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SMITHSONIAN IN STITUTION
UNITED STATES NATIONAL MUSEUM
PROCEEDINGS
OF THE
UNITED STATES NATIONAL MUSEUM
VOLUME 114
NUMBERS 3467-3475
UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON : 1964
Publications of the United States National Museum
The scientific publications of the United States National Museum include two
series, Proceedings of the United States National Museum and United States National
Museum Bulletin.
In these series are published original articles and monographs dealing with the
collections and work of the Museum and setting forth newly acquired facts in the
fields of anthropology, biology, geology, history, and technology. Copies of each
publication are distributed to libraries and scientific organizations and to special-
ists and others interested in the various subjects.
The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with the
publication date of each paper recorded in the table of contents of the volume.
In the Bulletin series, the first of which was issued in 1875, appear longer,
separate publications consisting of monographs (occasionally in several parts)
and volumes in which are collected works on related subjects. Bulletins are
either octavo or quarto in size, depending on the needs of the presentation.
Since 1902 papers relating to the botanical collections of the Museum have been
published in the Bulletin series under the heading Contributions from the United
States National Herbarium.
Frank A. Taytor,
Director, United States National Museum.
CONTENTS
Pages
Bonp, Gorman M. Geographic variation in the thrush 3
Hylocichla ustulata. One figure. No. 3471, published
March, 6,-1963.3¢ . -.o-- . 373-387
DreisspacH, R. R. New Eneeien e Epica) ae a
Auplopus, from the Americas south of the United States
(Hymenoptera: Psammocharidae). Thirteen plates. No.
3468, published March 19,:1963 >. warcenousi> > myedagina 20-211
New species: Auplopus shannoni, A. semialatus, A. splendens,
A. femurrubrus, A. villosus, A. nebulosus, A. opacus, A. minus,
A. zeteki, A. panamensis, A. sapphirus, A. exilis, A. earinus,
A. grossus, A. guatemalensis, A. aquilus, A. incognitus, A. oli-
varus, A. purpureus, A. violaceus, A. blatteus, A. roseus, A.
nigriculus, A. lineatus, A. quartus, A. venetus, A. minusculus,
A. magnus, A. callainus, A. buscki, A. medius, A. hidalgoensis,
A. fuscus, A. albifrons, A. semirufus, A. abnormalis, A. niger,
A. woodi, A. anthracinus, A. clypeatus, A. aeruginosus, A. argen-
tinus, A. marginalis, A. atratus, A. amoenus, A. vulcanensis, A.
gaumeri, A. kathryni, A. fulgidus, A. dietzi, A. minutus, A.
bequaerti, A. gertschi, A. hondurensis, A. hispidus, A. bermuden-
sis, A. cyaneus, A. ferrugineus, A. argentinensis, A. batesi, A.
argutus, A. pratens, A. aurarius, A. lasios, A. puniceus, A.
alarius, A. striatus, A. eriodes, A. brasiliensis, A. coracinus, A.
schausi, A. stagei, A. malinus, A. cordobensis, A. flavicrus, A.
subaurarius, A. ater, A. pratentis, A. pygidialis, A. editorialis,
A, viridulus.
Furnt, Outver S., Jr. Studies of Neotropical caddis flies, I:
Rhyacophilidae and Glossosomatidae (Trichoptera). Eight
figures. No. 3473, published April 16,1963... . . . 453-478
NEW species: Atopsyche clarkei, A. neolobosa, Mortoniella
apiculata, M. angulata, M. hodgesi, Mexitrichia aries, M. atenu-
ata, M. elongata, Protoptila bicornuta.
Hassroucx, Franx F. Moths of the family Acrolophidae
in America north of Mexico (Microlepidoptera). Two
hundred and nineteen figures. No. 3475, published
terabor tn, FORA tate nical syistet. chsiele Sune moleelsios 20m Cue
New species: Acrolophus acornus, A. bicornutus, A. klotsi, A.
baldufi, A. luriei, A. vanduzeei, A. forbesi, A. juxtatus, A.
chiricahuae, A. sinclairi, A. parvipalpus, A. serratus, A. secu-
latus, A. macrophallus, A. vauriet.
New subspecies: Acrolophus griseus capitatus, A. macrogaster bt-
pectinicornus, A. m. unipectinicornus, A. m. laminicornus, A.
laticapitanus heinrichi, A. 1. clarkei, A. sinclairi sinclairi, A. s.
nelsont.
III
IV CONTENTS
Pages
New combination: Acrolophus griseus griseus, A. macrogaster
macrogaster, A. laticapitanus laticapitanus, A. l. occidens, A. l. o.
form leopardus.
New name: Acrolophus pseudohirsutus.
Horrman, Ricuarp L. A revision of the North American
annelid worms of the genus Cambarincola (Oligochaeta:
Branchiobdellidae). Seventy-nine figures. No3470,
published"March’6,°1 963s 2° INS ae, a oie | eee
New species: Cambarincola ouachita, C. restans, C. mesochorea, C.
holti, C. shoshone, C. virginica, C. osceola, C. ingens, C. fallaz,
C. holostoma, C. heterognatha, C. demissa.
Howven, Henry F., and Cartwricut, Oscar L. Scarab
WI
beetles of the genus Onthophagus Latreille North of Mexico
(Coleoptera: Scarabaeidae). Eleven figures and nine
plates. No. 3467, published January 9,1963 ...... . . 1-185
New species: Onthophagus batesi, O. subtropicus, O. cavernicollis,
O. monticolus, O. alluvius, O. knulli, O. schaefferi, O. arnetti, O.
brown.
New subspecies: Onthophagus polyphemi sparsisetesus, O. orpheus
pseudor pheus.
New combination: Onthophagus strattulus floridanus.
Kissinger, Davin G. Weevils of the genus Maemactes.
One figure. No. 3474, published March 19,1963. . . . 479-486
New species: Maemactes punctatus, M. imitator.
New combination: Maemactes cribratus.
Osraztsov, Nichotas 5S. Some North American moths of
the genus Acleris (Lepidoptera: Tortricidae). Seven
figures and eighteen plates. No. 3469, published May 7,
1963 cn aye ‘
213-270
New species: Acleris macdunnought, A. santacrucis, A. klotst, A.
clarkei, A. capizziana, A. incognita, A. disputabilis.
New subspecies: Acleris emargana blackmoret.
New combination: Acleris ptychogrammos, A. braunana, A.
kearfottana, A. semiannula, A. implexana, A. senescens, A.
walkerana, A. robinsoniana, A. gloverana, A. maximana.
New status: Acleris implexana form gallicolana.
New combination and status: Acleris logiana placidana.
RANDALL, Joun E. Review of the hawkfishes (family
Cirrhitidae). Sixteen plates. No. 3472, published May
28, 1963 . : » Je et Ghats aipcceaciieys 389-451
New genus: [socirrhitus.
New species: Paracirrhites typee, P. xanthus, P. nisus, P. bicolor,
Cirrhitichthys serratus, C. falco.
Proceedings of
the United States
National Museum
SMITHSONIAN INSTITUTION - WASHINGTON, D.C.
Voiume 114 1963 Number 3467
SCARAB BEETLES OF THE GENUS
ONTHOPHAGUS LATREILLE NORTH OF MEXICO
(COLEOPTERA: SCARABAEIDAE)
Henry F. Howpen! and Oscar L. CartwricHrT?
—————
The purpose of this paper is to facilitate the identification of the
species of Onthophagus of the United States and Canada and to pre-
sent information on their habits and life histories. Twenty-three
species and subspecies described from North America, two species
introduced from Europe and Africa, and eleven previously unrecog-
nized species are included. A key to the species, bibliographical
references, complete new descriptions, photographs of both sexes,
and maps of distribution are given. Important nomenclatural
changes are presented.
Acknowledgments
As a staff member of the University of Tennessee, Howden was
encouraged to make four field trips to Florida and Alabama to study
distribution and biology of the group. Later the Entomology
Research Institute of Canada supported three trips to Texas, Arizona,
and Mexico in 1958, 1959, and 1960. The Smithsonian Institution
sent Cartwright, as a staff member of the United States National
Museum, to study types at the Museum of Comparative Zoology at
1 Entomology Research Institute, Canada Department of Agriculture, Ottawa, Canada.
2 Division of Insects, United States National Museum, Smithsonian Institution, Washington, D.C.
1
2 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
Harvard, the Academy of Natural Sciences of Philadelphia, and
Purdue University at Lafayette, Ind. Through the assistance of
the American Philosophical Society, longer field trips were made by
Howden to Texas in 1954 and Arizona in 1956, and by Cartwright to
Arizona in 1956. We are grateful for this support.
We also acknowledge our indebtedness and extend our thanks to
numerous institutions and individuals for the generous loan of speci-
mens. ‘The letters in parentheses in the following list of collections
studied are abbreviations used in the text when citing material; the
name of the curator responsible for the loan of the specimens follows
the abbreviation: Academy of Natural Sciences of Philadelphia
(ansp), J. A. G. Rehn; American Museum of Natural History (amMnu),
Mont A. Cazier; California Academy of Sciences (cas), H. B. Leech ;
Canadian National Collection (cnc), W. J. Brown; Carnegie Museum,
G. E. Wallace; Chicago Natural History Museum (cnum), Henry S.
Dybas; Cornell University (cu), Henry Dietrich; Illinois Natural
History Survey (Inus), M. W. Sanderson; Iowa State College (1s),
J. L. Laffoon; Museum of Comparative Zoology (mcz), P. J. Darling-
ton; Ohio State University (osu), J. N. Knull; Oregon State College
(osc), P. O. Ritcher; San Jose State College, J. Gordon Edwards;
United States National Museum (usnM); University of Arizona (ua),
Floyd Werner; University of California, Davis (ucp), A. T. McClay;
University of Kansas (uxa), R. H. Beamer; University of Michigan
(umich.), T. H. Hubbell; University of Nebraska, W. T. Atyeo; Uni-
versity of Texas (urex), H. J. Reinhard; University of Tennessee
(uT); and Utah State Agricultural College, G. F. Knowlton.
Individuals who kindly loaned material for study from their per-
sonal collections included: B. Benesh, (ss), L. J. Bottimer (xs),
B. K. Dozier, D. K. Duncan, R. J. Frederick (rsF), C. A. Frost,
B. Malkin, G. H. Nelson, F. H. Parker, Mark Robinson, W. Rosen-
berg (wR), G. B. Vogt, R. E. Woodruff (REw), and F. N. Young
(FNY).
We are especially indebted to E. B. Britton of the British Museum
(Natural History) who loaned type material from the Biologia Cen-
trali-Americana specimens, to H. Freude of the Sammlung des Bayer-
ischen Staats in Munich for the loan of specimens from the Harold
and Sturm collections, to K. Delkeskamp of the Zoologische Museum
der Humbolt-Universitit in Berlin for the loan of Harold specimens,
and to A. Villiers of the Muséum National d’Histoire Naturelle in
Paris for comparing specimens with type material from the Harold
and Laporte collections.
Mrs. Anne Howden aided the project in many ways, both in the
field and in the typing of the manuscript. Photographs were taken
by Jack Scott of the Smithsonian staff.
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 3
Review of literature
The genus Onthophagus was established by Latreille in 1802. Since
that time parts of the genus have received considerable attention.
The American species north of Mexico were discussed by Horn in 1875.
In this work he listed only five species; actually he included five species
under one of these names and started much of the confusion of later
authors. In 1881, he described three more. In 1887, Bates described
and listed many of the species occurring in Mexico and in Central and
South America. In Blatchley’s 1910 work on the Coleoptera of
Indiana, eight species and varieties are listed; incorrect placement of
several of these continued and compounded the subsequent confusion.
In 1914, Schaeffer included 18 species in ‘‘A Short Review of the North
American Species of Onthophagus,”’ the most recent comprehensive
paper on the American species north of Mexico. The list of the
Onthophagus of the world published by Boucomont and Gillet in 1927
greatly facilitated the study of the group. Subsequently a number of
new North American species were described by Brown (1927; 1929a;
1929b). These papers were followed by a synopsis of the Mexican and
Central and South American species by Boucomont in 1932.
Boucomont’s paper was intended to be used mainly to identify the
Mexican and South American forms, but the inclusion of many species
from the United States, in both keys and footnotes, made it useful
for all the Americas.
The habits of our North American species are still almost unknown.
Sim, in 1930, discussed the habits of Onthophagus subaeneus (under
the name cribricollis). In 1935, Lindquist mentioned some of the
habits of O. pennsylvanicus and O. alluvius (under anthracinus). 2-2) - Sie oe ae
Punctures of pygidium deere eae Aches Walter Basclle males and
females without any indication of pronotal protuberance. . .. 25
Pygidium apically convex, shining, distinctly punctate; nearly impunc-
tate and alutaceous basally; pronotal punctures usually lacking
distinct margins, fairly uniform in size; Texas and Arizona. . 24
Pygidium almost flat, very shallowly, indistinctly punctate, alutaceous
almost to apex; pronotal punctures with distinct margins, often
appearing annular; small shallow nonsetate secondary punctures
scattered among large punctures; Big Bend region of Texas and
mountains of northeastern Mexico.
monticolus, new species (p. 61)
Posterior half of metasternum medially impunctate or with one or
two shallow punctures; Texas. . . . . alluvius, new species (p. 65)
Posterior half of metasternum medially with a few very large punctures
near midline; Arizona... . . . . knulli, new species (p. 69)
Dull brownish-black species with pronotal punctures generally the same
size, usually all with setae; New Hampshire to Florida and South
Dakota to Texas ... . . .pennsylvanicus Harold (p. 82)
Shining black species with pronotel punctures of two sizes, very small
punctures lacking setae scattered among the large punctures; Vir-
ginia to Texas. ... . . . . Oklahomensis Brown (p. 80)
Large species over 7.5 mm. in STenatht with unicolorus elytra; west of
the Mississippi River from Arkansas to Texas. ........ 27
Species under 7.5 mm, or having spotted or bicolored elytra. . . . 28
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 13
27(26). Pronotum shining green, frontal carina of male extending almost to
eye; female frontal carina wide, gradually elevated from middle
to the rounded ends which drop sharply to surface of head, the ends
nearer eye than middle; in bat caves, Arkansas to Texas.
cavernicollis, new species (p. 32)
Pronotum with at most a greenish cast; male frontal carina extending
not more than one-half distance from middle to eye; female frontal
carina with each side elevated to a sharp tubercle, the tip of each
tubercle nearer the middle than to the eye; Texas.
subtropicus, new species (p. 30)
28(26). Head and pronotum dull black, elytra largely brown speckled with black;
length 6.7 to 9.5 mm.; male with median horn on vertex behind eyes;
female usually with eral prontal hump behind head; southern
Canada and northern United States except Central Plains area.
nuchicornis (Linnaeus) (p. 123)
Charactersnotasabove. .. . Mit ce ak eo
29(28). Pronotum shining between paneer aoe dal ror High Binck py of
usually distinctly punctate with at least apical half shining. . . 30
Pronotum duli brownish black, alutaceous between setigerous punctures;
pygidium shallowly punctate, usually alutaceous at least to apical
third; elytra dull black with scattered small round yellowish spots,
particularly along apical margin; eastern United States to Kansas
and Oklahoma, rarely Texas and Arizona.
tuberculifrons Harold (p. 85)
30(29). Punctures of pronotum lacking tubercles at anterior margins; males
without hornsonthehead..... hist <.oveptstea 1
Punctures of pronotum with small fabercled in rant of each seta; pate
and apex of elytra brownish yellow; head of male with two slender
horns extending upward behind the eyes; Vermont to Florida, west to
Nebraska and Texas.
striatulus striatulus (Palisot de Beauvois) (p. 41)
31(30). Pygidium bicolored, rarely allyellow. .. . ck Surcn ier toe
Pygidium uniformly dark; pronotum and coe enim greenish brown
to black; elytra with a reddish-yellow spot just inside each humerus
and frequently smaller round discal spots scattered elsewhere but
always showing on the fifth interval first; Texas.
schaefferi, new species (p. 88)
32(31). Base of pronotum finely margined only at middle; pronotum black or
brownish black, occasionally greenish black; anterior angles of prono-
tum and frequently lateral and anterior margins yellow; yellow of
elytra often forming stripes; apex of elytra frequently yellow but
usually broken by a dark marginal spot at end of fourth interval;
fifth interval never with discal spots, rarely with base and apex yellow;
AG ZAGNs «iP ctaks . .. . . landolti texanus Schaeffer (p. 91)
Base of Parana eee margined; pronotum green or coppery,
anterior angles not lighter in color; elytra usually with irregular
yellow areas at base and apex and with at least a few small round
spots on disc; Arizona. . ... . . . . hépfneri Harold (p. 95)
33(2). Tubercles of pronotum very conspicuous. Clypeus of males triangularly
produced upward at middle, clypeus of females ae rounded; if not,
elytra DICOloreds. yeu 3) = ; eee ack tea,
Tubercles of pronotum iubderave With Paoderate. vague punctures at
their bases; clypeus of both sexes broadly rounded or slightly emargi-
HC AAE wilt eewind. basher lewhern tisrulo ananes 34
14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
34(33). Surface of pronotum and (or) elytra finely alutaceous. . . . . 35
Surface of pronotum and elytra between tubercles smooth and shine:
in prairie dog burrows; Oklahoma, New Mexico.
eynomysi Brown (p. 57)
35(34). Uniformly brown or black; eyes flat and narrow, 6 to 7 facets wide;
anterior pronotal angles sharply rounded, a cae angles more
broadly rounded, 140° to 145°. . ... shies yah feo
Pronotum greenish, elytra black; eyes Toucoab le convex, 10 facets
wide; anterior pronotal aire broadly rounded, posterior angles
more sharply rounded, 130°. Length 6.6 to 8.6 mm.; southern
Arizonas “= . . . arnetti, new aoecien (p. 98)
36(385). Pronotum of rate! mith flat rojiecting protuberance, its wide shallow,
usually angular emarginate anterior edge wider than base, and its
external angles rounded; females with carina of vertex distinctly
bent posteriorly at middle; female pronotal protuberance distinct,
sharply defined; western Texas to Arizona.
browni, new species (p. 101)
Head of male with two upright diverging slender horns in front of the
high angulate anterior margin of pronotum; females with carina of
vertex nearly straight; thoracic protuberance of female very weak,
poorly defined; Texas, Colorado to Arizona. . velutinus Horn (p. 105)
37(33). Pronotum bright shiny green or bluish; elytra usually bicolored, green
with yellow base and apex; Pennsylvania to Florida and Louisiana.
concinnus Laporte (p. 108)
Color feebly shining to dull uniform dark green, blue, or black, sometimes
with brown spots at apex of elytra. . . . ol als ree eS
38(37). Elytral intervals triserially tuberculate; male without basal cephalic
NOLS sa". * SO,
Elytral intervals ipisenially mahciaveccuberculater green, setae, or black;
male with short acute basal horn directed upward and outward behind
each eye; Kansas to Texas and Louisana.
medorensis Brown (p. 112)
39(38). Pronotal setae fine and long, length much greater than distance between
elongate oval tubercles; east of Rockies.
hecate hecate (Panzer) (p. 115)
Pronotal setae short, inconspicuous, scarcely longer than distance
between small round tubercles; usually brown spots at apex of elytra;
Florida to South Carolina. . . . heeate blatchleyi Brown (p. 120)
Onthophagus coproides Horn
Puate 1, Fiaures 4 anp 5
Onthophagus coproides Horn, 1881, p. 75.—Henshaw, 1885, p. 87.—Schaeffer,
1914, p. 293.—Leng, 1920, p. 248.——Dawson, 1922, p. 178—Boucomont
and Gillet, 1927, p. 205.—Boucomont, 1932, p. 297.
Onthophagus cuboidalis Bates, 1887, p. 79.—Boucomont, 1932, p. 297—Black-
welder, 1944, p. 211.
Mae masors.—Length 11.5 to 14.0 mm., width 7.1 to 7.3 mm.
Dorsally and ventrally brown to piceous, with legs same color. Cly-
peus transverse, somewhat quadrangular; lateral anterior angles
sharply bent and sometimes extending beyond the lateral margins of
the genae; clypeus gradually reflexed between the anterior angles;
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 15
margin slightly thickened and slightly emarginate or truncate me-
dially; clypeal disc closely, shallowly, rugosely punctate, the punctures
fine to moderate in size, separated by less than a diameter, many
confluent; clypeal carina fine, low, evenly elevated, relatively straight,
impunctate. Frons slightly convex medially; shallowly, rugosely
punctate but less closely than clypeal disc; delimited laterally from
the gena by a definite sutural line. Carina of the vertex very strong,
bowed forward slightly, almost straight, lowest medially, elevated
at each end into a strong horn near the eyes, the carina and horns
shallowly punctate anteriorly; genae not distinctly flared laterally,
extending no further than the anterior angles of the clypeus, punctures
as on clypeus.
Pronotum completely margined, anterior angles rounded and flat-
tened, lateral margins bent downward sharply just in front of the
middle. Pronotum convex and tumid anteriorly, not overhanging
the head; the tumosity widely and bluntly shaped like an inverted
V, the arms of the V extending posteriorly as well-defined ridges,
sharply declivous in front of the ridge and weakly concave from lat-
eral end of the ridge down to the margin opposite the eye; the median
line weakly impressed over posterior third. The tumosity with a
scattered group of 10 to 15 coarse intermixed punctures high on each
side of the anterior face; the surface otherwise from closely, moder-
ately punctate anteriorly (the punctures separated by less than a
diameter) to gradually much finer over the posterior part of the disc
which is smooth and shining between the punctures. LElytral striae
shallowly punctate; intervals slightly convex, generally smooth and
shining, with scattered fine punctures; a few setae evident posteriorly,
largely in the punctures of the sutural interval.
Pygidium shining, moderately punctate, the punctures each bear-
ing a short yellow seta. Ventrally the thorax coarsely punctate
laterally, the punctures becoming smaller and finer medially; meta-
sternum smooth and shining, with median line finely, slightly im-
pressed; laterally the punctures bearing long thin dark reddish setae.
Abdomen medially impunctate, the last three segments each bearing
a row of small setigerous punctures laterally; last segment lighter in
color and narrowed medially to receive the pygidium. Forelegs
lengthened, the tibia considerably longer and thinner than in the
female, with four teeth, the basal one often obsolete; the margin
between the teeth smooth, differing in this respect from the other
species which have the margin serrated. Middle and hind femora
with a few large coarse punctures apically; scattered fine punctures
over the entire ventral surface.
MALE AND FEMALE MINORS.—These show only a vague anterior
pronotal carina and are quite similar in every way except in the shape
16 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 114
of the frontal carina. In the male the carina is slightly arcuate for-
ward and evenly elevated, with rounded ends. In the female the
frontal carina is sharply angulate anteriorly at the middle, and the
ends are also sharply angulate and prominent. The front tibia is
slightly wider in the female.
Frema.es.—Length 11 to 11.3 mm., width 6.5 to 7.2mm. Differing
from male majors in the following respects: Clypeus almost evenly
arcuate, anteriorly broadly slightly emarginate and only slightly
reflexed; laterally not extending beyond the genae; disc flat and not
concave as in male major, surface moderately rugosely punctate; a
low clypeal carina separating the clypeus from the frons and extending
laterally to the juncture with the genae. Frons moderately coarsely,
almost rugosely punctate as are the genae; carina of the vertex
strongly developed and strongly angularly bowed forward medially,
ending on each side in a long horn as described for the male major.
Pronotum with outline, punctation, convexity, and tumosity similar
to the male major. The elytra are similar also. The pygidium is
slightly convex apically, shining, and with scattered close setigerous
punctures. Ventrally the major differences from male majors are the
more pronounced thoracic punctures, the even length of the last
abdominal segment, and the shorter, thicker foretibia. The four
teeth on the foretibial margin are broader than in the male, but the
margin is still smooth and nonserrated. Females show more varia-
tion than is usually found in this sex, the horns at the ends of the
frontal carina in fully developed individuals being longer than in most
males and the high connecting carina being angularly bent far forward
with a short but noticeable longitudinal carina extending back from
the angle.
Typre.—Lectotype, present designation, a large female in Academy
of Natural Sciences of Philadelphia, Type 3569. Horn stated that
the three specimens before him were males, but examination shows
all are females.
TyPE LOCALITY.—Santa Fe Canyon, New Mexico (700 ft.).
SPECIMENS EXAMINED.—43.
DistrrputTion.—(See fig. 1, p 17.)
New mexico: Santa Fe Canyon, Water Canyon (5000 ft.), Clouderoft, Fort
Wingate, Las Vegas, El Porvenir (San Miguel Co.), Torrance, 6 mi. south of
Thoreau (McKinley Co.) from stomach of Ambystoma tigrinum. ARIZONA: Chi-
ricahua Mts., Douglas, Prescott, 8 mi. south of Showlow, Springerville (Apache
Co.), General Springs (Coconino Co.), Woolaroec. coLorapo: Colorado Springs.
NEBRASKA: Halsey.
Remarxks.—Onthophagus coproides, the largest North American
Onthophagus, can be distinguished by the nonserrated margin between
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 17
the tibial teeth of the foretibia, by the high carina of the vertex which
ends over each eye in a long horn in both sexes, by the smooth, finely
punctate basal portion of the pronotum, by the minute punctures
of the elytral intervals, and by the dark brown to piceous overall color,
Though coproides appears to be widely distributed in the higher
mountains of Arizona, New Mexico, and Colorado, it has also been
taken in Nebraska at a lower elevation. The scarcity of spec-
imens in collections indicates a restricted habitat. Quite likely
Figure 1. Distribution of species of Onthophagus:
X coproides Horn © striatulus floridanus Blatchley
@ = striatulus striatulus (Palisot de %* landolti texanus Schaeffer
Beauvois)
the species occurs in some rodent nest or animal burrow. It is
one of the few species about which little or nothing is known.
Among closely allied Mexican species, a female cotype of O. hippo-
potamus Harold loaned by the British Museum differs in having
(1) much denser pronotal punctation which is only very slightly
finer even in the basal areas, (2) the pronotal protuberance rounded,
not angulate in front as in coproides, and (3) the frontal carina on
the head straight and evenly elevated its entire length, not angulate
forward medially and not elevated into horns at the ends.
633411—62——2
18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Onthophagus cochisus Brown
Puiate 2, Figures 9 anp 10
Onthophagus cochisus Brown, 1927, p. 132—Boucomont, 1932, p. 317—Leng
and Mutchler, 1933, p. 38.
Mate mMaJsors.—Length 8.5 to 9.8 mm., width 5.3 to 6.2 mm., color
black. Head with clypeus reflexed anteriorly, the disc appearing
slightly concave; margin anteriorly broadly arcuate, laterally sharply
angulate with sides almost parallel; disc coarsely punctate, punctures
becoming more crowded laterally and posteriorly; clypeal carina
obsolete, indicated only by a slight convexity medially between the
clypeus and frons. Frons coarsely and densely punctate as is the
gena; carina of vertex low, bowed anteriorly, ending shortly behind
each eye in a slightly elevated sharp ridge. Vertex coarsely
punctate for a short distance behind the carina, the surface then
becoming smoothly alutaceous. Genae with lateral margins extending
a short distance beyond the lateral margins of the clypeus, but
with the sides only slightly arcuate, nearly parallel.
Pronotum completely margined, poorly so posteriorly; anterior
angles almost forming right angles; lateral pronotal margin bent
abruptly upward approximately 1 mm. behind the anterior angles,
forming a sharp, nearly tuberculate angulation. Pronotum widest
at the middle, approximately 1 mm. behind the angulation, convex,
similar in shape to the related Mexican species O. chevrolati Harold;
the anterior tumosity rising almost perpendicularly behind the anterior
margin, with face of the tumosity broadly rounded and laterally con-
cavely arcuate to the small lateral tubercles. Surface of the pronotum
smoothly, shallowly concave between the tumosity, tubercles, mar-
ginal angulations, and anterior angles; discal surface of pronotum
coarsely, densely punctate; anteriorly the surface of the tumosity
shining, with the punctures large and often bearing short setae;
posteriorly the surface between the punctures dull and alutaceous,
with the punctures smaller and bearing short but distinct setae.
Elytral striae punctate, scarcely impressed; intervals, except for
sutural interval, with from three to five irregular rows of small shiny
tubercles, each with a short black seta at its base; surface alutaceous
between the tubercles.
Pygidium alutaceous except for smooth apex, the entire surface
with scattered shallow punctures, each with a short seta. Ventral
surfaces of thorax shining, laterally coarsely punctuate, most of the
punctures bearing long fine reddish setae; metasternum finely punc-
tate along the shallowly impressed median line; abdominal segments
finely alutaceous, laterally sparsely punctate; last segment piceous in
color, narrowed medially, emarginate to receive the pygidium. Fore-
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 19
legs moderately elongate; foretibia slender and bent in the apical
third, tibial margin serrate above and between the four teeth. Femora
with ventral surface having a mixture of scattered large and minute
punctures, the coarse punctures usually bearing long setae.
Mate minors.—Length 6.1 to 7. mm., width 3.4 to 4 mm. _ Dif-
fering from the male majors in the following characteristics: Clypeus
rounded, not flared laterally, reflexed and slightly truncate anteriorly;
disc flat, coarsely, evenly, rugosely punctate, posterior clypeal carina
obsolete; frons coarsely punctate to shallowly, moderately punctate,
laterally and posteriorly alutaceous; carina of vertex evenly bowed,
highest medially, the sharp lateral portions of the carina near the eyes
greatly reduced; gena with sides broadly rounded, extending slightly
beyond the margins of the clypeus. Pronotum evenly convex, coarsely
punctate, less so near the anterior angles; surface alutaceous between
the punctures, an occasional small anterior smooth area being the
only indication of the median tumosity of the male major; lateral
pronotal margin only slightly bent in anterior third, not at all angulate.
Elytra not significantly different. Pygidium of male minor quite
convex, but otherwise similar to male major. Ventral surface similar
except for the foretibia, which is slightly bent in the apical half and
is shortened and thickened, but to a lesser degree than in the female.
Fremaes.—Length 8.5 to 9.5 mm., width 4.8 to 5.1mm. Differing
from the male majors in the following respects: Clypeus longer and
narrower, anteriorly reflexed, truncate or slightly emarginate, sides
laterally obliquely arcuate; disc flat, rugosely punctate, posteriorly
delimited by a low arcuate carina which is highest medially; frons very
coarsely, almost rugosely punctate; carina of vertex pronounced me-
dially, obsolete near the eyes, in some specimens thickened or indented
at the median line; vertex behind the carina with scattered coarse
punctures, surface finely alutaceous. Pronotum similar to that of
male minor; the median tumosity slightly more pronounced, as is
the angulation in the anterior third of the pronotal margin. Elytra
similar except that the tubercles on the intervals are more pronounced.
Pygidium similar. Ventral surface showing the major difference
in the foretibia, which is straight, short, and thickened in width,
and in the last abdominal segment, which is not emarginate medially.
Typr.—Academy of Natural Sciences of Philadelphia.
TypPE LocALity.—Pinery Canyon, Chiricahua Mts., Cochise Co.,
Arizona.
SPECIMENS EXAMINED.—346.
DistriBuTion.—(See fig. 2, p. 20.)
UniTED STATES: ARIZONA: Pinery Canyon (7000 ft.), Onion Saddle (7000 ft.),
Rustler Park (8400 ft.), Southwestern Research Station (5 mi. west of Portal),
and Cave Creek Canyon (5400 ft.), all in the Chiricahua Mts.
Mexico: CHIHUAHUA: Guerrero.
20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
Figure 2. Distribution of species of Onthophagus:
@ pennsylvanicus Harold © cochisus Brown
Remarks.—In the United States this uncommon species appears
to be restricted to the higher elevations (5000 to 8000 ft.) of the Chiri-
cahua Mountains of southeastern Arizona. Jt seems to be a fairly
general feeder as an adult, being taken on carrion and under fresh
horse droppings. Howden took 64 specimens on July 3 under leaf
litter at Onion Saddle in the Chiricahua Mountains. The specimens
were kept alive and attempts were made to rear the specimens in
flower pots. Both horse and cow dung were supplied, but owing to
higher laboratory temperatures or other unknown factors, no larval
cells were formed.
O. cochisus Brown can be separated from other American species
north of Mexico by its large size, its dull black alutaceous dorsal
surface, and the combination of a densely coarsely punctate pronotum
coupled with the conspicuously tuberculate elytral intervals.
Type material of closely allied species from south of our borders
was examined and the following differences noted:
O. hippopotamus Harold is slightly larger, is shining, and has punc-
tate rather than tuberculate elytral intervals.
O. totonicapanus Bates is much smoother, with shallow pronotal punc-
tures and practically no elytral tubercles, those showing being scarcely
visible and very widely scattered.
O. chevrolati Harold also is smoother in appearance, with pronotal
punctures finer, not as close nor as deep, and elytral tubercles much
finer and less conspicuous.
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 21
O. retusus Harold is also much less closely and evenly punctate,
with punctures smaller and with elytral tubercles and striae finer.
O. cyanellus Bates has pronotal punctures very noticeably of two
sizes, not uniform as in cochisus, and with very much finer tubercles;
it is deep blue rather than black.
Onthophagus batesi, new species
PLATE 2, FIGURES 6 AND 7
Onthophagus incensus (Say), Howden, 1955, p. 264.
Hototypr.—Male major, length 8 mm., width 5 mm. Shining, black,
with legs and marginal areas of head and pronotum reddish brown.
Anterior margin of clypeus abruptly, strongly reflexed; the apex
moderately, widely emarginate-truncate; lateral margins not reflexed.
Head wider than long, flat, without clypeal carina; genae weak, widely
rounded; the eyes convex, 11 facets wide, two-fifths as wide as long;
vertex with two long, widely separated, very slightly diverging ver-
tical horns; the horns nearly straight, weakly flattened transversely,
with a very distinct U-shaped carina extending down one horn,
across the front, and up the other; punctures of head fine, quite
evenly distributed in front of carina, separated generally by 2 to 3
diameters, smooth behind carina except at extreme occiput.
Pronotum finely, completely margined; anterior angles moderately,
sharply rounded but not acute, posterior angles broadly rounded;
median protuberance high, slightly bulbous, nearly vertical, grad-
ually rounding into the relatively shallow, vertical cavity on each
side which receives the cephalic horn; surface very finely punctate
laterally and medianly over protuberance; lateral cavities, disc, and
basal areas very minutely punctate, almost impunctate. Elytra shin-
ing; striae fine; strial punctures scarcely crenating the intervals which
are almost flat, all very finely, quite closely and confusedly punc-
tate, the punctures separated by 3 to 4 diameters. Pygidium nearly
flat, alutaceous except at apex; moderately, coarsely punctate each
side of a slightly elevated impunctate median line; the punctures
separated by 1 to 2 diameters, those in the basal angles and adjacent
lateral areas bearing setae about as long as distance between punc-
tures; setae decumbent and pointing inward and downward toward
apex; marginal bead extending upward at apex to a short toothlike
process (not evident in other specimens). Legs reddish brown, ante-
rior tibia ending in a blunt, slightly up-turned tooth above the apical
spur. Club of antennae fuscorufous.
Mate minors.—Length 6 to 9.5 mm., width 4 to 5 mm. Male
minors tend to resemble the female in elevation of clypeal apex,
length of cephalic horns, pronotal protuberance, and lateral cavities;
aD, PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
many almost duplicate the female except for a trace of a tubercle at
each end of the frontal carina, an apical tooth on anterior tibia, and
a narrowed middle of the terminal segment of the abdomen.
AtLotyPE.—Female, length 7 mm., width 4 mm. The clypeus
differs from that of the male as follows: Relatively longer; the ante-
rior margin slightly truncate and only very weakly reflexed; the
clypeal carina evenly elevated throughout, not higher at middle; the
frontal carina evenly elevated, straight except for ends slightly arcuate
posteriorly; surface of the clypeus transversely rugose-punctate; the
punctures between the rugae moderately coarse, separated by 1 di-
ameter or less, and except for the very fine punctures of the more
gentle posterior slope of the clypeal carina and the area back of the
frontal carina, the remaining surface similarly closely, moderately
coarsely, and deeply punctate. The pronotum with a similar but very
weak median protuberance and traces of the lateral cavities; the
punctation in the anterior angles generally more noticeable, being
about as on the head, gradually finer medially across the anterior,
and gradually much finer and sparser over the disc to the base where
the fine punctures are generally separated by 4 to 5 diameters. Other
characters very similar to male except that the pygidium is more
widely shining apically, the anterior tibia lacks the apical tooth, and
the terminal abdominal segment is not narrowed medially.
Typr.—USNM 65681.
Type Locatity.—La Union, El Salvador.
SPECIMENS EXAMINED.—161.
DistTRIBUTION.—(See fig. 3, p. 24.)
UNITED STATES: TEXAS: 30°’, 29 2, Brownsville, June 1, 1954, H. F. Howden
(Howden, usnm);1 9, Brownsville, June 21, 1955, Allen (usNnM).
Mexico: 1 0’, 1 9, no locality data, (USNM). cHIAPAS: 1 o’, Cerro Huaco,
Sept. 19, 1949, G. Halffter (cnc); 1 9, Chiapas, Pacific slope Cordilleras (800-—
1000), L. Hotzon (usnm); 2 7 o', 1 9, Tuxtla, Dr. Berendt (USNM). MORELOS:
1 &, Cuernavaca, July 1955, N. L. H. Krauss (usnm); 1 o, 3 9 9, Cuernavaca,
August 1955, cow dung, N. L. H. Krauss (usnm); 1 o&, 1 9, Cuernavaca, June
10, 1957 (usnm); 1 o&, Cuatla, July 20, 1956, A. W. Vasquez (USNM). JALIsco:
1 o, L. Chapala, July 1940, L. W. Saylor (cnc). sAaN Luis Potosi: 1 o,
El Salto, Aug. 3, 1949, L. J. Bottimer (tus); 5 &@o', 2 9 9, El Salto de Agua,
Aug. 13-14, 1958, H. F. Howden (cnc); 2 9 9, Tamazunchale, Aug. 15, 20, 1956,
A. W. Vasquez (usNM). yucaTAN: 1 o&, 3 9 9, Temax, Gaumer (USNM).
JALIsco: 1 9, Guadalajara, Aug. 22, 1903 (usnM); 2 ? 2, Chapala, Aug. 13-16,
1949, L. J. Bottimer (13s). VERACRUZ: 1 9, Veracruz, Apr. 1, 1955, J. Camelo
(usnM);1 o&, 2 9 9, Orizaba (Veracruz?), H. T. Osborn (usnm); 1 co, Orizaba
(Veracruz?), Salle coll. (usnm); 7 oo", 12 2 2, Lake Catemaco, Aug. 8-16, 1960,
H. F. Howden (cnc); 1 &, 1 9, Cotaxtla, July 6, 1957, W. W. Gibson (usnm);
20,1 9, Cotaxtla, Oct. 25, 1957, W. W. Gibson (usnm); 1 o’, Cotaxtla, Sept.
11, 1957, Campo Exp., W. W. Gibson (cnc); 1 &@, Cotaxtla, Nov. 11, 1957, W. W.
Gibson (USNM). OAXACA: 1 o, Oaxaco, Hoege (UsNM);1 o’, 2 9 9, Tuxtepec,
J. Camelo G. (usnm);2 2 2, Tuxtepec, J. Camelo G., 1934 (usnm);2 oc", 2 2 9,
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 23
Tuxtepec, J. Camelo G., November 1932 (usnm); 1 9, Tuxtepec, J. Camelo G.,
Nov. 7, 1934 (usNM). GUERRERO; 2 9 9, 3 mi. north of Mexcala, Aug. 23-24,
1958, H. Howden (cnc). conma: 2 9 9, Colima, L. Conradt (usnm); 1 9,
Volcan de Colima, L. Conradt (usNM).
CanaL Zone: 1 of, Corozal, June 20, 1937, R. Bliss (usnm); 1 oc, Gamboa,
E. E. Frick (cnc); 1 o, without locality, June 1944, K. E. Frick (cnc); 1 ?, Barro
Colorado Id., P. Rau (usnm);1 9, Tabernilla, July 1907, August Busck (usnm).
GUATEMALA: 2 9 9, no locality, (UsnM); 1 ?, south of Geronimo, Champion
(usnM); 1 o, Palin, May 1924, W. M. Mann (usnm);1 9, Lake Thiel, 1925, S.
Sebastian Retalnuleu (UsNM).
Honpvuras: 1 oc’, San Pedro Sula, W. M. Mann (usnM); 1 o, Carmelina,
W. M. Mann (usnM);1 9, La Ceiba, W. M. Mann (usnm);1 9, San Juan Pueblo,
W. M. Mann (usnm).
Ex Sauvapor: 2 oc’, 2 2 9, San Salvador, May 11-15, 1958, O. L. Cart-
wright (USNM); 1 o, 2 9 9, San Salvador, June 24, 1958, O. L. Cartwright
(usnM); 2 ? 9, San Salvador, June 9, 1958, O. L. Cartwright (usnm); 2 9 9,
San Salvador, June 17, 1958, O. L. Cartwright (usnm); 1 9, San Salvador,
June 7, 1958, O. L. Cartwright (usnM); 2 oo", 2 9 9, San Salvador, June 1-5,
1958, O. L. Cartwright (usnm); 1 o’, San Salvador, June 20-23, 1958, O. L.
Cartwright (UsNM);1 9, San Salvador, June 21, 1958, O. L. Cartwright (usNnmM) ;
6 2 ¢, San Salvador, May 1, 1957, P. A. Berry (usnM);1 7, 1 9, San Salvador,
June 21, 1958, L. J. Bottimer (Lys); 1 o, San Salvador, June 10, 1958, L. J.
Bottimer (LJB); 1 o, San Salvador, June 15, 1958, L. J. Bottimer (LB); 2 7d,
2 9 9 (includes holotype and allotype), La Union, May 30, 1958, O. L. Cart-
wright (UsNM); 1 o', La Union, May 30, 1958, L. J. Bottimer (Lys); 1 9, La
Palma, June 24, 1958, O. L. Cartwright (usnM); 1 co’, Tonocatepeque, June 20,
1958, L. J. Bottimer (LB); 1 @, 1 9, Voledn Conchagua, June 27-29, 1958, L. J.
Bottimer (tyB); 1 9, Santa Tecla, Apr. 29, 1954, P. A. Berry (usnM); 1 o, La
Ceiba, Vera Wellborn (usnM); 1 <’, no locality; 1 @#, 1 9, Lake Olomega, Dept.
San Miguel, July 14, 1925, R. A. Stirton (cnc, Howden); 2 9? 9, Mt. Caca-
guatique, Dept. San Miguel, Dec. 22, 1925, R. A. Stirton (cnc, Howden); 1 9°,
Sonsonate, Aug. 26, Fredk. Knab (usnm).
Nicaraaua: 1 o, Managua (8 mi. north), July 28, 1958, S. E. Neff and E.G.
Matthews (uUsNM).
Costa Rica: 2 fo’, 5 9 9, Hamburg farm, Reventazén above Limén, Aug.
1, 1928, F. Nevermann (usNM);1 ?, Hamburg farm, Reventazén, May 23, 1935,
F. Nevermann (usnM); 1 9, Hamburg farm, Reventazén, July 28, 1926, F.
Nevermann (usnM); 1 o, Esparta, Sept. 18, 1905, Fredk. Knab (usNnm); 1 0,
no data (USNM).
Remarks.—0O. batesi is normally black, occasionally with faint
ereenish lustre, and with legs reddish brown. Quite often traces of
reddish brown are found narrowly along the base and sides of the pro-
notum and head. Reddish-brown (teneral?) specimens are also found.
The coppery reflections seen in the type and other specimens of
O. incensus have not been observed in batest. The antennal club
is a lighter, redder brown in batesi, more fuscous in incensus.
This species is more finely punctate than is incensus Say, the male
horns are united by a rather sharp continuous carina down one horn,
arcuate across the front, and up the other, the female clypeal carina
is evenly elevated and not higher in the middle as in incensus, the
24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Ficure 3. Distribution of species of Onthophagus:
% batesi, new species © subopacus Robinson
@ oklahomensis Brown
frontal carina sometimes breaks downward suddenly, rather angu-
larly at the ends, the eyes are slightly shorter and wider, and in batesi
the pygidium usually shows a more or less impunctate, slightly ele-
vated median line. If the specimen is turned in reflected light, the
even curvature of the pygidium is seen to be slightly broken dorso-
ventrally by the midline. Many of the male minors show traces of
the clypeal carina.
The Brownsville, Tex., specimens were collected in shallow burrows
under fresh horse droppings in the area known as the “palm jungle”’
on the McCormick ranch 7 miles east-southeast of the city.
Onthophagus incensus Say
Puate 1, Figures 1-3
Onthophagus incensus Say, 1835, p. 173.—Lacordaire, 1856, p. 109.— Harold,1880,
p. 30.— Bates, 1887, p. 66—Boucomont and Gillet, 1927, p. 206.—Boucomont,
1932, p. 308.
Onthophagus curvicornis Latreille var. incensus (Say), Boucomont, 1932, p. 308.
Mae magors.—Length 7 to 10 mm., width 4 to 6 mm. Black,
shining, sometimes with greenish or coppery lustre at base of head
and anteriorly on the pronotum. Anterior margin of clypeus rather
widely, shallowly emarginate and sharply reflexed; lateral margins
only feebly or not at all reflexed. Head without carinae, slightly
wider than long, the genae prominent but obtusely rounded; disc
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 25
very slightly convex, basally with two long, widely separated verti-
cal horns slightly curved forward and weakly bowed laterally, the
horns somewhat flattened transversely with rounded edges, not con-
nected by a basal carina; punctures of head very fine medially,
separated by two or more diameters, somewhat larger laterally,
particularly on the genae.
Pronotum finely, completely margined; anterior angles sharply
rounded but not acute, posterior angles broadly rounded; median
protuberance vertical in front, quite high, the broad rounded sum-
mit with a very weak longitudinal, median depression, laterally rounded
into a strong vertical cavity each side to receive the cephalic horns.
Punctures fine in anterior angles and median groove of protuber-
ance, about as on genae, gradually finer away from these areas to
the disc and base; the disc sometimes almost impunctate, lateral
foveae strong. Elytra shining, rarely very finely alutaceous; striae
fine, the punctures slightly crenating the intervals; intervals weakly
convex, all finely, quite closely, confusedly punctate, the scattered
punctures separated by 1 to 2 diameters. Pygidium weakly, evenly
convex, very finely alutaceous, and not as shining as upper surfaces;
punctures close, very slightly larger and deeper than on elytra, occa-
sionally with a few in rows between transverse wrinkles. Legs dark,
almost black, the anterior tibia with a blunt, slightly upturned tooth
at the tip above the spur; club of antennae fuscous.
Mae minors.—Variation is toward the female in reduction in
the length of the cephalic horns, the pronotal prominence, and the
accompanying lateral cavities. The only noticeable difference be-
tween the least developed male seen and a female was the complete
absence of the clypeal carina, the presence of the anterior tibial tooth,
and the narrowed apex of the terminal abdominal segment.
Frema.tes.—Length 7.5 to 10 mm., width 4.75 to 6mm. _ Differing
from males as follows: Punctation usually more noticeable, closer,
and deeper on the upper surface; no tooth or tubercle above the spur
of the anterior tibia; the terminal abdominal segment not narrowed
apically; clypeus longer, rounded, and not noticeably emarginate-
truncate or reflexed; clypeal carina well developed and noticeably
higher at middle; frontal carina also well developed but usually slightly
lower and posteriorly angulate at middle; anterior prominence of
pronotum weak, similar to that of male, but widely depressed medi-
ally and laterally reduced to rounded tubercles.
Typr.—Museum of Comparative Zoology, Harvard College,
Cambridge, Mass.
Typr Locautity.—“Mexico.” Barber (1928) stated, “It is probable
that all of his (Say’s) Mexican forms were collected along the old road
between Veracruz, Jalapa, Mexico City, and Tacuba.”’
26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
SPECIMENS EXAMINED.—196.
DIstRIBUTION.—
UNITED StTaTESs: HAWAII: Oahu: Pohakea Pass, Schofield Plateau; Hawaii:
Kona, Kealakekua.
MEXICO: ME&XICO, D.F. HIDALGO: Jacala. VERACRUZ: Jalapa, Cérdoba, Ori-
zaba, south of Veracruz, Banderilla.
GuaTEMALA: Senahti (Alta Verapaz), Capetillo.
Ex Sautvapor: Monte Cristo.
Costa Rica: Meseta (central Costa Rica), Aurora Farm (Estrella Valley),
La Carpintera, Coronado, San José, Coliblaneo, Tablaco, Zarzero, Turrialba,
Voleén Irazt.
Panama: Voledén Chiriqui, Boquete (Chiriqui Province), Bambito (Chiriqui
Province), Bugaba.
BroLtocy.—Taken in numbers in sparse woods near Jacala, Mexico,
in cow and horse dung. Burrows in clay soil were 4 to 8 inches deep.
REMARKS.—Say’s description of O. incensus was published post-
humously in the Boston Journal of Natural History for May 1835.
The holotype female (pl. 1) is in the Museum of Comparative Zoology
at Harvard; it is in fair condition but lacks mouth parts, pygidium,
and part of the legs. The pronotum of the type specimen is a very
dark green, with reddish-brown lateral spots below the fovea and with
narrowly reddish-brown anterior margin opposite the head area be-
tween the eyes; the carinae of the head and the pronotal prominence
are faintly but distinctly coppery; the elytra are dark blue-black.
The narrow brownish anterior margin of the pronotum is found in
other Mexican specimens also. The clypeal carina is noticeably
higher at middle.
The cephalic horns in fully developed males are only rarely com-
pletely connected basally and then by a low swelling or obtuse carina,
straight across from one to the other. In batesi the connecting carina
is nearly always present even in the male minors, curves arcuately
from one horn to the other, and is sharply ridged on top.
This species which is very closely related to bates?, new species,
has not been taken in mainland United States but has been intro-
duced into Hawaii. It was introduced in 1923 by H. T. Osborn from
material collected at Morelos, Mexico, to assist in control of the horn-
fly, Siphona iritans (Linnaeus), and was recovered on the island of
Hawaii in 1934 and on Oahu in 1940 (see Swezey, 1935; 1940).
Some females of curvicornis Latreille from Colombia are super-
ficially almost identical with incensus Say; however, curvicornis is
usually larger and the fully developed male pronotal protuberance
extends forward in a sharply pointed but somewhat flattened cone.
Females of O. acuminatus Harold also resemble those of icensus
and batesi but are much smaller. The male clypeus in acuminatus
anteriorly is pointed. O. nitidor Bates is slightly smaller and is a
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT OT
brighter, shining green, with black clypeus; pronotal protuberance
in both sexes not as wide.
Onthophagus brevifrons Horn
Puate 2, Figures 12 anv 13
Onthophagus brevifrons Horn, 1881, p. 76.—Henshaw, 1885, p. 87.—Shaeffer,
1914, p. 300.—Leng, 1920, p. 49—Boucomont and Gillet, 1927, p. 204.—
Boucomont, 1932, p. 315.—Howden, Cartwright, and Halffter, 1956, p. 6.
Mae masors.—Length 8.6 to 10.3 mm., width 5.3 to 5.7 mm.
Color uniformly brown, piceous, or black; head and pronotum shin-
ing, occasionally with violaceous luster, elytral intervals dull, finely
alutaceous. Head with clypeus reflexed, more so anteriorly than
laterally, anterior edge vaguely emarginate (Horn’s Kansas specimens)
or truncate, rounded in worn specimens; lateral margin delimited from
gena by faint notch, gena scarcely flared, broadly arcuate; disc of
clypeus and genae with scattered large shallow punctures, clypeus
often rugosely punctate laterally, base of clypeus and anterior portion
of frons tumid, the clypeal carina barely indicated by an impunctate
line across the tumosity. Frons coarsely punctate, the punctures half
the size of the larger clypeal punctures. Surface of clypeus, genae,
and frons between the punctures smooth and shining. Carina of
vertex low, bowed anteriorly on each side of a raised median point, and
ending laterally near the posterior margin of the eyes; scattered coarse
punctures behind the carina, smooth areas between the punctures very
finely alutaceous.
Pronotum margined anteriorly and laterally, lateral margins no-
ticeably bent downward in anterior third; pronotum convex, with a
large, rounded, anterior tumosity rising abruptly behind the anterior
margins and ending on each side in a small raised ridge. Below the
ridge on each side of the tumosity a shallowly concave area extending
to the bend in the lateral pronotal margins and forward to the anterior
angles; surface of the concavity finely alutaceous near the anterior
angles, otherwise the surface smooth and shining between the pronotal
punctures. Punctures of two sizes, large, deep, sharply delimited
circular punctures separated by 1 to 2 diameters, and, between these,
scattered small well-defined punctures less than one-fourth the
diameter of the larger ones; both types of punctures most numerous on
the face and along the top of the tumosity, becoming more widely
separated posteriorly and near the anterior angles; large punctures on
the face of the tumosity often bearing setae.
Elytral striae coarsely punctate, shallowly impressed; intervals with
numerous small shining tubercles with fine setae at their bases, the
surface between the tubercles finely alutaceous, tubercles on the third,
28 PROCEEDINGS OF THE NATIONAL MUSEUM vou, 114
fourth, and fifth intervals forming two irregular rows. Pygidium
convex hear apex, surface finely alutaceous with scattered poorly
defined punctures usually bearing minute setae. Underside of thorax
with coarse punctures laterally, the punctures bearing moderately long
reddish setae; impunctate or with very fine punctures medially.
Abdominal segments bearing scattered punctures laterally, last seg-
ment moderately emarginate medially. Foreleg scarcely longer than
in female, not slender or bent in anterior third; four teeth of the tibia
broad and stubby; tibial margin only serrate or finely denticulate
above the last tooth, not serrate on or between the teeth. Ventral
surfaces of femora with a few widely scattered coarse punctures and
with more numerous but still scattered, fine punctures.
Mate minors.—Length 7.2 to 8.4 mm., width 4.1 to 4.7 mm.
Similar to females in clypeal outline and pronotal configuration.
Clypeal carina present but not as pronounced as in females; frontal
carina low, nearly straight, highest in middle instead of at ends as in
females. Similar to male majors in other respects.
Frematrs.—Length 7.1 to 9.6 mm., width 4.0 to 5.2mm. Differing
from the male majors in the following respects: Head with broadly
arcuate clypeus, shallowly reflexed anteriorly and laterally, anterior
margin slightly emarginate; disc flat, transversely rugosely punctate;
clypeal carina broadly raised, highest medially, laterally becoming
merely a raised line between the clypeus and genae. Frons and genae
coarsely, irregularly punctate, some of the punctures bearing fine
reddish setae. Carina of vertex distinct, lowest at the midpoint, rising
evenly (in the majority of specimens) to the point where the carina
terminates near the base of each eye; in a few of the diminutive fe-
males the carina straight across, not raised laterally, but in the larger
specimens the carina distinctly elevated near each eye, horns not
present, the elevated portions being merely the ends of a very obtuse
V-shaped carina. Behind the carina the vertex bears some scattered
coarse punctures, with impunctate areas finely alutaceous.
Pronotum moderately convex, broadly tumid in the large females,
only slightly so in small specimens; tumosity, when developed, approx-
imately as wide as head with only faint concave areas on each side;
lateral margins of the pronotum only vaguely bent in anterior third;
surface of pronotum bearing large and small punctures similar to
male major, many of the large punctures of the disc bearing short
inconspicuous setae. Elytra and pygidium similar to those of male
major. Ventral surface generally with more pronounced punctures
than in male. Last abdominal segment not narrowed medially.
Foretibia slightly shorter and wider than in male major but not
greatly different, outer margin serrate or denticulate only above the
four tibial teeth and not between them.
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 29
Typr.—Lectotype, present designation, a male in Academy of
Natural Sciences of Philadelphia, Type 3570.
Tyrer Locauity.—‘‘Plains of Kansas.”
SPECIMENS EXAMINED.—101 and fragments.
DISTRIBUTION.—(See fig. 4.)
Kansas: ‘Plains of Kansas.”?’ ARIZONA: Paradise, base of Pinal Mts., Globe,
Prescott, Yavapai Co., Portal. tTExas: (Cited without locality by Horn, 1881,
D.) 70) =
Remarks.—This species can be distinguished by its large size,
black or brown color, and shining, usually tumid pronotum. The
head may occasionally have greenish or violaceous reflections, but
the pronotum is never distinctly green, as stated by Horn (1881, p.
76). Pronotal disc with numerous, sharply delimited circular coarse
and fine punctures. Elytral intervals tuberculate, flat areas finely
alutaceous, not smooth and shining. Foretibia not greatly differing
between the sexes; tibial margin of both with four broad teeth, the
margin serrate basally, but smooth between the teeth. This species
may be distinguished from the following species, subtropicus, new
species, by the shape of the carina on the vertex of the female.
Because there were no specimens available to Schaeffer for study
when he presented his review of the genus in 1914, he merely repeated
Horn’s description. The present description of the male major is
in part based on one of Horn’s Kansas cotypes which was carefully
Ficure 4. Distribution of species of Onthophagus:
@ brevifrons Horn EA subtropicus, new species
©. cavernicollis, new species + depressus Harold
30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
examined by Howden. About 20 of the specimens seen apparently
were collected in 1920 by H. H. Kimball at Paradise, Ariz.; 41 were
collected in wood rat nests at Portal, Ariz., in September 1960 by
Howden. ‘The writers found a few specimens and fragments a mile
east of Portal in wood rat nests in late June 1956. The Arizona spec-
imens are almost identical with Horn’s Kansas specimens except
that the eyes are narrower and the specimens appear darker, with only
occasional traces of greenish lustre on the head of a female.
Although no recently collected specimens from Kansas have been
seen, the Kansas label of the type series (collected by Dr. H. A.
Brons) should be accepted because at least two species of wood rats
occur in the state.
The unconnected distribution can at present be explained only
by the seemingly restricted habits of O. brevifrons, the only biological
information being that they occur in the soil under the large stick
nests of the wood rats (Neotoma species).
Onthophagus subtropicus, new species
PuaTE 2, Figures 8 anp 11
Hototypr.—Male, length 8 mm., width 4.5 mm. Head and pro-
notum shining, dark blackish green; elytra black, alutaceous, weakly
shining. Clypeus arcuate, margin moderately reflexed over anterior
half; disc densely, shallowly, moderately punctate, with some anas-
tomosing and tendency to rugosity near edges; clypeal carina rather
high and distinct over middle third of a raised area between the
interior ends of the distinctly carinate sutures between clypeus and
genae. Frons closely, moderately punctate, a few coarser punctures
laterally near the frontal carina; frontal carina moderately high,
strongly sinuate, the middle and ends arcuate posteriorly and not as
high at middle as near the ends; somewhat finer punctures behind the
frontal suture, separated 1 diameter or less, otherwise minutely alu-
taeous. Genae widely rounded, margin joining clypeus without a
break, surface closely coarsely punctate. Eyes wider than in brew-
Jrons, two-fifths as wide as long, about 7 facets wide.
Pronotum margined anteriorly and laterally, basally very finely
so for a short distance at middle; disc very convex, high in front
with a broad vertical concavity on each side; upper edges of the
concavities sharp, almost cariniform for a short distance; anterior
angles not sharply rounded; surface of disc everywhere with close,
mixed, moderately coarse and fine punctures, the coarse punctures
separated by a diameter or more, the fine punctures everywhere in
between; surface otherwise smooth and shining except for alutaceous
sculpture narrowly across the base. Elytra strongly alutaceous,
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 31
striae shining; strial punctures distinct, slightly crenating the intervals;
intervals flat with irregular rows of widely spaced fine tubercles.
Pygidium alutaceous throughout, with scattered, moderately fine,
shallow punctures slightly closer laterally and apically. Ventral sur-
faces smooth and shining; metasternum with very fine, scattered punc-
tures at middle and moderate punctures forward and to sides; terminal
abdominal segment with a single transverse row of setigerous punc-
tures scarcely larger than those of the preceding segment; segment nar-
rowed apically to receive the pygidium. Forelegs slightly longer and
thinner than in the female.
A.LotyPE.—Female, length 8 mm., width 4.5 mm. Shining; head
black in front of clypeal carina, green behind; pronotum dark bluish
green, elytra black. Clypeus narrowly reflexed anteriorly, gradually
less so to genae; broadly truncate-emarginate at middle; disc moder-
ately, transversely punctate-rugose, most of the punctures anasto-
mosing and more or less indistinct; clypeal carina with narrow band
of close fine punctures on anterior slope, carina almost angularly
higher at middle. Frons quite closely punctate, the punctures sepa-
rated by 1 to 2 diameters; punctures fine anteriorly to moderately
coarse at frontal carina; frontal carina weakly arcuate posteriorly, low
and sharp medially, elevated each side to a high, acutely tipped
tubercle; tips of the tubercles slightly nearer the middle than to the
edges of the eyes; occiput, moderately finely punctate, more coarsely
so around the tubercles to the eyes. Genae closely coarsely punctate,
many of the punctures anastomosing; margin evenly obtusely rounded
to clypeus with only a tiny notch marking the juncture.
Pronotum finely margined, faintly and very finely so basally; disc
strongly convex, a wide, vague prominence in front with a shallow ver-
tical concavity on each side; anterior angles not sharply rounded ; surface
everywhere with close mixed moderate and fine punctures, tbe moder-
ate punctures separated by 1 to 2 diameters; surface otherwise smooth
except for very fine alutaceous sculpture narrowly along the base.
Elytra alutaceous except at shoulders; striae shining and sparsely,
shallowly punctate; intervals flat, the first with one, the second with
two and part of a third, and the next four with two irregular rows of
fine tubercles, separated by four to six or more times their diameters;
tubercles overhanging setigerious punctures but the setae so short as
to be almost invisible. Pygidium apically convex and shining, aluta-
ceous basally; scattered fine punctures becoming closer and deeper
apically. Ventral surfaces smooth and shining; metasternum with a
distinct, shallowly impressed midline, some fine scattered punctures
at middle, and moderate to coarse punctures forward and to sides.
Abdominal segments finely alutaceous, each with transverse row of
very fine tubercles overhanging very fine setae; setae relatively short
32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
and scarcely longer than the distance between them; apical segment
with an irregular band of coarse punctures separated by about a di-
ameter. Legs normal for females of the genus.
Typr.—USNM 65682.
Type LocaLiry.—Laguna Madre, 25 mi. southeast of Harlingen,
Tex.
SPECIMENS EXAMINED.—3.
DistRiputTion.—(See fig. 4, p. 29.)
Texas: Holotype, o, Laguna Madre, 25 mi. southeast of Harlingen, Sept. 20,
1945, D. E. Hardy and V. L. Woolley, nest of Neotoma micropus Baird (Robin-
son coll., usnm). Allotype, 9, Brownsville, Mar. 12, 1908 (Jones and Pratt
coll., usNM). Paratype, 9, Kingsville (C. T. Reed coll., CU).
Remarks.—This species is very closely related to O. brevifrons and
cavernicollis and is intermediate in size. In brevifrons, the color is all
black; in cavernicollis, the largest species, the pronotum is a bright,
shiny green; in subtropicus, the pronotum is not so shiny and is a
bluish or blackish dark green. The females of subtropicus may be
separated from the other two species by the shape of the frontal carina,
which is elevated on each side to a sharp median tubercle, the tips of
the tubercles being nearer the middle of the carina than to the eyes.
In brevifrons, this carina is wider and slopes evenly from the middle to
elevated rounded ends which are nearer to the eyes than to the middle
of the carina; the carina ends on each side in a sharp vertical drop to
the head surface. In subtropicus, the tubercles slope evenly away
from their tips in both directions. In cavernicollis, the frontal carina
is similar to brevifrons. The eyes of cavernicollis are larger and wider,
being a third as wide as long, 7 to 8 facets wide; those of subtropicus
are two-fifths as wide as long, 6 to 7 facets wide; those of brevifrons
are sometimes no more than a fourth as wide as long and only 4 to
5 facets wide. In brevifrons, there is a greater disparity in the sizes
of the pronotal punctures, with the coarse punctures closer together.
The holotype may be a male minor.
The type locality (given on the label as 25 mi. southeast of Harlin-
gen) was placed by a speedometer reading. Dr. Hardy informed us
the exact locality would be on the Laguna Madre about 10 miles north-
west of Port Isobel in an area used by Harlingen Air Force Base for
its gunnery school.
Onthophagus cavernicollis, new species
PLATE 3, FIGURES 22 AND 23
O. brevifrons Horn, 1881, p. 76, in part.
Hotoryrr.—Male major, length 11 mm., width 6.4 mm. Head
and pronotum shining, bright green; elytra weakly shining, black;
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 33
pygidium shining, dark green; ventral surfaces, excluding tibiae, black
with a more or less shining, greenish luster; tibiae dark brown with
only a faint greenish cast. Clypeus widely reflexed anteriorly, less
so laterally; margin broadly arcuate anteriorly (not even vaguely
emarginate as is often the case in unabraded brevifrons), becoming
sharply rounded laterally near the genae; disc concave near the cen-
ter, with coarse punctures shallow and usually separated by approxi-
mately their own diameters. Clypeal carina obsolete, the base of
the clypeus and the anterior portion of the frons medially forming
a low rounded hump; frons more finely, closely punctate than clypeus.
Gena with arcuate margin slightly flared, surface coarsely punctate.
Carina of vertex raised medially to a point, low and bowed on each
side, terminating near the base of the eyes, similar to that described
for brevifrons; scattered coarse punctures behind the carina, with
smooth surfaces finely alutaceous.
Shape of pronotum nearly identical with that of the male major
of brevifrons, the only differences in the pronotum of cavernicollis
being in the posterior, vaguely indented median line and in the punc-
tures (also a mixture of large and small ones) which are not as sharply
delimited and appear slightly smaller and shallower. Most of the
large punctures bearing short, inconspicuous, reddish setae. The con-
vexity of the pronotum seemingly more pronounced than normal
in brevifrons, but this apparent variation is perhaps an illusion caused
by the overall larger size of cavernicollis. The elytra likewise very
similar to those of brevifrons, differing only slightly by having smaller
tubercles on the intervals. Pygidium with scattered irregular punc-
tures bearing short, fine setae; the surface basally alutaceous, becom-
ing somewhat shiny and rugose near the apex, the smooth areas dull
green. Ventral surfaces and legs lacking any characteristics, other
than color, that would readily separate cavernicollis from brevifrons.
Mates minors.—No specimen seen.
AuLtotyPe.—Female, length 11.2 mm., width 6 mm. Differing
from the holotype male major in the following respects: Clypeus
arcuate but feebly emarginate anteriorly, clypeal margin scarcely
reflexed; disc transversely rugose, more closely so than in females of
brevifrons; clypeal carina pronounced, highest medially, becoming
obsolete by the genae. Frons coarsely punctate behind the carina,
punctures not so sharply delimited as in brevifrons; genae with margin
arcuately curving inward by the eyes, anteriorly not delimited from
the clypeal margin. Carina of vertex very similar in shape to that
described for the female of brevifrons but raised terminally to form
small horns, the carina extending at least two-thirds the distance from
the midline to the eye but still terminating nearly twice as far from
6383411—62——_3
34 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
the eye as in brevifrons; a few scattered coarse punctures behind the
carina, the smooth areas finely alutaceous.
Pronotum quite convex, tumid anteriorly, but considerably less
so than in the male major; the tumosity more rounded but still similar
to that of the male, and the lateral concavities much reduced, extend-
ing neither to the lateral margins nor to the anterior angles; margin
only slightly bent in the anterior third; punctures slightly more pro-
nounced than in the male, but still smaller and more scattered than in
brevifrons; midline of pronotum slightly depressed posteriorly and
with fewer coarse punctures. Elytra not noticeably different from
those of the male. Pygidium rather convex apically and more shining
than in the male but otherwise similar. Ventrally, differences from
the male are noted in the shorter and wider foretibia and in the last
abdominal segment, which is not emarginate in the female.
Variation.—In the males the length varies from 9.5 to 11 mm., the
width from 5.7 to 6.4 mm., and in the females the length varies from
9.1 to 11.2 and the width from 5.4 to6 mm. The 10 specimens seen
exhibit remarkably few noteworthy differences. The brightness of
the greenish color of the pronotum is slightly more pronounced in
some of the specimens, but in all it is quite evident. In the smaller
specimens, both male and female, the pronotal punctures are slightly
more pronounced than in larger specimens of the same sex. The
male paratypes have the carina of the vertex more sharply indicated
laterally than in the holotype, but otherwise the carinae are similar.
Type.—USNM 65683.
Type Locatiry.—Waglers Cave, Harrison, Ark.
SPECIMENS EXAMINED.—10.
DistrRiputTion.—(See fig. 4, p. 29.) Holotype, o’, and allotype, 9,
Harrison, Waglers Cave, Ark., Apr. 13, 1935, in bat dung, J. M.
Valentine. And the following paratypes:
ARKANsas: Washington Co., Apr. 24, 1988 (INHS). oKLAHOMA: 1 9, Tahle-
quah, Adair Co., Sept. 26, 1954, O. C. Schomberg (1uB). TEXas: Vicinity “cave
without name” near Boerne, Kendall Co., July 30, 1948, G. E. Ball (AMNH); no
locality (mcz); 1 Q (ANSP). MiIssouRI: 4 oc’, 29 9, Bat Cave, Liking, June
20, 1956, Condé (Muséum National d’Histoire Naturelle, cnc).
Remarks.—Onthophagus cavernicollis can be distinguished from
other North American species by its large size and bright, shining
green pronotum. Except for size and color, the characteristics listed
for separating O. brevifrons will also separate this species from the other
North American forms. Coloration, size, the smaller pronotal
punctures, and the shape of the carina on the vertex of the female
will separate cavernicollis from either brevifrons or subtropicus.
In addition to the characteristics given above, the habitat in which
the majority of specimens have been taken seems to set the species
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 35
apart from the other North American forms. The eight specimens
bearing biological data were all taken associated with bat dung in
caves. Collecting the specimens in this extremely unusual niche
might be considered chance except for the occurrence of a related
Mexican species on bat dung in caves. This latter species was de-
scribed, compared with brevifrons, and its biology discussed by How-
den, Cartwright, and Halffter (1956). There are probably other
species with similarly odd habits. Such an unusual habitat helps
explain why 0. carvernicollis has been so long undescribed and why
there are still so few specimens known.
The specimen from Texas, without definite locality, in the collec-
tion of the Academy of Natural Sciences of Philadelphia, bears a blue
paratype label, No. 3570.3, and is presumably one of the original
cotypes of brevifrons Horn.
Onthophagus polyphemi polyphemi Hubbard
PuatEe 3, Figures 14 anp 15
Onthophagus polyphemi Hubbard, 1894, p. 311.—Henshaw, 1895, p. 22.—Schaef-
fer, 1914, p. 293.—Leng, 1920, p. 248.—Boucomont and Gillet, 1927, p. 207.—
Cartwright, 1939, p. 285.—Howden, Cartwright, and Halffter, 1956, p. 10.
Mae mMasors.—Length 5.5 to 6.9 mm., width 3.5 to 4.1 mm.
Dorsal color very dark reddish brown to black, the head and pronotum
sometimes slightly darker than the elytra. Clypeus gradually, slightly
reflexed anteriorly, truncate, occasionally barely emarginate, later-
ally slightly arcuate, joining almost evenly with the margin of the
gena; disc bearing from 15 to 30 small scattered tubercles, at the
base of each tubercle a long reddish seta. Posterior margin of clypeus
delimited by a long, very distinct, evenly elevated, strongly anteriorly
arcuate carina. Front of head behind clypeal carina nearly flat
with a dozen or more widely scattered, setose tubercles; genae scarcely
extended laterally, their sides often almost parallel; plane of genae
slightly lower than frons and gradually sloping downwards, surface
smooth sometimes with two or more setigerous tubercles. Carina
of vertex nearly straight, obsolete medially (type series), or distinct
for its entire length, being highest laterally; when absent medially,
it has the appearance of two small separated carinae, one above each
eye (as noted in Hubbard’s original description); behind the frontal
carina an irregular row of six or more tubercles each with a seta.
Pronotum convex, margined anteriorly, laterally, and obsoletely
at middle of base. Pronotal protuberance represented by a broad
anterior swelling slightly wider than the head, vague at the middle,
more distinct laterally, the steep declivity weakly concave high up
at the sides above the anterior angles; anterior angles with scattered,
moderate, setigerous punctures, finer and fewer toward middle of de-
36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
clivity; each puncture with a small tubercle in front of the seta, the
tubercles disappearing laterally and posteriorly and the punctures
becoming much coarser and farther apart, with a few fine punctures
intermixed, the surface otherwise smooth and shining. Elytral striae
vaguely punctate, intervals smooth and shiny and, except for the su-
tural one, each with a double row of minute tubercles having setigerous
punctures at their bases. Pygidium feebly convex, with scattered,
coarse, setigerous punctures, the punctures becoming smaller apically;
surface between the punctures finely alutaceous basally, apically al-
most smooth.
Ventral surfaces and legs reddish brown to black. Antennae
reddish brown, the club slightly lighter in color. Proepisternum only
slightly excavated to receive antennae. Mesothorax and lateral areas
of prothorax and metathorax coarsely setigerously punctate and finely
alutaceous. Median area of metathorax smooth and shiny, with
scattered nonsetigerous punctures, often with a vague sulcus along
the median line. Abdomen except for first segment with a basal row
of minute setigerous punctures extending across each segment, last
segment narrowed medially. Forelegs approximately the same length
as those of female, tibia stoutly and conspicuously quadridentate;
outer margin smooth between the teeth, occasionally vaguely serrate
above them; conical projection lacking above apical spine. Femora
of all legs with numerous setae in an irregular row at their anterior
and posterior edges, the flattened ventral surface with scattered
coarse setigerous punctures.
Mats mrnors.—Length 4.7 to 6. mm., width 2.7 to 3.7 mm. Very
similar to male majors. Anterior clypeal margin slightly less re-
flexed but of approximately the same shape; elypeal carina and frontal
carina less pronounced than in male majors but of the same con-
figuration. Pronotum considerably less convex with pronotal pro-
tuberance scarcely evident, disc often with minute scattered punctures.
Except for slightly more pronounced punctures and setae on elytra
and other parts of body, the male minors do not differ conspicuously
from the male majors.
Fremates.—Length 5.5 to 6.4 mm., width 3.0 to 3.6 mm. Very
similar to male minors, the major difference being in the slightly more
elongate shape of pygidium and in the last abdominal segment which
is not narrowed medially. Little indication of a pronotal protuber-
ance; the punctures and setae usually more in evidence than in the
male. Little apparent difference in other aspects (carinae, legs, etc.)
between the male minors and the females.
Typr.—Lectotype, present designation, USNM 1300, a female
specimen collected at Crescent City, Fla., July 15 by H. G. Hubbard;
deposited in USNM in 1896 as type.
BEETLE GENUS ONTHOPHAGUS—HOWDEN, CARTWRIGHT 37
TypE LocaLity.—Crescent City, Fla.
SPECIMENS EXAMINED.—159.
DistriBuTION.— (See fig. 5.)
Fioripa: Crescent City, 4 mi. north of High Springs, Gainesville, Leesburg,
Lutz, Miami, Stemper. souTH CAROLINA: Tillman.
Remarks.—This moderate-sized species is quite distinct from other
American Onthophagus. It can be separated by its uniformly shining,
dark brownish-black dorsal surface, the presence of small tubercles
with setigerous punctures at their bases on the head and near the an-
terior pronotal angles, the smooth shining dise of the pronotum, the
smooth margin of the fore tibia between the four teeth, and the lack
of pronounced sexual differences, the male majors having only a widely
rounded pronotal protuberance and the forelegs being the same length
as those of the female. Pronounced tubercles, setae, and punctures
on the head, pronotal angles, and elytra will separate typical poly-
phemi from the west Florida, Alabama, Mississippi subspecies sub-
sequently described.
O. polyphemi polyphemi and its subspecies have been collected only
in the burrows of the gopher tortoise, Gopherus (Xerobates) polyphemus.
Adult specimens have been collected in March, June, July, and August,
appearing freshly emerged in March. Hubbard (1894, p. 305) stated,
“T did not find this beetle in the few galleries examined in the winter,
and it was probably in pupa at that season.”’ In July it was not rare.
Se heal PTA haga
\Tr A
; I ny”
goo
qa oa :
Ficure 5. Distribution of species of Onthophagus:
O knausi Brown @ polyphemi polyphemi Hubbard
X — hopfneri Harold ©. polyphemi sparsisetosus, new subspecies
38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
One of the burrows produced 21 specimens. ‘‘Its larva was not seen.”’
At the present writing the larva is still unknown.
Collecting even the adults is a difficult job, best accomplished by at
least partial excavation of the gopher burrow. In the spring it is
often unnecessary to excavate the burrow completely, the Ontho-
phagus being found 4 to 8 feet inside the entrance in the sand at the
sides of the burrow. Excavation of the complete burrow is quite
difficult because they are often 18 to 20 feet long and may reach a
depth of 8 to 12 feet. Hubbard, who first investigated the insects
associated with the gopher tortoise, wrote (1894, p. 303), ‘“The exca-
vation was in the loose yellow sand of our pine woods subsoil, and
when my exploration was completed, so large a pit had been dug that
a coach and span of horses might have been swallowed up in it.”? The
authors found Hubbard’s statement very true, the only detail seem-
ingly omitted that, when possible, the gopher seems to terminate its
burrow under the roots of a tree, adding to the already difficult job
of excavation.
Occasionally specimens may be procured by the simple process of
placing a trap can containing fermenting malt and propionic acid
inside the entrance of the burrow and then sealing the burrow with
cardboard, paper, sticks, and sand. ‘The next day, if the can is not
filled with sand by the tortoise, it often yields a number of the insects
residing in the burrow.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 4
Ficures 10-12.—Acleris species: 10, 4. clarkei, new species, holotype o, Cle Elum,
Kittitas County, Wash., Apr. 9, 1931 (J. F. Gates Clarke); USNM. 11, A. clarkei, new
species, allotype 9, Aweme, Manitoba, Apr. 18, 1905 (N. Criddle); USNM. 12, 4.
senescens (Zeller), variety. co, Half Moon Bay, Calif., Feb. 6, 1940 (W. H. Lange);
USNM.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 5
16
Ficures 13-16.—Acleris nigrolinea (Robinson): 13, Lectotype o’, no data; AMNH. 14,
Genitalia of the lectotype (slide 547—-Obr.). 15, &, Aweme, Manitoba, Oct. 20, 1904
(N. Criddle); AMNH. 16, &, St. Hilaire, Quebec, May 24; AMNH.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 6
Ficures 17-20.—Acleris disputabilis, new species: 17, Holotype co, Goldstream, British
Columbia, Oct. 16, 1902; USNM._ 18, Paratype, o, Duncans, Vancouver Island, British
Columbia, Sept. 17, 1913 (A. W. Hanham); USNM. 19, Paratype, o7, Victoria, British
Columbia, Apr. 15, 1910 (A. J. Croker); AMNH. 20,07, Spring Creek, 4000 ft., near Baker,
Baker County, Oreg., May 7, 1953 (J. H. Baker); AMNH.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 7
24
Figures 21—24—Acleris species: 21-23, 4. disputabilis: 21, 9, San Mateo, Calif., Mar.
6, 1942 (G.E. Pollard); AMNH; 22, 9, Hymers, Ontario, April; USNM; 23, 9, Salida,
Colo., Apr. 7, 1888; AMNH. 24, Species near 4. disputabilis, 9, Southwestern Research
Station of the American Museum of Natural History, 5400 ft., 5 miles west from Portal,
Cochise County, Ariz., Mar. 31, 1956 (Cazier and Ordway); AMNH.
PROC. U.S. NAT. MUS. VOL. 114
OBRAZTSOV, PLATE 8
Ficures 25-28.—Acleris maximana (Barnes and Busck): 25,
o, Robson, British Co-
lumbia, May 2, 1938; AMNH. 26, o&, Glenwood Springs, Colo.; AMNH. 27, 9,
Quamichan Lake, Vancouver Island, British Columbia, Apr. 4, 1916; USNM. 28, 9,
Plumas County, Calif., Apr. 16-23; USNM.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 9
; ae
%
&
31
Ficures 29-31.—Male genitalia of Acleris species: 29, 4. santacrucis, new species, holo-
type (slide prepared by A. Busck on Feb. 10, 1933) (see pl. 2, fig. 4). 30, 4. walkerana
McDunnough). Western Greenland, 69°45’ E., Aug. 22-25, 1949 (Mission P. E. Victor
H. de Lesse, 1949); slide 1-Obr.; Muséum National d’Histoire Naturelle. 31, 4. glov-
erana (Walsingham). Holotype, Sheep Rock, Siskiyou County, Calif., Sept. 3, 1871
(Walsingham); slide 5353; BM.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 10
Ficures 32-35.—Female genitalia of Acleris species: 32, A. santacrucis, new species. al-
lotype (slide 1-Obr., Jan. 28, 1959), Santa Cruz, Calif., Oct. 29, 1932 (Tilden); USNM. 33,
‘The above, detail of caudal part. 34, 4. implexana (Walker), holotype (slide 5355),
St. Martins Falls, Albany River, Hudsons Bay, Canada, 1844 (Barnston); BM. 35,
A. walkerana (McDunnough). Western Greenland, 69°45’ E., Aug. 22-25, 1949 (Mission
P. E. Victor H. de Lesse, 1949); slide 2-Obr.; Muséum National d’Histoire Naturelle.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 11
ye 7”
38 39 bacey
Figures 36-39.—Genitalia of Acleris nigrolinea (Robinson): 36, & (slide 565—Obr.), Aweme,
Manitoba, Apr. 4, 1904 (N. Criddle); AMNA. 37, o& (slide 564—Obr.), Aweme, Mani-
toba, Oct. 19, 1904 (N. Criddle); AMNH. 38, 2 (slide 557—Obr.), Aweme, Manitoba,
Mar. 25, 1905 (N. Criddle); AMNH. 39, 9 (slide 559-Obr.), Aweme, Manitoba, Apr.
26, 1905 (N. Criddle); AMNH.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 12
Ficures 40, 41.—Male genitalia of Acleris disputabilis, new species: 40, Satus Creek, 10
miles from ‘Toppenish, Wash., Apr. 14, 1956 (A. I. Good); slide 540-Obr.; AMNH. 41,
Slide 391—Obr. (see pl. 6, fig. 20).
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 13
Ficures 42, 43.—Male genitalia of Acleris disputabilis, new species: 42, Slide prepared by
A. Busck, Jan. 21, 1924 (see pl. 6, fig. 18). 43, No data; slide prepared by A. Busck,
Nov. 13, 1924; USNM.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 14
Figures 44-47.—Female genitalia of Acleris disputabilis, new species: 44, Salida, Colo.
(W. S. Foster); slide prepared by A. Busck, Nov. 14, 1924); USNM. 45, Slide 553—Obr.
(see pl. 7, fig. 23). 46, Jemez Springs, N. Mex., Apr. 8-15; slide 2—Obr., Nov. 2, 1959;
USNM. 47, Paratype, Victoria, British Columbia, Nov. 22 (A. W. Hanham); slide 563—
Obr.; AMNH.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 15
Ficures 48-50.—Male genitalia of Acleris maximana (Barnes and Busck): 48, Slide 556—Obr.
(see pl. 8, fig. 26). 49, Sierra Nevada, Calif.; slide 542-Obr.; AMNH. 50, Near Alpine
Ranger Station, 9,500-13,000 ft., Gunnison County, Colo., July 5, 1957 (F. and P. Rindge);
slide 185—Obr.; AMNH.
PROC. U.S. NAT. MUS. VOL.114 OBRAZTSOV, PLATE 16
Ficures 51-53.—Male genitalia of Acleris maximana (Barnes and Busck): 51, Wagon
Camp, Mount Shasta, Calif., June 12, 1939 (G. H. and J. L. Sperry); slide 552—Obr.;
AMNH. 52, Rampart, Alaska; slide prepared by A. Busck, Jan. 8, 1924; USNM. 53,
Fraser Mills, British Columbia, Sept. 11, 1921 (L. E. Marment); slide prepared by A.
Busck, Oct. 10, 1923; USNM.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 17
LD
te
Figures 54-56.—Male genitalia of Acleris maximana (Barnes and Busck): 54, Wellington,
British Columbia, April (G. W. Taylor); slide 544-Obr.; AMNH. 55, Wellington, British
Columbia (G. W. Taylor); slide 539-Obr.; AMNH. 56, Pennsylvania; slide 534—-Obr.;
AMNH.
PROC. U.S. NAT. MUS. VOL. 114 OBRAZTSOV, PLATE 18
99 60
Ficures 57-60.—Female genitalia of Acleris maximana (Barnes and Busck): 57, Slide
prepared by A. Busck, Oct. 4, 1924 (see pl. 8, fig. 28). 58, Wellington, British Columbia,
Mar. 24, 1903; slide prepared by A. Busck, Nov. 8, 1923; USNM. 59, Spring Creek, 4,000
ft., near Baker, Baker County, Oreg., June 10, 1953 (J. H. Baker); slide 424-Obr., AMNH.
60, Wellington, British Columbia, Mar. 28, 1903 (G. W. Taylor); slide 568-Obr.; AMNH.
Proceedings of
the United States
National Museum
SMITHSONIAN INSTITUTION - WASHINGTON, D.C.
Volume 114 1963 Number 3470
A REVISION OF THE NORTH AMERICAN
ANNELID WORMS OF THE GENUS CAMBARINCOLA
(OLIGOCHAETA: BRANCHIOBDELLIDAE)
By Ricwarp L. Horrman'
Introduction
The annelid worms of the oligochaete family Branchiobdellidae are
for several reasons of more than casual interest to the zoologist.
Owing to their singular mode of life as commensal or subparasitic
inhabitants of freshwater crayfish throughout the Northern Hemi-
sphere, they have departed in numerous respects from the more
typical families of the order, and have developed such a leech-like
habitus and appearance that for many years the Branchiobdellidae was
thought to be a family in the Hirudinea. The problem remains
unanswered: is the similarity of the two groups due to convergence
influenced by environmental factors, or do the oligochaetes represent
a way station along the evolutionary path taken by leeches? The
life history has been worked out for none of the species, nor do we
know more than a few inferential details about the distribution of any
of them. Branchiobdellids should compel the interest of investigators
1 Department of Biology, Radford College, Radford, Virgina.
271
22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
if for no other reason than the position of the group on the borderline
between commensalism and true parasitism, which ought to provide
good clues about the evolution of the latter. It is obvious, however,
that such studies can hardly be made until the classification of the
group has been worked out, and specific identities firmly established.
So far, no extensive work has been done in this direction.
The present study was rather in the nature of a test to see if the
species of a widespread and somewhat heterogeneous branchiobdellid
genus might be defined on the basis of such characters of the repro-
ductive systems as have been found reliable in many other groups of
invertebrate animals. Traditionally, genera as well as species in
the Branchiobdellidae have been based on such characters as body
form, shape of the jaws, even size of the animals. Clearly it would
not take long to exhaust the possible combinations of such features,
and by 1950 the taxonomy of American branchiobdellids had reached |
virtually an impasse. The major consideration underlying this
investigation, therefore, has been the determination of what con- —
stitutes a species in the genus Cambarincola, and the development of —
coherent diagnoses by which these units may be subsequently iden- |
tified. I feel that these objectives have been achieved with a fair —
measure of success.
An inquiry into characters of systematic value reveals that the —
most important appear to be the gross form as well as histological
nature of the spermiducal gland and the adjoining prostate gland,
and that these structures afford the basis for division of the genus
into sections and groups. The two largest sections seem mainly to
be composed of structurally generalized species on one hand and ~
presumably more specialized forms on the other. That a satisfactory —
grouping of species into a system approximating the evolutionary or
phylogenetic trends within the genus has been established is by no-
means asserted, however. That future studies will impose changes
and modifications seems inevitable, yet a start must be made!
The species of Cambarincola, as now defined on the basis of the —
male reproductive organs, are remarkably constant for soft-bodied
animals, and well-preserved material can be identified with virtually
100 percent confidence. Most of the species that are known from
sufficient material are structurally uniform over their entire area of
distribution, and the geographic ranges are in all cases entirely
consistent with established biotic or physiographic provinces. The ©
continent-wide distributions published by previous workers are
largely the result of misidentifications.
In addition to the purely taxonomic aspect of this work, it has been
possible to draw some inferences from present knowledge of structure |
ANNELID GENUS CAMBARINCOLA—HOFFMAN 273
and distributions concerning evolution in the group as well as the
possible routes by which branchiobdellids settled the North American
land-mass. I suspect that the worms came to this continent probably
in late Cretaceous times on primitive astacine crayfish from eastern
Asia, and that these crustaceans may have spread eastward across the
continent—giving rise to the more specialized cambarine genera in
eastern North America rather than in Mexico as postulated by students
of the Decapoda. The present discontinuity in the distribution of
Cambarincola, and the isolated, relict status of its most primitive
forms, are thought to be the result of fragmentation of the old habitat
conditions by climatic changes in the late Tertiary.
ACKNOWLEDGMENTS
This study of Cambarincola was made at the suggestion, and under
the sympathetic guidance of Dr. P. C. Holt. I take great pleasure in
acknowledging the extent of his personal interest and concern in
facilitating and furthering my investigations, always more in the role
of longtime friend than that of major professor. ‘The work was done
while I was a candidate for the Ph. D. degree in the Graduate School
of the Virginia Polytechnic Institute. Financial support was fur-
nished by a research grant (G—4439) to Dr. Holt from the National
Science Foundation.
I am likewise under an obligation to Dr. Fenner A. Chace, Curator
of the Division of Marine Invertebrates, U.S. National Museum, who
provided working space and the opportunity to study type specimens,
and later kindly loaned the bulk of the Museum’s holdings in branchio-
bdellids for study.
It is finally necessary to acknowledge the contribution of numerous
collectors who have obtained the material with either a direct or in-
direct interest in branchiobdellids, particularly Mr. C. W. Hart, who
made a special effort to obtain topotypes of Astacobdella philadelphica.
Review of the Literature
The first American species of Cambarincola to be described was
placed in the genus Astacobdella, a group of Palearctic leeches (Leidy,
1851). Since that time, the generic concept has been slowly but
progressively refined.
In 1894, J. P. Moore described several new species of branchio-
bdellids from eastern United States, and in his paper discussed Leidy’s
earlier name under the combination Branchiobdella philadelphica,
and published an outline drawing of a specimen. The majority of
Moore’s work on this group involved species of other genera, although
in a brief reference in his paper on the leeches of Illinois (1901) he
274 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
created the new combination Bdellodrilus philadelphicus. 'There is,
of course, little close affinity between philadelphica and Bdellodrilus
illuminatus.
Thus, by the beginning of the present century, only a single species
of Cambarincola had been described among the total of six branchio-
bdellids then known from America. In 1912 a new impetus to the
study of the group came in the form of two important papers. One
of these was a synopsis of the entire family by Umberto Pierantoni,
which listed all of the species known to him up to that time, and
included the description of a new species, Branchiobdella americana,
from Texas and North Carolina. This form is almost certainly a
species of Cambarincola as the genus is defined today. Almost
simultaneously, a short paper was published by Max M. Ellis in which
Cambarincola was established as a new genus for the reception of the
new species macrodonta from Colorado.
The description of macrodonta is a good one, and includes illustra-
tions of the jaws and a partial diagram of the male reproductive
system. Comparison was made largely with philadelphica, which was
one of the originally included species, and which was differentiated
largely on the basis of peristomial characters.
During the next several years, Ellis accumulated data from his own
collections and that of the U.S. National Museum. In 1918 he pub-
lished a brief annotated list of branchiobdellids obtained in northern
Michigan, and provided a key for their identification. This treatment
included Cambarincola philadelphica and a new species called
C. vitrea, although the latter was not formally described and the nota-
tion was made that a complete description was then in press. In
the following year appeared Ellis’s major work, a summary of the
branchiobdellid material of the U.S. National Museum. This paper
included a discussion of variability in various systematic characters,
photographs of entire animals, and the descriptions of several new
genera and species. A key was given for the known forms of
Cambarincola, with notes and records on established species and full
descriptions of C. vitrea, C. inversa, and C. chirocephala, the last two
names published for the first time.
Following the appearance of this useful paper, Ellis diverted his
interests into other channels, and the systematic study of our branchio-
bdellid fauna languished for several decades. During the last years of
the 1930’s, Clarence J. Goodnight took up the study of the group,
publishing several papers between 1939 and 1943. His major work is _
a synopsis of the North American branchiobdellids, which appeared
in 1940. Here he accounted for 21 species in 9 genera, with 4 of the |.
species being described as new. Insofar as Cambarincola was con- 4:
cerned, Goodnight admitted the genus in virtually the same sense as |
ANNELID GENUS CAMBARINCOLA—HOFFMAN 975
did Ellis. It was placed in the new subfamily Cambarincolinae with
7 other genera and, in the key to these genera, placed next to Xiro-
nogiton, solely on the basis of having an “‘accessory sperm tube’
(actually this structure is not present in Xironogiton at all). Cam-
barincola was then divided into two subgenera following the initial
dichotomy in the key of Ellis (1919, p. 256), with species in which the
“upper lip’ is entire composing the nominate subgenus, and those
with lobed peristomata forming a new subgenus Coronata which was
set up for philadelphica and chirocephala. In the subgenus Cam-
barincola, the typical species macrodonta was associated with inversa
and vitrea of Ellis, and a new species named C. elevata. Recent study
has indicated that both inversa and elevata are members of quite
different genera.
Subsequent to the appearance of his monograph, Dr. Goodnight
published three additional short papers in which new species of
Cambarincola were described: C. floridana in 1941, C. meyeri in 1942,
and C. macrocephala in 1943. In his 1941 paper on the branchio-
bdellids of Florida, Goodnight listed four species for the State.
In 1947, the study of branchiobdellids was taken up by Perry C.
Holt, and his Master’s thesis, published in 1949, treated the com-
parative morphology of two species (Xironogiton instabilius and
Cambarincola philadelphica) with respect to the male reproductive
systems. From the detailed findings of this study, it became evident
that the sexual organs, previously only casually mentioned in print,
represent a source of the first magnitude for taxonomic characters.
Holt’s doctoral dissertation dealt primarily with other branchiobdellid
genera, but included notes and studies on Cambarincola, and two of
the new species recognized were subsequently published under the
names C. branchiophila (1954) and C. macbaini (1955).
With the recent accumulation of extensive collections of specimens
from many localities, it has become possible to undertake extensive
systematic studies on American branchiobdellids, and as a preliminary
step in an overall general program, the holotype and paratypes of
Cambarincola macrodonta were restudied and the genus and species
redefined in a short paper published by Holt and Hoffman (1959).
Two of the most important papers recently published by Holt
treat the taxonomy and morphology of the genera Ceratodrilus
(1960a) and Ellisodrilus (1960b).
Names and Type Specimens
In general, the overall validity and stability of any systematic
revision increases proportionately to the number of type specimens
examined. ‘This is nowhere more true than in the case of the family
Branchiobdellidae, inasmuch as very few of the published descriptions
276 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
are at all explicit concerning details which now must be known for
the identification of species in the family.
I have discovered that Cambarincola has become something of a
catch-all name which has been stretched to cover a wide diversity of
species since its original proposal. So far, 13 species have been named
in the genus, listed chronologically as follows:
C. philadelphica (Leidy, 1851) C. floridana Goodnight, 1941
C. macrodonta Ellis, 1912 C. meyert Goodnight, 1942
C. vitrea Ellis, 1918 C. macrocephela (sic) Goodnight,
C. chirocephala Ellis, 1919 1943
C. inversa Ellis, 1919 C. gracilis Robinson, 1954
C. okadai Yamaguchi, 1933 C. branchtophila Holt, 1954
C. elevata Goodnight, 1940 C. macbaini Holt, 1955
Of the above names, I have seen the type specimens of all but two:
philadelphica and okadai. If Joseph Leidy prepared slides of his
specimens, their present location is unknown. ‘The specific concept
of philadelphica here adopted was derived from topotype specimens,
kindly obtained in Philadelphia by Mr. C. W. Hart, which agree in
every particular with the original description. Information on C.
okadai has been limited to what can be deduced from the description.
Types of the remaining 11 species are in the collection of the Division
of Marine Invertebrates, U.S. National Museum, and have been
studied through the kindness of Dr. Fenner A. Chace.
On the basis of the generic concept developed during the course of
this study, it has been determined that several of the foregoing species
do not belong in Cambarincola as now defined. ‘These are:
C. inversa Ellis: This species differs from C. macrodonta in details
of the bursa and in lacking a prostate gland, as well as in the form
of the jaws. Only the type slide has been available for study, but it
is evident that inversa belongs to a different, as yet undefined genus.
C. elevata Goodnight: The internal structure cannot be seen in the
poorly mounted holotype, but the recent examination of topotype
material shows that this species likewise is not congeneric with
macrodonta, and is being included instead in a new genus being readied
for publication by Dr. Holt.
C. macbaini Holt: This species, and some others as yet unnamed,
differs strikingly from the cambarincoloid forms in having a very
long, filiform, protrusible penis, and forms the basis of another new
genus soon to be proposed.
With these deletions, the number of valid forms in the genus to the
present is reduced to 10. The examination of a great number of
specimens in the Holt collection has disclosed a fair number of the
described species (6 of the 10) and even more undescribed forms, of —
which 12 are named in the following pages. The type specimens of ©
ANNELID GENUS CAMBARINCOLA—HOFFMAN OTF
these species are deposited in the U.S. National Museum, with para-
types retained by Dr. Holt.
In addition to the names in the preceding list, one other must be
recognized as a probable member of the genus. ‘This is the species
described in 1912 as Branchiobdella americana Pierantoni. The de-
scription is not adequate for recognition of the species, and was
probably composite in being based on specimens from Texas and from
North Carolina. Until Pierantoni’s material can be restudied and a
lectotype designated, the status of this name remains in doubt, but
the chances are good that it is based on specimens of Cambarincola.
Material Examined
The following summary of Cambarincola is based upon the exami-
nation of about 900 collections of branchiobdellids, some 800 of which
contained species of this genus. Over 1100 individual slide mounts
have been examined, each with an average of about eight worms per
slide. Thus, approximately 9000 specimens of Cambarincola have
actually been examined closely for structural details, and yet another
thousand have been handled while sorting preserved material prior
to making slides.
At least 90 percent of all this material is in the private collection of
Dr. Perry C. Holt, now housed at the Virginia Polytechnic Institute.
In addition, I have been able to examine the portion of the collection
of Max M. Ellis which was turned over to the U.S. National Museum.
These two sources constitute virtually all the specimens of branchi-
obdellids now available in North America.
Of the 22 species now accounted within the limits of the genus, I
have been able to examine the types of all but two, as well as the type
material of various other species described as Cambarincola but
properly referable to other genera. All the existing holotypes are in
the National Museum, paratypes of about half of the species are in
the Holt collection.
Methods of Preparation and Study
Although the structure of the reproductive systems undoubtedly
provides the best criteria by which species may be defined and their
affinities established, it requires far more careful observation and
study than has previously been expended in the mere determination
of body form and jaw shape; the aspiring student of the Branchiobdel-
lidae must be warned that the group is not one that can be compre-
hended with a few weeks of attention. The preparation of material
for study is not difficult, but observation of the internal characters of
even well-mounted specimens is almost always tedious. Probably a
278 PROCEEDINGS OF THE NATIONAL MUSEUM voL. 114
year of continuous study is the minimum prerequisite for any sort
of competence in the systematic study of the group.
Only well-preserved and accurately labeled material should be
utilized. 80 or 90 percent alcohol (which may include 1 percent of a
stock formalin solution) is suggested for initial preservation, and the
collector should avoid overcrowding the jar with crayfish. Too many
of the crustaceans in one container dilute the preservative and cause
maceration of the worms. Isopropyl alcohol may be used, full
strength, for preservation, although it is usually only about 70 percent
in strength, and fewer crayfish can be placed in a given volume.
Material so preserved may be left indefinitely in jars with the host
animals or isolated and restored in tiny shell vials plugged with cotton
and packed in larger jars. Each collection may be assigned a different
number which relates the specimen vial to collection data entered in
a book or on index cards. Either the collection number alone, or a
tiny label printed with a fine pen, will suffice to be included in the vial.
Cataloging systems are limited only by the ingenuity of the individual,
but in practice, the simpler a system the better.
In curating material, one removes the crayfish from the jars with
enough vigorous shaking to dislodge any worms that may be still
attached to the exoskeleton. It is also desirable to search the branchial
chambers and gill filaments for gill-inhabiting forms, either by cutting
open the carapace with stout scissors or by extracting the gills with a
curved-tip forceps inserted between the carapace and bases of the legs.
The gill material is then placed in a dish of alcohol and picked apart
under a low-power dissecting microscope. This done, the majority of
the alcohol is decanted carefully from the collecting jar to avoid loss
of any of the bottom sediment which is then transferred to a flat glass
dish and examined with a dissecting microscope.
In a given collection there may be but a single species or as many as
four, each representing a different genus. As a rule these species will
differ enough in habit form to facilitate easy separation, although in
some cases identification even to genus is difficult until slides are pre-
pared. In practice it is advisable to select good specimens of each
form for mounting, all of which can be processed together. From the
flat-bodied forms such as the species of Xironogiton and Xironodrilus,
it is desirable to choose specimens which are least curled or twisted.
From the cylindrical forms, which normally preserve in a sort of
crescent-shaped profile, one should select those which are most nearly
straight in the plane of the dorsoventral median body axis. Since
knowledge of the jaw form is important, it is useful to prepare at least
one specimen flattened dorsoventrally, or to remove the head from a
specimen and split it along one side to allow for subsequent spreading
at the time of mounting. This operation is easily done with a fine-
ANNELID GENUS CAMBARINCOLA—HOFFMAN 279
tipped scalpel or fragment of razor blade mounted in the end of a
matchstick. In very large specimens (over 5.0 mm. long when pre-
served), it is often difficult to observe the reproductive system in whole
mounts, and with a little practice it is easy to bisect the body between
segments VI and VII, and then dissect out the male organs to be
dehydrated and cleared separately.
Processing can easily be done in small glass dishes or similar con-
tainers. For ordinary whole mounts it is satisfactory to take speci-
mens from the collecting jar into 95 percent alcohol, then through one
change of absolute alcohol into clove oil. About 10 minutes in each
stage is sufficient. All the material from each collection can be car-
ried along in the same container, the fluids being added from dropping
bottles and removed by a fine-tipped pipette.
Specimens may be mounted singly on each slide, or all of those which
look alike mounted together, although this sometimes results in
several species on a slide. I have used fairly thick balsam in a drop
at the middle of the slide, subsequently spread out by the addition of
a little clove oil around its base. Cleared specimens are removed
from the clove oil with forceps or a bent needle and placed in the
balsam, with specimens about equally divided according to which side
is uppermost (to insure having at least one with the reproductive
systems on the upper side). If dissected heads are involved, they
can be spread in thicker balsam with the jaws uppermost and the
coverslip applied quickly.
Slides in the Holt collection are prepared with the mount in the
middle, leaving space for the catalog number and/or locality data
label at the right end, and for an eventual determination label at the
left. Following identification, an index card may be made out with
the species name, number, and collection data, and filed in syst2matic
order. Since a slide may contain several species, it is preferable that
slides be stored in simple numerical order.
In the study of specimens, the use of medium high power magnifica-
tion is mandatory, with a range of from about 60 to 600 diameters
most desirable. Most of the present study has been made with a
fluorite oil immersion objective, 40 <,N.A. 100, corrected to a working
distance of 1.5 mm., permitting the examination of relatively thick
preparations inaccessible to the ordinary high dry and oil-immersion
objectives.
Drawings are easily made with a camera lucida attachment, and
measurements with a calibrated ocular micrometer.
In many instances, the student will find it impossible to determine
single specimens with confidence. Contents of the gut may be so
Opaque as to conceal the sex organs, or the jaws may be in an un-
satisfactory position. Specimens preserved in weak alcohol may be
280 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 114
macerated internally. Sometimes thick or opaque specimens can be
utilized by turning the slide over and focusing on the other side of the
animal, or by dissolving the balsam away in clove oil or xylol and
remounting.
Very smal] animals, or those with complicated reproductive systems,
must sometimes be imbedded in paraffin and sectioned. Well-known
histological techniques, using a hematoxylin-eosin stain, suffice for
serial sections. I have found no advantage in staining specimens for
whole mounts.
Often it will be found necessary to derive specific characters from
several specimens, there being few mounts in which all the desired
details can be seen on a single animal.
Taxonomic Characters
The taxonomic utility of various anatomical systems in the Bran-
chiobdellidae has been discussed in the literature at least four times.
Insofar as earlier workers were concerned, general body form and
shape of the jaws provided sufficient basis for the recognition of species,
an opinion which unfortunately has survived in some quarters nearly
down to the present time. That internal anatomy might furnish
characters of really fundamental importance was first intimated by
the stil] unsurpassed account of Bdellodrilus wluminatus by J. Percy
Moore in 1895. Almost two decades later, Ellis (1912) relied to some
extent on the form of the male reproductive systems in the diagnosis
of his Cambarincola macrodonta, but in his subsequent work of 1919,
Ellis largely reverted to nonsexual characters in the definition of
genera and species. In this practice he was followed by C. J. Good-
night, and not until 1949 was the study of reproductive morphology
revived by Perry C. Holt. Subsequent work by Holt, and that done
under his direction by the present investigator, has been predicated
upon the assumption that characters of the male reproductive systems
provide the most reliable indices of evolution within the group, and,
therefore, the best means for the definition of species and genera.
Ellis (1919) devoted considerable attention to the form of the jaws,
the nomenclature of their dentition, and their mode of evolution, as
well as to pharyngeal diverticula, intersegmental septa, and form of
the gut. No further treatment of taxonomic characters appeared
until 1935, when Yamaguchi provided a detailed consideration of
various useful details, including the body form and ornamentation,
jaw structure, and the internal characters mentioned by Ells.
Yamaguchi also discussed variations in the reproductive systems,
but not in as much detail as might be desired.
Goodnight’s monograph of 1940 reviewed the work of both Ellis
and Yamaguchi, as well as several of their predecessors, and en-
ANNELID GENUS CAMBARINCOLA—HOFFMAN 281
deavored to evaluate the systematic value of the previously used
characters. Goodnight, however, added little to our knowledge of
this general subject, and his references to the structure of the sexual
organs are meager. Finally, on the basis of his own knowledge of
branchiobdellid anatomy, Holt (1953) published a review of previous
work, and introduced information compelling greater attention to the
reproductive systems.
The following discussion of anatomical characters which appear to
be of taxonomic significance summarizes the points of interest which
have been considered during the study of Cambarincola and related
genera. It has been found necessary to review each character regard-
ing its development in the family at large, and then particularly as it
occurs within the more circumscribed limits of Cambarincola.
I. Bopy Form
Branchiobdellids vary greatly in form, but the typical appearance
is that of an elongate fusiform cylinder, with a distinctly set-off head
and a caudal sucker, both of variable dimensions but normally smaller
than the head. In general each body segment is composed of two
subsegments, of which the posterior is usually a little larger in diam-
eter than the other. Several variations from this form occur: in
Xironogiton, Xironodrilus, and some related genera not yet formally
segregated from the latter, the body is strongly compressed dorso-
ventrally, broadest near the caudal end. In Xironogiton the body
outline is unusual in being very abruptly narrowed anteriorly, having
something of the shape of a tennis racket. In the poorly known
genus Triannulata, each body segment is said to have three subseg-
ments instead of two, although an approach to that condition occurs
in the large species Cambarincola macrocephala. A terete body such
as described in the first sentence characterizes most species of bran-
chiobdellids, including all Cambarincola, and is probably the primitive
condition in the family. Flattened bodies are doubtless specializa-
tions, although genera so formed are not always specialized in other
particulars.
II. Bopy S1zz
Owing to the normally extensive degree of contractability char-
acteristic of most oligochaetes, the subject of body size and propor-
tions is not an easy one to treat satisfactorily. Branchiobdellids in
life are able to double or halve their normal length during motion,
and preserved and contracted specimens give an inadequate idea of
the living animal. Yet with proper preservation, contraction is not
excessive and species can be compared in general terms of relative
size. Known species in the family range from about 1 mm. up to
18 mm. or more in length. Intraspecific variation tends to increase
282 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
with increase in size; smaller species usually remain within very close
limits to their average. The range in size in Cambarinecola is as great
as for the rest of the family, with no particular affinities being re-
flected by the development of very large or very small forms, both of
which occur in groups unrelated by other characters. Both size ex-
tremes appear to occur only in mountainous regions, whereas species
of moderate dimensions tend to be widespread in lowland areas. In
general, so far as Cambarincola is concerned, body size is often a good
specific character, usually associated with other structural features,
and is sometimes useful in identification.
III. Bopy ORNAMENTATION
The vast majority of branchiobdellids are basically similar in ex-
ternal appearance, but in several forms the segments are provided
with dorsal elaborations in the form of fleshy digitiform lobes of
various length, or even large median processes which are distally
branched. Species so ornamented have traditionally been thrown
into the genus Pterodrilus without any consideration of other char-
acters, and that “genus” has become a sort of assemblage of incon-
gruous forms most of which are quite unrelated.
In many species the peristomium is produced into four lobes of
variable size and length on the dorsal side. These structures reach
their maximum development (among American species) in the genus
Ceratodrilus, becoming nearly as long as the head. In Cambarincola,
lobation of the peristomium occurs in various forms, and has been
used and misused in the past for separation of species and subgenera,
even though the significance of the character has been misunderstood.
Cambarincola philadelphica has been considered by both Ellis and
Goodnight to be very variable in this respect, but in actuality the
variation ascribed to that species is due to a confusion of several dis-
tinct species under one name. Well defined peristomial lobes (‘‘ten-
tacles’’) occur in association with otber characters and reflect specific
distinctness of worms so endowed. They do not alone indicate rela-
tionships, however, for they occur in several groups of the genus
Cambarincola as well as in Ceratodrilus and Stephanodrilus.
IV. Jaws
Branchiobdellids are the only Oligochaeta having sclerotized
mouthparts (in the form of a dorsal and a ventral piece) of variable
size and shape, but generally provided with caudally directed denta-
tions or cusps on the posterior margins. (The name “jaw” is used
only provisionally for the buccal armature of branchiobdellids, with
the realization that it is certainly not homologous even with the
mouthparts of polychaetes.) Ellis (1919) postulated that the
ANNELID GENUS CAMBARINCOLA—HOFFMAN 283
primitive condition involves subequal jaws in which the cusps are
similar in size and shape, and that specialization influences enlarge-
ment of the median tooth of the upper jaw. He devised a system
of nomenclature to reflect size order of the teeth, and is responsible
for the term ‘dental formula” by which the number of teeth can be
stated numerically (e.g., 5-4), indicating that the upper jaw has
a median tooth and two more on each side while the lower jaw has
two paramedian teeth plus one lateral cusp on each side. However,
other arrangements are common, particularly in Oriental species.
The American Ceratodrilus has small transverse jaws each with 6
to 8 equal dentations, and genera such as Xironogiton are likewise
small-jawed but usually with a smaller number of teeth. Ellis studied
variation in the jaws of what he considered to be Cambarincola
philadelphica and found considerable individual variation, although
it is now known that he had confused several species under that name.
Insofar as Cambarincola is concerned, there is considerable latitude
in the ranges of variation, individual and geographic as well as specific.
Individual variation is largely a function of age, with small specimens
of a species having proportionately larger jaws in relation to head
size, with more acute dentations. Older specimens tend to have the
lateral cusps obscured or worn away. Geographic variation affects
the relative length of the jaws in such species as Cambarincola witrea.
The typical jaw arrangement in the genus is one of essentially
equal-sized jaws, the dorsal with a large median tooth and two
or four small lateral cusps, and the ventral jaw with two large
paramedian terminal teeth and a pair of small lateral dentations.
Various departures from this basic pattern occur, however, aifecting
both the relative size and dental formulae, and a rough classification
can be drawn up as follows:
Jaws isomorphic, having the same outline in dorsal aspect, and the dental formula
5-5 or 38-3. In lateral aspect, the jaws are mirror images. The species
so characterized can be called ‘“Shomognathous,’’ and they occur sporadically
in different sections of the genus. C. fallax is a good example.
Jaws anisomorphic, being dissimilar in armature, the dental formula being
5-4 or some other combination of odd and even numbers. This general
class is further divisible according to the relative size and shape of the teeth.
Teeth virtually subequal, presumably the primitive condition, occurring in
C. vitrea and some related forms. Such species are ‘“homodontous.”
Teeth dissimilar, the median dorsal and paramedian ventral dentations enlarged,
this being the most typical condition in the genus, and can once more
be divided according to the relative size of the jaws:
Jaws similar in size in both lateral and dorsal aspects, this including the
majority of species such as C. philadelphica, macrodonta, ingens, etc.
Jaws markedly dissimilar in size, the dorsal jaw up to four or five times as
bulky as the ventral. This is presumably an advanced stage of speciali-
zation, and species so characterized may be called ‘‘heterognathous.”
284 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 114
V. Mate REPRODUCTIVE SYSTEM
The various species of Branchiobdellidae depart but slightly from
a basic organization of the reproductive organs. The function of
producing spermatozoa and collecting and conveying them to the
exterior is accomplished by the same organs and structures in all
members of the family, although the individual parts of the system
are subject to variations which, when considered collectively, afford
a fair opportunity for systematic diversification. The genital systems
of species in the genera Bdellodrilus (Moore, 1895b), Xironogiton
and Cambarincola (Holt, 1949), and Ceratodrilus and Ellisodrilus
(Holt, 1960a, 1960b) have already been elucidated in print. The
treatment of Ceratodrilus is of particular interest in providing a
standardized and morphologically correct terminology to replace
the various haphazard names which have been used or misused by
previous workers. The following terminology is that developed by
Holt (1960a). The sequence of the reproductive organs proceeds in
order from the testes outward. Reference may profitably be made
to the accompanying diagrammatic representation (figs. 1 and 2) of
the arrangement.
ADD
ai
EB
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SEGMENT: |
ea) is SIN Ga ee
Ficure 1.—Diagrammatic representation of the organs, other parts, and nomenclature of
the reproductive systems in Cambarincola. (For explanation of abbreviations see
figs. 2-5, p. 288.)
ANNELID GENUS CAMBARINCOLA—HOFFMAN 985
Testes. The spermatogenic tissue of branchiobdellids is con-
centrated into one or two pairs of testes located on the posterior face
of segmental septa 4/5 and 5/6, or only on 5/6. As arule the testes are
perceptible as such only in immature animals, sexually mature adults
show the condition in which the gonads have liberated morulae into the
coelom, the fluid of which becomes filled with masses of spermatozoa
and disintegrating blastophore material. As a rule, however, the
sperm masses are easy to observe in most whole mounts, particularly
as they tend to become oriented in clusters before the openings of the
funnels, and provide evidence concerning the distribution of the testes.
At least one genus (Branchiobdella) is provided with testes only in
segment v1, a condition upon which Goodnight (1940b) has founded a
subfamily Branchiobdellinae. That this difference is a fundamental
one has been established by the recent examination of a European
species of the genus, which has revealed additional peculiarities of the
reproductive system as well. All the North American genera which
have been studied in detail have two pairs of testes, and thus belong to
Goodnight’s subfamily Cambarincolinae. With Branchiobdella still
very poorly known taxonomically, it is premature to postulate which
of these two conditions is a primitive one.
Funnels. Spermatozoa are collected from the coelomic spaces of
the testicular segments by the modified ends of the efferent ducts,
which assume the general shape of a laboratory thistle-tube funnel—
a subglobose enlargement, a subterminal constriction, and a slightly
flared free margin. The entire structure is composed of a single layer
of densely ciliated epithelial cells, and apparently varies in size and to
some extent in shape among different species and genera. Normally
the funnels are located in the lower posterior portion of the segments,
their openings directed dorsolaterad and easily detected by the
densely aggregated spermatozoa. Owing to the difficulty of obtaining
precise measurements from whole mounts, and the general similarity
of the funnels throughout the genus Cambarincola, their possible
taxonomic utility has not been carefully investigated.
Efferent ducts. Each funnel is attached to a slender tubular
efferent duct, which proceeds ventromesad into the vicinity of the
bursa in segment vi. Here it unites with that of the opposed funnel,
forming the deferent duct. The efferent ducts serving the funnels of
segment v penetrate the septum of segments v/vi near its ventral
margin, mesad to the funnels. Those confined to segment vi are to
be found in the ventral portion immediately caudad to the bursa.
Deferent ducts. These conduits, formed by the union of a pair of
the smaller efferent ducts, are histologically similar to them but
average considerably larger in diameter. Originating in the general
region of the atrial portion of the bursa, the deferent ducts proceed
286 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
more or less directly dorsad to merge with the ental end of the spermi-
ducal gland, either more or less abruptly, or by way of an attenuated
lobe of the latter organ. Holt (1954) has remarked considerable
difference in the diameter of the deferent ducts between C. branchio-
phila and C. philadelphica, suggesting that in sectioned material it
may be possible to note size differences of taxonomic significance.
Spermiducal gland. The union of the two deferent ducts brings us
to an abruptly larger and more intricate portion of the sperm transfer
system. This is a rather plump and heavily glandular organ of very
variable size, shape, and proportions, composed internally of a single
layer of glandular epithelium. It is located in all species on one side
of segment vr, lying in the coelomic cavity between the gut and body
wall and readily visible in well-cleared whole mounts. In all species of
Cambarincola, the spermiducal gland is oriented in a distinctly oblique
plane across the segment, with the ental end cephaloventrad in loca-
tion, and the ectal end caudodorsal. In contracted specimens the
orientation may be dorsoventral; eversion of the bursa tends to draw
the ectal end ventrad, bending the gland into a C shape or alining it
horizontally.
The precise function of the spermiducal gland remains in some
doubt, although from its histological character we can assume it
contributes some material beneficial to the passage of spermatozoa or
to the accomplishment of sperm transfer. Anatomically, it is to be
considered a specialized section of the deferent ducts both prior to,
and ectad of their commissure into a single conduit. In some genera,
such as Branchiobdella, the deferent ducts enter into the gland quite
near its ectal end, with the main bulk of the organ greatly prolonged
entally and variously coiled in the coelom. In Cambarincola and its
related genera, the gland is basically Y or T shaped, the ental branches
forming the places of entry of the deferent ducts, and this arrangement,
from a morphological standpoint, is probably a reflection of the
primitive form in early branchiobdellids. Departure from this
arrangement seems to occur chiefly in the genera which are concom-
itantly specialized in other respects as well; in the spermiducal gland
we have, therefore, a useful sort of yardstick for gaging affinities.
Assuming that a Y-shaped gland is a generalized condition, we can
likewise essay an arrangement of the species of Cambarincola into
eroups or series of progressive specialization, culminating with the
gland taking a simple tubular form with little or no lobing at the
entry of the deferent ducts so that the Y shape is essentially lost.
This morphological progression is one frequently employed in the
following classification of the genus. When the entry of the ducts
is marked by an acuminate production of the gland, so that the
transition is a gradual one, the gland is regarded as “lobed” and the
ANNELID GENUS CAMBARINCOLA—HOFFMAN 287
lobes are called anterior or posterior deferent lobes according to the
position of the deferent ducts which enter them. The posterior
ducts collect spermatozoa from the sixth segment, the anterior ducts
from the fifth.
Prostate gland. This tubular structure is the “accessory sperm
tube” of Ellis and other workers, but its function is almost certainly
at least analogous to that of the prostate gland in other animal
eroups. Anatomically it originates at the junction (in most cases)
of the spermiducal gland and the ejaculatory duct, and extends
entally along the gland, both closely invested with a common (peri-
toneal) membrane. The histology of the prostate varies considerably.
In some species it is superficially almost identical with the spermiducal
gland in appearance, as regards both the size and apparent compo-
sition of the cells. In others there is a remarkable difference, one
visible even under low power magnification, in that the prostate is
made up of very large vacuolated cuboidal epithelial cells apparently
with little or no secretory contents.
The actual size and shape of the prostate gland varies to a con-
siderable extent, chiefly in diameter and length with respect to the
spermiducal gland, and this variation is often of considerable diag-
nostic importance. It is of course necessary to make certain that
the true dimensions are determined by a careful observation—the
prostate is often partially concealed or its ental end turned away
from the major axis, creating a much foreshortened effect. In at
least one case, the diameter of the prostate in comparison with that
of the spermiducal gland provides the major basis for separation of
two species of the genus.
Entally the prostate ends blindly, although in the group in which
it is histologically distinct from the spermiducal gland, there is a
terminal development in the form of a clear bulb generally about
the same diameter as the prostate proper, but occasionally somewhat
rudimentary (in primitive species) or quite enlarged in specialized
forms. The relationship of terminal bulb to vacuolated epithelial
cells is so constant that the one is prima facie evidence of the other
even when conditions prohibit direct observation of both!
Within the family Branchiobdellidae, the prostate gland apparently
typifies the group of genera clustered around Cambarincola and may
therefore serve a useful purpose as a tribal or subfamilial character.
It is considered to be, in its simplest form, an outpocketing of the
spermiducal gland which has, in the course of evolution, become
histologically differentiated and altered in size and shape (from a
broad short process to a long, slender, tubular one). I suspect that
in at least one specialized branch of this part of the family, that which
has produced Ceratodrilus, the tendency has been toward consolidation
653871—63——_2
288 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Figures 2—5.—Cross-sections of male reproductive organs. 2, Reproductive systems of
Cambarincola shoshone, showing typical orientation and location of the major organs
within segments v and v1; 3, section through bursa of C. ingens in normal position;
4, cross-section through bursa of C. ingens in everted position; 5, cross-section through
spermiducal gland, prostate, and part of ejaculatory duct in C. philadelphica. Histology
semidiagrammatic, but size and shape of individual cells shown correctly. Abbreviae
tions: A, atrium of bursa; ADD, anterior deferent duct; ADL, anterior deferent lobe
ANNELID GENUS CAMBARINCOLA—HOFFMAN 289
of the prostate back into the spermiducal gland, so that all remaining
of it in existing species of that genus is the bulbar tip, occurring as
a protuberance on the dorsal side of the gland near its ental end.
Ejaculatory duct. The spermiducal gland discharges to the outside
through a tubular duct with distinctly muscular walls which is un-
doubtedly capable of spasmodic or peristaltic action, and which is
appropriately named the ejaculatory duct (Holt, 1949). Both the
structure and distribution of this duct vary greatly in the family.
In certain genera (e.g., Xironogiton), it is quite thin-walled, probably
the primitive condition. In at least one undescribed species referable
to Xironodrilus in the sense of Ellis, it is absent altogether. In
Cambarincola macbaini (for which a new genus is being proposed
elsewhere), it is greatly enlarged and filled with a mass of convoluted
tubing. From the morphological standpoint, the duct is simply a
segment of the main sperm conducting passage; we should therefore
expect the least modified condition to reflect the generalized or
primitive state. Insofar as Cambarincola in the strict sense is con-
cerned, the ejaculatory duct is basically similar through all the species—
of about the same proportionate length and with a moderately muscu-
lar wall. Ectally it merges gradually into the penial sheath of the
bursa.
Bursa. This, the outermost differentiated portion of the male
reproductive system, is fundamentally a muscular invagination of
the ventral body wall—invaginated to permit concealment of the
penis well within the body, and muscular to achieve extrusion of the
penis during copulation. The accompanying illustrations (figs. 3
and 4) afford some idea of the composition of the bursa, which is
divided into two major anatomical parts: the penial sheath and the
atrium.
In virtually all branchiobdellids the bursa is a subglobose to pyri-
form organ opening to the exterior at the midventral part of segment
vi. Typically it projects mesad into the coelom or is directed some-
what caudad. Its ental half is taken up largely by the penial sheath
(fig. 3, PS) which is the somewhat differentiated muscular area
surrounding the ectal end of the ejaculatory duct and the virtually
continuous penis. The latter is very variable in structure through
the family, taking the form of a simple truncate cone which terminates
the ejaculatory duct, or becoming gradually modified into a much
of the spermiducal gland; B, bursa; EB, ental bulb; ED, ectal duct; EJD, ejaculatory
duct; EP, ental process; P, penial part of bursa; PDD, posterior deferent duct; PDL,
posterior deferent lobe of the spermiducal gland; PR, prostate gland; PS, penial sheath
of bursa; SDG, spermiducal gland. The lower case letters a—d in figures 3 and 4 are
located at the same anatomical position in both drawings.
290 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
longer structure capable of being itself everted independently of the
bursa. The evolutionary trend in this particular is obvious enough.
In Cambarincola, the penis is invariably of the short cone type,
and does not, in repose, extend ectad beyond the major band of
circular muscle which approximately divides the bursa into halves.
During copulation it is carried to the exterior by eversion of the
bursa, which thrusts the penis and associated penial sheath down
through the atrium and to the position it occupies as a median projec-
tion in a circular, distally concave structure formed by the lining of
the atrium. This displacement of the atrium during eversion can be
appreciated by comparison of the guide letters @ through d in figures 3
and 4, showing that the inner wall of the atrium becomes the outer
surface of the everted bursa. As the penis is, in this genus, capable
of only limited penetration of the spermatheca of the copulatory
partner, it is evident that the animals are obliged to achieve and
maintain a very close ventral contact during the process of sperm
transfer.
Eversion of the bursa is accomplished partly by contraction of
the circular muscles of the ental half, and doubtless also in part by
internal pressure achieved by contraction of the segmental muscles.
Summary of the male sex organs. Holt (1949) has shown that
the entire male system, with the exception of the funnels and possibly
the efferent ducts, is ensheathed by peritoneum, and al] of the organs
so invested are probably derived from the body wall. The interested
student is referred to this paper for a more detailed treatment of the
reproductive system. For the present, I consider them chiefly from
the standpoint of their taxonomic utility.
It is postulated that the primitive arrangement for the family
consists of two pairs of testes, funnels, and efferent ducts, one pair
each in segments v and v1; two deferent ducts which merge into a
Y-shaped glandular enlargement, the spermiducal gland—which
probably lacks a prostate, and which does not extend entally beyond
the entry of the deferent ducts; a thin-walled, muscular ejaculatory
duct, and a simple, fusiform, eversible bursa which carries a short,
conical, unmodified penis in its ental half.
Except for the presence of a prostate, the animal which comes
closest to answering this description is a species of Cambarincola, to
be described in a following section. Species which are almost anti-
thetical to the foregoing are Jikewise known, but are members of
North American genera as yet undescribed. However, a general
progression away from most of the stipulated conditions can be found
within the confines of the genus Cambarincola, suggesting its status
as a dynamically evolving genus which has nonetheless retained, in
ANNELID GENUS CAMBARINCOLA—HOFFMAN 291
a relict status, species which approximate the postulated form of the
ancestral stock.
VI. Femace REepRopUCTIVE SySTEM
The female sex organs are much less intricate than those of the
male system, and are of considerably less significance in arriving at
groupings of species. Perhaps of greatest utility is the general form
of the spermatheca in providing supplementary points of difference
between related forms.
Ovaries. The ovaries of branchiobdellids are located in the coelom
of segment vir, and offer little for systematic use. The relative
amount of their development, of course, provides a measure of the
maturity of a specimen. In some species of the family, segment vir
is rather distinctly the largest of the body units, and may enlarge
anteriorly to somewhat overlap on the posterior part of segment vt.
Insofar as Cambarincola is concerned, however, no appreciable
differences are apparent either in the ovaries or the segment in which
they occur.
Spermatheca. The spermatheca, throughout the family, so far
as known, is a tubular structure formed by invagination of the mid-
ventral body wall of segment v, and of variable size and shape. It is
composed of several anatomically and histologically distinctive sec-
tions reflecting different aspects of the function of sperm reception,
storage, and discharge. In some genera, of which most are still
undescribed, the ectalmost part of the spermatheca is an enlarged,
muscular structure reminiscent of the bursa, and possibly discharging
a similar function. In Pterodrilus and Cambarincola, at least, the
spermatheca begins ectally with a thin-walled, muscular duct which
proceeds dorsolaterad around one side of the gut, terminating in a
variously enlarged, fusiform to globose reservoir or ental bulb, the
function of which is storage of spermatozoa. Histologically this part
of the spermatheca differs appreciably from the duct in being only
slightly muscular, and in gross appearance it is frequently semi-
transparent or clear-walled. The diameter of this portion is subject
to much variation, depending largely on whether or not it happens
to contain sperm masses; however, the ratio of its length to that of the
ectal duct seems to be relatively constant, and thus is available for
diagnostic use. In various members of the genus, the ental bulb is
terminated by an abruptly smaller lobe or process, the ental process.
This is composed of deeply staining, small epithelial cells containing
much granular material. These cells are doubtless secretory in func-
tion. As the ental process appears in general to be present and best
developed in specialized members of the genus, it may be provisionally
considered an evolutionary improvement in the spermathecal struc-
292 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
ture. Perhaps it provides some sort of secretion which facilitates
longer retention of viable sperm. Generally speaking, in species
which have been adjudged conservative or primitive on the basis of
other characters, the spermatheca tends to have a long, slender ectal
duct and a small, globose ental bulb without an ental process. More
specialized species reduce the length of the ectal duct, enlarge the ental
bulb, and add a glandular ental process.
Relationships Within the Family
So far, not enough is known of the branchiobdellids to permit the
formulation of any lasting concepts about the affinities of genera to
each other. A major difficulty les in the fact that heretofore the
classification has been based on characters which probably are of little
phylogenetic significance, and preexisting genera have been hetero-
geneous as well as very poorly known. With the establishment of
some comparatively severe generic definitions (only comparable,
however, to those used in other phases of systematic zoology), it
seems possible to summarize general impressions and ideas accumu-
lated during this study of Cam barincola.
Although the Branchiobdellidae is a normally homogeneous family
as regards general organization of body form and organ systems,
histology, and way of life, a very considerable variation affects the
structure of the reproductive systems. A basic dichotomy was made
by Goodnight (1940) in recognizing two subfamilies according to the
distribution of male germinal tissue. For species in which testes, or
the morulae which they produce, occur only in segment VI, Goodnight
proposed a subfamily Branchiobdellinae; for those in which sperma-
tozoa are produced in segment v as well as vi, he provided the name
Cambarincolinae. This difference, of course, had already been used
by Pierantoni (1912) as a generic character, and does appear to be a
fundamental distinction, particularly as it is reinforced, at least in the
one species of Branchiobdella which T have examined, by correspond-
ingly important differences in organization of the larger and more
ectal parts of the male reproductive system.
Another basically important anatomical distinction between genera
emphasized by several workers is the nature of the outlet of the
anterior nephridia. In some genera the two ducts open to the out-
side independently of each other; in others there is a commonly shared
nephridiopore. However, the nature of this relationship is often (Gn
fact usually) difficult to detect with complete confidence, and we are
denied the use of literature reports based on the study of whole mounts.
Tn cases where the nephridiopore has been carefully studied by the use
of serial sections, it does afford strong presumptive evidence of
relationships.
ANNELID GENUS CAMBARINCOLA—HOFFMAN 293
The basic similarities in structure, shape, arrangement, and his-
tology of the reproductive systems, however, certainly represent the
best reflection of affinities. That this statement is true within the
confines of genera is strongly supported by the findings outlined in the
systematic treatment which follows. ‘The thin, and usually sub-
jectively drawn, line between species groups and genera permits, I
think, an extension of the principle into so-called higher categories.
From the preceding discussions of characters, it will be recalled
that within the limits of Cambarincola one finds a fair amount of
variation within the reproductive system both in the gross and histo-
logical appearance of the individual organs. The casual observer
might suspect, perhaps with justification, that by placing major
emphasis upon different organs one could arrive at several entirely
different classifications. This is not only theoretically true, it is a
difficulty which has been a source of vexation since this investigation
was begun. The solution has been an arbitrary one, influenced in
no small measure by the more or less unconscious accumulation of
small impressions which collectively result in a conscious allocation
of species by the totality of their characters. After several abortive
classifications had been drawn up and found wanting, I struck upon
the one which immediately appeared satisfactory and this, which has
met the test of having to accommodate additional and unforeseen
species, is the one here used. The organization of the bursa is given
pre-eminence, within limits, in the definition of the genus.
In the preceding section I have discussed something of the vari-
ability of this structure among various branchiobdellids, and pointed
out that a surprising amount of diversification is to be found in such
a basically simple arrangement. There is now known to be a number
of species which share the fundamental “cambarincoloid” organization
of bursa, ejaculatory duct, spermiducal gland, prostate gland, two
deferent ducts, and four efferent ducts and funnels, in addition to
a terete body form and a generally similar appearance. However,
a detailed study of the bursa shows that on the basis of its several
modifications, these species can be classified into groups, in which
the component species are obviously quite similar and related in small
details as well as overall facies. That bursa structure is a character
of major importance is attested by the homogeneity of these groups,
which are certainly entitled to be called genera by any definition
but the most inclusive. It is now altogether likely that ‘“(Camba-
rincola” in the usage of Ellis will be found to correspond roughly to
the bounds of a subfamily[!] in terms of modern classification.
We have, then, to consider basically a number of species of North
American branchiobdellids in which the body is cylindrical, the
nephridiopore single, the spermatheca not divided or branched,
294. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
segments v and vi testicular, and the male sex organs consist of the
elements listed in the preceding paragraph. Discounting poorly
described species, there are about 18 forms, which Goodnight placed
in the 3 genera Pierodrilus, Cirrodrilus, and Cambarincola. Of these
three, Cirrodrilus has been disposed of (Holt, 1960a) in a careful paper
which restores the correct name Ceratodrilus to the American species
and elucidates the anatomy of the genus. This genus differs from
all others in the collective characters of (1) eversible penis, (2) virtual
absence of the prostate gland, (8) great elaboration of segmental and
peristomial epidermal tentacles.
Cambarincola in the usage of Ellis and Goodnight is of course
heterogeneous. ‘The genus in the strict sense includes only those
species which do not have segmental ornamentation, in which the
bursa, but not the small penis, is eversible (or extrusible), the ejacula-
tory duct is not strongly modified, and the prostate gland is present
and functional. This combination of characters rules out three
known species originally described in the genus: inversa Ellis, elevata
Goodnight, and macbaini Holt. Each of the last two named repre-
sents a distinct generic type, now being defined and readied for publi-
cation by Professor Holt. We have studied the types of C. inversa
and regard it, too, as worthy of generic rank, but action is deferred
pending the acquisition of fresh material for sectioning.
Pterodrilus is likewise heterogeneous. The species alcicornus and
distichus (with two additional undescribed forms) are basically very
similar internally. P. mezxicanus is still unknown save from the
poorly preserved type specimen. But P. durbini of Ellis is remarkably
different, and belongs to the recently described genus Hilisodrilus
(Holt, 1960b), along with a second, previously unknown related species,
E. clitellatus.
With all of these eliminations and realinements which have resulted
from a close comparative study of important internal organs, we are
left with a genus Cambarincola in a strict sense (which nonetheless
is now known to contain no less than 21 species), and a genus Ptero-
drilus with two known and two more undescribed forms; and insofar
as internal structure is concerned, these two genera might be con-
sidered identical.
Here the element of arbitrary decision has its hour upon the stage.
Some authors who may work with branchiobdellids at a future time
will perhaps desire to combine the two genera under the older name
Pterodrilus. My personal feeling is that although the two groups
are beyond peradventure very closely related, the evidence suggests
that the species of Pterodrilus represent an extreme specialization
of some early embranchment of the Philadelphica section of Camba-
rincola. Evolutionary recency is suggested by the small size of the
ANNELID GENUS CAMBARINCOLA—HOFFMAN 295
species, greatly reduced jaws, moderate to elaborate development of
segmental ornamentation, and extreme enlargement of the prostate
both with respect to its size in comparison with the spermiducal
gland, and as regards the great size of the individual cuboidal cells.
Critics of a narrowly defined genus might object that Pterodrilus
differs less from the Philadelphica section of Cambarincola than the
latter does from the Mesochorea section. But this is a matter of
personal preference, and something which must be settled by the test
of future usage. The recognition as a genus of a specialized offshoot
of some diverse genus has ample historical precedent. The lizard
genus Uta is generally recognized as only a modification of one of
the groups of Sceloporus, and doubtless a long list of similar evidence
could be marshalled in support of the Pterodrilus-Cambarincola
relationship.
Unfortunately, as regards other genera, relationships are not so
clear. The tendency toward development of an eversible (and
ultimately an extrusible) penis seems clearly a specialization, but
one which had perhaps been achieved independently and at different
times. Within the group of genera which are so endowed, there
exists considerable variation as regards presence or absence of the
prostate, modification of the ejaculatory duct, and modification of
the spermatheca. Since these genera are composed largely of very
small worms, with every indication that a great many remain to be
found and studied, it would be premature to venture any opinion on
their affinities.
Whether the genera Triannulata and Stephanodrilus (the latter in
the sense of Goodnight’s usage of it for a Californian species) differ
from Cambarincola is something which remains for future settlement.
Phylogenetic Considerations
The following remarks constitute an attempt to summarize the
inferences which can be reasonably drawn from our present state of
knowledge of the genus Cambarincola. It seems relatively safe to
assume that probably most of the more common species of the genus
have been described, although a number of localized forms undoubt-
edly remain to be discovered.
Some criteria have been set up for the evaluation of certain diag-
nostic characters against the standard of a hypothetical ancestral
condition (cf. pp. 281-291). On the basis of these criteria, it is possible
to consider some species as primitive and some as specialized in the
two largest sections of the genus. By restricting comparisons to
the members of a given section, rather than the genus as a whole,
we find that the presumptive conservative forms tend to be scarce,
296 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
localized, and on or near the periphery of the range of the group or
section.
Most of the United States, east of the 105th Meridian, is occupied
by at least two species of Cambarincola, although as a rule the area
shared by any two given species is not extensive (mesochorea and
vitrea have the greatest territory in common). The number of species
tends to increase in certain regions—to five in the Ozark area and to
seven in southwest Virginia (the Pacific Northwest, still largely
unknown, must be omitted from consideration but it probably also
has a large number of species).
Now the interesting inference to be drawn from known distributions
in the genus is that—in comparison with other members of their
respective subgeneric section—none of the species of the Appalachian
fauna can be considered as primitive, such forms rather occurring
in lowland regions where two or three widespread forms of Cambarin-
cola are the dominant and often the only branchiobdellids. In
short, morphologically primitive species do not occur in the regions
having the richest branchiobdellid fauna at the present time.
It will be recalled from a previous section (p. 290), that the general-
ized species of Cambarincola represent to a considerable extent the
hypothetical form and structure of the familial archetype, and that
members of the Mesochorea section of the genus lack specializations
in virtually all of the major diagnostic characters. It is therefore
particularly significant that these species are, in a sense, now known
largely as relicts strung out along a highly probable route of migration
taken by cambarine crayfish in their occupation of eastern North
America. The two most primitive known species of the genus are
endemic to the Ozark region. . \
)
Ficures 39-42.—Structural details of Cambarinccla philadelphica (Leidy). 39, Lateral
aspect of typical specimen; 40, jaws of the same specimen, dorsolateral aspect; 41,
jaws of another specimen, lateral aspect; 42, reproductive systems, specimen from
Frederick Co., Maryland.
Minnesota, and south through the Appalachians and Piedmont to
the Carolinas. An additional form appears to inhabit the Catskill
region and adjacent areas, this is generally similar to the typical
form but is apparently consistently smaller over a distinct geographic
range. At several localities it occurs together with the large phila-
delphica. Whether it represents (1) a depauperate northern phase
of the main population, (2) a distinct sibling species, or (3) merely
an assemblage of young specimens, remains to be worked out by
another investigator able to collect and make field studies in New
York.
In central Kentucky and Tennessee occurs a form in which the
peristomium, particularly the dorsal half, is hypertrophied and
frequently widely flared, giving the head a campanulate appearance.
This is probably a perfectly good subspecies, but the details of its
overlap with philadelphica in eastern Kentucky ought to be worked
ANNELID GENUS CAMBARINCOLA—HOFFMAN 345
out prior to a formal recognition. The problem is complicated by the
nearby occurrence of chirocephala which also appears to intergrade
with philadelphica.
Representative specimens of each of the three forms of this species
are illustrated, and material is separated appropriately in the lists of
specimens examined. There is little or no appreciable difference in the
reproductive systems.
AFFINITIES.—The species of the Philadelphica subgroup are all
closely related and by no means easy to separate. A complication is
introduced by the certainty that additional forms will be discovered
and defined, particularly in the Appalachian region. Of the named
Ficures 43-46.—Structural details of Cambarincola chirocephala Ellis. 43, Lateral aspect
of typical specimen, Benton Co., Arkansas; 44, dorsal aspect of jaws, specimen from
Benton Co., Arkansas; 45, lateral aspect of jaws, specimen from Logan Co., Arkansas;
46, lateral aspect of reproductive systems, specimen from Logan Co., Arkansas.
forms, philadelphica, chirocephala, and macrodonta are most alike, and
future work may indeed show them to be only components of a wide-
ranging polytypic species. Some of the Kentucky material listed
under chirocephala shows a remarkable similarity in almost every
respect to philadelphica, and I have been able to demonstrate gradual
east-west clinal variation in both the jaws and spermiducal gland of
chirocephala. That these two species intergrade seems almost cer-
tain. Perhaps the establishment of an arbitrary ratio of jaw widths
will help define the ranges of the two, as well as their intermediates.
The relationships of philadelphica to macrodonta are also very close.
The entire peristomium of the latter is a good distinction between
well-preserved material of the two, and if present impressions are cor-
rect, the smaller bursa (less than the spermiducal gland in diameter) of
macrodonta should aid in recognition of the species. C. macrodonta
apparently occurs in South Dakota, philadelphica in Wisconsin.
346 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Obviously the northern part of the Mississippi Valley is an area of
some importance in establishing whether or not these two species
overlap or intergrade.
Distrisution.—Northeastern United States, from New York west
to Wisconsin, south through the Appalachian system as far as South
Carolina and Tennessee. Material of the typical form of the species
has been seen from the following collections:
MARYLAND: FREDERICK COUNTY: Catoctin Creek between Brunswick and Point
of Rocks on Md. Hy. 464, October 12, 1952, L. B. Holthuis (PCH 600).
MINNESOTA: WRIGHT COUNTY: Clearwater River at Minn. Hy. 152, in the town
of Clearwater, July 20, 1958, Holt (PCH 790).
NortH CAROLINA: WATAUGA COUNTY: Campus pond at Boone, June 24, 1948,
Mike Wright (PCH 128).
New YORK: FRANKLIN COUNTY: Chateauguay River just west of Chateauguay
on U.S. Hy. 11, May 21, 1951, D. W. Crocker and A. Gustafson (PCH 634).
LEWIS county: Mohawk River at West Leyden, August 22, 1952, Crocker (PCH
552); outlet of Brantingham Lake, 3.5 miles north of Lyons Falls, May 21, 1951,
Crocker and Gustafson, (PCH 639). tTIoGa county: Catatonk Creek at Candor,
August 29, 1952, Crocker (PCH 569). tTompxins county: Owasco Inlet, 3 miles
south of Groton, May 18, 1951, E. C. Raney (PCH 628). wWwyomine county:
Cattaragus Creek at Arcada, August 29, 1952, Crocker (PCH 570).
PENNSYLVANIA: PHILADELPHIA COUNTY: Wissahickon Creek in Philadelphia,
September 2, 1957, C. W. Hart (PCH 695).
SoutH CAROLINA: YORK couNTY: 4.0 miles south of Rock Hill on 8.C. Hy. 72,
April 20, 1958, Holt (PCH 748).
VIRGINIA: ALLEGHANY county: Dolly Ann Creek, 2 miles east of Covington on
U.S. Hy. 60, June 18, 1948, G. Ailstock (PCH 115). amuerRst county: Pedlar
River, 6.3 miles northwest of Forks of Buffalo on U.S. Hy. 60, November 11, 1947,
Horton H. Hobbs (PCH 82). augausta county: 7.6 miles north of Steeles Tavern
on U.S. Hy. 11, May 11, 1947, Hobbs (PCH 71). BEprorp county: Little Otter
River, 3.1 miles east of Bedford on U.S. Hy. 460, Holt and Bobb (PCH 73).
BRUNSWICK COUNTY: 2.9 miles north of Edgerton on Va. Hy. 140, May 31, 1949,
Holt and Bobb (PCH 217). BucKINGHAM couNTY: 9.6 miles south of Sprouse’s
Corners on U.S. Hy. 15, November 9, 1946, Hobbs and Hoffman (PCH 36).
CAMPBELL county: Evington, July 13, 1947, Hobbs (PCH 106). cRraiIG couNTY:
1.5 miles south of Paint Bank on Va. Hy. 311, June 24, 1948, Hobbs (PCH 97).
CHESTERFIELD COUNTY: Small stream below the lake in Camp Shawandasee, near
Chesterfield Courthouse, May 14, 1949, Holt (PCH 211). FLUVANNA coUNTY:
1.5 miles south of Palmyra on U.S. Hy. 15, November 10, 1946, Hobbs and Hoff-
man (PCH 114). Grayson county: Stream in pasture north of High Rock,
northwest of Independence, July 17, 1949, Holt (PCH 260). HANOVER COUNTY:
3.1 miles north of Ashland on U.S. Hy. 1, June 3, 1949, Holt and Bobb (PCH 230).
HIGHLAND COUNTY: Shaw’s Creek, 6.6 miles east of McDowell on U.S. Hy. 250,
September 27, 1946, Hobbs (PCH 113). Louisa county: 25.5 miles east of
Charlottesville on U.S. Hy. 250, July 2, 1948, Holt and Bobb (PCH 121).
MONTGOMERY county: Trillium Dale at Blacksburg, November 17, 1957, Holt,
Riggin, Hoffman (PCH 873); Bottom Creek, 0.5 mile northeast of Otey on Va.
Hy. 637, May 22, 1958, Holt and Hoffman (PCH 874). NELSON couNTyY: North
Fork of Rockfish River, 5.4 miles south of Afton on Va. Hy. 151, September 1,
1946, Hobbs (PCH 112). rocksBRipGE county: Tributary to Buffalo Creek, 5.3
miles south of Lexington on U.S. Hy. 11, May 11, 1947, Hobbs (PCH 75); 4.3
ANNELID GENUS CAMBARINCOLA—HOFFMAN 347
Ficures 47-56.—Structural details of three species of Cambarincola. 47-50, C. chiro-
cephala: 47, Male reproductive system, Carroll Co., Arkansas; 48, jaws, caudolateral
aspect, same specimen; 49, mzle reproductive system, Parke Co., Indiana; 50, jaws,
lateral aspect, same specimen. 51-54, C. philadelphica, heads in lateral aspect: 51,
Small northern form, Tompkins Co., New York; 52, typical form, Montgomery Co.,
Virginia; 53, amplistomate form, Jackson Co., Kentucky; 54, amplistomate form,
Macon Co., North Carolina. 55, 56, C. macrodonta: paratypes, Boulder Co., Colo-
rado, both specimens show the characteristic entire peristomium.
miles north of Glasgow on U.S. Hy. 501, May 11, 1947, Hobbs (PCH 78). rocx-
INGHAM couNTY: Swift Run, 3 miles east of Elkton, December 12, 1946, Holt
(PCH 14). scorr county: Troublesome Creek, 8.2 miles west of Gate City on
U.S. Hy. 58, June 16, 1950, Hobbs and Hart (PCH 366).
West Virainia: East River, date and collector not stated (USNM 17705)
[probably in Mercer County, southwest of Glen Lyn].
Specimens of the small New York phase (either a depauperate
form of philadelphica or possibly a different species) have been
examined from the following collections:
New YorK: CHENANGO couNTy: Otego Creek, just east of Oneonta, August
27, 1952, Crocker (PCH 561). onrrpa county: Tributary to the N.Y. State
Barge Canal just west of Utica, August 23, 1952, Crocker (PCH 536); Oswego
River at Rome, August 22, 1952, Crocker (PCH 583). tTroga county: Catatonk
Creek at Candor, September 6, 1959, L. R. McManus (PCH 928). TOMPKINS
county: Owasco Inlet, 3 miles south of Groton, May 18, 1951, E. C. Raney
(PCH 628). WARREN county: Stony Creek, in the town of Stony Creek, August
19, 1952, Crocker (PCH 546).
Material of the form with enlarged peristomium, discussed in the
foregoing text, has been examined from the following localities:
KENTUCKY: JACKSON CouUNTY: 1.8 miles south of Bond, July 29, 1958, Holt
(PCH 831). woure county: 1.7 miles west of Compton on Ky. Hy. 15, July 31,
348 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
1958, Holt, (PCH 846); large stream 5.1 miles east of Compton on Ky. Hy. 15,
July 31, 1958, Holt (PCH 847).
NortH CAROLINA: MACON couNTY: Spillway of Lake Ravenel, Highlands,
July 8-15, 1958, Hoffman (PCH 875); Buck Creek drainage 3.5 miles northwest
of Highlands, August 3, 1958, Hoffman (PCH 884).
TENNESSEE: CUMBERLAND county: White Creek, 0.25 mile northwest of
Alloway, May 3, 1959, Holt and Ford (PCH 903).
Cambarincola chirocephala Ellis
Fiaures 43-50, 57
Cambarincola chirocephala Ellis, 1919, Proc. U.S. Nat. Mus., vol. 55, p. 263,
figs. 18, 19—Goodnight, 1940, Illinois Biol. Monogr., vol. 17, no. 3, p. 37
(in part, [ll., Ind., Ky., Mo., ?Ala. records); 1940, Rep. Reelfoot Lake Biol.
Station, vol. 4, p. 171 (?Tenn. record).—Holt and Hoffman, 1959, Journ.
Tennessee Acad. Sci. vol. 34, p. 103.
TyPE SPECIMEN.—Holotype, USNM 177138, from Orconectes virilis
collected at Rolla, Phelps County, Missouri, by J. Barley. This
specimen is mounted flattened dorsoventrally, showing the jaws
clearly; the reproductive systems are obscured.
Diaanosis.—A small member of the Philadelphica group, recog-
nized by the distinctly disparate anisomorphic jaws; by the reduced
size of the spermiducal gland and prostate (both often shorter than
the bursa); and by the elongate, slender, gradually enlarged sper-
matheca.
Description.—Body small, ranging from 1.5 to 2.5 mm. in length,
the greatest diameter at segment vir; prosomites of all segments
larger than metasomites. Diameter of segment 1 less than that of
head; segments rx and x reduced, the caudal sucker abruptly enlarged,
equaling or surpassing diameter of head.
Head of normal size and form for the genus, larger than segment 1,
as long as the first three body segments combined. Peristomium
conspicuously set off by a basal constriction, flared, its margin pro-
vided with four small lobes on the dorsal side and two broad, lower
lobes on the ventral.
Jaws disparate in both dorsal and lateral aspects, anisomorphic;
broadly triangular. Dorsal jaw largest, 1.5 to 2 times as broad as
the ventral, the median tooth very conspicuous, lateral cusps small
or obscure. Ventral jaw subtrapezoid, with two small paramedian
apical teeth and a median sinus, and a smaller sublateral cusp on
each side halfway to the base.
Male reproductive system moderate in size, extending dorsal
through about three-fourths of segment v1. Bursa elongate, pyriform,
2 or 3 times as long as its greatest diameter, the latter more than
half the length of the ventral part of the segment. Atrial portion
of bursa somewhat longer than the penial sheath. Ejaculatory
duct long and slender, two-thirds to three-fourths as long as the
ANNELID GENUS CAMBARINCOLA—HOFFMAN 349
Ficure 57.—Distribution of the three species of the Philadelphica subgroup. ©, C. phila-
delphica; ©, “‘amplistomate” variety of philadelphica; (, C. macrodonta; @, C. chiros
cephala.
bursa, subequal in length to the prostate gland. Spermiducal gland
relatively small, subequal to or shorter than length of bursa, its
ental end recurved caudomesad prior to entry of the anterior deferent
duct. Prostate relatively large, nearly as long and thick as sperm-
iducal gland, the terminal bulb large and conspicuous.
Spermatheca long and slender, the ectal half curving laterad
around the gut, ental half more or less abruptly enlarged to 1.5 to 2
times the diameter of ectal portion, the apex bluntly acuminate, not
set off as an evident ental process. Total length of spermatheca
somewhat greater than diameter of segment Vil.
VaRIATION.—Individual variation in this species is very slight,
involving chiefly differences in size of the body. The proportions
of the reproductive tracts remain constant for a given locality.
There is, however, some evidence that the sexual organs are subject
to the influence of geographic factors affecting variation. There
appears to be an east to west decrease in the spermiducal gland
length as indicated by the illustrations (figs. 47, 49). In Indiana
350 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
and Kentucky material, this organ is fairly robust and somewhat
longer than the bursa, while in specimens from Arkansas it is reduced
in overall size and is shorter than the bursal length. The prostate
remains unaffected over this range, so that in the southwest it becomes
porportionately longer and broader with respect to the spermiducal
gland. Presumably this variation is of at least potential systematic
interest, but whether it is evenly clinal in nature, or broken at some
intermediate area (such as the Mississippi River) cannot be ascer-
tained on the basis of available material. Specimens from Illinois
are presently not at hand for study, and for the time being the matter
must rest at this point.
There appears to be a similar east to west cline in jaw structure,
with the disparity in size becoming most distinct westward into
Kansas and Oklahoma. Kentucky material tentatively referred to
chirocephala can scarcely be distinguished from philadelphica on the
basis of size of the jaws.
Distripution.—Cambarincola chirocephala appears to be basically
autochthonous to the Ozark highlands, now reaching its greatest
abundance in northern Arkansas and Missouri. It extends, however,
eastward into Indiana and Kentucky where its range meets that of
philadelphica, with which it perhaps intergrades. Most of the
Indiana localities are in the Wabash River system; one, interestingly
enough, is in the Maumee River system which now drains northeast
into Lake Erie. ‘The presence of chirocephala in this drainage reflects
either fairly recent stream piracy in the region or transgression of the
low divides by crayfish—either possibility seeming equally likely.
It will be a matter of interest to establish additional records for the
species in northwestern Ohio and eastern Indiana. Goodnight
(1940, p. 38) has reported this species from a number of midwestern
localities as well as from some very unlikely stations in New York
and Virginia, and such extralimital records must be presumed to
have been based upon misidentifications.
MATERIAL EXAMINED: 22 slides, from the following localities:
ARKANSAS: BENTON COUNTY: Sugar Creek at crossing of U.S. Hy. 62, July 6,
1958, Holt (PCH 768). cARRoLL county: South Fork of Dry Creek, 2.8 miles
east of Green Forest on U.S. Hy. 62, July 6, 1958, Holt (PCH 766). crRiTTEN-
DEN coUNTY: 14.6 miles south of Marked Tree on U.S. Hy. 68, July 5, 1958,
Holt (PCH 758). FruLTon county: Salem, July 29, 1941, Horton H. Hobbs
(PCH 310). LoGan county: West fork of Mill Creek, 4.4 miles west of Dela-
ware on Ark. Hy. 22, July 30, 1941, Hobbs (PCH 189). NEWTON coUNTY:
Buffalo River, 14 miles south of Harrison on Ark. Hy. 7, July 29, 1941, Hobbs
(PCH 175). sHAaRP couNTy: 3.2 miles southeast of Hardy, July 29, 1941,
Hobbs (PCH 176); about 9 miles west of Hardy, July 6, 1958, Holt (PCH 760).
INDIANA: ALLEN county: St. Mary’s River at Fort Wayne, Max M. Ellis
(USNM 17706). MoNRoE county: Bloomington, May 1915, Will Scott (USNM
ANNELID GENUS CAMBARINCOLA—HOFFMAN 351
17709). PARKE couNTY: 5.6 miles west of Bellmore on U.S. Hy. 36, July 26,
1958, Holt (PCH 807).
KANSAS: HARPER COUNTY: 10.7 miles west of the county line on Kans. Hy.
14, July 8, 1958, Holt (PCH 776).
KENTUCKY: CLARK COUNTY: 9.7 miles east of Winchester on Ky. Hy. 15,
July 30, 1958, Holt (PCH 839). sackson county: 0.1 mile south of the Owsley
County line on Ky. Hy. 30, July 29, 1958, Holt (PCH 834). MapIsoNn county:
Otter Creek, 9.3 miles north of Richmond on U.S. Hy. 227, July 30, 1958, Holt
(PCH 841). ows.tEy county: Traveler’s Rest, July 29, 1958, Holt (PCH 835).
POWELL couNTy: 1.4 miles east of Slade on Ky. Hys. 11 and 13; also Natural
Bridge State Park, both July 29, 1958, Holt (PCH 836, 837).
MIssouRI: PHELPS COUNTY: Rolla, J. Barley (USNM 17713, the holotype).
WASHINGTON couNTy: Irondale, August 28, 1931, Robert Rice (PCH 168).
OKLAHOMA: COMANCHE COUNTY: Blue Beaver Creek in Fort Sill, June 6, 1959,
J. W. Berry (PCH 905).
TENNESSEE: HUMPHREYS CouNTY: Hurricane Creek, 10.2 miles east of Waverly
on U.S. Hy. 70, July 5, 1958, Holt (PCH 756).
Remarxs.—Goodnight (1940, p. 37) has emphasized the pronounced
elevation of prosomites over metasomites as diagnostic of the species,
and most specimens are so formed, but contracted individuals of
many branchiobdellids likewise assume a distinctly annulated ap-
pearance. The larger size of the dorsal jaw is a good character for
recognition of chirocephala over most of its range, but it must be
recalled that in Kentucky this character loses must of its significance.
The affinities of chirocephala are discussed in connection with
C. philadelphica; in brief, the likelihood of a genetic continuum between
the two seems good. However, the actual details of this relationship
are not clear at this time. Whether chirocephala represents the fairly
recent modification of a westwardly migrating philadelphica-stock or
whether it is an allopatrically differentiating species now radiating
from its place of origin and intergrading with the original parent
population remains to be established.
The name chirocephala (Gk. chiros, hand, and cephalos, head) pre-
sumably refers to the dorsal lobation of the peristomium, but it is
not particularly appropriate since none of the material examined
is more distinctly lobed than most specimens of even C. philadelphica,
and in no way approximates the conspicuous tentaculation of fallax
and some other species. Ellis himself misidentified the Indiana
material cited as philadelphica, the records being originally published
in the same paper as the description of chirocephala! The Indiana
worms were merely mounted in such a way that the jaws could not
be seen in dorsal aspect.
Cambarincola macrodonta Ellis
FIGURES 37, 38, 55, 56, 57
Cambarincola macrodonta Ellis, 1912, Proc. U.S. Nat. Mus., vol. 42, p. 481, figs.
1-3; 1919, Proc. U.S. Nat. Mus., vol. 55, p. 257 (Colorado records only?).—
302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Hall, 1914, Proc. U.S. Nat. Mus., vol. 48, p. 190.—Goodnight, 1940, Illinois
Biol. Monogr., vol. 17, no. 3, p. 31 (description only); 1940, Rep. Reelfoot
Lake Biol. Station, vol. 4, p. 171 (?South Dakota record).—Holt and Hoff-
man, 1959, Journ. Tennessee Acad. Sci., vol. 34, p. 97, figs. 1-6 (redescription
of species).
TypE SPECIMENS.—Holotype and two paratypes, USNM 53794,
from Cambarus diogenes collected at Boulder, Boulder County,
Colorado, by Max M. Ellis.
Draanosis.—A moderately small member of the genus, differing
from other members of the Philadelphica group by the combination
of the slender, elongate outline of both body and head, the large
anisomorphic subequal jaws; entire peristomial margin; relatively
large bursa in comparison with the long, slender spermiducal gland,
and shortened prostate gland. The ectal spermathecal duct is much
longer in relation to size of the ental bulb than in the other related
species.
Derscription.—A distinctly slender and graceful worm, the body
only slightly thicker at maximum diameter than at the narrowest part,
tapering more distinctly caudally down to the smal] caudal sucker, its
diameter less than that of either head or segment 1. Segments less
than twice as long as broad, the length almost equally divided into
prosomite and metasomite, the former not at all larger in diameter
than the latter.
Head rather long and narrow, about equal to first three segments
combined, its greatest diameter a little greater than that of segment I.
Peristomium set off by a basal constriction, not evidently flared, the
margin divided into dorsal and ventral halves but neither half with
lobes or tentacles.
Jaws large and massive, subtriangular in dorsal aspect, aniso-
morphic, heterodont, the dental formula 5-4; dorsal jaw slightly
larger than ventral in lateral aspect, its median tooth subequal to the
paramedian teeth of the ventral jaw.
Male reproductive system of moderate size, occupying half of one
side of the coelom of segment vi or less, the bursa elongate pyriform,
its greatest diameter near the ental third instead of near the midlength
as in related forms; ejaculatory duct short, about equal to bursal
diameter. Spermiducal gland slender and elongate, oriented almost
horizontally in striking contrast to the oblique dorsoventral position
taken by the gland in most other related species, increasing slightly in
diameter entally, with the ental fourth of the length bent a little
ventrad. Deferent lobes small or absent. Prostate about two-
thirds as long as spermiducal gland and from one-fourth to one-half
its diameter, extending along the dorsal side entally about as far as the
entry of the posterior deferent duct.
ANNELID GENUS CAMBARINCOLA—HOFFMAN 353
Spermatheca composed of three parts: An elongate, slender,
muscular walled ectal duct, which extends about halfway up the side
of segment v; a subglobose or ovoid ental bulb; and a small terminal
ental process composed of glandular cells.
VARIATION.—The material which was studied from three localities
in Colorado is relatively homogeneous as regards structure of the
jaws and reproductive systems, variations in these details being
largely a reflection of difference in size of the worms. It is quite true
that branchiobdellids are very variable in size after maturity, suggest-
ing continuous growth through life, and actual measurements carry
less significance than ratios of the measurements.
Various combinations of measurements~’as functions of some
standard have been plotted graphically, and most of these show that
the material of macrodonta examined maintains a very constant set of
proportions despite changes in overall size. One suggestive detail
involves relative head size. Three specimens from Fort Collins,
Colorado, yield a head diameter to body diameter ratio of .60, .61, and
.62. In eleven other specimens from Boulder and Black Wolf Creek,
Colorado, the same ratio ranges from .71 to .99, with an average of .83.
These values, plotted along the horizontal axis of a chart with body-
length intervals on the vertical axis, separate out into two discrete
groups. Unfortunately, it cannot be ascertained whether we are
dealing here with true geographic variation or with the effects of
preservation. Within a fairly wide latitude, the degree of contraction
or distention of a branchiobdellid after preservation is influenced by
the strength of the alcohol used. The foregoing example is introduced
to remark the likelihood that with uniformly preserved worms in
good series, a future student of the group will be able to cope with the
problems inherent in the study of soft-bodied, muscular animals.
AFFINITIES.—Cambarincola macrodonta is without doubt closely
related to C. philadelphica, and if the two occurred as sympatric or
adjacent allopatric forms, it would be difficult to separate them with
confidence. In well-preserved specimens, the peristomial character
is most useful, but some individuals of philadelphica often do not show
the peristomial lobes, and it is easy to understand how various other
species have been misidentified by previous workers as macrodonta.
On the basis of my own limited knowledge of the species, I would
judge it to be an isolated, conservative remnant of the old late Tertiary
pre-philadelphica stock which has become isolated in the foothills of
the Rockies by recent climatic events which have produced the now
semiarid nature of the Great Plains.
DisrrisutTion.—This species has been recorded from 12 States
raiging from New Mexico to Virginia, from South Dakota to Louisiana.
The specimens which I have been able to re-examine, those identified
354 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 114
by Ellis in the U.S. National Museum collection, fall into three
categories: Very small, obviously immature worms; adult but mis-
identified specimens; and, finally, a few slides of worms conspecific
with the type specimen. The juveniles are at present unidentifiable
with certainty. Misidentified adults include C. mesochorea and some
other species which cannot be confidently identified but which are not
macrodonta on jaw shape. Undoubted specimens of the species are
listed as follows:
COLORADO: BOULDER couNTY: Boulder, September 1915, Max M. Ellis
(USNM 17667). LAaRimEeR county: Fort Collins, L. C. Bragg (USNM 17662).
yuMA county: Black Wolf Creek, near Beecher’s Island, October 1915,
B. Jaffa (USNM 17664).
In addition to these Colorado records, the specimen cited by Ellis
from Las Vegas, New Mexico (USNM 17661), appears to be a
macrodonta, but is so heavily stained that the sex organs cannot be
seen. There is nothing from a geographic point of view to preclude
the specimen being macrodonta.
Specimens from Muldon and Agricultural College, Mississippi, are
very much like Colorado material in every respect, yet I hesitate
to admit them to the list of macrodonta localities, at least until more
material from Mississippi or from intermediate areas comes to hand.
Cambarincola meyeri Goodnight
Figures 67, 68
Cambarincola meyert Goodnight, 1942, Trans. American Microse. Soc., vol. 61,
no. 3, p. 272, figs. 1-3—Holt and Hoffman, 1959, Journ. Tennessee Acad.
Sci., vol. 34, p. 108.
TYPE sPECiAMEN.—Holotype, USNM 20597, from Cambarus bar-
tonii collected in Raven’s Creek, near Lexington, Fayette County,
Kentucky, by Marvin C. Meyer (date unknown).
Dracnosis.—A small to moderate-sized member of the Philadel-
phica group characterized particularly by the (?) entire peristomium,
stout reniform spermiducal gland with a short slender prostate, and
peculiar form of the spermatheca.
Description.—Length of holotype about 3.5 mm. Body of nor-
mal proportions, segmental diameter about three times the length
at midbody; segments tv—vr of essentially the same size; prosomites
elevated above level of metasomites; caudal sucker quite small, its
diameter less than that of head or of segment 1.
Head of moderate size, as long as first three body segments and
slightly wider than segment 1. Peristomium set off by a basal con-
striction, slightly flared but the margin entire except for being divided
into a dorsal and ventral half; no evidence of lobes visible,
ANNELID GENUS CAMBARINCOLA—HOFFMAN 355
Jaws fairly small for size of head, heteromorphic, the dorsal jaw
with a large median tooth and two very small cusps at its base on
each side; ventral jaw with two large paramedian teeth and a single
small cusp at the outer base of each. Jaws virtually identical in
size, the dorsal very slightly wider at the base.
Male reproductive system characterized by the small globose bursa
(its diameter less than that of the spermiducal gland), by the very
slender, tubular prostate which is less than half as long as the sper-
miducal gland, and by the stout, acutely reniform shape of the
latter, its ental half curved strongly caudad and accentuating the
region of the posterior deferent duct.
Spermatheca composed of three major portions: 1, An enlarged,
muscular ectal region about half as large as the bursa; 2, a strongly
constricted cervical region of the ectal duct; and 3, a greatly
expanded, thin-walled, subglobose ental bulb.
VaRIATION.—The species is known only from two specimens.
Goodnight’s original description must have been based upon the
paratype which he retained, for the holotype is somewhat larger
than the published dimensions, its dorsal jaw 0.11 mm. in width
instead of 0.07 as stated by Goodnight for his specimen. Having
studied but the single specimen, I cannot say anything further on
the subject of variation.
AFFINITIES.—C. meyert is undoubtedly a member of the Phila-
delphica subgroup, but differs from the others at least in the slen-
der, short prostate and strongly curved, heavy spermiducal gland.
It is possible that Goodnight’s observation on the peristomium will
be confirmed, to constitute another diagnostic feature. Had no
specimens been available for study, my inclination would have been
to dismiss the name as probably a junior synonym of philadelphica
or one of its localized races.
Goodnight states that ‘“Cambarincola meyer is closely related to
Cambarincola vitrea Ellis, but differs in the structure of the upper
jaw.” I agree to a relationship between the two, but only as mem-
bers of the same genus; actually they seem to be very dissimilar and
fall into different groups of the Philadelphica section.
Distrirpution.—Known definitely only from the type locality.
Remarks.—The status of this species is by no means as well-
established as might be desired. I have at hand a large series of
well-preserved specimens from the vicinity of Livingston, Overton
County, Tennessee (Holt, leg.), which agree in virtually every detail
with the holotype of meyeri, not only in size and shape of the body,
but also in small details of the reproductive systems. The concord-
ance is such that conspecificity with the type specimen is almost
assured. Yet there appears to be a discrepancy in the jaw struc-
356 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
ture—isomorphic and bipentadont in the Tennessee worms, aniso-
morphic and pentatetradont in meyeri. Two likelihoods can be
considered: 1, The original types of meyert may have been compos-
ite. Goodnight’s account of the jaws was based on the paratype.
Unfortunately, the dentition cannot be made out with certainty in
the holotype, owing to its orientation on the slide; 2, the types of
meyeri may be aberrant specimens with respect to jaw structure, if
we assume that both are anisomorphic.
The question is one which can be settled only by the study of a
series of fresh specimens from the type locality. On the basis of
present knowledge of the group, it seems utterly unlikely that virtual
identity in form of the sex organs would be contraverted by a basic
difference in jaw structure.
For the present, however, I refrain from identifying the Tennessee
worms as meyer, remarking only their great similarity to the type of
that species, and commending the matter to someone having the
opportunity to secure topotypes of the species.
Cambarincola fallax, new species
Ficures 58-60, 62, 63
Cambarincola philadelphica (in part) Ellis, 1919, Proc. U.S. Nat. Mus., vol. 55,
p. 262.—Goodnight, 1940, Illinois Biol. Monogr., vol. 17, no. 3, p. 38.—Holt,
1949, Journ. Morphology, vol. 84, p. 535 et seq.
?Cambarincola okadai Yamaguchi, 1933, Proc. Imp. Acad., vol. 9, no. 4, p. 191;
1934, Journ. Fac. Sci. Hokkaido Imp. Univ., vol. 3, no. 3, p. 190.
TYPE SPECIMENS.—Holotype and four paratypes, USNM 29945,
from Cambarus longulus subsp. collected in Maiden Spring Creek,
about 1 mile east of Wardell, Tazewell County, Virginia, on June 19,
1959, by R. L. Hoffman. Additional paratypes from the same
collection, PCH 904.
Driaenosis.—A moderate to large species of the Philadelphica group
characterized by the combination of homognathous, pentadont jaws
and conspicuous elongate peristomial tentacles.
Dezscription.—A moderate to fairly large species, up to about
4.0 mm. in length. Body form rather slender, without distinct
enlargement in diameter in going caudad to the middle of the length.
Prosomites about twice as long as metasomites and very distinctly
larger in diameter, imparting a pronounced annulate body profile.
Segments 11 to vir usually of about equal diameter.
Head moderate in size, about as long as first three body segments
combined, its diameter about equal to that of segment 11, equal to
or slightly larger than diameter of caudal sucker. Peristomium
large, set off by a deep basal constriction, almost half the total head
length; the dorsal half often a little flared, with four distinct blunt
ANNELID GENUS CAMBARINCOLA—HOFFMAN 307
Ficure 58-61.—Two species of the Fallax subgroup, external appearance. 58, Cambar-
incola fallax, new species, paratype from Tazewell Co., Virginia, with peristomium
extended; 59, specimen from Seneca Lake, New York; 60, specimen from Pike Co.,
Georgia. 61, C. holostoma, new species, paratype from Highland Co., Virginia, showing
the characteristic entire peristomium.
elongate tentacles of varying length. Ventral half of peristomium
shorter, set off from dorsal by a deep lateral sinus each side, subdivided
into two broad lobes.
Jaws about 10 percent of the head length, isomorphic, equal in size,
heterodont; the median teeth distinctly larger than the lateral cusps
which are nonetheless very distinct even in lateral aspect, dental
formula 5-5, 5-3, or 3-3, the smaller figures occurring chiefly in old or
large specimens.
Male reproductive system typical for the group, the more ectal
organs of moderate size and occupying about two-thirds or less of
the coelom on one side of segment vi. Bursa elongate pyriform,
its greatest diameter at midlength; penial sheath abruptly merging
into the much narrower ejaculatory duct; latter half as long as bursa.
Spermiducal gland oriented almost dorsoventrally, extending down
to or beyond level of penial sheath, generally subreniform in outline
with an occasional enlargement homologous to the posterior deferent
lobe of other species; diameter of gland about equal to that of bursa,
but somewhat smaller in occasional specimens. Prostate gland long,
slender, its diameter half that of spermiducal gland near their junc-
ture; not extending entally as far as apex of the latter.
3508 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Spermatheca rather small, generally similar to that of OC. phila-
delphica, composed of a slender ectal duct and an ovoid or fusiform
ental bulb located about halfway up one side of segment v, no glan-
dular ental process detected.
VariaTion.—Individual variation in body form and peristomial
shape is shown in figures 58-60. The two most similar worms are
from opposite extremes of the range.
The male reproductive system varies somewhat more than in most
other members of this genus, particularly with respect to size of the
spermiducal gland, but nothing has been noted to indicate any sort
of geographic dispersion. Figure 62 shows the typical proportions
of the larger organs. The spermatheca is likewise somewhat vari-
able, particularly the appearance of the ental bulb. This, however,
is pretty clearly a reflection of the degree of distention by its contents.
Distripution.—Known from numerous localities throughout the
Appalachian uplift and adjacent Piedmont from western New York
south as far as central western Georgia. In the southern part of
the range fallax appears to be the most abundant member of the genus,
further north the records are more scattered and C. philadelphica
becomes dominant. Material has been examined from the following
localities:
GEORGIA: FANNIN county: 5.7 miles south of Morganton on Ga. Hy. 60,
November 6, 1958, K. Simonds (PCH 909). murray county: Holly Creek,
10.6 miles north of the Gordon County line on U.S. Hy. 411, April 16, 1958, Holt
(PCH 737). pike county: Large stream on outskirts of Zebulon, April 17, 1958,
Holt (PCH 740). wurre county: Small ravine, 1 mile south of Helen, July 11,
1958, Hoffman (PCH 877).
NortH CAROLINA: CHEROKEE couNTY: Beaver Creek, 0.5 mile northwest
of Andrews, June 9, 1959, Simonds (PCH 912). cnuay county: 3 miles south
of Tuni Gap on the Hayesville-Andrews road, June 5, 1959, Simonds (PCH 918).
TENNESSEE: MONROE couNTY: Small woodland stream, 7 miles north of
Madisonville on U.S. Hy. 411, April 16, 1958, Holt (PCH 738). PoLK county:
2.3 miles south of Ocoee on U.S. Hy. 411, June 9, 1959, Simonds (PCH 907).
VIRGINIA: ALBEMARLE County: Tributary to Rivanna River near Stony Point,
April 25, 1947, Holt and Hobbs (PCH 45, 46). BucKINGHAM couNTY: 9.6 miles
south of Sprouses Corners on Va. Hy. 15, November 9, 1946, Hobbs and Hoffman
(PCH 36). cHARLOTTE couNTY: 8.8 miles south of Keysville on Va. Hy. 15,
November 9, 1946, Hobbs and Hoffman (PCH 38). ames county: Sinking
Creek at Va. Hy. 700, near Newport, July 3, 1950, Holt, Tuten, and Kizer (PCH
407). PATRICK county: Shooting Creek, 1.6 miles south of the Franklin County
line on Va. Hy. 40, April 13, 1953, Hobbs (PCH 723). TrazewELL county: Maiden
Spring Creek, about 1 mile east of Wardell, June 19, 1959, Hoffman (PCH 904,
type series).
New York: TomPKINS county: Owasco Inlet, 3 miles south of Groton, May 18,
1951, E. C. Raney (PCH 628). County not located: Reeder’s Creek, 1949, L. C.
Goldstein (PCH 245).
ANNELID GENUS CAMBARINCOLA—HOFFMAN 359
67 2
“
LA ie
ea ee
Ficures 62-68.—Structural details of three species of Cambarincola. 62, C. fallax, new
species, reproductive systems, specimen from Giles Co., Virginia; 63, dorsal aspect of
jaws, same specimen, each jaw tilted slightly in opposite directions, dental formula
actually 5-5. 64, C. holostoma, new species, reproductive systems, paratype from
Highland Co., Virginia; 65, the same, jaws in dorsolateral aspect, same locality; 66,
the same, jaws in lateral aspect, same locality. 67, C. meyeri Goodnight, male re-
productive system, from holotype, Fayette Co., Kentucky; 68, the same, spermatheca
from holotype, the specimen considerably flattened in mounting.
In addition to the foregoing records, there are doubtless many
published localities for C. philadelphica which really apply to this
species.
Cambarincola holostoma, new species
Ficures 61, 64-66
Type sPeEcIMENS.—Holotype and four paratypes, USNM 29946,
from Cambarus bartonii and C. longulus collected in Crab Run, 4 miles
west of McDowell, Highland County, Virginia, on U.S. Hy. 250, by
L. B. Holthuis, October 25, 1952. Topoparatypes, PCH 599.
Dracnosis.—A member of the Falilax subgroup characterized by
the long, slender body, the prosomites of which are not distinctly
raised; by the completely entire, flared peristomium; and by the
slender, elongate, fusiform spermatheca.
663871—63——_7
360 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Description.—A small worm reaching a maximum length of,about
2.0 mm. in preserved specimens. Body form slender, the diameter
increasing gradually to segment vi which is subequal in bulk to seg-
ments vir and viir; caudal sucker somewhat larger than preceding
segment and about as broad as the peristomium. Prosomites up to
three times as long as the metasomites, but not of greater diameter.
Head about as long as the first three body segments combined, and
equal in diameter to segment 111, the peristomium set off by a very
strong basal constriction and distinctly flared, its margin entire,
without any trace of division into dorsal and ventral halves or into
smaller lobes. Head otherwise not visibly segmented externally.
Jaws similar, dental formula 3-3 with the median tooth long and
acute, the general appearance very similar to the jaws of C. fallax
but the formula perhaps more often 3-3 than in that species.
Male reproductive system basically similar to that of other species
of the Philadelphica group. Bursa rather long, equaling the length
of the spermiducal gland, the two subequal in diameter. Ejaculatory
duct modest in size, its length less than the diameter of the spermiducal
gland. Latter of normal proportions, without evident Jobation at
entry of deferent ducts. Prostate long, slender, about half the
diameter of spermiducal gland, which it joins slightly entally of the
entry of the ejaculatory duct.
Spermatheca slender and elongate, curving laterad and dorsad
around the gut, and becoming slightly wider but maintaining about
the same diameter almost to its end at a point near the middorsal
area of the segment, a distinct ental bulb not being well-developed.
VarIATION.—In the small amount of material examined from three
localities, some of it not well-preserved, there appeared to be little or
no variation in the diagnostic characters of the species.
AFFINITIES.—The relationships of this form with C. fallax and
O. philadelphica, with both of which it is sympatric, are unquestion-
able. It differs from both, however, in characters of the peristomium
and spermatheca. Closer relationship with fallaz is postulated on the
basis of jaw form, here considered to be a more fundamental character
than peristomial lobation.
Disrrisution.—Aside from the type locality, this species is known
from two collections from western Virginia, in the James and upper
Potomac River drainages.
_ VIRGINIA: CRAIG COUNTY: 1.5 miles south of Paint Bank on Va. Hy. 311, June 24,
1948, Horton H. Hobbs (PCH 97). RockinaHam county: Swift Run, 3 miles
east of Elkton on U.S. Hy. 33, December 12, 1946, Holt (PCH 14).
Remarks.—Further knowledge of the distribution of this localized
form may be of interest in providing an insight into the factors influenc-
ing its speciation.
ANNELID GENUS CAMBARINCOLA—HOFFMAN 361
Ficures 69-73.—Structural details of two species of Cambarincola. 69, C. heterognatha,
new species, paratype in lateral aspect; 70, the same, lateral aspect of jaws; 71, the same,
reproductive systems. 72, C. demissa, new species, body profile of paratype, Wise
Co., Virginia. Part of body wall shown cut away to indicate the size of the reduced
reproductive organs in situ; 73, the same, reproductive systems in lateral aspect, same
specimen.
The collection localities, all of which I have seen, are rather small,
swift, mountain brooks, somewhat different in character from the
larger and perhaps more placid streams in which fallax occurs most
abundantly. Conceivably the peristomium of holostoma represents
the development of (or retention of) a more efficient holdfast device
than the dissected and lobed mouth of fallax, a matter which would
certainly enhance the origin and maintainence of specific differences
by ecological factors.
The problem is recommended to someone having the opportunity to
study branchiobdellids and their distribution in the upper James River
system of western Virginia.
HETEROGNATHA GROUP
A separate group must be erected to represent on a coordinate
standing the very unusual and highly specialized new species described
362 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
below. In many of its structural features, such as the disparate jaw
size and shape, small body, reduced size of the male reproductive
system, and particularly the shortened prostate gland, this species
shows a combination of evolutionary specialities which occur only
singly in various other forms of the genus.
Cambarincola heterognatha, new species
FicureEs 69-71, 74
TypE sPECIMENS.—Holotype and paratype, USNM 29947, from
Cambarus sp. collected in a tributary to Big Wilson Creek, 4 miles
south of Mouth of Wilson on Va. Hy. 16, Grayson County, Virginia,
by Horton H. Hobbs, Jr., and C. W. Hart, June 14, 1950. Additional
paratypes indicated in the list of specimens examined.
Diacnosis.—Immediately recognizable by the remarkably dis-
similar jaws alone. In addition, the male reproductive system is re-
duced in size and the prostate gland is less than half the length of the
spermiducal gland.
Description.—A moderate-sized species, body length up to about
3.0 mm., somewhat fusiform in body outline, the greatest diameter at
segments vi and vit, the least at x.
Head of normal size and proportions, about as broad as long, its
diameter less than that of segment 1; peristomium only a little flared,
without marginal lobes; head indistinctly subdivided into two halves
by a slightly median constriction. Jaws very dissimilar in size, the
upper triangular with a large median projection and about eight times
the bulk of the lower (!) which is about equally quadrate in shape with
two small paramedian cusps.
Body outline fusiform, segments vi and vit largest, each somite
divided by a distinct complete constriction, the prosomites the larger
of the two subsegments and not, or only slightly, elevated over the
level of the metasomites.
Male reproductive system rather small, extending only halfway
up the side of segment vi. Bursa fusiform, widest near the middle,
about 1.5 times as long as broad, somewhat constricted at entry of
ejaculatory duct. Latter of moderate length and rather slender, less
than a third the bursa diameter. Spermiducal gland subreniform,
slightly broader entally, its diameter equal to or slightly greater than
that of bursa, the length a little greater. Prostate small and slender,
less than half as long as spermiducal gland, generally about a third as
long, the terminal bulb distinct.
Spermatheca elongate and slender, abruptly constricted at the mid-
length, the ental half with a rounded tip or a slight ental process, the
organ extending dorsad nearly to the dorsal side of the coelomic cavity.
ANNELID GENUS CAMBARINCOLA—HOFFMAN 363
Figure 74.—Distribution of Cambarincola heterognatha, new species, an abundant Appalach-
ian endemic species. Each spot represents collections for one county.
VarraTion.—There is little or no appreciable variation in this
species as regards size of body, shape of jaws, and general details of
the sex organs. There is some individual divergence in the relative
length of the prostate gland, which may vary anywhere from a third
to half the length of the spermiducal gland, irrespective of locality.
The single collection from Kentucky is interesting in that the prostate
of all specimens studied is basally much thicker than noted elsewhere
in the range of the species, attaining a diameter at its base at least
half that of the spermiducal gland. The Kentucky locality is con-
siderably removed from the main distribution of the species, and this
minor difference may reflect some significant microevolutionary de-
velopment due to isolation. Aside from this one departure, hetero-
gnatha must be considered a very homogeneous species despite its
considerable geographic range.
Disrripution.—The southern Appalachians, from northwestern
Virginia and adjacent West Virginia, south and west to central eastern
parts of Tennessee and Kentucky. Specimens have been examined
from the following localities:
KENTUCKY: ADAIR couNTy: 8.9 miles east of Columbia on Ky. Hy. 80, July
28, 1958, Holt (PCH 827).
TENNESSEE: CLAIBORNE COUNTY: 3 miles southwest of New Tazewell on Tenn.
Hy. 33, April 16, 1951, Horton H. Hobbs and W. R. West (PCH 540). cumBEr-
LAND couNtTy: Daddy’s Creek, between Crossville and Pikeville, August 1950,
Holt (PCH 419). wasHINGTON couNTy: Hartsell Cove, Buffalo Mountain,
364 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
August 1953, Holt (PCH 495); Sinking Creek in Horse Cove, January 10, 1954,
Holt (PCH 580).
NortH CAROLINA: WATAUGA COUNTY: 2 miles south of Vilas on N.C. Hy. 194,
June 14, 1950, Hobbs and Hart (PCH 350).
VIRGINIA: AUGUSTA CoUNTY: South River, 2.4 miles south of Waynesboro,
September 3, 1948, Hobbs (PCH 271). crara county: 1.4 miles west of New-
castle on Va. Hy. 311, June 23, 1948, Hobbs (PCH 99). FRANKLIN COUNTY:
Smart View Picnic Area, Blue Ridge Parkway, September 14, 1958, Holt and
Hoffman (PCH 891). GmxEs country: Cascades of Little Stony Creek, June 25,
1952, Holt (PCH 492, paratypes). GRrAYsoN county: Tributary to Big Wilson
Creek, 4 miles south of Mouth of Wilson, June 14, 1950, Hobbs and Hart (PCH
339, types). LEE county: Hardy Creek, 11.1 miles west of Jonesville on U-S.
Hy. 58, June 16, 1950, Hobbs and Hart (PCH 371). RocKINGHAM couUNTY:
Swift Run, 3 miles east of Elkton, December 1946, Holt (PCH 4). smyvH
county: White Top Mountain, September 5, 1951, John T. Wood (PCH 531).
TAZEWELL CouNTY: Bluestone River, 11.2 miles northeast of Tazewell on U.S.
Hy. 460, June 18, 1950, Hobbs and Hart (PCH 393, paratypes); Burkes Garden,
June 30, 1947, R. L. Hoffman and H. I. Kleinpeter (PCH 532); Maiden Spring
Creek, 1 mile east of Wardell, June 19, 1959, Hoffman (PCH 904). wasHINGTON
county: 9.5 miles south of Abingdon on U.S. Hy. 11, April 14, 1951, Hobbs and
West (PCH 432); 4.8 miles south of Abingdon, April 14, 1951, Hobbs and West
(PCH 435); North Bristol, January 1, 1954, W. A. Whittaker (PCH 581). wyTHE
county: Reed Creek, 3.3 miles southwest of Wytheville on U.S. Hy. 11, April 14,
1951, Hobbs and West (PCH 439).
West VIRGINIA: GREENBRIER COUNTY: Dry Creek, east side of Kates Mountain
at White Sulphur Springs, July 3, 1947, Hobbs (PCH 294). PENDLETON COUNTY:
5.8 miles east of Franklin on U.S. Hy. 33, July 30, 1949, Hobbs and Word (PCH
275). WYOMING couNTy: Barker Creek, 5.3 miles south of Tralee, July 12, 1947,
Hobbs and Wilson (PCH 95).
From the standpoint of major drainage systems, the vast majority
of the preceding records lie within the basins of the upper Tennessee
and Kanawha rivers, and southwest Virginia clearly seems to be the
center of abundance for this species. Peripherally, the records are
distinctly more spotty, although in equally well-collected areas.
Towards the northeast, the species occurs in the James and Potomac
drainage systems.
The single known locality for Kentucky lies in a region which was
intensively collected by Dr. Holt during July 1958, a fact which
permits the inference that heterognatha may exist in central Kentucky
only as a relict, and this fact, together with the generally sporadic
distribution of species suggests that the range may be in the process
of contraction, perhaps as a result of post-Pleistocene increase in
temperature in the southern Appalachians.
Nonetheless, it must be emphasized that the distribution of het-
erognatha is by no means well-known, and many new localities doubt-
less remain to be established.
Remarxks.—It has already been observed that hetferognatha is
endowed with a remarkable combination of presumably evolutionary
specializations. It is one of the easiest of branchiobdellids to recog-
ANNELID GENUS CAMBARINCOLA—HOFFMAN 365
nize, owing to the enormous dorsal jaw which dominates the entire
anterior half of the head. In no other known form in the family is
heterognathy carried to such an extreme.
Additionally, the spermatheca offers very good specific characters,
particularly its length and pronounced median constriction which
are evident in every specimen examined.
Presumably heterognatha is a highly modified member of the general
philadelphica-chirocephala stock, which began to evolve its characters
at a very remote time, or has been able to develop them more rapidly
than other species of the genus.
DEMISSA SECTION
A separate section seems necessary to reflect the status of
Cambarincola demissa, a very disjunct species which has little relation-
ship with other members of the genus.
Judged from body shape, small head and jaws, and general appear-
ance, this species is perhaps one modified as a gill-inhabiting form, and
therefore especially liable to various structural concomittances of a
specialized habitat. Unfortunately, we know nothing definite about
the preferred microhabitat of the species.
Perhaps the most unusual feature of this animal is the marked
reduction in the size of the reproductive systems, indicated in outline
in the habit sketch of an entire worm (fig. 72). All of the normal
organs are present, with the fine structure characteristic of the genus,
but both spermiducal and prostate glands are very small proportion-
ately, and the latter—although histologically not differentiated—
terminates in a small clear bulb. The penial sheath of the bursa
is much smaller, in relation to the atrium, than in any other member
of the genus.
DEMISSA GROUP
A monotypic group with the characters of the section. The only
species is a moderately small, corpulent-looking worm, probably
branchiophilus, known only from extreme southwestern Virginia.
Cambarincola demissa, new species
Figures 72, 73
TYPE SPECIMENS.—Holotype and four paratypes, USNM 29948, from
Orconectes erichsonianus and O. juvenilis collected in a tributary to the
Powell River at Big Stone Gap, Wise County, Virginia, by Horton H.
Hobbs, Jr., and C. W. Hart on June 17, 1950.
Diacenosis.—A moderate-sized (2.8-3.2 mm. long) cuneate species
of Cambarincola, differing from all other members of the genus in the
exceptionally small sex organs, the prostate gland likewise differing
366 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 114
from all others in terminating in a small clear bulb although not
histologically differentiated from the spermiducal gland.
Descrietion.—BKody strongly enlarged caudally, segments vr and
vil about twice as wide as segment 1, and tapering very abruptly to
the fairly large caudal sucker which is as broad as the head or segment
1. Segments very short, as little as a third of the body diameter at
segment VI; prosomites twice as long as metasomites but not elevated
above them.
Head as long as first three body segments, but smaller in diameter
and thus continuing the anterior attenuation of the body; peristomi-
um about a third of total head length, set off by a strong basal constric-
tion, its margin apparently broadly lobed but not extended into
projections or tentacles (all material slightly macerated). An addi-
tional more posterior constriction of the head occurs and imparts a
trisegmented appearance to the head in lateral aspect.
Jaws relatively quite small, about 7.0 percent of the head length,
the dorsal jaw slightly the larger and longer, with a distinct large pro-
jecting median tooth; ventral jaw likewise with a median tooth and
subsimilar in general form to the dorsal, at least in lateral aspect (none
of the available material is mounted in a way to show dorsal or ventral
surfaces of the jaws, but careful observation indicates that the dental
formula is probably 3-3 or perhaps even 1-1).
Male reproductive system very small, confined to the ventrolateral
portion of the coelom of segment v1, extending dorsal less than half-
way up one side of the segment. Bursa small, cordate, the atrial
portion making up most of its bulk, the penial sheath confined to the
ental fourth of the bursa and very small by comparison with that in
other species. Ejaculatory duct moderately long, its length about
equal to that of the bursa or spermiducal gland, its wall muscular but
of normal thickness. Spermiducal gland and prostate collectively
only about as large as the bursa, their histological structure similar
(small, glandular, basophilic cells), but the prostate terminates in a
small clear bulb presumably homologous to that so characteristic of the
Philadelphica section. Spermiducal gland short and broad, at most
only half again as long as the diameter; prostate slender but much
shorter than spermiducal gland. Latter entally-rounded, without
evident lobation at the entries of the small, slender deferent ducts.
Spermathecae equally reduced, extending less than halfway up one
side of segment v, the shape somewhat fusiform, expanding laterally
from the small ectal portion and maintaining essentially the same
size to the abruptly acuminate ental tip.
VARIATION.—Owing to the small amount of available material, all
of which is slightly macerated from the initial preservation in weak
alcohol, it is not possible to dwell at any length on the observed varia-
ANNELID GENUS CAMBARINCOLA—HOFFMAN 367
tion. Because of the small size of the sex organs, which are not par-
ticularly confined by the gut as in most other species, the spermiducal
gland and prostate in particular are liable to considerable freedom of
motion in the coelom, and preserved specimens show much variation
in the shape of these two structures due to a difference in perspective.
The spermatheca seems to have a constant shape, as does the general
body outline. The range of this species may be so limited that geo-
graphically influenced variability will be found negligible.
RELATIoNSHIPS.—There is no other species in the genus with which
demissa can be compared. C. branchiophila of the Mesochorea section
is known to be a gill form, and has small jaws and a cuneate body
form, but its sex organs are of normal size and the major parts of the
bursa correctly proportioned. It is difficult to derive demissa from
either of the other two major sections of the genus, although if com-
pelled to make a choice I should tentatively place it much closer to the
Mesochorea section as representing the culmination of evolutionary
tendencies in that ensemble. As this form is probably localized among
the high mountains of southwest Virginia, there is every reason to
presume that many other endemic species remain to be discovered,
and some of these may cast some light on the affinities of this curious
and disjunct little worm.
Distrisution.—Aside from the type locality, C. demissa is known
only from the following locality:
VIRGINIA: TAZEWELL cCouNTY: Bluestone River, 11.2 miles east of Tazewell on
U.S. Hy. 460; June 18, 1950, Hobbs and Hart (PCH 393).
Remarks.—The possibility of any host specificity on the part of
this species is negated by the fact that the type series came from a
collection of two species of Orconectes, the other known material from
a collection of two species of Cambarus. Presumably the factors
influencing the distribution of demissa are those of simple geography
and water conditions rather than the occurrence of the crayfish hosts.
I cannot, at this time, imagine what might favor the selection re-
sulting in the drastically reduced size of the reproductive systems,
unless it be that residence within the branchial chambers of crayfish
affords survival with a reduced number of progeny. Perhaps knowl-
edge of the ecology of the species will shed some light on this interest-
ing evolutionary problem.
The name demissa (Latin, humble, unimposing, modest) seems
appropriate in view of the general form of the animal as well as its
very moderately scaled reproductive systems.
Species of uncertain systematic position
Under this heading I include two species which are known to be
referable to Cambarincola, but which, for one reason or another, cannot
368 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
I> 2
ae
((
ee
ae
ee
oa Su LE Q VAG 79
Ficures 75-79.—Structural details of two species of Cambarincola. 75, C. floridana Good-
night, body in lateral aspect, camera lucida drawing from holotype; 76, the same,
jaws in dorsal aspect, from freehand sketch. 77, C. gracilis Robinson, body outline
of holotype; 78, the same, male reproductive system of holotype; 79, the same male
reproductive system, with bursa everted, paratype from British Columbia.
at present be allocated to a group or even to a section with assurance.
The type specimens of both have been studied and drawn; the data on
hand are summarized and presented at this time for the benefit of
future workers who may be able to collect at the type localities and
establish the identities of the names.
Cambarincola floridana Goodnight
Fiaures 75, 76
Cambarincola floridana Goodnight, 1941, Trans. American Microsc. Soc., vol. 60,
p70.
Typr spECIMENS.—Holotype, USNM 20570, from Procambarus
fallax collected in Taylor County, Florida, by Horton H. Hobbs.
This specimen is mounted laterally, but the reproductive system
cannot be made out with certainty.
RemarKks.—The bursa of this species is of considerably greater
size, in relation to the other organs, than in most other species of the
genus. Otherwise the general configuration suggests that floridana
may be a specialized member of the Philadelphica section.
ANNELID GENUS CAMBARINCOLA—HOFFMAN 369
Cambarincola gracilis Robinson
Fiaures 77-79
Cambarincola gracilis Robinson, 1954, Journ. Parasitology, vol. 40, p. 466, figs.
1-4.—Holt and Hoffman, 1959, Journ. Tennessee Acad. Sci., vol. 34, p. 103.
TYPE SPECIMEN.—Holotype, USNM 26110, from Pacifastacus
klamathensis collected at Whitman College, Walla Walla County,
Washington, by A. G. Rempel. Paratypes: USNM 26111, from the
Klamath River, Siskiyou County, California, and USNM 26112,
Burnaby, British Columbia.
Remarks.—This species was described and illustrated in some detail,
the account being more meaningful than most of the existing descrip-
tions. Not only the body form and jaws were drawn and discussed,
but the form of the reproductive systems as well, and a reasonable
comparison was made with C. inversa and C. macrodonta. It is un-
fortunate that equal care in the diagnosis of new species was not
expended by several of Miss Robinson’s predecessors.
I have examined the type material in the U.S. National Museum,
and cannot improve upon the original description except to note that
the bursa is of the typical cordate Cambarincola-form, and not a
eradual enlargement of the ejaculatory duct as indicated in Robinson’s
figure 1. Her figure 3, a reconstruction from serial sections, was made
from a worm with everted bursa.
The reduced size of the male sex organs is perhaps of some diagnostic
importance, the appearance in situ being reminiscent of that of C.
demissa. Miss Robinson failed to record the histological appearance
of the spermiducal gland and prostate, and, unfortunately, I neglected
to note the same detail when examining the type.
A re-examination of material of gracilis can readily establish the
status and taxonomic position of the species. Assuming that the
prostate is not differentiated (it is known to have no terminal bulb),
the species will fall into the Mesochorea section and either the Branchi-
ophila group or a new one of its own.
C. gracilis was recorded from ten localities in California, Oregon,
Washington, and British Columbia, a relatively extensiverange. Yet
curiously enough, Goodnight (1959) refers gracilis to a category of so-
called scarce, localized species in his most recent synopsis, while rank-
ing in the main part of the key some of his own species known from
but a single locality.
The paratype from California cannot be studied with respect to the
reproductive system. That from British Columbia, however, is well-
mounted, and appears to be correctly identified with the holotype.
Drawings made from the holotype and paratype are presented as an
aid for future recognition of the species.
370 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 144
Literature Cited
Evuis, Max M.
1912. €
d
Ficure 4.—Hydrobiosinae genus A: a, larval head and pronotum, dorsal view; 4, larval
mandibles, dorsal; c, femur through tarsal claw of larval foreleg, posterior face; d, larval
prosternum, ventral; ¢, left anal proleg of larva, lateral.
NEOTROPICAL CADDIS FLIES I—FLINT 463
Atopsyche species 3
This larva is very similar to that assigned to alconura, but differs in
lacking the ventral dark marks on the head.
Larva.—Length 10 mm. Coloration as in alconura, except central
stripe on frontoclypeus narrower and no dark marks ventrally.
MarteriAu.—Venezuela: Rio Cobre of the Catatumbo system be-
low La Grita, Mar. 31, 1942, L. P. Schultz, 2 larvae (USNM).
Hydrobiosinae genus A
FIGURE 4
In the material from Ecuador are two hydrobiosine larvae that differ
in many structural characteristics from the larvae of Atopsyche. The
most important differences are the fusion of the pronotal halves, the
reduction of the prosternite to a minute sclerite, and the disappearance
of a segment in the forelegs.
No genus other than Atopsyche is known to occur in Ecuador, so I
can not even speculate as to the generic placement of these larvae.
Larva.—Length 6 mm. Head yellowish, muscle scars darker,
posterior margin black (fig. 4,¢@). Labrum as in Atopsyche. Maxilla
about the same, except lateral part of stipes almost completely mem-
branous except at seta and mesal part strongly sclerotized toward base
and closely united with cardo. Labium as in Atopsyche, except dor-
sally where mesal strap lacks teeth and hypopharyngeal rods with only
5 or 6 teeth on strongly angulate apical portion. Mandibles with only
a single dorsal subapical tooth (fig. 4,5). Pronotum yellowish, pos-
terior and lateral margins black; lacking mesal suture (fig. 4,2). Pro-
sternum very small (fig. 4,d), in width only one-fifth of the distance
between legs. Forelegs chelicerate, but femur not twisted; femoral
process much narrower than that of Atopsyche; apparently tibia and
tarsus fused (at least only 1 segment obviously present); claw extend-
ing beyond tip of femoral process (fig. 4,¢). Abdomen generally as in
Atopsyche. Anal proleg lacking basoventral spine; claw sharply curved
ventrad, basal segment with a broad curved spine arising apicoven-
trally and a normal seta arising laterally at its base (fig. 4,e).
MareriaAL.—Kcuador: Stream 5 miles south of Antisana, 13,500 ft.,
Apr. 28, 1958, R. W. Hodges, 1 larva; stream 11 miles west of Pujili,
12,500 ft., Mar. 15, 1958, R. W. Hodges, 1 larva.
Family Glossosomatidae
The subfamily Glossosomatinae is present in the New World only
in the Nearctic region; the Protoptilinae, also widely distributed in
North America, is the only subfamily present in the Neotropics.
464 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
Hight genera have been recognized for the South and Central American
species. The wing venation used here is explained in Ross (1956,
figs. 365-367).
Key to Neotropical Genera of Glossosomatidae
1. Wings, especially hind, reduced in size and venation . . Merionoptila Schmid
Wings not reduced ..... SE NCHO fo A Bie Ue mee
2. All four branches of M present in Porewine | . . . Culoptila Mosely
Only two or three branches of M present in forewing ......... 3
3. \) Two branches‘of.. Min forewing) <0. 2h: eons ee. Go SO ee
Mireeibranchesiofvisinwtorewangs cas. 1) ee eee se) 6) ees See
4. Cu, branched apically in hindwing. .... . . . . Mortoniella Ulmer
Cu; unbranched in hindwing . . . em
5. Branching of M in forewing at pbaath same fleck as ranching of R445.
Scotiotrichia Mosely
M branching much nearer wing margin than Ry; . . . Mexitrichia Mosely
6. Rg; branching nearer forewing margin than Roi; . . . . . Canoptila Mosely
Rapand R5.3 branching at nearly ‘same level-;*. = .2..*'5. 63 eee a
7.. Cuyin hindwing branching apically...) a | Matoptila Mosely
Cuysimple:):.. ay .2erdevieoe 3 ser ark we as Syeacoat hb, spec 44:Pretopiila Banks
The larvae of the Protoptilinae are very poorly known at present.
Ross (1944) has described some particulars of the larva and pupa of
Protoptila lega Ross, and Flint (1962) the immatures of Matrioptila
geanae (Ross). The larvae of Mortonielia angulata, M. apiculata, M.
hodgesi, and Meritrichia aries are described herein. The larvae of
these species and of Protoptila alexanderi Ross differ strikingly from
those of the Glossosomatinae in the structure of the anal claw. The
larvae of the glossosomatines have one large ventral hook and 1 or 2
dorsal accessory teeth. In the protoptilines the dorsal accessory
teeth apparently are divided longitudinally, hence there are 2 or more
pairs of accessory teeth, one row on each side of the ventral central
hook. In addition to the differences in the anal claw, there are dif-
ferences between the subfamilies in the apical spurs of the tibiae,
the sclerotization of the thoracic notae, and the symmetry of the
mandibles.
The larvae appear to show generic differences, although criteria
for the separation of the species have yet to be found. The tarsal
and anal claws, apical spurs of the tibiae, and ninth tergite show some
of the most marked generic differences found so far.
The following key is presented only as a guide to the characters
found in the previously mentioned species and will probably require
much modification when more species and genera become known.
Key to the Genera of Protoptiline Larvae
1. Tarsal claws trifid; ninth tergite with 3 pairs of setae; 3 pairs of accessory
anal teeth: 24. 7 see Wi ve iy iia ad. @ MatrieptilatRess
NEOTROPICAL CADDIS FLIES I—FLINT 465
Tarsal claws with a single apical hook; ninth tergite with 2 pairs of long
SOLHLO Wee OE bm Mie tobi hk oo 8 eSB A rei gla Pc ontel stPhe clue. cya) a4 = 2
2. Anal claw with 2 pairs of well-developed accessory teeth; seta of tarsal claw
short and very broad (fig.7,e) -°. . . 1 ke we Mortoniella Ulmer
Anal claw with 3 or more pairs of accessory teeth; seta of tarsal claw nor-
EL ERS SO an ees I) OOD EL ORR hee vet 3
3. Ninth tergite concave apically; seta of tarsal claw arising from base of cuticu-
Pani rent? (TB d ost 2:04) Ensen roe) whehen Mexitrichia Mosely
Ninth tergite convex apically; seta of tarsal claw arising from posterior face
Be cUiicolr Ouigrowul (fig. 7,7)... . . . . «ss Protoptila Banks
Genus Mortoniella Ulmer
Mortoniella Ulmer, Notes Leyden Mus., vol. 28, pp. 95-97, 1906.
Ulmer described the genotype, bilineata, from Ecuador in 1906.
Subsequently he added a second species, albolineata, from Brazil.
The latter, however, was removed from the genus by Mosely in 1939
and placed tentatively in Antoptila. Martynov (1912) described
tranquilla from Peru; however, this description, based on a female
and unaccompanied by any illustration, is unrecognizable. Such is
the present state of our knowledge of the genus.
In the Cornell University collection is a male specimen from
Ecuador that perfectly matches Ulmer’s description and figures of
the genotype and permits the genitalia to be refigured. A second
species present in both the adult and the immature material from
Ecuador is congeneric on both genitalic and venational characters.
Two additional species of Mortoniella are described from pharate
adults that agree closely in genitalic structure. Mewsitrichia wygod-
zinskit Schmid from Argentina apparently also belongs in Mortoniella
on genitalic considerations, though Schmid’s comments on the venation
indicate that Cu, in the hindwing is unforked in this species. If
Schmid’s comments are accurate, Mewitrichia Mosely may have to
be synonymized with Mortoniella, because the genitalic differences
alone do not seem to justify a generic separation. Until more
species and their venation become known, however, I prefer to regard
the two genera as distinct.
Larvae of the three new species described here are known to me.
They are all similar, in fact, as yet I can find no specific differences,
and they are easily separated from the larvae of the other genera.
The structure of the tarsal and anal claws is characteristic. The
anal claw has only 2 pairs of accessory teeth, and the seta on the
tarsal claw has become greatly modified into a short, broad, thumblike
process. It must be admitted, however that Mezxitrichia aries, on
which the larval differences are based, may not be congeneric with
the genotype and when larvae of other species in this genus become
known these apparent differences may not prove to be valid.
466 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
Tentatively then, the genus Mortoniella differs from Mezitrichia in
that: Cu, in the hindwing is forked; in the male genitalia the tenth
tergum is elongate, the ventral surface of the aedeagus is often
sclerotized in the form of paired processes, and there are comparatively
simple basoventral processes; in the larvae there are only 2 pairs of
accessory teeth on the anal claw, and the seta on the tarsal claw is a
short, broad, thumblike process; and in the pupa the distal tooth on
the inner margin of the mandibles is long and arises far from the apex.
Mortoniella bilineata Ulmer
Ficure 5,a
Mortoniella bilineata Ulmer, Notes Leyden Mus., vol. 28, pp. 97-98, 1906.
A figure of the lateral aspect of the male genitalia is given from a
cleared specimen to supplement the figures of Ulmer. The only
difference to be noted is in the tenth tergum which Ulmer shows to
be slightly longer and slightly different in shape. Otherwise the
genitalia, venation, and coloration are an excellent match with the
original description.
Matertau.—Ecuador: Huigra, 4500 ft., June 13, 1914, 1 0,
Parish (CU).
Mortoniella apiculata, new species
Ficures 5,b-d; 6; 7,a-e
This species is closely related to angulata from which it differs
most conspicuously in the expanded apex of the dorsal process of the
aedeagus and in the greater proportionate length of the genital capsule.
Aputt.—Length of forewing 5 mm., length of body 4.5 mm.
Wings of specimens in alcohol brown, with a pale streak in membrane
at anastomosis; venation as shown (fig. 5,c). Male genitalia (fig. 5,6):
Ninth segment almost quadrate in lateral view, very oblique; tenth
segment with an elevated basomesal area, extending about half length
of lateral arms which are separated by a deep narrow mesal incision
(fig. 5,d); dorsal process of aedeagus angulate dorsally shortly before
apex, apex widened into a thin triangular structure; lateral processes
of aedeagus simple, slightly angulate rods extending about two-
thirds the length of dorsal process; ventrally to aedeagus a pair of
short rods basally and 2 paired processes near midlength, ventralmost
pair longer than dorsal pair; venter of aedeagus sclerotized in form
of paired spatulate process.
Holotype o, allotype, paratype 9: Ecuador: 1 mile east of Papal-
lacta, Jan. 29, 1958, R. W. Hodges (USNM Type 66018). Additional
paratypes: Ecuador: 6
moderately sclerotized, U-shaped pouch receiving and supporting ©
bases of harpes. Lateral aspect: narrowing dorsocaudad to point
of articulation with lateral arm of tegumen.
Tegumen with lateral arm quite broad, gradually narrowing
cephalad to point of articulation with vinculum, approximate ventral
half punctate, dorsal half glabrous, ventral margin heavily sclerotized
and sinuate, dorsal margin moderately sclerotized and sublinear;
dorsal area broad, glabrous, rather weakly sclerotized but not emar-
ginate mesad.
Harpe simple, slightly variable. Lateral aspect: width medium,
sublinear or with apical half curving somewhat ventrad; costa and
sacculus fused, comprising approximate basal half of harpe, broadest
in apical portion at point of attachment of arm of transtilla, gradually
narrowing basad to rather narrowly rounded basal extremity, dorsal
margin very heavily sclerotized; glabrous except for heavily punc-
tate and setose, considerably expanded, subdigitate, caudal portion
of sacculus; cucullus partially separated from costa and sacculus by
dorsal constriction and ventral area of reduced sclerotization, com-
prising apical half of harpe, commonly curving somewhat ventrad
and mesad, markedly constricted dorsad near base, major apical
portion rather broad, heavily punctate and setose ectad and entad
except for glabrous dorsal portion of basal two-thirds, apex broadly
and evenly to irregularly rounded.
Transtilla with arm approximately one-third as long as harpe,
slender, linear to sublinear, well sclerotized, glabrous, rather closely
subparallel with dorsal margin of costa, terminating considerably
distad of basal extremity of harpe, apex slightly expanded and rounded.
Uncus prominently bifid, somewhat variable but very distinctive.
Dorsal aspect: base large, heavily punctate and setose, partially set
off from tegumen by lateral areas of reduced sclerotization; lateral
margins well sclerotized, sublinear to sinuate, converging distad;
angle of bifurcation narrowly rounded to acute; furcae with basal
portions approximate, broad, strongly narrowing distad, heavily punc-
tate and setose. Lateral aspect: furcae abruptly directed ventrad to
ventrocephalad at angle of 90 degrees or more, broadly expanded,
laterally flattened, subtriangular, punctate and setose, closely sub-
parallel to weakly divergent ventrad, gradually narrowing to acute
apices.
Gnathos paired, curving ventrocaudad, lateral and apical portions
well sclerotized, large mesobasal portion membranous; arms flattened,
densely scobinate entad, apical portions overlapping, apices rather
narrowly rounded.
Anellus membranous, unarmed, juxta absent.
NORTH AMERICAN ACROLOPHIDAE—HASBROUCK 561
Aedeagus approximately three-fifths as long as harpe, considerably
flattened dorsoventrad, sublinear in dorsal and ventral aspects, ap-
proximate apical third curving somewhat ventrad in lateral aspect,
asymmetrical in all aspects, glabrous, base broadly and irregularly
expanded laterad and ventrad, basal half tubular, apical half opening
broadly and irregularly dextrad; apical portion consisting simply of
one or two, elongate, slender, moderately sclerotized, dorsosinistral
strips terminating acutely or subacutely.
Vesica large, membranous, armed with two prominent clusters of
cornuti: cornuti variously sized, mostly large, heavily sclerotized and
darkened, linear, with apices finely acute; basal cluster largest, located
dorsodextrad near base of vesica, containing 4-13 cornuti; apical
cluster smaller, located dextrad nearer apex of vesica, containing 2-8
cornuti. The number of cornuti in each cluster varies from specimen
to specimen and thus constitutes the chief intraspecific variation to be
found in the genitalia of this species (see the tabulation, below, for
examples of this variation).
In the @ genitalia of all the individuals of popeanellus that have
been examined, the presence of two distinct clusters of cornuti in the
vesica has proved to be a consistent and positive specific character.
However, these cornuti commonly vary in size and number between
the two clusters of any one individual as well as between the corre-
sponding clusters of any two individuals. Thus, they furnish a good
example of an intergrading, intraspecific variation in regard to their
sizes, numbers, and ratios between the basal and apical groups. This
variation is illustrated by a sampling of 18 specimens listed in the
following table.
Tape 1.—Intraspecific variation of cornuti in A. popeanellus
Cornutt
Specimen Basal Apical Total
1 13 5 18
2 10 6 16
3 9 2 11
f 8 a 15
5 8 6 14
6 8 6 14
7 8 3 11
8 7 8 15
9 7 7 14
10 fi 6 13
11 7 5 12
12 iG 5 12
13 7 2 9
14 6 5 11
15 6 3 9
16 5 4 9
17 4 5 9
18 4 4 8
562 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 114
The basal cluster may contain as few as 4 cornuti to as many as 13, the
apical cluster may contain from 2 to 8, and the total for both clusters
ranges from 8 to 18. The basal cluster usually contains the most
cornuti, but occasionally its number is equaled or even exceeded by the
number in the apical cluster. Individually, the cornuti vary in size
from minute to very large. ‘This variation is essentially nongeographic.
Typr.—Two type specimens in the Academy of Natural Sciences of
Philadelphia.
TypE LOCALITY.—‘‘Texas.”’
DistRIBuTION.—This species has a very wide range. It has been
reported from northern South America, Central America, and Mexico,
as well as from the United States where it is quite common except in
the northwestern states and the extreme north and west. It has been
recorded from Arizona, New Mexico, and Texas northward through
Oklahoma, Kansas, and Nebraska and eastward to Pennsylvania,
New Jersey, North Carolina, and Florida.
SPECIMENS EXAMINED.—606, from 106 localities. Large series con-
taining both sexes are to be found in the collections of the author,
the American Museum of Natural History, and the University of
Kansas. At least one quarter of all the specimens accumulated for
this study eventually proved to be A. popeanellus, and the number of
specimens available for study has made it advisable to reduce the
distributional data for this common species simply to the localities
and months of occurrence:
ALABAMA: Gurley (July); Huntsville (July); Valley Head (Aug.). ARIzoNa:
Chiricahua Mountains (July); Douglas (Aug.); San Bernardino Ranch, Cochise
Co. (Aug.); Tombstone (Aug.); Tueson (July). Arkansas: Camp Chaffee
(Aug.); Hope (Aug.). FiLorma: Key Largo (Aug.); Sanford (June); Winter
Park (May, June). Groraia: Bainbridge (Sept.); Catoosa Co. (Aug.); Clarke
Co. (Aug.); Screven Co. (July, an unusually small ?); Tallapoosa (July). IL-
LINOIS: Champaign-Urbana (June and July, both sexes very common at light);
Dubois (July); Havana (June); Homer Park, Homer (June, July); Lawrenceville
(Sept.); Murphysboro (Aug.); Oakwood (July); Palos Park (July); Parker (July).
Peoria (June, July); Putnam County (June, July, Aug.); Quincy (June).
InpIANA: Putnamville (July); Tremont (June). Iowa: Denison (June); Des
Moines (July, Aug.); Linn Co. (no date); Sioux City (June, July). Kansas:
Barton County (June); Bourbon Co. (no date); Caldwell (May, June, Aug.);
Clark Co. (May, June); Decatur Co. (July); Douglas Co. (June and July, 2 0d
with mites on abdomen); Gove Co. (no date); Linn Co. (no date); Manhattan
(June); Medora (June); Ness Co. (July); Onaga (July); Republic Co. (July);
Riley Co. (July); Scott City (June); Scott Co. (June, 1 o& with mite on eye);
Sheridan Co. (no date); Topeka (no date). Kenruckxy: Harrodsburg (Aug.).
Mississippi: Starkville (July); A. & M. College, State College (Aug.). Mu1ssourt:
Brentwood (June); Kirkwood (May, June, July); Mineola (July); Ranken (June) ;
St. Louis (July); Webster Groves (July); Willard (June). Nrsrasxka: Lincoln
(June, July, Aug.). New Jersey: Vineland (no date). New Mexico: Artesia
(July); Carlsbad Caves, Carlsbad (July); Chaves Co. (July); Deming (July);
Eddy Co. (July); Las Cruces (Aug.); Lordsburg (Aug.); Mesilla (July); State
NORTH AMERICAN ACROLOPHIDAE—HASBROUCK 563
College (July, Aug., Sept.). Norra Carouina: Hickory (July, Aug.); Kinston
_ (Aug.); Maxton (May); Raleigh (June, July, Aug., Sept.); Wilkesboro (Aug.).
_ Oxto: Granville (July). Oxkuanoma: Bartlesville (July); Clinton (June); Eagle-
town (June). PENNsyLVANIA: Finleyville (July); Manoa, Delaware Co. (July).
_ SourH Caronina: Myrtle Beach (June, July). TENNEssEE: Memphis (Aug.);
| Monteagle (July, Aug.); Nashville (July). Texas: Brownsville (June); Burnet
Co. (July); Christoval (June); College Station (Oct.); Comfort (May, Sept.);
Corpus Christi (Sept., Oct.); Davis Mountains (Aug.); Devils River, near Del
Rio (May); Eastland Co. (April); Kingsville (June); Marathon (July); New
Braunfels (May); Palo Duro State Park, Randall Co. (Aug.); Richmond (June);
San Angelo (June). Virainta: Salem (Aug.).
Remarks.—A. popeanellus, one of the first three acrolophids
described from the United States, is undoubtedly our commonest
and most widely distributed species and is one of the larger and more
robust acrolophids occurring in North America. It also exhibits
considerable variation in size and coloration. These four factors
explain why popeanellus is also the most heavily synonymized North
American acrolophid.
A. popeanellus is closely related to klotsi, the two comprising a smal]
species group in general related to those acrolophids having elongate
labial palpi, setose eyes, laminate antennae, bifid uncus, and paired
gnathos. However, popeanellus and klotsi may be distinguished
from each other and from all other acrolophids on the basis of their
aedeagi and unci. The latter organ, in the popeanellus-klotsi group, 1s
distinctive in having its furcae abruptly directed or curved very
strongly ventrad. In popeanellus, these furcae, easily observable in
dried and undissected specimens, are characteristically expanded
into subtriangular plates. The genital characters of popeanellus
are consistent throughout my very large series and they are quite
distinct from those of all the other species treated here.
I have not examined the type specimens of this species. Busck
(1903), in his report on Clemens’ types of Tineina in the Academy of
Natural Sciences of Philadelphia, noted:
Anaphora popeanella Clemens. Two types, both rubbed, one unspread and
without abdomen; the other spread, and lacking the head and left wings. Clemens’
No. 11; alar exp., 28 mm. Like the present conception of Anaphora popeanella,
as determined by Walsingham and synonymous with scardina Zeller, and with
agrotipennella Grote. Aspecimen compared with the type is in the U.S. National
Museum. Habitat: Eastern United States.
Darlington located one of these type specimens at Philadelphia, and
described it (in litt., 1946) as: ‘“‘popeanella Clem. Type. Right wings
only, poor condition, much rubbed, sex not determined.” The com-
bined information that I have been able to gather in regard to pope-
anellus leaves no doubt in my mind as to the proper identity and cor-
rect concept of this species. The specimens of popeanellus in the U.S.
National Museum agree with my previous concept of this moth. In
564 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
addition, at the Museum are several slide preparations of # genitalia,
labeled ‘‘popeanellus Clem.,’”’ agreeing with my figures of this species.
Clemens apparently named this moth after Captain Pope. Wals-
ingham (1887, pp. 155-156) designated this species as the type of
Clemens’ genus, Anaphora, and ten years later (1897) reported that
the larvae of popeanellus attack the roots of Trifolium pratense in
April and May, and that this insect also occurs in the West Indies
(Puerto Rico).
Grote described the &@ of Anaphora agrotipennella as a new species
in July 1872. The locality given was ‘‘Central Alabama” where Grote
reported the moth to be very common in June and July. An alar
expanse of 27 mm. was listed.
Following the description Grote stated:
I have only seen males of this species, in which the ornamentation of the fore
wings above recalls that of various species of Agrotis, such as A. jaculifera, etc.
I have tried to recognize in this species A. Popeanella, Clemens, from Texas, but
I have failed to reconcile his description with my specimens, which are not “‘luteous
or yellow along inner margin.”’ In A. agrotipennella, at the extremity of the
median ochrey shade subterminally, are a few black scale points. These can hardly
be the same as the row “of dark brown spots’”’ of Popeanella. Neither can I, from
the description, consider the differences of colour and ornamentation as produced
by any defect in the condition of Dr. Clemens’ specimens.
Grote, describing the 9 of Anaphora agrotipennella in August 1872,
further attempted to differentiate his species from Clemens’ popeanella.
When Walsingham (1887) placed it as a synonym of popeanella, he
stated:
An examination of my extensive series shows that both varieties belong to the
same species. The anal appendages do not differ, and intermediate variations
of colouring are noticeable. There can be no doubt that they differ only in the
extension of the pale colour of the fold in the direction of the dorsal margin.
Walsingham also added that Zeller had evidently regarded the two as
synonymous. Since 1887, agrotipennella has been generally treated
by writers as a synonym of popeanellus, although Barnes & McDun-
nough (1917) and McDunnough (1939) maintained it as a distinct
and valid species in their checklists.
I have not seen the type specimen, which is in the collection of the
Academy of Natural Sciences, but Darlington, in 1946, reported it to
be in perfect condition.
All the specimens labeled ‘“‘agrotipennellus’”’ that I have ever seen
proved, upon examination of their genitalia, to be examples of
popeanellus. Likewise, there is no indication in Grote’s descriptions
that his agrotipennella could not be popeanellus. In addition, there
is no North American species available, save popeanellus, to rep-
resent Grote’s descriptions of agrotipennella. It thus seems best to
consider agrotipennellus an old synonym of popeanellus (Clemens)
with Walsingham receiving the credit for sinking it.
NORTH AMERICAN ACROLOPHIDAE—HASBROUCK 565
Zeller described Anaphora scardina as a questionably new species
in 1873. At the conclusion of his discussion of Clemens’ genus
Anaphora, Zeller said of his own series of specimens: “Whether I
have Clemens’ species among my own before me, I cannot decide
with certainty from his descriptions; therefore I am giving the latter
under separate names.’’ He then proceeded to describe scardina
and bombycina at considerable length. That Zeller was uncertain
of the validity of his species, scardina, is evidenced by the fact that
immediately beneath its designation at the head of the original
description he inserted ‘‘? Popeanella Clemens’ rather than listing
it as “n. sp.” He gave the ‘“Vaterland” or locality as Texas and
Carolina, where popeanellus is quite prevalent, and also mentioned
that the largest example of all in his collection was from an unknown
locality, thus indicating the possibility of a mixture of species. Cham-
bers (1878), following Zeller’s note of caution, gave the listing ‘A.
scardina, Zell. =? popeanella, Clem.” Walsingham (1887), after
studying Zeller’s specimens, placed scardina in the synonymy of
popeanella, where it has since consistently appeared.
However, the type material for scardina, now in the British Muse-
um (Natural History), presents a considerably different situation.
One of the three photographs furnished by Tams is labeled “‘scardina
Zell., type &”” and easily but inconclusively passes for an adult popea-
nellus. Likewise, a second photograph labeled “seardina Zell.”’ rep-
resents a ventral view of the complete < genitalia of a specimen which
is clearly and unmistakably popeanellus. On the other hand, the third
photograph, also representing a set of genitalia and labeled “scardina
Zell., type o,” certainly does not represent popeanellus, nor does it
agree with any other known North American species of Acrolophus.
Hence, it is quite probable that Zeller’s scardina is actually a valid
species of Acrolophus, although not referable to popeanellus or any
other North American species. It also seems fairly certain that Zel-
ler’s original series of scardina contained at least two species of Acrolo-
phus: one, popeanellus, from Texas (also the locality of Clemens’ orig-
inal specimens of popeanellus) and Carolina; the other, apparently
designated by Zeller as the type of scardina, from parts unknown.
The latter may well be the large example of unknown origin mentioned
in Zeller’s original description.
Although scardina has long been considered an old synonym of
popeanellus, the type specimen indicates it does not belong to that
segment of the genus occurring north of Mexico. The name is spelled
scardinus to agree grammatically with its present genus, Acrolophus.
Walsingham (1887) described Anaphora morrisoni as a new species
on the basis of two oc" collected in Florida by Herbert Knowles
Morrison. The alar expanse given was 18 mm. He apparently
566 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 114
named the species after the collector, who died in 1885. Since 1887,
morrisont has consistently appeared in the literature as a distinct
species. Walsingham’s illustrations of the o genitalia of morrisoni,
showing the uncus and the cucullus of the harpe in both lateral and
dorsal aspects, are essentially the same as his corresponding figures of
the o& genitalia of popeanellus. On the preceding page of his revision,
Walsingham had separated the two species as follows:
1. Points of the uncus distinctly separate—popeanella.
2. Points of the uncus closely approximate—morrisont.
This difference is easily absorbed in the range of intraspecific variation
exhibited by popeanellus.
The type o& is at the British Museum (Natural History), from
which tams has sent two photographs, labeled ‘‘morrisoni Wals., type
3.’ These show, through a dorsal view of the pinned moth and a
ventral view of its entire genital capsule, that it is simply a small but
typical Floridian specimen of popeanellus. Thus, morrisont (Walsing-
ham) should be considered a new synonym of popeanellus (Clemens).
Beutenmiiller’s manuscript name,