior margin of first to third dorsal abdominal segments
distinctly rounded, posterior margin. of eighth dorsal abdominal
segment sinuate. Metasternum behind transverse suture shortened
medially.
Winged female: Proportional length of dorsal abdominal seg-
ments (third to eighth):: 88: 38: 44: 47: 48: 43. Seventh dorsal
abdominal segment slightly widened posteriorly, posterior margin
sinuate; seventh connexival segment straightly narrowed apically,
apex with a bundle of black and long hairs. Eighth segment with
posterior margin rounded. Metasternum behind transverse suture
lengthened laterally.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh ):: 33: 35: 89: 89: 45: 51: 45. Connexivum
flattened out laterally.
Rhagovelia hambletoni Drake and Harris
1933. Rhagovelia hambletoni Drake and Harris, Proc. Biol. Soc, Washington,
vol, 46, p. 48. :
Wingless male: Pronotum with posterior margin broadly sinuate,
feebly produced posteriorly in the middle. Mesonotum with pos-
terior margin approaching to anterior margin of first dorsal abdom-
inal segment. Proportional length of dorsal abdominal segments
(first to eighth) :: 18: 23: 28: 24: 24: 31: 58: 89. Seventh connexival
segment not extending beyond posterior margin of seventh dorsal
abdominal segment. Metasternum behind transverse suture sub-
equal in length throughout, slightly lengthened near leg base.
Wingless female: Connexivum strongly reflexed and the ventral
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 927
side well-exposed above. Proportional length of dorsal abdominal
segments (first to eighth):: 20: 26: 80: 32: 85: 45: 58: 45 ( para-
type). Metasternum behind transverse suture as in male.
Rhagovelia imitatrix Bacon
(Pl. 5, figs. 12a, b)
1948. Rhagovelia imitatrix Bacon, Jour. Kansas Ent. Soc., vol. 21, no. 3, p. 76.
Winged female: Apex of pronotum reaching posterior margin of
first dorsal abdominal segment. Forewings greyish black, with a
lower rather large apical cell. Proportional length of dorsal abdom-
inal segments (first to eighth):: 15: 82: 42: 46: 41: 45: 50: 52.
Posterior margin of first dorsal abdominal segment well defined in
the middle. Second segment with longitudinal carinae obliterated
posteriorly; anteromesal impression shallow, anterolateral and pos-
terolateral impressions small, clearly separated from each other;
intersegmental groove between second and third segments shallow,
sparsely longitudinally rugose. Third segment without longitudinal
carinae; anteromesal impression shallow but distinct, anterolateral
and posterolateral impressions subcontiguous, intersegmental groove
between third and fourth segments almost as in the preceding one.
Seventh dorsal abdominal segment transverse, its posterior margin
rather deeply concave; seventh connexival segment with lateral
margin rounded. Eighth segment with posterior margin broadly
rounded. Metasternum behind transverse suture equal in length
throughout.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 29: 88: 40: 40; 41: 41: 51: 51 (allotype).
Metasternum behind transverse suture shortened medially.
Wingless male: Posterior margin of pronotum roundly concave,
feebly produced in the middle. Posterior margin of mesonotum
obliterated. Proportional length of dorsal abdominal segments
(first to seventh):: 18: 21: 24; 24: 25: 29: 53 (holotype). Meta-
sternum behind transverse suture shorter at middle, inclined pos-
teriorly.
Rhagovelia janeira Drake
(Pl. 6, figs. 19a, b)
1958, Rhagovelia janeira Drake, Proc. Biol. Soc. Washington, vol. 66, pp. 151-
152.
Winged male: Pronotum broadly rounded apically, reaching the
middle of second dorsal abdominal segment. Forewings fuscous,
lower apical cell is formed and relatively large. Proportional length
Tue Untiversiry SCIENCE BULLETIN
928
of dorsal abdominal segments (second to seventh) :: 25: 30: 81: 29:
32: 60. First dorsal abdominal segment with the median elevated
area well defined posteriorly. Second dorsal abdominal segment
with longitudinal carinae thick and rounded laterally; anteromesal
impression obliterated, anterolateral impression transverse and ob-
liquely placed, posterolateral impression small and clearly separated
from anterolateral impression; intersegmental groove between sec-
ond and third segments well impressed, its anterior margin zigzag.
Third segment with anteromesal impression well-impressed and
rounded, anterolateral and posterolateral impressions poorly devel-
oped and contiguous; longitudinal carinae obliterated before they
reach the middle of the segment; intersegmental groove between
third and fourth segments with anterior margin zigzag. Connexi-
vum horizontally spread laterally. Metasternum behind transverse
suture subequal in length throughout.
Wingless male: Posterior margin of pronotum concave. Meso-
notum with posterior margin obliterated. Dorsal abdominal seg-
ments straightly narrowed posteriorly. Proportional length of dorsal
abdominal segments (first to seventh):: 19: 28: 23: 23: 24: 28: 52
(holotype). Mesosternum behind transverse suture slightly length-
ened laterally.
Winged female: Seventh dorsal abdominal segment distinctly
longer than wide; seventh connexival segment with lateral margin
slightly sinuate at middle, narrowed posteriorly. Eighth segment
a little shorter than the preceding segment, posterior margin feebly
rounded,
Wingless female: Pronotum with apex narrowly rounded, much
more strongly produced posteriorly than in male. First dorsal ab-
dominal segment with anterior margin strongly elevated and strongly
produced in the middle, sinuate on either side of the middle. Con-
nexivum strongly reflexed, completely covering sixth to eighth dorsal
abdominal segments. Mesonotum behind transverse suture short,
subequal in length throughout.
Rhagovelia longipes Gould
(Pl. 5, figs. 12a, b)
1931. Rhagovelia longipes Gould, Univ. Kansas Sci. Bull., vol. 20, p. 35.
Pronotum with apex not reaching first dorsal abdominal segment.
Forewings fuscous, veins almost black, with one large apical cell.
Proportional length of dorsal abdominal segments (first to eighth) ::
15: 23; 22: 25; 23: 27: 48: 39. Connexivum rather strongly reflexed
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 929
laterally. First dorsal abdominal segment with median elevated
area defined laterally by a faint suture. Second dorsal abdominal
segment with longitudinal carinae strongly divergent posteriorly,
not reaching to posterior margin; anteromesal impression shallow,
anterolateral and posterolateral impressions small, contiguous by a
depression; intersegmental groove between second and third seg-
ments represented by a series of round and small impressions. Third
dorsal abdominal segment with faint carinae running obliquely,
not reaching to posterior margin; anteromesal impression obliter-
ated, anterolateral and posterolateral impressions are represented by
a contiguous laevigate, narrow impression along inner margin of
third connexival segment; intersegmental groove between third
and fourth segments well marked and with a series of small tu-
bercles. Posterior margin of seventh dorsal abdominal segment
strongly sinuate.
Wingless male: Posterior margin of mesonotum reaching an-
terior margin of first dorsal abdominal segment. Proportional length
of dorsal abdominal segments (first to seventh) :: 17(?): 18: 20: 20:
22; 29: 57.
Winged female: Seventh dorsal abdominal segment transverse,
slightly narrowed posteriorly, lateral margin sinuate; seventh con-
nexival segment with lateral margin rounded. Eighth segment with
posterior margin subtruncate. Mesosternum behind transverse su-
ture level, equal in length throughout, slightly lengthened near leg
base.
Wingless female: Posterior margin of mesonotum subtruncate, not
reaching to first dorsal abdominal segment. Proportional length of
dorsal abdominal segments (first to eighth) :: 25: 26: 30: 31: 31: 33:
41: 48.
Rhagovelia modesta Bacon
1956. Rhagovelia modesta Bacon, Univ. Kansas Sci, Bull., vol. 38, pt. I, p. 731.
Wingless male: Posterior margin of pronotum substraight, slightly
produced posteriorly in the middle, Proportional length of dorsal
abdominal segments (first to seventh):: 15: 15: 18: 22: 21: 27: 50.
Metasternum behind transverse suture equal in length throughout.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 21; 18: 28: 31: 36: 39: 58. Metasternum
behind transverse suture slightly lengthened medially.
Tue Universrry ScrENCE BULLETIN
Rhagovelia paulana Drake
(PI. 6, figs. 18a, b)
1953. en ihe paulana Drake, Proc. Biol. Soc. Washington, vol. 66, pp. 149-
150.
Winged female: Pronotum reaching posterior margin of first
dorsal abdominal segment. Forewings dark fuscous, forming a
small apical cell, and it is often obsolete. Proportional length of
dorsal abdominal segments (first to eighth) :: 20: 32: 80: 82: 29: 27:
46: 35. First dorsal abdominal segment with median elevated area
not well defined laterally, its posterior margin substraight. Second
dorsal abdominal segment with longitudinal carinae slightly di-
vergent posteriorly; anteromesal impression obliterated, antero-
lateral and posterolateral impressions rather small and clearly sepa-
rated from each other; intersegmental groove between second and
third segments distinctly impressed, posterior margin of the segment
on either side of intersegmental groove rounded. Third dorsal ab-
dominal segment without carinae, with a small anterolateral impres-
sion; intersegmental groove between third and fourth segments
distinct, trisinuate on its anterior margin. Seventh dorsal abdominal
segment a little longer than wide; seventh connexival segment
strongly sinuate in the middle, reaching almost to the middle of
eighth segment. Eighth segment distinctly wider at base than
seventh segment, posterior margin gently rounded. The last geni-
tal segment elongate, rounded apically. Metasternum behind trans-
verse suture subequal in length throughout.
Wingless male: Pronotum with posterior margin broadly sinuate.
Mesonotum almost reaching to posterior margin of metanotum.
Metanotum subequal in length throughout. Proportional length of
dorsal abdominal segments (first to seventh):: 21: 22: 22: 21: 21:
24: 47 (holotype). Lateral margin of eighth segment slightly
rounded.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 23: 31: 80: 27: 27: 27: 47: 40 (allotype).
Connexivum behind second segment strongly reflexed on the dor-
sum. Mesosternum behind transverse suture as in winged female.
Rhagovelia plana Drake and Harris
1933. Rhagovelia plana Drake and Harris, Proc, Biol. Soc. Washington, vol. 46,
p. 49.
Wingless male: Posterior margin of pronotum feebly concave,
posterolateral margin feebly sinuate. Mesonotum with posterior
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 931
margin obliterated. Abdomen substraightly narrowed apically, Pro-
portional length of dorsal abdominal segments (first to eighth)::
15: 20; 20: 25: 23: 29: 50 (paratype). Metasternum behind trans-
verse suture subequal in length throughout.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 20: 25: 26: 30: 33: 39: 45: 37. Connexivum
strongly reflexed throughout.
Rhagovelia plumbea Uhler
1894. Rhagovelia plumbea Uhler, Proc. Zool. Soc. London, p. 217.
Wingless male: Pronotum with posterior margin feebly trisinuate.
Mesonotum with posterior margin broadly rounded. Metanotum
broadly exposed behind mesonotum. Proportional length of dorsal
abdominal segments (first to eighth):: 20: 21: 20: 20: 20: 22:
39: 18. Oblique row of pubescence on mesosternum is lacking.
Metasternum behind transverse suture strongly shortened medially,
posterior margin of seventh ventral abdominal segment strongly
roundly emarginated.
Wingless female: Posterior margin of pronotum trisinuate. Meta-
notum well exposed behind mesonotum. Proportional length of
dorsal abdominal segments (first to eighth):: 28: 24: 24: 24: 28:
28: 87: 82. Connexivum strongly reflexed and folded on the dor-
sum, Metasternum behind transverse suture shortened medially
as in male,
Remarks: This species has the following peculiar characteristics:
(1) Posterior margin of pronotum trisinuate.
(2) Metanotum well exposed behind mesonotum.
(3) A paired oblique row of pubescence on mesosternum is
lacking.
(4) Metasternum behind transverse suture strongly shortened
medially.
Rhagovelia salina (Champion)
1898. Trochopus salinus Champion, Biol. Centr. Amer., Het., vol. 2, p. 140.
Wingless male: Posterior margin of pronotum substraight. Pos-
terior margin of mesonotum feebly sinuate leaving well exposed
metanotum behind. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 30: 80: 30: 30: 32: 37: 62: 42. Oblique
row of long pubescence on mesosternum lacking. Metasternum be-
hind transverse suture subequal in length throughout.
932, Tur Universiry ScrENCE BULLETIN
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 82: 85: 35: 35: 35: 43: 57: 47. Connexivum
more strongly reflexed than in male.
This species is easily distinguished from another marine form
Bh, plumbea Drake and Harris in the following points besides the
characteristics given by Bacon:
(1) Narrower body.
(2) Difference in proportional length in the seventh and eighth
dorsal abdominal segments.
(3) Difference in the metasternum behind transverse suture.
This species, however, shares a common characteristic with Rh.
plumbea in lacking the oblique row of long pubescence on the
mesosternum.
Rhagovelia spinosa Gould
(Pl. 5, figs. 14a, b)
1931. Rhagovelia spinosa Gould, Univ. Kansas Sci. Bull., vol. 20, p. 43.
Winged male: Pronotum reaching near posterior margin of sec-
ond segment, bluntly rounded apically. Forewings pale fuscous,
veins nearly black, with two small well-defined apical cells. Pro-
portional length of dorsal abdominal segments (second to seventh )::
98: 21: 23: 23: 28: 49. First dorsal abdominal segment with median
elevated area feebly bisinuate on posterior margin. Second dorsal
abdominal segment with longitudinal carinae roundly divergent
posteriorly; anteromesal and anterolateral impressions well-marked
and connected by the depression along posterior margin of first
segment, posterolateral impression small. Third dorsal abdominal
segment with longitudinal carinae less distinct than those on second
segment, but almost reaching to posterior margin of third segment;
anteromesal, anterolateral and posterolateral impressions contiguous
by distinct depression connecting them; intersegmental groove be-
tween second and third segments and that between third and fourth
segments are well marked, with a series of round impressions on
them. The last connexival segment with lateral margin slightly
sinuate.
Wingless male: Posterior margin of pronotum obliterated. Pro-
portional length of dorsal abdominal segments (first to seventh) ::
17: 20: 20: 20: 22: 22: 43 (holotype). Connexivum slightly reflexed.
Winged female: Seventh dorsal abdominal segment slightly nar-
rowed posteriorly; seventh connexival segment with apical margin
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 9383
sinuate. Eighth segment about as long as the preceding segment,
paratergite well defined and large.
Wingless female: Connexivum more strongly reflexed than in
male, but does not hide tergites beneath. Proportional length of
dorsal abdominal segments (first to eighth):: 21: 25: 26: 31: 35:
38: 52: 45. Metasternum behind transverse suture subequal in
length throughout.
Remarks: This species is characteristic among the representatives
of angustipes group in the following points:
(1) Male both in winged and wingless forms is armed with a
prominent tubercle at base of seventh ventral abdominal segment.
(2) Third dorsal abdominal segment in winged form occasionally
with distinct longitudinal carinae which almost reach to posterior
margin of the same segment.
(3) Spiracle on seventh segment placed distinctly closer to an-
terior margin than to posterior margin.
(4) Well-defined and large paratergite on eighth abdominal seg-
ment in winged female.
Rhagovelia tantilla Drake and Harris
(Pl. 6, figs: 17a, b)
1938. Rhagovelia tantilla Drake and Harris, Proc. Biol. Soc. Washington, vol.
46, p. 49.
Winged female: Pronotum reaching the middle of second dorsal
abdominal segment. Forewings fuscous, with one closed small
apical cell. Proportional length of dorsal abdominal segments (first
to eighth) :: 12: $1: 27: 27: 88: 87: 48: 51. Second dorsal abdominal
segment with longitudinal carinae not reaching to posterior margin
of the same segment; anteromesal impression obsolete, anterolateral
and posterolateral impressions are subcontiguous; intersegmental
groove inconspicuous, with a pair of transverse laevigate spots lo-
cated behind posterior end of longitudinal carina on intersegmental
groove. Third segment without longitudinal carinae, anterolateral
and posterolateral impressions continuous, inconspicuous. Con-
nexivum reflexed vertically. Eighth segment with posterior margin
broadly rounded. Metasternum behind transverse suture short,
slightly shortened medially.
Wingless male: Posterior margin of pronotum broadly concave.
Mesonotum fused with metanotum posteriorly. Proportional length
of dorsal abdominal segments (first to eighth) :: 24: 24: 25: 26; 33:
38: 48: 48. Metasternum behind transverse suture slightly short-
934 Tue Universitry ScreENCE BULLETIN
ened medially. First and second dorsal abdominal segments strongly
elevated, posterior margin of first segment broadly rounded,
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth ):: 28: 25: 25: 26: 35: 40: 45: 48. Connexivum
strongly reflexed. Metasternum behind transverse suture shortened
medially.
Rhagovelia tenuipes Champion
(Bh. 6; Age. Lb Ay b)
1898, Rhagovelia tenuipes Champion, Biol. Centr. Amer., Het. vol. 2, p. 137.
Winged male: Pronotum reaching first dorsal abdominal segment.
Forewings purplish black, apical cell not formed. Proportional
length of dorsal abdominal segments (first to eighth):: 15: 30: 26:
26: 27: 32: 51: 46. First dorsal abdominal segment with median
elevated area inclined posteriorly. Second dorsal abdominal seg-
ment with longitudinal carinae divergent posteriorly; anteromesal
impression shallow, anterolateral and posterolateral impressions
well marked; intersegmental groove between second and_ third
segments obscurely longitudinally rugose. Third segment with
longitudinal carinae poorly developed, not reaching the middle of
the segment; anteromesal impressions contiguous; intersegmental
groove between third and fourth segments well marked, evanescent
laterally. :
Wingless male: Pronotum with posterior margin broadly sinuate,
posterior margin of mesonotum indistinguishably fused with meta-
notum. Proportional length of dorsal abdominal segments (first
to seventh):: 19: 22: 22: 24: 25: 28: 50. Connexivum strongly re-
flexed. Metasternum as in winged male.
Winged female: Seventh dorsal abdominal segment longer than
eighth segment, Posterior margin of eighth segment broadly
rounded. Metasternum behind transverse suture short, slightly
lengthened near leg base.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 25: 30: 82: 34: 86: 42: 47: 48.
Rhagovelia velocis Drake and Harris
1935. Rhagovelia velocis Drake and Harris, Proc. Biol. Soc. Washington, vol.
A8, p. 86.
Wingless male: Pronotum with posterior margin broadly sinuate,
posterior margin of mesonotum indistinguishably fused with meta-
notum. Proportional length of dorsal abdominal segments (first
to seventh:: 25: 28: 24: 23: 25: 31: 53. Anterior margin of first
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 935
segment feebly sinuate, feebly produced in the middle. Metaster-
num behind transverse suture shortened in the middle,
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 29: 81: 82: 80: 32: 40: 52: 45. Metaster-
num as in wingless male.
Rhagovelia versuta Drake and Harris
(Pl. 3, figs. 8, 4; pl. 6, figs. 16a, b)
1935. Rhagovelia versuta Drake and Harris, Proc. Biol. Soc. Washington, vol.
48, p. 87.
Winged female: Pronotum bluntly rounded apically, covering
first dorsal abdominal segment beneath. Forewings black, with
purplish tinge, veins with conspicuous hairs, with one rather ob-
scurely defined apical cell, subcosta reaching the middle of upper
margin of forewing. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 22: 40: 40: 44: 46: 45: 61: 52. Second
dorsal abdominal segment with longitudinal carinae divergent and
narrowed posteriorly, with longitudinal rugosities behind posterior
margin of median elevated area of first dorsal abdominal segment,
anteromesal impression obsolete, anterolateral and posterolateral
impressions separated; intersegmental groove between second and
third segments with distinct longitudinal rugosities. Third segment
with longitudinal carinae narrower, traceable as far as middle of
the segment or sometimes almost reaching posterior margin of third
segment; anterolateral and posterolateral impressions contiguous;
intersegmental groove between third and fourth segments is repre-
sented by a series of round impressions, evanescent laterally. Seventh
connexival segment elongate subtriangular, with a bundle of long
black hairs at apex. Eighth segment with apex broadly rounded.
Metasternum behind transverse suture short,
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 30: 35; 33: 31; 82: 40: 70. Metasternum
as in winged female.
Winged male: Proportional length of dorsal abdominal segments
(third to seventh ):: 32: 84: 81: 87: 61. Connexivum reflexed in the
same degree as in female,
Wingless male: Posterior margin of pronotum broadly sinuate.
Posterior margin of mesonotum sinuate. Proportional length of
dorsal abdominal segments (first to seventh):: 25: 32: 30: 27: 29:
34: 64, Anterior margin of first dorsal abdominal segment feebly
bisinuate. Posterior margin of eighth segment broadly rounded.
Metasternum as in winged male.
936 Tur Universrry ScreNcE BULLETIN
Rhagovelia viriosa Bacon
1956. Rhagovelia viriosa Bacon, Univ. Kansas Sci. Bull., vol. 38, pt. I. p. 751.
Wingless male: Posterior margin of pronotum concave. Posterior
margin of mesonotum reaching to anterior margin of first dorsal
abdominal segment which is feebly sinuate. Proportional length of
dorsal abdominal segments (first to eighth) :: 80: 83: 33: 35: 86: 42:
80: 70. Metasternum behind transverse suture lengthened laterally
near leg base.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh) :: 80: 40: 40: 50: 53: 60: 75. Anterior margin
of first dorsal abdominal segment straight. Metasternum behind
transverse suture subequal in length throughout.
Group CHARACTERISTICS OF THE ABRUPTA Group
The following three species out of seven species treated by Bacon
were available for the study of winged forms: Rh. abrupta Gould,
Bh. trista Gould, Rh. vivata Bacon.
In all three species the distinct longitudinal carinae occur on the
second dorsal abdominal segment, the carinae on the third segment,
when present, are poorly developed (Rh. abrupta) or traceable as
a series of fine tubercles (Rh. trista). The forewing venation is
essentially the same as that in the angustipes group. Two apical cells
are always formed in the proximal half of the wing and the sub-
costa never develops beyond the basal half of the anterior margin
of the wing. The pronotum in winged forms is bluntly rounded
apically as in the angustipes group.
The proportional length of the sixth and seventh dorsal abdominal
segments in winged males ranges from about 2:3 to about 7:8 except
for Bh. lucida in which the third segment is much longer than the
second one (43:29).
In the intermediate leg the third tarsal segment is always longer
than the second one.
In the hind leg the femur is distinctly longer than the tibia ex-
cept for Rh. lucida in which the femur is just slightly longer than
the tibia (100:97).
The following characters are also peculiar to this group:
(1) Slender body except for Rh. lucida.
(2) Pronotum short, but longer than in the angustipes group, its
posterior margin always roundly produced posteriorly.
(3) The anterior margin of first dorsal abdominal segment
roundly produced anteriorly.
937
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA
(4) The seventh dorsal abdominal segment in female, in both
winged and wingless forms, is elongate except for Rh. lucida.
Rh. lucida apparently deviates strongly from other species of
this group in many characters, as noted above, and this will be fur-
ther noted in the description of this species.
The abrupta group is identical in the characters of the dorsal ab-
dominal segments and wing venation with the angustipes group,
but differs from it as follows:
(1) The pronotum in wingless forms is longer and its posterior
margin roundly produced posteriorly.
(2) The hind femur is longer than the hind tibia in all species.
(3) The metasternum behind the transverse suture is inclined
posteriorly.
Rhagovelia abrupta Gould
(Pl. 8, fig. 75 pl. 7, figs. 20a, b)
1933. Rhagovelia hungerfordi Gould, Ann. Ent. Soc. America, vol. 26, p. 467.
Winged female: Forewings totally purplish black, veins darker,
with two well-defined apical cells, of which the lower one is larger
than the upper one. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 16: 87: 42: 48: 56: 64; 75: 49. Longi-
tudinal carinae thick, roundly divergent posteriorly in second seg-
ment; anteromesal impression shallow, continuous with antero-
lateral impression by groove along posterior margin of first seg-
ment, posterolateral impression small, distinctly separated from
anterolateral impression; intersegmental groove between second
and third segments well defined, finely granulated. Third seg-
ment with longitudinal carinae obsolete behind the middle of the
segment; three laevigate impressions continuous by groove along
posterior margin of second dorsal abdominal segment and inner
margin of third connexival segment; intersegmental groove between
third and fourth segments well defined and laevigate. Seventh
dorsal abdominal segment with posterior margin straight; seventh
connexival segment with lateral margin straight, apex densely
clothed with dark and long hairs. Eighth segment with posterior
margin broadly rounded.
Wingless female: Proportional length of dorsal abdominal seg-
| ments (first to eighth):; 35: 39: 40: 45: 60: 71: 80: 50. Sixth,
seventh and eighth dorsal abdominal segments longer than wide.
Winged male: Proportional length of dorsal abdominal segments
(third to eighth):: 85: 35: 38: 45: 75: 95. Metasternum behind
transverse suture inclined posteriorly and shortened medially.
938 THE Universrry Science BULLETIN
Wingless male: Proportional length of dorsal abdominal segments
(first to eighth ):: 32: 35: 83: 82: 35: 41: 68: 80. Anterior margin of
first dorsal abdominal segment slightly roundly produced anteriorly.
Metasternum as in winged male.
Rhagovelia lucida Gould
1931. Rhagovelia lucida Gould, Univ. Kansas Sci. Bull., vol. 20, p. 36.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 38: 28: 28: 24: 25: 32: 65. First dorsal ab-
dominal segment rounded both on anterior and posterior margins.
Metasternum behind transverse suture inclined posteriorly, slightly
shortened medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 50: 30: 80: 26: 23: 35: 65:57. First dorsal
abdominal segment roundly produced anteriorly. Seventh segment
inclined posteriorly. Connexivum subvertically erected. Meta-
sternum behind transverse suture level, slightly shortened medially.
Remarks: Abnormally long first dorsal abdominal segment, con-
siderably deviated proportional length of sixth and seventh dorsal
abdominal segments in both sexes, hooklike apex of clasper, much
deviated proportional length between hind femur and tibia, much
longer third antennal segment than second segment are character-
istic of this species which breaks close relationship with any other
representative of this group.
Rhagovelia torquata Bacon
1948. Rhagovelia torquata Bacon, Jour. Kansas Ent. Soc., vol, 21, no. 3, p. 83.
Wingless male: Proportional length of dorsal abdominal segments
(first to eighth) :: 28: 30: 30: 30: 32: 38: 50: 45 (holotype). First
dorsal abdominal segment produced anteriorly,
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 34: 87: 89: 88: 43: 57: 65: 42. Metaster-
num behind transverse suture inclined posteriorly, shortened me-
dially.
Rhagovelia trepida Bacon
1948. Rhagovelia trepida Bacon, Jour. Kansas Ent. Soc., vol. 21, no. 8, p. 84.
Wingless male: Proportional length of dorsal abdominal segments
(first to eighth) :: 86: 88: 83: 32: 32: 82: 88: 65 (holotype). Meta-
sternum behind transverse suture inclined posteriorly, slightly short-
ened medially. All ventral abdominal segments are longitudinally
elevated in the middle,
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 989
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 87: 40: 85: 87: 50: 38: 60: 42 (allotype).
Metasternum behind transverse suture not inclined posteriorly. Last
three dorsal abdominal segments as in Rh. trista. Seventh dorsal
abdominal segment elongate, narrowed apically. Eighth segment
with posterior margin broadly rounded.
Rhagovelia trista Gould
(Pl. 7, figs. 22a, b)
1931. Rhagovelia trista Gould, Univ. Kansas Sci. Bull., vol. 20, p. 45.
Winged female: Forewings greyish fuscous, with two well-defined
apical cells. Proportional length of dorsal abdominal segments
(second to eighth):: 87: 32: 34: 40: 47: 52: 40. First dorsal ab-
dominal segment with median elevated area roundly sinuate on pos-
terior margin. Second segment with longitudinal carinae narrow
and rounded; anteromesal impression obliterated, anterolateral and
posterolateral impressions clearly separated; intersegmental groove
between second and third segments without rugosity. Third seg-
ment with a faint series of fine tubercles disposed obliquely behind
posterior end of longitudinal carinae on second segment, antero-
lateral and posterolateral impressions contiguous. Seventh dorsal
abdominal segment longer than wide, widened posteriorly, longer
than eighth segment; seventh connexival segment with lateral margin
straight. Eighth segment with posterior margin broadly rounded.
Metasternum behind transverse suture inclined posteriorly, subequal
in length throughout.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 30: 83: 83: 36: 42: 52; 58: 45 (allotype).
Anterior margin of first dorsal abdominal segment substraight.
Apical three segments much as in winged form, Connexivum sub-
vertically erected. Metasternum behind transverse suture as in
winged female. .
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh) :: 28: 80: 27: 27: 30: 38: 45 (paratype). Meta-
sternum behind transverse suture inclined posteriorly, slightly short-
ened in the middle.
Rhagovelia vivata Bacon
(Pl. 7, figs. 21a, b)
1948, Rhagovelia vivata Bacon, Jour. Kansas Ent. Soc., Vol. Bly no: 8; 9.) 86.
Winged male: Forewings fuscous, veins darker, with two distinct
apical cells as in Rh. abrupta. Proportional length of dorsal abdomi-
nal segments (second to eighth ):: 42: 38: 37: 47: 52: 65: 47. Second
940 Tue Universiry Science BULLETIN
dorsal abdominal segment with longitudinal carinae narrow, reach-
ing posterior margin of second segment; anteromesal impression
obliterated, narrowly impressed along posterior margin of first seg-
ment on either side of carinae, anterolateral and posterolateral im-
pressions well sculptured and subcontiguous; intersegmental groove
between second and third segments finely granulated. Third seg-
ment without carinae. Anterolateral and posterolateral impressions
smaller and continuous; intersegmental groove between third and
fourth segments finely granulated. Metasternum behind transverse
suture inclined posteriorly, shortened medially. Seventh ventral
abdominal segment about as long as two preceding segments com-
bined.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh) :: 31: 34: 33: 28: 35: 41: 63. Metasternum behind
transverse suture inclined posteriorly, shortened medially.
Winged female: Seventh dorsal abdominal segment with anterior
margin rounded, posterior margin sinuate, about 14 times as long
as eighth segment (60:42), slightly narrowed apically; seventh con-
nexival segment with lateral margin straight. Eighth segment with
posterolateral margin feebly rounded.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 31: 33: 35: 37: 47: 60: 70: 47 (allotype).
First dorsal abdominal segment straight or feebly produced.
Group CHARACTERISTICS OF THE Hirriprs Group
The only representative of this group Rh. hirtipes Drake and
Harris has the following peculiar characters which are not shared by
any one group of the genus Rhagovelia:
(1) Longitudinal carinae occur only in the second dorsal ab-
dominal segment and reach to the middle of the segment.
(2) The median elevated area of the first dorsal abdominal seg-
ment is obscurely defined laterally.
(3) The last genital segment is bent back and completely tele-
scoped on the ventral surface of the eighth abdominal segment.
(4) The seventh dorsal abdominal segment is twice as long as the
eighth segment in the wingless female.
(5) The third and fourth dorsal abdominal segments in the
winged female is strongly depressed laterally.
(6) The dorsum is longitudinally carinate in the middle after
the first two segments in the female.
(7) The connexivum after the fifth segment is strongly reflexed,
narrowed and lustrous.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 941
(8) The dorsal surface of the abdomen in winged forms is much
the same as in the wingless form in both sexes.
(9) The shape of the male clasper is much different from that of
any other species of this genus.
This species, however, has the following characters in common
with the majority of the angustipes group:
(1) The second tarsal segment is about as long as the third one
in the intermediate leg.
(2) The femur is shorter than the tibia in the hind leg.
(3) The metasternum behind the transverse suture is level.
Rhagovelia hirtipes Drake and Harris
(Pl. 7, figs. 23a, b)
1927. Rhagovelia hirtipes Drake and Harris, Proc. Biol. Soc. Washington, vol.
AO, p. 136.
Winged female: Pronotum with apex rounded, clothed with long,
dark brown hairs. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 27: 39: 40: 89: 42: 45: 63: 43, First seg-
ment with median elevated area rather narrow, laterally obliquely
defined. Second segment with longitudinal carina thick, reaching
just to the middle of second segment; anteromesal impression is
represented by a small, round impression; anterolateral and postero-
lateral impressions contiguous along inner margin of second con-
nexival segment. Third and fourth segments strongly and widely
depressed on either side of median longitudinal axis. Seventh seg-
ment much longer than eighth segment (63:43), posterior margin
feebly sinuate. Eighth segment with posterior margin rounded
and armed with two pairs of conspicuous bundles of long, black
hairs directed more or less downwardly. Connexivum in basal
five segments oblique, then strongly reflexed, narrowed and _ pol-
ished posteriorly, clothed with long brown hairs. Seventh con-
nexival segment near apex with rather conspicuous bundle of hairs.
The last genital segment turned completely downwards and folded
on the ventral surface of eighth abdominal segment. Metasternum
behind transverse suture level, subequal in length throughout,
slightly shorter in the middle.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 53: 42: 87: 44: 49: 61: 73: 387. Second
dorsal abdominal segment with posterior region subvertically in-
clined posteriorly. Third and fourth segments depressed laterally
as in winged form. Connexivum vertically reflexed after fourth seg-
THe Universiry Science BULLETIN
942
ment. Posterior margin of eighth segment densely clothed with pale
brown hairs. The last segment as in winged form.
Winged male: Proportional length of dorsal abdominal segments
(first to seventh):: 19: 40: 40: 34: 39: 43: 63. Connexivum less
strongly reflexed than in female. Third and fourth dorsal abdominal
segments without conspicuous lateral depression as in female.
Metasternum behind transverse suture inclined posteriorly, strongly
shortened medially. Seventh ventral abdominal segment shallowly
depressed on either side of middle, posterior margin nearly straight.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 28: 89: 87: 85: 37: 40: 60, posterior margin of
each segment except for sixth and seventh segments bisinuate.
Metasternum and ventral side of abdomen as in winged male.
Group CHARACTERISTICS OF THE ELEGANS Group
The following three species of this group were available for the
study of winged forms: Rh. insularis Champion; Rh. elegans Uhler;
Rh. uncinata Champion.
Pronotum in winged forms without apical process, with obscure
longitudinal ridge throughout the entire length. Pronotum in wing-
less forms broadly rounded apically, sometimes not completely cov-
ering the mesonotum beneath (Rh. insularis).
Forewings with two well-developed apical cells formed at ib
basal region of distal half of the wing.
Longitudinal carinae occur always on the second and third seg-
ments, continuous and fine; intersegmental groove finely and longi-
tudinally rugulose.
The following characters are also peculiar to this species group:
(1) The species is of large size.
(2) The winged forms are relatively abundant.
(3) The dorsum of the abdomen of the wingless female tapers
rather evenly to the apex.
(4) The anterior margin of the first dorsal abdominal segment is
roundly produced,
(5) The metasternum behind the transverse suture is inclined
posteriorly, the transverse suture being more or less sinuate on either
side of the middle.
(6) The anterior margin of metasternum always slightly pro-
duced anteriorly,
(7) The proportional length between the sixth and seventh ab-
dominal segments in winged males range from 53:67 in Rh. insularis
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 943
to 55:67 in Bh. uncinata; that in wingless males from 53:73 in Rh.
elegans, to 53:67 in Rh. insularis. The proportional length between
the seventh and eighth segments in winged females range from
55:45 in Rh. insularis, to 63:51 in Rh, uncinata, that in wingless fe-
males from 62:50 in Rh. elegans and Rh. uncinata, to 43:48 in Rh.
insularis.
(8) The seventh connexival segment with inner margin is always
produced inwardly in the middle, its lateral margin rounded in the
winged female.
(9) In the intermediate leg the third tarsal segment is distinctly
longer than the second one.
(10) In the intermediate leg the tibia is more than twice as long
as the second tarsal segment.
(11) In the hind leg the femur and the tibia are subequal in
length.
Rhagovelia elegans Uhler
(PL. 8, figs. 25a, b)
1894, Rhagovelia elegans Uhler, Proc. Zool. Soc. London, p. 216.
Winged male: Forewings almost black, with two large apical
cells. Proportional length of dorsal abdominal segments (first to
seventh) :: 23: 53: 57: 62: 60: 67: 80. First segment emarginated
on posterior margin. Second segment with median longitudinal
carinae thin, roundly produced laterally at middle, continuous with
those on third segment; anteromesal impression continuous with
anterolateral impression by depression along posterior margin of
first segment, anterolateral and posterolateral impressions separated
but connected by depression along inner margin of second con-
nexival segment; intersegmental groove between second and third
segments finely and longitudinally rugulose, its anterior margin
produced. Third segment with longitudinal carinae slightly di-
vergent posteriorly; anteromesal, anterolateral and posterolateral im-
pressions continuous; intersegmental groove between third and
fourth segments finely longitudinally rugulose, its anterior margin
slightly produced in the middle. Metasternum behind transverse
suture as in Rh. insularis.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh) :: 43: 50: 50: 52: 53: 53: 78,
Winged female: Seventh dorsal abdominal segment about as long
as wide, posterior margin slightly sinuate; seventh connexival seg-
ment with inner margin rather strongly produced inwardly, lateral
944 Tue Universiry Science BULLETIN
margin rounded, sparsely clothed with long hairs. Eighth segment
with posterior margin broadly rounded.
Wingless female: Seventh dorsal abdominal segment about as
long as wide, posterior margin slightly sinuate; seventh connexival
segment with inner margin rather strongly produced inwardly, lat-
eral margin rounded, sparsely clothed with long hairs. Eighth seg-
ment with posterior margin broadly rounded.
This species is closely related to Rh. insularis, but may be distin-
guished from the latter in the following points in addition to the
characters given by Bacon:
(1) All spiracles from third to seventh segments in this species
are placed midway between anterior and posterior margins.
(2) First dorsal abdominal segment with median elevated area
longer in this species.
Rhagovelia insularis Champion
(Pls. 1, 2; pl. 8, figs. 1, 2; pl. 8, figs. 27a, b)
1898. Rhagovelia insularis Champion, Biol. Centr. Amer., Het., vol. 2, p. 185.
Winged male: Pronotum with apex reaching basal region of sec-
ond dorsal abdominal segment, narrowly rounded. Forewings dark
fuscous, veins black, with two distinct apical cells formed beyond
the middle of the wing. Proportional length of dorsal abdominal
segments (first to eighth):: 13: 51: 51: 51: 51: 58: 67: 57. First
dorsal abdominal segment short, with median elevated region
roundly concave on posterior margin. Second segment with longi-
tudinal carinae narrow, rather strongly produced laterally in apical
half; anteromesal impression obsolete, anterolateral and postero-
lateral impressions separate from each other; intersegmental groove
between second and third segments finely rugulose. Third segment
with longitudinal carinae less strongly divergent posteriorly than
in second segment; anteromesal impression distinct, anterolateral
and posterolateral impressions are continuous; intersegmental
groove between third and fourth segments finely longitudinally
rugulose. Posterior margin of seventh segment nearly straight.
Connexivum rather flattened, lateral margin of each segment more
or less rounded. Metasternum behind transverse suture strongly
inclined posteriorly and shortened in the middle, with a distinct
longitudinal elevation in the middle.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh) :: 42: 48: 48: 48: 48: 53: 65, First dorsal abdominal
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 945
segment roundly produced anteriorly. Metasternum as in winged
male.
Winged female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 17: 57: 58: 56: 55: 58: 55: 45. Seventh
dorsal abdominal segment with lateral margin sinuate in posterior
half, posterior margin strongly sinuate; seventh connexival segment
with lateral margin feebly rounded. Eighth segment with postero-
lateral margin feebly rounded, posterior margin feebly sinuate. Con-
nexivum reflexed. Metasternum behind transverse suture strongly
inclined posteriorly, strongly and straightly shortened medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 42: 43: 50: 50: 48: 50: 53: 48. Pronotum
not reaching posterior margin of mesonotum.
Rhagovelia uncinata Champion
(Pl. 8, figs. 26a, b)
1898. Rhagovelia uncinata Champion, Biol. Centr. Amer., Het., VOho2, Pi 1Bo,
Winged male: Forewings dark fuscous, veins black, with two
well-defined apical cells. Proportional length of dorsal abdominal
segments (first to seventh):: 20: 51: 48: 55: 51: 55: 63, Second
segment with longitudinal carinae slender, gently roundly produced
laterally in the middle; anteromesal impression small but distinct,
anteromesal and anterolateral impressions connected by depression
along posterior margin of first segment, posterolateral impression
clearly separated from anterolateral impression, oval in shape; in-
tersegmental groove finely longitudinally rugulose; disk obscurely
transverse-obliquely rugose. Third segment with longitudinal cari-
hae continuous with those on second segment, fine, gently roundly
produced laterally in the middle; anteromesal impression distinct;
anterolateral and posterolateral impressions subcontiguous; inter-
segmental groove finely longitudinally rugulose; disk obscurely
transversely rugose. Metasternum with transverse suture slightly
sinuate on either side of the middle; the area posterior to transverse
suture strongly inclined posteriorly, its caudal margins straight,
oblique and meeting at an angle medially. All spiracles from third
to seventh segments located midway between anterior and posterior
margin of each segment.
Winged female: Proportional length of dorsal abdominal seg-
ments (third to eighth):: 50: 56: 56: 63; 63: 51. Seventh dorsal
abdominal segment widened posteriorly, lateral margin sinuate
31-3378
946 Tae University Science BULLETIN
at middle; seventh connexival segment with lateral margin rounded.
Eighth segment with posterolateral margin sinuate, posterior mar-
gin slightly rounded. Lateral margin of seventh and eighth segments
sparsely clothed with long hairs.
Wingless female: Pronotum almost covers mesonotum beneath.
Proportional length of dorsal abdominal segments (first to eighth) ::
40: 47: 48: 50: 52: 57: 62: 50. First dorsal abdominal segment with
anterior margin roundly produced anteriorly. Connexivum strongly
reflexed. Metasternum behind transverse suture strongly inclined
posteriorly, strongly shortened medially.
Rhagovelia merga Bacon
1956. Rhagovelia merga Bacon, Univ. Kansas Sci, Bull., vol. 38, pt. I, p. 774.
Wingless male: Proportional length of dorsal abdominal segments
(first to sixth):: 40: 58: 50: 49: 49: 55 (holotype). First dorsal ab-
dominal segment with anterior margin nearly straight.
Group CHARACTERISTICS OF THE CRASSIPES GROUP
The following species were available for the study of winged
forms:
Rh. crassipes Champion Rh. palea Bacon
Rh. horrida Bacon Rh. perfidiosa Bacon
Rh. jubata Bacon ° Rh. sinuata Gould
Rh. nitida Bacon Rh, varipes Champion
In this group Rh. nitida deviates considerably in some characters
from other species of this group defined by Bacon, as will be noted
in connection with the description of this species. When this spe-
cies is excluded the group becomes increasingly homogeneous as is
characterized below.
Pronotum in winged form without apical projection except for the
female of Rh. crassipes. Pronotum entirely covers mesonotum in
wingless forms.
Forewings always with two large apical cells formed in distal
half of the wing.
Longitudinal carinae fine but well defined, always occur on sec-
ond and third segments; anterolateral and posterolateral impres-
sions are always clearly separated.
Connexivum never folded on the dorsum in wingless forms. Sev-
enth dorsal abdominal segment about as long as wide or wider than
long; seventh connexival segment gradually narrowed apically,
never strongly produced posteriorly. Proportional length of sixth
SUPPLEMENTARY STUDY OF THE Genus RHAGOVELIA 947
and seventh dorsal abdominal segments in winged male ranges
from 35:55 in Rh. perfidiosa to 78:85 in Rh. crassipes, that in wing-
less male from 30:52 in Rh. perfidiosa to 58:65 in Rh. williamsi.
Proportional length of seventh and eighth segments in winged fe-
male ranges from 87:85 in Rh. varipes to 78: 60 in Rh. crassipes,
that in wingless female from 85:70 in Rh. varipes to 48:70 in Rh.
williamsi.
In intermediate leg third tarsal segment is always longer than
second one; tibia is always more than twice as long as second tarsal
Segment,
In hind leg femur always longer than tibia.
Rhagovelia crassipes Champion
(Pl. 9, figs. 28a, b, c)
1898. Rhagovelia crassipes Champion, Biol. Centr. Amer. Het., vol. 2, p. 183.
Winged female: Pronotum with apex erected, acutely pointed,
with a small process at base of apical process. Forewings fuscous,
veins darker, with two well-defined elongate cells in distal half.
Proportional length of dorsal abdominal segments (first to eighth )::
25; 55; 70: 78: 75: 82: 78: 60. First dorsal abdominal segment with
median elevated area feebly sinuate on posterior margin. Second
segment with longtitudinal carinae slightly divergent posteriorly,
forked into two arms on intersegmental groove, embracing partly
a shallow, round depression, obscurely depressed along posterior
margin of first segment, anteromesal impression small, anterolateral
impression transverse, posterolateral impression placed in the de-
pression, clearly separated from anterolateral impression; inter-
segmental groove shallow, ill-defined. Third segment with longi-
tudinal carinae arising from inner apical arm of the carinae on
second abdominal segment, forked posteriorly, inner branch is more
conspicuous; intersegmental groove between third and fourth seg-
ments shallow; anteromesal laevigate impression absent, antero-
lateral and posterolateral impressions shallow, placed in obscurely
defined depression along third connexival segment. Seventh dor-
sal abdominal segment a little wider than long, subquadrangular
in shape; seventh connexival segment gradually narrowed apically,
inner margin sinuate, lateral margin rounded and densely clothed
with brown hairs which become longer posteriorly. Eighth seg-
ment with paratergite clothed with long hairs on dorsal surface;
Posterior margin of eighth segment broadly rounded. Posterior
margin of seventh and eighth ventral abdominal segments with
948 Tue Universiry ScreENCE BULLETIN
small black tubercles at sides. The last genital segment gradually
narrowed apically, apex narrowly rounded. Metasternum behind
transverse suture without distinct median longitudinal elevation,
inclined posteriorly, longer laterally near leg base.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth ):: 52: 60: 70: 70: 75: 80: 80: 60. Connexivum
a little more strongly reflexed than in winged male. Abdomen on
ventral side as in winged female.
Winged male: Pronotum with the apex less strongly produced pos-
teriorly. Proportional length of dorsal abdominal segments (first
to seventh):; 25: 55: 68: 73: 72: 75: 85. Seventh dorsal abdominal
segment strongly widened posteriorly, lateral margin sinuate. Ven-
tral abdominal segment with median longitudinal elevation. Meta-
sternum behind transverse suture strongly inclined posteriorly,
strongly shortened in the middle. Dorsolateral and ventrolateral
margins of seventh abdominal segment with a series of minute black
spines directed inwardly.
Wingless male: Pronotum completely covers mesonotum. Pro-
portional length of dorsal abdominal segments (first to seventh):
58: 73: 73: 72: 71: 80: 95. Anterior margin of first dorsal abdominal
segment roundly produced anteriorly. Abdomen on ventral side
as in winged female.
Rhagovelia amazonensis Gould
1931. Rhagovelia amazonensis Gould, Univ. Kansas Sci. Bull., vol. 20, p. 15.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh) :: 31: 87: 35: 30; 34: 37: 56 (holotype ). Pronotum
entirely covering mesonotum. Connexivum reflexed laterally at an
angle of about forty-five degrees. Metasternum behind transverse
suture inclined posteriorly and shorter at middle. Without median
longitudinal ridge on venter. Seventh and eighth ventral abdominal
segments without black spines.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 35: 87: 40; 88: 48: 47: 55: 48 (allotype).
Anterior margin of first dorsal abdominal segment broadly rounded
and produced anteriorly. Apex of seventh connexival segment
and lateral margin of eighth dorsal abdominal segments with thick
bundle of black setae. Metasternum as in wingless male.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 949
Rhagovelia castanea Gould
1931. Rhagovelia castanea Gould, Univ. Kansas Sci. Bull., vol. 20, p. 19.
Wingless male: Pronotum entirely covers mesonotum. Propor-
tional length of dorsal abdominal segments (first to seventh):: 44:
46: 41: 40: 43: 48: 76 (holotype). Connexivum slightly reflexed
laterally. Metasternum behind transverse suture shorter at middle.
Venter without median longitudinal carina. Seventh and eighth ven-
tral abdominal segments without a row of black spines on posterior
margin.
Wingless female: Pronotum as in wingless male. Proportional
length of dorsal abdominal segments (first to eighth):: 45: 43: 41:
43: 49: 59: 72: 59 (allotype). Eighth segment with posterior mar-
gin broadly rounded.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth) :: 52: 60: 70: 70: 75: 80: 80: 60. Connexivum
a little more strongly reflexed than in winged male. Abdomen on
ventral side as in winged female.
Winged male: Pronotum with apex less strongly produced pos-
teriorly. Proportional length of dorsal abdominal segments (first
to seventh):: 25: 55: 68: 73: 72: 75: 85. Seventh dorsal abdominal
segment strongly widened posteriorly, lateral margin sinuate. Ven-
tral abdominal segments with median longitudinal elevation. Meta-
sternum behind transverse suture strongly inclined posteriorly,
strongly shortened in the middle. Dorsolateral and ventrolateral
margins of seventh abdominal segment with a series of minute black
spines directed inwardly.
Wingless male: Pronotum completely covers mesonotum. Pro-
portional length of dorsal abdominal segments (first to seventh)::
58: 73: 73: 72: 71: 80: 95. Anterior margin of first dorsal abdominal
segment roundly produced anteriorly. Abdomen on ventral side
as in winged female.
Rhagovelia femoralis Champion
1898. Rhagovelia femoralis Champion, Biol. Centr. Amer., Het., vol. 2, p, 188.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 40; 48: 46: 44: 45: 51: 80. Connexivum
reflexed at angle of forty-five degrees laterally. Venter with distinct
median longitudinal elevation clothed with long hairs throughout
second to sixth segments. Seventh ventral abdominal segment
without row of black spines on posterior margin.
Tue Universiry ScreNCE BULLETIN
Rhagovelia horrida Bacon
(Pl. 10, figs. 88a, b)
1956. Rhagovelia horrida Bacon, Univ. Kansas Sci. Bull., vol. 38, pt. I, p. 787.
Winged female: Pronotum with apex acute, reaching first dorsal
abdominal segment. Proportional length of dorsal abdominal seg-
ments (first to eighth):: 25: 50: 58: 59: 59: 61: 61: 51. First dorsal
abdominal segment with posterior margin shallowly concave. Sec-
ond dorsal abdominal segment with longitudinal carinae well
defined, slightly divergent and dilated posteriorly; anteromesal
impression obliterated; anterolateral and posterolateral impressions
clearly separated from each other; intersegmental groove between
second and third segments long and ill defined posteriorly, longi-
tudinally rugose. Third segment with longitudinal carinae extend
into intersegmental groove, with a shallow impression on either
side of longitudinal carinae at apical end on each segment; inter-
segmental groove between third and fourth segments shorter than
that between second and third segments, longitudinally rugose;
disk on either side of longitudinal carinae obscurely transversely
rugose. Seventh dorsal abdominal segment wider at base than
long; seventh connexival segment with lateral margin rounded;
seventh ventral abdominal segment with a row of black spines
directed inwardly on posterior margin. Eighth with posterolateral
margin broadly rounded, ventral side with a mass of black tubercles
at sides. Metasternum behind transverse suture shortened medially.
Wingless female: Pronotum. completely covers mesonotum,
broadly rounded apically. Proportional length of dorsal abdominal
segments (first to eighth):: 40; 49: 52: 54: 58: 62: 64: 53 (allo-
type). Anterior margin of first dorsal abdominal segment sinuate
on either side of nearly straight and produced median area.
Winged male: Pronotum as in winged female. Proportional
length of dorsal abdominal segments (first to seventh):: 25: 52:
55: 56: 56: 59: 74. Seventh dorsal abdominal segment simply
widened posteriorly; lateral margin of seventh connexival segment
with long hairs. Ventrolateral and dorsolateral margins of seventh
and eighth abdominal segments with a row of black spines.
Wingless male: Pronotum completely covers mesonotum. Pro-
portional length of dorsal abdominal segments (first to seventh)::
88: 47: 47: 43: 44: 49: 71 (holotype). Metasternum behind trans-
verse suture strongly shortened medially, with distinctly elevated
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 951
median longitudinal carina which extends down to sixth ventral
abdominal segment. Dorsolateral and ventrolateral margins of
seventh and eighth segments as in winged male.
Rhagovelia jubata Bacon
1948. Rhagovelia jubata Bacon, Jour. Kansas Ent. Soc., vol. 21, p. 78.
Winged female: Pronotum broadly rounded apically. Forewings
with two basal cells white at base, then changes into orange, wing
veins fuscous. With distinct longitudinal carinae on second and
third dorsal abdominal segments. Seventh connexival segment
narrowed posteriorly as in Rh. horrida or Rh. crassipes, with dark
brown hairs near apex.
Since only the allomorphotype is winged the structure beneath
the wing was not observed carefully. However, distinct carinae
on second and third dorsal abdominal segments were clearly seen
by lifting the wing.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 50: 46: 45: 43: 51: 54: 62: 82. Anterior
margin of first dorsal abdominal segment rather strongly roundly
produced.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 45: 48: 43: 44: 46: 50: 70. Connexivum
rather strongly reflexed laterally. Venter with distinct longitudinal
carina throughout metasternum behind transverse suture to sixth
segment. Seventh ventral abdominal segment with minute black
spines on posterolateral angle.
Rhagovelia nitida Bacon
(Pl. 10, figs. 35a, b)
1948. Rhagovelia nitida Bacon, Jour. Kansas Ent. Soc., vol, 21, p. 79.
Winged female: Pronotum at apex obtusely rounded, reaching
the middle of second abdominal segment. Forewings nearly black,
with two large apical cells. Proportional length of dorsal abdom-
inal segments (first to eighth):: 22: 53: 50: 60: 63: 68: 73: 38.
First dorsal abdominal segment with posterior margin broadly
concave. Second dorsal abdominal segment with longitudinal
carinae narrow but well-defined, rounded near base, usual three
laevigate impressions clearly separated from each other, located on
the depression continuous throughout along posterior margin of
first segment and inner margin of second connexival segment;
intersegmental groove short but well-defined, longitudinally ru-
952
Tue Universiry Science BULLETIN
gose. Third segment with longitudinal carinae narrow but well-
defined, divergent posteriorly, anteromesal impression small,
depressed along posterior margin of third segment, anterolateral
and posterolateral impressions subcontiguous; intersegmental
groove between third and fourth segments short but well defined.
Seventh dorsal abdominal segment elongate, narrowed apically,
posterior margin sinuate; seventh connexival segment with lateral
margin gently rounded and densely clothed with rather short hairs.
Eighth segment about half as long as seventh segment on dorsal
side, posterior margin nearly straight, with short hairs throughout.
The last segment bent downward and invisible from above.
Metasternum behind transverse suture inclined posteriorly and
shortened medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 42: 42: 47: 51: 58: 60: 74: 48 (allotype).
Ventral side of abdomen without median longitudinal carina. Con-
nexivum feebly reflexed laterally.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 46: 44: 43: 44: 47: 49: 75 (holotype).
Connexivum almost horizontally spread laterally. First dorsal ab-
dominal segment roundly produced anteriorly. Metasternum be-
hind transverse suture almost vertically inclined posteriorly. Basal
ventral abdominal segments with distinct median longitudinal
carina, Seventh and eighth ventral abdominal segments without
a row of black spines on posterior margins.
This species is characterized by the following points:
(1) The last genital segment is bent downwardly and placed
on the ventral surface of the eighth segment.
(2) The seventh dorsal abdominal segment almost twice as long
as the eighth segment in winged females.
(3) The shape of the male clasper is much different from that
of other species of the crassipes group.
(4) In the hind leg, the tibia is distinctly longer than the femur.
Rhagovelia ornata Bacon
1948. Rhagovelia ornata Bacon, Jour. Kansas Ent. Soc., vol. 21, p. 80.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh ):: 58: 62: 62: 62: 65: 69: 100. Metasternum
behind transverse suture strongly shortened in the middle, inclined
posteriorly, with median longitudinal elevation which extends down
to basal ventral abdominal segments. Seventh ventral abdominal
segment with posterior margin without black spines.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 953
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 60: 65: 72: 75: 80: 87: 97: 65 (paratype).
Metasternum as in wingless male. Median longitudinal elevation
on venter less pronounced than in male. Connexivum almost
vertically reflexed.
Rhagovelia palea Bacon
(Pl. 9,-figs,, Sla,b)
1956. Rhagovelia palea Bacon, Univ. Kansas Sci. Bull., vol. 88, pt. I, p. 795.
Winged female: Pronotum with apex bluntly rounded, apical
region sparsely clothed with long hairs. Forewings pale brown,
veins white, with a wide white stripe on upper basal cell, with two
large closed apical cells. Proportional length of dorsal abdominal
segments (third to eighth):: 48: 50: 53: 57: 62: 55. First segment
with median elevated area sinuate on posterior margin. Second
segment with longitudinal carinae well defined, almost straightly
divergent posteriorly; anteromesal impression obsolete, obscurely
depressed along posterior margin of first segment on either side
of carinae, anterolateral impression obliquely placed, posterolateral
impression small, separated from anterolateral impression, but
placed on the same shallow depression along inner margin of
second connexival segment; intersegmental groove well defined,
longitudinally rugose. Third dorsal abdominal segment with longi-
tudinal carinae almost straightly divergent posteriorly; anteromesal
impression well marked, depressed along posterior margin of sec-
ond segment on either side of median longitudinal carinae, antero-
lateral and posterolateral impressions clearly separated from each
other; intersegmental groove well defined and longitudinally rugu-
lose. Seventh dorsal abdominal segment widened posteriorly,
posterior margin sinuate, seventh connexival segment with inner
margin sinuate, lateral margin rounded, clothed with long hairs.
Eighth segment with posterior margin almost straight, posterolateral
margin rounded; paratergite with fine hairs. The last genital
segment small, rounded apically, clothed with fine hairs. Meta-
sternum behind transverse suture inclined posteriorly, shortened
medially.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 50: 42: 42: 43: 50: 54: 63: 56 (allotype).
Connexivum reflexed laterally at angle of about forty-five degrees.
First dorsal abdominal segment with long dark brown hairs di-
rected posteriorly.
954 Tse UNiversiry SCIENCE BULLETIN
Wingless male: Pronotum entirely covers mesonotum. Propor-
tional length of dorsal abdominal segments (first to seventh):: 40:
44: 35: 35: 35: 42: 75. Connexivum horizontally spread laterally.
Metasternum behind transverse suture strongly inclined posteriorly,
shorter in the middle. Venter with obscure median longitudinal
elevation extending from base to sixth segment. Seventh ventral
abdominal segment sparsely decorated with minute black spines on
lateral area and along posterior margin at sides.
Rhagovelia perfidiosa Bacon
(Pl. 9, figs. 29a, b)
1948. Rhagovelia perfidiosa Bacon, Jour. Kansas Ent. Soc., vol. 21, p. 81.
Winged male: Pronotum with apex broadly rounded. Forewings
pale fuscous, whitish at base, veins darker and thick, with two large
apical cells. Proportional length of dorsal abdominal segments
(second to seventh):: 82: 32: 82: 85: 35: 55. All dorsal abdominal
segments are transverse. Second dorsal abdominal segment with
longitudinal carinae slightly roundly widened at middle, finely
longitudinally rugose along posterior margin of first dorsal ab-
dominal segment throughout; anterolateral and posterolateral im-
pressions clearly separated; intersegmental groove between second
and third segments finely longitudinally rugulose. Third segment
with longitudinal carinae less strongly rounded than in second
segment; intersegmental groove between third and fourth segments
finely longitudinally rugose; anterolateral and posterolateral im-
pressions contiguous, Metasternum behind transverse suture in-
clined posteriorly, shortened in the middle. Seventh ventral
abdominal segment with posterior margin without black minute
spines. Median longitudinal elevation on ventral surface obscure.
Wingless male: Pronotum just reaching posterior margin of meso-
notum. Proportional length of dorsal abdominal segments (first
to seventh):: 82: 81: 28: 24: 25: 30: 52 (holotype). First dorsal
abdominal segment with anterior margin slightly roundly pro-
duced. Ventral surface of body as in winged male.
Winged female: Proportional length of dorsal abdominal seg-
ments (fifth to eighth):: 86: 40: 48: 45. Seventh dorsal abdominal
segment slightly widened posteriorly; seventh connexival segment
with apex acutely pointed, reaching the middle of eighth abdominal
segment, with a bundle of dark brown setae arising ventrally from
near apex. Eighth segment with posterolateral margin broadly
rounded. The last genital segment small, conical apically.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 955
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth ):: 37: 31: 32: 28: 33: 40: 50: 42, Connexivum
subvertically erected. Metasternum behind transverse suture shorter
than in male.
Rhagovelia relicta Gould
1981. Rhagovelia relicta Gould, Univ. Kansas Sci. Bull., vol. 20, p. 39.
Wingless male: Proportional length of dorsal abdominal segments
(first to sixth):: 33: 40: 35: 35: 37: 40 (holotype). Ventral longi-
tundinal ridge of abdomen obliterated.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 38: 45: 40: 89: 43: 48: 58: 50 (allotype).
Connexivum vertically reflexed throughout.
Rhagovelia robusta Gould
1931. Rhagovelia robusta Gould, Univ. Kansas Sci. Bull., vol. 20, p. 41.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 41: 51: 48: 44: 45: 50: 71 (holotype). Meta-
sternum behind transverse suture strongly inclined posteriorly,
strongly shortened in the middle. Ventral abdominal segments
strongly depressed laterally and with distinct median longitudinal
carina from base down to sixth segment. Seventh ventral abdominal
segment without black spines on posterior margin.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):; 40: 47: 45: 45: 51: 59: 64: 51 (allotype).
Metasternum behind transverse suture strongly inclined posteriorly,
shorter in the middle. Connexivum slightly reflexed.
Rhagovelia scitula Bacon
1956, Rhagovelia scitula Bacon, Univ, Kansas Sci, Bull., vol, 88, pt. I, p. 804.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh ):: 38: 45: 87: 88: 40: 44: 63, Metasternum
behind transverse suture inclined posteriorly, shortened in the
middle. Seventh ventral abdominal segment without black spines
on posterior margin.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 40: 50: 45: 45: 52: 55: 72: 52. Meta-
sternum as in wingless male.
Tue UNtversiry SCIENCE BULLETIN
Rhagovelia sinuata Gould
(Pl. 9, figs. S8la, b)
1931. Rhagovelia sinuata Gould, Univ. Kansas Sci. Bull., vol. 20, p. 42.
Winged female: Pronotum rounded apically. Forewings dark
fuscous, lighter in upper basal cell, veins black. Proportional
length of dorsal abdominal segments (second to eighth):: 55: 53:
55: 58: 62: 63: 55. First dorsal abdominal segment with median
elevated area strongly sinuate on posterior margin; deeply depressed
along posterior margin of first segment on either side of the carinae.
Second segment with longitudinal carinae narrow but well marked,
strongly divergent at base, then parallel-sided; posterolateral im-
pression deep and well marked, distinctly separated from antero-
lateral impression; intersegmental groove between second and
third segments shallow but well defined, longitudinally rugose.
Third segment with longitudinal carinae divergent posteriorly;
anteromesal impression shallow but distinct; anterolateral and
posterolateral impressions are clearly separated; intersegmental
groove shallow but well defined, longitudinally rugose, with a
transverse shallow and distinct impression behind the posterior end
of longitudinal carinae on third segment. Seventh dorsal abdominal
segment slightly dilated at basal one third, distinctly wider at base
than long in the middle, (90:63). Seventh connexival segment
narrowed apically, its lateral margin rounded, apex acutely pointed
and reaching the middle of eighth segment, apical margin sinuate.
Eighth segment with apical margin broadly rounded, with a bundle
of long brown hairs on the dorsal surface of eighth connexival
segment. Metasternum behind transverse suture strongly inclined
posteriorly, strongly and straightly narrowed medially. Ventral
surface of abdomen without longitudinal carina.
Wingless female: Pronotum entirely covers mesonotum. Pro-
portional length of dorsal abdominal segments (first to eighth)::
43: 53: 49: 53: 55: 60: 67: 57. Metasternum as in winged female.
Winged male: Pronotum broadly rounded on apical margin.
Proportional length of dorsal abdominal segments (first to sev-
enth):: 40; 50; 45: 43: 48: 52: 80. Metasternum as in winged
female. Ventral side of abdomen without median longitudinal
carina. Seventh segment without black spines on posterior margin.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 40: 50: 45: 43: 48: 52: 80. Metasternum
and ventral side of abdomen as in winged male. Connexivum
spread horizontally laterad.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 957
Rhagovelia varipes Champion
(Pl. 10, figs. 82a, b)
1898. Rhagovelia varipes Champion, Biol. Centr. Amer., Het., Vol. ¥, p. 13S.
Winged female: Pronotum broadly rounded apically. Forewings
fuscous, veins black, with distinct two apical cells. Proportional
length of dorsal abdominal segments (second to eighth):: 81: 80:
90: 87: 85: 87: 85. First dorsal abdominal segment with median
elevated area posteriorly well defined and substraight. Second seg-
ment with median longitudinal carinae fine but well defined, feebly
produced laterally at middle; anteromesal impression obliterated,
obliquely depressed along posterior margin of first segment on
either side of the carinae, posterolateral impression round, clearly
separated from anterolateral impression; intersegmental groove ob-
scurely defined and obscurely longitudinally rugulose. Third seg-
ment with longitudinal carinae feebly produced laterally at middle;
anteromesal impression small and shallow; anterolateral and pos-
terolateral impressions located on shallow depression along inner
margin of third segment; intersegmental groove between third and
fourth segments shallow, obscurely defined anteriorly. Seventh
dorsal abdominal segment with posterior margin broadly sinuate;
seventh connexival segment with lateral margin rounded, slightly
narrowed posteriorly, apical margin sinuate. Eighth segment about
as long as the preceding segment in the middle, posterior margin
broadly rounded and densely clothed with long dark brown hairs,
denser and longer laterally. The last genital segment small, rounded
apically. Metasternum behind transverse suture strongly shortened
in the middle, distinctly longitudinally carinate on basal two seg-
ments. Spiracle on seventh segment placed closer to posterior mar-
gin than to anterior margin.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 47: 72: 71: 76: 76: 81: 85: 70. Meta-
sternum as in winged male. Fine median longitudinal carinae on
venter as in winged female.
Winged male: Proportional length of dorsal abdominal segments
(first to seventh) :: 27: 75: 65: 73: 67: 67: 80. Metasternum as in
winged female. Fine median longitudinal carina extends down to
basal one fourth of fourth segment. Spiracle on seventh segment
placed a little closer to anterior margin than to posterior margin.
Seventh ventral abdominal segment without black spines on pos-
terior margin.
958 Tae Universtry ScieNCE BULLETIN
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 53: 85: 75: 65: 67: 65: 100. Connexivum ver-
tically erected. Metasternum, ventral longitudinal carina and loca-
tion of spiracles as in winged male.
Rhagovelia whitei (Breddin )
1898. Rhagovelia whitei Breddin, Jahrb. Nat. Ver. zu Magee, p. 14.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 35: 48: 43: 43: 48: 51: 68. Metasternum be-
hind transverse suture inclined posteriorly. Median longitudinal
ridge distinct down to sixth segment.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 35: 48: 43: 43: 48: 51: 57: 49. Connexi-
vum reflexed at angle about forty-five degrees.
Rhagovelia williamsi Gould
1931. Rhagovelia williamsi Gould, Univ. Kansas Sci. Bull., vol. 20, p. 47.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 35: 48: 48: 43: 48: 51: 68. Metasternum behind
transverse suture inclined posteriorly. Median longitudinal ridge
distinct down to sixth segment.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 35: 48: 43: 43: 48: 51: 57: 49. Connexivum
reflexed at angle of about forty-five degrees.
Group CHARACTERISTICS OF THE CoLLARIs Group
The following species were available for the study of winged
forms:
Rh. acuminata Bacon Rh. planipes Gould
Rh. armata (Burmeister ) Rh. scabra Bacon
Bh. collaris (Burmeister ) Rh. tayloriella Kirkaldy
Rh. impensa Bacon
Longitudinal carinae fine and well defined, parallel-sided except
for Rh. collaris and Rh. tayloriella in which the carinae are divergent
posteriorly and tend to be obliterated.
Forewings with two large apical cells formed in distal half of
the wing; subcosta extends apically as far as apical one fourth.
Pronotum in winged male develops into a long process; con-
spicuous projection at base of apical process directed downwardly.
Proportional length of sixth and seventh dorsal abdominal seg-
ments in winged male ranges from 44:70 in Rh. collaris to 80:92 in
Rh. scabra, that in wingless male ranges from 58:85 in Rh. solida to
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 959
67:76 in Rh. armata. Proportional length of seventh and eighth
dorsal abdominal segments ranges from 75:60 in Rh. planipes to
85:60 in Rh. acuminata, that in wingless female ranges from 88:98
in Rh. scabra to 82:59 in Rh. planipes.
The following characters are also peculiar to this group:
(1) In the intermediate leg the tibia is always more than twice
as long as the second tarsal segment; third tarsal segment is always
longer than the second one.
(2) In the hind leg the femur is considerably longer than the
tibia except in Rh. collaris and Rh. tayloriella in which they are
subequal to each other.
(3) The eighth segment with the paratergite in winged females
with a bundle of hairs arising ventrally.
(4) The apex of the seventh connexival segment acutely pro-
duced except in Rh. collaris and Rh. tayloriella.
(5) In wingless females the dorsum of the abdomen is narrow.
(6) The intermediate femur of female is flattened beneath at
base.
(7) The anterior tibia of the male is usually greatly dilated and
excavated within.
Rh, collaris and Rh. tayloriella in common apparently deviate in
some characters from other representatives of this group.
Rhagovelia acuminata Bacon
(Pl. 12, figs. 40a, b, c)
1956. Rhagovelia acuminata Bacon, Univ. Kansas Sci, Bull., vol. 88, pt. I, p. 817.
Winged male: Pronotum acutely pointed apically, obscure longi-
tudinal carinae running in the middle throughout the entire length
of pronotum. Forewings fuscous, veins thick and black. Propor-
tional length of dorsal abdominal segments (first to seventh ):: 23:
51: 68: 66: 70: 74: 92. First dorsal abdominal segment with median
elevated area well defined laterally and posteriorly. Second seg-
ment with longitudinal carinae parallel-sided, thin, sparsely irregu-
larly denticulated on lateral margin; anteromesal impression
obsolete, anterolateral and posterolateral impressions shallow and
clearly separated, shallowly depressed along posterior margin of
first segment on either side of the carinae; intersegmental groove
between second and third segments shallow. Third segment with
longitudinal carinae feebly rounded at middle, extending posteriorly
into intersegmental groove and black, lateral margin sparsely
irregularly denticulated; anteromesal, anterolateral and_postero-
960 Tue University SCIENCE BULLETIN
lateral impressions shallow and separated from each other. Sev-
enth dorsal abdominal segment strongly widened posteriorly; sev-
enth connexival segment elongate subtriangular in shape, feebly
rounded in the posterior half of lateral margin, its inner margin
sinuate; posterior margin of seventh ventral segment straight in the
middle, then oblique and slightly rounded laterally. Metasternum
behind transverse suture inclined posteriorly, strongly shortened
in the middle. Second ventral abdominal segment depressed.
Ventral surface of abdomen obscurely longitudinally carinate in
the middle from posterior region of metasternum to sixth segment.
Seventh ventral abdominal segment with posterior margin armed
with a series of small, black spines laterally.
Wingless male: Pronotum completely covering mesonotum. Pro-
portional length of dorsal abdominal segments (first to seventh )::
36: 55: 58: 60: 70: 74: 93 (holotype). First dorsal abdominal seg-
ment with anterior margin gently roundly, produced anteriorly.
Ventral surface of abdominal segments as in winged male.
Winged female: Pronotum produced posteriorly as a long process
in female, this process is also provided with a projection at base
and with a shallow cavity cephalad to it. Proportional length of
dorsal abdominal segments (fourth to eighth):: 79: 85: 90: 85: 60.
Seventh dorsal abdominal segment a little less than 1% times as long
as eighth segment; seventh connexival segment with lateral margin
feebly rounded and narrowed posteriorly, with brown hairs through-
out, these hairs become increasingly longer posteriorly; apex
strongly acutely pointed and produced posteriorly, distinctly ex-
tending beyond posterior margin of eighth segment; posterior
margin of seventh ventral abdominal segment feebly bisinuate.
Eighth segment provided with a conspicuous bundle of long, black
hairs arising from ventral side. Ventral side of abdomen without
depression at base. Metasternum behind transverse suture length-
ened at middle, posterior margin of metasternum bisinuate. Second
and. third ventral abdominal segments with median longitudinal
elevation in anterior half. Seventh ventral abdominal segment with
posterior margin with small black tubercles on either side of the
base of ninth segment.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 43: 52: 64: 80: 91: 95(?): 75(?). Con-
nexivum almost vertically reflexed throughout. Second dorsal ab-
dominal segment strongly inclined posteriorly. First dorsal ab-
dominal segment with anterior margin broadly rounded and
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 961
produced anteriorly, clothed with long yellowish brown hairs di-
rected posteriorly. Posterior margin of seventh ventral abdominal
segment with a mass of black tubercles at sides.
Rhagovelia armata (Burmeister )
(Pl. 12, figs. 89a, b, c)
1885. Velia armata Burmeister, Handb. d. Ent., Bd. 2, p. 212.
Winged female: Pronotum develops posteriorly into a long proc-
ess, directed upwards at the angle of forty-five degrees at base,
then almost horizontally tapering into acute tip, with a light
yellow process directed downward at base, this basal process is
hollowed out on caudal side. Forewings purplish black, unicolor-
ous, with two large apical cells. Proportional length of dorsal
abdominal segments (first to eighth):: 27: 47: 64: 73: 80: 80: 77:
60. First dorsal abdominal segment with median elevated area
straight and well defined posteriorly. Second dorsal abdominal
segment with longitudinal carinae parallel-sided, lateral margin
with irregular obscurely defined denticulations, depressed along
posterior margin of first segment on either side of longitudinal ca-
rinae; anteromesal impression obsolete, anterolateral and postero-
lateral impressions distinct and large but separated from each other;
intersegmental groove rather shallow, well-defined anteriorly and
ill-defined posteriorly. Third segment with longitudinal carinae
subparallel-sided, lateral margin with irregular, obscure denticula-
tions; anteromedial impression well marked and black, anterolateral
and posterolateral impressions subcontiguous; intersegmental groove
between third and fourth segments black. Fifth to seventh dorsal
abdominal segments each almost as long as wide. Seventh con-
nexival segment acutely pointed, its apical margin sinuate, not
reaching to the apex of eighth segment, posterior margin of sev-
enth segment on ventral side feebly bisinuate; seventh ventral
abdominal segment with posterior margin with a mass of few
small, black tubercles at sides. Eighth connexival segment with a
bundle of long, black hairs arising from ventral side. Last genital
segment densely haired, gradually narrowed apically. Metaster-
num behind transverse suture lengthened at middle, posterior
margin feebly bisinuate.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 42: 48: 52: 63: 80: 85: 88: 67. First dorsal
abdominal segment with anterior margin broadly rounded and
produced, Second and third segments inclined posteriorly. Fourth,
fifth, and sixth segments roundly produced anteriorly. Connexivum
962 Tue University ScreENCE BULLETIN
almost vertically reflexed. Metasternum as in winged female.
Seventh ventral abdominal segment with posterior margin armed
with small black tubercles at sides.
Winged male: Metasternum behind transverse suture shortened
in the middle, posterior margin broadly sinuate. Abdomen on
ventral surface strongly depressed in basal region.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 40: 55: 52: 53: 56: 67: 76. Metasternum
and seventh ventral abdominal segment as in winged male.
Rhagovelia collaris (Burmeister )
(Pl, 11, figs: 87a, b,c)
1835. Velia collaris Burmeister, Handb. d. Ent., Bd. 2, p. 212.
Winged female: Pronotum with apical process rather thick, di-
rected upwardly at the angle of about forty-five degrees, then
directed slightly downward; the process at base of apical process
small and rounded. Forewings dark fuscous brown, veins black,
with two large apical cells. Proportional length of dorsal abdominal
segments (first to eighth):: 23: 88: 47: 47: 50: 52: 68: 50. First
dorsal abdominal segment well defined posteriorly. Second seg-
ment with longitudinal carinae roundly divergent posteriorly;
anteromesal impression distinct, anterolateral and posterolateral
impressions subcontiguous, distinctly depressed along posterior
margin of first segment on either side of carinae, the area within
the carinae transverse; intersegmental groove distinctly impressed,
well defined anteriorly, obscurely defined posteriorly. Third seg-
ment with longitudinal carinae divergent posteriorly, not quite
reaching posterior margin of the segment; anteromesal impression
small, anterolateral and posterolateral impressions continuous. Sev-
enth segment wider at base than it is long at middle (90:63);
seventh connexival segment narrowed apically, its lateral margin
broadly rounded, apex acutely rounded; posterior margin of sev-
enth ventral abdominal segment bisinuate, densely pubescent
throughout. Eighth segment purplish black except for the middle
yellow; lateral margin of eighth connexival segment almost straight,
with a bundle of long dark brown hairs arising mainly from the
ventral side, posterior margin of eighth dorsal abdominal segment
broadly rounded. The last genital segment subparallel, narrowed
apically, downward and densely clothed with dark hairs espe-
cially on lateral margin. Metasternum behind transverse suture
short, posterior margin broadly sinuate and shortened in the middle.
¥
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 963
Wingless female: Connexivum strongly reflexed and folded on
the dorsum completely after second segment.
Winged male: Pronotum simply and acutely pointed. Propor-
tional length of dorsal abdominal segments (first to seventh):: 25:
38: 41: 42: 42: 44: 70. Metasternum behind transverse suture and
basal two ventral abdominal segments depressed, posterior margin
of metasternum more strongly sinuate than in female. Obscure
longitudinal carinae running in the middle of metasternum behind
transverse suture and second to sixth ventral abdominal segments.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 45: 48: 48: 46: 47: 53: 77 (paratype). Meta-
sternum as in winged male.
This species is different from Rh. armata (Burmeister ) as follows:
(1) The longitudinal carinae are strongly divergent posteriorly,
and tend to be obliterated apically on the third segment.
(2) The last female genital segment is strongly bent downward.
(8) The posterior margin of the seventh connexival segment is
rounded.
Rhagovelia impensa Bacon
(PI. 11, figs. 88a, b, ¢)
1956, Rhagovelia impensa Bacon, Univ. Kansas Sci. Bull., vol. 38, pt. I, p. 827.
Winged male: Pronotum narrow and acutely pointed at apex.
Forewings dark fuscous, veins black, with two large apical
cells. Proportional length of dorsal abdominal segments (first
to seventh):: 25: 42: 50: 55: 58: 60: 75. Second segment
with longitudinal carinae slightly divergent posteriorly, with ob-
scure denticulation on lateral margin; anteromesal impression
obsolete, lateral impressions small, clearly separated, hardly
depressed along posterior margin of first segment, shallowly
depressed along inner margin of second connexival segment;
intersegmental groove well defined anteriorly, obscurely defined
posteriorly. Third segment with longitudinal carinae subparallel-
sided, slightly divergent posteriorly; anteromesal impression ob-
literated; anterolateral and posterolateral impressions separated,
depressed along inner margin of third connexival segment; inter-
segmental groove distinct anteriorly, obscure on posterior margin.
Metasternum behind transverse suture and basal two ventral ab-
dominal segments strongly depressed and with median longitudinal
Carinae; transverse suture on metasternum straight, posterior margin
of metasternum broadly roundly emarginated. Posterior margin of
964 Tue Unriversiry ScreENCE BULLETIN
seventh ventral abdominal segment with a series of minute black
spines. ;
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 85: 48: 44: 48: 47: 53: 80. Ventral side
of the body as in winged male.
Winged female: Pronotum with lower margin of apical process
roundly continuous with posterior margin of basal process, lower
margin of basal process rather weakly developed, apex of apical
process densely clothed with long hairs. Seventh dorsal abdominal
segment as long in the middle as wide at base; seventh connexival
segment reflexed laterally in apical half, acutely pointed. Eighth
segment about three-fourths as long as preceding segment, posterior
margin broadly rounded; paratergite produced laterally at basal
one third; disk of eighth segment with a pair of round cavities on
basal margin, apical half and the last genital segment strongly di-
rected downwardly. Lateral margin of the last genital segment
clothed with short but strong black hairs. Metasternum behind
transverse suture not inclined posteriorly, subequal in length
throughout except for leg base. Seventh ventral abdominal seg-
ment with posterior margin with black tubercles at sides. Eighth
ventral abdominal segment with posterior margin with a series of
minute black spines directed inwardly. ,
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to sixth) :: 40: 45: 45: 41: 55: 68. Connexivum strongly
reflexed, at more than ninety degrees after third segment. Posterior
margin on ventral side as in winged female.
Rhagovelia planipes Gould
(Pl. 12, figs. 41a, b, c)
1931. Rhagovelia collaris planipes Gould, Univ. Kansas Sci. Bull., vol. 20, p. 22.
Winged female: Pronotum with apical area strongly almost hori-
zontally produced posteriorly as a process, with a well-developed
basal process directed downward. Forewings fuscous, upper basal
cell whitish, veins black, with two large apical cells. Proportional
length of dorsal abdominal segments (first to eighth):: 27: 52: 63:
73: 73: 80: 75: 60. Second dorsal abdominal segment with longi-
tudinal carinae straight and slightly divergent posteriorly; obscurely
grooved along posterior margin of first dorsal abdominal segment;
anteromesal impression small and round, anterolateral and postero-
lateral impressions clearly separated; intersegmental groove well
defined and with obscure oblique elevations. Third segment with
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 965
longitudinal carinae parallel-sided, extending slightly into fourth
dorsal abdominal segment; anteromesal impression small and round,
depressed along posterior margin of second segment on either side of
longitudinal carinae; anteromesal impression small and round, an-
terolateral and posterolateral impressions separated from each other;
intersegmental groove between third and fourth segments rather
obscurely defined. Seventh dorsal abdominal segment a little wider
than long at middle (90:75); seventh connexival segment gradually
narrowed apically, apex acutely pointed and black, reaching pos-
terior margin of eighth segment. Eighth segment with posterior
margin broadly rounded; paratergite slightly elevated. Metaster-
num behind transverse suture short and slightly longer near leg
base. Posterior margin of seventh ventral abdominal segment with
several small black teeth laterally.
Wingless female: Proportional length of dorsal abdominal seg-
ments (second to eighth):: 58: 58: 62: 78: 80: 82: 59. Second and
third dorsal abdominal segments inclined posteriorly. Connexivum
reflexed at the angle a little less than ninety degrees. Metasternum
as in winged female.
Winged male: Pronotum narrowed apically and acutely pointed.
All dorsal abdominal segments transverse. Proportional length of
dorsal abdominal segments (fifth to seventh) :: 58: 58: 89. Seventh
dorsal abdominal segment strongly widened posteriorly; seventh
connexival segment subtriangular and relatively short; seventh ven-
tral abdominal segment with posterior margin deeply sinuate, with
small, black teeth laterally. Metasternum behind transverse suture
strongly inclined posteriorly, posterior margin broadly roundly
sinuate, shorter in the middle. Ventral surface of abdomen de-
pressed and with longitudinal carinae in the middle.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 45: 55: 55: 52: 52: 65: 82. Metasternum be-
hind transverse suture as in winged form.
Rhagovelia scabra Bacon
(Pl. 8, fig. 24)
1956, Rhagovelia scabra Bacon, Univ. Kansas Sci, Bull., vol. 38, pt. I, p. 831.
Winged male: Pronotum narrowed apically, acutely pointed.
Wings purplish fuscous, veins black, with two large apical cells.
Proportional length of dorsal abdominal segments (first to seventh ) ::
30: 57: 71: 78: 73: 80: 92. First dorsal abdominal segment with
posterior margin on either side of median elevation obliterated.
Tse Universtry ScreENCE BULLETIN
966
Second dorsal abdominal segment with longitudinal carinae sub-
parallel except at base slightly divergent, narrow, well-defined;
anteromesal impression obliterated, anterolateral and posterolateral
impressions clearly separated, slightly depressed along inner margin
of second connexival segment; intersegmental groove between sec-
ond and third segments well defined anteriorly, obscurely defined
posteriorly. Third segment with longitudinal carinae narrow, sub-
parallel except in intersegmental region; anteromesal impression
obliterated, anterolateral and posterolateral impressions separated,
depressed along inner margin of third connexival segment; inter-
segmental groove well defined anteriorly. Metasternum with trans-
verse suture straight, metasternum behind transverse suture broadly
sinuate on posterior margin and shorter in the middle, strongly de-
pressed. Second and third ventral abdominal segments strongly de-
pressed. Posterior margin of seventh ventral abdominal segment
broadly roundly emarginated, with a series of small black teeth on
either side of the middle.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 87: 62: 55: 55: 57: 63: 80 (holotype). Meta-
sternum and ventral side of abdomen as in winged male.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth ):: 42: 60: 65: 67: 88: 88: 98: 70. Second and
third dorsal abdominal segments strongly inclined posteriorly, apical
half of seventh and basal half of eighth dorsal abdominal segments
clothed with long black hairs. Metasternum with transverse suture
straight, posterior margin of metasternum feebly bisinuate. Con-
nexivum reflexed at more than ninety degrees after third segment.
Rhagovelia solida Bacon
1956. Rhagovelia solida Bacon, Univ. Kansas Sci. Bull., vol. 88, pt. I, p, 833.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 47: 55; 54: 54: 54: 58: 85 (holotype).
First dorsal abdominal segment broadly rounded and slightly pro-
duced. Metasternum behind transverse suture inclined posteriorly,
strongly shortened in the middle. Seventh ventral abdominal seg-
ment with posterior margin armed with a series of minute black
spines laterally, deeply sinuate.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to fifth):: 55: 62: 57: 78: 87 (allotype). Connexivum
reflexed at the angle more than ninety degrees after third segment.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 967
Third segment inclined posteriorly, Metasternum behind transverse
suture level.
Rhagovelia tayloriella Kirkaldy
(Pl. 11, figs. 36a, b, c)
1900. Rhagovelia tayloriella Kirkaldy, Ent., vol. 38, p. 72.
Winged female: Pronotum with apical process rather short, ro-
bust, raised upwardly at the angle a little less than forty-five
degrees, with a weakly developed process at base of apical process.
Forewings with a greenish tinge along the upper margin in basal
half. Second dorsal abdominal segment with longitudinal carinae
roundly divergent towards middle; anteromesal impression obso-
lete, anterolateral and posterolateral impressions clearly separated,
shallowly depressed along posterior margin of first dorsal abdom-
inal segment on either side of carinae, anterolateral and postero-
lateral impressions clearly separated; intersegmental groove between
second and third segments strongly produced and lengthened
anteriorly at middle. Third segment with longitudinal carinae less
distinct than those on second segment, but reaching posterior
margin of third segment, slightly divergent posteriorly; anteromesal
laevigate impression obsolete, instead distinctly depressed along
posterior margin of second dorsal abdominal segment on either
side of longitudinal carinae, distinctly depressed along inner margin
of third connexival segment, anterolateral and posterolateral im-
pressions separated. Disk bounded laterally by longitudinal ca-
rinae, transverse in both second and third segments; intersegmental
groove between third and fourth segments distinct. Seventh dorsal
abdominal segment about as long as wide at base; seventh con-
nexival segment slightly narrowed posteriorly, apex narrowly
rounded and produced, thickly clothed with long dark brown hairs;
seventh ventral abdominal segment with posterior margin feebly
bisinuate. Eighth segment with posterior margin broadly rounded;
paratergite in basal half strongly reflexed, with a bundle of long,
dark brown hairs arising from ventral side. The last genital seg-
ment inclined posteriorly, densely clothed with short hairs. Meta-
sternum behind transverse suture shorter at middle, depressed and
with median longitudinal carina. Second ventral abdominal seg-
ment depressed and with median longitudinal carina which is
continuous with that on metasternum.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to eighth):: 42: 42: 42: 46: 47: 63: 77: 55. Third
segment inclined posteriorly, fifth to seventh segments each longer
968 Tue Untversiry Scrence BULLETIN
than wide, posterior margin of seventh dorsal abdominal segment
densely clothed with long, black hairs. Connexivum reflexed at
about right angle. Metasternum behind transverse suture subequal
in length throughout, slightly shorter at middle. Seventh ventral
abdominal segment with posterior margin feebly bisinuate. Last
genital segment bent downward.
Winged male: Proportional length between sixth and seventh
segment:: 55: 70.
Wingless male: Proportional length of dorsal abdominal seg-
ments (first to seventh):: 35: 43: 41: 41: 45: 48: 70. Seventh
ventral abdominal segment without a series of black spines on
posterior margin.
Remarks: This species has characteristics in common with Rh.
collaris (Burmeister) as follows:
(1) The area bounded laterally by longitudinal carinae on the
second and third dorsal abdominal segments distinctly wider than
long.
(2) The male without a series of black spines on the posterior
margin of the seventh ventral abdominal segment.
(3) The hind femur and tibia are subequal in length.
(4) The last genital segment is strongly bent downward.
(5) The apex of the seventh connexival segment is not acutely
produced,
Group CHARACTERISTICS OF THE OpEsA Group
The following three species were available for the study of winged
forms: Rh. distincta Champion, Rh. obesa Uhler, and Rh. rivale
Bueno.
Pronotum in winged female with short and thick process apically;
pronotum in wingless forms rather short except for Rh. oriander.
Forewings with two large apical cells formed in distal half of
the wing.
Longitudinal carinae occur always on second and third dorsal
abdominal segments in winged forms.
Proportional length of sixth and seventh dorsal abdominal seg-
ments is about 2:3 in winged male of Rh. distincta and Rh. rivale;
that of wingless male ranges from 40:77 in Rh. obesa to 46:75 in
Rh. distincta. Proportional length of seventh and eighth segments
in winged female is 82:60 in Rh. distincta and 55:45 in Rh. obesa,
that in wingless female is 110:78 in Rh. distincta.
The following characters are also peculiar to this group.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 969
(1) The lateral margin of the seventh connexival segment in
winged females sinuate.
(2) The posterior margin of the seventh ventral abdominal seg-
ment with a series of minute black spines, except in Rh. rivale.
(3) The eighth segment has a conspicuous bundle of long setae
arising ventrally near the lateral margin in the female.
(4) The intermediate femur is dorsoventrally flattened in the
female.
(5) The hind femur is distinctly longer than the tibia except in
Rh. rivale, in which they are subequal to each other.
(6) The anterior tibia is never greatly dilated or excavated
beneath in the male.
As noted from the above Rh. rivale deviates distinctly from other
representatives of this group at least in two characters. These two
characters, i. ¢., the proportional length of the hind femur and tibia
and the absence of black spines on posterior margin of seventh
ventral abdominal segment are characters shared by Rh. collaris
and Rh. tayloriella of the collaris group. Rh. rivale, however, is
definitely a member of this group in the structure of the anterior
tibia upon which Bacon’s classification is based.
Rhagovelia distincta Champion
(Pl. 13, figs. 42a, b, c)
1877. Rhagovelia mexicana Signoret, Bull. Soc. Ent. Fr. (5): 7: p. 53 (no de-
oa ead 8 an : ‘
1898. Rhagovelia distincta Champion, Biol. Centr. Amer., Het., vol. 2, p. 185.
Winged female: Pronotum with apical process rather short and
thick, clothed with short black hairs, raised upward at an angle a
little less than forty-five degrees, upper margin of apex almost
horizontal, basal process weakly developed. Forewings dark fus-
cous, with greenish tinge on upper basal cell, veins black, two large
apical cells formed in distal half. Proportional length of dorsal ab-
dominal segments (second to eighth):: 50: 50: 55: 59: 63: 82: 60.
Longitudinal carinae on second dorsal abdominal segment slightly
divergent posteriorly, anteromesal impression obliterated, antero-
lateral and posterolateral impressions clearly separated from each
other; intersegmental groove between second and third segments
shallow but long, with obscure longitudinal elevation. Third seg-
ment with longitudinal carinae slightly divergent posteriorly, extend
slightly beyond intersegmental groove apically in one specimen
examined; anterolateral and posterolateral impressions clearly sepa-
rated from each other; intersegmental groove between third and
970 Tue University ScreNCE BULLETIN
fourth segments shorter, obscurely longitudinally rugose. Seventh
dorsal abdominal segment a little longer in middle than basal
width (82:75); seventh connexival segment narrowed apically, apex
narrowly rounded, reaching the middle of eighth segment; seventh
ventral abdominal segment feebly bisinuate at middle on posterior
margin. Eighth dorsal abdominal segment with apical margin
broadly rounded; paratergite ill defined posteriorly, with a bundle
of long black setae arising from ventral surface. Last genital seg-
ment short. Metasternum behind transverse suture level, short,
subequal in length throughout.
Wingless female: Proportional length of dorsal abdominal seg-
ments (first to seventh) :: 42: 50: 57: 71: 90: 110: 78. Connexivum
behind third segment folded on dorsum. Third dorsal abdominal
segment not strongly inclined posteriorly.
Winged male: Proportional length of dorsal abdominal segments
(first to seventh ):: 28: 43: 50: 45: 45: 50: 72. Metasternum behind
transverse suture inclined posteriorly, shortened medially, Seventh
ventral abdominal segment with posterior margin armed with a
series of minute black spines.
Wingless male: Mesonotum slightly exposed behind pronotum.
Proportional length of dorsal abdominal segments ( first to seventh) ::
37: 45: 43: 41: 41: 46: 75. Metasternum behind transverse suture
inclined posteriorly.
Rhagovelia knighti Drake and Harris
1927. Rhagovelia knighti Drake and Harris, Proc. Biol. Soc. Washington, vol.
40, p. 183,
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 29: 34: 31: 80: 30: 36: 67 (paratype). Third
dorsal abdominal segment not inclined posteriorly. Metasternum
behind tranverse suture rather strongly shortened in the middle.
Seventh ventral abdominal segment with posterior margin armed
with a series of black spines laterally.
Wingless female: Proportional length of dorsal abdominal seg-
ments (second to eighth):: 80; 33: 84: 29: 38: 47: 75. Connexivum
strongly reflexed and folded on the dorsum, Metasternum behind
transverse suture short, subequal in length throughout except for
near leg base slightly longer.
SUPPLEMENTARY STUDY OF THE GENUS: RHAGOVELIA 971
Rhagovelia obesa Uhler
(Pl. 18, figs. 44a, b, c)
1871. Rhagovelia obesa Uhler, Proc. Bost. Soc, Nat. Hist., vol. 14, p. 107.
Winged female: Pronotum with apex produced posteriorly as a
thick process, and dilate apex notched on apical margin. Forewings
fuscous, veins black, with two large apical cells. Proportional length
of dorsal abdominal segments (first to eighth) :: 22: 85: 32: 83: 40:
43: 55: 45. Second dorsal abdominal segment with longitudinal
carinae straight, slightly convergent posteriorly; anteromesal im-
pression obsolete, anterolateral and posterolateral impressions small
and clearly separated; intersegmental groove between second and
third segments rather obscure. Third segment with longitudinal
carinae straight, subparallel; anterolateral and posterolateral im-
pressions subcontiguous; intersegmental groove short, obscurely
longitudinally rugose. Seventh dorsal abdominal segment longer
at middle than basal width (55:50); seventh connexival segment
with lateral margin sinuate in apical half, apex narrowly rounded
and with short, black hairs, Eighth dorsal abdominal segment with
paratergite strongly reflexed, with a bundle of long and black setae,
posterior margin of eighth segment almost straight, posterolateral
angle broadly rounded. The last genital segment gradually nar-
rowed apically, apical margin rounded. Metasternum behind trans-
verse suture short and level.
Wingless female: Mesonotum well exposed posteriorly, posterior
margin of pronotum feebly sinuate. Proportional length of dorsal
abdominal segments (first to sixth):: 82: 82: 32: 28: 32: 47. Con-
nexivum strongly reflexed and the dorsum narrowly exposed. Sec-
ond and third segments slightly inclined posteriorly. Metasternum
behind transverse suture short, slightly lengthened at middle.
Wingless male: Pronotum short. Mesonotum widely exposed
posteriorly. Proportional length of dorsal abdominal segments
(first to seventh):: 28: $2: 82: 82: 85: 40: 77. Metasternum behind
transverse suture inclined posteriorly, shorter in the middle.
Seventh ventral abdominal segment with posterior margin armed
with a few black spines laterally.
Tue UNiversiry SCIENCE BULLETIN
Rhagovelia oriander Parshley
(Pl. 10, fig. 84)
19z2, Pe ite oriander Parshley, South Dakota State Ent. Tech. Bull., vol.
Pim 9 .
Wingless male: Pronotum with apex obtusely pointed, extending
slightly beyond posterior margin of mesonotum, but posterolateral
portion of the latter well exposed. Proportional length of dorsal
abdominal segments (first to seventh) :: 30: 34: 82: 30: 31: 34: 65.
Metasternum behind transverse suture short, shortest in the middle.
Posterior margin of seventh ventral abdominal segment with a series
of black spines laterally.
Wingless female: Pronotum produced as a thick process, with
weakly developed basal process. Connexivum strongly reflexed and
folded on dorsum for apical four segments, Metasternum behind
transverse suture subequal in length throughout, short.
Rhagovelia rivale Bueno
(Pl. 13, figs. 43a, b, c)
1924. Rhagovelia rivale Torre-Bueno, Trans. Amer. Ent. Soc., vol. 50, p. 247.
Winged male: Pronotum acutely pointed and raised apically.
Forewings fuscous, yellowish along lower margin, veins black, with
two well-defined apical cells. Proportional length of dorsal ab-
dominal segments (first to seventh):: 25: 38: 35: 84: 34: 40: 60.
Second segment with longitudinal carinae roundly divergent pos-
teriorly; anteromesal, anterolateral and posterolateral impressions
separated from each other, but located within the continuous
groove along anterior and lateral margins of second segment;
intersegmental groove between second and third segments short
but rather deep, longitudinally rugose. Third segment with longi-
tudinal carinae divergent posteriorly, anterolateral and postero-
lateral impressions subcontiguous; intersegmental groove between
third and fourth segments deep but short. Metasternum behind
transverse suture shorter at middle, posterior margin sinuate.
Seventh ventral abdominal segment without black spines.
Wingless male: Pronotum with apical margin feebly produced
posteriorly in the middle. Proportional length of dorsal abdominal
segments (first to eighth):: 27: 32: 30: 80: 80: 38: 58: 51. Meta-
sternum behind transverse suture short, subequal in length through-
out.
Winged female: Pronotum with apical process rather short,
strongly dilated apically, clothed with long dark hairs on apical
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 973
margin. Seventh dorsal abdominal segment a little longer in the
middle than basal width; seventh connexival segment with lateral
margin slightly sinuate in apical half, apex acutely pointed and
directed slightly laterad and with a few black, short but stiff hairs,
inner margin bisinuate. Eighth segment broad, apical margin
broadly rounded. The last genital segment directed slightly down-
ward, densely clothed with black hairs. Metasternum behind
transverse suture short, subequal in length throughout.
Wingless female: Posterior margin of pronotum broadly rounded.
Mesonotum broadly exposed behind pronotum. Proportional length
of dorsal abdominal segments (first to eighth) :: 87: 45: 42: 43: 46:
52: 68: 58, Connexivum strongly reflexed and almost folded on
the dorsum after fourth segment. Metasternum behind transverse
suture subequal in length throughout except for near leg base
where it is a little longer.
Group CHARACTERISTICS OF THE AINSLEI GROUP
All three species belonging to this group were available for the
study of winged forms.
Distinct longitudinal carinae in basal dorsal abdominal segments
always occur on the second and third segments. The carinae also
occur on the fourth segment in male Rh. ainslei. First dorsal
abdominal segment with the median elevated area rather narrow.
Forewings always with two long apical cells formed beyond the
basal half of the wing; subcosta develops as far as apical one fourth
of the wing. Pronotum of winged forms broadly rounded apically,
completely covering the mesonotum beneath, uppersurface of
pronotum with median longitudinal ridge running throughout the
entire length and with rather large impressions scattered on the
surface.
The following characters are also peculiar to this group.
(1) The dorsum of the abdomen is narrowed after the first
three segments.
(2) The seventh connexival segment in female forms is a small
triangular cell between the lateral margins of eighth segment.
(3) The eighth segment in the female has a bundle of black
setae directed more or less downward.
(4) The male in both winged and wingless forms is strongly
depressed in the basal ventral abdominal segments.
(5) The seventh ventral abdominal segment with a series of
minute black spines directed inwardly.
Tue Universiry SCIENCE BULLETIN
974
(6) The intermediate femur of the female is dorsoventrally
flattened.
(7) The intermediate tibia is less than twice as long as the sec-
ond tarsal segment.
(8) The second and third tarsal segments in the intermediate
legs are equal or subequal to each other in length.
(9) The hind femur is longer than the tibia, except in Rh. ainslei.
(10) The terminal genital segment is prolonged into a sharply
pointed process in both sexes,
Key To THE WINGED ForMs OF THE SPECIES OF 'THE AINSLEI GRouP
1. Hind femur shorter than tibia. Distinct longitudinal carinae occur on
the fourth segment at least in male. Apex of seventh connexival seg-
tient tn feniale rounded: te. we ee Rh. ainslei
WV’. Hind femur longer than tibia. Without distinct longitudinal carinae on
the fourth segment. Apex of seventh connexival segment in female
PCULEIY DOWER, rors err a nd crc ork Gee et ecm 2
2. Pronotum in female with the apical process vertically erected at base.
Longitudinal carinae on the basal abdominal segments convergent
posteriorly. Seventh connexival segment in female reflexed and pro-
duced above the eighth segment.......... 000i. in Rh. becki
2’. Pronotum in female with the apical process subhorizontally produced
posteriorly. Longitudinal carinae divergent posteriorly. Seventh
connexival segment not reflexed............. 00. ..05 Rh. gracilis
Rhagovelia becki Drake and Harris
(Pl. 14, figs. 47a, b, c)
1936. Rhagovelia becki Drake and Harris, Proc. Biol. Soc. Washington, vol. 49,
p. 106.
Winged female: Pronotum with a black low longitudinal ridge
running from anterior margin to apex, apical processlike projection
almost vertically erected at base, then bent posteriorly at about
forty-five degrees, extreme apex narrowly rounded. Forewings
fuscous, with greenish tinge at base, veins darker. Proportional
length of dorsal abdominal segments (second to eighth):: 42: 49:
57: 65: 72: 78: 54. Second dorsal abdominal segment with longi-
tudinal carinae divergent at base, then slightly convergent pos-
teriorly as far as posterior margin, obscurely grooved and with
obscure longitudinal elevation along median elevated portion of
first segment; anteromesal impression obliquely placed along longi-
tudinal carinae at base, anterolateral and posterolateral impressions
clearly separated from each other. Third dorsal abdominal seg-
ment with longitudinal carinae slightly convergent posteriorly;
intersegmental groove between second and third segments obscurely
longitudinally rugulose; anteromesal and anterolateral impressions
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 975
contiguous, posterolateral impression obliterated. Fifth and sixth
segments longer than wide. Seventh segment with posterior margin
feebly sinuate; seventh connexival segment narrowed apically, re-
flexed and produced above eighth segment, apex acutely pointed
and with short black hairs. Eighth dorsal abdominal segment with
apical margin broadly rounded, with a conspicuous bundle of black
setae arising ventrally. The last genital segment rather acutely
pointed at apex, densely clothed with short black hairs. Meta-
sternum behind transverse suture subequal in length throughout.
Wingless female: Pronotum with apex broadly rounded, without
longitudinal ridge. Connexivum subvertically reflexed throughout.
Proportional length of dorsal abdominal segments (first to eighth) ::
38: 45: 45: 50: 65: 76: 92: 58. Posterior margin of seventh dorsal
abdominal segment densely clothed with long and black setae.
Metasternum as in winged female.
Winged male: Apical process of pronotum as in winged female,
but much less strongly developed. Connexivum strongly reflexed.
Eighth dorsal abdominal segment thick and long, densely clothed
with white hairs, subparallel-sided, slightly dilated apically. Meta-
sternum behind transverse suture and second ventral abdominal
segment depressed and with median longitudinal elevation, pos-
terior margin of metasternum subangularly emarginated in the
middle.
Wingless female: Pronotum with apex rounded, completely cov-
ers mesonotum. Proportional length of dorsal abdominal segments
(first to seventh):: 35: 88: 87: 87: 42: 53: 83. Seventh ventral
abdominal segment with posterior margin armed with minute
black spines directed inwardly.
Rhagovelia gracilis Bacon
(Pl. 14, figs. 46a, b, c)
1956. Rhagovelia gracilis Bacon, Univ. Kansas Sci. Bull., vol. 38, pt. I, p. 861.
Winged female: Pronotum with a low black ridge from anterior
margin to apex; apex developed into a long simple process clothed
with long black hairs. Forewings with two distinct long closed
cells, fuscous, veins darker. Proportional length of dorsal abdom-
inal segments (fourth to eighth):; 42: 56: 55: 63: 42. First seg-
ment with median elevation rather narrow. Second segment with
longitudinal carinae well marked, roundly strongly divergent pos-
teriorly; anteromesal impression obliterated, anterolateral and
posterolateral impressions separated; intersegmental groove between
Tue Universiry Science BULLETIN
second and third segments well defined but rather shallow. Third
segment with longitudinal carinae strongly divergent posteriorly;
anteromesal impression obliterated, anterolateral and posterolateral
impressions reduced and continuous; intersegmental groove be-
tween third and fourth segments ill defined posteriorly. All dorsal
abdominal segments transverse. Seventh connexival segment
strongly narrowed in apical half, inner margin sinuate, apex acutely
pointed and reflexed with short black hairs. Eighth segment
with paratergite ill defined posteriorly, posterolateral margin of
eighth dorsal abdominal segment broadly rounded, with a bundle
of long, black bristles directed downward. The last genital seg-
ment subtriangular, clothed with fine long hairs. Metasternum
behind transverse suture short, slightly lengthened laterally.
Wingless female: Pronotum with a black longitudinal low ridge
reaching near to apex which completely covers mesonotum. Pro-
portional length of dorsal abdominal segments (first to eighth)::
40: 42: 41: 41: 56: 71: 92: 53. Metasternum behind transverse
suture lengthened laterally. Connexivum strongly reflexed and
folded on the dorsum in the middle.
Winged male: Pronotum with apex acutely pointed and slightly
produced posteriorly. Proportional length of dorsal abdominal seg-
ments (first to seventh):: 22: 32: 35: 40: 42: 45: 70. First to sixth
dorsal abdominal segments transverse. Metasternum behind trans-
verse suture and basal two ventral abdominal segments depressed
and with median longitudinal carinae. Seventh ventral abdominal
segment with a series of black minute spines directed inward at
sides.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 30: 34: 86: 36: 43: 51: 80 (holotype). Meta-
sternum behind transverse suture and two basal ventral abdominal
segments strongly depressed, with distinct median longitudinal
elevation.
Rhagovelia ainslei Drake and Harris
(Pl, 14, figs. 45a, b, c)
1983. Rhagovelia ainslei Drake and Harris, Proc. Biol. Soc. Washington, vol.
46, p. 50.
Winged male: Pronotum with distinct black median longitudinal
ridge running throughout the entire length of pronotum, apical
process weakly developed. Forewings with two distinct closed
cells, fuscous; veins darker. Proportional length of dorsal abdom-
inal segments (second to seventh):: 85: 35; 38: 38: 43: 65. Con-
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 977
nexivum vertically erected throughout. First dorsal abdominal seg-
ment with median elevated area well-defined, but comparatively
narrow. Second dorsal abdominal segment with longitudinal cari-
nae thick, rather strongly divergent at base; anteromesal impression
small, anterolateral and posterolateral impressions contiguous; in-
tersegmental groove between second and third segments shallow.
Third segment with longitudinal carinae roundly divergent in ante-
rior half then converge posteriorly; anteromesal impression distinct,
anterolateral and posterolateral impressions contiguous; interseg-
mental groove between third and fourth segments distinct and well
defined. Fourth segment with distinct though less well marked
longitudinal carinae almost reaching to posterior margin of the
same segment, this carina is not developed in one female examined.
Eighth segment thick and subparallel. Seventh ventral abdominal
segment with posterior margin armed with a series of minute black
spines directed inward at sides.
Wingless male: Proportional length of dorsal abdominal segments
(first to seventh):: 28: 31: 25: 80: 31: 40: 65. Connexivum ver-
tically reflexed. Anterior margin of first dorsal abdominal segment
straight. Eighth segment thick, parallel-sided. Metasternum be-
hind transverse suture subvertically inclined posteriorly, shortened
in the middle. Second to fourth segments depressed and with me-
dian longitudinal ridge.
Winged female: Pronotum with a black median longitudinal
ridge throughout the entire length, apex strongly developed sub-
horizontally as a long process and clothed with long, dark hairs.
Forewings fuscous, veins darker. Connexivum less strongly re-
flexed than in male. Seventh dorsal abdominal segment a little
wider at base than it is long in the middle; seventh connexival seg-
ment slightly narrowed apically, posterior margin broadly rounded,
forming a small transverse, subtriangular plate between lateral mar-
gin of eighth segment. Eighth connexival segment with inner mar-
gin obliterated posteriorly, with a bundle of black setae arising
ventrally. Metasternum behind transverse suture plane, shortened
medially.
Group CHARACTERISTICS OF THE SPINIGERA GROUP
All four species treated by Bacon were available for the study of
Winged forms.
Pronotum in winged male acute at tip, strongly produced as proc-
€ss apically in winged female.
32—38378
978
Tue Universiry SCIENCE BULLETIN
Forewings always dark fuscous and with greenish tinge in basal
half, with two large apical cells formed in distal half.
Longitudinal carinae always occur on second and third ab-
dominal segments; the carinae are also poorly developed on the
fourth segment (Rh. spinigera and Rh. ignota). First dorsal ab-
dominal segment with the median elevated area well defined both
laterally and posteriorly. Second segment always with well-defined
laevigate depression along the posterior margin of first segment;
posterolateral laevigate impression is clearly separated from antero-
lateral impression.
Proportional length between sixth and seventh dorsal abdominal
segments in winged male ranges from 42:57 in Rh. choreutes to 42:
67 in Rh. ignota, that in wingless male always about 4:7. Propor-
tional length between seventh and eighth dorsal abdominal seg-
ments in winged female is 56:52 in Rh. ignota and 60:44 in Rh.
formosa, that in wingless female is 63:52 in Rh, ignota and 77:42 in
Rh. spinigera.
Other group characters are as follows:
(1) The intermediate femur of female is transversely constricted
at middle.
(2) The dorsum of abdomen of the wingless female is narrow
after the first three segments. The connexivum is strongly reflexed
for the last four segments.
(3) The pronotum in wingless forms completely covers mesono-
tum.
(4) The last three abdominal segments in the female have con-
spicuous bundles of black hairs except in Rh. spinigera.
(5) The second antennal segment is considerably longer than the
third one.
(6) In the intermediate leg tibia is more than twice as long as
the second tarsal segment; third tarsal segment is distinctly longer
than the second one.
(7) In hind leg, the femur is longer than the tibia except in Rh.
choreutes.
(8) Abundance of winged forms.
Key Tro THE SPECIES OF THE SpINIGERA Group ON THE Basis OF THE
WinceD FEMALE
1. Eighth segment on ventral surface strongly longitudinally elevated in
Uinfcranatlols lt putghat rs (0 Ng ane AsG tga BU EAL igdaniy Soe tule agli er tet Rh. ignota
1. Eighth segment not elevated longitudinally in the middle of ventral
SUntahe-Te oy i ie oe eerie ods vs ee een Orde 2
2. Apical margin of seventh connexival segment and lateral margin of
Rh. formosa
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA 979
2. Apical margin of seventh connexival segment not continuously sinuate
with lateral margin of eighth segment..i,........6 4. iF
SLIT
\) Y
33a R.(R.) horrida
34 R.(R.) oriander
35a R.(R.) nitida
THE University Science BULLETIN
PLATE 11
Dorsau View or ABDOMINAL SEGMENTS AND LATERAL Virw OF PRONOTUM
Fic, 36a. Basal abdominal segments of Rhagovelia (Rhagovelia) tayloriella
Kirkaldy; 36b, apical portion of pronotum; 86c, apical abdominal segments of
winged female.
Fic. 87a. Basal abdominal segments of Rhagovelia (Rhagovelia) collaris
(Burmeister); 87b, apical portion of pronotum; 37c, apical abdominal segments
of winged female.
Fic. 38a. Basal abdominal segments of Rhagovelia (Rhagovelia) impensa
Bacon; 38b, apical portion of pronotum; 38c, apical abdominal segments of
winged female.
SUPPLEMENTARY STUDY OF THE Genus RHAGOVELIA
PLATE 11
37a R.(R.) collaris
83—3378
1009
Tue Universiry SCIENCE BULLETIN
PLATE 12
DorsaL Virw or ABDOMINAL SEGMENTS AND LATERAL View OF PRONOTUM
Fic. 39a. Basal abdominal segments of Rhagovelia (Rhagovelia) armata
(Burmeister); 39b, apical portion of pronotum; 39c, apical abdominal segments
of winged female.
Fic. 40a. Basal abdominal segments of Rhagovelia (Rhagovelia) acuminata
Bacon; 40b, apical portion of pronotum; 41c, apical abdominal segments of
winged female.
Fic. 41a. Basal abdominal segments of Rhagovelia (Rhagovelia) planipes
Gould; 41b, apical portion of pronotum; 41c, apical abdominal segments of
winged female. :
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA
PLATE 12
mis
39a R.(R,) armata
41a R.(R.) planipes
1011
Tue Universiry Science BULLETIN
PLATE 13
Dorsau View or ABDOMINAL SEGMENTS AND LATERAL VrEw OF PRONOTUM
Fic. 42a. Basal abdominal segments of Rhagovelia (Rhagovelia) distincta
Champion; 42b, apical portion of pronotum; 42c, apical abdominal segments of
winged female.
Fic, 48a. Basal abdominal segments of Rhagovelia (Rhagovelia) rivale
Bueno; 43b, apical portion of pronotum; 43c, apical abdominal segments of
winged female.
Fic. 44a. Basal abdominal segments of Rhagovelia (Rhagovelia) obesa
Uhler; 44b, apical portion of pronotum; 44c, apical abdominal segments of
winged female.
SUPPLEMENTARY StupyY OF THE GENUS RHaAcovEeLIA 1018
PLATE 18
ET ge)
42a R.(R.) distincta
. OTS A
Nec
A3a R.(R.) rivale
Aha R.(R.) obesa
83—8378
Tue Universiry Science BULLETIN
PLATE 14
Dorsau View or ABDOMINAL SEGMENTS AND LATERAL View or PRONOTUM
Fic. 45a. Basal abdominal segments of Rhagovelia (Rhagovelia) ainslei Drake
and Harris; 45b, apical portion of pronotum; 45c, apical abdominal segments of
winged female.
Fic. 46a. Basal abdominal segments of Rhagovelia (Rhagovelia) gracilis
Bacon; 46b, apical portion of pronotum; 46c, apical abdominal segments of
winged female. ‘
Fic. 47a. Basal abdominal segments of Rhagovelia (Rhagovelia) becki
Drake and Harris; 47b, apical portion of pronotum; 47c, apical abdominal seg-
ments,
SUPPLEMENTARY STUDY OF THE GENUS
PLATE 14
45a Re(Ro) ainslei
Ava R.(R.) becki
RHAGOVELIA
1015
1016 Tur Unriversiry SCrENCE BULLETIN
PLATE 15
Dorsau View or ABDOMINAL SEGMENTS AND LATERAL View or PRONOTUM
Fic. 48a. Basal abdominal segments of Rhagovelia (Rhagovelia) formosa
Bacon; 48b, apical portion of pronotum; 48c, apical abdominal segments of
winged female.
Fic. 49a. Basal abdominal segments of Rhagovelia (Rhagovelia) ignota
Drake and Harris; 49b, apical portion of pronotum; 49c, apical abdominal seg-
ments of winged female.
Fic, 50a. Basal abdominal segments of Rhagovelia (Rhagovelia) choreutes
Hussey; 50b, apical portion of pronotum; 50c, apical abdominal segments of
winged female.
Fic. 51a. Basal abdominal segments of Rhagovelia (Rhagovelia) spinigera
Champion; apical portion of pronotum; 51c, apical abdominal segments of
winged female.
SUPPLEMENTARY STUDY OF THE GENUS RHAGOVELIA
PLATE 15
51a R.(R.) spinigera
1017
THE UNIVERSITY OF KANSAS
SCIENCE BULLETIN
Vou. XXXVIII, Pr. 1] DrcEMBER 20, 1956 [No. 12
Concerning the Relationship of Certain Neotropical Gek-
konid Lizard Genera, with Comments on the Micro-
scopical Structure of Their Glandular Scales
BY
Epwarp H. Taytor and A. Byron LEONARD
Department of Zoology, University of Kansas
Axsstract: The classification of the gekkonid lizards is reviewed briefly.
The recent family classification of the gekkonid lizards by Garth Underwood
is discussed and followed as far as Western Hemisphere groups are concerned.
The absence of femoral pores in all endemic species of the Gekkonidae in the
Western Hemisphere is reported. The genera Peropus and Hemidactylus
are regarded as transported here by man. Specialized glandular scales on the
posterior ventral and subfemoral regions are discussed and a_ preliminary
microscopical study of the scales is offered. The name “escutcheon scales,”
proposed for these by Chapman Grant, is used. It is believed that these scutes
may represent the ancestral type of the femoral and preanal pores.
For a considerable time the gekkonid lizards have been of much
interest to the senior author who in his field exploration in various
parts of the world has collected representatives of some eighty
forms of these interesting saurians, several of which have been
described as new. His recent review of the lizard fauna of Costa
Rica has again opened the problem of the classification of the gek-
konid lizards and he found it necessary to make a cursory review
of those groups occurring in the Western Hemisphere, and to make
an attempt to re-evaluate the rank of the groups represented.
In the latter half of the 19th century the gekkonid lizards were
regarded as belonging to three families: the Uroplatidae, Euble-
pharidae and the Gekkonidae, This arrangement was used by
Boulenger in his British Museum Catalogue and by Cope in his
work on American Herpetology. It was likewise followed by
a majority of the herpetologists of their time.
Fiirbringer (1900) published a classification of the lizards recog-
nizing three families. However he placed the Uroplatidae in a
(1019)
1020 THe Universitry Science BULLETIN
“gens” different from the other two. The following year Gadow
(1901) considered all the gekkonid lizards under the suborder Geck-
ones placing them in a single family, Geckonidae, with three sub-
families—Uroplatinae, Eublepharinae, and Geckoninae. After a time
there was a tendency on the part of herpetologists to follow this ar-
rangement, but not without some variation. Thus Charles Camp
(1923) in his scholarly contribution on classification, reviewed the
Sauria, recognizing the Uroplatidae as a distinct family but treating
the Eublepharidae with the Gekkonidae.
One of the troublesome groups formerly associated with the
Gekkonidae is one comprising certain small Central and South
American saurians including the genera Sphaerodactylus, Gona-
todes, Pseudogonatodes, Coleodactylus and Lepidoblepharis. More-
over, one cause for this trouble was that certain forms in South
Asia (of different relationship ) were placed in the genus Gonatodes
erroneously, so that one had in a single genus both amphicoelous
and procoelous forms, thus rendering the character of the vertebrae
useless for the separation of families.
G. K. Noble (1921) in his study of Sphaerodactylus and allied
genera noted the characteristics of the group in Central and South
America, He found it necessary to remove this group of genera
from the Gekkonidae proper and associate it with the Euble-
pharidae. Moreover, he did not include amphicoelous Asiatic
forms in Gonatodes. Werner (1912) already had referred Lepi-
doblepharis to the Eublepharidae. Noble was convinced that these
genera of small saurians represented a natural group. Concerning
the vertebrae he states, “The procoelous vertebrae have been de-
veloped in this series quite independently of similar changes in
any other series.” Despite numerous other differences he associated
them with the Eublepharidae chiefly on the basis of this one char-
acter,
H. W. Parker (1926) treated this same group, adding to it an-
other genus, Coleodactylus. He subscribed to the Gadow classifi-
cation and all were placed in the Gekkonidae. The Zoological
Record of 1926 did not use the family name Eublepharidae and it
did not appear in the American Checklist of Stejneger and Barbour
of 1923.
Smith and Taylor in their Mexican checklist (1946) reluctantly
followed this usage although it was apparent to both authors that
the arguments were equally as strong for the retention as for the
abandonment of the family. Since they were not treating species
NeotrropicaL GekKoniw LizaARD GENERA 1021
of the Uroplatidae the status of that family was not their immediate
concern. As for the group delineated by Noble, his opinion as to
their relationship was accepted. The group entered Mexican terri-
tory only in the southern part and two diminutive species of
Sphaerodactylus were the only ones available to them in their col-
lections.
To the senior author in his studies of the Costa Rican fauna, the
group was more important since Costa Rica has representatives of
three genera and several species and the general characteristics of
the group were examined. It was noted that, aside from the pro-
coelous vertebrae, the paired parictals, sternum characters, and
the sternum-rib relationship pointed out by Noble, they lacked a
voice mechanism. The “clutch” consisted of a single egg at one
laying. Neither postanal sacs nor postanal bones were present in
either sex. It was noted that the males and females had no femoral
or preanal pores but that there were present in males a series of
specialized glandular scales in the posterior region of the venter.
In some species these also appeared in the femoral region, sug-
gesting their possible analogy to the true femoral and preanal pores.
He began a hasty search of the literature to ascertain whether
these characters had been described and was informed by his stu-
dent, Mr. Peter Chrapliwy, of the description by Maj. Chapman
Grant (1931) in a paper dealing, with West Indian members of the
genus Sphacrodactylus, and of the work by Noble and Klingel.
Grant who seemingly was the first to mention them in literature had
given the name of “escutcheon scales” to this group of glandular
scales. Noble and Klingel seem to have been the first and perhaps
the only subsequent workers to give consideration to these scales.
Aside from noting the escutcheon scales on certain species they
made a preliminary study of the microscopical structure ( Noble
and Klingel, 1932).
From the first we were convinced that the group unquestionably
merited at least subfamily recognition and proposed to deal with it
in this category. Shortly after, we received a paper by Garth
Underwood (Nov., 1954) * dealing on an extensive scale with the
anatomy and classification of the Gekkoes.
While it is somewhat futile to report our own proposals for
the arrangement of the American gekkonid lizards, they were as
follows:
* This paper was first received by us in May, 1955. 250.
NeorropicaAL Gexxonip Lizarp GENERA 1027
Within the depressed portions of the specialized scales, however,
another germinative epithelium (fig. 1, d; fig. 2, 7) composed of
large faintly staining cuboidal to rounded cells lies in direct con-
tact with the cornified cells of the basal epithelium. This second
germinative epithelium produces large, polygonal to depressed
rhomboidal cells containing dense accumulations of coarse granules
(fig. 1, c; fig. 2, 1) which are presumed to be secretory in nature.
For this reason this layer is here referred to as the “secretory”
epithelium. The cells of this layer fill the cuplike depression of
the scale, and often produce a distinct convexity on the outermost
surface. The granules of these cells stain faintly with Celestine
Blue, but with Mallory’s Triple Stain they become deeply pig-
mented (fig. 2, i). There is but little modification among these
granular cells, except a slight degree of depression near the outer
surface of the layer.
A third germinative epithelium rests on the granular cells of the
secretory epithelium (fig. 1, b; fig. 2, h); these cells are cuboidal to
rounded, and seem never to be columnar. They give rise to a layer
not more than three to five cells in thickness which quickly be-
comes cornified and squamose; we refer to this outermost non-
pigmented epithelium as the capping epithelium. This varies in
thickness, but everywhere presents the same general characteristics
and in preserved specimens as stated may be removed by rubbing
a finger over the surface.
The arrangement of three distinctive superimposed epithelia
in these scales not only provides a unique morphological feature
that serves to distinguish these lizards from others known to us,
but raises several intriguing biological problems. For example,
how these superimposed epithelia are nourished, situated as they
are above one or more layers of cornified cells of presumed im-
permeability, and certainly remote from the circulatory supply that
lies below the basal epithelium, presents a difficult problem. Fur-
thermore, the function of the secretory epithelium is unknown
to us. It seems obvious that these cells pass through a cycle of
biochemical stages, typical of secretory cells (compare fig. 1, ¢
with fig. 2, i) since in our material it can be observed that the gran-
ules become larger as the cells enlarge; in addition, the staining
properties of the cells when treated with Mallory’s stain is not con-
sistent; some cells stain deeply with fuchsin, others with aniline blue,
and certain cells contain granules some of which take the blue
stain, others the red. Still another problem is raised by the ob-
servation that the germinative layer of the secretory epithelium
|
|
|
|
'
|
i
i
Gy
t
|
i
|
i
1028 Tue Universirry Scrence BULLETIN
and that of the capping epithelium are frequently out of phase;
that is, when one seems active and well developed, the other seems
less so (compare fig. 1 with fig. 2 with reference to the germinative
layers). In some scales the capping layer seems to have desqua-
mated to the point of nearly exposing the underlying secretory
cells (fig. 1) but whether or not the secretory cells actually be-
come exposed at the surface is not revealed by any of our material.
Unfortunately for present purposes, all the material available to
us was collected by the senior author in a single season (summer )
of the year, so that presumed cyclic changes in the specialized
scales cannot be determined at this time.
The relationship of these scales to the preanal and femoral pores
is uncertain.* Their position on the body in the same general areas
as the pores suggests that they may subserve the same general pur-
pose. One might suspect that they represent a primitive stage in
pore development. The femoral and preanal pores themselves are
of several kinds and may be polyphyletic in origin. Further study
is necessary to ascertain the degree of relationship.
LITERATURE CITED
BouLENcER, Grorcr A.
1885. Catalogue of the lizards in the British Museum (Natural History )
Qnd ed. vol. 1, Geckonidaec, Eublepharidae, Uroplatidae, Pygo-
podidae, Agamidae. 1885, pp. I-XII; 1-486, pls. 1-32.
Camp, Cuaries Lewis
1923. Classification of the lizards. Bull. Amer. Mus. Nat. Hist., vol. 48,
art. XI, pp. 289-481, figs. 1-112.
Furerincer, M.
1900. Zur vergleichenden Anatomie des Brustschulterapparates und der
Schultermuskeln. Jenaische Zeit., Bd. 84, pp. 215-718, pls. 18-17.
Gapow, Hans
1923. Amphibia and Reptilia (The Cambridge Natural History, vol. 8)
1928, pp. L-XII, 1-668, figs. 1-181.
Garpner, Lynn W. and Grant, CHapMAN
1940. The herpetology of Jamaica, pt. 1, Amphibians; pt. 2, The Reptiles,
Bull. Inst. Jamaica, Sci. Ser., no. 1, 1940, pp. 1-148.
GRANT, CHAPMAN
1931. The sphaerodactyls of Porto Rico, Culebra, and Mona Island.
Jour. Dept. Agri. Porto Rico, vol. 15, no. 8, July, 1931, pp. 199-218,
pls, 20-24 (first report on the “escutcheon” group of specialized
scales).
. * The preanal pores on the gekkoes of the genera Naulti Hoplodactyl Racho-
dactylus, and perhaps others are arranged in patches or in several rows, The number
is excessive and may reach 80 or more in one or more species, At the moment none of
these lizards are available to us for examination, It is possible that these may throw some
light on the relationship of the escutcheon scales to true femoral and preanal pores,
NerotropicaAL Grexxkonip Lizarp GENERA 1029
1982. A new sphaerodactyl from Porto Rico. Jour. Dept. Agri. Puerto
Rico, vol. 16, no. 1, Jan. 1932, p. 81.
1936. Herpetological notes with new species from the American and
British Virgin Islands. 1936. Jour. Agri. Univ. Puerto Rico,
vol. 21, no. 4, Oct. 1987, pp. 508-522.
Nose, G. K.
1940. The bony structure and phyletic relations of Sphaerodactylus and
allied lacertilian genera, with the description of a new genus.
Amer. Mus. Novitates, no. 4, pp. 1-16.
1940. The herpetology of the Cayman Islands, Bull. Inst. Jamaica, Sci.
Ser. no, 2, 1940, pp. 1-56.
Description of “escutcheon scales’
dactylus.
Nosue, G. K., and Hasster, W. G.
1933. Two new species of frogs; five new species and a new race of lizard
from the Dominican Republic. Amer. Mus. Novitates, no, 652, Aug.
14, 1988, pp. 1-17.
Nose, G. K., and Kurcst, G. C.
1932. The reptiles of Great Inagua Island, British West Indies. Amer.
Mus. Novitates, no. 549, Aug. 11, 1982, pp. 1-25.
Discusses Grant’s discovery of escutcheon scales and gives data
on several species, and a brief description of their microscopic char-
>
in three species of Sphaero-
acter.
Parker, H. W.
1926. The neotropical lizards of the genera Lepidoblepharis, Pseudo-
gonatodes, Lathrogecko, and Sphaerodactylus, with a description
of a new genus. Ann. Mag. Nat. Hist., ser. 9, vol. 17, Mar. 1926,
pp. 291-801.
Smrra, Hoparr M., and Taytor, Envwarp H.
1950. An annotated checklist and key to the reptiles of Mexico exclusive
of the snakes, U. S. National Mus. Bull. 199, 1950, pp. I-IV,
1-258,
Smiru, MALCOLM
1933. Remarks on some Old World geckoes. Rec. Ind. Mus., vol. 85,
pp. 9-19.
1985. The fauna of British India including Ceylon and Burma ‘
Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. I-XII; 1-440, pl. 1,
figs. 1-94,
Taytor, Epwarp Harrison
1956. A review of the lizards of Costa Rica. Univ. Kansas Sci. Bull.,
vol. 88, pt. 1, pp. 1-322.
Treats of Sphaerodactylidae in Costa Rica.
WerRNER, FRANZ
1912. Reptilia, Lacertilia (EHublepharidae, Uroplatidae, Pygopodidae).
Das Tierreich, Lief, 33, pp. 1-83.
Publications of the University of Kansas Science Bulletin
Kansas University Quarterly
Volumes I to X are no longer available for distribution, since the supply is either ex-
*hausted or the numbers so reduced that they are kept for distribution in whole sets to libraries.
University of Kansas Science Bulletin
Volume
ee Nos. 1-4, weight 8 ounces. Nos. 5-9, weight, 12 ounces. Nos. 10-12,
weight, 6 ounces.
as Nos. 1-8, weight, 20 ounces. Nos. 4-9, weight, 11 ounces. Nos. 10-15,
weight, 20 ounces. :
Tc Nos. 1-6, weight, 83 ounces. Nos. 7-10, weight 25 ounces.
Wea es Nos. 1-6, weight, 83 ounces. Nos, 7-20, weight, 28 ounces.
Vas ce Nos, 1-11, weight, 83 ounces. Nos. 12-21, weight, 27 ounces.
Viv No. 1, weight, 27 ounces. Nos. 2-7, weight, 19 ounces.
VIPs Nos. 1-17, weight, 50 ounces.
Ville. Nos. 1-10, weight, 52 ounces.
dre Nos. 1-21, weight, 54 ounces.
a Nos, 1-15, weight, 17 ounces.
a. No. 1, weight, 20 ounces.
Oe Nos. 1-2, weight, 19 ounces.
LED is ee Pt. I, Nos. 1-9, weight, 12 ounces. Pt. II, Nos. 10-15, weight, 10 ounces.
ev Nos. 1-21, weight, 34 ounces.
XV......Nos. 1-6, weight, 18 ounces.
RV oak Nos, 1-6, weight, 14 ounces,
Vile cae Pt. I, No. 1, weight, 18 ounces. Pt. II, Nos. 2-7, weight, 8 ounces.
VELL id Nos, 1-18, weight, 38 ounces,
MIRA, Pt. I, Nos, 1-7, weight, 6 ounces. Pt. II, Nos, 8-14, weight 16 ounces,
DD Gna Pt. I, Nos. 1-6, weight, 11 ounces, Pt. II, Nos, 7-21, weight, 15 ounces.
Sale as Nos. 1-16, weight, 32 ounces,
Dei is Nos. 1-18, weight, 32 ounces.
Mees cas No, 1, weight, 40 ounces.
Mal Ve res Nos, 1-21, weight, 38 ounces.
MV es Nos. 1-22, weight, 43 ounces.
Vee Nos. 1-15, weight, 40 ounces.
Dor Vk Pt. I, weight, 20 ounces.
WAV. Pt. I, weight, 20 ounces. Pt. II, weight, 20 ounces,
XXIX......Pt. I, weight, 20 ounces, Pt. Il, weight, 20 ounces.
>, a Pt. I, weight, 15 ounces. Pt. II, weight, 17 ounces,
AA a Pt. I, weight, 15 ounces. Pt. II, weight, 24 ounces.
P.O, e, 4 1a Issued as a single volume, weight, 45 ounces.
XR Ot Pt. I, weight, 15 ounces. Pt. II, weight, 20 ounces,
MLV its Pt. I, weight, 80 ounces. Pt. II, weight, 25 ounces,
AAV Pt. I, weight, 50 ounces. Pt, II, weight, 50 ounces. Pt. III, weight 20
ounces,
XXXVI......Pt. I, weight, 50 ounces. Pt. II, weight, 50 ounces.
WV Pt. I, weight, 50 ounces, Pt. II, weight, 50 ounces,
uu
Sey CY
§9)
+e ge oe pe
+