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SMITHSONIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 


PROCEEDINGS 


OF THE 


UNITED STATES NATIONAL MUSEUM 


VOLUME 110 
NUMBERS 3416-3421 


UNITED STATES 
GOVERNMENT PRINTING OFFICE 
WASHINGTON : 1960 


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 
specialists 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, sep- 
arate 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. 

REMINGTON KELLOGG, 
Director, United States National Museum. 


CONTENTS 


Gurney, AsHuury B., and Brooks, A. R. Grasshoppers of 
the mexicanus group, genus Melanoplus (Orthoptera: 
Acrididae). Eighteen figures and five plates. No. 3416, 
published July 9, 1959 


New subspecies: Melanoplus bilituratus vulturnus. 


Hanbury, Cuaruss O., Jr. A revision of American bats of 
the genera Huderma and Plecotus. Twenty-seven figures. 
No. 3417, published September 3, 1959 Mest: 

Hersekovirz, Pum. Mammals of Northern Colombia, 
preliminary report No. 8: Arboreal rice rats, a systematic 
revision of the subgenus Oecomys, genus Oryzomys. 
Six figures and twelve plates. No. 3420, published 
February 24, 1960 


New subspecies: Oryzomys concolor bahiensis. 


Jounson, Puynurs T. The rodent-infesting Anoplura 
(sucking lice) of Thailand, with remarks on some related 
species. Seventy-five figures. No. 3421, published 
November 10, 1959 . pee 


New species: Enderleinellus corrugatus, Hoplopleura thurmanae, 
Neohaematopinus callosciuri, N. cognatus, N. capitaneus, N. 
kinabalensis, N. elbeli, Polyplax cannomydis. 


Kissinger, Davip G. A revision of the Apion subgenus 
Trichapion Wagner in the New World (Coleoptera: 
Curculionidae). Nineteen figures. No. 3418, published 
October 23, 1959 . 

New species: Apion (Trichapion) bettyae, A. (T.) acanonicum, 
A. (T.) adaetum, A. (T.) coryi, A. (T.) plectrocolum, A. (T.) 
ortotes, A. (T.) parcum, A. (T.) eaenum, A. (T.) enoplus, A. (T.) 
innocens, A. (T.) innocuum, A. (T.) latitator, A. (T.) mirandum, 
A. (T.) mirificum, A. (T.) evustum, A. (T.) nanulum, A. (T.) 
subsequens. 

SHort, J. R. T. A description and classification of the final 
instar Jarvae of the Ichneumonidae (Insecta, Hymenop- 
tera). Sixty-four figures. No. 3419, published October 
743) %el 215) a 


Pages 


1-93 


95-246 


513-568 


569-598 


247-389 


391-511 
Tir 


whi fy 
Afb otis! 
’ 


a” 


by) PRR 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 110 Washington : 1959 No. 3416 


GRASSHOPPERS OF THE MEXICANUS GROUP, 
GENUS MELANOPLUS (ORTHOPTERA: ACRIDIDAE) 


By Asauey B. Gurney! and A. R. Brooxs? 


INTRODUCTION 


Since the historic outbreaks of the Rocky Mountain grasshopper 
(Melanoplus spretus (Walsh)) in the 1870’s, the close relatives of that 
species, especially the one recently called MZ. mexicanus (Saussure), 
have been of great economic importance in the United States and 
Canada. The present study was undertaken to clarify the status of 
the various taxonomic entities in this complex, and we hope it will 
stimulate further clarifying studies. We have also attempted to 
summarize briefly the more important information on the biology of 
each species. 

For many years the status of spretus has been a puzzle to ento- 
mologists because no specimens have been collected since the early 
1900’s. Gradually the opinion developed that spretus disappeared 
because it was the migratory phase of a normally solitary grasshopper, 
and it was supposed that mezicanus was that species. Our study of 


1 Entomology Research Division, Agricultural Research Service, U. S. Department of Agriculture. 
2 Entomology Section, Canada Department of Agriculture Research Laboratory, Saskatoon, Saskatche- 
wan, Canada. 


1 
477119—59——_1 


2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


the aedeagus indicates that spretus is a distinct species. Any further 
evidence bearing on this opinion naturally is very desirable. 

It is chiefly on the basis of the differences in the aedeagus that 
mexicanus has been found to be distinct from the species commonly 
misidentified as M. mexicanus by American entomologists. The 
species most frequently misidentified as mezicanus is M. bilituratus 
(Walker), and the structure of the aedeagus further suggests that 
three subspecies merit recognition. 


Limits oF Group 


The mexicanus group consists of Melanoplus mezicanus and five 
closely related species. There are five subspecies in addition to the 
typical ones. Distribution is limited to North America, from southern 
Mexico to the Arctic. At this stage in our knowledge of the species 
groups of Melanoplus, the best that can be offered as a definition of 
the mexicanus group is a somewhat loose combination of the following 
characters: 


1. Pronotum with hind margin of the dorsal surface produced posteriorly, 
either broadly angulate or rounded (fig. 15). 

2. Tegmina usually covering most or all of abdomen, often extending beyond 
it, sometimes covering only about a third of the abdomen (pl. 5), but always 
elongate, never lobate. 

3. Male cercus (figs. 7, 8) varying from slender and somewhat rectangular to 
broadly and irregularly oval. 

4, Male subgenital plate tending to develop a dorsal lip at the apex, sometimes 
posteriorly produced, frequently with twin apical knobs (figs. 11, 16). 

5. Dorsal valve of aedeagus anterior to the apical portion of the main stem, 
usually well sclerotized, more or less concave laterally (figs. 1, 2, 3). Ventral 
valve of aedeagus relatively short, straplike, its apex usually visible in lateral 
view near the base of the dorsal valve, dorsad of the accessory lobe. Accessory 
lobe of aedeagus moderate to large, weakly sclerotized. 

The name of the mezicanus group is taken from the specific name 
that has been used most often in the literature in recent years, though, 
as now restricted, mexicanus is neither the most widely distributed 
nor the most important species. MM. bilituratus is the most widely 
distributed, and since the outbreak of spretus ended it has been the 
most important. Hebard (1935a, p. 57) recognized a borealis group; 
earlier (1928) he briefly discussed some members of the mexicanus 
group as he conceived it. It seems to us that the mezicanus group is 
sufficiently cohesive to be recognized as we are treating it, with 
borealis included, at least until the group relationships within Melano- 
plus are better understood. M. femur-rubrum (De Geer), genotype 
of Melanoplus, has not been included in this group, though the male 
cercus is very much like that of borealis borealis. The subgenital 
plate of femur-rubrum is not produced apically as in the members of 
this group. A distinctive character of femur-rubrum is a small 


GRASSHOPPERS—GURNEY AND BROOKS 3 


spinelike structure (x of fig. 1,2) borne near the base of the accessory 
lobe of the aedeagus; it does not seem homologous with the lateral 
projection from the dorsal valve in mezicanus. 


TABLE 1.—Checklist and index to specific and infraspecific names in the mexicanus 
group of Melanoplus 


(Page numbers refer to the present work; synonyms are shown in italics) 


Name Author and year Original genus Present status 
affinis (p. 14) Scudder, 1897 Melanoplus Syn. of bilituratus 
(Proposed bilituratus 
as distinct 
species.) 
affinis (p. 42) Scudder, 1897 Melanoplus Syn. of devastator 
(Proposed as 
form of M. 
devastator.) 
alaskanus (p. 45) Scudder, 1897 Melanoplus Syn. of bruneri 
angelicus (p. 14) Scudder, 1897 Melanoplus Syn. of bilituratus 
bilituratus 
arcticus (p. 14) Walker, 1870 Caloptenus Syn. of bilituratus 
bilituratus 
ater (p. 41) Scudder, 1897 Melanoplus Syn. of devastator 
atlanis (p. 14) Riley, 1875 Caloptenus Syn. of bilituratus 
bilituratus 
bilituratus (p. 12) Walker, 1870 Caloptenus Valid species 
borealis (p. 67) Fieber, 1853 Caloptenus Valid species 
bruneri (p. 45) Scudder, 1897 Melanoplus Valid species 
caeruleipes (p. 52) Cockerell, 1889 Caloptenus Syn. of spretus 


coeruleipes (p. 14) 


(Proposed as 


var. of spretus.) 


Cockerell, 1888 
(Proposed as 


var. of atlanis.) 


Melanoplus 


Syn. of bilituratus bilitu- 
ratus 


cockerelli (p. 75) Scudder, 1900 Melanoplus Syn. of borealis stupe- 
factus 
consanguineus Scudder, 1897 Melanoplus Syn. of devastator 
(p. 41) 
conspicuus (p. 42) Scudder, 1897 Melanoplus Syn. of devastator 
(Proposed as 
form of 
devastator.) 
defectus (p. 38) Scudder, 1897 Melanoplus Valid subspecies of 
bilituratus 
devastator (p. 41) Scudder, 1878 Melanoplus Valid species 
diminutus (p. 41) Scudder, 1897 Melanoplus Syn. of devastator 
dimidipennis Bruner, 1904 Melanoplus Syn. of bilituratus 
(p. 15) 
excelsus (p. 45) Scudder, 1897 Melanoplus Syn. of bruneri 
extremus (p. 67) Walker, 1870 Caloptenus Syn. of borealis borealis 
intermedius (p. 14) Scudder, 1897 Melanoplus Syn. of bilituratus bilitu- 
ratus 
junius (p. 68) Dodge, 1876 Pezotettix Syn. of borealis borealis 
latifercula (p.75) Caudell, 1903 Melanoplus Syn. of borealis stupe- 
factus 
mexicanus (p. 8) Saussure, 1861  Pezotettix Valid species 


4. PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


TaBLE 1—Checklist and index to specific and infraspecific names in the mexicanus 
group of Melanoplus—Continued 


Name Author and year Original genus Present status 
monticola (p. 68) Scudder, 1897 Melanoplus Syn. of borealis borealis 
obscurus (p. 42) Scudder, 1897 Melanoplus Syn. of devastator 

(Proposed as 
form of 
devastator.) 
palaceus (p. 78) Fulton, 1930 Melanoplus Valid subspecies of bore- 
alis 
parvus (p. 68) Provancher, Caloptenus Syn. of borealis borealis 
1876 
sapellanus (p. 75) Scudder, 1900 Melanoplus Syn. of borealis stupe- 
factus 
scandens (p. 68) Scudder, 1897 Melanoplus Syn. of borealis borealis 
scriptus (p. 14) Walker, 1870 Caloptenus Syn. of bilituratus 
bilituratus 
selectus (p. 8) Walker, 1870 Caloptenus Syn. of mexicanus 
septenirionalis Saussure, 1861 Pezotettix Syn. of borealis borealis 
(p. 67) 
sterranus (p. 15) Scudder, 1897 Melanoplus Syn. of bilituratus 
bilituratus 
spretis (p. 55) Thomas, 1865 Acridium Unrecognizable; see p. 55 
spretus (p. 52) Walsh, 1866 Caloptenus Valid species 
stupefactus (p. 75) Scudder, 1876 Pezotettix Valid subspecies of bore- 
alis 
typicalis (p. 42) Scudder, 1897 Melanoplus Syn. of devastator 
(Proposed as 
form of 
devastator.) 
uniformis (p. 41) Scudder, 1897 Melanoplus Syn. of devastator 
utahensis (p. 81) Scudder, 1897 Melanoplus Valid subspecies of bore- 
alis 
virgatus (p. 42) Scudder, 1897 Melanoplus Syn. of devastator 


vulturnus (p. 25) 


New subspecies of bili- 
turatus 


The species here treated in the mezxicanus group were distributed by 
Scudder (1897b) in five different “series” of Melanoplus, and M. 


borealis stupefactus (Scudder) was in the genus Podisma. 


Those 


“series” of Scudder are not comparable to the groups of Hebard and 
other modern workers. 


ANATOMY OF THE Mare GENITALIA 


Since the appearance of Hubbell’s (1932) important study, the 
concealed male genitalia of Acrididae have been utilized for distin- 
guishing species and subspecies to a considerable extent. Roberts 
(1941) coordinated the terms for the structures useful to taxonomists 
with those studied by morphologists, and proposed several new 
single-word terms. Dirsh (1956) carried the study further, comparing 
nearly 800 genera and modifying previous terminology by several 
changes and a number of new terms. 


GRASSHOPPERS—GURNEY AND BROOKS 5 


The male genital mass or assemblage, preferably known as the 
phallic complex, including the aedeagus and its supporting apodemes, 
as well as the epiphallus—located anterior to the aedeagus (fig. 2,b)— 
is situated at the end of the abdomen beneath the supra-anal plate 
and the pallium (membranous covering usually dorsad of the sub- 
genital plate). Taxonomically, the aedeagus and epiphallus are the 
most important organs of the phallic complex, and the critical parts 
of each are illustrated by M. femur-rubrum (fig. 1,h,%). 


MetTHODS AND TECHNICS 


A brief outline of the technic employed in examining the male 
phallic complex may be helpful. 

With freshly collected specimens it is a simple matter to slip back 
the pallium with a dissecting needle, thus exposing the aedeagus. 
The attaching muscles are easily disrupted sufficiently to permit the 
phallic complex to be lifted and pulled posteriorly. It is then exposed, 
attached to the subgenital plate, and is readily visible in the pinned 
specimen. However, for the most detailed study, removal of the 
phallic complex and its examination in alcohol are recommended. 
It may be preserved in a micro-vial, partly filled with glycerine, set at 
an angle beneath the grasshopper, with the pin passing through the 
cork of the micro-vial. The dry preparations still attached to 
specimens are adequate for sorting and for moderately critical study. 
In the case of dry grasshoppers, sufficient relaxation to permit expo- 
sure or removal of the phallic complex may be effected either by 
placing the specimens in a moist relaxing chamber for one or two days, 
or by dipping the end of the abdomen for a minute or two in water 
that is coming to a boil. Although very quick, the latter method 
frequently destroys the natural colors of the hind legs and results in 
crumpled tegminal apices; so the relaxing chamber is recommended if 
the future appearance of specimens is important. 

Because of the difficulty of identifying females to species in some 
cases, our maps and distribution records, except as noted, are based on 
males. ‘This practice has eliminated many uncertainties which other- 
wise would have entered into the distribution plotting for subspecies 
and for species of similar appearance which occupy the same areas. 
Lists of localities from which specimens have been examined, or full 
data on specimens, have been given only where they appear to be of 
considerable value. Localities are listed roughly in the order that they 
appear on the map, taken in rows from north to south, beginning in 
the east. For two or more localities very close together, only one spot 
is shown on the map. ‘The intensive field work of the junior author in 
Manitoba, Saskatchewan, and Alberta would, for some species, have 
permitted the inclusion of an almost solid mass of dots on the map, 


6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


so in those cases only limital and representative localities are shown. 

Tegmen (front wing or elytron) length is subject to inconsistencies 
unless a standard measurement is taken. Thus, it is measured “from 
the distal extremity of the tubercle formed by the junction of the 
subcostal and radial veins” (Proc. 4th Internat. Locust Conference, 
Cairo, p. 97, 1937) (also see Dirsh, 1953, for a treatment of standard 
measurements). We have tried to select specimens with tegminal 
apices intact, thus avoiding difficulties encountered when the tip of 
the tegmen is frayed (see Ramchandra Rao, Bull. Ent. Res., vol. 33, 
pp. 247-249, 1942). 


ACKNOWLEDGMENTS 


Most of the type specimens are located in the Museum of Compara- 
tive Zoology (MCZ), the Academy of Natural Sciences of Philadelphia 
(ANSP), and in the U. S. National Museum (USNM). We are much 
indebted to J. A. G. Rehn (of ANSP) and P. J. Darlington (of MCZ) 
for the privilege of studying the historical material in their charge, 
as well as for loans of important series from those collections. The 
courtesies extended by Mr. Rehn and by his colleague, Harold J. 
Grant, Jr., on several visits to Philadelphia by the senior author, are 
much appreciated. It has been especially helpful to examine many 
of the specimens preserved there which were studied by the late 
Morgan Hebard. To the following curators we would acknowledge 
deep appreciation for the loan of critical types or information about 
them: Max Beier (Naturhistorisches Museum, Vienna); N. M. 
Comeau (Musée de la Province, Quebec); Charles Ferriere (Museum 
d’Histoire Naturelle, Geneva) ; and David R. Ragge (British Museum). 
Other entomologists to whom we are very grateful for important loans 
are: T. H. Hubbell (University of Michigan); George P. Holland 
(Canadian National Collection, Ottawa); E. S. Thomas (Ohio State 
Museum); Henry Dietrich (Cornell University); S. W. Frost (Penn- 
sylvania State University); John R. Hilliard (University of Texas) ; 
F. W. Werner (University of Arizona); Edwin W. King (Clemson 
Agricultural College); R. E. Pfadt (University of Wyoming); George 
E. Wallace (Carnegie Museum); Paul D. Hurd (University of Cali- 
fornia); Neely Turner (Connecticut Agricultural Experiment Sta- 
tion); H. V. Weems (Florida Plant Board); B. B. Pepper (Rutgers 
University); R. D. Bird (Dominion Entomological Laboratory, 
Brandon, Manitoba); Frank T. Cowan, J. R. Parker and Fred Skoog 
(U. S. Department of Agriculture, Bozeman, Mont.); O. L. Barnes 
(U. S. Department of Agriculture, Tempe, Ariz.). 

Several other entomologists have cooperated in supplying informa- 
tion or other help, and acknowledgment of their assistance appears 
in our discussion of the various species. 


GRASSHOPPERS—GURNEY AND BROOKS 7 


Figures 1,a,d,e; 2 (all except f); 9,f,9,k-2; and 15 were made by 
Arthur D. Cushman, Entomology Research Division, U. 8. Depart- 
ment of Agriculture. All other drawings and the maps were made by 
the senior author. 


Systematic Treatment 


Key to species and subspecies of the mexicanus group of Melanoplus 
(Based on males) 


1. A distinct mesosternal swelling or ““hump”’ present (fig. 1,d,f,g). . .- . . 2 
Without a mesosternal “hump,” or with a weakly developed one (fig. 1,e). 7 
2. Dorsal valve of aedeagus (fig. 3,k) of a general circular shape, appearing thin 
and parchment-like, and with a triangular projection (x of fig. 3,k) on the 
dorsal rim; subgenital plate usually without twin apices well developed 
(fig. 11,a-1,2,4,5), occasionally with them distinct (fig. 11,a-3). (Distri- 
butions figs: 125,015), \spterisbnedens ie esterase . . bruneri Scudder 
Dorsal valve variously formed (fig. 3,a—-h) but not wath a generally circular 
shape as in fig. 3,k; subgenital plate normally with well-developed twin 
apices (fig. 9,f,h-2), best seen in posterior view (fig. 10,uw-z). . .... 3 
3. Dorsal valve of aedeagus strongly recurved laterally and posteriorly (fig. 3,a), 
the hornlike structure heavily sclerotized (pl. 2,p). (Distribution: fig. 14). 
spretus (Walsh) 
Dorsal valve (fig. 3,b-f) variously shaped, but not as in opposite category. 4 
4. Dorsal valve of aedeagus with a lateral projection (p) from its ventrobasal 
region (fig. 3,b-1, lateral view; fig. 3,b-2, lateroventral view). (Distribution: 
figs. 4, 6) . SpE SIe Certo tein tin ute. cis Nenie me mexicanus (Saussure) 
Dorsal valve without a Tavera projection from its ventrobasal region. . 5 
5. Dorsal valve of aedeagus (fig. 3,c) typically produced into a blunt hook or 
knob at lateroanterior angle (y), especially by a carina (x) which separates 
main lateral surface from a concave, heavily sclerotized ventroanterior 
marginal area (z). (Distribution: fig. 6). 

bilituratus vulturnus, new subspecies 
Dorsal valve (fig. 3,d-f) without a hook or knob at lateroanterior angle, and 
without a carina on main lateral surface, though the ventroanterior margin 
usually recurves as a lip (z of fig. 3,d) . THREE EAR toed tte CS OE + oti 6 
6. Dorsal valve of aedeagus appearing inflated, ‘igpiealln ith a concave anterior 
margin (x of fig. 3,d) and with a sinuate margin (z) of the recurved anterior 

flap (y), the latter wide. (Distribution: fig. 6). 
bilituratus defectus Scudder 
Dorsal valve with little appearance of inflation, without a concave anterior 
margin, and with a relatively straight margin of the recurved anterior flap, 
the latter of variable width, often narrow (fig. 3,e). (Distribution: figs. 
bays: (a) att Ree Caen Ces eae RERUN [rae SR bilituratus bilituratus (Walker) 
7. Cercus (fig.7,h) narrow; subgenital plate usually with twin apices (fig. 9,4-2), 
though variable and occasionally poorly developed (fig. 10,7); aede- 
agus with apex produced dorsally beyond the base of dorsal valve (figs. 
2,k; 3,2); dorsal valve extending primarily dorsoanteriorly. (Distribution: 
GG TA) po rsd as, (gnats fb bordel yeudartectay ony: «Xb. set devastater'Scudder 


8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Cercus (figs. 7,7; 8,a—e) usually less slender than above; subgenital plate most 
often without twin apices (fig. 11, b-f); apex of aedeagus scarcely extending 
dorsally beyond base of dorsal valve (fig. 3,7); dorsal valve extending 
primarily anteriorly, with a lateral twist. (Distribution: figs. 17,18). . 8 

8. Tegmina covering about one-half of abdomen (pl. 5,8,F). (Occurs at very 
high altitudes; distribution: fig. 18). . . borealis stupefactus (Scudder) 

Tegmina covering more than one-half & abdomen. ... «ae? Sale 

9. Subgenital plate with scarcely any or comparatively little oral or posterior 
development of the apex (fig. 16,d). (Distribution: figs. 17, 18). 

borealis borealis (Fieber) 

Subgenital plate with considerable dorsal or posterior development of the 


apex (fig. 16,e-g) . . . 3 2 LO 
10. Subgenital plate (fig. 16,f; ah 4 nie nee feenanienonels protaced poste- 
riorly. (Distribution: fig. 18) . . . . borealis utahensis Scudder 


Subgenital plate (fig. 16,e) erodnced! Honeeiie more than posteriorly. (Ab- 
domen often conspicuously recurved as in pl. 4,p.) (Distribution: fig. 18.) 
borealis palaceus Fulton 


Melaneplus mexicanus (Saussure) 
Ficursxs 2,c; 3,b; 4; 6; 7,a; 10,f,q; PuaTe 1, 


Pezotetitiz mexicana Saussure, Rev. Mag. Zool., ser. 2, vol. 13, p. 160, 1861 (male, 
female, Temperate Mexico). Lectotype, here designated: Male labeled 
“Cordova, t. c.’’? (printed), a separate green label ‘‘Melanoplus mexicanus 
Sauss.”’ (apparently in Saussure’s longhand). (Museum d’Histoire Naturelle, 
Geneva, Switzerland.) 

Caloptenus selectus Walker, Catalogue of the specimens of Dermaptera Saltatoria 
in the collection of the British Museum, pt. 4, p. 682, 1879 (female, Oajaca 
(sic), Mexico). Lectotype, one of two females, selected by Uvarov (1925, p. 
298). 

The condition of the lectotype of mezicanus is good, with all ap- 
pendages except antennae, right middle leg, and left hind tibia present. 
It is not an alcoholic specimen, and color is well preserved. Dark 
dorsal bands on hind femora are well developed, also the pale area of 
the lower third of the outer paginal area and the pink ventral surface. 
The hind tibia is deep pink. The aedeagus (dry on specimen) is typical 
of the species as shown by the series examined. ‘The cercus is some- 
what less robust than the one from Bledos, Mexico, illustrated (fig. 
7,a-1). The mesosternal hump is well developed. 

Charles Ferriere, curator of entomology at the Museum of Natural 
History in Geneva, has located 16 specimens (6 males, 10 females) of 
the original series of mezicanus. He kindly loaned 8 specimens, all 
bearing green name labels like that above. One female has the same 
locality label as the lectotype. Two females bear ‘Orizaba, Sumi- 
chrast,’”’ and one female ‘‘Orizaba, M. H. de Saussure.’’ Two males 
without locality labels and one female “‘Orizaba, reg. temp. Sumi- 
chrast”’ prove to be M. femur-rubrum (De Geer), showing that the 
original series was a mixed one. 


GRASSHOPPERS—GURNEY AND BROOKS 9 


It is probable that the green labels are not original ones, but later 
ones written by Saussure, since the generic name Pezotettiz is not 
used. The history of Saussure’s travels suggests that the specimens 
seen are part of the original series studied by Saussure. Burr’s 
obituary of Saussure (Ent. Record, vol. 17, pp. 167-170, 1905) records 
Saussure’s visit to the New World, and mentions an article published 
by him about the volcano of Orizaba in 1858. This volcano is adja- 
cent to Cérdoba (spelling now usually followed). Saussure (1870, 
p. 2) referred to Fr. Sumichrast (one of the collectors of mezicanus 
types) as his old aide and travel companion. The abbreviation “‘t. ¢.”’ 
is Spanish for “tierra caliente” (warm region), as indicated by Hebard 
(1932a, p. 281). However, the specimens may have been taken 
under temperate conditions, as Cérdoba, about 50 to 60 miles west 
of Veracruz, in the State of Veracruz, is at the base of Mount Orizaba. 
The wide variety of zonal conditions encountered when one descends 
from Mount Orizaba to Cérdoba is mentioned by Goldman (1951, 
p. 279). Also, Sumichrast, and apparently Saussure too, collected 
extensively at nearby Mirador (see Goldman, p. 276), and, depend- 
ing on the detail with which specimens were labeled, some of the 
original mezxicanus series may have originated there. Five males 
taken at La Cumbre, near Cérdoba, at an elevation of 6,000 feet, 
by Dr. H. R. Roberts in 1936, agree perfectly with the lectotype. 

Uvarov (1925, p. 298) placed selectus in synonymy. A male from 
Ocotlén, Oaxaca, which is near the type locality of selectus, has been 
examined by us and is considered typical of mezicanus. 

Two specimens, male and female (ANSP), from ‘littoral du Mexi- 
que,’ labeled as ‘‘Platyph. mezicanus Sauss.,’”’ apparently by Saussure, 
and as paratypes of Platyphymus mexicana Saussure by Hebard, are 
really Melanoplus mezxicanus. The incorrect name labels may have 
been prepared accidentally by Saussure; at any rate Platyphyma 
mexicanum, described by Brunner in 1861, is a very different insect 
with short elliptical tegmina. Hebard (1932a, p. 281) treated mezi- 
canum Brunner as a synonym of Pedies virescens Saussure. The two 
specimens noted probably are the ‘“‘paratypes’”’ mentioned under 
Melanoplus mexicanus by Hebard (1928, p. 279), since they are stand- 
ing under M. mezicanus in the Philadelphia collection. Walker’s 
Caloptenus mexicanus (1870, p. 682) was placed in the synonymy 
of Platyphyma mexicanum Brunner (there called Paradichroplus 
mexicanus) by Scudder (1897b, p. 19). 

The early literature dealing with Melanoplus mezicanus is scanty. 
Thomas (1873, p. 222) merely listed it (as Pezotettiz mexicana) from 
Mexico, and he paraphrased Saussure’s description. Bruner (1908, 
p. 301) stated that he saw specimens of M. devastator which were 
labeled, probably by Saussure, as Pezotettix mexicana with a query. 


10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Bruner further inferred that Saussure’s type series may have been 
mixed, and that another included species may have caused Brunner 
von Wattenwyl (1900, p. 257) to transfer mexicana to Dichroplus. 
Bruner felt that Brunner must have examined Saussure’s types. We 
have not seen the material which Brunner recorded from Colombia 
as Dichroplus mexicanus, but an error probably occurred. Scudder 
(1897b, p. 404) listed Pezotettizx mexicana as undetermined, but 
(p. 405) he suggested that it might be the same as Melanoplus atlanas. 
Kirby (1910, p. 513) correctly placed mezicana near atlanis Riley 
and spretus Thomas (true spretus, though incorrectly credited to 
Thomas), but elsewhere (p. 487) placed it in Trigonophymus. ‘The 
latter placement apparently refers only to the portion of Saussure’s 
type series which Brunner associated with Colombian specimens. 
Hebard (1917, p. 271) first brought the name mezicanus into general 
use when he applied it to the widespread grasshoppers which wenow 
recognize as a complex, and he placed atlanis of authors in synonymy, 
though retaining atlanis Riley for a population of eastern United 
States and vicinity. 

Descriptive Notes: A small to medium-sized member of the 
mexicanus group. Head with dorsal carinae of vertex more prom- 
inent and conspicuous, and closer together at anterior margins of 
eyes than in bilituratus, especially in male; antenna about one-third 
longer than head and pronotum combined (male), about equal to 
combined length (female) ; a small to medium-sized mesosternal hump 
present in male. 

MALE GENITALIA: Cercus with extremes as illustrated (fig. 7,a), 
usually approximating that of figure 7,a-2; furculae slender, taper- 
ing, usually divergent, about one-third as long as supra-anal plate; 
subgenital plate moderately upturned in lateral view, twin apices 
(posterior view) prominent, the intervening depression rather deep 
and well rounded; dorsal valve of aedeagus not extending dorsad 
nearly so far as apex of aedeagus, the anterior and dorsal margin 
covered by a light membrane, a pronounced lateral process (P of 
fig. 3,6) borne on ventral margin, near tip of ventral valve; accessory 
lobe large for group, larger than in bilituratus. 

FEMALE GENITALIA: As illustrated (fig. 10,f,¢); cercus triangular, 
apex well rounded. 

Cotoration: General color not distinctive, medium to dark for 
group, no rich yellow specimens seen. Hind femur with four dark 
transverse dorsal bands (including knee), inner paginal area irregu- 
larly tinged with pinkish, followed by solid pink below, outer paginal 
area dull brownish, paler along ventral margin and at base, ventral 
surface deep pink; hind tibia variable, either dark pink or ashy 
greenish gray. 


GRASSHOPPERS—GURNEY AND BROOKS 11 


VARIATION: Seven representative males, measured in millimeters, 
vary in pronotal length from 3.3 to 4.5 (average, 4.0), in length of hind 
femur from 9.5 to 13.0 (av. 11.3), and in length of tegmen from 13.2 
to 19.0 (av. 16.0). The smallest male is from Atoyac, Mexico, the 
largest ones from Mt. Alvarez, Mexico, and 18 miles north of Alpine, 
Tex. Three measured females vary in pronotal length from 4.3 to 
5.1 (av. 4.7), in length of hind femur from 12.4 to 13.5 (av. 12.8), 
and in length of tegmen from 17.0 to 21.0 (av. 18.7). The tegmina of 
most specimens exceed the hind femur by about the maximum width 
of hind femur, by somewhat more than that in a few specimens, while 
in one specimen (north of Tehuacdn) the tegmina reach only to the 
femoral apices. Most, but not all, specimens collected within the 
past 20 years have pink hind tibiae, but in most old specimens and a 
few recent ones the hind tibiae are ash or greenish gray. Apparently 
some variation occurs in nature, though killing agents and the condi- 
tions of preservation may be responsibile for some loss of color. 

The shape of the dorsal valve of aedeagus is variable in dry prepara- 
tions, due to shriveling. The membrane along the anterior margin 
then often becomes indistinct and the margin becomes irregular, 
sometimes indented. The lateral process is of uniform length, for the 
most part, but is below average length in the male from Alpine, Tex. 
The dorsal shoulder at the base of the apical ‘‘scoop”’ of the dorsal 
Ovipositor valve is somewhat variable, occasionally more angular 
than in fig. 10,q. 

Distripution: The map (fig. 4) shows the distribution of mexicanus, 
based on specimens examined. ‘Two lots of Texas specimens have 
been examined: 4 miles west of Ft. Stockton, Pecos Co., Aug. 8, 1955, 
J.R. Hilliard (27, 39); in Jeff Davis Co. 18 miles north of Alpine, June 
19,1939, F.B.Isely (1¢). Hebard (1917 ;1932a) listed other Mexican lo- 
calities, but only one Mexican State, Querétaro, additional to those on 
the map, is represented by those records. Itis probable that all or most 
localities represented by Bruner’s specimens (1908) were reported 
by Hebard because he acquired Bruner’s collection of North American 
Orthoptera. Hebard (1925b) did not report mezxicanus from Sinaloa, 
but collections from northwestern Mexico, in general, have not been 
extensive. More collecting is needed to determine the distribution 
of mexicanus in Texas, but the paucity of records suggests scarcity 
or localized occurrence. Tinkham (1948, p. 619) had no records 
other than to repeat the El Paso record of atlanis by Rehn and Hebard 
(1909). El Paso specimens have not been examined, but they probably 
represent WM. bilituratus defectus. 

Biotocy: No detailed information on the biology of mexicanus is 
available. The data on the specimens examined show occurrence in 
central Nuevo Leén at altitudes of both 2,200 and 7,000-8,000 feet, 


2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


in Guerrero at altitudes ranging from 5,000 to 8,000 feet, and in 
Michoacan from 6,000 to 8,000 feet. The series of about 35 Mexican 
specimens examined includes captures made in all months except 
January and April, and it is evident that there is more than one 
generation, perhaps continuous breeding, slowed down by unfavorable 
local conditions. 


Melanoptus bilituratus (Walker) 


This widespread, variable species is represented by most of the 
material which, in recent years, has been referred to Melanoplus 
mexicanus mexicanus. Exceptions are specimens now recognized 
as true MM. mezxicanus, and M. spretus. Specimens of M. 
bilituratus from the Far West occasionally have been confused with 
M. devastator, sometimes referred to as M. mezicanus devastator. As 
mapped by Newton and Gurney (1956-57) (as mezxicanus complez), 
bilituratus occurs almost throughout the United States, except for 
peninsular Florida. Our present maps (figs. 5, 6) are based on males 
the aedeagi of which have been examined to determine the subspecies 
concerned. 

While more detailed research may warrant the recognition of a larger 
number, the material now examined, which is representative of most 
areas, seems to justify only three subspecies. ‘There is some indication 
that ecological subspecies, recognized by color and size, do occur, 
especially in Canada and Alaska (see p. 20). Brooks (1958, p. 20) 
has recognized these differences by treating atlanis (Riley) as a valid 
subspecies; however, the senior author doubts the value of applying 
names to these questionable ecological subspecies at this stage in the 
study, and atlanis is regarded as a synonym. 

The reason that the three populations concerned are recognized 
as subspecies, and not species, is that in each case intergradation in 
structural characters of the aedeagus occurs. Except for material 
from ‘Texas, intergrading specimens are here discussed under the 
appropriate subspecies. Those from Texas are of sufficiently un- 
certain relationship that it seems best to discuss them in one place. 
Males from Gainesville and several other Texas localities as listed 
below are typical or essentially typical of bilituratus vulturnus. Some 
divergence, in that the carina of the dorsal valve of aedeagus is not 
well developed, is shown by some specimens from Austin and Wichita 
Falls. The male from Phantom Lake, Davis Mountains, appears 
typical of belituratus defectus, but one from nearby Marfa and those 
from Blanco County, Plainview, Lubbock, and Sweetwater are not 
typical bilituratus defectus and are best considered intermediates of 
uncertain relationship. The specimen from Kokernot Mountain, 
Alpine, Tex., is unusual in that the anterior margin of the dorsal 


GRASSHOPPERS—GURNEY AND BROOKS 13 


x anterior 
ancora, bridge y projection GpeKnof 
. 4 Delian, main stem 
1 N ! 
} 
__ lateral 
plate 
ventral 
valve --~ 
\ . 
posterior A accessory — 
projection lobe cad 


Ficure 1.—a, Sketch showing location (circle) of mesosternal hump in ventral view of 
thorax of certain species of Melanoplus. 6, Melanoplus bilituratus bilituratus, female, 
Medicine Lake, Mont., apical portion of abdomen (z, cercus). c, M. femur-rubrum, 
female, Poplar, Mont., same view as b. d-g, Males, showing lateral view of mesosternal 
area in lateral profile: d, M. bilituratus: 1, M. b. oulturnus, Marshall, Mo.; 2, M. 6. bili- 
turatus, Pullman, Wash. e¢, M. devastator: 1, 2, two specimens from Tulare, Calif. 
*, M. spretus, Minnesota, Oct. 6, 1876. g, M. bruneri: 1, East Spanish Peak, Colo. 
2, Fishers Peak, Colo.; 3, Mt. Moriah, Nev. h-i: M. femur-rubrum, male, Euclid; 
Minn.: h, dorsal view of epiphallus; 7, aedeagus: 1, lateral view; 2, dorsoposterior views 


valve is irregular (fig. 3,g); the specimen is not typical of any one of 
the subspecies, nor is it like the Marfa specimen. Additional material 
from western Texas is required to determine the relationship of the 
one or more populations present. 
Texas males of bilituratus examined are as follows: 
M. bilituratus vulturnus: Gainesville, Wichita Falls, Quanah, Dallas, College 
Station, Austin, Wimberley, 2 miles east of Cedar Creek, Bastrop Co. 
M. bilituratus defectus: Phantom Lake (Davis Mountains). 


Intermediates of uncertain relationship (M. bilituratus in a broad sense): Blanco 
Co., Plainview, Lubbock, Sweetwater, Marfa. 


14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


As mentioned under the respective subspecies, the occurrence of 
two annual generations has been reported for bilituratus defectus, 
and evidence suggesting two generations or a partial second genera- 
tion has been presented for bilituratus vulturnus and typical bilituratus. 
In some cases the evidence consists of a second peak of adult abun- 
dance during a year. In recent years several important discoveries have 
been made concerning diapause in the eggs of grasshoppers, as re- 
viewed by Lees (1955, pp. 7, 46, 68, 127). Further studies may 
disclose that the subspecies of bilituratus differ with respect to dia- 
pause. 

Entomologists frequently have had difficulty distinguishing be- 
tween the females of bilituratus (in the past usually called mezxicanus 
or atlanis) and femur-rubrum. With a little practice it usually is a 
simple matter to separate them by the cerci (fig. 1,6,c), which are 
much more slender and apically acute in femur-rubrum. 


Melanoplus bilituratus bilituratus (Walker) 
Figures 1,b,d-2; 2,e,f; 3,em; 5; 6; 7,f,9,l; 8,3,m; 9,9; 10,9,n; PuatTE 1,8,c. 


Caloptenus bilituratus Walker, Cat. Derm. Salt. B. M., pt. 4, p. 679, 1870 (male, 
female, Vancouver Island, Canada; not 2 females, see Uvarov, 1925, p. 298). 
Lectotype, designated by Uvarov (1925, p. 298): Male from Vancouver 
Island (BM). 

Caloptenus scriptus Walker, Cat. Derm. Salt. B. M., pt. 4, p. 680, 1870 (females, 
Vancouver Island, Canada). Lectotype, designated by Uvarov (1925, p. 
298): Female from Vancouver Island (BM). 

Caloptenus arcticus Walker, Cat. Derm. Salt. B. M., pt. 4, p. 681, 1870 (one female, 
“Arctic America. Presented by Dr. Rae’’). New synonymy. 

Caloptenus atlanis Riley, Seventh Ann. Rep. Nox. Benef. Ins. Missouri, p. 169, 
1875 (males, females, Boscawen, Merrimack Co., N. H.). New synonymy. 
Lectotype, here designated: Male labeled ‘‘N. H., type no. 1153, U.S. N. M., 
Type, Caloptenus Atlanis Riley” (USNM). 

Melanoplus atlanis, var. coeruleipes Cockerell, Entomologist, vol. 21, p. 301, 
1888 (near Templar Rock, Custer Co., Colo.). New synonymy. 

Melanoplus affinis Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 20, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 182, 171-172, pl. 12, fig. 2, 1897. 
(See Hebard, 1917, footnote, p. 251.) (Males, females, 8 localities in Utah, 
Wyoming, Washington, and British Columbia.) Newsynonymy. Lectotype, 
designated by Rehn and Hebard (1912, p. 79): Male from Salt Lake Valley, 
Utah (ANSP). 

Melanoplus angelicus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 22, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 202-203, pl. 13, fig. 10, 1897 
(2 males, Los Angeles, Calif.). New synonymy. Lectotype, designated by 
Rehn and Hebard (1912, p. 80): Male from Los Angeles, Calif. (USNM). 

Melanoplus intermedius Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 20, 
32, 1897; Proc. U. S. Nat. Mus., vol. 20, pp. 132, 172-174, pl. 12, figs. 3, 4, 
1897 (males, females, 5 localities in Colorado, Montana, Wyoming, Idaho, 
and Washington). New synonymy. Lectotype, designated by Rehn and 
Hebard (1912, p. 79): Male from Yellowstone, Mont. (ANSP). 


GRASSHOPPERS—GURNEY AND BROOKS 15 


Melanoplus sierranus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 21, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 193-194, pl. 13, fig. 1, 1897 
(males, females, Lake Tahoe, Placer Co., Calif.; Placer Co., Calif.; Truckee, 
Nevada Co., Calif.). Newsynonymy. Lectotype, designated by Rehn and 
Hebard (1912, p. 80): Male from Truckee, Nevada Co., Calif. (MCZ). 

Melanoplus dimidipennis Bruner, Colorado Agr. Exp. Sta. Bull. 94, pp. 66-67, 
1904 (one male, Fort Collins, Colo.). New synonymy. 

The unique type of dimidipennis and all lectotypes except those of 
bilituratus and scriptus have been examined. Uvarov (1925) synony- 
mized scriptus and also suggested (p. 299) that arcticus may represent 
M. femur-rubrum (De Geer), but Dr. David R. Ragge (in litt., Feb. 
22, 1957) has reported that the cercus of the unique female type of 
arcticus is much blunter than that of femur-rubrum, and he states 
definitely that the two names are not synonymous. He has found 
that except for pale color arcticus agrees very well with an Alaskan 
specimen of bilituratus which was sent for comparison; the wing 
lengths of the two specimens are almost identical. Since the Rae 
specimen described by Walker apparently came from the region of the 
Mackenzie and Slave Rivers (see Richardson, 1852, p. 476), and 
specimens of bilituratus from that region agree essentially with the 
Alaskan ones, we regard arcticus as a synonym. Hebard (1928, pp. 
279-280) placed affinis, atlanis, and intermedius as synonyms of 
M. mexicanus mexicanus; this was done without reference to the 
aedeagus. Hebard (1929, p. 391) placed dimidipennis as a synonym 
of brunerz; however, the aedeagus of the type shows that it is belituratus. 
The type of dimidipennis is small and unusually short-winged, with 
the external genitalia including features like both bruneri and 
bilturatus, but not fully typical of either. It probably is an abnormal 
individual, as Hebard suggested. Scudder (1879b, p. 179) placed 
coeruletpes as a synonym of atlanis. No type specimens of coeruleipes 
are known, and since Cockerell (1888) simply applied the name 
without a formal description to individuals of atlanis (as then so-called) 
with bluish hind tibiae, and only one nominal form of bilituratus occurs 
in the area concerned, coeruleipes is relegated to the synonymy. The 
spelling was given as “caeruleipes’’ by Cockerell (1889). 

DescrIPTIVE NotEs: A usually medium-sized, fully winged member 
of the mezicanus group. Head with dorsal carinae of vertex moder- 
ately prominent; mesosternal hump strongly developed in male, 
absent in female. 

MALE GENITALIA: Cercus variable (fig. 7,f), over most of eastern 
half of range about as in figure 7,f-1; in Utah, California, Idaho, 
Oregon, Washington, and coastal British Columbia averaging nar- 
rower (fig. 7,f-2-4); in Nevada often decidedly slender, especially 
from near Reno (fig. 7,f-5-6); furculae about one-third to nearly 
one-half as long as supra-anal plate, straight or divergent; subgenital 


16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


plate (fig. 10,w) with twin apices usually conspicuous, rarely poorly 
differentiated. Aedeagus with dorsal valve well sclerotized and with 
a narrow to rather wide marginal fold, anterior margin usually broadly 
rounded, rarely with traces of sinuation such as in bilituratus defectus; 
accessory lobe of medium size for group; epiphallus with lophus 
rarely showing anterior angulation in lateral view, and then poorly 
developed, usually erect, in dorsal view the apex of pean broad and 
semitruncate (figs. 2,e,f; 3,e; 8,7,m). 

FEMALE GENITALIA: Geneon (fig. 10,9) cae relatively 
blunt; dorsal valve of ovipositor with “scoop” (fig. 10,n) moderately 
rounded, the shoulder obtusely angular. 

VARIATION: The size of 11 representative males, measured in 
millimeters, varies in pronotal length from 3.4 to 5.1 (av. 4.3), in 
length of hind femur from 9.8 to 13.7 (av. 11.6), and in length of 
tegmen from 15.0 to 25.0 (av. 21.1). Eight measured females vary 
in pronotal length from 3.7 to 5.5 (av. 4.7), in length of hind femur 
from 12.0 to 14.0 (av. 12.8), and in length of tegmen from 17.0 to 
21.0 (av. 18.0). Eastern specimens average smaller than those from 
the West. 

Tegminal length is decidedly variable. Unusually long-winged 
specimens are occasionally encountered. Among them are a female 
from Doniphan Co., Kans., Sept. 9, 1940, with tegmina reaching 
9 mm. posterior to the hind femora, and two females taken in Fergus 
Co., Mont., Aug. 16, 1940, with comparable measurements of 6.5 
and 7.5mm. A series of six females and two males was found dead 
in a recently baited field at Fort Benton, Mont., July 14, 1940, by 
Parker and Butcher. The average length of the tegmina extending 
beyond the hind femora in the females of that series is 5.1 mm., with 


EXPLANATION OF FIGURE 2 


Lateral views of aedeagus, and (b only) dorsal view of aedeagus and epiphallus in natural 
position within phallic complex. 


Melanoplus bruneri, Ft. McLeod, Alberta, Canada. 

mexicanus, Mt. Alvarez, Mexico. 

mexicanus, Guerrero, Mexico. 

. bilituratus bilituratus, Placer Co., Calif. 

. bilituratus bilituratus, Victoria, Vancouver Island, British Columbia, Canada. 
. bilituratus, uncertain subspecific position, Guymon, Okla. 

. bilituratus, uncertain subspecific position, 18 miles north of Alpine, Tex. 

. bilituratus defectus, Mesilla Park, N. Mex. 

. bilituratus oulturnus, Stone Mountain, Ga. 

. devastator, lectotype. 

M. bilituratus defectus, Casa Grande, Ariz. 

Melanoplus sp., probably abnormal bilituratus oulturnus, Muscatine Co., Iowa. 


M. spretus, Nebraska. 


a, 


ong ee 
RESRSSRERS 


GRASSHOPPERS—-GURNEY AND BROOKS 17 


Ficure 2.—Explanation on facing page. 


477119—59—_2 


18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


a maximum of 8.5; for the males the average is 8.3, the maximum 11.0 
mm. The aedeagus of the shorter-winged of the two males is normal 
for bilituratus bilituratus, and the cercus is comparatively narrow. 
The other male, with a wider cercus (fig. 7,g), has the dorsal valve of 
the aedeagus rather irregular along the lateroanterior margin, but 
without any indication of spretus. A male with a similarly abnormal 
dorsal valve was taken in Flathead Co., Mont., July 8, 1917, and its 
tegmina exceed the hind femora by 5.5mm. Rehn (1952) noted the 
considerable variation shown by Alaskan specimens. The folded 
tegmina of some individuals examined by him scarcely surpassed the 
apices of hind femora, while those of other individuals exceeded the 
femora by a distance about equal to pronotal length. 

Rehn and Hebard (1906), under the names intermedius, atlanis, and 
bilituratus, discussed the variation observed in the cerci and other 
organs of specimens from various Western States. When recording 
this grasshopper from Oregon, Fulton (1930) used the name MM. 
mexicanus bilituratus, and he observed that cerci of males vary from a 
little less than twice as long as middle width to nearly three times as 
long as broad. He noted that a series from Malin had consistently 
narrower cerci than those from other Oregon localities. 

It is evident from rearing experiments under different conditions 
of temperature, relative humidity, and food (Parker, 1930) for 
bilituratus bilituratus (called mexicanus mezxicanus), that size, color, 
spotting of tegmen, length of tegmen, and color of hind tibia vary in 


EXPLANATION OF FicureE 3 
Except as noted, lateral views of aedeagus drawn from preparations in glycerine. 


a: Melanoplus spretus: 1, Arapahoe Peak, Colo.; 2, 3, neotype, posterior and lateral views 
of aedeagus, dry preparation. 

b: M. mexicanus: 1, 18 miles north of Alpine, Tex.; 2, Durango, Mex., lateroventral view 
of aedeagus (p, lateral projection). 

c: M. bilituratus oulturnus: 1, holotype, dry; 2, Priest Bridge, Md. (m, membranous fold; 
x, carina; y, hooklike apex; z, concave area). 

d: M. bilituratus defectus, Tempe, Ariz., reared (x, anterior margin; y, flaplike area of 
dorsal valve; z, sinuate margin of flap). 

e: M. bilituratus bilituratus, Wasaga Beach, Ontario, Canada. 

Ff: Intermediate between M. bilituratus bilituratus and M. bilituratus defectus, Baker, Nev., 
dorsal valve only. 

g: M. bilituratus, uncertain subspecific position, Kokernot Mt., Alpine, Tex., dorsal valve 

only. 

M. bilituratus defectus, Chandler, Ariz. 

M. devastator, Roseville, Calif. 

M. borealis borealis, Nachvak Bay, Labrador, Canada. 

M. bruneri, Mt. Moriah, Nev. (x, triangular projection of dorsal rim of dorsal valve). 

Same specimen as &, right half of aedeagus in dorsal view. 

M. bilituratus bilituratus, Vancouver Island, British Columbia, Canada, same view as J. 


Se. 


GRASSHOPPERS—GURNEY AND BROOKS 


Ficure 3.—Explanation on facing page. 


19 


20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


response to ecological conditions. For these reasons, and perhaps 
because of genetic factors as well, bilituratus bilituratus shows much 
diversity in superficial appearance. This is especially evident in 
regions where a number of comparatively distinct, large, and relatively 
stable ecological associations occur; such large and diverse areas occur 
across southern Canada and in the mountains. The junior author 
has attempted to segregate Canadian specimens on the basis of the 
above and other external characters. In the southern coastal areas of 
British Columbia and from Vancouver Island there are small to 
medium-sized, moderately dark, often more hairy specimens, the 
tegmina frequently barely covering the abdomen. In most of the 
interior of southern British Columbia (range lands) specimens of 
moderate size, more brownish, and with red hind tibiae occur. A 
single series taken at Westbank, British Columbia, in 1923 includes 
rather large, predominantly yellowish specimens with the tegmina 
only about long enough to cover the abdomen. In northern British 
Columbia, Northwest Territories, and in Alaska the type usually 
developed is very large, blackish, with reddish hind tibiae, and very 
long tegmina; obviously of the same type but somewhat smaller are 
specimens from the foothills of Alberta and from the forested regions 
of Alberta, Saskatchewan, and Manitoba. In the grasslands of 
southern Alberta, Saskatchewan, and Manitoba the predominant 
type is of moderate size and grayish, with the tegmina extending 
scarcely beyond the abdomen, and usually with bluish or buff hind 
tibiae; in the eastern, more humid part of this area (parklands) the 
form is similar but more brownish and more often with reddish hind 
tibiae, very like specimens from interior British Columbia. In 
Ontario and Quebec specimens are nearly intermediate between those 
from the forested area of the prairie provinces and from the eastern 
grasslands, usually have red hind tibiae, and are comparatively 
stable in appearance. In dry years, during periods of grasshopper 
abundance, and when there is a trend toward swarming, the grassland 
types of central Canada develop longer tegmina, are of increased size 
and paler color, and usually have bluish hind tibiae. By and large, 
in Canada, observations suggest that most if not all major ecological 
areas have a smaller, grayer, shorter-winged type during “poor 
grasshopper years’ and a larger, more yellow, and longer-winged 
type during ‘“‘good grasshopper years.” ‘These differences seem more 
noticeable in areas which generally are dry than in those where 
moisture is abundant. 

Genitalic characters of the populations in different parts of Canada 
do not differ significantly or consistently enough to warrant, at this 
stage of our knowledge, the recognition of separate subspecies. It 
would appear that the superficial appearance of the populations in 


GRASSHOPPERS—GURNEY AND BROOKS vA 


different areas is largely the result of surrounding conditions. Even- 
tually, studies may show that the recognition of additional named 
segregates is desirable. 

The dorsal valve of the aedeagus may have a narrow anterior flap 
and a rather shallow lateral concavity (figs. 2,f; 3,e), or approach the 
more swollen type found in intergrades with defectus (fig. 3,f), with 
wide anterior margin and deep lateral concavity. Practice is re- 
quired to become familiar with the variation, which apparently is 
partly individual and partly geographic. A very small male with 
abortive wings, collected at Clark,;Wyo., Aug. 7, 1937, has the dorsal 
valve rather stubby, without the usual details, and apparently im- 
perfect because of the failure of the individual to develop normally. 


Ficure 4.—Distribution of male specimens of Melanoplus mexicanus examined. 


The distribution of males of bilituratus bilituratus examined is shown 
in figures 5 and 6. 

BIoLoGY AND ECONOMIC IMPORTANCE: The great importance of M. 
bilituratus bilituratus, which represents the northern subspecies of what 
usually has been called M. mezicanus or M. atlanis, is well known. 
Important bulletins dealing with this grasshopper are by Herrick and 
Hadley (1916) and by Shotwell (1930, 1941). Although crop and 
range damage occur most often west of the Mississippi River, injury 
on a smaller scale occurs in the Eastern States. Washburn (1953) 
described a severe outbreak on two farms near Palmer, Alaska, in late 
July 1951, in which populations up to 300 per square yard were 


pap PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


recorded, and vegetables, oats and raspberries were seriously injured. 
Early July in that locality was the warmest in 35 years, and that 
condition may have brought about the early maturity and great 
abundance of this grasshopper. 

M. bilituratus bilituratus is decidedly migratory at times, and in 
areas of abundance is usually much more inclined to loose group 
movements than are most grasshopper species. Parker, Newton, 
and Shotwell (1955) have described the spectacular flights which in 
1938-1940 occurred in North Dakota, South Dakota, Montana, and 
portions of nearby States and Canada. These flights occurred during 
an outbreak cycle, and a general population increase had preceded the 
flights. Drake and Tauber (1946) discussed flights which entered 
Towa from the west in 1941 and earlier years. 

Corkins (1922) reported on extensive flights in 1920, mainly in 
Bottineau and Renville Counties, N. Dak., and he noted their re- 
semblance to the historic flights of spretus. By the use of an airplane 
one heavy swarm was examined and found to be most dense at an 
altitude of 500-800 feet, with a few individuals at 1,650 feet. 

Nymphs as well as adults occasionally migrate (Parker, et al., 
1955, p. 16; Shotwell, 1930, p. 24). 

Early papers (Herrick and Hadley, 1916; Shotwell, 1930) gave the 
impression that bilituratus bilituratus is almost omnivorous in its food 
habits. In recent years, however, entomologists have become keenly 
aware that many grasshoppers are far from being the general feeders 
which they were once thought to be, and several investigations 


Figure 5.—Distribution of male specimens of Melanoplus bilituratus bilituratus examined 
from Alaska and Canada. 


GRASSHOPPERS—GURNEY AND BROOKS 23 


suggest that bilituratus bilituratus has definite responses and prefer- 
ences concerning food plants. In cage studies conducted in eastern 
Wyoming, Pfadt (1949) found that adults preferred dandelion and 
wheat to all other plants which he tested, that tansymustard and 
dandelion gave the most nymphal growth, and that dandelion, wheat, 
and tansymustard were most favorable to egg production. In 
Manitoba, Smith, Hanford, and Chefurka (1952) fed certain plants 
through three successive generations (in cages) and recorded survival, 
rate of development, and the number of eggs laid. They found that 
wheat, barley, and dandelion were the most favorable food plants, 
and that Russian thistle and alfalfa were the least favorable. How- 
ever, Anderson and Wright (1952) found during Montana field studies 
that feeding occurred on a wide variety of plants, representing a 
wider host range than suggested by Pfadt’s cage studies. Hebard 
(1929, p. 391) recorded a pair of Melanoplus (probably bilituratus 
bilituratus) from Perth, Nev., which were very small, with malformed, 
reduced wings. They occurred in alfalfa and, although normal 
specimens were abundant, perhaps the pair in question fed entirely 
or mainly on alfalfa. Scharff (1954) reviewed the effects of food plants 
and weather. 

Environments occupied by bilituratus bilituratus are extremely 
variable, Hebard (1925a, p. 112) stating that it (alanis) ‘‘is equally at 
home in weedy waste land or on the sage-brush plains, its environment 
having a very decided effect on its general appearance.’’ The general 
preference for light, sandy soils has been mentioned by Morse (1920). 
In the grain belt (Shotwell, 1930) it frequents fence row areas and 
wheat-stubble fields, especially where Russian thistle grows. In 
southern Michigan, Cantrall (1943) concluded that this grasshopper 
is characteristic of, and prefers, a mixed grass-herbaceous habitat, and 
that in many other environments where the grasshopper appears it 
probably is a sporadic member of the fauna, since it wanders a great 
deal. 

Altitudinally, the range of bilituratus bilituratus captures extends 
from sea level in many places to the Alpine Zone (see Alexander, 1951, 
pp. 107-108, for critical comments on occurrence in the Alpine Zone 
of Colorado, also notes by Gurney, 1953, p. 317). Through the 
courtesy of Dr. George E. Wallace, Carnegie Museum, we have 
examined a rather long-winged male of bilituratus bilituratus taken by 
the late Dr. Walter Sweadner, Aug. 9, 1932, at an altitude of 7,200 
feet, imbedded in the Sperry Glacier, Glacier National Park, Mont. 
No further details are available. 

In New England bilituratus bilituratus occurs from the seashore to 
the summits of the highest mountains (Morse, 1920). Some altitudes 
on the labels of specimens examined are as follows: Junction of Moose 


VOL. 110 


PROCEEDINGS OF THE NATIONAL MUSEUM 


24 


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snupoixaw * 


GRASSHOPPERS—-GURNEY AND BROOKS 25 


Pass-Hope Highway and Moose Pass-Kenai Highway, Alaska, 1,800 
ft.; Signal Peak, Wash., 5,000 ft.; Wasatch Mts., Utah, 7,400 ft.; 
Chaffee Co., Colo., 8,000—8,500 ft. 

The classic report on the effects of temperature and moisture on 
this grasshopper is by Parker (1930). MacCarthy (1956) dealt with 
the correlation between adult populations and weather factors in 
Saskatchewan. 

Criddle (1918, 1924) and Shotwell (1930) are among those who have 
described the egg pods of biliiwratus bilituratus, and Cantrall (1943) 
gave a detailed account of the way the ovipositing female prepares 
the hole in the soil. Parker (1930) recorded one caged female which 
laid 15 egg pods, and the average for 30 females observed by him was 
8.8 pods. Other data on egg production were presented by Pfadt 
(1949) and Smith, Handford, and Chefurka (1952). In laboratory 
cages, using material from central Saskatchewan, Pickford (personal 
communication to A. R. B.) has obtained an average of 30 pods for 
each of 9 females with a calculated maximum of 43 pods (872 eggs) 
for one female of the lot. 

A description by Handford (1946) compared nymphs of bilituratus 
bilituratus with those of other species of Melanoplus. The seasonal 
history varies a great deal, depending on the weather (Shotwell, 1930, 
p. 23) and the location. The majority of adults studied by us were 
collected in July or August, but those collections undoubtedly reflect 
the habits of collectors almost as much as those of the grasshopper. 
Shotwell has noted oviposition as late as November. Cantrall (1943) 
found adults from early July until late October in Michigan, and he 
suspected that there were two generations. It is suspected that two 
years are required for the completion of the life cycle in parts of 
British Columbia (Canad. Insect Pest Rev., vol. 34, p. 273, 1956). 


Melanoplus bilituratus vulturnus, new subspecies 
Figures 1,d-1; 2,j,m; 3,c; 6; 7,d; 8,k; 9,f; 10,0; 15,b,c; PLate 1,p,£. 


Mate (holotype): Size medium for group; fully winged; general 
appearance as in M. bilituratus bilituratus. Pronotum not strangulate 
in dorsal view, the posterior margin with the sides forming an angle 
of about 120°, apex well rounded; median carina of prozona and meta- 
zona proportioned as to length as 21 : 24, in lateral view only slightly 
depressed at the principal sulcus; tegmen with portion extending 
posterior to the apex of the hind femur in proportion to tegminal 
length as 3.5:17.5; hind femur moderately swollen for group. 
Furculae slender, sharply tapered, scarcely divergent, extending one- 
third the length of supra-anal plate; cercus as in figure 7,d-1; sub- 
genital plate moderately upturned, with twin apices closely associated, 
evident but poorly separated. 


26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Aedeagus (dry) as in figure 3,c-1, the apex of the dorsal valve 
strongly curved laterally with anterior margin very heavily sclerotized 
and excavate along entire length, main portion of dorsal valve less 
heavily but well sclerotized, decidedly concave, dorsal marginal 
membrane inconspicuous. Epiphallus with apex of each lophus 
(fig. 8,k) broad in dorsal view, in lateral view with lophus erect, 
much elevated, the apical margin barely angulate anteriorly. 

Coloration: General coloration pale brown, with the customary 
markings of blackish, mottled with undertones of brown; head with 
fairly wide blackish brown postocular bar; pronotum mottled with 
brown above, laterally with irregular blackish longitudinal bar; tegmen 
lightly speckled with brown, markings very inconspicuous on apical 
half; hind femur with three fairly noticeable dorsal dark bars, these 
weakly suggested on external paginal area, ventral portion of paginal 
area yellowish, section between paginal area and ventral keel pink, 
ventral and mesal surfaces yellowish, with traces of pinkish, knee 
crescents black; hind tibiae pale, greenish gray mesally and yellowish 
brown externally, spines and apical half of spurs blackish brown. 

Measurements in millimeters: Body, 22.0; pronotum, 4.5; front 
femur, 3.5; hind femur, 12.0; tegmen, 17.5. Greatest width of 
pronotum (posterior, including lateral lobes in perspective from above), 
3.8; of hind femur, 2.9. 


EXPLANATION OF FiGuRE 7 


a-j: Lateral view of left cercus of male. 

a: Melanoplus mexicanus: 1, from Bledos, Mexico; 2, from Teotihuacan, Mexico. 

b: M. spretus: 1, neotype; 2, from Kansas, 1874. 

ce: M. bilituratus defectus, Casa Grande, Ariz. 

d: M. bilituratus oulturnus: 1, holotype; 2, from Quincy, Fla.; 3, from Independ- 

ence, Kans.; 4, from Nantucket, Mass. 

e: M. bilituratus defectus: 1, from Tempe, Ariz.; 2, from Camarillo, Calif.; 

3, from Mount Charleston, Nev. 

Ff: M. bilituratus bilituratus: 1, from Gun Lake, Mich.; 2, from Washington 
Territory; 3, from Spokane, Wash.; 4, from Eureka, Calif.; 5, from Placer 
Co., Calif.; 6, from Doyle, Calif. 

M. bilituratus bilituratus, with abnormal dorsal valve, Fort Benton, Mont. 
M. devastator: 1, from Clarkson, Calif.; 2, from Roseburg, Calif.; 3, from 
Sisson, Calif.; 4, from Grant’s Pass, Oreg.; 5, from Alpine, Calif. 
i: M. bruneri: 1, 2, from Ft. McLeod, Alberta, Canada (2 specimens); 3, from 
Chilcotin, British Columbia, Canada; 4, from Mt. Moriah, Nev. 
j: M. borealis borealis: 1, from Churchill, Manitoba, Canada; 2, from Laggan, 
Alberta, Canada; 3, from Matanuska, Alaska; 4, from Cummington, Mass.; 
5, from Grant, Colo. 
k,l: Ventral view of right cercus of male (basal end at bottom of page): 
k: M. devastator, Tacoma, Wash. 
l: M. bilituratus bilituratus, Dorris, Calif. 


Sale 


GRASSHOPPERS—GURNEY AND BROOKS 


J | ‘ 


Ficure 7.—Explanation on facing page. 


27 


28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Frmate (allotype): General form as in male, slightly more robust; 
differing as follows: Compound eyes less globose; longitudinal carinae 
of vertex more widely separated and less prominent; median carina 
of prozona and metazona proportioned as 21:26, in lateral view some- 
what more angulate at principal sulcus; mesosternal hump lacking; 
cercus as in bilituratus defectus (fig. 10,h); dorsal valve of ovipositor 
as in figure 10,0. 

Coloration: Somewhat darker than in holotype; hind tibia red. 

Measurements in millimeters: Body, 23.0; pronotum, 4.7; front 
femur, 3.2; hind femur, 13.0; tegmen, 18.0. Greatest width of 
pronotum, 4.2; of hind femur, 3.2. 

VARIATION: The size of 10 representative males measured (in milli- 
meters) varies in pronotal length from 3.5 to 5.0 (av. 4.4), in length 
of hind femur from 10.0 to 14.0 (av. 12.0), and in length of tegmen 
from 15.5 to 21.8 (av. 18.3). Six measured females vary in pronotal 
length from 4.2 to 5.3 (av. 4.7), in length of hind femur from 11.5 to 
14.0 (av. 12.8), and in length of tegmen from 16.5 to 23.5 (av. 18.5). 
Western specimens average larger than eastern ones. In most speci- 
mens the tegmen extends about 2 to 3 mm. beyond the apex of the 
hind femur, but in some it scarcely exceeds it by as much as 5 mm. 
The male cercus usually is essentially like that of the holotype, but 
the tiny apicoventral “tooth” frequently is absent, and general shape 
is somewhat variable (fig. 7,d-2,d-4). Very little variation in the 
shape of the epiphallus has been noted. 

Intergradation is shown by the shape of the dorsal valve of the 
aedeagus; in intermediate specimens the ventroanterior margin of the 
dorsal valve varies between the clearly excavate one of bilituratus 
vulturnus and the simply recurved margin of bilituratus bilituratus. 
Wet preparations often show membranous folds apically (“‘m” of 
fig. 3,c). Some intergrading specimens represent one of the two sub- 
species more strongly; others represent the other one, and some are 
fully intermediate. In some cases, as is true of material from Erie 
and Wood Counties, Ohio, and from Lafayette, Ind., fairly large 
series of males taken at the same time and place vary. This situation 
suggests that a mingling of genes of the two subspecies is occurring 
and that the populations are intergrading, though certain individuals 
appear nearly typical of one subspecies. 

A specimen collected at Muscatine, Iowa, Sept. 12, 1935, by R. L. 
King probably is bilituratus vulturnus, but the dorsal valve (fig. 2,m) 
is very unusual, recalling certain very long-winged Arizona specimens 
of bilituratus defectus. The specimen is very long-winged, the tegmen 
27 mm. long and extending 11.5 mm. posterior to the hind femur. 
In 1935 Iowa was experiencing a buildup of grasshoppers, but there 
is no record of extensive flights into the State then, though at other 


GRASSHOPPERS—-GURNEY AND BROOKS 29 


times “light swarms of what appeared to be nearly typical spretus 
entered Iowa from the northwest and settled down in the western 
and southwestern parts of the state’ (Drake and Tauber, 1946). 
The aedeagus of this specimen appears to be abnormal, but an 
explanation of this condition merits future attention. 

Special interest is attached to the material reared by Brett (1947) 
which he ascribed to M. mezxicanus and the most long-winged speci- 
mens of which he believed comparable to spretus, the latter considered 
by him a synonym. Specimens were deposited in the U.S. National 
Museum by Dr. Brett, and additional ones have been loaned by 
F, A. Fenton of Oklahoma A. & M. College, where Brett’s main 
reared series are preserved. He reared specimens in cages, mainly 
feeding head lettuce, alfalfa, corn, and sorghum. The largest indi- 
viduals reared were fed head lettuce at 100° F. and 35 percent and 
50 percent relative humidity. Some of them resembled spretus in 
body size and tegminal length, though none of them examined by us 
has the spretus-type of aedeagus. At the other extreme, individuals 
reared on alfalfa were comparatively small, and in many of those 
reared under cool, humid conditions the tegmina were poorly developed 
as well as short. 

The aedeagus of a total of 65 males from the Brett material has 
been examined, these specimens representing the extremes of temper- 
ature and relative humidity under which rearing occurred, as well 
as tegminal lengths varying from abortive to extending as much as 
6 mm. posterior to the hind femoral apices. Dr. Brett has informed 
us (in litt., Dec. 17, 1956) that his steck used at temperatures of 75° 
and 80° F. was collected at Bennet, Nebr. (about 11 miles southeast 
of Lincoln), that stock for rearings at temperatures above 80° was 
collected near Stillwater, Okla., and that no Kansas stock was used. 
Reared material from both the Nebraska and Oklahoma stocks is 
variable in the structure of the dorsal value of the aedeagus, and there 
are indications that both stocks are from tension areas where bilituratus 
bilituratus and bilituratus vulturnus influence occurs, but the great 
majority of males may readily be recognized as bilituratus vulturnus 
and we have not considered them intergrades. Males collected at 
Lincoln, Nebr., and Stillwater, Okla., are comparable to the reared 
series in aedeagal structures and variation. No reared specimen 
has an aedeagus like that of spretus, but several reared specimens 
have the dorsal valve well sclerotized and with little of the shape 
characteristic of adults of bilituratus vulturnus. It seems most likely 
that the aedeagus of these specimens failed to mature completely, 
though the cerci, supra-anal plate, and subgenital plate are, with 
few exceptions, the type normal for adult bilituratus vulturnus. The 
tegmina of these particular specimens are mostly abortive or mal- 


30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


formed, suggesting that the individuals had difficulty developing 
normally. 

Twelve males from Pittsburgh, Pa., have been examined, including 
four from a single area within the city, Forbes Field. As a whole, 
the aedeagus indicates an intergrading population, with no specimens 
fully typical of either bilituratus bilituratus or bilituratus vulturnus. 
There is variation, even within the series from Forbes Field, but most 
of the specimens are nearer bilituratus vulturnus. Although Presque 
Isle, Pa., is considerably north of Pittsburgh, the two males from 
there both are typical of bilituratus vulturnus. Perhaps a series would 
show intergradation. Jennings (1909) reported that though the 
flora of Presque Isle is more northern than southern in its affinities, 
some of the plants are typical of those centering in the lower Ohio 
basin and on the Piedmont Plateau. Possibly this grasshopper 
represents such a relationship. Jennings illustrated the many 
ecological habitats which occur on Presque Isle. 

The single male from Nantucket examined is typical bilituratus 
vulturnus, though a male from Marion, in the nearby Fall River area 
of Massachusetts, is intermediate. Other Massachusetts specimens 


EXPLANATION OF FicurRe 8 


a-e: Left cercus of male, lateral view: 

a: Melanoplus borealis palaceus: 1, holotype; 2, topotype; 3, 4, from Lake- 
view area, Mont. (2 specimens). 

b: M. borealis utahensis: 1, holotype; 2, from Ogden, Utah; 3, from Big 
Cottonwood Canyon, Utah. 

c: Intermediate between M. borealis utahensis and M. b. palaceus, 3 miles 
north of Mantua, Utah. 

d: M. borealis stupefactus: 1, lectotype; 2, from Los Pinos, Colo.; 3, from 
Mineral Co., Colo. 

e: Intermediate between M. borealis stupefactus, M. b. palaceus, and M. b. 
borealis, Tomboy, Colo. 

f-i: Outline of epiphallus in dorsal view: 

f: M. borealis borealis, Nachvak Bay, Labrador, Canada. 

g: M. b. utahensis, holotype. 

h: M. b. stupefactus, holotype of latifercula. 

i: M. bilituratus defectus, Casa Grande, Ariz. 

j-l (2): Outline of epiphallus of male in dorsal view, lophi only: 

j: M. bilituratus bilituratus, Tuttle, Idaho. 

k: M. bilituratus oulturnus, holotype, dry preparation. 

lz: M. devastator: 1, from Davis, Calif.; 2, from Sacramento Co., Calif. 

1(3)-m: Epiphallus in lateral view [/ (5)—m showing lateral outline of lophus only): 

I: M. devastator: 3, lectotype; 4, from Roseville, Calif.; 5, from Tacoma, 
Wash., dry preparation; 6, from Tulare, Calif., dry preparation; 
7, from Gibson, Calif.; 8, same specimen as 7, in lateroposterior view; 
9, from Lower Lake, Calif. 

m: M. bilituratus bilituratus, Milford, Calif. 


PAV LYN BRA 
DAP NSN 
Pet NS TS 


32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


are typical bilituratus bilituratus. Detailed field work in south- 
eastern Massachusetts might be very informative. Scudder (1878a) 
noted that Nantucket specimens (reported as atlanis) have a peculiar 
coloring and nearly all specimens have pale glaucous hind tibiae. 
Griscom and Folger (1948) reported that the limited flora and fauna 
of Nantucket include “many southern and relict species,’’ and the 
climate averages milder than that of Boston. A comparable situation 
probably occurs in the relation of Long Island to upstate New York. 
It is surprising that intermediate specimens or typical bilituratus have 
not been found in the Appalachian Mountains south of Pennsylvania. 
Specimens from Spruce Knob, W. Va., and Mt. Davis, Pa., show a 
slight trend toward bilituratus bilituratus, though they are not recog- 
nized as intermediates. 

Color variation in bilituratus vuliurnus runs through the usual range 
common to this group. About 18 percent of the specimens examined 
have pale or glaucous hind tibiae, the remainder red. Brett (1947, 
pp. 32-33) felt that food, temperature, and humidity are closely 
related to the color of the hind tibiae. Specimens reared by him on 
corn and head lettuce had a higher percentage of red hind tibiae than 
those reared on alfalfa. High temperature and humidity tended to 
produce red tibiae. On the other hand, King and Slifer (1955), after 
rearings from stock collected at Iowa City, Iowa, concluded that red 
hind tibiae “are dependent on the presence of a dominant, autosomal 
gene and that blue tibiae are produced when its recessive allele is 
present in the homozygous condition.””, Nymphs which Brett reared 
under crowded conditions tended to develop dark patterns character- 
istic of the gregarious phase of migratory grasshoppers, but adults 
resulting from such rearings were not distinctively colored, and they 
tended to have shorter wings than those reared under solitary con- 
ditions (Brett, 1947, p. 37). 


EXPLANATION OF FIGURE 9 


Dorsal view of supra-anal plate and furculae of males; f and h(2) show full view of apical 
portion of abdomen. 


a: M. bruneri: 1, from Ft. McLeod, Alberta, Canada; 2, from Mt. Moriah, Nev.; 3, from 
Crook Co., Wyo. 

b: Intermediate between M. borealis utahensis and M. b. palaceus, 3 miles north of Mantua, 
Utah. 

c: M. borealis borealis: 1, from Matanuska, Alaska; 2, from Chelsea, Iowa; 3, from Church- 

ill, Manitoba, Canada. 

M. borealis stupefactus: 1, from Los Pinos, Colo.; 2, from Mineral Co., Colo. 

M. borealis utahensis: 1, from Big Cottonwood Canyon, Utah; 2, from Ogden, Utah. 

M. bilituratus oulturnus, Stokes, S. C. 

M. bilituratus bilituratus, Pompey’s Pillar, Mont. 

M. devastator, 1, from San Bernardino, Calif.; 2, from Chico, Calif. 


TAT A 


GRASSHOPPERS—GURNEY AND BROOKS 


Figure 9.—Explanation on facing page. 
477119593 


34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Type: USNM 63543. 

TYPE LOCALITY: Stokes, Colleton Co., S. C. 

The holotype male was collected by Miss Lotta Bootle in September 
1924. The allotype (USNM) was taken at Columbia, S. C., Sept. 22, 
1923, by Philip Luginbill. 

There are 99 male and 36 female paratypes deposited in the U. S. 
National Museum, Canadian National Museum, Academy of Natural 
Sciences of Philadelphia, Museum of Comparative Zoology, University 
of Michigan, University of Arizona, Clemson Agricultural College, 
Florida Plant Board, and the collection of John R. Hilliard. The para- 
types are from the following localities: 

Norra Carouina: Weldon, Greensboro, Pilot Mt., Charlotte, Roan Valley, Mt. 
Mitchell, Asheville, Hendersonville, Black Mts., Valley of Black Mts., Balsam, 
Sunburst, Highlands. 

Soutu Carotina: Green Sea, Mullins, Cheraw, Florence, Manning, Columbia, 
St. Paul, Eutawville, Stokes, Clemson, Madison. 

GrorciA: Rabun Bald, Black Rock Mountain, Tallulah Falls, Louisville, Dublin, 
Stone Mountain, Atlanta, Locust Grove, Cedar Rocks (5 miles north of Jackson), 
Barnesville, Macon, Montezuma, Smithville, Calhoun, Sand Mountain (near 
Trenton). 

Fiorina: Gainesville, Alachua Co., Hamilton Co., Houston, Lake Miccosuke 
(Jefferson Co.), Quincy, Gadsden Co., Rock Bluff Landing (Liberty Co.), 
“Camp Torreya” (Liberty Co.), Jackson Co., Marianna, Chipola. 

ALABAMA: Auburn, Stokes, Selma, Greenville, Flomaton, Canebreak. 

MississipP1: Meridian, Perkinston, Jackson, Vicksburg. 

LovistANA: Harahan, Ruston, Sibley. 


Most of the specimens from additional localities shown on the map 
(fig. 6) for bilituratus vulturnus are typical, but some vary toward 
bilituratus bilituratus, though not sufficiently to be considered inter- 
mediates, and so are not designated as paratypes. However, males 
from other localities, as mapped, are considered intermediates between 
bilituratus bilituratus and bilituratus vulturnus. 

BIOLOGY AND ECONOMIC IMPORTANCE: In many localities bilituratus 
vulturnus becomes extremely abundant, and it is highly important 
economically, especially in the western part of its range. Kansas, 
Oklahoma, and Missouri are the chief States where it is very injurious. 
Wilbur and Fritz (1940) stated that during the 1933-39 outbreak in the 
bluestem region of eastern Kansas it was the most injurious crop infest- 
ing grasshopper, and they briefly described the buildup of its popu- 
lation during several drought years. 

Morse (1904, p. 42) referred to this grasshopper as ‘‘probably the 
most widely and generally distributed” of those found in the South- 
eastern States, but this opinion differs from that of Rehn and Hebard 
(1916, p. 237), who found bilituratus vulturnus (reported as atlanis) 


GRASSHOPPERS—GURNEY AND BROOKS 35 


“relatively infrequent in the Coastal Plain south of North Carolina 
and the lower Gulf drainage of southern Georgia, in addition to having 
never been definitely recorded from peninsular Florida.’’ Material 
examined suggests that in northern Florida bilituratus vulturnus is 
fairly widely distributed, though it is not generally abundant. Friauf 
(1953) examined a wide variety of grassland and other habitats on an 
area of about 2,180 acres in Putnam County, Fla., collecting an esti- 
mated 24,000 specimens of Orthoptera and Dermaptera during a period 
of more than 15 months in the field, yet he did not find bilituratus 
vulturnus. There is a paucity of records from the lower coastal plain 
of South Carolina and Georgia. The pertinent physiographic regions 
are shown on a map by Rehn and Hebard (1916, p. 95), and a com- 
parison of their collecting stations (map, p. 91) with those indicated 
by Morse (1904, p. 10) suggests that the latter did relatively less 
sampling of the coastal plain fauna in Georgia and South Carolina, 
while his collecting trip (made in 1903) did not include peninsular 
Florida at all. 

This grasshopper occurs most frequently in fields and uncultivated, 
open areas. Fox (1917) mentioned its tendency to congregate in 
Virginia in farm lands on sandy soils, and Isley (1938), in eastern 
Texas, classified it as a species of shallow soil. Like the northern 
subspecies, bilituratus vulturnus is found in many less typical environ- 
ments, such, for example, as the shale barrens of Virginia which 
Gurney (1941) described as the habitat of M. impudicus Scudder. 
Fox reported it quite scarce in the higher Appalachians, but our records 
and those of Hebard (1945) show that it is frequent in clearings at 
higher elevations. The following are among the altitudes cited on 
labels of specimens examined: White Top Mountain, Va., 5,500 ft.; 
Little Bald, Great Smoky National Park, Tenn., 4,800 ft.; Unaka 
Mountains, Tenn., 3,500 ft. 

As to food, Isley (1938) found that bilituratus vulturnus (reported 
as mexicanus) “selected rather widely among monocotyledons and 
dicotyledons showing a definite preference for crop plants.” 

In a report on the sculpturing and other detailed features of grass- 
hopper eggs by Tuck and Smith (1939), descriptions of mezicanus 
eggs probably are attributable to bilituratus vulturnus because the 
work was conducted mainly in eastern Kansas. 

Regarding the seasonal occurrence of adults, the following early 
and late dates from labels of specimens examined may be cited; 
Jackson Co., Ohio, June 12, 1932; Vinton Co., Ohio, Nov. 24, 1936; 
McNeill, Miss., May 8, 1925; Austin, Tex., April 15, 1956; College 
Station, Tex., June 17, 1920, Nov. 20, 1924; Weatherford, Okla., 


36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


June 6-7, 1938; Ft. Sill, Okla., Nov. 10, 1918; House Springs, Mo., 
Nov. 11, 1925; Clarksville, Tenn., Oct. 14, 1916. In Virginia, Fox 
(1917) concluded from observations of adult prevalence that there 
are two generations per year. He found that adults appear about 
June 1, reach a peak of abundance, and then in late July decline in 
numbers; about the middle of August he noted the appearance of a new 
group of adults. In Kansas, Wilbur and Fritz (1940) recorded a peak 
of abundance from the middle of June until late July; in 1934 there 
was a fall peak of adults between October 6 and November 9. 

Remarks: The name of this subspecies is from ‘“‘vulturnus ventus,”’ 
a Roman term for a south-southeast wind, chosen in allusion to the 
distribution of this subspecies. 


EXPLANATION OF FicuReE 10 


a-e: Epiphallus in lateral view, males: 
a: Melanoplus spretus, Neotype. 
b: M. bruneri: 1, from Ft. McLeod, Alberta, Canada; 2, from Mt. Moriah, 
Nev. 
c: M. borealis borealis, Nachvak Bay, Labrador, Canada. 
d: M. borealis stupefactus, holotype of latifercula. 
M. borealis utahensis, holotype. 
ercus of females: 
M. mexicanus, Cuernavaca, Mex. 
M. bilituratus bilaturatus, Hogeland, Mont. 
M. bilituratus defectus, San Carlos Indian Reservation, Ariz. 
M. devastator, Gazelle, Calif. 
M. bilituratus defectus, Chandler, Ariz. 
M. bruneri, Isle Royale, Mich. 
M. borealis borealis, lectotype of septenirionalts. 
m: M. borealis stupefactus, Mineral Co., Colo. 
n-p: Left dorsal valve of ovipositor: 
n: WM. bilituratus bilituratus, Hogeland, Mont. 
o: M. bilituratus oulturnus, allotype. 
p: M. devastator, Gazelle, Calif. 
q: Female apical abdominal structures of M. mexicanus, Cuernavaca, Mexico. 
r-t: Left dorsal valve of ovipositor: 
r: M. bruneri, Isle Royale, Mich. 
s: M. borealis borealis, lectotype of sepientrionalts. 
t: M. borealis palaceus, allotype. 
u-x: Apex of male subgenital plate in posterior view: 
u: M. bilituratus defectus, Chandler, Ariz. 
vo: M. spretus: 1, from Pueblo, Colo., July 8-9, 1877; 2, from Argentine Pass, 
Colo., July 16, 1877. 
w: M. bilituratus bilituratus, Poplar, Mont. 
x: M. devastator: 1, from Merced River, Yosemite National Park, Calif.; 
2, from Tehachapi, Calif.; 3, from Alpine, Calif.; 4, from Grant’s Pass, 
Oreg. 


8 


fm: Left 


oO 


m See FT SH 


GRASSHOPPERS—GURNEY AND BROOKS 


ome BN 


| 2 3 é 


Ficure 10.—Explanation on facing page. 


38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Melanoplus bilituratus defectus Scudder, new combination 
Ficures 2,1,1; 3,d,h; 6; 7,c,e; 8,2; 10,h,j,u; 16,0; Phares 2,A,B,C; 4,A. 


Melanoplus defectus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 20, 32, 
1897; Proc. U. S. Nat. Museum, vol. 20, pp. 132, 177-178, pl. 12, fig. 6, 
1897. (One male, one female, Grand Junction, Mesa Co., Colo.) Lectotype, 
designated by Rehn and Hebard (1912, p. 80): Male from Grand Junction, 
Colo. (ANSP). 


Hebard (1929, p. 390) placed defectus as a synonym of brunera, 
but the aedeagus of the lectotype shows that it belongs to the Grand 
Junction population of bilituratus, here considered to be encompassed 
within the limits of the southwestern subspecies. The lectotype is 
small, with a body length of 19.5 mm. The tegmina fail by about 
2 mm. to reach the tips of the hind femora. The dorsal valve of 
the aedeagus is poorly sclerotized and pale, but is comparable in 
shape to other specimens of this subspecies from southwestern Colo- 
rado. The optimum development of bilituratus defectus occurs in 
the central and southern portions of Arizona and New Mexico, and 
it is unfortunate that the type locality is not within that area. The 
lectotype shows closer relationship to the optimum development of 
bilituratus defectus than it does to bilituratus bilituratus. The term 
“intermediate” as used for this subspecies (including the explanation 
of fig. 6), connotes specimens showing intergradation between the 
optimum population of bilituratus defectus and bilituratus bilituratus. 
From a nomenclatural point of view those specimens are not inter- 
mediates comparable to those discussed elsewhere in this paper 
because the type locality of bilituratus defectus is outside the area of 
optimum development. 

The most distinctive character of bilituratus defectus is the dorsal 
valve of the aedeagus, which in specimens of optimum development 
has the anterior margin (x of fig. 3,d) concave and the lateral flap- 
like area (y) wide and with a sinuate margin (z). In some speci- 
mens, mainly those intermediate with bilituratus bilituratus (fig. 3,f), 
the lateral portion is not typically developed. (Also see second 
paragraph below.) ‘The male cercus usually is about as in figure 
7,e-2), though occasionally it is much broader (fig. 7,e-3). Especially 
in intermediate specimens from Baker, Nev., the cercus is narrower. 

VARIATION: The size of 13 representative males, measured in milli- 
meters, varies in pronotal length from 3.5 to 5.7 (av. 4.3), in length 
of hind femur from 9.8 to 14.5 (av. 12.4), and for length of tegmen 
from 11.5 to 22.5 (av. 17.8). Eight measured females vary in pro- 
notal length from 4.5 to 6.0 (av. 5.4), for length of hind femur from 
13.0 to 16.0 (av. 14.5), and for length of tegmen from 17.5 to 23.4 
(av. 21.0). Specimens from Ft. Wingate, N. Mex., and Kaibab, 


GRASSHOPPERS—GURNEY AND BROOKS 39 


Ariz., are small and dark, in marked contrast to the richly colored, 
more yellowish specimens reared on winter mustard at Tempe, Ariz. 
In average specimens of bilituratus defectus the tegmen extends about 
4 mm. beyond the hind femur. With the exception of material noted 
in the next paragraph, extremes among specimens examined are a 
male from Kaibab, Ariz., and one from Phoenix, Ariz., in which the 
tegmen extends 1.5 and 7 mm., respectively. 

Six males and 11 females from several Arizona localities are so 
long-winged and, in the case of males, have so unusual an aedeagus 
that they are discussed separately in order that all the data will be 
assembled for future reference if importance should become attached 
to them as a distinct population. The aedeagus in lateral view 
(figs. 2,/; 3,h) has the dorsal valve extending anteriorly and with 
little development of a concave lateral surface; however, the dorsal 
valve differs slightly in length and shape, and in one specimen the 
anterior apical margin is very weakly recurved in a suggestion of 
bilituratus defectus; another specimen (from Tumacacori National 
Monument) has a second right dorsal valve present, which approaches 
that of bihturatus defectus, although the left one and the other right 
one are essentially as figured. It appears that the aedeagus in this 
series is abnormal or not fully developed, and that the specimens are 
bilituratus defectus. Fred Skoog, collector of the Chandler specimens, 
has informed us that they were taken among bilituratus defectus 
where the population was 50 to 75 per square yard in an alfalfa 
field, and 500 to 1000 along the margins of the field. The few un- 
usually long-winged specimens found by him had only recently 
become adult, and their somewhat teneral condition may have had 
a bearing on the shape of the aedeagus. Other genitalic characters 
of these specimens fall within the normal variation of bzlituratus 
defectus; the following figures illustrate them: 7,c; 8,7,; 10,7,w; 16,6. 

The general appearance of the above specimens is as shown (plates 
2,c;4,A). Gurney (1953, pl. 7, fig. c) previously illustrated one of 
the females from Chandler, and Barnes (1956) referred to long-winged 
Arizona specimens; both authors applied the name M. mezxicanus 
mexicanus. The lengths of the tegmen and hind femur of these 
specimens are, respectively, as follows: Males, 26.0-31.0 (av. 27.90), 
13.3-14.8 (av. 14.02); females, 27.0-33.5 (av. 29.67), 15.0-17.2 (av. 
15.93). The extremes and averages of the distance from apex of 
hind femur to apex of tegmen are as follows: Male, 9.0-14.0 (av. 11.28) ; 
female, 8.0-13.5 (av. 10.04). 

The following label data accompany this long-winged material (all 
from Arizona): Casa Grande, swept from alfalfa, July 20, 1951, 
N. J. Nerney (1); Chandler, lush irrigated alfalfa, May 1945, 
F. Skoog (2c, 89); Tempe, alfalfa, Aug. 31, 1938, O. L. Barnes 


40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


(10%); Tempe, swept from alfalfa, June 8, 1950, N. J. Nerney (19); 
Theba, on alfalfa, Oct. 26, 1939, O. L. Barnes (1); Tumacacori 
National Monument (Pendleton Ranch), Sept. 9, 1939, E. R. Tinkham 
(1°); Yuma, alfalfa, Aug. 2, 1938, O. L. Barnes (19); Yuma, alfalfa, 
Nov. 11, 1936, O. L. Barnes (19). 

Of 311 specimens of bilituratus defectus examined, only two (one 
each from Las Vegas, N. Mex., and Lupton, Ariz.) have glaucous 
hind tibiae; the others are red. However, 13 of 57 specimens which 
are intermediate with bilituratus bilituratus have glaucous hind tibiae. 
None of these intermediate specimens is as long-winged and large as 
the most optimum specimens of bilituratus defectus. Several of the 
intermediates from southern Colorado and northern New Mexico are 
noticeably small and dark. 

The distribution of males of bilituratus defectus examined is shown in 
figure 6. The single Texas locality is Phantom Lake, Davis Moun- 
tains. 

BioLoGY AND ECONOMIC IMPORTANCE: This grasshopper is important 
economically and is most injurious in irrigated areas (Ball, Tinkham, 
et al., 1942; Harper, 1952). It is said by Harper not to have become 
a pest in the Imperial Valley of California until 1915 when farming 
there became extensive. Cultivated crops are injured more severely 
than grazing areas. 

Barnes (1955) compared five plants as food for bilituratus defectus 
in cage studies in Arizona, and his paper should be consulted for 
observations on mating, egg development, adult body dimensions, 
and weights of adults, as influenced by diet. Two annual generations 
usually are reported, and Barnes feels that a considerable difference 
in average size of individuals representing the two generations probably 
is due to the plants available as food. For instance, in 1944 the second 
generation in the Yuma, Ariz., area developed almost exclusively on 
dry alfalfa and Bermuda grass, and resulting adults were much smaller 
than first generation adults. A comparable difference in adult size 
at the San Carlos Indian Reservation, Ariz., was noted in the two 
generations in 1953. First generation nymphal diet consisted mainly 
of forbs, while the second generation had to develop mostly on grasses. 
It may be significant that all of the especially long-winged specimens 
discussed in detail, which bear habitat data, were associated with 
alfalfa. 

In southern California (Harper, 1952), bilituratus defectus first 
hatches by March or early April; then the second generation occurs 
in September and October. Some early and late dates on labels of 
adults examined are as follows: Yuma, Ariz., Jan. 5, 1937; Bard, 
Calif., Jan. 19, 1937; Yuma, Calif., Apr. 17, 1942; Saugus, Calif., 
Oct. 19, 1950; Zion National Park, Utah, June 18, 1953; same, Sept. 


GRASSHOPPERS—GURNEY AND BROOKS 41 


5, 1926; Peeples Valley, Ariz., Oct. 2, 1936; Chino Valley, Ariz., 
Nov. 23, 1936; Roll, Ariz., Nov. 12, 1943; same, Dec. 1, 1936. 

Altitudinally, bilituratus defectus ranges in distribution from about 
sea level in Imperial Co., Calif., to 11,800 feet on Mt. Charleston, Nev. 
Other high altitudes indicated on labels of specimens examined are 
the following: Magdalena Mts., N. Mex., 7,100 ft.; Lee Canyon, 
Spring Mountains, Nev., 8,600 ft.; Rim Rock Peak, Sandia Mts., 
N. Mex., 9,500-10,000 ft. The specimens from high elevations 
average somewhat smaller than specimens of optimum development 
from low elevations, and they are relatively dark in color, but in 
general they are not distinctive. 


Melanoplus devastator Scudder 
Ficures, l,e;, 2,4;:3,i;.7,h,k;.8,l;.9,h;/10,i,p,2; 14; Puate 3,8,c. 


Melanoplus devastator Scudder, Proc. Boston Soc. Nat. Hist., vol. 19, pp. 287, 288, 
1878. (Males, females, Shasta Valley, Sissons, Sausalito, Calif.; Lake Tahoe, 
Reno, Glen Brook, Nev.; Beaver Brook, Morrison, Colo.) Lectotype, here 
designated: Male labeled ‘‘Sept. 1, 1874, Sauzalito, Cal.,”’ ‘““Scudder’s Type,” 
“Mel. devastator typicalis Scudder’s Type, 1895,’ ‘Sc. reference Rev. 
Melanopli 1897, p. 198,” “‘M. devastator Scudd. Cab. 8. H. Scudder.” (ANSP). 
Sausalito is one of the original localities mentioned by Scudder (1878b), 
who said that the form named M., d. typicalis ‘‘best represents the original 
types of the species when first described,’ and this specimen is one so identified 
by him. 

Melanoplus ater Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 22, 32, 1897; 
Proc. U.S. Nat. Mus., vol. 20, pp. 133, 194-195, pl. 13, fig. 2, 1897 (two males, 
3 females, San Francisco, Calif.). Lectotype, designated by Rehn and Hebard 
(1912, p. 80): Male (ANSP). 

Melanoplus consanguineus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), 
pp. 21, 32, 1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 192-193, pl. 12, fig. 
10, 1897 (male, female, Sacramento Co., Calif., Sonora Co. in plate explana- 
tion, p. 409, error for Sonoma Co.; see Rehn and Hebard, 1912, p. 80). New 
synonymy. Lectotype, designated by Rehn and Hebard (1912): Male 
(USNM). 

Melanoplus diminutus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 21, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 190-191, pl. 12, fig. 9, 1897 
(males, females, San Francisco, Calif.; Monterey, Calif.). New synonymy. 
Lectotype, designated by Rehn and Hebard (1912, p. 80): Male from Mon- 
terey, Calif. (MCZ). 

Melanoplus uniformis Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 22, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 201-202, pl. 13, fig. 9, 1897 
(males, females, five California localities). New synonymy. Lectotype, 
designated by Rehn and Hebard (1912, p. 80): Male from Sacramento, 
Calif. (USNM). 


3 Labels such as this one, with ‘‘Scudder’s Type, 1895’ printed, were affixed to many of the specimens 
studied by Scudder in connection with his revision (1897b). Species described by other authors, as well as 
those by Scudder, and, where series were available, more than one specimen of the same species, were So 
treated. The principal function of this label today is to indicate a specimen studied by Scudder himself, 
and one presumably considered by him to be typical of the species named. 


42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Melanoplus virgatus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 22, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 133, 199-201, pl. 13, fig. 8, 1897 
(males, females, six California localities). Lectotype, designated by Rehn 
and Hebard (1912, p. 80): Male from Siskiyou Co., Calif. (MCZ). 

Melanoplus devastator affinis Scudder, Proc. U. 8. Nat. Mus., vol. 20, p. 199, pl. 13, 
fig. 6, 1897 (males, Sierra, Sacramento, and Los Angeles Counties, Calif., 
also “California, S. Henshaw.’’). Lectotype, here designated: Male labeled 
“Cal. (Ricksecker) 8. Henshaw,” ‘‘Mel. devastator affinis Scudder’s Type 
1895,” and figured by Scudder, 1897b (MCZ). 

Melanoplus devastator conspicuus Scudder, Proc. U. S. Nat. Mus., vol. 20, p. 199, 
pl. 13, fig. 7, 1897 (males, Sacramento, El Dorado, Amador, and Merced 
Counties, Calif.). Lectotype, here designated: Male labeled ‘Clarkson, 
Eldorado Co. Cal. 14/8, 85, Type No. 1160, U. S. N. M., Mel. devastator 
conspicuus Scudder’s Type, 1895’”” (USNM). (We have been unable to 
locate a Clarkson in El Dorado County, and the label may have been in error 
for Clarksona, Amador County, which is nearby. The male from Sacra- 
mento, Calif., figured by Scudder has not been found. El Dorado County is 
the second locality mentioned.) 

Melanoplus devastator obscurus Scudder, Proc. U. S. Nat. Mus., vol. 20, pp. 198- 
199, pl. 13, figs. 3, 4, 1897 (males, San Francisco, Calif.; Sierra, Placer, Marin, 
Sacramento, El Dorado, Alameda, Siskiyou, and Los Angeles Counties, Calif.). 
Lectotype, here designated: Male labeled “‘Sissons Cal., Drawn, S. H. Scudder 
Coll, Type 15439, Mel. devastator obscurus Scudder’s Type, 1895” and 
figured by Scudder, 1897b (MCZ). 

Melanoplus devastator typicalis Scudder, Proc. U. S. Nat. Mus., vol. 20, p. 199, 
pl. 13, fig. 5, 1897 (males, ‘‘nearly every county . . . from Shasta to San 
Diego and from Marin to Sierra,’”’ Calif.). (The sexes and number of speci- 
mens of this and the previous three entities were not given by Scudder, but 
male specimens were figured. In each case both sexes were before him, as 
shown by preserved specimens.) Lectotype, here designated: Male labeled 
“Tighe Calif.. Drawn, S. H. Scudder Coll, Type 15441, Mel. devastator 
typicalis Scudder’s Type, 1895” and figured by Scudder (1897b) (MCZ). 


Scudder listed Dr. Edward Palmer as the collector of the Tighe 
specimen. We have been unable to find Tighe on available maps, and 
for the information that Tighe was about 8 miles east of Ramona, 
Calif., are indebted to Rogers McVaugh, a botanist who has made a 
study of Palmer collecting localities. This location, about 45 miles 
northeast of San Diego, apparently was a ranch. Palmer’s un- 
published notes indicate that he was there July 5-6, 1875. For a 
biography and sketch of Palmer’s travels, see Safford (Pop. Sci. 
Monthly, vol. 78, pp. 341-354, 1911) and McVaugh’s (1956) recent 
book, wherein (p. 333) the locality is listed as Tigh(e)’s (Ranch or 
Station). 

All the lectotypes mentioned have been examined, in each case 
including the aedeagus and epiphallus, and they are considered to 
represent a single species, (4. devastator Scudder. Kirby (1910) placed 
d. affinis, d. conspicuus, d. obscurus, and d. typicalis as synonyms of 
devastator, and Caudell (1915) did likewise for virgatus and affirmed 
the synonymy of d. affinis and d. typicalis. He also considered 


GRASSHOPPERS—GURNEY AND BROOKS 43 


consanguineus a synonym of diminutus. M. d. affinis should not be 
confused with M. affinis, described by Scudder (1897b, p. 171), 
which preoccupies it. The types of ater, diminutus, and d. obscurus 
are small, very dark individuals; those of uniformis, virgatus, and 
d. conspicuus are noticeably large and yellowish; consanguineus is 
relatively small and moderately dark; d. affinis is medium-sized and 
yellowish; devastator and d. typicalis are medium-sized and moderately 
dark. 

DescriIpTivE Notes: A medium-sized, usually fully winged member 
of the mezxicanus group. Head with dorsal carinae of vertex about as 
in bilituratus bilituratus; mesosternal hump absent or weakly de- 
veloped in male, absent in female. 

MALE GENITALIA: Cercus elongate and slender for group, with a 
characteristic preapical knob extending ventrally and mesally on the 
mesal surface (fig. 7,k), typically as in figure 7,h-1, sometimes pro- 
portionally shorter, often with a small tooth near the ventroposterior 
angle (fig. 7,h-2,3); furculae sometimes more divergent than illus- 
trated (fig. 9,h-2); subgenital plate weakly to moderately upturned 
(pl. 3,8), in posterior view (fig. 10,7) with twin apices usually distinct, 
rarely (fig. 10,7-4) scarcely developed; aedeagus (figs. 2,k; 3,7) with 
dorsal valve lightly sclerotized, rather thin and parchment-like, the 
anterior margin with narrow fold, in dry specimens a basal lobe along 
ventroanterior margin usually evident; when dry the dorsal valve often 
is more concave and the margin is more recurved than when preserved 
wet; accessory lobe medium-sized to large for group; epiphallus 
(fig. 8,/-3,9) distinctive in that lophus is angulate in lateral view 
near apex, the angulation correlated with more or less acute apex in 
dorsal view (fig. 8,/-1,2). 

FEMALE GENITALIA: Cercus (fig. 10,7) relatively acute for group, 
not so robust and blunt as in bilituratus bilituratus; dorsal valve of 
ovipositor with ‘‘scoop”’ well developed. 

VaRIATION: The size of 10 representative males, measured in 
millimeters, varies in pronotal length from 3.7 to 5.3 (av. 4.3), in length 
of hind femur from 9.7 to 15.2 (av. 11.4), and in length of tegmen from 
12.0 to 21.8 (av. 14.8). Seven measured females vary in pronotal 
length from 3.6 to 5.9 (av. 4.8), in length of hind femur from 10.0 
to 15.5 (av. 12.8), and in length of tegmen from 10.6 to 23.1 (av. 
17.5). There is great variation in size, and small specimens are 
frequently encountered in the San Francisco, Calif., area, and in 
Oregon and Washington. The tegmina of the smaller specimens 
usually do not surpass the apex of the hind femur, and in some 
specimens they are as much as 2 mm. short of the femoral apex. 
In larger specimens with optimum development the tegmina fre- 
quently exceed the hind femora by about 3 mm, 


44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


DistriBuTion: Figure 14 shows the distribution of devastator as 
known to the authors. However, uncertainty concerning identifica- 
tions in the group during the past has been responsible for early 
published records which pointed to a much wider distribution. Both 
Essig (1926, p. 81) and Harper (1952) include Arizona in the dis- 
tribution, but Hebard (1935b, p. 304) stated that devastator does not 
occur there. Buckell (1922, p. 31) included devastator in his British 
Columbia list, based on a Vancouver Island record by Caudell. 
Caudell (1908, p. 77) recorded devastator conspicuus from Wellington, 
British Columbia, and a manuscript note written by him in 1926 
indicated that the record was based on a female specimen. The 
specimen, a female with red hind tibiae, is in the U. 8S. National 
Museum, and is bilituratus bilituratus. 

Male specimens of Melanoplus devastator have been examined from 
the localities shown in figure 14. 

BioLoGy AND ECONOMIC IMPORTANCE: When Scudder (1878b) 
originally described devastator, he stated that it probably ‘Ss the 
source of most of the damage to crops in California,” and it frequently 
has been of much importance there. The fullest early account is 
Coquillett’s (1892) report on the 1891 infestation, which includes 
biological notes. Woodworth (1902) stated that devastator was one 
of the two most destructive California grasshoppers in 1901. Brief 
later accounts are by Essig (1926) and Harper (1952). Fulton (1930, 
p. 634) referred to devastator as one of the ‘‘more strictly prairie species” 
of the Willamette Valley of western Oregon and universally present 
there during the proper season. 

A concise summary of the biology and economic importance of 
devastator by Wilson (1947) points out that it is the dominant species 
on much of the semiarid range in California, and that preventing mass 
migrations from natural habitats to cultivated crops is one of the 
critical control measures, in addition to preventing serious damage to 
range forage. Nymphs inhabit much of the range and feed on legumes, 
alfilarias, bromes, and hordeums, in the order named, as long as they 
remain succulent. Wilson, to whose excellent summary the reader is 
referred for more detailed information on habits and seasonal occur- 
rence, commented as follows on the physiological distinctness of 
devastator: 

By the second week of July the species has largely reached the adult stage. To 
the casual observer the adults resemble the lesser migratory grasshopper (M. 
mexicanus Sauss.). However, field and microscopic observations show that these 
two species differ in external anatomy, habitat, and seasonal history. M. devasta- 
tor is naturally found on the range, whereas M. mexicanus is found in California 
and Nevada, largely, if not entirely, in irrigated alfalfa fields. Dissections have 


shown that, unlike M. mexicanus and many of the other economic species that 
begin oviposition within 10 days after reaching the adult stage, M. devastator has a 


GRASSHOPPERS—GURNEY AND BROOKS 45 


preoviposition period ranging from 90 to 176 days, during which no ovarian 
development or copulation takes place. The breaking of this diapause, or period 
of arrested development, apparently depends on temperature and food. Cool 
fall weather, with rains sufficient to germinate the seed of annual plants, appears 
to be the determining factor in breaking the diapause. 


Middlekauff (1958) more recently summarized observations on the 
bionomics of devastator. 

The following are some early and late capture dates noted on 
specimens examined: Catirornia: San Bernardino, May 14; Kern 
Lake, Kern Co., May 22; San Luis Obispo, June 14; Monterey Co., 
Oct. 31; Roseville, Nov. 26; Red Bluff, Dec. 2. Wasuineron: 
Tacoma, Sept. 25. 


Melanoplus bruneri Scudder 
FieureEs 1,9; 2,a,b; 3,k,l; 7,2; 9,a; 10,6,k,r; 11,a; 12; 13; 16,h; Puare 3,a. 


Melanoplus brunert Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 19, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 132, 164-166, pl. 11, fig. 7, 1897 
(males, females, 12 localities in Alberta, Montana, Nebraska, Colorado, and 
Washington). Lectotype designated by Rehn and Hebard (1912, p. 79): Male 
labeled ‘‘Ft. McLeod, B. A. Aug. 82’’ (near southwest corner of Alberta; 
itinerary described by Bruner, 1883, who was given as collector by Scudder, 
1897b, p. 166) (ANSP). 

Melanoplus excelsus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 19, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 132, 166-167, pl. 11, fig. 9, 1897 
(males, females, above timber, 11,000 to 13,000 ft., Mt. Lincoln, Park Co., 
Colo.). Lectotype designated by Rehn and Hebard (1912, p. 79): Male 
(MCZ). 

Melanoplus alaskanus Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 20, 32, 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 132, 169-170, pl. 12, fig. 1, 1897 
(2¢°, 19, Alaska; Spilmacheen, British Columbia). Lectotype designated 
by Rehn and Hebard (1912, p. 79): Male from Alaska (USNM). 


Hebard (1929, p. 390) considered M4. defectus Scudder and M. di- 
midipennis Bruner synonyms of M. bruneri, but the types indicate 
that defectus is a subspecies of bilituratus, and dimidipennis a synonym 
of bilituratus bilituratus. 

Hebard (1928, p. 284; 1929, p. 390) indicated the synonymy of 
excelsus, which is borne out by the lectotype. The posterior surface 
of the subgenital plate is dimpled, but not vertically grooved nor 
incised at the apex, a condition included in the variation of bruneri, 
and the aedeagus is typical of bruneri. 

Walker (1910, 1920) indicated the synonymy of alaskanus, and the 
lectotypes of alaskanus and brunert demonstrate that synonymy. 
Earlier, Walker (1906) had recorded alaskanus from Vernon, British 
Columbia, with the identity questioned. In 1909 Walker discussed 
the variation shown by an Ontario series, as correlated with the 
differences in Scudder’s descriptions of the two supposed species 
(alaskanus and bruneri) and he then reduced alaskanus to a variety of 


A6 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


brunert. Pettit and McDaniel (1918) treated the Michigan population 
as adlaskanus. Essig (1926, p. 81) listed alaskanus as a synonym of 
bruneri. Hebard (1925a, p. 114) questioned the synonymy indicated 
by Walker, and later said (1928, pp. 281-282) he believed alaskanus 
to be a geographic race of mexicanus if nominal recognition is war- 
ranted. Rehn (1952) agreed with Hebard’s opinion. 

Because of the difference of opinion concerning alaskanus, it may be 
helpful to discuss the type in more detail. It is the specimen selected 
by Rehn and Hebard, and was received at the U. S. National Museum 
on Feb. 29, 1892, from T. C. Mendenhall of the U. S. Coast and Geo- 
detic Survey. Accession records show that the material was collected 
by J. E. McGrath during an expedition which was active along the 
Alaskan Boundary during September 1890-May 1891. McGrath 
(1893) has described the general experiences of the party which he 
led up the Yukon River, and (p. 185) has commented about the plenti- 
ful insects found near the river in summer. It appears that the type 
specimen was collected along the Yukon River somewhere between 
Ft. Yukon and the 141st Meridian. 

A male of bruneri, collected in 1948 near Fairbanks, Alaska, a 
locality at about the same latitude as the intersection of the Yukon 
River and 141st Meridian, confirms the presence of this species in 
Alaska, so there is no reason to doubt the source of the alaskanus 
lectotype. Hebard did not examine the aedeagus of the lectotype and 
so did not have this evidence which shows agreement with brunert. 

Although bruneri, excelsus, and alaskanus were all described in the 
same publication, the action of the first revisers (Hebard, 1928, for 
excelsus; Walker, 1910, for alaskanus) establishes the precedent of 
using bruneri as the valid name of this species. It is much the best 
known of the three names, and has page priority in Scudder (1897b). 
Though not required, page priority was recommended in the 1953 
Copenhagen Decisions of Zoological Nomenclature when other con- 
ditions are equal. 

Descriptive Notes: A usually medium-sized, fully winged member of 
the mexicanus group, most closely related to bilituratus, from which it 
differs mainly in male genitalia. Mesosternal hump variable (fig. 
1,9), averaging fully as prominent in male as in bilituratus, absent in 
female; tegmina usually reaching slightly beyond tip of abdomen, 
sometimes short of it. Females not always separable from bilituratus, 
but in any given locality where both species occur brunert females 
usually more robust and often shorter-winged; males usually recog- 
nizable by cercus and subgenital plate, but in doubtful cases the dorsal 
valve of aedeagus of bruneri is highly distinctive in its much more 
expansive and thin, striated, conformation. 

MALE GENITALIA: Cercus (fig. 7,7) variable, usually about as in 


GRASSHOPPERS—GURNEY AND BROOKS 47 


figure 7,i-2 more incurved along dorsal margin and more evenly 
rounded along ventroposterior margin than frequent in bilituratus; 
supra-anal plate (fig. 9,¢@) elongate, lateral elevations prominent; 
furculae usually reaching to middle of supra-anal plate or approxi- 
mately so; subgenital plate in lateral view with apical portion strongly 
upturned, sometimes rather abruptly so (fig. 16,4), often superficially 
accentuated by upward curvature of abdomen; posterior view of sub- 
genital plate usually as in figures 11,a-1 or 11,a-4, with apical margin 
entire and posterior surface grooved, frequently apical margin is 
dimpled (fig. 11,a-2), or there are well formed twin apices (fig. 11,a-3), 
and rarely margin is squarish and irregular (fig. 11,¢-5). Aedeagus 
with main stem relatively short; dorsal valve moderately sclerotized, 
thin, often appearing almost parchment-like, large and marked with 
striations; in dorsal view a triangular projection (x in fig. 3,k) of the 
well-sclerotized dorsal rim of the dorsal valve is distinctive; accessory 
lobe large for group; epiphallus as in figure 10,0. 

FEMALE GENITALIA: Cercus (fig. 10,k) relatively short and blunt, 
the dorsal margin often somewhat irregular; ovipositor with ‘‘scoop”’ 
of dorsal valve (fig. 10,7) strongly concave. 

Variation: The size of nine representative males, measured in 
millimeters, varies in pronotal length from 3.6 to 5.1 (av. 4.6), in 
length of hind femur from 9.0 to 13.0 (av. 11.9), and in length of 
tegmen from 12.0 to 19.5 (av. 16.6). Seven measured females vary 
in pronotal length from 3.6 to 6.0 (av. 4.8), in length of hind femur 
from 9.6 to 16.0 (av. 13.1), and in length of tegmen from 11.5 to 20.7 
(av. 17.1). The smallest specimens seen are from Mt. Moriah, Nev., 
the largest from Michigan. 

Cotoration: General coloration about as in bilituratus, varying 
from pale brown to blackish; hind femur usually with three external 
and dorsal dark bands, poorly developed in pale individuals; hind 
tibia varying from pale greenish yellow to red, usually pinkish to red. 

DistRiIBuTION: Several records of bruneri add significantly to the 
distribution based on male specimens examined (figs. 12, 13). Fulton 
(1930) reported it from Grant’s Pass, Oreg.; Hebard (1925a) listed 
Nipigon and Dwight (Ontario), Norrie and Oregon (Wisconsin), 
Gordon (Nebraska), and Englewood (South Dakota); Hebard (1932b) 
reported a specimen from Onawa, Iowa; and Hebard (1935a, p. 68) 
said that Beulah and Agua Fria Park are eastern limits in New 
Mexico. Stehr (1936) gave several Minnesota records which supple- 
mented those of Hebard (1932b; 1934a). 

BioLoGy AND ECONOMIC IMPORTANCE: On the high montane ranges 
of the Northwestern States and western Canada, bruneri is injurious 
to grazing areas. There have been fairly frequent references to the 
importance of bruneri in western Canada (Canad. Insect Pest Rev., 


48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


vol. 23, p. 2, 1945; vol. 24, p. 348, 1946; vol. 28, p. 255, 1950; vol. 33, 
pp. 10, 341, 1955). Treherne and Buckell (1924, p. 26) described it 
and Bradynotes chilcotinae Hebard (now Buckellacris chilcotinae 
chilcotinae) as the most common and injurious grasshoppers “‘in the 
lightly timbered areas of the summer range in the Chilcotin section 
(of B. C.) during 1920 and 1921.” In 1921 A. G. Ruggles reported 
to the Insect Pest Survey, U. S. Department of Agriculture, that 
damage by bruneri was occurring at Grand Rapids, Minn. The 
frequency of its capture on upland ranges in Montana suggests its 
importance there, as do comments on its abundance by Pepper, 
et al. (1953). 

An unusual infestation of bruneri occurred on Mt. Moriah, Nev., in 
the 1930’s. Material collected Sept. 2, 1937, and Aug. 16, 1939, is in 
the U. S. National Museum, and data on the infestation have been 
obtained from Lee M. Burge, Division of Plant Industry, Nevada 
Department of Agriculture. Mt. Moriah is located about 35 miles 
east of Ely, Nev., just north of Baker. The infested range is a large 
table area at an elevation of about 11,000 feet, bearing a valuable 
stand of bunch grasses and grazed over by sheep. Injurious popula- 
tions of bruneri existed there in 1936-1938. During the first 5 days 
of September 1937, poison bait was carried up to the sheep range by 
pack mules and spread by hand. Later in 1937 a survey indicated 
that approximately 2,000 acres carried a population of 20 adults per 
square yard. A visit made by Nevada entomologists during 1938 
coincided with a hail storm, and it was estimated that half of the 
large population then existing was killed by the hail and cold. 

No detailed food-plant studies of bruneri have been reported, but 
the general ecology is rather well known. Morse (1908) described 
the species (as alaskanus) as the dominant Melanoplus on Isle Royale, 
Mich., and recorded it from clearings, along trails, rock ridges, the 


EXPLANATION OF Ficure 11 
Posterior view of male subgenital plate (except ¢(4), which is ventroposterior view). 


a: Melanoplus bruneri: 1, 2 from Ft. McLeod, Alberta, Canada (2 specimens); 3, 4 from 
Mt. Moriah, Nev. (2 specimens); 5, from Crook Co., Wyo. 

b: M. borealis stupefactus: 1, lectotype; 2, 4, 5, 6, from Mineral Co., Colo. (4 specimens); 
3, from Los Pinos, Colo.; 7, from Silverton, Colo. 

c: M. borealis borealis: 1, from Nain, Labrador, Canada; 2, 3, from Cummington, Mass. 
(2 specimens). 

d: M. borealis palaceus: 1, 2, 4, from Lakeview area, Mont. (3 specimens); 3, topotype; 
5, holotype (x, rectangular vertical area). 

e: M. borealis utahensis: 1, holotype; 2, from Big Cottonwood Canyon, Utah; 3, from 
Ogden, Utah; 4, ventroposterior view of same specimen as 3. 

‘: Intermediate between M. borealis utahensis and M. borealis palaceus, 3 miles north of 
Mantua, Utah. 


GRASSHOPPERS—GURNEY AND BROOKS 


Ficure 11.—Explanation on facing page. 


477119 —59——__4 


49 


50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


‘ } 9 ? 
rs \ / 
—& y - \ 
Pye Sia 
VI 


Ficure 12.—Distribution of male specimens of Melanoplus bruneri examined from Alaska 
and Canada. 


beach heath zone, and other environments. Hebard (1925a, p. 114) 
stated that in the United States bruneri is strictly sylvan, living in 
brushy forest undergrowth, and that this contrasts with the preferred 
habitat in the Chilcotin District, British Columbia. In the latter 
area Treherne and Buckell reported that bruneri normally lives 
“among rank-growing grasses and plants growing under the aspen 
poplar groves which often form belts separating the open range from 
the timbered hills,” also that it sometimes advances into open glades 
and small mountain meadows where vegetation is thick and green. 
Probably on the basis of these observations, Buckell (1921), in a 
summary of ecological preferences of Chilcotin Orthoptera, classified 
brunert as a semi-sylvan species which extends into the geophilous 
campestrian area where the open range adjoins the forest. The 
described Chilcotin habitat seems comparable to the Lake Upsilon, 
N. Dak., habitat where Hubbell (1922) found bruneri “quite common 
in a dry clearing covered with low bushes and shrubs of various kinds 
(Corylus rostrata, young aspens, willows, birches, etc.) and tall her- 
baceous plants such as fireweed and goldenrod, interspersed with 
small, grassy areas.” In the prairie provinces of Canada the habits 
of bruneri are similar to those reported by Buckell (1921). In the 
Cypress Hills plateau of southeastern Alberta, as well as in the foot- 
hills of western Alberta, where the open grasslands are replaced rather 
suddenly by coarse plants, low shrubs, and aspen poplar leading to 
the evergreen forest, bruneri just as suddenly replaces bilituratus as 
the dominant species. Similarly, grassy clearings within the forest 
margin in the Prince Albert-Whitefox area of Saskatchewan are 


51 


GRASSHOPPERS—GURNEY AND BROOKS 


“SaTPEOO] saqvIg peru) wWory pouruexs tusundq Ssnpdouvjayy Jo suauutoads aye Jo uornqiysiq—¢] AUNTY 


Br 


52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


populated mostly by bilituratus while the margins of those clearings 
support bruneri. Hebard (1930) remarked on the very different 
habitat preferences of bruneri and bilituratus (mexicanus), and this 
distinction was also brought out by Handford (1946, p. 172). The 
‘rather hot and dry valley fields” where Fulton (1930) found brunerz 
in Oregon probably were adjacent to sylvan conditions. Specimens 
of bruneri (10 o’, 9 9) in the Philadelphia collections, from Red Banks 
in Logan Canyon, Utah, apparently are those referred to by Hebard 
(1936b, p. 173) as M. utahensis, said to be abundant in areas of 
“cow cabbage” (Wyethia amplexicaulis Nutt.) 

Nymphs of bruneri have been illustrated by Handford (1946), who 
found that they resembled darker nymphs, at least, of dbilituratus 
bilituratus (~mezxicanus) in every respect. 

In northern Michigan adults of brunert are not abundant until the 
beginning of August (Hebard, 1909). It has been suggested (Canad. 
Insect Pest Rev., vol. 33, p. 10, 1955) that since the 1954 outbreak 
in parts of British Columbia seemed related to one in 1952, while the 
1953 infestation was light, the life cycle there may occupy two years. 
Collection dates on the great majority of specimens examined are in 
August and late July. Some of the more limital dates are as follows: 
Francis Lake, British Columbia, July 5; Castleton, Utah, July 5; 
Chilcotin, British Columbia, Aug .1; 37 miles southeast of Fairbanks, 
Alaska, Sept. 1; Drain, Oreg., Sept. 11; Therma, N. Mex., Sept. 12; 
Gallatin Canyon, Mont., Oct. 3. 

Some of the altitudes (in feet) on labels of specimens examined are 
as follows: Evergreen, Idaho, 3,600; Strawberry Mts., Oreg., 4,450; 
McCall, Idaho, 5,000-5,050; Jackson Hole, Wyo., 6,500; Fishers Peak, 
Colo., 7,000-8,000; Cascade Canyon, Teton National Park, Wyo., 
9,000; southwest base of Delano Peak, Utah, 10,000; Mt. Wheeler, 
Ney., 10,000-10,600; Mt. Lincoln, Colo., 11,000—13,000. 


Melancoplus spretus (Walsh) 
Ficures 1,f; 2,n; 3,a; 7,6; 10,a,v; 14; 15,a,d; 16,a; Plate 2,p 


Caloptenus spretus Walsh, Pract. Ent., vol. 2, p. 1, 1866 (Highland, Kansas). 
Neotype here designated: Male labeled ‘‘Lawrence, Kans. June 18, 1877, 
Geo. F. Gaumer’”’ (USNM). 

Caloptenus spretus caeruleipes Cockerell, Entomologist, vol. 22, p. 127, 1889 
(probably Glencoe, Nebr., where observations by Dodge (1878) presumably 
were made). 

The condition of the neotype is excellent, with colors well preserved 
and all appendages present. The aedeagus and epiphallus have been 
exposed as dry preparations still attached to the body (see figs. 
3,a-2,3; 10,2). The subgenital plate and cercus are shown in figures 
16,a-2 and 7,b-1. Measurements (in millimeters) are as follows: 


GRASSHOPPERS—GURNEY AND BROOKS a} 


Length of body, 26.5; tegmen, 23.0; hind femur, 12.5; extent of 
tegmen posterior to apex of hind femur, 9.0; pronotum, 4.6. The 
E/F ratio (length of tegmen divided by length of hind femur; see 
comments on p. 6) is 1.840. The general coloration is light brown, 
the head with gena rather pale and postocular dark bar distinct; 
antenna reddish brown; dorsum of occiput with broken, indistinct 
longitudinal dark streaks; tegmen with dark flecks in basal half only; 
hind femur with three transverse blackish bars (exclusive of knees) 
primarily on mesal half of dorsal surface; hind tibia and tarsus pale 
red, spines, claws, and apical half of spurs black. 

Five additional males and one female examined bear the same data 
as the neotype. The locality where the neotype was collected, 
Lawrence (Douglas Co.), Kans., is about 65 miles south of the type 
locality, Highland (Doniphan Co.), Kans. George F. Gaumer, the 
collector of the neotype, was a special assistant to the U.S. Entomo- . 
logical Commission who worked in Kansas in 1877. He has reported 
(Gaumer, 1878) that on June 16, 1877, he observed locusts flying over 
the Wakarusa River (which is about 3 miles south of Lawrence). He 
observed them on June 18 also, and, though a precise locality is not 
described, it can be inferred that the same general locality was 
involved. The locusts were seen flying in a north-northeast direction, 
and at 5 p.m. many specimens dropped to the ground. This informa- 
tion published by Gaumer explains the probable background of the 
historic specimen chosen as neotype. 

Walsh (1866) did not give a formal description of spretus, though he 
gave various characters and compared it to M. femur-rubrum, and he 
attributed the specific name to Uhler, who Walsh said had given the 
locust the name Caloptenus spretus without describing it. Since no 
published proposal of the name by Uhler was made, the name dates 
from Walsh. Walsh had specimens which were sent to him by 
Prof. W.S. Robertson of the Indian Orphan Institute, Highland, Kans. 
Since he did not consider that he was describing spretus as new, pos- 
sibly specimens were not preserved, at any rate no such specimens 
nor further references to them have been found, in spite of an examina- 
tion of the principal historic collections of Melanoplus. In case 
specimens were preserved in the Walsh collection, they probably 
were destroyed in 1871, when the Walsh material was burned in the 
great Chicago fire (Burks, 1953, p. 16). Since no type is known to 
exist, in spite of efforts to locate one, selection of a neotype is advisable. 
This is especially true because spretus is of great historic importance. 
It has not been collected recently, and old specimens are scattered in 
many collections, and eventually the number of good examples may 
become scarce. 

The name caeruleipes was proposed by Cockerell (1889) for the 


VOL. 110 


OF “SOT 


PROCEEDINGS OF THE NATIONAL MUSEUM 


54 


(I6BI)Aajry wor 
Pajdopo snjaids 
JO uol\Ngiaysig 


x 


OTT 


uoibay Asosodway 
(coioy juaudwuadqns —..— 
uoibay juauowdag | " 
snjaids we wt 


JO,OISOABP ‘W + ‘Z| ‘BI4 


area ay! 


GRASSHOPPERS—GURNEY AND BROOKS 55 


variety “‘of spretus with blue hind tibiae.”” He referred to material 
reported by Dodge (1878), but no types are known to exist, and 
probably none was set aside. It was previously reported (Riley, et al., 
1878, p. 47) that some specimens of spretus had blue hind tibiae, and 
it appears clear that no nominal significance should be attached to 
such individuals. Scudder (1897b, p. 185) placed caeruleipes Cockerell 
in the synonymy of spretus, and Hebard (1925a, p. 113) referred to 
the synonymy. 

Melanoplus spretis—originally described in Acridiuwm and credited 
to Uhler by Thomas (1865) but in reality described by Thomas him- 
self because Uhler had published no description—is of uncertain 
identity and the source of some confusion in the literature. No 
type specimens of spretis were mentioned by Frison (1927) or Hebard 
(1927). Riley, et al. (1878, pp. 43-44), stated that there must have 
been some mistake in the description, and that the original speci- 
mens of spretis were destroyed soon after the paper was written. 
The name spretis occasionally has been used in the sense of the 
Rocky Mountain grasshopper, as when Rehn (1900) recorded it 
from Mexico, following which Hebard (1917; 1932a, p. 289) pointed 
out the misidentification of mezxicanus. Scudder (1897b, p. 185) 
treated spretis Thomas as a synonym of spretus Walsh, and Hebard 
(1934b, p. 197) did likewise. Earlier, however, Hebard (1925a, p. 113) 
considered spretis a synonym of atlanis Riley. He appears to have 
overlooked the fact that spretis antedated atlanis by 11 years! 
Thomas (1865) stated that spretis was quite common in Illinois, 
which we believe was untrue of spretus, and it would seem that 
Melanoplus bilituratus was involved. The measurements given by 
Thomas apply fully as well to spretus as to Illinois specimens of 
bilituratus, however. Thomas (1878) credited spretus (sic!) to him- 
self and referred to the Illinois State Agricultural Report. Because 
of uncertainties over the identity of spretis, and the confusion with 
spretus if both names were to be retained for distinct populations, it 
is best to treat Acridium spretis Thomas 1865 as unrecognizable. 

Seudder (1878b) was the first to assign spretus to the genus 
Melanoplus. 

Descriptive notes: A large, long-winged member of the mezi- 
canus group (pl. 2,p). Head with fastigium comparatively far in 
advance of compound eyes; the dorsal carinae of vertex less prom- 
inent and farther apart than in mezicanus; eyes less rounded and 
bulging than in mezicanus; especially in male; pronotum (fig. 15,@) 
with strangulate narrowing of sides in front of principal transverse 
sulcus, this conspicuous on lower portions of lateral lobes; in lateral 
view the principal sulcus lower than remainder of median carina; 
mesosternal hump well developed in male, absent in female. 


56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


rE 


se are SE ae ste cor ary 
f bs Laat apace? 


wel Kas eal 
Se 


Water fet ae 


Ficure 15.—a, }, Dorsal view of pronotum, with grid to show extent of constriction: a, M. 
Spretus, male, Pueblo, Colo.; b, M. bilituratus oulturnus, Wellington, Kans. c, d, Lateral 
view of: c, M. bilituratus oulturnus, Wellington, Kans.; d, M. spretus, male, Pueblo, Colo. 


GRASSHOPPERS—GURNEY AND BROOKS 5% 


Mate GeEniTatia: Cercus (fig. 7,5) broad for group, the apex 
oblique at ventral side; subgenital plate (fig. 16,@) strongly upturned, 
twin apices (posterior view) well developed; aedeagus (figs. 2,n; 3,a) 
with dorsal valve extending laterally more than anteriorly, and re- 
curved dorsally and posteriorly, conspicuously concave on posterior 
surface, the apex of valve (dorsally) membranous; accessory lobe 
comparatively elongate; apex of aedeagus dorsally produced beyond 
dorsal valve. 

FEMALE genitalia: Cercus slightly more robust and blunt than in 
mexicanus; dorsal valve of ovipositor with “scoop” well developed, 
the dorsal shoulder at base angular. 

CotoraTIon: General coloration relatively dark for group. Hind 
femur with four dark transverse dorsal bands (including knee), inner 
paginal area rich yellowish, occasionally tinged with pinkish, outer 
paginal area yellowish to light brown, paler along ventral margin, 
ventral surface yellowish, the outerhalf tinged with pink; hind tibia 
usually red; tegmen with conspicuous blackish flecks; wing trans- 
parent, with faint tinges of yellowish near base. (Brett, 1947, p. 33, 
reported that of 100 specimens of spretus he examined in the Bruner 
Collection, University of Nebraska, 85 percent had red hind tibiae 
and 15 percent glaucous or pale hind tibiae.) 

Variation: The size of 14 representative males, measured in milli- 
meters, varies in pronotal length from 4.2 to 5.5 (av. 4.8), in hind 
femur from 11.5 to 14.2 (av. 13.0), and in tegmen from 19.0 to 26.0 
(av. 22.8). Seven measured females vary in pronotal length from 
4.8 to 5.4 (av. 5.1), in hind femur from 13.2 to 15.2 (av. 14.2), and 
in tegmen from 23.0 to 25.5 (av. 24.6). No correlation between size 
differences and geographic distribution has been recognized. 

The extent to which the tegmen extends posteriorly beyond the 
apex of hind femur varies from 4.5 to 10 mm., with an average of 
about 7 mm. for males examined. For females, extremes of 5, and 
10, and an average of about 8 mm. occur. 

Measurements by Riley, et al. (1878, pp. 47-49), are not compa- 
rable to the present ones because the former concern the distances 
that various body structures exceed others. Faure’s (1933) measure- 
ments are expressed in averages only. His averages for length of 
tegmen are higher than ours, perhaps because more distance at the 
very base of the tegmen was included through not using the method 
later recommended by the International’ Locust Conference (see p. 6 
of the present paper). 

Of the males seen, there is none with pale hind tibiae which may 
not owe the lack of pigmentation to immersion in fluid. The hind 
tibiae of two or three of the. females seem to be pale from original 
coloration, though fading is suggested by some of the oldest specimens. 


58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Furculae vary from straight to markedly divergent. Very little 
variation in the male cercus has been seen. The aedeagus seldom 
varies much, except in appearance due to shriveling from drying. The 
apical portion of the main stem shrivels considerably and often is 
smaller and somewhat distorted when dry. The apical valve usually 
is heavily and darkly sclerotized, extending decidedly more laterally 
and posteriorly, like “horns,” in dry than wet preparations. In wet 
preparations the apical membrane of the dorsal valve is much more 
noticeable, decreasing the hornlike appearance. The dorsal valve 
occasionally is well anterior of the apex of the main stem, but fre- 
quently, especially in dry preparations, extends laterad of the apex. 
The apex of the ventral valve sometimes extends out from the main 
stem, but usually is tightly appressed to the anteriobasal portion of 
the aedeagus. The accessory lobe usually shrivels and has little dis- 
tinctive shape following drying. 

DistriputTion: The map (fig. 14) shows the distribution of males 
examined which have precise locality data. Males with only State 
labels are from Montana and New Mexico, as well as from eight other 
States and Provinces from which there are precise data. Females 
examined which add significantly to the distribution shown by males 
are from Taos Peak, N. Mex., only. The Permanent, Subpermanent, 
and Temporary Regions (fig. 14) in the sense of the U. S. Entomolog- 
ical Commission, are adapted from Riley (1891). Riley (1877) and 
Riley, et al. (1880), also mapped the distribution of spretus. It was 
frankly stated by Riley and his associates that various details as 
presented on their maps were based on fragmentary data and so should 
be taken as tentative only. Many early records of spretus are un- 
supported by specimens, so there is no way of checking identifications. 
It seems possible that some early Nevada records may be attributable 


EXPLANATION OF FicurE 16 
Subgenital plate of male, in lateral view. 


Melanoplus spretus: 1, from Argentine Pass, Colo., July 16, 1877; 2, neotype. 

M. bilituratus defectus, Casa Grande, Ariz. 

M. borealis stupefactus, Los Pinos, Colo. 

M. borealis borealis: 1, from Cummington, Mass.; 2, from Churchill, Manitoba, Canada; 

3, from Pikes Peak, Colo. 

e: M. borealis palaceous: 1, holotype; 2, topotype; 3, from Lakeview area, Mont.; 4, from 
Cedar Breaks, Utah. 

Ff: M. borealis utahensis: 1, holotype; 2, from Ogden, Utah; 3, from Big Cottonwood 
Canyon, Utah. 

g: Intermediate between M. borealis utahensis and M. b. palaceus: 1, 2, from 3 miles north 
of Mantua, Utah (2 specimens). 

h: M. bruneri: 1, from Ft. McLeod, Alberta, Canada; 2, from Chilcotin, British Columbia, 

Canada; 3, from Drain, Oregon; 4, from Mt. Moriah, Nev. 


LR SS 


GRASSHOPPERS—GURNEY AND BROOKS 59 


ees, 
Some. 


IOC 
awae, 
AS 
RIS Aw 


Ficure 16.—Explanation on facing page. 


60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


to the Nevada sage grasshopper, Melanoplus rugglesi Gurney. For 
instance, Riley, et al. (1880, p. 7), included the following paragraph: 


This year [1878], as well as 1877, was a locust year in this sparsely settled 
territory. July 11 great numbers appeared at Winnemucca, and disappeared the 
13th; on the 18th and 19th swarms again appeared. (United States Weather 
Report). 

Since the season of the year noted above is normal for rugglesi, and 
Winnemucca is in the same area through which swarms of rugglesi 
passed in the late 1940’s, also since rugglesi could have been mistaken 
for spretus by local observers, it seems quite possible that rugglesi is 
the species responsible for the report quoted. 

Specimens of Melanoplus spretus examined, with full locality label 
data except State names: 


Canapa: Rosebank, Manitoba, (1 @, 1 9);!Aweme, Manitoba, July {19, 1902, 
N. Criddle (1 @, 1 2); Fairfax, July, J. Fletcher (1 &#, 1 2);* Boissevain, 
Manitoba, Aug. 26, 1879 (1 @); Delorain (Manitoba), July 8, 1898 (1 #, 1 9); 
Manitoba, Aug. 28, 1877 (1 @); Arctic America, R. Kennicott, S. H. Scudder 
cn 

Minnesota: Glyndon, Aug. 1877 (1 9); Glyndon, Clay Co., July 1, 1875 
(1 2); Minneapolis, Aug. 16, 1877 (2 @); ‘‘Minnesota,” Oct. 6, 1876 (38 ¢,6 9). 

Norty Daxorta: Bismark, July 23, 1877, Geo. W. Sweet (4 o', 1 2); Clifford, 
Traill Co., July 25, 1891 (2 &); Larimore, July 29, 1901 (1 &); Turtle Mts.® 
(1 #, 2 9); “N. Dakota,”’ July, 1900 (1 ¢&). 

Montana: “Mont” (1 ¢, 19). 

Iowa: Ackley, Sept. 1, 1877 (2 @); Dallas Co., Aug., Allen (1 9); Harrison 
Co., Aug. 4, 1873 (1 co); Monona Co., Aug. 1873 (1 92); “Iowa,” June 1877 
(1 &). 

NEBRASKA: Lincoln, June 28, 1893, 1,150 ft., Bruner (1 @); Red Willow Post 
Office, Aug. 1874, John Byfield (1 &); ‘“‘Nebr.” 1868 (3 o, 1 92); “Nebr.” 
1874 (3 o&, 2 Q). 

Wromine: Alkali St(ation), 6,000 ft., July 27, 1877 (1 @, 1 9); Steele, “fiying 
from S. &S. W.,” July i879, C. A. H. McCauley (1 @); Steele, July 1878 (1 &); 
Steele, July 1879 (1 9); “Wyo. Terr.” (1 &). 

Ura: Salt Lake Valley, 4,300 ft., Aug. 1-4, 1877 (3 @); Salt Lake, Aug. 16 
(1 9); Salt Lake (1 9). 

Cotorapo: Argentine Pass, 13,000 ft., July 16, 1877 (2 =); Arapahoe Peak, 
11-12,000 ft., Packard (2 @); Boulder, July 19, 1877 (2 &); Colorado Springs, 
July 1878 (1 o), July 26, 1877 (2 ¢, 2 9), July 21, 1877 (1 9), Aug. 1, 1877 
(1 9), July 1877 (2 2); near Denver, L. Agassiz (1 @); Georgetown, 85—9,500 
ft., July 12-13, 1877 (1 &, 29); Mt. Lincoln, above timber, 11-13,000 ft., 
Aug. 138, 1877 (2 9); Pueblo, July 8-9, 1877 (1 &); White River, Aug. 6, 1877, 
Mrs. E. H. Danforth (1 9); “Colorado,” July 1877 (1 @); “‘Colorado,” H. V. 
Morrison (1 &). 

4 These specimens probably are part of the ‘‘enormous numbers” of spretus reported by Mitchener (1954, 
p. 50) to have been found at Fairfax, Manitoba, July 4, 1S01, by James Fletcher. 

5 Probably collected by Kennicott in the Winnipeg area of Manitoba, as discussed by Scudder (1897b, p. 
187), the Scudder name on the label probably referring only to the Scudder collection. 

‘These Turtle Mt. specimens probably sre from the same source as a large collection preserved at the 
University of Nebraska and reported by Brett (1947, p. 14) as collected by Lawrence Bruner in the early 
1880’s. O.S. Bare (in litt., Dec. 19, 1946) has informed us that M. S. Swenk once told him that the Turtle 
Mt. series was collected about 1890 


GRASSHOPPERS—GURNEY AND BROOKS 61 


New Mexico: Taos Pk., 13,000 ft. (1 9); ““N. Mexico,” July 21, 1877 (1 &), 
July 21, 1877 (1 9). 

Missouri: Sarcoxie, July 10, 1877 (2 9); Jackson Co., 1874 (2 9); Jasper 
Co., Nov. 12, 1876 (1 o', 1 9); Johnson Co., 1874 (1 @, 19); “N. Mo.,” 1868 
(1 &); “Central Mo.,” 1874 (1 9); Holt Co., 1868 (2 9). 

Kansas: Cawker City, 1874, N. B. Freeland (1 @, 19); Lawrence, June 18, 1877, 
Geo. F. Gaumer (6 o’, one the neotype, 1 9); Manhattan, Nov, 1876 (3 9), 
1874, W. C. Howard (2 @); Salina, Oct. 1877 (1 #@, 1 9); Douglas Co., Oct. 
1877, G. F. Gaumer (1 @, 1 9); ‘“‘Kans.,”’ 1874 (2 #, 2 2), 1876 (1 2), Nov. 
1876 (1 #, 2 9). 

Texas: Near Brenham, 1868, Thos. Affleck (1 9); Calvert, devouring kernels 
of corn, J. F. Moulton, Jr. (1 o@); Dallas, Boll? (1 &); Salado, May 1877, 
J. H. Myers (1 9); “Texas,” Belfrage 7 (1 6, 1 9); ‘‘Texas, common every- 
where, has been very destructive in this and adjoining counties, the past few 
years,” Belfrage (1 2); “‘Texas,” Pope, May 15 (1 9); “Texas” 1868 (1 9). 

Locality uncertain: An original Uhler specimen, probably sent to Scudder by 
Uhler, belonging to the Museum of Comparative Zoology, with a round pink 
disk, also the labels, ‘“‘Collection of P. R. Uhler’’; “S. H. Seudder Coll’’; ‘“‘Mel- 
anoplus spretus Scudder’s Type, 1895”; ‘‘Caloptenus spretus Uhler NRR mi- 
gratory” (1 o) (manuscript notes from the late H. S. Barber include a list of 
abbreviations given him by Uhler; NRR meant “North Red River’’; also see 
Scudder (1897b, p. 187)). 


BIoLoGy AND ECONOMIC IMPORTANCE: Prior to 1885 the literature 
dealing with the Rocky Mountain grasshopper probably was more 
extensive than that then available concerning any other single native 
American insect. Since there is no recently gathered information 
and it is impossible to sift facts from opinions in much of the volumin- 
ous early record, the biology of spretus will be reviewed briefly. The 
most important references on biology are those of Packard (1877), 
Riley (1876, 1877a, 1877b, 1891), and Riley, et al. (1878, 1880, 1883). 
There is a great deal of repetition in those reports, and Riley’s sum- 
mary (1891, pp. 9-26) is recommended. When Cyrus Thomas 
began the field study of spretus in 1869 (Riley, et al., 1880, p. 73), 
he believed from the first year’s observations that the species was 
essentially sedentary, but later he and the other commissioners 
decided that it was predominantly migratory. Areas inhabited were 
defined as follows (Riley, 1891, map; and elsewhere): Permanent 
Region, or native breeding grounds, where the species is always found 
in greater or less abundance; Subpermanent Region, which the species 
frequently invades, in which it can perpetuate itself for several 
years, but from which in time it disappears; Temporary Region, or 
that only periodically visited, and from which the species disappears 
within a year (see map, fig. 14). 

Riley, et al. (1878, 1880), supposed portions of Canada (then cus- 
tomarily referred to as Arctic America, or British America) to be an 


7G. W. Belfrage (1834-1882) and Jacob Boll (1828-1880), both well-known early collectors in Texas. Bel- 
frage collected most in Bosque and McLennan Counties. 


62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


important part of the Permanent Region, but it is not clear how 
important a part. Walker (1910, p. 336) summarized distributional 
findings of the U. S. Entomological Commission for Canada, so far as 
the ‘‘Regions” were concerned, but he added little new information. 
Criddle (1917), perhaps basing an opinion on observations during years 
long after the main outbreak period, said that ‘invasions of Canada” 
occurred only during dry seasons, the swarms arriving in July and 
August. Roe (1951), in his discussion of the buffalo, said that in the 
1870’s and 1880’s there was quite common ignorance of western 
Canada. It may have been easy to ascribe the source of something 
poorly understood (i. e., swarms of spretus) to a vast, distant region 
that was little known entomologically. 

Within the Permanent Region, spretus did not breed everywhere, 
but instead it did so in favorable spots. River bottoms, sunny slopes 
of uplands, and subalpine grassy areas among the mountains were 
considered to be favored egg-laying sites. After maturity was 
reached, flights within the Permanent Region were common. More 
devastating and consequently much more widely publicized, however, 
were the “invading swarms” which went primarily in a southerly 
or southeasterly direction (except when west of the Rocky Moun- 
tains). A day’s flight was estimated to be from 20 to 150 miles. 
Stops were usually made at night, but not always. During brief 
stops, or when settled in an area for egg-laying, the swarms ate nearly 
all crops and wild plants. Swarms might stop anywhere in the 
Temporary Region. Riley (1877b, p. 95) stated that in 1876 swarms 
left Montana about the middle of July and reached Texas by the 
end of September, thus covering 1,500 miles in 75 days, an average 
of 20 miles per day. 

Swarms developing from eggs laid in the Temporary Region usually 
migrated northward or northwestward. In the southern part of the 
infested area a partial second generation sometimes occurred, but 
spretus had essentially a single generation per year. The time of 
hatching, as a general rule, varied from the middle to end of March 
in Texas, to the middle of May or beginning of June in Montana. 
The young grasshoppers typically displayed gregarious habits soon 
after hatching and initial feeding near the hatching site. The nymphs 
were described as moving in “schools or armies,” generally ‘‘marching 
in a given direction until toward evening” (Riley, 1891, p. 21). 
Sometimes the nymphs migrated in a band a mile wide across the 
front (Riley, 1877b, p. 233). Usually, marching occurred only during 
the warm hours of the day. 

Melanoplus spretus was considered to be nearly omnivorous on both 
native and cultivated plants, with vegetables and cereals a “main- 
stay,”’ though at least in some instances sorghum and broom corn 


GRASSHOPPERS—GURNEY AND BROOKS 63 


were left untouched (Riley, et al. 1878, p. 251). No thorough analysis 
of preferences among the native food plants of the Permanent Region 
seems to have been made, leading Pfadt (1949, p. 39) to conclude 
that there is no information concerning the food-plants on which 
spretus originally developed. 

During the years when it was more or less prevalent, spretus varied 
considerably in its abundance and destructiveness. Riley (1891, p. 
10) gave a résumé of the relative abundance between 1818 and 1888. 
Bethune (1875) briefly described the early destructive outbreaks in 
Manitoba, and a recent summary is by Mitchener (1954). Some of 
the summaries of experience with grasshoppers in various States, such 
as that of Smith (1954) for Kansas and of Munro (1949) for North 
Dakota, have not always differentiated between various species, as 
of course it is impossible to do for many of the early years. Schle- 
becker (1953) has given perspective to the general subject of grass- 
hopper outbreaks, and has reviewed the period of great damage by 
spretus. The year 1866, when the Rocky Mountain grasshopper did 
great damage in Kansas, Nebraska, northwestern Texas, and also 
invaded western Missouri, is described as one of the first years in 
which it attracted national attention. The years of greatest abun- 
dance were 1873-1876, with 1874 the high point in destructiveness. 
In 1874, Colorado, Nebraska, and Kansas were overrun, and parts of 
Wyoming, the Dakotas, Minnesota, Iowa, Missouri, New Mexico, 
Oklahoma, and Texas were ravaged by swarms from Montana and 
Canada where they were abundant (Riley, 1891). As recorded by 
Riley, et al. (1880), and recalled by Bruner (1883), infestations 
dwindled in 1877. Widespread abundance of spretus never occurred 
again, though some injury in various years of the 1880’s is attributed 
to it, including an important local outbreak in Otter Tail Co., Minn., 
in 1888. None of the 1888 specimens is preserved, so far as known 
(Hebard, 1932b, p. 37). Riley, et al. (1883), mapped the reported 
hatching and flights of 1880 and 1881. Bruner (1893) reported 
spretus present in many parts of middle and eastern Nebraska in 1892, 
and quite common at Lincoln, Nebr. 

It is probable that the layers of spretus in Grasshopper Glacier, 
Park Co., Mont., were deposited during the major recorded swarming 
years for spretus, or in earlier years that were comparable, though not 
recorded. This glacier, located near the northeastern corner of 
Yellowstone National Park, has been discussed in detail by Gurney 
(1953). Flights of grasshoppers, probably aided by favorable air 
currents, occasionally pass over the glacier, even at its altitude of 
nearly 11,000 feet, and specimens landing on the snow are preserved 
when the necessary subsequent conditions of freezing and snowfall 
occur. Several species have been collected alive on the snow of the 


64 PROCEEDINGS OF THE NATIONAL MUSEUM YOL. 110 


glacier in recent years, but spretus is the only species identified from 
the remains imbedded in layers here and there in the glacier, and 
sometimes conspicuous on the vertical face at the foot. Preserved 
samples have long been supposed to be spretus, but during the period 
when spretus was considered a migratory phase of mezicanus, and 
before the critical features of the aedeagus were known, the samples 
were not identifiable other than as mexicanus (as then called). Study 
of the aedeagus of specimens now shows the samples in the U. S. 
National Museum to be spretus; unfortunately, this had not been 
done when the 1953 glacier paper was published. Prior to 1898, 
grasshoppers were known to be preserved in the glacier, and they were 
first examined and identified as spretus in 1914. The rather fragmen- 
tary samples now at the U. S. National Museum represent 13 speci- 
mens, from which two males taken in 1949 by Dr. J. R. Parker and 
one taken in 1952 by Dr. Irving Friedman have been dissected and 
found to be spretus. The dorsal valve of those males is well pre- 
served and typical. Fragments of grasshoppers collected from the 
central and bottom portions of the glacier in 1952 by Dr. Friedman, 
of the U. S. Geological Survey, were sent by him to W. F. Libby, 
then of the Institute for Nuclear Studies and Department of Chemis- 
try, University of Chicago, for age determination by the carbon—14 
method. Dr. Libby reported (in litt., Apr. 15, 1953) that the results 
indicated an age of 45 + 300 years and that the limits of error were 
45 + 600 years. He has since reported (Libby, 1954) the age as 
45 + 150 years. ‘The specimens are thus indicated as quite modern, 
and the likelihood is that the layers of preserved spretus are not more 
than 200 years old, perhaps much younger. 

Various writers at about the turn of the century remarked on the 
strange disappearance of spretus. Cooley’s (1904a) experience of not 
capturing a specimen during five years of collecting in all parts of 
Montana was typical. Except for the Grasshopper Glacier material, 
the only specimens collected since 1890 that we have examined are 
from Clifford, N. Dak., 1891; Lincoln, Nebr., 1893; Delorain, Mani- 
toba, 1898; ‘“North Dakota,” 1900; Larimore, N. Dak., 1901; Fairfax, 
Manitoba, apparently 1901; Aweme, Manitoba, 1902.8 Rehn and 
Hebard (1906, p. 408) and Hebard (1928, p. 283; 1929, p. 390) recorded 
three specimens (as spretus) which Hebard collected at Pike’s Peak 
and near Manitou, Colo., in 1904. The single male, from Pike’s Peak, 
on examination of the aedeagus, proves to be M. bilituratus bilituratus. 
Thus, spretus is not known to have existed since 1902. 

Why spretus disappeared and whether it is extinct are unknown. 
Riley, et al. (1883), included a long discussion by Lawrence Bruner, 


8 All these latest records are represented by males. Aweme, where Norman Criddle lived and worked 
for many years, is near Treesbank, and about 15 to 20 miles southwest of Carberry, Manitoba. 


GRASSHOPPERS—GURNEY AND BROOKS 65 


the foremost American field student of grasshoppers of his time, of 
factors of weather, natural enemies, and other influences thought to 
be related to spretus abundance, but the enigma remains. Cantrall 
and Young (1954) hinted at an influence by the American buffalo: 
“Tt is obvious that any disturbance in a habitat is accompanied by 
population fluctuations with the increase of some species and the 
decrease of others. The appearance of the migratory locust, Melano- 
plus mexicanus migratory phase spretus, following reduction of the 
great buffalo herds may be a classical example of the phenomenon.” 
We have examined the well-documented reports of Hornaday (1889) 
and Roe (1951) in an effort to determine whether there is a chrono- 
logical correlation between the disappearance of the buffalo and the 
abundance or disappearance of spretus. So far as the southern herd, 
centered in Kansas and adjacent States, is concerned, the last hunting 
on a large scale occurred during the winter of 1875-1876. In 1871, 
by which time much attention already had been attracted to the 
migration of spretus, millions of buffalo remained in Kansas, Okla- 
homa, and the Texas Panhandle. The northern herd, located mainly 
in Wyoming, Montana, and the Dakotas, was slaughtered later than 
the southern one. The years 1878-1883 were the biggest hunting 
years, with the last major hunt in the season (about October—Febru- 
ary) of 1882-1883. Therefore, in Montana, which was the heart of 
the Permanent Region for spretus, the peak of grasshopper outbreaks 
had passed (1877) before the buffalo herd was removed. After 
considering possible effects of the buffalo in feeding and trampling on 
the range, and the feeding by young nymphs on dry buffalo manure, 
it is difficult to see any relation other than near chronological coinci- 
dence between grasshopper and buffalo. 

Uvarov (1921) advanced the “phase theory” of locusts, which later 
became generally accepted as a fact, and Hebard (1925a, pp. 112, 113; 
1928, pp. 281-284) suggested that spretus might be the migratory 
phase of mexicanus. Faure (1933) concluded, following rearing experi- 
ments, that he had demonstrated that nymphs of bilituratus [mezi- 
canus] could change into spretus, thus establishing the phase explana- 
tion. However, the measurements of his laboratory-reared “‘spretus’’ 
are not fully typical of the historical specimens of spretus, and his 
demonstration was not conclusive (Hebard, 1936c, p. 48). Brett 
(1947) also believed that by regulating the food and laboratory 
environment of growing nymphs he had produced “spretus.”’ His 
material, samples of which have been deposited in the U. S. National 
Museum, shows considerable variation in wing length, but the aedeagi 
of all the males seen are ascribable to M. bilituratus vulturnus, and 
no approach to the aedeagus of spretus is shown. If spretus were a 


477119—59——_5 


66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


phase of bilituratus, it might be supposed that during the fairly ex- 
tensive flights of the 1930’s, as reported by Parker, et al. (1955), 
there would have been signs of spretus development. But such was 
not the case, and now, in the light of aedeagal characters here dis- 
cussed, it seems that bilituratus and spretus are distinct species. 
Nevertheless, for 25 years the view that spretus is the migratory phase 
of bilituratus [mexicanus] had considerable acceptance (Hebard, 1929, 
1931, 1936c; Gurney, 19538, p. 314, pl. 7, fig. p; Parker, et al., 1955). 

Brett (1947) felt that settlement of spretus breeding grounds for 
farming and grazing was responsible in some way for the disappearance 
of the grasshoppers. Impressed by short-winged specimens reared on 
alfalfa, he also stressed a possible correlation between the increase in 
alfalfa acreage and spretus decline. Pfadt (1949, p. 39) expressed 
doubt that alfalfa was a major factor and implied that the little 
available information concerning preferred native food plants of 
spretus is an inadequate basis for any general conclusions explaining 
the disappearance. We have failed to learn of any pronounced 
range-plant changes in Montana during the 1860’s, 1870’s, or 1880’s, 
so far as the general disappearance or replacement of plant species is 
concerned. (See Andrewartha and Birch (1954, esp. pp. 594-600) 
for a discussion of conditions related to population fluctuations.) 

The irregular periodicity of historic spretus swarms, the apparently 
nearly omnivorous appetite and very long-winged condition of 
swarming adults, and the marching behavior of nymphal bands 
suggest the occurrence of a migratory phase. If this view is correct, 
spretus probably had a solitary phase which occurred in relatively 
localized colonies, possibly adapted to a specialized environment. 
There is the example of Melanoplus rugglesi, which in the Great Basin 
attracted widespread attention for about 13 years (1939-1952) as a 
migratory phase. Previously, rugglest had been virtually unknown, 
and at present the solitary phase is scarcely found except by ento- 
mologists acquainted with the specialized habitat of the small localized 
colonies. A handicap in studying the possible correlation of weather 
conditions and spretus abundance is a shortage of reliable weather 
records, prior to 1886, for such Western States as Montana. 

Criddle (1917) discussed the former prevalence of spretus, and added: 
“At present, however, the insect seems to have vanished completely. 
Indeed, there are some who would place it with the Passenger Pigeon 
as an object of the past. It seems almost incomprehensible, however, 
that such can be the case. More probably the real permanent 
breeding grounds are more restricted than was supposed, and the 
locust will yet be located either by the discovery of its real haunts 
or by a new invasion following favorable weather conditions for 
breeding purposes.’”’ One immediately wonders whether a solitary 


GRASSHOPPERS—GURNEY AND BROOKS 67 


phase of spretus still exists, and if so what are the most likely breeding 
places. Montana, east of the Continental Divide, and North Dakota 
would seem likely possibilities. A solitary phase of spretus, if it 
exists, probably is smaller and shorter-winged than the _ historic 
specimens, and the aedeagus may be the only reliable distinguishing 
character. It is possible that specimens of a solitary phase now 
exist in collections, but if so they have not been recognized. Popula- 
tions of the mexicanus group, both preserved specimens and field 
samples, should have the aedeagus examined critically on the chance 
that new evidence bearing on the spretus problem will be disclosed. 


Melanoplus borealis (Fieber) 


The distribution of this species is indicated by the two maps of 
its subspecies (figs. 17, 18) and, for the United States, by Newton 
and Gurney (1956, map 56). As the specific name suggests, this is 
a boreal insect, which in the United States is primarily northern or 
occurs in mountainous areas. 

Mostly on the basis of tegminal length and the shape of the male 
subgenital plate, four subspecies are here recognized. Our study 
of the aedeagus has failed thus far to show differences in the aedeagus 
among the subspecies. However, Hebard (1935a, p. 67), in writing 
of what he called the borealis group, stated: “Examination of the penis 
now shows that this organ is of the same general type in all of the 
recognized races but shows distinct proportionate differences in some.” 
Some differences in the epiphallus are here reported, but the examina- 
tion of more material is required before their significance can be fully 
evaluated. 

Melanoplus borealis borealis (Fieber) 


FicurgEs 3,3; 7,3; 8,f; 9,c; 10,c,l,s; 11,c; 16,d; 17; 18; PLate 5,a,B,pD. 


Caloptenus borealis Fieber, Lotus, vol. 3, p. 120, 1853 (unstated number of 
specimens from ‘‘(Grénland) Nord. Cap. (Labrador) M. Hal. Fieb.’”’). Lecto- 
type here designated: Male from original Fieber series (according to Dr. 
Max Beier, in litt., Jan. 17, 1957) with the following labels: ‘‘Podisma 
borealis Fieb. Gronl.’’; “Mus. Caes. Vindebon.’’; ‘det. Holdhaus Melanoplus 
borealis Fieb.”’; ‘‘Par.”’; ‘23’? (Naturhistorisches Museum, Vienna). 

Pezotettix septentrionalis Saussure, Rev. Mag. Zool., p. 159, 1861 (unstated 
number of specimens from Labrador). Lectotype here designated: Female 
labeled “P. septentrionalis Sss. Type’’; ‘Labrador’; and with a lectotype 
label attached by Gurney (Museum d’Histoire Naturelle, Geneva). (The 
Saussure collection contains two series of septentrionalis, one consisting of 
lectotype, a second containing 17 specimens (15 from Labrador and 2 from 
“Hudson’’) (C. Ferriere, in litt., Jan. 23, 1957).) 

Caloptenus extremus Walker, Cat. Derm. Salt. B. M., Pt. 4, p. 681, 1870 (two 
females from “‘Arctic America”). Lectotype here designated: Female labeled 
Caloptenus extremus and as type, and with both hind legs present (British 
Museum). (The second original female lacks both hind legs. The lectotype 


68 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


also bears an accessions registration label, ‘51/49,’ which is associated 
with the following notebook data: “District watered by Mackenzie and 
Slave R. Presented by Sir John Richardson.” (D. R. Ragge, in litt., Jan. 
10, 25, 1957).) 

Pezotettix junius Dodge, Canad. Ent., vol. 8, pp. 9-10, 1876 (males, females, from 
Glencoe, Dodge Co., Nebr.). New synonymy. Lectotype here designated: 
Male labeled ‘‘Pez. Junius &G. M. Dodge Type; Collection C. V. Riley” 
(USNM). (A similarly labeled female is associated with the lectotype. 
Though no original locality label exists, Dodge stated that the species occurred 
at Glencoe, Nebr.) 

Caloptenus parvus Provancher, Nat. Canad., vol. 8, p. 110, 1876 (one male from 
Cap Rouge, Quebec [southern outskirts of Quebec City]). Lectotype here 
designated: The first of two males pinned in Provancher'’s first collection 
(Musée de la Province, Quebec). (N.—M. Comeau has reported (in litt., 
Jan. 8, 1957) that although a single specimen was originally mentioned, an 
1877 catalog mentioned two, and that both are now in very good condition.) 

Melanoplus extremus scandens Scudder, Proc. U.S. Nat. Mus., vol. 20, p. 289, pl. 1, fig. 
f, 1897 (unstated number of specimens from Big Horn Mountains, Wyo., 
and several localities in New England, as well as unstated localities ‘‘in the 
high north’). Lectotype here designated: Male labeled ‘‘White Mts. Alpine. 
Scudder, Drawn, Mel. extremus Scudder’s Type, 1895, Drawn for Ap- 
palachia, S. H. Scudder Coll., Type 15466” (MCZ). (Tegmina of lectotype 
extend 5 mm. behind the apex of abdomen. A female with similarly long 
tegmina bears the same data.) 

Melanoplus monticola Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 24, 
34, 1897; Proc. U. 8S. Nat. Mus., vol. 20, pp. 135, 290-291, pl. 19, fig. 5, 1897 
(two males, two females from ‘‘above timber line on Sierra Blanca, Colo- 
rado’”’). New synonymy. Lectotype designated by Rehn and Hebard 
(1912, p. 84): The figured male (MCZ). (Only fragments of the lectotype 
remain. The second original male is at Philadelphia (ANSP); its subgenital 
plate is about as in fig. 16,d-2, in lateral view, and in posterior view two prom- 
inences are developed, much as in fig. 11,b-4.) 


The following four specimens of borealis were loaned from Vienna 
by Dr. Beier in addition to the lectotype: One nymph labeled as from 
Greenland; a female labeled Labrador, 1870; a female from Okak 
Islands, Labrador, 1838; a female from Nain, Labrador, 1832. The 
last two specimens bear labels reading ‘‘Coll. Br. v. W. Labrador, 
Coll. Sommer.’”’ M. C. Sommer was a Danish handler and collector 
of insects who died in 1868, from whom Brunner is recorded as ob- 
taining Orthoptera, and it seems probable that these are the female 
specimens from Labrador recorded by Brunner (1861, p. 223) as 
Caloptenus borealis. At the same time he recorded material from 
Valdivia [Chile], the latter doubtless being a misidentification. 
Brunner’s specimens may not have been available to Fieber in 1853, 
hence the lectotypic selection of the male which does not provide a 
clear type locality, though it unmistakably is the species currently 
called Melanoplus borealis (aedeagus exposed, preserved in dry 
condition). 


GRASSHOPPERS—GURNEY AND BROOKS 69 


There is no record of borealis from Greenland since Fieber. Henriksen 
and Lundbeck (1917) and Henriksen (1939) did not report it, and 
S. L. Tuxen of Copenhagen, who is familiar with current work on 
Greenland, has assured us (in litt., Jan. 26, 1957) that the Fieber 
record must be incorrect. No Acrididae have been taken on Green- 
land by Danish entomologists who in summer collect there regularly. 

Likewise there is no subsequent record of borealis from North Cape 
or elsewhere in Norway (Ander, 1949a, 1949b; Knaben, 1943), 
although Scudder (1897b, p. 272) assumed that Fieber’s locality ‘‘Nord. 
Cap.” is the one in Norway. Since Fieber included borealis in a 
treatment of European Orthoptera, and did not enclose the locality 
in parentheses, as he did the two non-European localities, it is fair to 
assume that he regarded North Cape, Norway, as one of his localities. 
On the other hand, there is North Cape, Prince Edward Island, and 
Cape North, Nova Scotia, which may have been unknown to him 
though ports of call for North Atlantic shipping. There is a strong 
probability that none of Fieber’s specimens originated in either 
Norway or Greenland. Therefore, no definite type locality can be 
recognized; the lectotype probably came from Labrador or nearby. 
No information is available about any Fieber material of borealis 
which may have been deposited at Halle, Germany; the main Fieber 
collection is in Vienna. 

The synonymy of septentrionalis was indicated by Scudder (1897b, 
p. 270) and later by Blatchley (1920, p. 424). Blatchley also treated 
extremus and parvus as synonyms of borealis. Essig (1926, p. 81) 
incorrectly listed extremus as a synonym of brunert. Hebard (1925a, 
p. 111) treated extremus as a synonym of borealis borealis, and he also 
placed scandens as a synonym of borealis junius. Scudder (1897b, 
p. 287) synonymized parvus under eztremus. His interpretation 
recognized parvus as the northern form with red color prominent on 
the hind legs rather than the pale type as represented by the synony- 
mous junius. It is in agreement with Provancher’s description of 
parvus, which refers to the red color on both hind femur and hind 
tibia. Hebard (1925a, p. 111) may have overlooked this feature of 
parvus, and thus was incorrect in regarding parvus as identical with 
junus. Both junius and parvus were described in 1876,-in journal 
issues of January and April, respectively, though actual dates of 
mailing have not been ascertained by us. The dates and the identity 
of parvus are important if in the future the southeastern population 
should be restored to subspecific rank. 

Scudder utilized what he believed to be differences in the shape of 
the apical part of the male subgenital plate to separate borealis, 
extremus and monticola, but his differences are not constant in series 
of specimens. His scandens consisted of individuals mainly from 


70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


high altitudes and was characterized by tegmina reaching somewhat 
beyond the apices of the hind femora, while in gunius the tegmina 
usually did not quite reach those apices. Hebard (1930, p. 397) 
noted that monticola is separated from typical borealis largely by 
shorter wings and tegmina, but he indicated that some far northern 
specimens would be assigned to borealis monticola if that character 
was followed, and he clearly was impressed by the local variation 
occurring in the Northwest. For the most part, specimens from the 
northeastern United States and southeastern Canada have the lower 
surfaces of the hind femora yellowish, and the hind tibiae are yellowish 
or dirty pale brown, while specimens from further north and the 
Rocky Mountains usually have the lower femoral surfaces and the 
hind tibiae reddish. Series from Ontario and Manitoba show inter- 
mediate conditions, and as a whole the populations represented by 
qunius and monticola do not seem sufficiently defined for their recogni- 
tion to serve a useful purpose. Hebard (1929, p. 387) said that 
borealis monticola was the least well defined of four races of borealis 
which he recognized, and (1932b, p. 37) he stated that gunius is a 
“very weakly defined race.” Later (Hebard 1934a, p. 105), he noted 
the difficulty of separating borealis junius and borealis borealis in 
northern Minnesota and suggested that junius might be merely a 
pallid color phase developed in response to a more temperate environ- 
ment. White and Rock (1941) recorded both borealis borealis and 
borealis junius from Alberta, and later (1945) recorded borealis 
monticola also. In 1945 they discussed variation and the problem of 
trying to justify the recognition of these subspecies. Blatchley (1920) 
and Morse (1920) have discussed the variation shown by borealis. 
Scudder (1897b, pp. 270, 272) placed Caloptenus arcticus Walker in 
the synonymy of borealis, with some uncertainty, and Blatchley (1920) 
did so definitely. This is now considered untrue, as here discussed 
under M. biltturatus bilituratus (p. 15). 

In Canada the junior author has recognized two races or ecological 
forms within borealis borealis. Specimens from Labrador, James Bay, 
Northwest Territories, northern Quebec and Ontario, northern 
Manitoba, Saskatchewan and Alberta, and from British Columbia are 
heavier, darker, more discolored, with more evident spotting on the 
tegmina, and consistently higher tegmina/femur ratios in both long- 
and short-winged forms than the more southern counterparts. The 
form from the southern or grassland portion of the prairie provinces 
is smaller, generally pale brownish yellow, with tegminal spotting 
much reduced, and frequently with the hind femur entirely pale. 

A broad zone of transition between the two forms is evident in 
eastern Saskatchewan and in Manitoba, the smaller, paler form 


GRASSHOPPERS—GURNEY AND BROOKS 71 


occurring in southern (aspen-willow association) localities, the larger, 
darker form in the north (aspen-spruce association). Both forms 
contain long-winged and short-winged individuals, often together in 
one population; the short-winged type usually predominates but 
occasionally (as in the Cypress Hills in 1952) the long-winged type 
appears more numerous. 

DescrirTivE Notes: A small to medium-sized, fully winged member 
of the mexicanus group; closely related to borealis stupefactus, from 
which it differs primarily in wing length, and to borealis palaceus, 
from which it differs mainly in the less strongly upturned male sub- 
genital plate. Males with tegmina usually varying from slightly 
short of the apex of the abdomen to slightly beyond the abdomen, 
occasionally a little short of reaching supra-anal plate or at the other 
extreme with as much as one-fourth of the tegmen reaching beyond the 
abdomen. Females with tegmina varying from a condition in which 
about two-thirds of the abdomen is covered to one in which the 
tegmina extend beyond the abdomen by nearly one-fourth of their 
length. Posterior third of male abdomen occasionally strongly re- 
curved dorsally, especially in specimens from northern Rocky Moun- 
tains, but usually abdomen is horizontal or only moderately recurved. 

VariaTion: The size of 12 representative males measured (in 
millimeters) varies in pronotal length from 3.7 to 4.7 (av. 4.2), in 
length of hind femur from 9.7 to 12.2 (av. 10.8), and in length of teg- 
men from 8.4 to 16.4 (av. 12.6). Eight females vary in pronotal 
length from 4.1 to 5.5 (av. 4.8), in length of hind femur from 10.2 to 
12.7 (av. 11.4), and in length of tegmen from 11.0 to 16.6 (av. 13.0). 
The largest specimens are from the northwest, particularly the North- 
west Territories and Alaska. 

Mate GENITALIA: Cercus (fig. 7, 7) variable, usually about as in 
figure 7,j-1; supra-anal plate with lateral elevations strongly raised; 
furculae usually strongly tapered, separation evident to base, and 
scarcely reaching middle of supra-anal plate (fig. 9,c-1), sometimes 
variable (fig. 9,c-2, c-3); subgenital plate in lateral view with dorsal 
margin varying from essentially straight or slightly upturned apically 
(fig. 16,d-1) to moderately upturned, in posterior view with margin 
usually entire, rarely with an indication of twin apices (fig. 11,c-1); 
posterior surface of subgenital plate rather evenly rounded, without 
development of the rectangular-shaped disk usually characteristic of 
borealis palaceus, as in fig. 11,d-5. Aedeagus with main stem relatively 
short; dorsal valve moderately sclerotized but not parchment-like as 
in bruneri, in dorsal view lacking the triangular projection shown for 
bruneri (fig. 3,k,1); accessory lobe relatively large and frequently 
directed somewhat laterally; epiphallus as drawn (fig. 8,/). 


72 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


FEMALE GENITALIA: Cercus (fig. 10,/) averaging more slender, the 
dorsal ‘‘scoop” of the dorsal ovipositor valve less excavate than in 
bruneri, but females of the two species not always separable. 

Coxtoration: General coloration about as in bilituratus; hind femur 
usually unbanded except dorsally on mesal side where three dark 
transverse bands usually occur, external bands sometimes present, 
as in specimens from Hudson Bay area which have the pale areas along 
the dorsal surface of the hind femur noticeably yellowish in fresh 
specimens; ventral surface of hind femur and lateral surface ventrad of 
paginal area usually reddish except in specimens from the Plains 
States eastward and in southern Ontario and Quebec, reddish area of 
femur normally correlated with red hind tibia. 

Males of borealis borealis from the Big Horn Mountains, Wyo., and 
Fremont Co., Idaho, show a trend toward borealis palaceus, and they 
suggest intergradation although they are not fully intermediate. 
Males from Smithers, British Columbia, show a noticeable develop- 
ment of the dorsal lip at the apex of the subgenital plate. 

DistrispuTion: The principal recorded distribution of M. borealis 
borealis, additional to what is shown on our maps (figs. 17, 18), is that 
noted by Morse (1920) for Maine, and Hebard’s statement (1931, p. 
184) that borealis junius is known from Mountain Grove in the Ozark 
Mountains of south-central Missouri. J. W.H. Rehn (1939) recorded 
borealis from Grande Miquelon and St. Pierre, near Newfoundland. 

Male specimens of Melanoplus borealis borealis have been examined 
from the localities shown in figures 17 and 18. 

BioLOGY AND ECONOMIC IMPORTANCE: This subspecies is significant 
as a rangeland pest, and occasionally it attacks crops, but it is not 
of major importance. Information from the Bozeman, Mont., lab- 
oratory of the Entomology Research Division, U. S. Department of 
Agriculture, indicates that this is one of the important grasshoppers 
in the Targhee National Forest, Fremont Co., Idaho, and the Bridger 
National Forest, Wyo. It is reported to have been the dominant 
grasshopper in 1952 in the northern Cariboo and the Prince George 
Districts of British Columbia (Canad. Insect Pest Rev., vol. 30, 
p. 307, 1952; Neilson, 1953), as well as injuring alfalfa, grain, and 
vegetable gardens in northern Saskatchewan (Canad. Insect Pest 
Reyv., vol. 30, p. 150, 1952). 

Buckell (1921) characterized borealis in the Chilcotin District of 
British Columbia as an humicolous hygrophile, an ‘inhabitant of the 
tall rank vegetation beneath birch and willow on the borders of 
streams.” He found it a sluggish, richly colored insect. ‘Tolerance 
for a variety of habitats is shown by W. J. Brown’s experience (in 
litt., Mar. 1, 1949) of collecting it at Churchill, Manitoba, in areas 
supporting trees, though at Reindeer Depot he collected it on gently 


GRASSHOPPERS—GURNEY AND BROOKS 73 


Ficure 17.—Distribution of male specimans of Melanoplus borealis borealis examined from 
Alaska and Canada. 


rolling, treeless, Arctic tundra that consisted of damp, cold sphagnum 
which thawed only from 11 to 19 inches during summer. Plants 
present there included Labrador tea, blueberry, grasses, sedges, and 
others. Readers interested in the limits of the true Arctic Zone are 
referred to Kimble and Good (1955) and Munroe (1956). 

Caudell (1900) suspected borealis of being localized in Alaska, after 
the collector of the specimens he reported, T. Kincaid, found it in 
a sphagnum swamp at Kukak Bay, but nowhere else in the surround- 
ing countryside. The only habitat notes from Labrador which have 
been seen are those of Scudder (1897b, p. 272), who reported that 
a collector described finding borealis abundant in luxuriant plants 
beside a mountain brook. 

In the United States and southern Canada borealis has been found 
most often in cool, damp situations, as discussed by Blatchley (1920) 
and Morse (1920), but, while it appears to enjoy bogs and areas of 
sedges and other plants found in mountain meadows, it also occurs 
in moist open pastures and mowed fields in New England (Gurney 
1935). Cantrall (1943) considered it a characteristic inhabitant of 
“the leatherleaf stratum of the bog habitat” in southern Michigan. 
In the prairie provinces of Canada the subspecies inhabits tall rank 
vegetation on the borders of streams and ponds within a forested 
area. Similar situations in the open grasslands are not occupied. 
The distribution is therefore local in grassland areas. 

The biology of borealis borealis has not been studied fully. Nymphs 
are described by Handford (1946). It has been suggested that in 


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PROCEEDINGS OF THE NATIONAL MUSEUM 


74 


GRASSHOPPERS—GURNEY AND BROOKS Vi) 


parts of British Columbia two years are required for a complete life 
cycle (Canad. Insect Pest Rev., vol. 34, p. 273, 1956). Seasonally, 
it has been considered an early form by Morse (1920) and Cantrall 
(1943). The following are extreme dates on the labels of specimens 
examined: Warren Woods, 8S. Dak., May 28; Cummington, Mass., 
June 12—Aug. 16; Mt. Greylock, Mass., July 3, Aug. 17; Bloomington, 
Ind., June 16; Sawmill Bay, Northwest Territories, July 14; Chilkat 
Pass, British Columbia, July 13; Reindeer Depot, Northwest Terri- 
tories, Aug. 3, Aug. 16; Fairbanks, Alaska, July 4, Sept. 2; Nain, 
Labrador, Aug. 4; Hopedale, Labrador, Sept. 6; Big Horn Mts., 
Wyo., Aug. 15. Morse (1921) reported collections in Maine from 
June 6 to Aug. 25, and Alexander (1951, p. 109) reported that both 
nymphs and adults (as M. b. monticola) were taken in August from 
several high Colorado peaks. 

This subspecies is known from many mountainous areas in the 
southern part of its range, but it is not confined to high altitudes. 
Localities near sea level in Labrador, in the bogs at Mer Bleue,® 
Ontario, at about 1,400 feet in Cummington, Mass., and at various 
localities in Michigan, Minnesota, and Iowa show that fact. The 
following are some of the altitudes, shown on labels, at which it has 
been collected: Chilkat Pass, British Columbia, 3,500 ft.; Yoho Val- 
ley, British Columbia, 5,000 ft.; Yellowstone National Park, 6,800 
ft.; Pikes Peak, Colo., 12,000 ft. 


Melanoplus borealis stupefactus (Scudder) 
Ficures 8,d,h; 9,d; 10,d,m; 11,6; 16,c; 18; Puate 5,5,F. 


Pezotettiz stupefactus Scudder, Rep. Surv. W. of 100th Mer., App. JJ, p. 283, 
1876 (1 male, 3 females, nymphs, Taos Peak, Sangre de Cristo Mountains, 
N. Mex.). Lectotype designated by Hebard (1929, p. 387): Male (USNM). 

Melanoplus cockerelli Scudder, Psyche, vol. 9, pp. 124-125, 1900 (1 male, 1 female, 
summit of range between Pecos and Sapello Rivers, N. Mex., near head- 
waters of Pecos). Lectotype designated by Hebard (1929, p, 388): Male 
labeled ‘‘Top of range betw. Sapello and Pecos Rivers. Alt. 11,000 ft. 
Aug.” (MCZ). 

Melanoplus sapellanus Scudder, Psyche, vol. 9, p. 125, 1900 (1 male, 3 females, 
same locality as cockerelli). Lectotype designated by Hebard (1929, p. 388): 
Male (MCZ). 

Melanoplus latifercula Caudell, Proc. U. 8S. Nat. Mus., vol. 26, pp. 803-804, 
pl. 55, fig. 4, 4a, 1903 (1 male, Cumbres, Colo.) (USNM). 


Hebard (1929) indicated the synonymy here shown, which is sup- 
ported by our study of the four types. Scudder (1897b) placed 
stupefactus in the genus Podisma, and then when he described cocker- 
ell1 he failed to recognize stwpefactus and considered it a relative of 


§ A large area containing many sphagnum bogs, near the southeastern city limits of Ottawa. F. A. 
Urquhart and E. G. Munroe have informed us that Carlsbad Springs is near the eastern extremity of the 
bog area, and that Hawthorne is at the northwest corner. Auer (1930, fig. 1) shows Mer Bleue as some 
10 to 15 miles east of Ottawa. 


76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Melanoplus dawsoni (Scudder). His sapellanus, with data identical 
to those of cockerelli, was based largely on a male with imperfectly 
developed genitalia. 

Descriptive Notes: A short-winged, flightless subspecies averaging 
smaller than borealis borealis and with the furculae usually wider than 
in the latter. 'Tegmina broadly lanceolate, usually overlapping nar- 
rowly but sometimes not quite attingent, covering from scarcely more 
than one-third to slightly more than one-half of abdomen (about 1% 
to 1% times length of pronotum); wings vestigial. Posterior third of 
abdomen moderately to considerably recurved dorsally. 

MALE GENITALIA: Cercus (fig. 8,d) proportionately broader than in 
the other subspecies (distorted by drying in lectotype of stupefactus, 
fig. 8,d-1); furculae usually as in figure 9,d-2, seldom a little more 
tapering, and sometimes broader (fig. 9,d-1) ; subgenital plate in lateral 
view with posterior portion of dorsal margin upturned as a lip about 
to the extent in figure 16,c, not sinuate, posterior margin sometimes 
more nearly vertical and less oblique than in the figured specimen, in 
posterior view the dorsal margin variable (fig. 11,0) but usually with 
at least weakly indicated twin apices. Aedeagus as in borealis borealis; 
epiphallus with rather deep emargination of anterior margin between 
ancorae, lophi lanceolate in dorsal view, rather highly elevated and 
with profile of posterior portions evenly rounded in lateral view (fig. 
10,d). 

FEMALE GENITALIA: About as in borealis borealis, the cercus averag- 
ing more robust (fig. 10,m). 

Cororation: Typically dark for the species; hind tibia and ventral 
surface of hind femur, reaching slightly onto external paginal area, 
dark red; dark transverse bands of hind femur visible dorsally, 
variable on external surface. 

VARIATION: The size of eight representative males, measured in 
millimeters, varies in pronotal length from 3.6 to 4.5 (av. 4.0), in 
length of hind femur from 9.0 to 10.8 (ay. 9.7), and in length of tegmen 
from 4.7 to 7.0 (av. 5.9). Three measured females vary in pronotal 
length from 4.4 to 4.7 (av. 4.5), in length of hind femur from 11.2 to 
11.3 (av. 11.3), and in length of tegmen from 6.5 to 7.4 (av. 6.9). 

Hebard (1929, p. 389) reported 29 males, 27 females, and 2 nymphs 
from Tomboy, Colo., also a nymph from Trout Lake, San Miguel Co., 
Colo., which he regarded as intermediate between borealis monticola 
and borealis stupefactus. Specimens from the Tomboy series and a male 
from White River Forest, Colo., have the tegmina covering about 
three-fourths of the abdomen, the furculae averaging narrower than in 
typical borealis stupefactus, the dorsal lip of the apex of the male 
subgenital plate higher and without twin apices developed. These 
specimens average larger than borealis stupefactus, and have an average 


GRASSHOPPERS—GURNEY AND BROOKS V7 


tegminal length of 9.5 mm. in males, 10.8 mm. in females. We regard 

them as intermediate between b. borealis, 6. stupefactus, and 6. palaceus. 

Disrrisution: Hebard’s record from Gothic, Colo. (1929), is a 
northern extension of the distribution (fig. 18) based on specimens 
examined. 

Specimens of Melanoplus borealis stupefactus have been examined 
from the following localities: 

Cotorapo: Mineral Co.; Silverton, San Juan Co.; Lizard Head; peak north of 
Cumbres, Cumbres Range, Conejos Co.; Los Pinos, Conejos Co.; Cumbres. 

New Mexico: Taos Peak; Truchas Peak; Beulah; top of range between Sapello 
and Pecos Rivers; Santa Fe Baldy; head of Nambe Creek, Sangre de Cristo 
Range. 

The following material is intermediate between Melanoplus borealis 
stupefactus, M. b. borealis, and M. b. palaceus: 

Cotorapvo: Tomboy, Marshall Basin, San Juan Range (about 2 to 3 miles north- 
east of Telluride; A. B. G.), 11,400 ft., Sept. 2, 1921, Rehn and Hebard (12 0’, 
15 9); White River Forest, Aug. 1933 (1 0”). 

The male from Trout Lake, San Miguel Co., Colo., collected by 
Hebard Sept. 4, 1921, at 9,700 feet, and reported by him (1929, p. 
389) as intermediate between borealis monticola and 6b. stupefactus has 
tegmina covering slightly more than one-half of the abdomen. We 
regard it as 0. stupefactus. 

BroLoGy AND ECONOMIC IMPORTANCE: Most of the known biological 
data on this grasshopper have been summarized by Hebard (1929, pp. 
388-389; 1935a, p. 67). It is of no recognized economic importance, 
and there have been no detailed studies of its bionomics. Distribu- 
tion is mainly if not entirely in the Arctic Alpine and Hudsonian 
zones in northern New Mexico and southwestern Colorado. The 
lowest altitude at which it has been recorded is 8,500 feet at the head 
of Chupidero Creek, N. Mex., and the highest 12,629 feet at the 
summit of Santa Fe Baldy, N. Mex. 

Small numbers of specimens have been recorded well above timber 
line, there occurring in patches of green vegetation, but it has been 
found abundant on mountain slopes and in mountain meadows of 
lush grasses and other plants such as marsh marigold (Caltha lepto- 
sepala Hooker). At each of two localities (Los Pinos, Colo.; head of 
Chupidero Creek, Sangre de Cristo Range, N. Mex.), Hebard reported 
more than 50 adult specimens take by himself and Rehn, while on 
Neff Mountain, Cumbres Range, Colo., he reported their collecting 
more than 100 adults. Collection dates range from July 24 to Sep- 
tember 11, and nymphs have been observed throughout July and 
August. Hebard found about two-thirds of the specimens at the 
summit of Santa Fe Baldy still immature on August 1. Many nymphs 
of the intermediate population at Tomboy, Colo., were taken on Sept. 
2, 1921, by Rehn and Hebard. 


78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Melanoplus borealis palaceus Fulton 
Frcurss 8,a; 10,t; 11,d; 16,e; 18; PuatE 4,p. 


Melanoplus borealis palaceus Fulton, Ann. Ent. Soc. Amer., vol. 23, pp. 619-620, 
fig. 1,r-u, 1930 (5 males, 8 females, Upper Klamath Marsh, Oreg.). The 
holotype male and allotype female were designated by Fulton (USNM). 

Descriptive NoTEs: A usually fully winged subspecies, averaging 
slightly larger than borealis borealis, moderately larger than 6. 
stupefactus, and slightly smaller than 6. utahensis. Characterized 
especially in the male by the strongly upturned and recurved apical 
third of the abdomen, and by the subgenital plate which apically is 
considerably produced dorsally though only moderately produced 
posteriorly. Male tegmina usually reaching about to apex of 
abdomen and apices of hind femora, sometimes a little shorter; female 
tegmina averaging a little shorter, rarely slightly exceeding hind 
femora. 

Mate GENITALIA: Cercus usually as in figure 8,a-2, somewhat 
variable (fig. 8,a-3,4), and in holotype (fig. 8,a-1) its lateroposterior 
margin rather irregular; supra-anal plate and furculae of holotype 
and of most specimens much as in figure 9,d, in a few specimens the 
furculae somewhat shorter and less widely separated at their apices; 
subgenital plate in lateral view (fig. 16,¢) somewhat variable, the 
apical portion of dorsal margin much elevated, in posterior view the 
dorsal margin (fig. 11,d) usually entire, sometimes with traces of 
emargination, rarely with a median depression creating twin knobs, 
the posterior surface of subgenital plate usually with a roughly rec- 
tangular vertical area which is sometimes weakly excavate (x of fig. 
11,d-5); one paratype (Univ. Michigan) with subgenital plate less 
elevated and with rectangular area of posterior surface less well 
defined than in holotype. Aedeagus essentially as in borealis borealis; 
epiphallus in dorsal view usually about as in figure 8,f, in holotype 
the apices of lophi somewhat more lanceolate, in lateral view lophus 
usually noticeably flattened at top near apex, much as in figure 10,e. 

FEMALE GENITALIA: About as in borealis borealis; cercus of allotype 
essentially like figure 10,/, in some specimens slightly more acute; 
dorsal ovipositor valve with “scoop” of allotype (fig. 10,t) and several 
other specimens more deeply excavate than usual in b. borealis, but in 
other females not clearly different, margins of ‘‘scoop” slightly irregu- 
lar, seldom with a deep notch as in allotype. 

VaRIATION: The size of seven representative males, measured in 
millimeters, varies in pronotal length from 4.2 to 4.8 (av. 4.5), in 
length of hind femur from 10.8 to 11.7 (av. 11.2), and in length of 
tegmen from 11.0 to 14.5 (av. 12.6). Seven measured females vary 
in pronotal length from 4.9 to 5.9 (av. 5.3), in length of hind femur 


GRASSHOPPERS—GURNEY AND BROOKS 79 


from 12.3 to 14.8 (av. 13.4), and in length of tegmen from 11.7 to 15.3 
(av. 14.3). 

Cotoration: Averaging rather dark for group; hind femur usually 
with transverse bands evident only dorsally, ventral surface, part of 
mesal surface and external surface ventrad of paginae usually reddish, 
rarely yellowish; hind tibia reddish. 

Specimens intermediate between borealis palaceus and 6. utahensis 
are discussed under the latter. Three males and one female from 
Teton Co., Wyo., 5 males from Togwatee Pass (about 30 mi. east of 
Jackson Hole), Wyo., and 2 males from Yellowstone National Park, 
Wyo. (south side of Mt. Washburn, 8,600 ft.; Camp Cowan, Fireside 
River, 7,100 ft.), are considered intermediate between b. palaceus and 
b. borealis because of the shape of the male subgenital plate. Males 
typical of b. borealis also have been seen from Yellowstone National 
Park, and it is evident that the latter is a tension area. Males from 
southern Utah are not fully typical of 6. palaceus, and they seem to 
vary toward 6. borealis. Three males from Hackamore, Calif., also 
are atypical. The apex of the subgenital plate of those males is 
dorsally produced less than in typical palaceus, and there is scarcely 
any development of a rectangular area on the posterior surface of 
the subgenital plate. The number of localities from which 6. palaceus 
is known are too few and too scattered to permit a complete under- 
standing of the relationships of this subspecies. 

DisrrisutTion: Male specimens of Melanoplus borealis palaceus 
have been examined from the following localities: 

Montana: Lakeview area. 
Wromine: Cokeville. 
Uran: La Sal Mts., Grand Co.; Puffer Lake, Tushar Mts.; East Fork of Merchant 

Valley, Tushar Mts.; Beaver; Cedar Breaks. 

Wasuineton: Bonaparte Lake, Okanogan Co. 
OrEecon: Upper Klamath Marsh; Bly; Bulls Prairie, Warner Mts. 
CauirorniA: Hackamore, Modoe Co. 

BIOLOGY AND ECONOMIC IMPORTANCE: The types of M. borealis 
palaceus were “found near the pine woods on the border of a large 
meadow land known as Upper Klamath Marsh, situated on the 
plateau” east of Crater Lake, Oreg. (Fulton, 1930). A male and 
female from Bly, in Klamath Co., Oreg., about 50 miles southeast of 
the type locality, are labeled “Swp. mdw.,’”’ presumed to mean swampy 
meadow. ‘There is little information about altitudinal preferences of 
this grasshopper beyond the general upland character of all the areas 
where collections have occurred. In the La Sal Mountains, the most 
easterly locality where palaceus has been taken, various habitats are 
available, as discussed by Tanner and Hayward (1934). At Upper 
Klamath Marsh, Oreg., adults have been taken on June 23 and August 


80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


11, and in the Lakeview area, Mont., from July 4 to August 24. 
The Bonaparte Lake, Wash., record is based on a male taken on 
July 14 at an altitude of 4,000 feet. Three males and one female 
from Hackamore, Calif., were taken July 17. The Hackamore area 
is characterized by swampy places in which wildlife refuges have 
been established, but there is no record of the habitat from which the 
specimens were taken. Specimens taken by Rehn and Hebard in the 
Tushar Mountains, Utah, were at altitudes varying from 8,200 to 
9,000 feet, while on the Markagunt Plateau at Cedar Breaks, Utah, 
captures occurred at 10,400 feet. Dates of these Utah captures by 
Rehn and Hebard ranged from August 20 to August 31. 

Because of the scanty information about the environment in most 
of the localities where palaceus has been collected, a brief description 
of the Lakeview area, Mont., is appropriate. For these notes we are 
indebted primarily to Frank T. Cowan, who loaned an unpublished 
manuscript prepared by the late H. M. Jennison, when, as a member 
of a research team under the direction of J. R. Parker in the early 
1930’s, he studied the vegetation of the Centennial Valley. Lakeview 
is a small village in the Centennial Valley, in extreme southeastern 
Beaverhead County. This high mountain valley is 40 to 45 miles long 
and 3 to 8 miles wide, and is bounded on the south by the Centennial 
Mountains. The lowest level of the valley floor, through which the 
Red Rock River flows westward, is about 6,400 feet, and Red Rock 
Pass, at the east end of the valley, is about 7,000 feet. In the spring, 
run-off from melting snow in the mountains provides many wet areas 
in the meadows of wild hay, and it also supports several shallow lakes. 
Specimens of palaceus are noted in the Bozeman laboratory records as 
having been taken in two environments, one dominated by wiregrass 
and the other by bluegrass. 

Wiregrass (Juncus balticus) is dominant in extensive swales, or areas 
of low marshy ground. In an average year the swales are wet with 
slowly moving water until July, and some arms remain wet until late 
July or early August. The bluegrass areas include two or more species 
of Poa, as well as species of Hordeum and Puccinella. Greasewood 
(Scarcobatus) also occurs. These bluegrass areas are much drier than 
the wiregrass swales, but often are adjacent to the latter or appear as 
islands in their midst. They are green and succulent in spring and 
early summer, but by mid-July the grasses usually have matured and 
are light straw-colored or almost white. Both these environments are 
considered steps in the development of the grassland climax, as con- 
trasted with the sagebrush climax in other parts of the valley. 


GRASSHOPPERS—GURNEY AND BROOKS Sl 


Melanoplus borealis utahensis Scudder, new combination 
FicureEs 8,b,g; 9,e; 10,e; 11,e; 16,f; 18; PLatE 4,8 


Melanoplus utahensis Scudder, Proc. Amer. Philos. Soc., vol. 36 (154), pp. 19, 32’ 
1897; Proc. U. S. Nat. Mus., vol. 20, pp. 132, 167-168, pl. 11, fig. 10, 1897 
(1 male, Salt Lake Valley, Utah). 

The unique male type, cited by Rehn and Hebard (1912, p. 79), is 
labeled ‘S. L. Val. Utah 8-30’; “L. Bruner Collector’; “Drawn”; 
“Mel. utahensis Scudder’s Type, 1895.” Also, the specimen bears a 
Bruner manuscript label and more recently attached type labels and 
comments on type locality (USNM). 

Neither precise type locality nor the year of the type’s capture is 
definitely known. When Hebard examined the type, probably in the 
late 1920’s, he attached a label reading “From a western canyon of the 
Wasatch Mts., back from Ogden.”” A male and female from a series 
of four specimens from Ogden in the Philadelphia (ANSP) collection 
were labeled by Hebard as topotypes. Each of these bears a printed 
label “Ogden, Utah,” and one male is labeled “‘Melanoplus utahensis 
Bruner, Type spec.,”’ apparently in Bruner’s handwriting. The pair 
labeled as topotypes by Hebard bear the further notation “Taken w. 
type,” added by Hebard. Neither in Scudder’s revision nor elsewhere 
have we found any information relating to these four specimens. It 
may be that Bruner did not send them to Scudder for inclusion in the 
revision and that, although Bruner gave them a manuscript name, 
they were not studied again until his North American collection be- 
came available to Hebard. Scudder referred to the Bruner manuscript 
name when wtahensis was described, and the unique type has a Bruner 
name label like the one on the Ogden male. These labels may have 
led Hebard to assume that the whole series was taken together. 

Unless more information supporting Hebard’s opinion becomes 
available, it seems logical to conclude from the itinerary discussed by 
Bruner (1890) that the type was collected near Salt Lake City in 
1890, and that the Ogden specimens are not topotypes. In a report 
to C. V. Riley dated Sept. 1, 1890, Bruner described a trip made in 
August, primarily to Idaho. Near the end of the trip he went to 
Ogden and Salt Lake City, ‘‘at both of which points collections were 
made.’’ Thence, he went directly home to Lincoln, Nebr.; by train. 
Because of the date on the specimen, August 30, it is probable that 
the specimen was collected during Bruner’s last day in the field. 
Since Big Cottonwood Canyon, where utahensis has been collected, 
is within a few miles of Salt Lake City, that or a nearby area may 
well have been the locality which Bruner visited. The 1890 trip is 
the only one for which a record apparently satisfying the known 
data has been found. The fact that the specimen was deposited in 

47711959 6 


82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


the Riley collection, as recorded by Scudder, suggests that it was 
taken while on a Federal assignment such as the one in 1890. 

M. borealis utahensis, considered by Hebard (1928, p. 281) probably 
to be a distinct species closely related to M. bruneri, is shown by the 
aedeagus to belong instead to the borealis line of development. 

Descriptive Notes: A nearly fully winged subspecies of medium to 
large size in relation to other subspecies. Characterized especially in 
the male by the posteriorly prolonged (as well as elevated) subgenital 
plate. Tegmina failing by about 2 to 3 mm. to reach the tips of 
hind femora and the end of the abdomen. Posterior third of abdomen 
strongly upturned. 

MALE GENITALIA: Cercus typically as in figure 8,b-1-2, sometimes 
with the apex more obliquely rounded (fig. 8,b-3); furculae broad, in 
holotype about as in figure 9,e-1, sometimes more tapering; subgenital 
plate in lateral view extremely prolonged, somewhat variable in 
shape (fig. 16,f), in posterior view with dorsal margin varying from 
twice emarginate (fig. 11,e-1-3) to entire. Aedeagus as in borealis 
borealis; epiphallus (fig. 8,g) with lophi somewhat oblique anteriorly 
in dorsal view, in lateral view (fig. 10,e) rather flattened anteriorly. 

FEMALE GENITALIA: Cercus about as in borealis borealis (fig. 10,1); 
Ovipositor essentially as in b. borealis except that “scoop”’ of dorsal 
valve averages slightly more excavate, with apex more upturned. 

CoLoraTIon: Type with narrow yellowish green longitudinal stripes 
on lateral carinae of pronotum, lower half of lateral lobe of pronotum 
and lower part of head similar; metepisternal stripe conspicuously 
yellowish; tegmen brown, unmarked; hind femur with external 
paginal area light brown, paler toward base, dorsal surface with 
three distinct dark bars, in addition to dark brown knee, external 
surface ventrad of paginae pink, mesal surface tinged with pink; 
hind tibia pink. Specimens from Big Cottonwood Canyon lighter- 
colored than type, tegmina and dorsum of pronotum light brown, 
most of remainder of body yellowish brown, hind tibia weak pinkish. 

Variation: Measurements (in millimeters) of the holotype are: 
Body, 27.0 (Scudder’s measurement) ; pronotum, 6.1; hind femur, 14.3; 
and tegmen, 17.0 Specimens from Ogden average larger than those 
from Big Cottonwood Canyon. Measurements of the Ogden speci- 
mens are: Males (2), pronotum, 5.3, 5.5 (av. 5.4), hind femur, 12.0, 
12.0 (av. 12.0), tegmen, 15.3, 18.8 (av. 17.1); females (2), pronotum, 
5.7, 6.0 (av. 5.9), hind femur, 13.7, 14.7 (av. 14.2), tegmen, 12.5, 
15.5 (av. 14.0). Measurements of representative Cottonwood Canyon 
specimens are: Males (3), pronotum, 4.6 to 5.0 (av. 4.8), hind 
femur, 11.4 to 11.6 (av. 11.5), tegmen, 11.3 to 11.8 (av. 11.5); females 
(2), pronotum, 5.1, 5.3 (av. 5.2), hind femur, 12.4, 12.5 (av. 12.5), 
tegmen, 12.3, 12.5 (av. 12.4). 


GRASSHOPPERS—GURNEY AND BROOKS 83 


The series from 3 miles north of Mantua, Utah, which was labeled 
in the Philadelphia (ANSP) collection by Morgan Hebard as atypical 
Melanoplus utahensis, is here considered intermediate between borealis 
utahensis and b. palaceus. External male genitalia are as illustrated 
(figs. 8,c; 9,6; 11,f; 16,9), and the subgenital plate suggests the inter- 
mediate relationship of the population. The lophi of the epiphallus 
are apically lanceolate in dorsal view, well flattened at the top in 
lateral view. The general color of these specimens is dull yellowish 
brown; hind tibiae are tinged with pink. The general color suggests 
that of the Red Banks series of brwnert and may reflect comparable 
ecological conditions. 

Measurements (in millimeters) of representative Mantua speci- 
mens are: Males (5), length of pronotum, 5.9 to 6.4 (av. 6.2), hind 
femur, 12.4 to 14.2 (av. 13.5), tegmen, 12.5 to 15.5 (av. 14.3); females 
(2), length of pronotum, 6.5, 7.0 (av. 6.8), hind femur, 14.8, 15.5 (av. 
15.2), tezmen, 15.0, 16.0. (av. 15.5); 

Disrrrsution: This grasshopper appears to be a relatively localized 
subspecies of the northern Wasatch Mountains, and to be more or 
less surrounded by M. borealis palaceus rather than being distributed 
linearly adjacent to M. 6. palaceus. ‘There may be altitudinal differ- 
ences also, so that material from additional localities is likely to 
clarify the situation. Full data on all specimens examined are as 
follows: Salt Lake Valley, Utah, Aug. 30, L. Bruner (10%, holotype) ; 
Big Cottonwood Canyon, Wasatch Mts., Utah, 7,400 ft., Canadian 
zone, Aug. 19, 1924, Rehn and Hebard (807, 82); Ogden, Utah 
(207, 29). 

Intermediate between Melanoplus borealis utahensis and M. 6. 
palaceus: 3 miles north of Mantua, Box Elder Co., Utah, Wasatch 
Mts., 5,800 ft., Aug. 9, 1928, Rehn & Hebard. (1867, 179, 1 juv. ¢&, 
apparently 5th instar). 

BIoLOGY AND ECONOMIC IMPORTANCE: We are indebted to Rehn 
and Hebard for the only notes on the biology of this poorly known 
grasshopper. In Big Cottonwood Canyon, Utah, on Aug. 18, 1924, 
they found it in the Canadian zone at 7,400 feet, present ‘in moderate 
numbers in lush herbage near stream on flat with much aspen and 
spruce” (Hebard, 1936b, p. 173). Hebard also gave notes on a 
grasshopper taken at Red Banks in Logan Canyon, and ascribed to 
utahensis, but, as previously mentioned under M. bruneri, the only 
specimens we have seen with corresponding data prove to be bruneri. 


84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


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KNABEN, NILS 
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GRASSHOPPERS—GURNEY AND BROOKS 89 


KozHAntsuHikoy, I. V. 

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90 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


PacKARD, A. 8S. 

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GRASSHOPPERS—GURNEY AND BROOKS Ol 


Rean, J. W. H. 

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Rivey, C. V.; Packarp, A. 8., Jr.; anp THomas, Crrus 

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1941. A comparative study of the subfamilies of the Acrididae (Orthoptera) 
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1951. The North American buffalo, a critical study of the species in its wild 

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SCHLEBECKER, JOHN T. 

1953. Grasshoppers in American agricultural history. Agricultural History, 

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issued as a separate with pp. 498-515.) 


92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


1878a. Remarks on Calliptenus and Melanoplus, with a notice of the species 
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1878b. Brief notice of the American species of Melanoplus found west of the 
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SHOTWELL, RoBeErT L. 

1930. A study of the lesser migratory grasshopper. U.S. Dep. Agr. Tech. 
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1941. Life histories and habits of some grasshoppers of economic impor- 
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1952. Some effects of various food plants on Melanoplus mexicanus mexi- 

canus (Sauss.). Canadian Ent., vol. 84, pp. 113-117. 
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1954. An analysis of 100 years of grasshopper populations in Kansas (1854 

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1936. Some notes and records of Minnesota Orthoptera. Ent. News, vol. 
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TANNER, Vasco M., anp Haywarp, C. LYNN 

1934. <A biological study of the La Sal Mountains, Utah. Report No. 1 
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TINKHAM, E. R. 

1948. Faunistic and ecological studies on the Orthoptera of the Big Bend 
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TREHERNE, R. C., AND BucKELL, E. R. 

1924. Grasshoppers of British Columbia. Canadian Dep. Agr. Bull. 39, 

new ser., pp. 1-47, 18 figs., 3 pls. 


GRASSHOPPERS—GURNEY AND BROOKS 93 


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WALKER, FRANCIS 
1870. Catalogue of the specimens of Dermaptera Saltatoria in the collection 
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WatsH, BENJAMIN D. 
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1953. A grasshopper outbreak near Palmer, Alaska, 1951. Journ. Econ. 
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1941. New records of Acrididae from Alberta. Canadian Ent., vol. 73, p. 
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1945. <A contribution to the knowledge of the Acrididae of Alberta. Sci. 
Agr., vol. 25, pp. 577-596. 
Wizzvr, D. A., anv Fritz, R. F. 
1940. Grasshopper populations of typical pastures in the bluestem region 
of Kansas. Journ. Kansas Ent. Soc., vol. 13, pp. 86-100. 
Witson, C. C. 
1947. Control of the devastating grasshopper in California. Bull. Cali- 
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Woopwokty, C. W. 
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1-36, 17 figs. 
1913. Guide to California insects. 360 pp., 361 figs. 


U. S. GOVERNMENT PRINTING OFFICE: 1958 


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a, Melanoplus mexicanus, male, La Cumbre, Mexico. B, c, .M. bilituratus bilituratus, 
males: B, from Vancouver Island, British Columbia, Canada; c, from Claydon, Saskatche- 
wan, Canada. pb, £, M. bilituratus vulturnus, females: p, from Stony Man Mountain, 
Shenandoah National Park, Va.; ©. from Independence, Kans. 


PROC. U.S. NAT. MUS. VOL. 110 GURNEY AND BROOKS, PLATE 2 


A-C, Melanoplus bilituratus defectus: a, male from Pendleton’s Ranch, 4 miles south of 
fumaeacori, Ariz.; 8, male from Tempe, Ariz., reared: c, female from Chandler, Ariz. 
T1Z.; J pe, 
b, M. spretus, female from Iowa, 1873. 


PROC. U.S. NAT. MUS. VOL, 110 GURNEY AND BROOKS, PLATE 


,, Melanoplus brunert, male, from Sheridan County, Wyo. 8, ¢, M. devastator: B, male, 
from Folsom, Calif.; c, female from El Dorado County., Calif. 


PROC. U.S. NAT. MUS. VOL. 110 GURNEY AND BROOKS, PLATE 4 


A, Melanoplus bilituratus defectus, male, from Chandler, Ariz. B, M. borealis utahensis, 
male, from Ogden, Utah. _c, male, intermediate between M. borealis utahensis and M. borealis 
palaceus, from 3 miles north of Mantua, Utah. ov, M. borealis palaceus, male, holotype. 


GURNEY AND BROOKS, PLATE 5 


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Vol. 110 Washington : 1959 No. 3417 


A REVISION OF AMERICAN BATS 
OF THE GENERA EUDERMA AND PLECOTUS 


By Cartes O. Hanp ey, JR. 


Introduction 


Confusion and damage to nomenclatural stability are consequences 
of taxonomic revisions that do not provide adequate explanations or 
justifications for proposed changes in nomenclature. They are ig- 
nored, accepted with reservation, or blindly followed by those who 
have occasion to use them. Confusion of this sort involves a wide- 
spread American bat, now referred to in literature either as Cory- 
norhinus or Plecotus. The purpose of this paper is to clarify the status 
of this bat and its American and Eurasian relatives (particularly 
Euderma), to assay the evolution of the group, to map geographic 
distributions, and to bring together the literature on natural history of 
the group. 

The genera Euderma and Plecotus are vespertilionid bats with ex- 
ceptionally large ears. They are known colloquially as mule-eared 
bats, jack-rabbit bats, or, simply, big-eared bats, or long-eared bats. 
Some species are referred to as lump-nosed bats. Plecotus, as here 
understood, includes the American Jdionycteris and Corynorhinus as 
subgenera. 

Perhaps the earliest published reference to an American big-eared 
bat was that of Clayton (1694, p. 124) in his remarks on the “‘beasts 
of Virginia”: ‘Batts, as I remember at least two sorts; one a large 

95 


96 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


sort with long Ears, and particularly long stragling hairs. The other 
much like the English, something larger I think, very common.”’ 

There is now an extensive American literature on Huderma and 
Plecotus, but most reports contain little more than records of speci- 
mens. The only revisionary works have been those of H. Allen (1864, 
1894), Miller (1897), and G. Allen (1916). Most of our knowledge 
of the anatomy and physiology of the group is included in the papers 
of Larsell and Dow (1935) and Pearson, Koford, and Pearson (1952). 
The few life history studies are generally superficial; those of Dalquest 
(1947), Pearson, et al. (1952), and Twente (1955) are notable excep- 
tions. Much yet remains to be learned about Huderma and the Ameri- 
can populations of Plecotus. 

These bats occur in most temperate portions of the Northern 
Hemisphere. They are generally scarce. In America, Huderma is 
known by 16 specimens, the subgenus IJdionycteris is known by less 
than a dozen specimens, all but two of them collected since 1955; and 
perhaps no other North American bat with such an extensive geo- 
eraphic range as the subgenus Corynorhinus has been collected so in- 
frequently. Species of the latter subgenus are known to occur from 
the Atlantic to the Pacific and from Canada to the Isthmus of Tehuan- 
tepec in Mexico. This wide geographic range is deceptive, however, 
for these bats display narrow limits of tolerance in their choice of 
habitat. Within the bounds of their ranges are large areas in which 
they apparently do not occur, and others in which they occur only 
sparingly. There are few areas in which they are common. 

MATERIALS 

The scarcity of Euderma and Plecotus in America is evident from 
the small number of museum specimens listed in the early reviews. 
H. Allen’s 1864 monograph included only 18 specimens, and his 1894 
monograph only 19. For Miller’s 1897 revision, 43 examples were 
available. G. Allen’s 1916 review was based on a total of 126 speci- 
mens. 

The material from which I have drawn my conclusions concerning 
these bats consists of four dry skins and skulls and a single specimen 
preserved in alcohol representing Huderma; three specimens of the 
subgenus Jdionycteris, dry skins and skulls; 25 dry skins and skulls, 
two skeletons, and 32 specimens preserved in alcohol, representing at 
least five races of the subgenus Plecotus; and about 800 specimens (see 
species accounts for type of preservation) representing all the named 
forms of the subgenus Corynorhinus. 

In the course of the study I have examined enough specimens to 
define Huderma and American Plecotus, but have not attempted to 
assemble all available specimens. Those examined are in the collec- 
tions of the following: 


AMERICAN BATS—HANDLEY 97 


American Museum of Natural History 


(AMNH) 

Albert Schwartz (private collection) 
(AS) 

California Institute of Technology 
(CIT) 


Carlsbad Caverns National Park (CC) 

Charleston (South Carolina) Museum 
(ChM) 

Carnegie Museum (CM) 

Chicago Natural History Museum 
(CNHM) 

W. Gene Frum (private collection) 
(GF) 
Instituto de Biologia, Universidad Na- 
cional Auté6noma de México (IB) 
Illinois Natural History Survey 
(INHS) 

University of Kansas 
Natural History (KU) 

Louisiana State University Museum 
of Zoology (LSU) 

Harvard University Museum of Com- 
parative Zoology (MCZ) 

University of California Museum of 
Vertebrate Zoology (MVZ) 

Oklahoma A. & M. College (OAM) 


Museum of 


Ohio State Museum (OSM) 
School of Tropical and Preventive 


Medicine, Loma Linda, Calif. 
(STPM) 

Texas A. & M. College (Texas Co- 
operative Wildlife Collection) 
(TCWC) 


Texas State Department of Health, 
Austin, Tex. (TSDH) 

Texas Technological College (TT) 

University of Arkansas Department of 
Zoology (UAZ) 

University of Illinois Museum of Nat- 
ural History (UI) 

University of Kentucky Zoology De- 
partment (UK) 

University of Michigan Museum of 
Zoology (UMMZ) 

University of Oklahoma Museum of 
Zoology (UOMZ) 

U.S. National Museum (including 
U.S. Fish and Wildlife Service’s 
Biological Surveys collection) 
(USNM) 

University of Utah Museum of Zool- 
ogy (UU) 


I express sincere thanks and appreciation to the owners of private 
collections and to the museum authorities who kindly loaned speci- 
mens for my study. To those at the American Museum of Natural 
History and at the Museum of Comparative Zoology, Harvard Univer- 
sity, I am particularly grateful. I am indebted to Seth B. Benson, 
Museum of Vertebrate Zoology, University of California, and to 
Rollin H. Baker and E. Raymond Hall, Museum of Natural History, 
University of Kansas, for the loan of specimens set aside for their 
personal study. Special thanks are due to John A. Sealander, Univer- 
sity of Arkansas, for allowing me to utilize specimens under his care 
in the description of a new subspecies, and for his kindness in depositing 
the type specimen in the U.S. National Museum; and to Aurelio 
Malaga Alba, Pan American Sanitary Bureau, for his cooperation in 
securing much needed Mexican specimens, one of which has served 
as the type of a new subspecies. The Fish and Wildlife Service, 
U.S. Department of the Interior, has generously allowed me to utilize 
from its files the unpublished field notes of its collectors. 

The following individuals contributed information and other assist- 
ance for which I am grateful: R. W. Barbour, F. L. Burnett, W. W. 
Dalquest, W. G. Frum, A. F. Ganier, Helen Gaylord, B. P. Glass, 
Woodrow Goodpaster, R. M. Goslin, Iracy O. Handley, R. H. Hand- 


98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ley, H. B. Hitchcock, Patricia Isham, E. V. Komarek, Barbara 
Lawrence, R. E. Mumford, Thelma Reid, C. C. Sanborn, William 
Schaldach, Jr., Albert Schwartz, P. M. Smith, P. W. Smith, Arthur 
Stupka, Robert Traub, Bernardo Villa R., and F. N. Young. 

I am thankful to the persons who reviewed the manuscript and 
offered helpful criticism and advice: A. M. Elliott, C. W. Hibbard, 
EK. T. Hooper, Remington Kellogg, D. H. Johnson, H. W. Setzer, 
A. H. Smith, and C. F. Walker. 


MeETHODS 


In tables 7-18 cranial and external measurements, in millimeters. 
are given for Huderma maculatum, Plecotus auritus, Plecotus phyllotis, 
and for all the races of the subgenus Corynorhinus. Females are 
significantly larger than males in these bats. Accordingly, measure- 
ments for the two sexes are tabulated separately. Only adults, as 
determined by ossification of the finger joints and appearance of the 
skull, were measured. 

Measurements of Huderma are from the following sources: Ashcraft 
(1932, p. 162), Durrant (1952, pp. 59, 66), Nicholson (1950, p. 197), 
Parker (1952, p. 480), Stager (1957, p. 260), and four specimens 
measured by me. 

External measurements of Plecotus, except of the forearm, are those 
taken by the collectors. JI measured the forearm (greatest length of 
radius-ulna including carpals) on the preserved specimens. All cranial 
measurements (fig. 1) were taken with dial calipers and the aid of a 
binocular microscope in the following manner: 

Greatest length: Distance, on the diagonal, from the anteriormost border of the 
premaxilla to the posterior limit of the supraoccipital; incisors are excluded. 
Zygomatic breadth: Greatest distance between the outer borders of the zygomatic 

arches. 

Interorbital breadth: Least diameter of the interorbital constriction of the 
frontals. 

Brain case breadth: Distance, measured above the auditory bullae, between the 
lateral limits of the parietals. 

Brain case depth: Distance between the dorsal extreme of the skull and the 
ventral limit of the occipitosphenoid plate; auditory bullae are excluded. One 
bar of the calipers lies flat on the occipitosphenoid plate; the other is in 
contact with the top of the brain case. 

Maxillary tooth row: Distance between the anterior edge of the canine and the 
posterior border of the third molar. 

Postpalatal length: Distance from the anterior limit of the mesopterygoid fossa 
to the ventral lip of the foramen magnum. 

Palatal breadth: Distance between the lateral extremes of M3. 

Presence or absence of a secondary cusp in I! was indexed according 
to the following scale (fig. 2): 


AMERICAN BATS—HANDLEY 99 


0, Secondary cusp absent; no trace. 
1, Secondary cusp incipient, in the form of a shoulder, not separated from primary 
cusp by a notch. 
2, Secondary cusp present, separated from primary cusp by a notch. 
Frequently, the degree of development of the secondary cusp of I! 
is not the same on both sides. When absent in one tooth and incipient 
in the other, it was indexed as 0.5; when absent in one and present 
in the other, as 1; when incipient in one and present in the other, 
as 1.5; etc. 
The degree of development of the secondary cusp in each population 
was determined by averaging individual indexes. Thus, a population 


GREATEST LENGTH 


BRAIN CASE 
DEPTH 


— BRAIN CASE 
BREADTH 
ZYGOMATIG 

BREADTH INTERORBITAL 


BREADTH 


PALATAL 
BREADTH 


POSTPALATAL LENGTH 


Ficure 1.—Cranial measurements of Euderma and Plecotus. 


100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Vays 


6) 1 2 


Ficure 2.—Degrees of development of the secondary cusp in the first upper incisor 
(anterolateral view). 


with an incisor cusp index of 0 lacks the secondary cusp. An index 
of 1 indicates some development of the secondary cusp, and so on. 

Throughout the text, relative proportions are expressed as per- 
centages of the greatest length of the skull. 

Color was measured in diffused white fiuorescent light, without 
direct natural light. Capitalized color terms in the text are from 
Ridgway (1912). There appears to be no sexual dichromatism or 
seasonal variation in coloration. If such exists, it is so slight that, 
except in cases of extremely worn pelage, it can be disregarded in 
comparisons. 

The type of preservation of the specimens examined is indicated 
in the species accounts by the following abbreviations: a, alcoholic; b, 
bones (skeleton) ; and s, skin and skull. 


DENTAL NOMENCLATURE 


The nomenclature of cusps and the numerical designation of teeth, 
as used in this paper, are outlinediin figure 3. The naming of the 
individual teeth requires further eculenvGon 

The complete primitive dentition of adult placental mammals is a 


set of 44 teeth: 
3-3 1-1 4-4 3-3 
Lee ee ey 
The nearest approach to this in bats is a set of 38 teeth, lacking one 
upper incisor and one upper and one lower premolar in each jaw: 


3-3 ae 


cx a Pe 3, MS 


Incisors: Which of the incisors it is that is always missing in 
Chiroptera is a matter of controversy. Most authors have believed 
without much doubt that the consistently missing incisor is the first. 


According to Miller (1907, p. 27) this is indicated, as shown by Winge, by the 
correspondence of the two upper teeth with the two outer of the lower jaw when 
the maximum set is present, and also, even more strongly, by the general tendency 
throughout the group for the premaxillaries to become reduced, particularly 
along the inner edge. This would inevitably result in eliminating that part of 
the bone in which the first incisor grows. 


AMERICAN BATS—HANDLEY 101 


PARACONE PROTOCONE 
(PROTOCONE) (DEUTEROGONE) 

MESOSTYLE HY POCONE 
(TRITOCONE) (TETAR TOCONE) 

PARACONE PROTOCONE 

ME TACONE HYPOCONE 


PARASTYLE ————__ 
m2 
MESOSTYLE 


1ST GOMMISSURE 
2ND GOMMISSURE 
3RD COMMISSURE 


METASTYLE 


CINGULUM 


METACONID 


(TRITOGONID) 
PROTOCONID PARACONID 
PROTOGONID METACONID 
HYPOCONID ENTOCONID 


Ss \-———- rriconio 
| cecal 


Figure 3.—Nomenclature of teeth of Plecotus. Right upper dentition (above) and left 
lower dentition (below). 


There are many objections to the belief that the permanently 
missing incisor is I’, and no really substantial evidence as to which, 
if either, of the others it may be. However, what evidence there is 
favors the theory that the missing tooth may be I’. 

The deciduous dentition does not contribute information on the 
lost tooth. No more than two upper incisors have been observed in 
any normal chiropteran deciduous dentition. For the Vespertilionidae, 
the formula is the same in all of the genera for which it is known: 
=22- 


. 2-2 1-1 2-2 
di 3-3” de = dp 3-2 


102 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


On the basis of an abnormal supernumerary deciduous incisor in 
Myotis lucifugus, Stegeman (1956, p. 60) postulated that the normally 
missing tooth may beI?. G. Allen (1916, p. 335) described a specimen 
of Plecotus townsend (USMN 150273) with three upper permanent 
incisors. The “extra’’ incisor was the outermost of the three and 
differed in form from the other two. Allen speculated that this 
might represent a long lost I. 

The uniform absence of a tooth throughout a group as large and 
as varied as the Chiroptera compels the impression that the absence 
is a characteristic dating from the dawn of development of the group 
and subsequently shared by all derivatives. It is doubtful that 
deviations from such a deep-seated characteristic, such as the devia- 
tions reported by Allen and by Stegeman, can be assumed to represent 
a reversion to an even more primitive long lost characteristic. 

An example of a clearly abnormal extra incisor may be seen in 
a specimen of P. mericanus (KU 29858). In place of the right outer 
upper incisor there are two closely approximated twin teeth, neither 
of which resembles the normal outer incisor in the left premaxilla. 

Investigation of the factors now operating to cause reduction and 
loss of the remaining incisors in Recent bats may give clues to the 
incisor missing throughout the order. As noted above, Miller, 
following Winge, supposed that reduction of the inner portion of the 
premaxillary bone left the innermost incisor without a place for its 
root. However, although all bats lack one upper incisor, not all bats 
have the tooth-bearing portions of the premaxilaries reduced. There- 
fore, if the same incisor is missing in all bats, then reduction of the 
premaxillaries can not always account for the loss. 

Actually, several evolutionary trends may account for crowding 
and subsequent reduction and loss of incisors. In forms with an 
extensible tongue, the action of the tongue may be a responsible agent. 
In others, a narrowing of the rostrum and mandibles, an enlargement 
of the canines, or a reduction of the tooth-bearing portions of the 
premaxillaries may be a cause. 

The result of these processes is always the same. It is not the 
reduction and loss of the innermost of the two remaining incisors, but 
a crowding of both toward the canine. A consequence of this crowding 
is conflict of the outermost incisor with the occluding lower canine. 
Further alternative consequences are reduction of the lower canine, 
diminution or obliteration of the outermost upper incisor, anterior 
or posterior displacement of the outermost incisor, rotation of the 
outermost incisor to a narrower axis, elevation of the rostrum, or 
depression of the mandible. All of these conditions may be observed 
in Recent bats. It is important to note that in all Chiroptera it is 
invariably the outer of the two remaining incisors that is reduced in 


AMERICAN BATS—HANDLEY 103 


size and apparently is the first to disappear. It is most reasonable 
to assume that these same factors may have operated to eliminate the 
missing incisor. The first to go then, at a very early stage of 
chiropteran development, was probably I°. This is in agreement with 
Andersen’s (1908, p. 205) and Thomas’ (1908, p. 348) criticisms of 
Miller’s conclusions. 

PREMOLARS: Since the number of premolars in each maxilla and 
mandible in all known bats is at least one less than the primitive 
placental number, it is assumed that one has been lost from each jaw. 
Thomas (1908, p. 347) argued convincingly that the missing tooth is 
the second. Miller (1907, p. 28), without proof, arbitrarily assumed 
it to be the first. Evidence at hand possibly supports Miller’s thesis. 
A specimen of Plecotus townsend (USNM 81647) has four premolars 
in one mandible; the other ramus is normal. ‘The extra tooth is on 
the lingual side of the tooth row, adjacent to the normal anterior 
premolar and forcing that tooth to the labial side of the row. Both 
teeth touch the canine. The extra tooth is smaller than the normal 
anterior premolar and rather like P; in size and shape. If it were to 
be assumed that this extra tooth reflects a primitive condition in 
which the normal mandible bore four premolars, then it would be 
necessary, because of its position, to assume that the extra tooth is P. 
However, the argument refuting similar evidence for the missing 
incisor can be applied with equal propriety here. 

Besides lacking, as do all bats, P2 in maxilla and mandible, Huderma 
and Plecotus also lack another maxillary premolar. In this instance 
the missing tooth is probably P?. An examination of Myotis in this 
connection proves instructive. This genus normally lacks only a 
single premolar, assumed to be the second, in each jaw. In some forms 
the remaining premolars normally are not even crowded (e.g., MM. 
lucifugus). However, other forms (e.g., MM. occultus) frequently lack 
two premolars in each jaw. The tooth that is variable in occurrence 
is the third premolar. In 15 specimens of M. occultus P* was present 
in 4, absent in 11; P; was present in 14, absent in 1. P* normally is 
absent in Plecotus, but occasionally is present (USNM 265387, 
297711). In these specimens P* is present in only one maxilla, is 
similar to P! but is smaller, and is wedged between P! and P* (postero- 
internal to P!, on the lingual border of the tooth row). The “‘extra”’ 
tooth in each instance is similar in form and position to P*® of Myotis 
occultus. ‘These teeth perhaps represent the normal third premolar 
and not merely abnormal accessory teeth. They thus may be 
authentic reversions to a primitive condition. 

In addition, the third mandibular premolar normally is lacking in 
Euderma. In Plecotus it is present, but is smaller than either P, or Py. 
In the subgenus Corynorhinus it is commonly squeezed between the 


104 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


larger teeth so that its cross-sectional outline is distorted. Frequently 
it is displaced to the lingual side of the tooth row. 

DENTAL FORMULAE: Based on the foregoing conclusions, the dental 
formulae of the bats discussed in this paper may be written as follows: 


Se Myotis 

ee S38 Plecotus 

ae : = 34 Euderma, Barbastella 

aa *: = oe = 32 Histiotus, Laephotis 

= - — = =30 Otonycteris, Nyctophilus, Pharotis 
oa : ae 3 =28 Antrozous 


Supraspecific Nomenclature and Relationships 
History OF NOMENCLATURE 


SUPRAGENERIC NOMENCLATURE: Several genera of the family 
Vespertilionidae have exceptionally large ears: Plecotus (including 
Corynorhinus and Idionycteris, now regarded as subgenera), Huderma, 
Histiotus, Laephotis, Otonycteris, Nyctophilus, Pharotis, and Antrozous. 
The relationships of these genera have been variously interpreted. 
Typical of the opinion of his time, Dobson (1875, p. 348) included all 
known genera in his ‘Group Plecoti’’ which was the equivalent of a 
subfamily. As late as 1897 Miller (1897, p. 41) employed the name 
Plecotinae, of subfamily rank, to include the American big-eared 
vespertilionids Plecotus (subgenus Corynorhinus) and Euderma. For 
Antrozous he erected a separate subfamily, Antrozoinae. Later 
Miller (1907, p. 197) abandoned the subfamily Plecotinae and included 
its genera in the subfamily Vespertilioninae. At the same time he 
adopted the terminology of Peters (1865) and placed Antrozous in the 
subfamily Nyctophilinae. 

The large ears and auditory bullae and strong zygomata—the 
principal characteristics common to the 10 genera and subgenera 
listed above—may not indicate kinship. The classification of Tate 
(1941, p. 590; 1942, p. 225), based on other characters, probably 
better expresses the natural relationships of these genera and sub- 
genera: 

Subfamily Vespertilioninae 
Group Myotini 
Genera: Plecotus 
Corynorhinus 


Idionycteris 
Euderma 


AMERICAN BATS—HANDLEY 105 


Group Pipistrellini 
Genera: Histiotus 
Laephotis 
Group Nycticeini 
Genus: Otonycteris 
Subfamily Nyctophilinae 
Genera: Nyctophilus 
Pharotis 
Antrozous 

Tate (1942, p. 229) supposed the assemblage of big-eared bats of 
the group Myotini to be simply a specialized derivative of the Myotis 
stock, presenting an odd combination of primitive and specialized 
characters, but not being sufficiently differentiated to warrant the 
erection of a special category. 

GENERIC NOMENCLATURE: Initially confused with the exclusively 
South American genus Histiotus (J. Allen, 1891, p. 195), the bat 
H. maculatum was soon recognized as the representative of a distinct 
genus, Huderma (H. Allen, 1892, p. 467). Subsequently, the validity 
of Huderma has not been questioned. 

The status of the names Idionycteris, Plecotus, and Corynorhinus 
has been less clear. Jdionycteris has remained so little known since 
its description by Anthony in 1923 that there has been no speculation 
on its systematic position beyond Anthony’s supposition that its 
nearest relative is Plecotus auritus. The nomenclatural history of 
the Eurasian Plecotus has been one of stability. Subsequent to the 
recognition of the genus in 1818 by E. Geoffroy Saint-Hilaire, the 
big-eared vespertilionines of Europe and temperate Asia have been 
recognized consistently by the name Plecotus. 

No such stability characterizes the nomenclatural history of the 
bats to which the name Corynorhinus has been applied. Rafinesque 
used Vespertiio for the first named form in 1818. Later authors 
(Lesson, 1827; LeConte, 1831; Cooper, 1837) discerned a likeness 
between European and American big-eared bats and used the name 
Plecotus for both groups. Following this was a brief period of belief 
that the American big-eared bats and Eurasian barbastelles were 
related, and the name Synotis was used for both (Wagner, 1855; 
H. Allen, 1864). H. Allen gave the American big-eared vespertilionines 
independent generic status for the first time in 1865 with the name 
Corynorhinus. This position was confirmed by Miller (1897, 1907) 
and more recently by Tate (1942). In the 75 years following its 
conception, the name Corynorhinus has been in almost universal usage 
for the American forms exclusive of Euderma. A major exception 
was Dobson’s (1878) conclusion that Corynorhinus is a synonym of 
Plecotus. Beginning with Simpson (1945), there has been a new 
trend away from the independent use of the name Corynorhinus and 


106 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


a tendency to synonymize it under the name Plecotus. However, 
there seems to have been no study of specimens to support the con- 
tention that Corynorhinus and Plecotus are congeneric, or to determine 
the relation of Idionycteris to them. 


CLASSIFICATION OF Bic-EARED Bats 


While agreeing with the basic outline of Tate’s (1942, p. 225) 
classification of vespertilionid big-eared bats, I prefer a slightly 
modified version: 

Subfamily Vespertilioninae 
Group Myotini 
Genera: Euderma 


Plecotus 
Subgenera: Jdionycteris 
Plecotus 
Corynorhinus 


Group Pipistrellini 
Genera: Histiotus 
Laephotis 
Group Nycticeini 
Genus: Otonycteris 
In my opinion, the degree of difference between the nominal genera 
Idionycteris, Plecotus, and Corynorhinus is of subgeneric or specific 
rather than generic grade. Jdionycteris is a basal or relict form, 
whereas Plecotus and Corynorhinus represent more advanced or later 
evolutionary stages. Corynorhinus has evidently differentiated more 
rapidly than Plecotus, and has approached the point of generic dis- 
tinction from its near relatives. Inasmuch as these forms represent 
different stages of evolution and since they form a disjunct series, at 
least one of the steps of which contains more than one species, it seems 
best to regard the degree of difference between the three forms of as 
subgeneric rather than specific magnitude. Evidence in support of 
this thesis is presented in the following pages. 


CHARACTERS AND COMPARISONS 


Cranial characters relating Huderma and Plecotus and distinguishing 
them from other big-eared bats (Histiotus, Laephotis, Otonycteris, 
Nyctophilus, Pharotis, and Antrozous) of the family Vespertilionidae 
are: 


Rostrum relatively weak, narrow, and shallow. 

Brain case elongated and vaulted in the frontal region. 

Zygomata parallel (in dorsal view) and not bowed out posteriorly. 

. Dentition weak (teeth not robust). 

. Cusps of teeth high and sharp-pointed. 

. P, not so high as Mj. 

. Teeth reduced in number only slightly from the chiropteran maximum, 


NOT RWN HE 


AMERICAN BATS—HANDLEY 107 


The bats of the genera Huderma and Plecotus form a close-knit unit, 
morphologically and, apparently, phyletically. Huderma and Plecotus 
appear to be more closely related to one another than either is to any 
other vespertilionid genus, yet the differences separating them are 
clearly of generic grade (table 1). 

The nearest relative of Huderma and Plecotus appears to be Barba- 
stella (p. 118). Also closely related are Myotis and Pvrpristrellus 
and their closest allies. Similarities most striking are the lack of 


-BARBASTELLA BARBASTELLUS USNM 142583 SWITZERLAND 


4213Q ARIZONA 


PLEGOTUS (IDIONYGTERIS) PHYLLOTIS AMNH 62260 TAMAULIPAS 


Gaol FX KE 


PLEGOTUS (PLEGOTUS) AURITUS MGZ 36987 GERMANY 


PLECOTUS (CORYNORHINUS) TOWNSENDIL MGZ 40876 WASHINGTON 


Ficure 4.—Skulls of Barbastella, Euderma, and Plecotus (Barbastella about 4 times natural 
size, others about 3 times natural size). ‘Traced from photographs, 


108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


shortening of the rostrum and lack of dental specialization (loss of 
teeth and cusps). In these features the resemblance of Huderma and 
Plecotus to Myotis is greater, to Pipistrellus less. This relation is 
further substantiated in the postcranial skeleton, where Huderma and 
Plecotus have some characters of Myotis (species compared: lucifugus, 
thysanodes, yumanensis), and some of Pipistrellus (species compared: 
javanicus, subflavus), but in most features they are more like Myotis. 
On the basis of dental, osteological, and external features, it appears 
that Huderma and Plecotus should stand between Myotis and Pipi- 
strellus in the classification of Chiroptera. 

Genus Euperma: Huderma maculatum is more specialized than any 
species of Plecotus. It has carried auditory modification—the most 
spectacular character of the two genera—to the greatest extreme, 
and in dental characters it is unique. By elongation, in addition to 
enlargement, of the auditory bulla it has departed from the usual 
vespertilionid pattern. Externally too, Huderma has exceeded Plecotus 
and, indeed, all other vespertilionids in auditory specialization—in 
extreme enlargement of the auricle, in the simplification of the tragus 
by elimination of the posterior basal lobe, and in the attachment of 
the tragus to the posterior basal lobe of the auricle as well as to the 
anterior basal lobe so that the auricle forms a more perfect funnel 
to the auditory meatus. 

Hall (1934) pointed out important differences between the body 
skeletons of Euderma and Plecotus. Notable are the unusual length 
and shape of the presternum and the shape and angle of the acromion 
process of the scapula in Huderma. 

The anterior portions of the tooth rows show great specialization 
in Huderma. In the upper jaw I’ and P! are reduced; in the lower the 
canine is reduced (though provided with a prominent accessory cusp), 
P; has been eliminated, and P, is single-rooted. The posterior portions 
of the tooth rows, on the other hand, are morphologically primitive in 
character: There is a trace of a hypocone cusp on M! and on M?, the 
metacone of M? is well developed, and P, has a metaconid cusp. With 
the exception of the elongated brain case and bifid median postpalatal 
prominence, other features of the cranium are primitive as far as the 
genera Huderma and Plecotus are concerned: The rostrum is weak, the 
supraorbital region is sharply ridged, the temporal ridges do not 
coalesce, and the zygomata are strong and are provided medially with 
a large postorbital expansion. 

The nostrils are unspecialized, retaining the primitive basic vesper- 
tilionid shape, as seen in Myotis and Prpistrellus. 

The evolutionary significance of the bold black and white color 
pattern of the fur is not known. Such a pattern crops up sporadically 
in the orders of the class Mammalia, but is notably rare in Chiroptera. 


AMERICAN BATS—HANDLEY 109 


Among the vespertilionids it is seen only in Huderma maculatum and in 
the rare African Glauconycteris superba (Hayman, 1939, 1946), which 
resembles Huderma maculatum in being black and white but has a more 
elaborate pattern. Hayman even suggested a similarity of the pattern 
of Glauconycterts superba to that of the spotted skunk, Spilogale. 
Some other species of Glauconycteris have a similar pattern but with 
subdued brown and buff tones rather than black and white. Scoto- 
manes ornatus of southeastern Asia has a pattern of white spots and 
stripes on the pelage, but the ground color is bright reddish rather than 
black. In some other forms, Kerivoula picta and Myotis formosus for 
example, the flight membranes, rather than the fur, are patterned 
with contrasting colors (Wroughton, 1912, p. 1195, pl. a). 

Despite numerous evidences of specialization, Huderma retains 
many primitive traits. It appears to occupy an evolutionary position 
below the more abundant and more progressive, although less special- 
ized, Plecotus, with which it shares its geographic range. An early 
derivative of the Euderma-Plecotus stock, Euderma maculatum has 
traveled a road of independent specialization to the point where it 
is possibly overspecialized. 

Genus Puecotus: From the study of Recent and fossil material, 
I infer that the hypothetical ancestor from which the subgenera 
Idionycteris, Plecotus, and Corynorhinus were derived probably had 
the following characteristics: Tooth rows not crowded; I' bicuspidate; 
I? simple, about two-thirds the height of I'; upper canine strong and 
exceeding P* in height; P! robust, considerably exceeding cingulum of 
canine in height; P* with full hypocone surface, a large anterior cingu- 
lum, and a protocone cusp; M! and M? without trace of a hypocone 
cusp; M?* with prominent metacone and a fourth commissure; lower 
incisor series strongly imbricated and increasing in size from J, to I;; 
lower canine equaling protoconid cusp of M, in height; P, much more 
robust than P;; P, double-rooted; lower molars with internal cusps 
almost equaling external cusps in height. Rostrum narrow and 
arched; anterior nares not enlarged; supraorbital region sharply 
ridged; temporal ridges not forming a sagittal crest; zygoma strong 
and expanded in middle third; basial pits not developed; median 
postpalatal prominence absent. Nostril with full cornu and lacking 
posterior elongation; muzzle glands not enlarged; auricle large, with 
a simple, complete anterior basal lobe, a small accessory anterior basal 
lobe, and transverse ribs not reaching the posterior margin of the 
auricle. 

From this generalized or primitive pattern, derivations, either in 
the form of reduction or simplification or in the form of elaboration, 
are to be regarded as traits of specialization or progressiveness. In 


110 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


no instance are these bats sufficiently well known to determine the 
teleological significance of these traits. 

The differences distinguishing the subgenera Idionycteris, Plecotus, 
and Corynorhinus mainly concern the external configuration of the 
nostril, muzzle, and ears (table 2). Dental, cranial, and skeletal 
differences are numerous but slight (table 3). In the following dis- 
cussion, Plecotus townsendw is given as a typical example of the 
subgenus Corynorhinus. Unless otherwise noted, statements regard- 
ing it apply equally to the other Recent species of the subgenus, 
P. mexicanus and P. rafinesquii. The subgenera Plecotus and Idio- 
nycteris are monotypic, represented respectively by the species P. 
auritus and P. phyllotis. Inasmuch as P. phyllotis is known to me 
only from three specimens, statements concerning it must be regarded 
as highly tentative. 

The dental formula is the same in the three subgenera, and, as may 
be seen from the following comparisons, the individual teeth must be 
closely scrutinized in order to detect differences. 

The secondary cusp of I! is absent or variable in P. townsendii, 
usually present in P. mexicanus, and always present in P. rafinesquit, 
P. phyllotis, and P. auritus. The space between IJ? and the canine is 
frequently, but not always, greater, and P! averages larger in P. auri- 
tus than in the other species. FP* is consistently wider than long in 
P. phyllotis and P. townsendi; longer than wide in P. auritus. Be- 
cause the metacone of M?® is somewhat reduced in P. auritus, the 
third commissure of M? is equal to or shorter than the second in most 
individuals, whereas in P. phyllotis and P. townsendii the third com- 
missure is normally equal to or longer than the second. P, is usually 
more robust in P. auritus and P. phyllotis than in P. townsendii, 
considerably exceeding P; in size. P, is double-rooted in P. auritus; 
single-rooted in P. phyllotis and P. townsendii. In P. phyllotis the 
internal cusps of the mandibular molars are almost as high as the 
external cusps; in the other species of Plecotus the internal cusps are 
lower. 

An analysis of these observations on dentition reveals four features, 
which, because each is consistent in at least one of the subgenera, may 
be regarded as having primary taxonomic significance. These fea- 
tures are the simplification in I’, M?, and P, respectively, and the 
specialization in P*. P. phyllotis exhibits the generalized (unsimpli- 
fied and unspecialized) condition in three features, is specialized in 
one (P,); whereas P. auritus is specialized in two (P* and M?); and 
all of the species of the subgenus Corynorhinus are specialized in one 
(P,) and one of the species in another (I'). Three other features— 
crowding of the upper incisor series and diminution of the size of P! 
and P;—are inconsistent, but may be of some significance. In these 


AMERICAN BATS—HANDLEY 


Euderma 


Median postpalatal process a bifid 
prominence. 


Auditory bullae roughly elliptical 
in outline. 


Brain case relatively long: post- 
palatal length averages 40-41% 
of greatest length. 

Presternum longer than wide. 

Upper incisors stand in line with 
toothrow. 

I! almost as small as I’. 

P! minute; not exceeding cingulum 


of canine in height. 


Hypocone cusp barely indicated in 
M! and M?. 


Lower canine small; not exceeding 
P, in height. 


Anterointernal cusp of lower ca- 
nine almost equal to primary cusp 
in height. 


TasBLE 1.—Morphological characters distinguishing Euderma from Plecotus 


Plecotus 


Median postpalatal process absent 
or a simple spine. 


Auditory bullae roughly circular in 
outline. 


Brain case relatively short: post- 
palatal length averages 34-38% 
of greatest length. 


Presternum wider than long (un- 
known in Plecotus phyllotis). 


Upper incisors stand inside tooth- 
row. 


I! a third higher and twice as wide 
as J?, 


P! larger; exceeding cingulum of 
canine in height. 


Hypocone cusp not indicated in M! 
and M?. 


Lower canine large; much exceeding 
P, in height. 


Anterointernal cusp of lower canine 
small and greatly exceeded by pri- 
mary cusp. 


P; absent. 


P, with well defined metaconid 
cusp. 


Tragus lacking basal lobe. 


Tragus united with posterior basal 
lobe of auricle. 


Coloration black and white in bold 
pattern. 


497256—59——__2 


P3 present. 


P, lacking metaconid cusp. 


Tragus with prominent basal-lobe. 


Tragus and posterior basal lobe of 
auricle not united. 


Coloration drab brownish, essen- 
tially without pattern. 


111 


112 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 
TaBLE 2.—Comparison of external characters of the subgenera of Plecotus (Idionyc- 


teris most generalized, Corynorhinus most specialized, Plecotus intermediate) 


Idionycteris 


Nostril unspecialized. 


Muzzle glands not en- 
larged (?). 


Auricle with anterior 
basal lobe complete. 


Accessory anterior basal 
lobe of auricle devel- 
oped into a projecting 
lappet. 


Transverse ribs on auri- 
cle fade out before 
reaching posterior 
border. 


Second phalanx of third 
digit longer than first 
phalanx. 


Calear keeled. 


Interfemoral membrane 
attached to tip of last 
caudal vertebra. 


Plecotus 


Nostril with posterior 
elongation; cornu re- 
duced. 


Muzzle glands slightly 


enlarged. 


As in Idionycteris. 


Accessory anterior bas- 
al lobe of auricle 
slightly developed. 


Transverse ribs on au- 
ricle run without in- 
terruption to poste- 
rior border. 


Second phalanx of third 
digit equal to or 


shorter than first 


phalanx. 
Calcar not keeled. 


Interfemoral mem- 
brane attached to 
base of last caudal 
vertebra. 


Corynorhinus 


Nostril with posterior 
elongation; cornu 
absent. 


Muzzle glands greatly 
enlarged. 


Auricle with anterior 
basal lobe reduced. 


Accessory anterior 


basal lobe of auricle 
absent. 


Transverse ribs on au- 
ricle interrupted by 
vertical rib near 
posterior border. 


As in Idionycteris. 


As in Plecotus. 


As in Idionycteris. 


features P. phyllotis is generalized in one (P,) and is dubiously special- 
ized in the others; P. auritus is generalized in all; and P. townsendit 
is specialized in all. 

Likewise, in cranial details specialization is greatest in P. town- 
sendii, least in P. phyllotis (fig. 4). In the more primitive species, P. 
phyllotis and P. auritus, the supraorbital region is sharply ridged; in 
P. townsendu it is smoothly rounded [also in the Lower Pleistocene 
P. crassidens; faintly ridged in the Middle Pleistocene P. (Coryno- 
rhinus) alleganiensis]. The temporal ridges converge to form a median 
sagittal crest in P. auritus and P. townsendii, but remain apart in P. 
phyllotis and to a lesser degree in P. alleganiensis. 


AMERICAN BATS—HANDLEY 


TABLE 3.—Comparison of crania of the subgenera of Plecotus (Idionycteris most 


generalized, Corynorhinus most specialized, Plecotus intermediate) 


Idionycteris 


Supraorbital region 
sharply ridged. 


Zygoma relatively thick 
and strong; postorbi- 
tal expansion in mid- 
dle third of arch. 


Median postpalatal 
process absent. 


Basial pits absent. 


Brain case relatively 
shallow: averages 
33% of greatest 
length. 


Brain case relatively 
broad: averages 53% 
of greatest length. 


Rostrum flattened, with 
median concavity. 


P4 wider than long. 

Metacone of M? not re- 
duced; 3rd commis- 
sure equal to or long- 


er than 2nd. 


P, single-rooted. 


Plecotus 


As in Idionycteris. 


As in Idionycteris. 


Median postpalatal 
process a poorly de- 
veloped spine. 


As in Idionycteris. 


Brain case relatively 
shallow: averages 
32% of greatest 
length. 


Brain case relatively 
narrow: averages 
49% of greatest 
length. 


Rostrum arched, lack- 
ing median concav- 


ity. 
P4 longer than wide. 
Metacone of M? re- 
duced; 8rd commis- 


sure usually equal to 
or shorter than 2nd. 


P, double-rooted. 


Corynorhinus 


Supraorbital region 
smoothly rounded 
or faintly ridged. 


Zygomarelatively thin 
and fragile; post- 
orbital expansion in 
posterior third of 
arch. 


Median postpalatal 
process a prominent 
spine. 

Basial pits prominent. 

Brain case relatively 
deep: averages 35- 
38% of greatest 
length. 

Brain case relatively 
narrow: averages 
48-50% of greatest 
length. 


As in Idionycteris. 


As in Idionycteris. 


As in Idionycteris. 


As in Idionycteris. 


113 


Broadening and anterior bulging of the brain case in P. phylilotis 
and increase in depth of the brain case in P. townsendii may be re- 
garded as specializations. Shortening, broadening, and flattening of 
the rostrum together with enlargement of the anterior nares in P. 
townsendu are specializations that are somewhat paralleled in P. 


114 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


NU 


PLECOTUS (GORYNORHINUS) RAFINESQUI!I 


USNM 297218 LATERAL 
LOBE 
VENTRAL 
LOBE 
POSTERIOR 
PLECOTUS (CORYNORHINUS) TOWNSENDII LOBE 
CM 18760 


ae Sf 


PLEGOTUS (PLEGOTUS) AURITUS 
USNM 18487 


Figure 5.—Comparison of the presternum of three species of Plecotus. Ventral (left) and 
lateral (right) aspects. Camera lucida sketches. 


phyllotis, although the anterior nares are not enlarged in the latter 
species. 

The zygomatic arches show progressive reduction and weakening 
through the series from relatively heavy and strong in P. phyllotis 
and P. auritus to relatively light and weak in P. townsendii. Also, 
in the latter species there is a shift of the postorbital expansion of the 
zygoma, from its primitive median position to a posterior location. 

The median postpalatal process, absent in P. phyllotis, is slightly 
developed in P. auritus, and is a prominent spine in P. mexicanus and 
in some forms of P. townsendw. The lateral grooves (‘‘basial pits’’) 
commencing in the basioccipital and ending abruptly in the basisphe- 
noid are prominent in P. townsendvi; they are ill-defined or absent in 
the more primitive P. phyllotis and P. auritus. Slight elongation of 
the auditory bullae is a specialization peculiar to P. phyllotis. The 


AMERICAN BATS—-HANDLEY 115 


typical vespertilionid bullar shape, as seen in P. auritus and P. town- 
sendii, is roughly circular. 

The postcranial skeleton is unknown in P. phyllotis and differences 
in this respect between P. auritus and P. townsendii are few. In the 
latter species the proximal articular surface of the radius is expanded, 
and the presternum has the ventral lobe enlarged and the posterior 
lobe laterally expanded (fig. 5). P. auritus has departed from the 
usual phalangeal pattern of Plecotus and EHuderma, in which the second 
phalanx of the third digit is longer than the first (by a slight elongation 
of the first phalanx and a considerable shortening of the second), 
rendering not only the proportion different but the combined length 
less (fig. 6). The calcaral keel, present in P. phyllotis, is lacking in 
the other species of Plecotus. 

Hamilton (1949, p. 100) pointed out distinctions between the 
bacula of P. rafinesquu and P. auritus (as described and figured by 
Matthews, 1937, p. 222). However, intrageneric discrepancies of 
similar magnitude seem also to exist among the several species of 
Myotis, Prpistrellus, and Lasiurus as described and figured by these 
same authors. Krutzsch and Vaughan (1955, p. 99) described the 


PLECOTUS (CORYNORHINUS) 
USNM 297218 


ae 
II 
4 
4 
Ce 
7 
7 Pd 
PLEGOTUS (PLEGOTUS) AURITUS al: f 
USNM 18487 7 
7 7 
2 7 
oO TTS 
Vv Iv 


Figure 6.—Pectoral appendages of Plecotus, showing relative proportions of first and 
second phalanges of third digit. 


RAFINESQUI! 


116 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


baculum of P. townsendw as being similar to that of P. rafinesqutt. 
Pearson, Koford, and Pearson (1952, p. 281) noted that the subgenera 
Plecotus and Corynorhinus “possess in common two pairs of Cowper’s 
glands, a feature not found in other vespertilionids. Likewise, 
seminal vesicles apparently are lacking in both of these genera [sub- 
genera].” 

Of all the morphological features distinguishing the species of 
Plecotus, those of the muzzle and external ears have been supposed to 
be of the greatest taxonomic significance. 

Enlarged muzzle glands in P. townsendw protrude as grotesque 
lumps two to three millimeters high on either side of the muzzle 
(fig. 7). There are similar glandular areas on the muzzle of P. auritus, 
but they do not project as in P. townsendii. They may not be con- 
spicuous in P. phyllotis; there is no external trace of them on three dry 
study skins. Dalquest and Werner (1954, p. 156) have observed that 
the facial glands of P. rafinesquii are histologically similar to those of 
other vespertilionids, particularly Myotis and Pipistrellus. The 
prominent muzzle glands (or ‘‘nose lumps’’) are solid masses of skin 
covered sebaceous glandular tissue. There are in addition, in 
the facial area, nests of sebaceous glandular cells, sudoriferous glands, 
and a sublingual gland of the mucous type. The submaxillary gland 
apparently has been forced out of the facial area by the extensive 
development of the other glandular tissues. 

A peculiar posterior elongation of the nostril in P. auritus gives the 
nostril a crescentic shape (fig.8). The cornu surrounding the narial 
opening is slightly reduced from the primitive vespertilionid condition, 
as seen in Myotis and Pipistrellus, and there is a conspicuous shallow 
basin extending posteriad from the posterolateral corner of the 
opening and separated from it by a septum. A similar posterior 
elongation of the nostril in P. townsendii is partially concealed by the 
overlapping muzzle glands. In this species the cornu is reduced to 
the point of obliteration, and the narial opening is squarish in outline 
and is bordered by smooth pale-colored skin resembling the lining of 
the nostril. As in P. auritus, the narial opening proper and the 
posterior elongation are separated by a septum. In the available 
specimens of P. phyllotis the nostrils appear to approximate the 
primitive vespertilionid type, with full cornua and without posterior 
elongations. 

The snout is truncate in P. auritus and P. townsendvi rather than 
acute as in Huderma maculatum and most other vespertilionines (pos- 
sibly acute in P. phyllotis). 

A lappet projecting over the forehead from the anterior base of the 
auricle is the chief characteristic of P. phyllotis. This appendage is 
an enlargement of the accessory anterior basal lobe of the auricle (fig. 


AMERICAN BATS—HANDLEY oul WA 


9), a lobe that is present, although small and inconspicuous, in P. 
auritus but absent in P. townsendw. Modification also occurs in the 
primary anterior basal lobe, where variation ranges from reduction in 
P. townsendii to no modification in P. phyllotis and enlargement in 
P. auritus (fig. 9). 

Thus, with regard to the dental, osteological, and external charac- 
teristics of these bats, it appears that P. (dionycteris) phyllotis is the 
most generalized and primitive member of the group; P. (Coryno- 
rhinus) townsend is most specialized and progressive; and P. (Plecotus) 
auritus is intermediate (but nearer P. townsendir). 

Comparison of Plecotus with other vespertilionine genera confirms 
the observation that the differences among its species are, at most, 
of no more than subgeneric magnitude. Myotis, Pipistrellus, and 


PLEGOTUS (CORYNORHINUS) 


TOWNSENDII 
USNM 148550 


ra 
PLEGOTUS (PLEGOTUS) AURITUS 


USNM 8660I 


MYOTIS VEL 
USNM 102472 


Ficure 7.—Muzzles of Plecotus, compared in anterolateral aspect with the muzzle of 
Myotis, a generalized vespertilionid. Camera lucida sketches. 


118 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Eptesicus are not suitable for such a comparison because of the prob- 
ability that they are polyphyletic aggregations, and not natural 
groups. Most of the other vespertilionine genera are either monotypic, 
consist of slightly differentiated species, are of disputed validity, or 
are not sufficiently well known. The American genus Lasiurus is a 
notable exception, being well known, widely distributed, and contain- 
ing several well-differentiated species. Included in the genus are the 
red bats (LZ. borealis, etc.), hoary bats (L. cinereus, etc.), and yellow 
bats (ZL. ega, etc.). The magnitude of the differences distinguishing 
these species equals or exceeds the magnitude of the differences dis- 
tinguishing the species Plecotus phyllotis, P. auritus, and P. townsendii 
(compare tables 2, 3, and 4). 

Stratus oF Genus BarpasteLua: Miller (1907, p. 224) pointed out 
the likeness of Barbastella to EHuderma and Plecotus, whereas Tate 
(1942, p. 230) tried to show that any relationship between Barbastella 
and these genera isremote. Tate believed that Huderma and Plecotus 
are derivatives of the Myotis stem, and he supposed that Barbastella 
came from the Pipistrellus stem. 

The strongest resemblances of Barbastella to Pipistrellus are in 
tooth formula, and in the posterior shortening of the palate, which 
extends only a short distance past M*. Like Pipistrellus, but also 
like Myotis as well, Barbastella has small, relatively simple auditory 
bullae; fragile, much-reduced zygomata without postorbital expan- 
sions; and a short angular process of the mandible. The few resem- 
blances of Barbastella to Pipistrellus and its relatives seem to lose 
much of their significance, however, in view of the many similarities 
of Barbastella to Euderma and Plecotus: 

. Rostrum relatively narrow and weak. 

. Brain case elongated and vaulted in frontal region. 

. Zygomata (in dorsal aspect) parallel and not bowed out posteriorly. 
. Coronoid process of mandible high. 

. Dentition weak. 

. Cusps of teeth high and sharp. 

Lower incisors graduated in size (I; large to I, small). 

. I; with prominent extra internal cusp (total of 4 cusps). 


. Lower canine with prominent anterointernal basal cusp. 
. Pg not so high as M,. 


SCHOANOORWN HE 


— 


The resemblance of the teeth in Barbastella, Euderma, and Plecotus 
is particularly striking. In shape and proportions, the teeth of these 
genera are almost identical. ‘The facts that one of the upper pre- 
molars (P!) has been crowded out of the tooth row and that one of 
the lower premolars (P;) has been eliminated in Barbastella are not 
sufficient in themselves to indicate that Barbastella is closely related 
to Pipistrellus. (Note that Huderma also has the same dental for- 
mula as Pipistrellus.) 


AMERICAN 


BATS—HANDLEY 


119 


Tasue 4.—Morphological characteristics of bats of the genus Lasiurus 


red bats 
(L. borealis, ete.) 


Size small (forearm 
37-44 mm.) 


Lateral wings of pre- 
sternum equal to 


body of presternum 
in width. 


Presternum about as 
long as wide. 


Auditory bullae not 
enlarged. 


Rostrum relatively 
short. 


Sagittal crest very 
weak. 


Coronoid process 
medium height. 


P! usually present. 


Hypocone slightly re- 
duced on M! & M?., 


M3 very reduced. 
P, double-rooted. 


M; talonid reduced. 


hoary bats 
(L. cinereus, etc.) 


Size large (forearm 
46-57 mm.) 


yellow bats 
(L. ega, etc.) 


Size large (forearm 
44-57 mm.) 


Lateral wings of pre- 
sternum equal to 
body of presternum 
in width. 


Presternum much 
longer than wide. 


Auditory bullae 
slightly enlarged. 


Rostrum medium. 


Sagittal crest weak. 


Coronoid process low. 


P! usually present. 


Hypocone much re- 
duced on M! & M?. 


M3 reduced. 
P, single-rooted. 


M; talonid only 
slightly reduced. 


Lateral wings of pre- 
sternum consider- 
ably broader than 
body of prester- 
num. 


Presternum about as 
long as wide. 


Auditory bullae not 
enlarged. 


Rostrum relatively 
long. 


Sagittal crest strong. 

Coronoid process 
high. 

P! always absent. 


Hypocone slightly re- 
duced on M! & M2. 


M3 slightly reduced. 
P, double-rooted. 


M; talonid much re- 
duced. 


Barbastella is unique in the degree of posterior extension of the 
anterior nares, which reach so far back that the vomer is exposed 


as a Median spine. 


the subgenus Corynorhinus. 


Its muzzle glands are almost as large as those in 
A pair of deep grooves extend from the 


nostrils to the upper lip. The auricles are peculiarly short and broad 


and are connected across the forehead. 


In each there is a conspicuous 


subapical lobe on the posterior border, and the posterior basal lobe 
is attached nearer the angle of the mouth than in Luderma or Plecotus. 


120 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


PLEGOTUS (CGORYNORHINUS) 
TOWNSEROI 

USNM 149550 

(RIGHT MUZZLE GLAND 


REMOVED TO EXPOSE FULL Is nae, 
EXTENT OF POSTERIOR 


aia a Sooty 


MUZZLE GLAND 


PLECOTUS (PLEGOTUS) 
AURITUS 


POSTERIOR ELONGATION 


USNM 16386 SEPTUM 


NARIAL OPENING 


CORNU 


MYOTIS VELIFER 
USNM 102472 


Ficure 8.—Muzzles of Plecotus, compared in anterodorsal aspect with the muzzle of 
Myotis, a generalized vespertilionid. Camera lucida sketches. 


The tragus is relatively long, is broad at the base, and tapers abruptly 
to a pointed tip. 

Barbastella is a relative of Huderma and Plecotus that lacks the ex- 
treme auditory specialization characterizing these genera. In this 
respect, Barbastella must be more primitive than Huderma and Plecotus. 
On the other hand, in most cranial details Barbastella is similar to 
Euderma and Plecotus, and in some features such as fragile zygomata, 
large anterior nares, posterior nares lying far forward, and short tooth 
rows it has surpassed them in degree of specialization. 

If the assumption is correct that Barbastella is closely related to 
Euderma and Plecotus, then it follows that most of the cranial char- 
acters shared by the three genera must have been established before 
auditory specialization occurred. Further, the Barbastella stock must 


AMERICAN BATS—HANDLEY 1 PA 


PLEGOTUS (CORYNORHINUS) 


TOWNSENDI 
USNM 148550 


PLEGOTUS (PLEGOTUS) AURITUS 
USNM 86667 


TRANSVERSE 
RIBS 


ANTERIOR BASAL 
LOBE 


TRAGUS ACCESSORY ANTERIOR 


BASAL LOBE 
POSTERIOR 
BASAL LOBE CONNECTING 
MEMBRANE 


PLECOTUS (IDIONYCTERIS) 
PHYLLOTIS 
AMNH 62260 


MCZ 5943 


Ficure 9.—Variation in the auricle and its appendages in Plecotus. Free-hand sketches. 


have been derived very early and had a long subsequent period of in- 
dependent development and specialization. 


Systematic Treatment 


Key to the genera Euderma and Plecotus 


Posterior basal lobe of auricle attached to base of tragus; auditory bulla elongated 

| (elliptical in outline); I! almost as small as I?; hypocone cusp indicated on M! 
and M?; lower canine small, but its anterointernal cusp greatly enlarged; Ps 
| absent; P4 with well defined metaconid cusp; coloration black and white in 
j bold pattern... . . . . . Euderma (p. 122) 
) Posterior basal lobe of Ruricle Renarared for irae by a notch; auditory bulla 
not elongated (circular in outline); I’ a third higher and twice as wide as [?; 

| hypocone cusp not indicated on M! and M?; lower canine large, but its antene 
internal cusp small; P; present; P, lacking metaconid cusp; coloration drab 
| brownish, essentially without pattern. ...... . . . Pleeotus (p. 127) 


P22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Genus Kuderma H. Allen 


Histiotus J. A. Allen, 1891, p. 195. (Part.) 
Euderma H. Allen, 1892, p. 467. 

Type species: Histiotus maculatus J. A. Allen. 

DistriBuTIon: Southwestern United States and probably north- 
western Mexico. 

Description: Supraorbital region sharply ridged (fig. 4); temporal 
ridges not coalescing posteriorly to form a sagittal crest; brain case 
exceptionally elongated; zygoma relatively heavy, with postorbital 
expansion in middle third of arch; median postpalatal process a bifid 
prominence; auditory bulla roughly elliptical in outline; presternum 
longer than wide. 

Upper incisors in line with tooth row; I’ simple, without accessory 
cusps (fig. 2), except occasionally near cingulum, and small, only 
slightly larger than I’; I? close to or touching upper canine; P! minute; 
hypocone cusp barely indicated on M! and M?; third commissure 
longer than second and metacone relatively well developed in M?; 
lower canine reduced in thickness, not exceeding P, in height, and 
possessing a prominent anterointernal secondary cusp almost equaling 
primary cusp in height; P; about one-half the size of P,, and not 
crowded; P; absent; P, single-rooted and possessing a well defined 
metaconid cusp. 

No large glandular masses on muzzle (fig. 7); nostril opening 
small, of primitive vespertilionid type (fig. 8), provided with a cornu, 
and not extended posteriorly or opening upward; auricle and tragus 
very large (fig. 9); tragus not provided with a basal lobe, but united 
with posterior basal lobe of auricle; anterior basal lobe of auricle 
complete; transverse ribs on auricle extend to posterior border of 
auricle; second phalanx of third digit longer than first phalanx (fig. 6); 
calcar not keeled; interfemoral membrane attached to base of last 
caudal vertebra; coloration black and white in bold pattern; mem- 
branes and ears grayish. 


Euderma maculatum J. A. Allen 


Histiotus maculatus J. A. Allen, 1891, p. 195. 
Euderma maculata H. Allen, 1894, p. 61. 

Hototypr: AMNH 3920/2991; young adult skin (skull lost sub- 
sequent to original description) ; ‘“‘caught on a fence” in March 1890, 
by Thomas Shooter; near Piru, Ventura County, Calif. [probably 
mouth of Castac Creek, Santa Clara Valley, 8 miles east of Piru, 
Los Angeles County, Calif. (Miller, 1897, p. 49)]. 

DistRiBuTION: Southwestern United States and probably north- 
western Mexico (fig. 10). West to Washoe County, Nev.; Mariposa, 


AMERICAN BATS—HANDLEY W3 


Kern, and Los Angeles Counties, Calif. South to Riverside County, 
Calif.; Yuma County, Ariz.; (?Magdalena, Sonora?); Dona Ana 
County, N. Mex. East to Dona Ana County, N. Mex.; San Juan and 
Salt Lake Counties, Utah; Yellowstone County, Mont. North to 
Yellowstone County, Mont.; Canyon County, Idaho; Washoe County, 
Nev. Zonal range: Lower Austral to Transition. Altitudinal range: 
189 feet below sea level in Riverside County, Calif., to 4,045 feet in 
Mariposa County, Calif. Only 16 specimens are known to be 
preserved in museums. 

Description: As indicated above. Adult coloration: Upper parts 
blackish with large, roughly circular, white spots on the shoulders 


Figure 10.—Distribution of Euderma maculatum J. A. Allen. Solid symbols, specimens 
in museums; open symbols, other records; encircled solid symbol, type locality. 


124 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


and at the base of the tail, and patches of cottony white hair at 
the posterior base of each auricle; all hairs black at base, except 
those at posterior bases of auricles. Underparts with hairs white 
tipped, the white incompletely concealing the black bases. Membranes 
pale grayish brown, auricles and tragi yellowish brown. 

MEASUREMENTs: See tables 7, 13. 

Remarks: USNM 269842, not reported previously, was captured 
9 miles east of Roll, Yuma County, Ariz., on 5 August 1940, and was 
submitted to the National Museum by Mrs. W. A. Jones, Box 3, Roll, 
Ariz. It is an adult, probably a male, mummified, with the skull 
still inside the skin. In Mrs. Jones’ words, the details of its capture 
were as follows: 

Dear Sirs. Am sending a bat of a kind I haven’t seen before. ... The 
bat was captured the fifth of August, here at my home. It was a very hot day, 
and the bat kept coming and lighting on the screened porch. My son [Weldon 
Allen, Jr.], nine years old, caught it and brought itin the house. We gave it water. 
It was very thirsty. It lived about three hours after catching it. ... I wish 
you could have seen the bat when it was alive. It was a pretty thing in an odd 
way. 

Hasirat: Huderma has seldom been collected or observed alive by a 
mammalogist, despite the fact that it inhabits one of the most in- 
tensively collected areas of the world. At least five of the records for 
it can be attributed to small boys, and many of the other records stem 
from persons neither scientists nor naturalists. Use of mist nets and 
other trapping devices has failed to produce specimens. 

Euderma appears to be an inhabitant primarily of open or scrub 
country. There is only one record of occurrence in a forest, in 
Yosemite Valley, Calif. (Parker, 1952, p. 480); however, Vorhies 
(1935, p. 225) quoted Herbert Brown as stating that Arizona Indians 
assured him it was a forest dweller. Usually it has been found in or 
on houses—in biological laboratory (Miller, 1903, p. 165; Hall, 1935, p. 
148) ; in residence (Nicholson, 1950, p. 197); on porch (Hardy, 1941, p. 
293; Mrs. W. A. Jones, 1940, in litt.; Benson, 1954, p. 117); under 
eaves of house (Durrant, 1935, p. 226; Ashcraft, 1932, p. 162); clinging 
to side of brush house (Vorhies, 1935, p. 225); in driveway near house 
(Tucker, 1957, p. 406). Less frequently it has been observed in caves 
or cave-like structures—in moist cave (Hardy, 1941, p. 293); in 
springhouse dug into hillside (Parker, 1952, p. 481); in mine tunnel 
(Vorhies, 1935, p. 225); in root cellar (Hall, 1946, p. 158). One that 
was being attacked by yellow jackets when found perhaps had tried 
to creep into a crevice or tree hole where these insects nested (Parker, 
1952, p. 480). Several specimens have been in or near water— 
floating in puddle, the overflow from a railway water tank (J. Grinnell, 
1910, p. 317); floating in shallow reservoir (Benson, 1954, p. 117); 


AMERICAN BATS—HANDLEY 125 


damp and bedraggled, one foot from river’s edge (Parker, 1952, p. 
480); hanging from rock projection in closed springhouse (Parker, 
1952, p. 481); hibernating above large pool of water in cave (Hardy, 
1941, p. 293). One was found hanging [impaled?] on a fence (J. A. 
Allen, 1891, p. 195) and one with injured wing was found clinging to 
the lower side of a rock at the base of a cliff (Hall, 1939, p. 103). 

Five of the specimens were dead, two were obviously injured, and 
all but one of the remainder died within a few hours after capture. 
Several that were obtained alive showed great thirst for water, and at 
least 10 of the live individuals were exposed to daylight where found. 
Of the 22 recorded occurrences, 13 were about houses. Six of the 13 
were clinging to the outside of the structures. 

The normal habitat of this species is unknown. The foregoing facts 
suggest departures from its normal habitat in response to stimuli of 
rather frequent occurrence. A narrow tolerance in habitat selection 
by the spotted bat could explain the lack of natural collections and 
the frequency of abnormal ones. Suppose that the bat’s habitat were 
very restricted and were such that it might frequently change to the 
bat’s disadvantage. Suppose for instance that Huderma maculatum 
requires absolute privacy from other bats and requires moist places 
for roosting in its normally arid surroundings. This might restrict it 
largely to moist crevices, which because of their limited extent might 
occasionally dry up and force the bat out, to alight, thirsty and dying 
from desiccation, on porches, sides of houses, in driveways, etc. 

The spotted bat fits very well Amadon’s description (1953, p. 464), 
of relict species: 

Relicts are species that tend to become more and more restricted both geo- 
graphically and ecologically, because they are unable to compete successfully with 
other species. . . . On continents, relicts often survive by becoming more and 


more specialized, through a process of natural selection, to a narrow ecological 
niche where they do manage to retain a competitive advantage. 


Bruavior: Most recorded individuals appear to have been solitary 
wanderers. Although Huderma maculatum has been observed at all 
seasons, seven of the 17 occurrences for which the date is recorded are 
for the months of August, September, and October, the period of post- 
breeding season wandering characteristic of many bats. Only at 
Yosemite Valley, Calif., where two specimens were collected 20 years 
apart, has more than one specimen been secured, although Hardy 
(1941, p. 293) was told that four had been found hibernating in a 
Utah cave. C. Hart Merriam was told by ranchers in the Vegas 
Valley, Nev., that “a very large bat ‘with ears like a jackass and a 
white stripe on each shoulder’ is abundant in that place in the summer” 
(Miller, 1897, p. 49). 

Two reports suggest possible association of Huderma with other 


126 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


species of bats. Vorhies (1935, p. 225) quoted Mexican informants as 
saying “‘there were many bats in the drifts [side shafts of a mine] and 
among them was the one spotted one.”” Hardy (1941, p. 293) observed 
Myotis subulatus and Plecotus townsendii in a Utah cave in which 
Euderma had been reported. 

Apparently Huderma rests suspended by the feet with head down 
(Parker, 1952, p. 480); an informant of Hardy (1941, p. 293) reported 
hibernating individuals suspended by the thumbs, with head up. 
Parker (1952, p. 480) recorded that a captive specimen walked with 
wrists and feet, and could thus travel over horizontal surfaces with 
considerable facility. Durrant (1935, p. 226) was told that a captive 
moved about its cage in pursuit of flies held with forceps. 

When the bat is alert the normal carriage of the ear is erect, directed 
slightly forward, with tip bent slightly back. When disturbed, the 
bat folds ears, but not the tragus, down against the side of the neck, 
much in the fashion of Plecotus townsendit (Parker, 1952, p. 480). 

Some individuals were docile, with little inclination to resist handling 
(Parker, 1952, p. 481), but one was said to have “had plenty of fight’’ 
when molested (Hall, 1939, p. 103). One was being attacked by one 
or more ‘yellow jackets,” and had sustained injury to the humeral 
areas of both wings when it was found. It was attempting to fend off 
the insects with its feet (Parker, 1952, p. 480). 

The voice of this bat has been described as a “soft, extremely high- 
pitched, metallic squeak”’ (Parker, 1952, p. 481); a hissing noise and 
rat-like squeak (Hall, 1939, p. 103); and a typical bat chirp (Durrant, 
1935, p. 226). Also, it clicks its teeth together (Hall, 1939, p. 103) 
and makes ‘‘a queer grinding noise by gnashing its teeth” (Ashcraft, 
1932, p. 162). 

The report of an individual that flew through an open door or win- 
dow into a house at about 11 p.m. (Nicholson, 1950, p. 197) indicates 
that Huderma, like Plecotus auritus, may be nocturnal rather than 
crepuscular. 

Foop: Although several captive individuals eagerly accepted water 
to drink, few showed interest in various types of food. One ate cottage 
cheese (Parker, 1952, p. 481). Another, force-fed at first, later readily 
took flies offered with forceps. It accepted five flies at each of two 
daily feeding periods. When flies were offered it wrinkled up its nose, 
chirped, and advanced as rapidly as possible to seize the fly. It also 
eagerly pursued empty forceps. This individual lived for three weeks 
in captivity. Its death was accidental (Durrant, 1935, p. 226). Food 
habits under natural conditions are not known. 

Data on migration, hibernation, reproduction, and molt are lacking 
for this species. 

Loca NAMEs: ‘“‘Spotted”’ bat is the name in most general usage, 


AMERICAN BATS—HANDLEY 127 


although Hardy (1941, p. 293) referred to the use of the name “‘pinto”’ 
bat in Utah, and Vorhies (1935, p. 225) reported that Mexicans knew 
it as ‘‘pinto” bat in Sonora. Nevadan ranchers likened its ears to 
those of a jackass (Miller, 1897, p. 49) and Grinnell (1910, p. 318) 
likened its dorsal color pattern to a “‘death’s-head.”’ Two boys who 
discovered one on the side of a brush house mistook it, at first glance, 
for a young rabbit (Vorhies, 1935, p. 225), while a boy who saw one 
crawling on the ground first thought it was a tarantula spider (Parker, 
1952, p. 480). People who saw one with its ears folded back thought 
it was “some sort of horned beast’”’ (Parker, 1952, p. 480). 
SPECIMENS EXAMINED: Five, from the following U.S. localities: 
New Mexico: Dona Ana County: Mesilla Park, la (USNM). Arizona: 
Yuma County: 9 miles east of Roll, ls (USNM); 4 miles south of Yuma, Is 


(MCZ). Cautrornia: Los Angeles County: Mouth of Castac Creek, 8 miles east 
of Piru, ls (AMNH). Uran: Salt Lake County: Salt Lake City, Is (UU). 


ADDITIONAL RECORDS (asterisk indicates specimen preserved in a 
museum) : 

MEXICO: Sonora: Near Magdalena (Vorhies, 1935, p. 225). 

UNITED STATES: Arizona: Maricopa County: Tempe* (Stager, 1957, p. 
260). Catirornia: Kern County: Red Rock Canyon*, 30-40 miles north of 
Mohave (Hall, 1939, p. 103). Mariposa County: Yosemite National Park’, 
Yosemite Valley, 4,045 feet, 2 specimens (Ashcraft, 1932, p. 162; Parker, 1952, p. 
480). Riverside County: Mecca*, 189 feet below sea level (J. Grinneil, 1910, 
p. 317). San Bernardino County: China Lake*, 4 miles south of Inyo County line 
(Stager, 1957, p. 260); Granite Wells, near Pilot Knob, Barstow area (Parker, 
1952, p. 481); Twentynine Palms* (Benson, 1954, p. 117). NEevapa: Clark 
County: Vegas Ranch, Vegas Valley (Miller, 1897, p. 49). Esmeralda County: 
Fish Lake Valley (Hall, 1946, p. 158). Washoe County: Reno* (Hall, 1935, p. 
148). Uraun: Garfield County: Hatch (Hardy, 1941, p. 293). Kane County: 
Crocodile Cave, 4 miles north of Kanab (Hardy, 1941, p. 293). San Juan County: 
5 miles northwest of Monticello* (Benson, 1954, p.117). Ipano: Canyon County: 
15 miles southwest of Caldwell* (Tucker, 1957, p. 406). Montana: Yellowstone 
County: Billings* (Nicholson, 1950, p. 197). 


Genus Plecotus EK. Geoffroy Saint-Hilaire 


The synonymy is given under the subgenera. 

Types specizs: Vespertilio auritus Linnaeus. 

Distripution: Temperate parts of North America, Eurasia, and 
northern Africa (Palaearctic and Nearctic regions) (fig. 11). 

Description: Supraorbital region sharply ridged or smoothly 
rounded (fig. 4); temporal ridges remain apart or coalesce to form 
sagittal crest; brain case relatively short; zygoma relatively heavy or 
light, with postorbital expansion in middle or posterior third of arch; 
median postpalatal process absent or a spine-like prominence; auditory 
bulla roughly circular in outline; presternum wider than long. 

497256—59 3 


128 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


E 7 , 
Ficure 11.—Distribution of Plecotus, showing coincidence with the North Temperate Zone 


(here defined as a region having an average annual temperature between 35° and 


70° F.). 


Upper incisors not in line with remainder of tooth row (lingual); 
I’ simple or bifid (fig. 2), much larger than I’; I? not touching canine; 
P! small; hypocone absent and protocone reduced posteriorly in M? 
and M?; size of third commissure and metacone variable in M?’; lower 
canine much exceeding P, in height, and possessing a well defined, al- 
though small, anterointernal basal cusp; P; and P; small and crowded 
or not; P, single-rooted or double-rooted and lacking a well defined 
metaconid cusp. 

Muzzle provided with dorsolateral glandular masses which in some 
forms rise above the muzzle as peculiar lumps (fig. 7); nostril openings 
small and of primitive vespertilionid type or large, opening upward, 
and peculiarly extended posteriorly (fig. 8); auricle and tragus large 
(fig. 9); tragus provided with a prominent basal lobe, and not united 
with posterior basal lobe of auricle; anterior basal lobe of auricle 
normal, expanded, or reduced; transverse ribs on auricle variable in 
posterior extension; second phalanx of third digit longer or shorter 
than third phalanx (fig. 6); calear keeled or not; interfemoral mem- 
brane attached to base or tip of last caudal vertebra; coloration brown, 
without pattern, except for ventral pallor; membranes and ears 
brownish. 

Key to subgenera of Plecotus 


1. Nostril unspecialized; accessory basal lobe of auricle developed into a projecting 
lappet; calcar keeled; brain case broad (53 percent of greatest length). 
Subgenus Idionycteris (p. 129) 

Nostril with posterior elongation and reduction of cornu; accessory basal lobe 
of auricle absent or only slightly developed; calcar not keeled; brain case 
narrow (averaging 48-50 percent of greatest length). ...... Jai a4 


| 


| 
| 
| 
| 


AMERICAN BATS—HANDLEY 129 


2. Supraorbital region sharply ridged; postorbital expansion of zygoma in middle 
third of arch; basial pits absent; P4 longer than wide; 3d commissure of M® 
equal to or shorter than 2d; Ps double-rooted; muzzle glands'slightly enlarged; 
anterior basal lobe of auricle complete; 2d phalanx of 3d digit shorter than 
isiphalank 34.0. . . . . . Subgenus Plecotus (p. 132) 

Supraorbital region emoothiy Pounded or faintly ridged; post orbital ex- 
pansion of zygoma in posterior third of arch; basial pits prominent; P4 
wider than long; 3d commissure of M? equal to or longer than 2d; P single- 
rooted; muzzle glands greatly enlarged; anterior basal lobe of auricle re- 
duced; 2d phalanx of 3d digit longer than Ist phalanx. 

Subgenus Corynorhinus (p. 136) 


Subgenus Idionycteris Anthony 


Plecotus J. A. Allen, 1881, p. 184. (Part.) 
Corynorhinus G. M. Allen, 1916, p. 352. (Part.) 
Idionycteris Anthony, 1923, p. 1. 

Typr sprcius: Jdionycteris mexicanus Anthony. 

Distrispution: Northeastern Mexico and southwestern United 
States (fig. 12). 

Description: Supraorbital region sharply ridged (fig. 4); temporal 
ridges do not coalesce to form a sagittal crest; rostrum broad for a 
Plecotus, flattened, and with pronounced middorsal concavity; brain 
case relatively broad and shallow; zygoma relatively thick and strong 
in appearance, with postorbital expansion in middle third of arch; 
median postpalatal process absent; basial pits absent; auditory bulla 
slightly elongated in outline. 

I! with accessory cusp (fig. 2), near base of tooth; P! small, barely 
exceeding cingulum of canine in height; P* wider than long; space 
between posterointernal edge of P* and anterior edge of M!' only 
about one-third the lingual length of P*; third commissure of M? 
equal to or longer than second commissure (fourth commissure may be 
barely indicated); metacone of M®? fairly prominent; P; much larger 
than P3; P, single-rooted. 

Glandular masses on muzzle not noticeably enlarged (fig. 7); nostril 
of primitive vespertilionid type, with well-defined cornu (fig. 8); 
auricle with anterior basal lobe complete, but not prolonged beyond 
anterior edge of auricle (fig. 9); accessory anterior basal lobe enlarged 
and produced beyond edge of auricle as a “lappet’”’; transverse ribs 
on auricle may or may not extend to posterior edge of auricle;! tragus 
relatively broad and short; second phalanx of third digit longer than 
first phalanx (fig. 6); forearm averages relatively short; calcar keeled; 
interfemoral membrane with 12 to 13 transverse ribs and attached 
to tip of last caudal vertebra. 


1In the type of mezicanus the ribs fade out indeterminently near the posterior border, leaving this area of 
membrane clear; in the type of phyllotis the ribs apparently end about 3 mm. from the posterior border, at 
which point they break into a multiplicity of wrinkles which extend to the border; in USNM 302900, from 
Arizona, the ribs definitely extend to the posterior border of the auricle. 


130 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Plecotus phyllotis G. M. Allen 
Plecotus auritus J. A. Allen, 1881, p. 184. 
Corynorhinus phyllotis G. M. Alien, 1916, p. 352. 
Idionycteris mexicanus Anthony, 1923,? p. 1. 
Plecotus phyllotis Dalquest, 1953, p. 63. 
Tdionycteris phyllotis Handley, 1956, p. 53. 

Hototyre: MCZ 5943; adult female, skin and skull; collected 
24 March 1878 by Edward Palmer; San Luis Potosi, [probably near 
the city of San Luis Potosi or near Rio Verde (J. A. Allen, 1881, 
p- 193)], Mexico. 


DistripuTion: Known only from San Luis Potosi, Tamaulipas, 
and Arizona (fig. 12). Further collecting may show this species to 
have a geographic range similar to that of Plecotus (Corynorhinus) 
mexicanus (fig. 15). 

Description: As in subgeneric description. Adult coloration: 
Mass effect of upperparts between Warm Buff and Cinnamon-Buff; 
hair bases Bone Brown, sharply distinguished from hair tips; a 
conspicuous cottony tuft of whitish hairs surrounds the posterior 
base of each auricle. Tips of hair of underparts between Pale 
Ochraceous-Buff and Light Buff, sharply distinguished from the 
Bone Brown to Fuscous-Black hair bases. Face covered with short 
hairs; anterior external border of auricle with many long, curly 
hairs; hairs long, lax, and dense on remainder of body. Size large 
for genus; tragus appearing shorter than in other species of Plecotus 
and almost as broad as in Plecotus auritus. 

MBASUREMENTS: See tables 7, 13. 

Remarks: Until Cockrum’s capture of a specimen in southeastern 
Arizona in 1955 (Cockrum, 1956b, p. 546), this species had been 
known only by two Mexican specimens which had served individually 
as types for Corynorhinus phyllotis G. M. Allen and IJdionycteris 
mexicanus Anthony. ‘These names have been shown to be synony- 
mous (Handley, 1956, p. 53). The two Mexican specimens are 
almost identical. Compared with them, the example from Arizona 
is larger, its tragus is more broadly rounded (more blunt) distally, 
and the bat is more pallid throughout. Dorsally its fur is more 
buffy, less yellowish brown, and its ears and membranes are more 
grayish, less brownish. These differences may represent geographic 
variation, but it hardly seems worthwhile to apply a new subspecific 
name until additional specimens, now available, are studied. 

Hasirat: Very little is known of the natural history of this species. 
The Arizona specimen was taken at 1:00 a.m. on 30 May 1955 in a 
mist net stretched across an artificial swimming pool on the lawn of 


2 Holotype: AMNH 62260; adult female, skin and skull; collected 17 June 1922 by W. W. Brown; Miqui- 
hauna [= Miquihuana?], Tamaulipas; collector’s No. 2. This locality is on the eastern flank of the Sierra 
Madre Oriental, at about 5,000 feet elevation, near the upper edge of the Lower Austral Zone (Goldman, 
1951, p. 263). 


AMERICAN BATS—HANDLEY 131 


Figure 12.—Distribution of Plecotus phyllotis G. M. Allen. Solid symbols, specimens 
examined; encircled solid symbol, type locality. 


the Southwestern Research Station of the American Museum of 
Natural History (Cockrum, 1956b, p. 546). This is a riparian com- 
munity in a region where the oak-juniper association dominates. 
Subsequently, additional specimens of P. phyllotis have been 
netted in the same area, but data on them have not been published. 
SPECIMENS EXAMINED: Three, from the following localities: 


MEXICO: San Luis Porosf: 1s (MCZ). Tamavurpas: Miquihauna 
[= Miquihuana?], ls (AMNB). 

UNITED STATES: Arizona: Cochise County: Southwestern Research 
Station, 5 miles west-southw est of Portal, 5,400 feet, ls (USNM). 


132 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


Subgenus Plecotus E. Geoffroy Saint-Hilaire 


Vespertilio Linnaeus, 1758, p. 32. (Part.) 

Macrotus Leach, 1816, p. 5. (Nomen nudum.) 
Plecotus E. Geoffroy Saint-Hilaire, 1818, pp. 112, 118. 
Plecautus F. Cuvier, 1829, p. 415. 


Tyre Spseciss: Vespertilio auritus Linnaeus. 

DistrirpuTion: Temperate parts of Eurasia and northern Africa. 
Norway, Great Britain, Ireland, Portugal, and Canary Islands east 
to Kamchatka, and Japan; north to Kola Peninsula, lat. 60-62°N. 
in Siberia, Sakhalin, and Kamchatka; south to Canary Islands, 
French Morocco, Tunisia, Sudan, Israel, northern Iran, northern 
Pakistan, Nepal, Sinkiang, Szechwan, Hopeh, Korea, and Japan 
(fig. 13). Altitudinal range, sea level to 10,500 feet. 

Description: Supraorbital region Sigel ridged (fig. 4); tem- 

poral ridges coalesce to form a sagittal crest; rostrum narrow com- 
ae with other subgenera of Plecotus, arched, and without mid- 
dorsal concavity; brain case relatively narrow and shallow; zygoma 
relatively thick and strong in appearance, with postorbital expansion 
in middle third of arch; median postpalatal process a poorly devel- 
oped spine; basial pits absent; auditory bulla circular in outline. 

I! with accessory cusp (fig. 2); P! relatively large, considerably 
exceeding cingulum of canine in height; P* longer than wide; space 
between posterointernal edge of P* and anterior edge of M? almost 
equals lingual length of P*; third commissure of M® equal to or shorter 
than second commissure; metacone of M? not prominent (reduced to 


Ril ty sell 


Ficure 13.—Distribution of Plecotus auritus Linnaeus. Solid symbols, type localities. 
1, P. a. auritus Linnaeus; 2, P. a. teneriffae Barrett-Hamilton; 3, P. a. meridionalis 
Martino; 4, P. a. christiet Gray; 5, P. a. mordax Thomas; 6, P. a. wardi Thomas; 7, P. a. 
puck Barrett-Hamilton; 8, P. a. homochrous Hodgson; 9, P. a. kozlovi Bobrinskii; 10, 
P. a. ariel Thomas; 11, P. a. sacrimontis G. M. Allen. 


AMERICAN BATS—HANDLEY 133 


vanishing point in some specimens) ; P; much larger than P;; P, double- 
rooted, almost as long in anteroposterior diameter at the cingulum as 
the tooth is high. 

Glandular masses on muzzle slightly enlarged (fig. 7); nostril with 
slightly reduced cornu and with posterior elongation (fig. 8); auricle 
with anterior basal lobe complete and projected noticeably beyond 
anterior edge of auricle (fig. 9); accessory anterior basal lobe small 
and not projected beyond edge of auricle; transverse ribs on auricle 
extend without interruption to posterior border of auricle; tragus 
relatively broad and long; second phalanx of third digit equal to or 
shorter than first phalanx (fig. 6); forearm averages relatively short; 
calcar not keeled; interfemoral membrane attached to base of last 
caudal vertebra. 

MEASUREMENTS: See tables 7, 13. 

Remarks: According to Ellerman and Morrison-Scott (1951, p. 
181), the subgenus includes only one species, P. auritus. They list 11 
geographic races (fig. 13). 

Literature on the natural history of Plecotus auritus is extensive. 
The following data were gleaned from English sources, mostly from 
Alcock and Moffat (1901), Barrett-Hamilton (1910-1911), Matthews 
(1952), and Millais (1904). For reference to the excellent studies in 
Germany, see Hisentraut (1937). 

Hasirat: Plecotus auritus is distinctly arboreal, characteristically 
hunting in the foliage of large trees. Ash trees seem to be particularly 
favored, but oaks and other deciduous species also may be frequented. 
However, it is not unusual to see P. auritus hawking around the upper 
shoots of hedgerows and willows, and sometimes it descends to forage 
among the lower shrubs and flowers in gardens. Rarely it hunts low 
over meadows, and has been observed to alight on cow dung to devour 
the insects attracted there. 

P. auritus commonly hangs up by day in the summertime in old 
boathouses, cowsheds, church attics, and similar places; behind 
window shutters and under eaves of old houses; in tree holes; and even 
on trunks of trees, sheltered only by the roughness of the bark. This 
species may utilize separate sites as daytime sleeping places and 
night-time dining-halls or lounges. Barns and outbuildings are 
favored for the latter, but tree-trunks are also used. Accumulations 
of insect remains in such places betray the nightly visits of the bats. 

Winter roosts are principally in caves, where crevices as well as open 
chambers may be utilized. Attics of old houses and occasionally 
hollow trees are also used. 

Brnavior: In some parts of its range, as in Britain, P. auritus may 
be one of the commonest and most widely distributed bats, yet its 
habits render it less conspicuous,than many of the others. It is 


134 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


nocturnal and not crepuscular, beginning to forage 30 to 40 minutes 
after sunset. Then it moves in such a delicate, mothlike fashion in 
and out of foliage that it is not easily detected. It has been suggested 
that to see this species the observer should station himself at dusk 
under a tree whose foliage is not too dense to be seen through (e.g., an 
ash) and watch closely for the bats to appear among the branches 
overhead. Close attention is necessary, for the movements of the 
long-ear may be so stealthy that the presence of even five or six 
individuals hunting simultaneously in a single tree may be overlooked. 
Furthermore, the observer should choose fine weather for his vigil 
inasmuch as P. auritus seems to be much affected by atmospheric 
conditions. 

Undoubtedly this species catches some flying insects, but much of 
its prey is picked from foliage. It hovers somewhat like a humming- 
bird to pick insects from leaves or flowers, or it plunges into the middle 
of a spray and remains for several seconds clinging to the twigs. It 
has been observed climbing among branches, on tree trunks, and on 
walls, presumably in search of food. 

Flight from tree to tree or roost takes a special form. When the 
destination is near at hand the bat darts through the air with a swift, 
direct flight. When the destination is more distant, the bat makes a 
plunging descent to within a few inches of the ground and makes off 
in a jerking zig-zag fashion. So sudden and irregular are its move- 
ments that its course is difficult to follow. 

In direct flight the ears appear to be held erect, but when the bat 
threads its way among foliage in search of insects they are thrown 
forward so as to resemble a “proboscis” or are curled downward and 
forward so as to suggest “‘cheek pouches.’ In sleep the ears are 
bent backwards until they le close along the body, partially covered 
by the wings, or they are curled down and forward almost like the 
horns of a ram. In the former position the tragus always points 
forward, in the latter it may be reflexed. The ears may be folded or 
unfurled slowly, or snapped up suddenly. The action of the two ears 
can be synchronized or independent. 

P. quritus alights in an upright position, clinging first to an object 
with its thumbs. It soon reverses its position and hangs head down- 
ward. In climbing or walking, the limbs are moved alternately, the 
head bobs to and fro, and the bat advances in a curious jerky fashion 
but with surprising speed. Preparing to fly, the bat raises its head, 
pricks forward its ears, and looks about with a seemingly intelligent 
air. When forced to do so, P. auritus swims rather well. 

The long-ear has a variety of call notes. The commonest are a 
high-pitched chirping squeak used in ordinary communication and in 


AMERICAN BATS—HANDLEY 135 


quarrels, and a shrill, long-drawn, querulous, childlike note uttered 
when the bat is disturbed or surprised. There are also buzzing 
sounds, and a short, melodious, trilling, birdlike sound used for 
communication, both in flight and at rest. 

Moffat (1922, p. 107) observed mating both before and after 
hibernation. The single young is born in June or July. Females 
commonly form nursery colonies of up to 50 or 100 or more indi- 
viduals during spring and summer. Large winter aggregations are 
infrequent. Solitary hibernating individuals are often found. The 
long-ear is ordinarily rather quarrelsome in disposition, attacking 
and biting bats of other species that venture too near its roosting 
spot. It has been observed to attack and drive away the pipistrelle 
from its feeding area. Occasionally mixed colonies of P. auritus and 
Pipistrellus pipistrellus and various species of Myotis have been 
encountered. 

Ryberg (1947, p. 71) tested the homing ability of the long-ear. 
Two of nine banded individuals returned 32 miles from the point of 
release to their home territory. 

P. auritus is easily maintained in captivity. It tames quickly, 
eats readily, and remains in good health for some time. It is clean 
in its habits, grooming itself carefully after feeding. It seems more 
intelligent than other bats and is playful, alert, and lively, even in 
daytime. 

Flower (1931, p. 161) recorded the survival of a P. auritus in 
captivity for 1 year, 2 months, and 2 days. The maximum recorded 
ages of banded, wild individuals are 5 years, 4 months (Ryberg, 
1947, p. 78) and 5 years, 6 months (Verschuren, 1956, p. 5). 

Parasirss: Stiles and Nolan (1931, p. 713) listed 27 genera of 
parasites known to have infested P. auritus. 

Foop: The long-eared bat is primarily a moth-eater. Gould 
(1955, p. 400) reported Lepidoptera of 22 species in its diet. These 
varied in wing spread from 30 to 67 millimeters (average 45). Re- 
mains of several species of Coleoptera and Diptera have been found 
in its droppings. In captivity it seems to prefer mealworms, but 
will also accept flies and raw meat. 

HisperNATION: In the British Isles the hibernating season for 
P. auritus extends from about the middle of October to early April. 
This species is a light sleeper, and if disturbed will generally wake, 
chatter, and attempt to bite if touched. It sometimes emerges in 
winter from its hibernating quarters when the weather is mild, and 
it commonly shifts during the winter from spot to spot within its 
place of hibernation without venturing out into the open. 


136 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Subgenus Corynorhinus H. Allen 


Vespertilio Rafinesque, 1818, p. 446. (Part.) 

Plecotus Lesson, 1827, p. 96. (Part.) 

Synotus Wagner, 1855, p. 720. (Part.) 

Corynorhinus H. Allen, 1865, p. 173. 

Corynorhynchus Peters, 1865, p. 524. (Nomen nudum.) 
Corinorhinus Dobson, 1875, p. 348. 


Tyrer species: Plecotus macrotis LeConte. 

DisrRinuTIoN: Temperate North America in Lower Austral and 
Lower Sonoran to Canadian life zones from Virginia, Ohio, Illinois, 
Kansas, South Dakota, Idaho, and British Columbia south to the 
Gulf of California and the Gulf of Mexico and through the Mexican 
Highlands to the Isthmus of Tehuantepec (fig. 27). 

Description: Supraorbital region smoothly rounded or faintly 
ridged (fig. 4); temporal ridges remain apart or coalesce to form a 
sagittal crest; rostrum broad compared with other subgenera of 
Plecotus, flattened, and with slight middorsal concavity; brain case 
relatively narrow and deep; zygoma relatively thin and fragile in 
appearance, with postorbital expansion in posterior third of arch; 
median postpalatal process a prominent spine; basial pits prominent; 
auditory bulla circular in outline. 

I! with or without accessory cusp (fig. 2); P! small, barely exceeding 
cingulum of canine in height; P* wider than long; space between 
posterointernal edge of P* and anterior edge of M! only about one- 
third lingual length of P*; third commissure of M? equal to or longer 
than second commissure; metacone of M? fairly prominent; P, only 
slightly larger than P3; Py single-rooted, much shorter in antero- 
posterior diameter at the cingulum than the tooth is high. 

Glandular masses resembling thumbless mittens, rise from sides of 
muzzle (fig. 7); nostril without cornu and with posterior elongation 
(fig. 8); auricle with anterior basal lobe reduced to a small lap of 
membrane near tragus (fig. 9); accessory anterior basal lobe absent; 
transverse ribs on auricle interrupted by vertical rib near posterior 
border of auricle; tragus relatively narrow and long; second phalanx 
of third digit longer than first phalanx (fig. 6); forearm averages 
relatively long; calcar not keeled; interfemoral membrane attached 
to tip of last caudal vertebra. 


Key to species of the subgenus Corynorhinus 


1. M3 with 4th commissure almost as long as 8rd (Pleistocene species). 
P. tetralophodon (p. 140) 
M3 with 4th commissure barely indicated or absent. . .......-. 2 
2. Supraorbital region faintly ridged, those ridges continuous with the temporal 
ridges, which do not coalesce to form a sagittal crest (Pleistocene species). 
P. alleganiensis (p. 137) 


AMERICAN BATS—HANDLEY 137 


Supraorbital region not ridged; extension of temporal ridges into the inter- 
orbital region variable; temporal ridges normally coalesce to form a sagittal 
crest (Recent species) . a Ua) wae Renee A 3 

3. Tips of ventral hairs ne or whitish, Sreeoly contrasted with blackish beeea 
median postpalatal process triangular in shape, with a broad base; rostrum 
weak and much depressed; I! with prominent secondary cusp; anterointernal 
cusp of cingulum of P4 usually present (Southeastern United States). 

P. rafinesquii (p. 151) 

Tips of ventral hairs brownish or buff, often not sharply contrasted with 
slate, gray, or brownish bases; median postpalatal process usually styliform, 
with a narrow base (occasionally triangular); rostrum strong and not 
depressed (east of Great Plains and in Pacific Northwest), or variable; I! 
with or without secondary cusp; anterointernal cusp of cingulum of P4 
absent (east of Great Plains), or variable ........ Ye Oke 4 

4. Coloration of dorsum dark sooty-brown, with scant contrast ipetwean bases 
and tips of hairs; greatest length of skull usually less than 15.7 mm. (9) 
or 15.5 mm. (co); maxillary tooth row usually less than 4.9 mm.; I! usually 
with secondary cusp; tragus usually less than 13 mm. long; interfemoral 
cross-ribs usually less than 9 (Mexico). . . . . P. mexicanus (p. 141) 

Coloration of dorsum (in Mexico) yellow-brown, with sharp contrast between 
bases and tips of hairs; greatest length of skull usually more than 15.7 mm. 
(9), or 15.5 mm. (co); maxillary tooth row usually more than 4.9 mm.; I! 
usually simple; tragus usually more than 13 mm. long; interfemoral cross- 
ribs usually more than 9 (Appalachians, Ozarks, western United States, 
southwestern Canada, Mexico)... .. . . . P. townsendii (p. 165) 


PLEISTOCENE SPECIES 
Plecotus alleganiensis Gidley and Gazin 
Corynorhinus alleganiensis Gidley and Gazin, 1933, p. 345. 


Hoxtotypr: USNM (Vert. Paleont.) 12412; skull with incomplete 
dentition and lacking auditory bullae, zygomatic arches, and man- 
dibles; collected 1912-1915, by J. W. Gidley; Cumberland Cave, 
Allegany County, Md. 

DistriBuTIon: Known only from the Pleistocene fauna of Cumber- 
land Cave, Md. (fig. 14). 

Description: Skull similar in size and proportions to those of the 
Recent species P. rafinesquii and P. townsendii, but brain case possibly 
shallower and wider; supraorbital region slightly ridged; temporal 
ridges strongly marked, continuous with supraorbital ridges, and 
converging posteriorly, but not coalescing to form a sagittal crest; 
dorsal profile of rostrum almost straight; upper dentition (P'—M® 
known) appears to be identical to that of Recent material; lower 
dentition (Ii—M; known) similar to that of Recent species, but 
teeth average slightly more robust. 

MeaAsuREMENTS: Holotype, in millimeters: Greatest length, 15.9+ ; 
interorbital breadth, 3.9; brain case breadth, 8.5; brain case depth, 
5.5; maxillary tooth row, 4.9+; postpalatal length, 5.9+; palatal 
breadth, 5.9. 


138 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 14.—Type localities of the Pleistocene species: 1, Plecotus alleganiensis Gidley and 
Gazin; 2, Plecotus tetralophodon Handley. 


Comparisons: When Gidley and Gazin described Plecotus al- 
leganiensis from the Pleistocene deposits of Cumberland Cave, Md., 
they had little comparative material of Recent species from nearby 
areas. I have compared the type and other cranial fragments of 
Plecotus alleganiensis with the now abundant material of P. townsendi 
from West Virginia and of P. rafinesquii from the southeastern 
United States. While some of the characters ascribed to P. al- 
leganiensis by Gidley and Gazin are not as distinctive as they supposed, 
still there are sufficient differences to warrant separation of P. al- 
leganiensis from P. townsendii and P. rafinesquii. 

The faint supraorbital ridges and the prominent, independent, 
temporal ridges distinguish P. alleganiensis from the Recent species. 
In both specimens of P. alleganiensis that have the middorsal portion 
of the skull intact, the supraorbital and temporal ridges are continu- 
ous, and the temporal ridges converge posteriorly to within about 
0.75 mm. of each other, but do not merge to form a sagittal crest. 
In Recent species of the subgenus Corynorhinus the temporal ridges 
sometimes extend into the interorbital region, but there is no supra- 
orbital ridge. A sagittal crest usually is present in Recent species 
(exceptions noted are a specimen of P. mexicanus (KU 29911) from 


AMERICAN BATS—HANDLEY 139 


Mexico and specimens of P. townsendii (AMNH 142023, USNM 
297710) from West Virginia; in these the temporal ridges are contigu- 
ous but not coalesced). 

The presence of supraorbital ridges is interpreted as a primitive 
characteristic. They are present in Huderma maculatum, Plecotus 
auritus, and P. phyllotis. They are absent in P. mezxicanus, P. 
rafinesquii, and P. townsendii. Separate temporal ridges, also con- 
sidered to be a primitive characteristic, are observed in Huderma macu- 
latum and in Plecotus phyllotis; they unite to a common crest in the 
other Recent species of the genus Plecotus. 

Other cranial characters relate P. alleganiensis closely to the Recent 
species P. rafinesquit and P. townsendiv. The rostrum is relatively 
broad and flattened; the anterior nares are enlarged; the rostral 
profile is almost straight (normally, though not invariably, concave 
in P. townsendii virginianus, and normally almost straight in P. 
rafinesquii). The palate and basicranium are similar to those of 
Recent species of the subgenus Corynorhinus; basial pits are prominent. 

General proportions of the skull of P. alleganiensis are the same as 
in P. t. virginianus; the brain case may be a trifle shallower. Depth 
of the brain case is difficult to assess in the only specimen in which 
it is intact, however, for the posterior part of the skull has been 
compressed (the occipital condyles are in the plane of the basioccipital- 
basisphenoid plate, the foramen magnum is in almost the same plane, 
and the supraoccipital, rather than being almost vertical to the main 
axis of the skull, has its lower margin tilted forward about 30°). 

The upper dentitions of Recent P. ¢. virginianus and fossil P. 
alleganiensis are indistinguishable. None of the 27 mandibular frag- 
ments of P. alleganiensis is complete, but all parts of the lower jaw 
are represented. Differences between the mandibles of P. alleganien- 
sis, P. townsendii, and P. rafinesquii are slight. The masseteric 
(coronoid) fossa is usually deeper and better defined in the fossil, 
and the mandibular foramen lies more posteriorly and lower, nearer 
the base of the angular process; the canine averages larger in diameter 
at the cingulum; P,; averages slightly larger; and the molars average 
slightly broader—in essence, the dentition of P. alleganiensis is slightly 
more robust. 

P. alleganiensis differs markedly from the other Pleistocene species, 
P. tetralophodon, in having the skull apparently broader, the supra- 
orbital region ridged, the temporal ridges farther apart, and the fourth 
commissure of M® barely indicated. 

Rewationsuips: P. alleganiensis shares many morphological fea- 
tures with the two Recent species of the genus that now inhabit the 
eastern United States. It could have been directly ancestral to either 
or both of these species. Relationship of P. alleganiensis to the 


140 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


Mexican Pleistocene®species, P. tetralophodon, appears to be more 
remote. 

SPECIMENS EXAMINED: Four cranial fragments and 27 fragments of 
mandibles from Cumberland Cave, Md. 


Plecotus tetralophodon Handley 
Corynorhinus tetralophodon Handley, 1955b, p. 48. 


Hotorype: CIT (Vert. Paleont.) 192/2989; well preserved skull 
with worn teeth (lacking mandibles, auditory bullae, hamular proc- 
esses, all incisors, right canine, and the minute premolar, P', from 
each maxilla); collected by Chester Stock in Pleistocene deposits of 
San Josecito Cave, near the town of Aramberri, southern Nuevo 
Leén, Mexico, elevation 7,400 feet. 

DistRipuTION: Known only from the Pleistocene fauna of San 
Josecito Cave, Nuevo Leén, Mexico (fig. 14). 

Description: Similar to Recent species of the subgenus Coryno- 
rhinus in most cranial details. Anterior nares relatively small and 
rounded in outline (dorsal view) ; brain case relatively shallow; supra- 
orbital ridges lacking; temporal ridges prominent and converging 
posteriorly, so that they meet but do not completely merge; inter- 
maxillary notch relatively small; extension of palate posterior to M? 
relatively short; median postpalatal process styliform; basial pits 
deep and well defined. Tooth rows crowded; teeth relatively fragile 
(not robust); canine with small internal cingular cusp; anterointernal 
cingular cusp of P* only slightly indicated; M*® with well developed 
fourth commissure, almost equaling third commissure in length. 

MEASUREMENTS: Holotype, in millimeters: Greatest length, 15.6; 
zygomatic breadth, 8.2; interorbital breadth, 3.4; brain case breadth, 
7.7; brain case depth, 5.3; maxillary tooth row, 5.0; postpalatal 
length, 5.9; palatal breadth, 5.7. 

Comparisons: A well developed fourth commissure of M? distin- 
guishes P. tetralophodon from related species. This commissure is 
barely indicated in other New World species of Plecotus and in 
Euderma maculatum; there is no trace of it in the Old World Plecotus 
auritus, in which even the third commissure of M? is reduced. 

The degree of shallowness in the cranium of P. tetralophodon (brain 
case depth equals 34 percent of greatest length) is rarely equaled in 
Recent specimens of the subgenus Corynorhinus. It is observed in 
less degree in Huderma maculatum, Plecotus auritus, P. phyllotis, and 
possibly in P. alleganiensis. 

Absence of a sagittal crest is a character shared with Huderma 
maculatum, Plecotus alleganiensis, and P. phyllotis. The temporal 
ridges usually are even farther apart in those forms, however. A few 
specimens of P. mexicanus and P. townsendii resemble P. tetralophodon 
in this respect. 


AMERICAN BATS—HANDLEY 141 


Rerationsuirs: Plecotus mexicanus and P. townsendri now inhabit 
the region where the fossil P. tetralophodon was found. P. tetralopho- 
don must have been closely related to these species, for the general 
conformity of the skull is similar in all three; probably all came from 
a common stock. I do not believe that P. tetralophodon is directly 
ancestral to either of these Recent species, however. It is less like 
P. rafinesquii and P. alleganiensis. 

SPECIMEN EXAMINED: The holotype. 


RECENT SPECIES 
Plecotus mexicanus G. M. Allen 


Corynorhinus macrotis pallescens Miller, 1897, p. 52. (Part.) 
Corynorhinus megalotis mexicanus G. M. Allen, 1916, p. 347. 
Corynorhinus rafinesquii mexicanus Miller, 1924, p. 83. (Part.) 
Plecotus rafinesquii mexicanus Dalquest, 1953, p. 64. (Part.) 
Corynorhinus mexicanus Handley, 1955c, p. 148. 


Ho.totypr: USNM (Biol. Surv. Coll.) 98285; adult female, skin 
and skull; collected 25 August 1899, by E. W. Nelson and E. A. Gold- 
man; near Pacheco [Sierra de Brefia, 8,000 feet], Chihuahua, Mexico; 
collector’s No. 13955. 

Distrisution: The higher and more humid parts of the Sierra 
Madre Occidental, the transverse volcanic belt of central Mexico, 
and the Sierra Madre Oriental (fig. 15). North to El Tigre Moun- 
tains, Sonora; near Pacheco, Chihuahua; and 22 miles south-southeast 
of Monterrey, Nuevo Leén. South to Pétzcuaro, Michoacén; Monte 
Rio Frfo, 28 miles east-southeast of Ciudad México; and Jico, Vera- 
cruz. Zonal Distribution: Mostly Transition and Upper Austral. 
Altitudinal range from 4,800 feet to 10,500 feet; usually above 6,000 
feet. 

Description: Adult coloration: Tips of hairs of upperparts vary 
from Verona Brown to Fuscous, bases from Bone Brown to Fuscous- 
Black; degree of differentiation between tips and bases of hairs slight; 
burnishing of hair tips absent or only slightly evident in most speci- 
mens; hairs at posterior base of auricle paler than remainder of dor- 
sum. Bases of hairs of underparts vary from Benzo Brown to Fuscous, 
tips are about Pale Pinkish Cinnamon. Immature coloration: Dor- 
sum smoky brown, similar to coloration of some adults. 

Size small for subgenus; lobes of presternum slightly expanded; 
forearm averages relatively long; brain case averages relatively wide 
and deep, sloping up abruptly above the short, depressed rostrum; 
anterior nares, viewed from above, usually relatively small and angu- 
lar in posterior outline; median postpalatal process varies from styli- 
form to triangular; auditory bulla averages relatively small. First 
uppers incisor usually with’ a® prominent accessory cusp and always 


142 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 
{ i] 
1 i 
‘ 1 
i 1 
AS | s 
Se 1 I 
Bae 1 pag eA Taian be a a ee ~ 
_ ' SN 
seine Px: Wes. Eee || lah! Ne 
7°) ES} ‘ 
g ar S 
=a \ 
o— ie: 
— \ 
~~ 
\ 
Ss 
erate) \ wore 
=a ~ 5 NS 
= “Sal ~ 
= si \ 
+ yh Ae 
———— / \ 
\ : 
Xe 


oe 


Figure 15.—Distribution of Plecotus mexicanus G. M. Allen. 


Solid symbols, specimens 
examined; open symbols, other records; encircled solid symbol, type locality. 


with at least a trace of such a cusp; upper canine somewhat reduced in 
size; P* usually with a small anterointernal cingular cusp. Tragus 


(possibly also auricle) averages relatively short; cross-ribs on inter- 
femoral membrane average eight. 


MEASUREMENTS: See tables 8, 14. 


Comparisons: Compared with Plecotus townsendii australis and 
P. t. pallescens, P. mexicanus is darker, with less contrast between 
bases and tips of dorsal hairs; usually fewer cross-ribs on the inter- 
femoral membrane; smaller tragus on the average; smaller skull; 


AMERICAN BATS—HANDLEY 143 


deeper brain case; shorter, weaker, and more depressed rostrum; 
smaller auditory bullae; actually and relatively shorter maxillary 
tooth row; and more consistently bilobed first upper incisor. There 
is at least an indication of an accessory cusp on I’ in all specimens of 
P. mexicanus that I have examined; few specimens of P. t. australis or 
P. t. pallescens have even a trace of it. 

Retationsures: There is a strong resemblance between P. mezxi- 
canus and P. rafinesquii. Similarities include short, weak, depressed 
rostrum; deep brain case; bilobate first upper incisor; and possession 
of a small anterointernal cingular cusp on P*. 

P. mexicanus differs from P. townsendit in somewhat the same man- 
ner as does P. rafinesquit. This parallel may not be purely coinci- 
dental. Perhaps both P. merxicanus and P. rafinesquii were isolated 
from a common stock (of which P. townsendii is the Recent descendant) 
at about the same time. However, P. mericanus is not differentiated 
from P. townsendii quite as well as is P. rafinesquit. This may indi- 
cate more complete or longer isolation for P. rafinesquii, or partially 
parallel evolution between P. mezxicanus and P. townsendii. 

In coloration P. mexicanus resembles typical P. t. townsendii of the 
Pacific Northwest. Apparently this parallel can be traced to similar 
environmental stimulae. P. ¢. townsendii inhabits cool, humid, 
coastal lowlands, and P. mexicanus inhabits cool, relatively drier 
highlands. 

Remarks: Since 1890 it has been known that bats of the subgenus 
Corynorhinus inhabit Mexico. The first specimen was referred by 
J. A. Allen (1890) to Plecotus townsendii, then thought to be mono- 
typic. Miller (1897) referred all Mexican specimens to a new race, 
P. macrotis pallescens, restricting the name townsendii to populations 
in the Pacific Northwest. Later, G. M. Allen (1916) referred all 
Mexican material to a new race, P. megalotis mexicanus, restricting 
pallescens to the relatively arid portions of the western United States. 
Subsequent authors, with the exception of Burt (1938) for bats of 
northern Sonora, have applied the name P. rafinesquit mexicanus to all 
Mexican populations of the subgenus Corynorhinus. Present evi- 
dence, however, indicates that there are in Mexico two species to 
which the names P. townsendii and P. mezicanus apply. 

Evidence that these are distinct species has been overlooked, 
although examples of each have been in museum collections for more 
than 60 years. Reasons for this oversight are several: 

1. Populations of P. townsendii just north of the range of P. mexicanus in 
Arizona converge toward P. mexicanus in size and in coloration, giving an illusion 
of geographic intergradation between the two species. 

2. The two species are not strikingly different, although they are more distinct 
than some species in other genera of bats, such as of Myotis. 

497256—59_4 


144 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


INTERFEMORAL CROSS—RIBS 
(NUMBER OF) 


P. MEXICANUS 
SPECIMENS !N SAMPLE: TI 
LESS THAN 9 RIBS IN 7T6% 


FREQUENCY 


PLT, AUSTRALIS 
SPECIMENS IN SAMPLE: 26 
MORE THAN 9 RIBS IN 88% 


MAXILLARY TOOTH ROW 
(LENGTH OF) 


P, MEXICANUS 
SPECIMENS IN SAMPLE: 70 


LESS THAN 4.9MM. IN B6% 


> 
o 
Zz 
w 
> 
So 
w 
i 
ve 


P.T. AUSTRALIS 
SPECIMENS IN SAMPLE: 23 
MORE THAN 4.9MM. IN 91% 


ZYGOMATIC BREADTH 


P. MEXIGANUS 
SPECIMENS IN SAMPLE: 65 
LESS THAN 8.5MM. IN 95% 


FREQUENCY 


P. T. AUSTRALIS 
SPECIMENS IN SAMPLE: 20 
MORE THAN 8.5MM. IN 70% 


GREATEST LENGTH OF SKULL 


P. MEXICANUS 
SPECIMENS IN SAMPLE: 68 


LESS THAN 15.7 MM. IN 94% 


FREQUENCY 


P.T. AUSTRALIS 
SPECIMENS IN SAMPLE: 21 
MORE THAN 15.7MM. IN 95% 


Ficure 16.—Comparison of frequency of occurrence of certain measurements in adult 
female P. mexicanus (solid symbols) and P. #. australis (open symbols), including all 
measurable individuals studied. 


497256 O - 59 (Face p, 144) 


MONTANA S|) OKLAHOMA VIRGINIA 
KANSAS WEST VIRGINIA 


NORTHERN) 
a 
WASHINGTON CENTRAL |END, 
OREGON ARIZONA KAS ARKANSAS 


| 


25 


lz0 


16.5 


16.0 


155 


15.0 


MALE 
tet OD LY See MACE 
8 — NUMBER OF SPECIMENS ee 
IN SAMPLE 
OBSERVED RANGE 
OBSERVED MEAN 


2 STANDARD ERRORS O 
EITHER SIDE OF MEA! 


2 STANDARD DEVIATION 
ON EITHER SIDE OF ME 


LEGEND 


FIGURE 17.—Gllecotus mexicanus 
ic. 


TANA i 
MON’ pote is Nas pre Lia uenco HIDALGO DURANGO OKLAHOMA VIRGINIA 
RORTHERN ANU COAHUILA KANSAS — WEST VIRGINIA 
ne ZACATECAS OAXACA NUEVO LEON 


WASHINGTON CENTRAL PACHEGO. 
, SAN LUIS BIG BEND, 
OREGON ARIZONA CHIHUAHUA JALISCO CHARCAS VERACRUZ Potosi phy ; ARKANSAS 


N+to 


O 


‘ 
’ 


15.5 


15.0 


—---- FEMALE 


8 — NUMBER OF SPECIMENS 14.5 
IN SAMPLE 
OBSERVED RANGE 
OBSERVED MEAN 


2 STANDARD ERRORS ON 
EITHER SIDE OF MEAN 


2 STANDARD DEVIATIONS 
ON EITHER SIDE OF MEAN 


LEGEND 


497256 © = 59 (Face p, 144) 


Ficure 17.—Greatest length (in mm.) of skulls of specimens of Plecotus townsendii (upper set of figures) and Plecotus mexicanus 
(lower set of figures), arranged in geographic sequence from northwest to south, thence to northeast. 


i i, whostoa ATHAS 44) Ria Ra Aves AMARSHTL 
a (on SATALLAV BLED elm COMME 
CAZSACS DADSTAIAS: VO CE ay 


Sit |, GOHNOSE Big be 
ADBAHOWE RACRGDS | JAL > RUM AUHEHO Se anoning 


— 


- ; : 


aWOITAIVaG 
“—- ctl sue aaHTIS ~~ 


?«. 
' 


a sAzcossaqe loetlede Geofeerarins) idagnad) ie tr 0A 
4 | spon aidqargorghmickogoana:{ertugit-todsensiol) 


A | | ! ae) 

i ae aay 1 

i “a 7 ; H yA s 
— is \ } 7 iD : 


he, A 
AD A ' 


AMERICAN BATS—HANDLEY 145 


3. No other two species of the subgenus were known to be sympatric. 

4, Relatively few specimens have been available for study. Until recently, 
both species were represented only in the U.S. National Museum. The Texas 
Cooperative Wildlife Research Unit has had an example of each species since 1942; 
the Museum of Zoology, University of Michigan, has had both since 1946 (skins 
and skulls of P. townsendii; P. mexicanus in alcohol); and the Museum of Natural 
History, University of Kansas, has had both since 1948. 

5. With the exception of the type series of P. mexicanus (preserved as dry skins 
and skulls), all early collections of Mexican big-eared bats in the National Museum 
were preserved in alcohol, and in a random selection of specimens for removal 
of skulls, only P. mexicanus happened to be chosen. The presence of P. town- 
sendit, thus, was not detected. 


When Mexican material was assembled from several museums for 
this study, variation was found to be slight in some series, great in 
others. The entire sample segregated into two groups (figs. 16, 17), 
each of which included both sexes and various ages. The amount of 
variation far exceeded that observed in samples from other parts of 
the range of the subgenus Corynorhinus (fig. 17). With one exception, 
all specimens could be referred without hesitation to one group or 
the other on the basis of a dozen diagnostic characters. Each speci- 
men had most of the characters of its group, i.e., any specimen with 
dark coloration also had small tragi and few cross-ribs on the inter- 
femoral membrane, etc. (fig. 16). 

The two forms have distinct although broadly overlapping geo- 
graphic ranges (fig. 18). The range of P. mexicanus—the cooler, 
moister, higher elevations of the Sierra Madre Occidental, the 
transverse volcanic belt of central Mexico, and the Sierra Madre 
Oriental (Goldman and Moore, 1946)—is strikingly similar to the 
ranges of the dark colored races of the harvest mouse, Reithrodon- 
tomys megalotis, outlined by Hooper (1952, p. 51) and to that of the 
white-footed mouse, Peromyscus difficilis (Osgood, 1909, p. 179). 
The range of the other form, P. townsendiz, is in part complementary. 
It inhabits the lower elevations of the arid plateau and desert ranges 
of north-central Mexico north of the transverse volcanic belt, and 
the arid valleys of Jalisco, Morelos, and Oaxaca south of the trans- 
verse volcanic belt. In addition, it occurs together with P. mexicanus 
in the southern extremity of the Sierra Madre Occidental (Sierra de 
Valparaiso, 8,200 feet) and in the transverse volcanic belt. (Santa 
Rosa, 9,500 feet, Convento de Acolman, and Lago Texcoco, 7,500 
feet. However, it should be noted that the latter localities are rela- 
tively arid, despite their high elevation (Davis, 1944, p. 371; Gold- 
man, 1951, p. 146). 

This pattern of distribution might exist without intergradation 
between the two forms if they were: (1) Subspecies with seasonal 
migrations; (2) ecologically isolated subspecies; or (3) distinct, 
partially sympatric, species. 


146 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


0}, 
ae 


————— 


CI %, 
tee es 
e 

fo 

“J 
= 


pbssiogresess: 


“Len 


Ficure 18.—Sympatry of Plecotus mexicanus (triangular symbols) and Plecotus townsendit 
(circular symbols) in Mexico. 


Seasonal movement is not apparent in available samples (table 5). 
P. mexicanus is found in northern and southern parts of its range in 
both winter and summer. P. townsendii has been taken at high eleva- 
tions in the transverse volcanic belt (in the range of P. meaicanus) 
in both summer and winter, has been found south of the belt in spring, 
late summer, and fall, and north of the belt at all seasons. 

The theory of conspecificity becomes less tenable when seasonal 
movement of one or the other of the subspecies cannot be demon- 
strated. The phenomenon of overlapping ends of a chain of sub- 


AMERICAN BATS—HANDLEY 147 


Tas.E 5.—Seasonal distribution of P. mexicanus and P. townsendii in Mexico, as 
determined from specimens for which collection date is known 


P. mexicanus 


North of transverse volcanic belt In transverse volcanic belt 
February: Mojarachic January: Las Vigas 
February: Monterrey June: “Morelos” 

August: El Tigre Mts. July: Jico 

August: Pacheco July: Monte Rfo Frio 
December: Sierra de Valparaiso July: PAtzcuaro 


November: Santa Rosa 
November: Zacualpilla 


P. townsendiz 


North of transverse volcanic belt In or south of transverse volcanic belt 


January:  Yoitepec April: San Andrés 
February: San Juan July: Convento de Acolman 
April: Gen. Cepeda August: Oaxaca 

April: Sta. Helena September: Guadalajara 

June: San Buenaventura September: Cuernavaca 

July: Zimapan November: Charcas 

September: Bella Unién November: Santa Rosa 


October: Presa de Guadalupe 
November: Bledos 

November: Muralla 

December: Sierra de Valparaiso 


species, reacting to one another as species rather than as related 
subspecies, however, occurs with some frequency in mammals, al- 
though it is more conspicuous in birds. It might apply in this instance 
if it could be proved that both P. mexicanus and the Mexican popula- 
tions of P. townsendii (P. t. australis) intergrade to the northward 
with other populations of P. townsendii (P. t. pallescens). Then it 
could be assumed that the extremely arid environment of eastern 
Chihuahua and western Coahuila, where big-eared bats are not known 
to occur, has served as a barrier, sufficiently isolating related popula- 
tions to the east and west of it so that they reacted to each other as 
species when both segments invaded areas to the south of the deserts. 

Mexican populations of P. townsendii intergrade northward with 
other populations of P. townsendii. The zone of intergradation is 
broad, extending from northern Coahuila, through the Texas Pan- 


148 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


handle. There is no evidence in existing samples of intergradation 
between mezicanus and townsendii. 

The geographic ranges of the two forms overlap in northern Sonora; 
a sample from El Tigre Mountains contains typical individuals of 
both forms, and Burt (1938, p. 26) may have had both species in 
material from Sfric. It is true that populations of P. townsendii 
become smaller in body size and darker in color toward the Mexican 
border, but there is no accompanying increase in the frequency of 
the accessory cusp of I’, reduction in the number of interfemoral 
cross-ribs, reduction in contrast between bases and tips of dorsal 
hairs, disproportionate decrease in the size of the auditory bullae, 
the auricles and tragi, or the average length of the maxillary tooth 
row, increase in the average depth of the brain case, or depression of 
the rostrum. In other words, populations of P. townsendii, neighbor- 
ing P. mexicanus on the north, show a convergence of some morpholog- 
ical characters toward P. mexicanus, perhaps in response to similar 
environmental stimulae. Lack of convergence in other characters, 
however, indicates a different genetic structure in the two forms. 

Dice (1940) used a concept of ecological subspecies to explain the 
discontinuous distribution of dark-colored subspecies on isolated 
mountain peaks surrounded by lowlands inhabitied by pale-colored 
subspecies. Dice supposed that the several isolated dark-colored 
populations might have originated in situ as independent derivations 
from the pale-colored stock, presumably in response to similar environ- 
mental conditions. According to this hypothesis, some subspecies 
may be ecological units, rather than units of similar heredity. 

Were it not for the mixed samples (Sierra de Valparaiso and Santa 
Rosa) and absence of intergradation between the two forms, the 
concept of ecological subspecies could be an adequate explanation for 
the distribution of the Mexican bats of the subgenus Corynorhinus. 
It could be assumed that the alternative forms developed wherever 
certain ecological conditions prevailed. Under this hypothesis, inde- 
pendent development of the ‘‘mexicanus’’-type might be expected 
wherever relatively cool, moist conditions of environment occur, and 
the ‘“australis’”-type would be expected under conditions of greater 
warmth and aridity. The distribution of both forms then would be 
largely independent of geography and would be related primarily to 
ecology. The mixed samples could be accounted for as chance min- 
gling of the two forms during the nonbreeding season as a consequence 
of the interfingering of humid and arid environments. Existing 
samples appear to be too numerous and too well distributed, however, 
to have failed to show evidence of intergradation, if such exists. 

It thus appears that the hypothesis of conspecificity is In no wise 
applicable to the Mexican populations of the subgenus Corynorhinus. 


AMERICAN BATS—HANDLEY 149 


Provided that the data from available samples have not been misinter- 
preted, the two forms in consequence must be considered distinct but 
partially sympatric species. 

A few specimens—such as KU 44759 from near Hacienda La Mari- 
posa, Coahuila, which has the coloration, size, large auricle, and shal- 
low brain case of P. townsendiz, but the short tooth row, short tragus, 
and few interfemoral cross-ribs of P. mexicanus—may be hybrids. In 
view of the probable close relationship between the two species, it is 
surprising that such specimens appear to be so infrequent. 

Geographic variation in cranial characters in P. mericanus seems 
insignificant; even individual variation in these characters is of small 
magnitude. Geographic and individual variation in color is slight. 
The average mass effect of the dorsum in the fine series from Las Vigas 
is smoky, blackish brown, with the hair tips so dark as to be hardly 
distinguishable from the dark hair bases. The dark extreme appears 
almost black to the casual glance, while the pale extreme is brownish, 
with sooty tones less pronounced, and with the hair tips slightly paler 
than the hair bases. Specimens from the northwestern extremity of 
the range of the species are somewhat paler than those from the south- 
east; dark extremes are similar in the two regions. 

The fur is usually rather short and woolly, but specimens from 
10,500 feet in Estado de México (TCWC 2823), 8,200 feet in Zaca- 
tecas (USNM 91931), and west-central Chihuahua (USNM 265660) 
have long, laxfur. This does not appear to be a seasonal phenomenon, 
for short-haired individuals have been taken in January, February, 
August, November, and December. The long-haired specimens were 
taken in February, July, and December. These differences in pelage 
are not obviously related to age or sex. 

Hasirat: Apparently this species is partial to forested mountainous 
regions, where it roosts in caves, mine tunnels, and_ buildings. 
Warmer, less humid, and more desert-like districts in the same general 
area are inhabited by P. townsendii. P. mexicanus has been reported 
in the following habitats: 

Summer: Flying about rain pool in opening in pine-fir forest on Monte Rio 
Frio (Davis, 1944, p. 380); in ranch house on lower border of oak forest (5,500 
feet) above Jico (E. W. Nelson, field notes); hanging from roof of small, damp, 
north-facing grotto in vicinity of mixed forest of pine, oak, and madrojio near 
Pdtzcuaro (E. W. Nelson, field notes; Goldman, 1951, p. 195). 

Winter: In old mine near Santa Rosa, near summit of Sierra de Guanajuato, 
upper slopes of which are characterized by light rainfall and second-growth forest 
of oak, alder, madrofio, Crataegus, and wild cherry (E. A. Goldman, field notes; 
Goldman, 1951, p. 146). Hibernating at 8,200 feet in Sierra de Valparafso in 
cold, damp cavern through which strong currents of air passed from one entrance 


to another; upper slopes of sierra forested with pine, juniper, cypress, oak, mad- 
rofio, and manzanita, frequently interrupted by wet, grassy meadows (E. A. Gold- 


150 PROCEEDINGS OF THE NATIONAL MUSEUM von. 110 


man, field notes; Goldman, 1951, p. 290). In shallow cave in region of oak and 
pine near Mojarachic (Knobloch, 1942, p. 297). 


Benavior: Information on behavior is scant. E. W. Nelson (field 
notes) remarked of a Patzcuaro specimen that: 

When first taken in hand its long ears were bent spirally back and downward 
close against the sides of the head exactly like a pair of little recurved horns. 
While examining the animal with surprised curiosity these horns suddenly ex- 
tended and became a pair of huge upright ears. While examining these ears at 
first I noted a fine regular waviness to the inner margin of the coiled ear which 
disappeared when the ear was extended. 

The field notes of HE. A. Goldman tell of a group of big-eared bats 
found in the Sierra de Valparaiso in December, 1897: 

IT found eight specimens of this species hibernating in a cave at 8,200 feet 
altitude. Six of them in formalin were unfortunately lost along with my tank 
by my hunter while descending to the town of Valparaiso. They were hanging 
as usual by their feet, from the roof of the cave and at considerable distances 
from each other—6 to 10 or 12 feet. Their eyes were closed and their bodies cold 
and stiff. Their wings were hanging very close to the sides of their bodies. Their 
long ears had the margins rolled inwards and they were curled over backward and 
downward somewhat like the horns of a sheep and were pressed close to the sides 
of the head. When first taken from the cave the only signs of life that I noticed 
in them was a very slight moving of the feet, and they would hook them to my 
finger. I tied them all up together in a [handkerchief?] and carried them to 
camp. Their blood soon began to circulate, their bodies became warm and they 
revived sufficiently to be able to chatter and bite at each other and my fingers. 
Six hours after they were taken from the cave they could craw] about but although 
uninjured none of them were able to fly or at least made no effort to do so. The 
cave was cold and damp and a strong current of air passed in at one entrance and 
out at another. 

Foop: Not known. 

Mieration: Like other species of Plecotus, P. mexicanus is not 
known to make extensive migrations (see table 5 and p. 146). 

Hisernation: According to Goldman’s 1897 field notes (above), 
this species may become very lethargic under certain conditions. 

Repropuction: Dalquest noted on specimen labels that no embryos 
were found among a large group of females collected at Las Vigas in 
January. Adult males (10) and adult females (53) were together in 
this colony. Adults of both sexes (3 males, 10 females) likewise were 
found together at Santa Rosa in November. A group of nine females, 
accompanied by five immatures, almost fully grown, but with finger 
joints not completely ossified, were collected near Pacheco on 25 
August. Solitary immature individuals of slightly younger age were 
found on 14 July at Jico and at P&tzcuaro. 

Mott: Two of nine adult females collected near Pacheco on 25 
August were beginning to molt on chest and shoulders (new hairs still 
short and prostrate). Two of five immatures collected with this 
group were beginning to molt on the belly. 


AMERICAN BATS—-HANDLEY 151 


SPECIMENS EXAMINED: A total of 104 from the following localities 
in Mexico: 

Cuinvanua: Near Pacheco [Sierra de Brefia, 8,000 feet], 14s (USNM); Moja- 
rachic [= Mafuarachic?], Rayon District, ls (USNM). Guanasuato: Santa Rosa 
[9,500 feet], 18a (USNM). Mé£xico: Monte Rio Frio, 10,500 feet, 28 miles east- 
southeast of Ciudad México, ls (TCWC). MicnHoacikn: 2 miles north of Patz- 
cuaro, 7,100 feet, 1a (USNM). Moretos: No exact locality, la (USNM). 
NurEvo LE6n: 22 miles south-southeast of Monterrey, 4as (MCZ). Sonora: 
Santa Marfa Mine, El Tigre Mts., la (UMMZ). VeEracruz: Jico [5,500 feet], 
la (USNM); 2.5 miles east of Las Vigas, 8,500 feet, 59bs (KU), 4s (GF); 6 miles 
west-southwest of Zacualpilla, 6,500 feet, 3s (KU). Zacatrcas: Sierra de Val- 
parafso [13 miles west of Valparaiso, 8,200 feet], ls (USNM). 

ADDITIONAL RECORDS: The following records from Mexico probably 
refer to this form: 


PurBLa: Hacienda de Miguel Sesma, 2 miles northwest of Esperanza (Schwartz, 
1954, in litt.). Sonora: Sdric (Burt, 1938, p. 26). 


Plecotus rafinesquii Lesson 


Synonymy is given under the subspecies. 

DistRIBUTION: Southeastern United States (fig. 19). 

Description: Adult coloration: Hairs of upperparts blackish at 
base and yellowish brown to reddish brown distally, the two areas 
sharply demarked; hairs at posterior bases of ears usually not different 
from those of remainder of dorsum. Mass effect darkens with wear. 
Hairs of chest and belly black or blackish at base, white or whitish at 
tip, the division between the two colors sharp; hairs of chin and throat 
like remainder of underparts or with buffy wash encroaching from sides 
of neck; hairs bordering membranes whitish throughout. Immature 
like adult, except much darker throughout, the Avellaneous tips of 
the dorsal hairs and the whitish tips of the ventral hairs failing to 
conceal the blackish hair bases. 

Size medium for the subgenus; presternum with all lobes, but espe- 
cially the ventral lobe (almost as wide at tip as lateral lobe), somewhat 
expanded. Rostrum weak and much depressed, rendering facial profile 
relatively straight; anterior nares relatively small, and posterior border 
with somewhat angular shape; intermaxillary notch of palate relatively 
narrow posteriorly; median postpalatal process triangular in shape, 
with broad base; brain case averages relatively deep for subgenus. 
I’ with prominent secondary cusp; P* normally has small anterointernal 
cingular cusp. 

Comparisons: Plecotus rafinesquii may be distinguished from geo- 
graphically contiguous races of Plecotus townsendii by numerous char- 
acters. When all populations of P. townsendii are considered, however, 
the separation is not so striking. Apparently, only coloration will 
distinguish all the individuals of the two species, all populations 
considered. P. rafinesquii always has the bases of the ventral hairs 


152 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 19.—Distribution of Plecotus rafinesquii Lesson: 1, P. r. macrotis; 2, P. r. rafines- 
quit. Solid symbols, specimens examined; open symbols, other records; encircled solid 
symbols, type localities. Cross-hatching indicates known areas of intergradation between 
subspecies. 


black or blackish and the tips white or whitish, with the two colors 
sharply contrasted, whereas P. townsendii, on the other hand, always 
has the bases of the ventral hairs slate, gray, or brownish and the tips 
brown or buff, with the two colors often not sharply contrasted. 

A secondary cusp on the inner upper incisor is present in all speci- 
mens of P. rafinesquii. It is absent (or represented only by a slight 
shoulder) in all specimens of the geographically adjacent P. t. virgini- 
anus. It is absent in most but not all examples of the geographically 
more remote P. t. australis, P. t. pallescens, and P. t. townsendii. It is 
usually present in geographically adjoining P. t. ingens. The secondary 
cusp is present in all specimens of P. mezicanus. 

In P. rafinesquii there is usually, though not always, a small antero- 
internal cingular cusp on P*. This cusp was not observed in specimens 
of P. t. ingens or P. t. virginianus, but is usually present in P. t. town- 


AMERICAN BATS—HANDLEY 153 


sendii and P. mexicanus and sometimes occurs in P. ¢. australis and 
P. t. pallescens. 

The median postpalatal process is triangular in shape, with a broad 
base, in P. rafinesquit. It is normally narrower at the base and more 
styliform in P. townsendii. In some examples of P. ¢. australis and 
P. mexicanus the process is almost needle-like. The broad-based form 
is encountered in some individuals of all races of P. townsendii, however. 

The intermaxillary notch of the palate is narrower posteriorly in 
P. rafinesquii than in most individuals of P. townsendit. Likewise, 
the anterior nares are relatively small in P. rafinesquii and have the 
posterior border angular in outline. Geographically adjacent races of 
P. townsendii have the nares larger and the posterior border rounded 
in outline. The nares in the more remote P. t. townsend and P. t. 
australis are generally large and rounded, but in P. mezicanus and 
P.t. pallescens they are relatively small and angular as in P. rofinesquit. 

Another characteristic of P. rafinesqui that is geographically vari- 
able in P. townsendu is the appearance of the rostrum. In P. rafine- 
squirt the rostrum is typically weak and much depressed. On the other 
hand, adjacent races of P. townsendii, P. t. ingens and P. t. virginianus, 
as well as P. t. townsendvi, have the rostrum strong and not depressed. 
The rostrum usually is not so weak or depressed in P. t. australis and 
P. t. pallescens as in P. rafinesqun. P. mexicanus, showing these 
characters in the extreme, is similar to P. rafinesquit. 

Skeletons are not available for all of the races of either species, but 
it appears that the presternum may have taxonomic worth. In speci- 
mens examined, all lobes, but particularly the ventral lobe, were more 
expanded in P. rafinesquv thanin P. townsendi (fig. 5). The presternum 
of P. mexicanus is intermediate between these extremes. 

RELATIONSHIPS: Plecotus rafinesquv is the most primitive living 
member of the subgenus Corynorhinus. It is similar in many respects 
P. mexicanus (see p. 143). It is more distinct from geographically 
contiguous races of P. townsendw (ingens and virginianus) than it is 
from some of the geographically more remote races (pallescens and 
townsendit) of that species. This indicates a considerable period of 
isolation for P. rafinesquii and a subsequent extension of range to 
bring the more progressive P. townsendit into geographical contact with 
P. rafinesquii. Isolation for P. rafinesquii is further confirmed by the 
fact that primitive traits (e.g., secondary cusp on I', anterointernal 
cingular cusp on P*, triangular shape of median postpalatal process, 
small anterior nares and intermaxillary notch) which are constant or 
nearly so in P. rafinesquii are normally variable in P. townsendit. 
The coloration of P. rafinesquii, the only character separating all 
individuals of the Recent species of the subgenus Corynorhinus, most 


154 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


resembles the coloration of the more primitive subgenera [dionycteris 
and Plecotus. 

Remarks: Referring to a visit to the “lower parts of the Ohio, the 
Wabash, Green River, Barrens, Prairies, and the States of Indiana, 
Illinois, etc.,’’ Rafinesque (1818, p. 446) described nine species of bats, 
ten mice and shrews, and a number of snakes and fish. Four of the 
nine species of bats and a number of the mice and shrews cannot be 
related to species known to occur in the lower Ohio Valley. The de- 
scription of one of the bats, Vespertilio megalotis, suggests what we 
now know as Plecotus: “‘Vespertuio megalotis. R. (Big-eared bat.) 
Tail three-eighths of total length, body dark gray above, pale gray 
beneath, ears very large, duplicated, auricles nearly as long. Length 
4 inches, breadth 12 inches.”’ Most of Rafinesque’s contemporaries 
thought his V. megalotis was related to Eurasian Plecotus. Lesson 
(1827, p. 96) paraphrased Rafinesque’s description of V. megalotis 
and renamed it Plecotus rafinesquii. He stated that it was ‘perhaps 
only a variety of our long-eared bat [European Plecotus].”’ 

After a brief period of use, V. megalotis and P. rafinesquii fell into 
obscurity, and Plecotus macrotis of LeConte (1831, p. 431) came into 
universal usage as the name for the big-eared bat of the eastern 
United States. 

A. H. Howell’s (1909, p. 68) collection of a second species of big- 
eared bat at Burkes Garden in the Allegheny Mountains of Virginia 
in 1908 led to G. M. Allen’s resurrection of the name V. megalotis. In 
his revision of the genus (now subgenus) Corynorhinus, Allen (1916, 
p. 339) wrote: 

The discovery of a Corynorhinus (szc) distinct from C. macrotis, from extreme 
western Virginia, westward, in the eastern United States was wholly unexpected. 
It is the eastern representative of the desert-colored pallescens of west-central 
United States, from which it chiefly differs in its somewhat darker, more drab, 
coloration. 

On the basis of specimens from Virginia, Kansas, and Colorado, 
Allen supposed that this Plecotus, distinct from P. macrotis, had a 
range continuous across the Ohio and upper Mississippi valleys from 
Virginia to Kansas and Colorado. His Kansas and Colorado specimens 
were somewhat paler than those from Virginia, indicating, Allen 
thought, intergradation with the western pallescens. He presumed 
that specimens from Kentucky and Indiana, recorded in literature as 
P. macrotis, had been misidentified, and were actually representatives 
of the newly discovered second species. With the range that he 
envisioned for the second species, Allen’s choice of Rafinesque’s old 
name, Vespertilio megalotis (type locality, lower Ohio River), as the 
name for it was a logical step. In the absence of material from the 


AMERICAN BATS—HANDLEY 155 


type region, Allen considered specimens from Burkes Garden, Va., 
to be typical. 

That Vesperiilio megalotis Rafinesque is a junior homonym of Vesper- 
tilio megalotis Bechstein (1800, p. 622) escaped Allen’s notice. Bech- 
stein’s V. megalotis is based on Le Vaillant’s (1796, p. 279) description 
of a bat that came into his tent while he was one or two day’s journey 
north of the Orange River in Great Namaqualand, South-West Africa 
(about lat. 28° S.). There is little doubt that Bechstein’s Vespertilo 
megalotis is a member of the megadermatid genus Lavia. The descrip- 
tion, except for the measurements, is an accurate one for that genus. 
No specimen of this family has been reported subsequently south of 
lat. 15° S: 

Thomas (1916, p. 127) pointed out the unavailability of the name 
V. megalotis for an American bat and proposed the substitution of 
P. rafinesquir Lesson in its place. Subsequently the two eastern 
American species of Plecotus have been known as P. macrotis (South- 
eastern States) and P. rafinesquu (North-Central States). Collections 
of Plecotus from north of the Ohio River have never been critically 
examined (e.g., Lyon, 1936, p. 86). 

Numerous specimens of big-eared bats are now available from the 
type region of Lesson’s P. rafinesquii: the Ohio Valley and upper 
Mississippi Valley States of Tennessee, Kentucky, Illinois, Indiana, 
Ohio, and West Virginia. All of those specimens are the “‘C. macrotis’’ 
of G. M. Allen. Big-eared bats from a small portion of the Appala- 
ehian Highlands in western Virginia, eastern West Virginia, and east- 
ern Kentucky are the “C. rafinesquii” of G. M. Allen. Likewise, 
specimens from the Ozark Highlands of northwestern Arkansas, and 
from the gypsum cave area of south-central Kansas, Oklahoma, and 
northern Texas are the “C. rafinesquit’”’ of G. M. Allen. Instead of 
having a continuous distribution from Virginia to Kansas as Allen 
supposed, his “C. rafinesquii”’ apparently is isolated in the central 
Appalachians, and reappears again to the westward only in the Ozark 
Highlands and beyond. 

Further, Lesson’s description of P. rafinesquii, ‘“Pelage dark gray 
above, and pale gray below ... .” comes closer to the yellowish 
brown above, grayish white below of Allen’s ‘“C. macrotis’ than it 
does to Allen’s “C. rafinesquit”’ which is reddish brown above, buffy 
below. In this connection, Neill’s observation (1953, p. 383) on the 
appearance of a live example of P. macrotis is pertinent: 

In life, the dorsal hairs were dark, grayish-black with light, horn-colored tips. 
These light tips were very inconspicuous, however, and to casual inspection the 


upper surfaces were grayish-black. The ventral hairs were dark gray with whitish 
tips which were more evident posteriorly. 


156 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


My own observations of living specimens of the mid-Appalachian big- 
eared bat (‘‘C. rafinesquiz”’ of G. M. Allen) show it to be as brown in 
appearance when living as when made up as a study skin. It could 
not be described as “‘dark gray above’”’ under any circumstances. 

Thus, as first pointed out by G. M. Allen (1916, p. 339), there are 
two species of Plecotus in the eastern United States. Current applica- 
tion of names and delineation of geographic distributions for the two 
species are incorrect, however. One species inhabits the southeastern 
and central United States. The names P. rafinesquii Lesson (=V. 
megalotis Rafinesque) (type locality, “Lower Ohio Valley’) and P. 
macrotis LeConte (type locality, ‘“Georgia’”’) are available for it. The 
earlier name is P. rafinesquii Lesson. The second species inhabits 
the central Appalachian Highlands, the Ozark Highlands, and the 
western United States, southwestern Canada, and Mexico. The 
earliest name for this species is P. townsendii Cooper (type locality, 
“Columbia River’) (Handley, 1955c, p. 147). 

Although the geographic ranges of P. rafinesquii and P. townsendii 
overlap in the Appalachian Highlands and are at least contiguous in 
the Ozark Highlands, the two forms have never been taken at the 
same locality. Frum (1948, p. 418) secured specimens of typical P. 
rafinesquti near the region inhabited by P. townsendzi in West Virginia. 
Barbour (1957, p. 141) collected typical P. townsendii within the range 
of P. rafinesquii in Powell County, Ky. Welter and Sollberger (1939, 
p. 79) reported P. townsendii from nearby Rowan County, Ky. How- 
ever, this identification is questionable since no specimen was pre- 
served (Barbour, 1957, p. 141). Typical P. rafinesquii (MCZ 6062) 
has been collected on the Osage River in Arkansas. This locality 
cannot be pinpointed but may be within the range of P. townsendit. 
A skeleton (KU 5645) of P. rafinesquii from Crawford County, Ark. 
(Black, 1936, p. 30), was secured near collecting stations of P. town- 
sendii, but at a lower elevation, in the Lower Austral zone. 

P. rafinesquii is polytypic. One geographic race inhabits the At- 
lantic and Gulf Coast regions, the other the area west of the Appala- 
chians and north of Alabama, Mississippi, and Louisiana. lLesson’s 
name P. rafinesquii can be used for the interior race, and LeConte’s 
P. macrotis for the coastal race. 

Hasirat: Plecotus rafinesquit occurs in a forested region largely 
devoid of natural caves. It has been observed most frequently in 
buildings. Natural roosting places are hollow trees, crevices behind 
loose bark, and similar arboreal retreats. Lowery (1943, p. 224) found 
it in hollow trees (as well as in buildings) in Louisiana; a specimen in 
the U.S. National Museum was collected in a hollow cypress in the 
Dismal Swamp of Virginia; and Harper (1927, p. 280) was told of one 
that had been found under dry leaves in a cypress swamp in Georgia. 


AMERICAN BATS—HANDLEY 157 


In some regions where natural caves occur, P. rafinesquit has been 
found in buildings but not in caves, despite diligent search. Thus, 
the only specimen from Florida Caverns State Park was taken in a 
building (Sherman, 1945, p. 201). Hibbard (1954, in litt.) found the 
species only in the attic of an old house at Mammoth Cave, Ky. 

Where P. rafinesquiz occurs with some frequency in caves, it usually 
has been found in small caves, or near the entrances of larger caverns. 
P. M. Smith (1954, in litt.) found an individual 25 feet from the 
entrance of a small cave in Indiana and another 50 feet from the en- 
trance of a small limestone cave in Ohio. Ganier (1954, in litt.) found 
numerous individuals 30 to 100 feet from the entrance of a small cave 
on a wooded mountainside, 400 feet above a valley floor in Tennessee. 
Hahn (1908, p. 145) observed this species in Indiana only in dim 
light near cave entrances and believed (1909, p. 619) that it is not 
truly a cave dweller. However, all records from the northern fringe 
of the range of this species are from caves or subterranean excavations 
and, it should be noted, all are for the cooler months of the year 
(October to May), indicating, possibly, a choice of some other roosting 
place during the warmer months. 

P. rafinesquir also inhabits other cave substitutes. Neill (1953, 
p. 282) reported one that was found clinging to the wall of an artificial 
limestone grotto, built for ornamental purposes and overgrown with 
ferns and mosses so as to resemble natural limestone caverns which 
are numerous around Silver Springs, Fla. Moore (1949a, p. 60) 
recorded finding big-eared bats on two occasions in a concrete culvert 
(66X75 feet) at the bottom of a ravine, some 40 feet deep, which 
probably had water running through it the year around (Florida). 
Goodpaster and Hoffmeister (1952, p. 365) found big-eared bats on 
several occasions in western Tennessee in an open cistern, 20 feet 
deep, built in the shape of a bottle, with open top about 2% feet in 
diameter extending about 3 feet above the ground, and containing 
about an inch of water at the bottom. In warm weather the bats 
hung near the top; in cold weather near the bottom. Harper (1927, 
p. 280) noted that a Plecotus had been found in a turpentine still in 
the Okefenokee Swamp. 

P. rafinesquii roosts both in occupied and abandoned buildings. 
In the Great Smoky Mountains, Stupka (1954, in litt.) found this 
species to be a regular inhabitant of a house that had been abandoned 
for about 15 years. Two or three rooms and the attic were utilized 
by the bats. He reported others that had been found in a schoolhouse 
(in summer) and in a Civilian Conservation Corps camp building. 
In the same region, the Komareks (1938, p. 148) found Plecotus in 
the attic of an abandoned schoolhouse and on the chimney, above 
the hearth, in an occupied cabin. Barbour (1957, p. 141) located a 


158 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


colony in the attic and storerooms of a large log building in a recreation 
camp. In North Carolina, Brimley (1945(?), p. [10]) reported it in 
an “old house,” and Odum (1949, p. 188) had a specimen from an 
occupied dwelling. Some South Carolina specimens have come from 
a barn (Schwartz, 1954, in litt.) and from a closet in an occupied 
dwelling (ChM specimen labels). Harper (1927, pp. 279-281) 
recorded that residents of the Okefenokee Swamp had observed 
Plecotus most frequently in buildings—on several occasions during 
services in a meeting-house, in occupied dwelling houses (on wall 
near ceiling, and on chimney), in a shanty, in a commissary, and 
beneath the flooring of a railway telephone booth. In Florida 
big-eared bats have been discovered in the headquarters building of 
an Indian reservation (E. V. Komarek, 1954, in litt.) and in the base- 
ment of an occupied dwelling in an urban area (Moore, 1949b, p. 50). 
Alabama records cite the ceiling of a small round gatehouse and the 
garret of an old abandoned grist mill (A. H. Howell, 1921, p. 28). 
In Louisiana, Vernon Bailey (field notes) found it in the attic of a 
large plantation house, and Louisiana State University specimen 
labels record it from occupied farm houses. 

Beruavior: P. rafinesqui seems to be rather unsociable. Solitary 
individuals are more commonly encountered than clusters of indi- 
viduals. Summer colonies have been recorded as follows: 5 gravid 
females, Houma, La., 12 May (Vernon Bailey, field notes); 20 or 25 
(3 collected were gravid females) in small cave in Warren County, 
Tenn., 31 May (Ganier, 1954, in litt.); breeding colony (females and 
young) in Breathitt County, Ky., 14 June (Barbour, 1957, p. 141); 
3 females and 1 small young collected near Baton Rouge, La., 17 June 
(LSU specimen labels). Five of unspecified sex were taken at 
Taylorsville, N.C., on 25 August (Brimley, 1945(?), p. [10]); and a 
group of seven, of both sexes, were collected at Cades Cove, Tenn., 
on 15 July (Stupka, 1954, in litt.). 

Winter aggregations (usually consisting of both sexes) are more 
numerous, but even at this season solitary individuals are frequently 
encountered. Stupka (1954, in litt.) collected 11 in a house near 
Gatlinburg, Tenn., on 17 November, but the bats were scattered in 
two or three rooms and the attic; seven were collected in Georgetown 
County, S.C., on 18 December (ChM specimen labels); Ganier 
(1954, in litt.) has reported small groups in caves of Warren County, 
Tenn., on 19 March and in early September; and in western Tennessee 
Goodpaster and Hoffmeister (1952, p. 365) found several big-eared 
bats hanging in a clump in an open cistern on 2 September (15 were 
collected) and on 13 March found numerous Plecotus in the same 
cistern, hanging individually and in two or three small clumps (25 
specimens taken). 


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AMERICAN BATS—-HANDLEY 159 


Many references to P. rafinesqui specify that no other species 
of bats were found in the same building or cave. Stupka (1954, in 
litt.) mentioned that the abandoned house near Gatlinburg, Tenn., 
in which he found big-eared bats for several years, was shared with 
wood rats (Neotoma). Pipistreilus was found on four occasions and 
Plecotus twice in a Florida culvert, but not on the same dates (Moore, 
1949a, p. 60). 

The sleeping P. rafinesquiti hangs head downward with its ears 
curved backward and flattened against the sides of its neck. In this 
position the anterior edge of the ear is uppermost and forms a, half 
ellipse. The posterior portion of the ear folds along its transverse 
ribs like an accordion. When the bat wakes the ears are usually 
straightened slowly (Hahn, 1909, p. 619). 

Residents of the Okefenokee Swamp told Harper (1927, p. 280) of a 
peculiar motion of the ears of these bats. One was discovered on 
the wall of a commissary when it attracted attention by “‘waving”’ its 
ears. Another that was seen several times in a railway telephone 
booth “wagged”’ its ears alternately. Several other informants also 
mentioned ‘‘ear-wagging.”’ 

P. rafinesquw must be strictly nocturnal, for it has never been 
observed in twilight flight. 

Hahn, (1908, p. 177) experimented with the ability of bats to avoid 
obstacles. His subjects were mostly Myotis and Pipistrellus but 
included one Plecotus rafinesqui. The big-eared bat struck vertical 
wires 52 out of 102 chances, while in contrast Myotis and Pipistrellus 
struck on 24 to 25 percent of the chances and Eptesicus on 67 percent. 
In these experiments there may have been a relationship between 
wingspread of bat and success of avoidance. Hahn noted that the 
flight of captive Plecotus was swift and steady and that both Myotis 
and Plecotus in attempting to escape from a captivity room repeatedly 
flew against window panes and screens with great force. 

Several references to P. rafinesquii indicate a variable agility in 
avoiding capture. One that began flying about during a service (17 
July) in the meeting-house on Billy’s Island in the Okefenokee Swamp 
was caught by hand by a member of the congregation (Harper, 1927, 
p. 280). Goodpaster and Hoffmeister (1952, p. 365) found (25 May) 
a Plecotus hanging in the upper part of a cistern that had an opening 
at the top 2 feet in diameter. When disturbed the bat first flew to 
the bottom of the cistern and there circled until forced to attempt to 


_ escape through the top. It was then captured by hand. Hahn (1909, 


| 
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| 
| 
| 
if 


p. 619) saw two of these bats on 22 February in a cave at Mitchell, 

Ind. They avoided capture and flew out of the cave into the cold 

winter air. A solitary individual in a mine tunnel in Illinois (5 Octo- 
497256—59 5 


160 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ber) avoided three persons and escaped out of the tunnel (Elder, 
1946, p. 434). An immature that Hamilton (1930, p. 308) found 
injured on a Kentucky roadway may have struck a vehicle. 

On the authority of a local informant Harper (1927, p. 280) reported 
the voice of P. rafinesqua as a “‘fizzling noise.” 

Foop: The food habits of Plecotus rafinesquii appear to be un- 
recorded. Specimens that Hahn (1909, p. 619) kept in captivity 
refused food and soon died. 

Mieration: There is no evidence of migration by this species. It 
has been found at all seasons throughout its range except in the ex- 
treme northern part, where it has been discovered only in winter 
(October to May). 

Hisernation: Whether Plecotus rafinesquit becomes lethargic in 
the winter in all parts of its range is not certain. At least under some 
circumstances it does truly hibernate. Goodpaster and Hoffmeister 
(1952, p. 365) found (13 March) big-eared bats hibernating near the 
bottom of an open cistern. ‘The bats were cold and sluggish and made 
no attempt to escape when placed ina cage. A February observation 
in the Okefenokee Swamp reported by Harper (1927, p. 280) suggests 
injury or hibernation. On the other hand, specimens found in No- 
vember in a culvert near Satsuma, Fla., had to be captured with a 
net (Moore, 1949a, p. 60). One that Elder (1946, p. 433) found in 
a mine in southern Illinois on 5 October was very alert and escaped 
out of the mine when it was approached. Two that Hahn (1909, 
p. 619) saw on 22 February in a Mitchell, Ind., cave escaped capture 
and flew out into the cold air. 

Repropuction: The season of mating and length of gestation are 
not known in this species. ‘The young usually are born in late May 
or early June, shed their milk dentition by mid-July, and reach full 
size and assume the adult pelage in August or early September. 

With regard to a female captured near Satsuma, Fla., on 5 Novem- 
ber, Moore (1949a, p. 60) reported that her ‘“‘vulva was perforate, 
reddish, and swollen. Her teats were dry and her uterus was empty.” 
Schwartz noted on specimen labels that testes were prominent in all 
males captured near Andrews, N.C., on 19 and 20 January. Good- 
paster and Hoffmeister (1952, p. 365) observed that bats taken in 
hibernation in Obion County, Tenn., on 13 March began mating 
after warm surroundings had overcome their lethargy. 

Vernon Bailey (field notes) collected females containing single 
well-developed embryos in the attic of a large plantation house near 
Houma, La., on 12 May. On 31 May, in a Tennessee cave, Ganier 
(1954, in litt.) found females with pregnancy so advanced that birth 
of the one young of each seemed imminent. Barbour (1957, p. 141) 


AMERICAN BATS—HANDLEY 161 


reported a nursery colony in Kentucky in which the young averaged 
66 mm. in total length on 22 June. 

A specimen (LSU 1846) with milk teeth still in place, M? and M? 
not fully erupted, no trace of postjuvenile molt, and total length of 
71 mm., was collected near Baton Rouge, La., on 17 June. AMNH 
70517 with some milk teeth still in place, M?® not fully erupted, no 
trace of postjuvenile molt, and total length of 84 mm. was collected 
near Quicksand, Ky., on 8 July. LSU i397, collected on 28 June in 
Nachitoches Parish, La., has mature dentition, no trace of post- 
juvenile molt, and total length of 88 mm. Individuals almost full- 
grown (total length 93, 98, 99, and 100 mm.), and molting to a post- 
juvenile pelage, have been collected in South Carolina and Alabama 
on 24 July, 17 August, and 22 August. However, UI 853, collected 
on 2 September in Tennessee, although almost full-grown (total 
length 101 mm.), shows no trace of the postjuvenile molt. 

Mott: Molt is evident on the head, neck, and underparts, but not 
on the back of ChM 33.188.3 (24 July); on head, neck, underparts, 
and rump of ChM 33.218.3 (17 August); on neck and upper back, and 
almost completed on the belly of ChM 33.218.2 (17 August); and on 
all parts of the body of USNM 180198 (22 August). 

Molting in adults has been noted only in two specimens, both of 
which show molt on all parts of the body: USNM 201714, male, 
Alabama, 4 July; and ChM 33.218.4, female, South Carolina, 17 
August. 

Plecotus rafinesquii macrotis LeConte 

Plec{otus] macrotis LeConte, 1831, p. 431. 

Plecotus lecontit Cooper, 1837, p. 72, pl. 3, fig.5. (Substitute name for macrotis.) 

Vesp[ertilio] macrotis Lesson, 1842, p. 23. (Not Vespertilio macrotis Temminck, 
1840, p. 218=Pipistrellus imbricatus Horsfield.) 

S[ynotus] Lecontit Wagner, 1855, p. 720. 

Synotus macrotis H. Allen, 1864, p. 63. (Not Synotus macrotis Wagner, 1855, p. 
719=Pipistrellus imbricatus Horsfield.) 

Clorynorhinus] macrotis H. Allen, 1865, p. 174. 

Corynorhinus rafinesquit Handley, 1955c, p. 147. 

Hototyrer: None designated. Type locality subsequently fixed 

(Miller, 1897, p. 51) near the LeConte plantation, 5 miles south of 
Riceboro, Liberty County, Ga., (see p. 162.) 
_ Disrrisution: Southeastern United States (fig. 19). Northeast- 
ward to Dismal Swamp, Va. Southwestward to Terrebonne, West 
- Baton Rouge, and Natchitoches Parishes, La.; and McCurtain and 
LeFlore Counties, Okla. Inland to Dismal Swamp, Va.; Bertie and 
Wayne Counties, N. C.; Darlington County, S. C.; Autauga and Green 
Counties, Ala.; Kemper County, Miss., and Crawford County, Ark. 
Zonal distribution, Lower Austral. Altitudinal distribution, sea level 
to about 500 feet. 


162 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Description: As for the species. In adults, hairs of upper parts 
blackish basally; tips between Sayal Brown and Snuff Brown. Hairs 
of underparts blackish basally, with a slight reddish tinge; tips 
whitish with a yellowish buff cast. 

MEASUREMENTS: See tables 9, 15. 

Comparisons: Compared with P. r. rafinesquii, P. r. macrotis has 
coloration darker, more brownish dorsally; mass effect of the under 
parts less clear white, more yellowish; and mesopterygoid fossa and 
postdentary part of palate averaging narrower. 

Remarks: Body size averages small in populations in southern 
Alabama, Mississippi, and Louisiana (tables 9, 15). Specimens from 
other parts of the range of P. r. macrotis (Atlantic coast and Arkansas, 
Oklahoma, and northern Louisiana average larger in body size, and 
resemble P. r. rafinesquit in this respect. There is no corresponding 
variation in coloration. Morphologically large and morphologically 
small populations of P. r. macrotis are similarly colored and are equally 
distinct from P. r. rafinesquit. 

Intergradation between P. r. macrotis and P. r. rafinesquit is shown 
in numerous specimens. Some from Sevier County, N.C., dorsally 
are like typical P. 7. rafinesquit, but some show traces of the yellowish 
ventral coloration characteristic of P. r. macrotis. Individuals from 
Marshall and Weaverville, N.C., and Huntsville, Ala., are more per- 
fectly intermediate in all details of coloration, but are nearer P. r. 
rafinesquit. A specimen from Leighton, Ala., approaches Gulf Coast 
populations of P. 7. macrotis in size, but is like P. r. rafinesquit in colora- 
tion. Specimens from central and northern Louisiana are unac- 
countably variable in coloration but average nearer P. r. macrotis. 
Characters of P. 7. macrotis apparently are most accentuated in coastal 
plain populations. Those of P. r. rafinesquii are found west of the 
Appalachians, from Tennessee northward. 

Although there is no assurance that the specimens upon which 
LeConte based his description of Plecotus macrotis came from his 
plantation near Riceboro, it is reasonably certain that they came from 
Georgia. The original description stated the range simply as the 
“United States,” but in a later note on macrotis, LeConte (1855, p. 
437) specified that it “inhabits Georgia.” There are two LeConte 
specimens of Plecotus in the U.S. National Museum, perhaps the very 
specimens upon which the description of macrotis was based. Both — 
are skulls only, labeled simply “United States.”” Both could very 
well have come from the same coastal locality, possibly from the Le- _ 
Conte Plantation. Since coastal Georgia is an area probably typical 
of the coastal race of P. rafinesquii, and lacking evidence that Le- 
Conte’s description was based on specimens from elsewhere, I follow 
Miller (1897, p. 51) in designating the vicinity of the LeConte Planta- | 


AMERICAN BATS—HANDLEY 163 


tion, 5 miles south of Riceboro, Liberty County, Ga., as the type 
locality of P. r. macrotis. 

SPECIMENS EXAMINED: A total of 56 from the following U.S. local- 
ities: 

Vireinia: Nansemond County or Norfolk County: Lake Drummond, Dismal 
Swamp, 1ls(USNM). Sours Carourna: Charleston County: Hampton Plantation, 
Santee, 4s (ChM). Darlington County: Society Hill, 28 (USNM). Dorchester 
County (?): Summerville, 1s (ChM). Georgetown County: Darlington Plantation, 
9s (ChM). WNo ezact locality, 23 (USNM). Groratia: Early County: Kesler, la 
(MCZ). ‘United States’? [=Liberty County: 5 miles south of Riceboro?], 2s 
(USNM). Fiorina: Hendry County: Big Cypress Seminole Indian Reservation 
Headquarters, 27 miles east-southeast of Immokalee, 1s (USNM). Marion 
County: Silver Springs, la (USNM). Atasama: Autauga County: Autaugaville, 
2a (USNM). Mississippi: Hancock County: Bay St. Louis, 1a (USNM). West- 
ville, Is (USNM). Lovistana: East Baton Rouge Parish: Baton Rouge, and 3 
miles south of Baton Rouge, 7s (LSU), 1s (USNM). Madison Parish: Tallulah, 
2a (USNM); Waverly, 1s (LSU). Natchitoches Parish: Kisatchie, 3s (LSU). 
Terrebonne Parish: 9 miles southeast of Houma, 3s (USNM). West Baton Rouge 
Parish: Lobdell, ls (USNM). Arxansas: Crawford County: Mulberry, 1b (KU). 
Sevier County: Near Horatio, ls (OAM). County uncertain: “Osage River,” la 
(MCZ). Oxtanoma: McCurtain County: 2.5 miles west of Smithville, 5s; UOMZ). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: 


NortH Carourna: Bertie County: No exact locality (Brimley, 1905, p. 22), 
Wayne County: Goldsboro (Brimley, 1905, p. 22). SouraH Carouina: Beaufort 
County: Hardeeville (Miller, 1897, p. 52). Georata: Charlton County: Okefe- 
nokee Swamp (Harper, 1927, p. 279). Ware County: Okefenokee Swamp (Harper, 
1927, p. 280). Fuoripa: Alachua County: Micanopy (H. Allen, 1864, p. 65). 
Jackson County: Florida Caverns State Park, near Marianna (Sherman, 1945, 
p. 201). Levy County: Gulf Hammock (Pearson, 1954, p. 474). Orange County: 
Zellwood (Moore, 1949b, p. 50). Putnam County: 1.6 miles north of Satsuma 
(Moore, 1949a, p. 60). ALABAMA: Greene County: Eutaw (H. Allen, 1864, p. 65). 
Hale County: Greensboro (Miller, 1897, p. 52). Mussissrppr: Kemper County: 
No exact locality (H. Allen, 1864, p. 65). Louisiana: Lincoln Parish: Ruston 
(Lowery, 1943, p. 224). Orleans Parish: New Orleans (Lowery, 1936, p. 18). 
Rapides Parish: Alexandria (Dalquest & Werner, 1954, p. 156). OxkLAHOMA: 
Le Flore County: Houston (Blair, 1939, p. 102). 


Plecotus rafinesquii rafinesquii Lesson 


Vespertilio megalotis Rafinesque, 1818, p. 446. (Not Vespertilio megalotis 
Bechstein, 1800.) 

Plecotus rafinesquii Lesson, 1827, p. 96. 

Plecotus macrotis Garman, 1894, p. 58. 

Corynorhinus macrotis Butler, 1895, p. 86. 

Corynorhinus rafinesquii Handley, 1955c, p. 147. 

Houotyren: None designated. Type locality: “. . . lower parts of 
| the Ohio, the Wabash, Green River, Barrens, Prairies, and the states 
of Indiana, Illinois, etc.’’ (Rafinesque, 1818, p. 446). Herein 
restricted to Mount Carmel, Wabash County, Ill. (see p. 164). 


164 PROCEEDINGS OF THE NATIONAL MUSEUM YO. 110 


DistrisuTion: East-central United States (fig. 19). North to 
Nicholas County, W. Va.; Adams County, Ohio; Putnam County, 
Ind.; Wabash and Alexander Counties, Ill. West to Putnam County, 
Ind.; Alexander County, Iil.; and Obion County, Tenn. South to 
Obion County, Tenn.; Colbert and Madison Counties, Ala.; Towns 
County, Ga.; and Macon, Transylvania, and Alexander Counties, 
N.C. East to Alexander County, N.C. and Nicholas County, 
W. Va. Zonal distribution: Upper Austral and lower portions of 
Transition. Altitudinal distribution: From 400 feet above sea level 
in western Tennessee to 3,850 feet in western North Carolina. 

Description: As for the species. In adults, hairs of upperparts 
blackish basally; tips between Sepia and Buffy Brown. Hairs of 
underparts black at base, almost pure white at tip. 

MEASUREMENTS: See tables 9, 15. 

Comparisons: For comparison with P. r. macrotis, see page 162. 

Remarks: Rafinesque’s specimens, which led to the description of 
P. rafinesquii by Lesson, are not known to be still in existence. 
This material is said to have been secured on the ‘‘. . . lower parts 
of the Ohio, the Wabash, Green River, Barrens, Prairies, and the 
states of Indiana, Illinois, ete.” (Rafinesque, 1818, p. 446). The 
big-eared bat is not known to occur as far north as the Green River, 
and it has not been taken on the barrens or prairies of Illinois, Indiana, 
or Kentucky. Neither has it been taken on the banks of the Ohio 
River, although it undoubtedly occurs there. However, there are 
specimens in the U.S. National Museum secured in 1896 and 1903 at 
Mount Carmel on the Wabash River in Illinois, a region specifically 
mentioned by Rafinesque. In addition, the species is also known to 
have occurred at other localities in Illinois and Indiana. Conse- 
quently, I propose that the type locality of Plecctus rafinesquii rafines- 
quit Lesson be restricted to Mount Carmel, Wabash County, Ill. 

Plecotus rafinesquit seems to be rare near the northern limits of its 
range. Despite repeated investigations, the caves and mines of 
Illinois have produced but two recent records (Elder, 1946, p. 4383; 
and Smith and Parmalee, 1954, p. 204); Indiana, two (P. M. Smith, 
1953, specimen submitted to U.S. Nat. Mus.; and R. E. Mumford, 
1954, in litt.); Ohio, one (Goslin, 1954, p. 480); West Virginia, one 
(Frum, 1948, p. 418); and Kentucky, two (Barbour, 1957, p. 141; 
and Hibbard, 1954, in litt.). 

Plecotus may have occurred in Pennsylvania within historic times, 
but it has not been found there by recent investigators (Mohr, 1954, 
p. 21). Harrison Alien (1864, p. 64) stated that ‘I am informed by 
Prof. Baird that specimens of a Synotus, probably of this species 
[Synotus macrotis], were received some years ago by the Smithsonian 
Institution, from Meadville, Pa., but that they have become in some 


AMERICAN BATS—HANDLEY 165 


way misplaced and are not now to be found.” I have checked the 
early catalogs of the U.S. National Museum and have failed to find 
any record of big-eared bats from Pennsylvania. In 1857 and again 
on two occasions in 1861 specimens were received from J. T. Thuckston 
of Meadville, Pa. These were Condylura, Myotis, Tamiasciurus, 
Peromyscus, and Microtus. If big-eared bats were received from the 
same source they apparently were never cataloged. 

SPECIMENS EXAMINED: A total of 50 from the following U.S. local- 
ities: 

NortH Carouina: Buncombe County: Weaverville, 1s (USNM). Cherokee 
County: 4 miles east-southeast of Andrews, 1s (AS). Madison County: Marshall, 
ls (MCZ). Swain County: 14 miles northeast of Andrews, 5s (AS); Forney 
Creek, 2,400 feet, 9 miles northwest of Bryson City, 1s (USNM). Gzorarta: 
Towns County: Young Harris, ls (USNM). Auasama: Colbert County: Forster’s 
Mill on Town Creek, 5 miles east of Leighton, ls (USNM). Madison County: 
Monte Sano, 1,600 feet, southeast of Huntsville, Is (USNM). TENNESSEE: 
Blount County: Cades Cove, 1,750 feet, 1s (USNM). Campbell County: Well 
Springs, Powell Valley, ls (OSM). Obion County: 10 miles northeast of Tipton- 
ville, 6s (UI). Sevier County: Sugarlands area, 1,900 feet, 3 miles south of 
Gatlinburg, 13ab (USNM). Warren County: McMinnville, 1s (CNMH). 
Kentucky: Breathitt County: Noble, 5s (UK); Quicksand, Is (AMNH). 
Inuinois: Alexander County: One-half mile north of Olive Branch, 1s (INHS). 
Wabash County: Mount Carmel, 2a (USNM). Inprana: Lawrence County: 
Mitchell, Upper Twin Cave, 1s (UI), ls (USNM). Washington County: Near 
Salem, ls (OSM). Onto: Adams County: Waggoner Ripple Cave, Green Town- 
ship, ls (OSM). West Virainita: Nicholas County: Collison Cave, near Mt. 
Lookout, 1s (CM), 2s (GF). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: 


NortH Carouina: Alexander County: 10 miles northwest of Taylorsville (G. M. 
Allen, 1916, p. 351). Macon County: Highlands, 3,850 feet (Odum, 1949, p. 188). 
Transylvania County: Pink Beds, head of South Fork of Mills River, 3,300 feet, 
Pisgah National Forest (Oberholser, 1905, p. 9). THNNESSEE: Franklin County: 
“Southeast portion’? (Ganier, 1954, in litt.). Sevier County: Greenbrier Cove 
(Komarek and Komarek, 1938, p. 148). Warren County: Near Hubbard’s or Bat 
Cave, 8 miles north of Beersheba Springs (Ganier, 1954, in litt.). Kenruckxy: 
Edmonson County: Mammoth Cave (Hibbard, 1954, in litt.). Warren County: 
Bowling Green (Garman, 1894, p. 58). Iniinois: Alezander County: Eleo (Elder, 
1946, p. 433). Inprana: Putnam County: 5 miles southwest of Greencastle 
(Butler, 1895, p.86). Washington County: Near Saltillo (Mumford, 1954, in litt.). 


Plecotus townsendii Cooper 
Synonymy is given under the subspecies. 


Distrispution: Central Appalachian Highlands, Ozark Highlands, 
western United States, southwestern Canada, and Mexico south to 
the Isthmus of Tehuantepec (figs. 22-24). 

Description: Adult coloration: Hairs of upperparts gray or slate 
at base; color of tip varying from pale cinnamon brown to blackish 


166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


brown; division between colors of tip and base often not sharp; hairs 
at posterior bases of ears frequently paler than those of other parts of 
dorsum. Mass effect darkens with wear. Hairs of chest and belly 
slate, gray, or brownish at base, brownish or buff at tip, the division 
between the colors often not sharp; hairs of chin and throat like 
remainder of underparts or with darker colored tips; hairs bordering 
membranes with bases colored like tips. Immature like adult, except 
darker or duller throughout. 

Size varies from medium to large for the subgenus; forearm varies 
from medium, relative to skull length, to long; presternum with lobes 
not expanded (ventral lobe narrower at tip than lateral lobe) (fig. 5). 
Rostrum varies from weak and depressed to strong and not depressed; 
anterior nares relatively small to relatively large, and posterior 
border rounded, or with angular shape; intermaxillary notch usually 
relatively broad; median postpalatal process usually styliform, with 
narrow base (sometimes triangular); auditory bulla averages relatively 
large; brain case averages relatively shallow for subgenus. I! with 
secondary cusp varying from prominent to absent (fig. 2); P* with 
small anterointernal cingular cusp varying from present to absent. 
Tragus (possibly also auricle) averages relatively long; cross-ribs on 
interfemoral membrane average 10 or more. 

ComParRIsons: See pages 138, 140, 142, and 151. 

ReELATIoNsHIps: Of the three Recent species of the subgenus 
Corynorhinus, P. townsendii has the largest geographic range and the 
greatest environmental tolerance. Its geographic range almost spans 
the continent from east to west and extends through 35 degrees of 
latitude. It inhabits broadleaf deciduous forests, coniferous forests, 
prairies, and desert scrub; cool, humid, rain forests and hot, arid, 
deserts; coastal lowlands and high mountains; areas with great 
annual temperature variation and areas with slight annual temperature 
variations. 

The other species of the subgenus are much more restricted: P. 
mexicanus to the cool, humid, highlands of northern and central 
Mexico; P. rafinesquii to the warm, humid, lowlands of the south- 
eastern United States. 

P. mexicanus and P. rafinesquti appear to be genetically stable and 
only slightly variable, whereas P. townsendii is so variable that several 
geographic races are recognized. P. townsendit appears to be the 
most plastic, progressive species of the genus Plecotus. 

Remarks: Plecotus townsendii was long known only by the Colum- 
bia River specimens upon which Cooper based his 1837 description. 
Beginning with Harrison Allen in 1864, P. townsendii was confused by 
all authors with P. rafinesquit (=P. maerotis) until the two species 
were recognized as distinct by G. M. Allen in 1916. Allen continued 


AMERICAN BATS—HANDLEY 167 


the confusion of names, however, believing P. rafinesquit and P. 
macrotis to be the names applicable to the two species. It is now 
evident (p. 156) that both of these names apply to a single species of 
the southeastern United States, and that P. townsendii is the name 
to be used for the wide-ranging species inhabiting parts of Canada, the 
United States, and Mexico. 

Hasirat: In a species so widely distributed as P. townsendii, the 
formulation of a generalized description of its habitat preferences is a 
difficult matter. For the brief description of such a distributional 
pattern, the life zones of Merriam are the most useful of the numerous 
ecological unit concepts that have been proposed. Such units as the 
biotic provinces, faunal areas, biotic areas, and biotic districts of 
American authors (Cockrum, 1952, pp. 9-15) are useful in small scale, 
as in describing the distribution of an animal in a county or a state, 
but they are not adapted to definition of a range continental in scope. 

P. townsendit is an inhabitant chiefly of the Upper Sonoran and 
Transition life zones, ranging in some places into the cooler Canadian 
zone and in others into the warmer Lower Sonoran zone. Eastern 
populations occupy parts of the humid subdivisions of the Upper 
Sonoran (Upper Austral) and Transition zones. 

This species appears to be absent or generally uncommon in prairie 
and extreme desert habitats but may be relatively common in a 
variety of others. Dalquest (1947, pp. 21-24) provided excellent notes 
on habitats of the species in California (west of the Great Valley). 
Typical are: Cultivated valleys bordered by broad-leafed trees and 
dense thickets of brush; nearby hills with extensive grassy slopes, 
groves of oaks, areas of chaparral, and forests of coniferous trees and 
madrofia. Oak-covered hills just below the juniper and pifon belt. 
Coastal lowland supporting dense, ocean-side vegetation, such as 
brush and lush annuals. 

Johnson, et al. (1948, p. 343) in the Providence Mountains of the 
Mohave Desert found this bat in caves and tunnels near the boundary 
between the yucca belt of the lower slopes and the pifion-juniper belt 
of the upper slopes of the mountains. In the Pocatello region of 
Idaho, Whitlow and Hall (1933, p. 243) observed Plecotus in mine 
tunnels in the juniper belt lying between the Douglas fir-aspen zone 
of the mountain tops and the sagebrush zone of the valleys. 

Most Mexican records for P. townsendiz are from arid regions. 
Thus, Hooper (1955, p. 5) found this species at San Andrés where the 
hills were grass covered and the nearby mountains were arid, mostly 
grassy, but with pines and deciduous oaks scattered over the upper 
slopes, and forming thickets and forest on the canyon floors. How- 
ever, Goldman (field notes) collected P. townsendii together with P. 
mexicanus at more humid localities in Guanajuato and Zacatecas where 


168 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


oak, pine, juniper, cypress, madrofia, and mansanita were prominent. 

Bailey (1936, p. 387) noted big-eared bats in the pine-fir-hemlock- 
broadleaf deciduous forest of western Oregon. Findley and Negus 
(1953, p. 237) reported a specimen captured at the edge of a spruce-fir 
forest in Colorado. 

In Kansas, Oklahoma, and Texas, Plecotus occurs in gypsum caves 
in the zone of meeting of tall-grass and short-grass prairies. Ozark 
and Appalachian populations inhabit caves mostly in oak-hickory 
forests. Originally, there were few natural openings in these forests, 
and even today cultivated areas are mostly restricted to narrow bands 
along valley bottoms. Some Appalachian caves frequented by big- 
eared bats are situated in unbroken forest several miles from meadow 
and farmland. 

Roostine situs: P. townsendii apparently is a true cave bat, for 
unlike P. rafinesquit it has not been known to roost in trees. Many 
populations of the western United States and Mexico have found to 
their liking such man-made structures as abandoned mine tunnels and 
pits, wineries, missions, stage stations, etc., and less frequently, 
occupied human dwellings. Kast of the Rocky Mountains, P. town- 
sendii has been recorded only in caves; never in houses. In some areas 
P. townsendii has been quick to appropriate man-made structures as 
roosting places. For example, Townsend (1839, p. 325) observed 
that it was already a rather common inhabitant of the Hudson’s Bay 
Company fur storehouses of the Columbia River district, not long 
after the construction of the first buildings in that region. 

Pearson, et al. (1952, p. 317) theorized that since natural caves are 
few in California, and since most known colonies of Plecotus in that 
area inhabit man-made structures, it must have been rare in California 
before the advent of civilization. However, beyond a certain point, 
increasing the number of roosts does not seem to increase the popula- 
tion. Large numbers of mine tunnels dug in the western foothills 
of the Sierra Nevada in the last half of the nineteenth century were 
subsequently abandoned. Despite the prevalence of apparently suit- 
able roosts, P. townsendii does not seem to be more numerous in that 
part of California than elsewhere. 

Roosts may be classified according to the use made of them by the 
bats. For P. townsendii there are at least four categories: Winter 
hibernating roosts, occupied by both sexes; summer nursery roosts, 
utilized by females and young; ordinary summer roosts, inhabited 
mostly by males; and summer night-time roosts, used probably by 
both sexes. With regard to the latter, Dalquest (1947, p. 26) has 
noted that P. townsendii, in common with several other species of 
bats, retires to a convenient cave or building to hang up and rest 
after feeding flights at night. By taking advantage of this habit, 


AMERICAN BATS—HANDLEY 169 


a collector may utilize the night-time roost as an effective trap. 
Two small buildings at Lebec, Calif., served as bat traps for Dalquest 
on 4 June 1945. At midnight the traps held a Plecotus townsendii, 
6 Myotis yuwmanensis, and 2 Myotis volans. Earlier in the night 
these same traps caught 11 Myotis yumanensis, 3 Myotis volans, 
1 Myotis californicus, and 4 Antrozous pallidus. Three Myotis 
thysanodes were taken later in the traps. Often the night-time 
retreats are uninhabited by bats during daylight hours. 

There is much variation in the selection of specific microhabitats 
within the roosting structures chosen by various kinds of bats. Many 
authors have noted the preference of P. townsendii for an open ceil- 
ing, wall, or beam rather than a crack or crevice as a roosting site. 
Apparently this species seldom crawls away from the spot where 
it first lands, and thus normally is found hanging in places open 
enough to be reached by flying. In Kansas caverns Twente (1955b, 
p. 721) observed that this bat characteristically roosted on very rough 
walls. Wind currents and consequently temperature fluctuations were 
often great in such places. However, the bats usually rested in slight 
concavities or behind projections on the rough walls. In these pockets, 
air movement was slight and the temperature relatively constant in 
contrast to the gross conditions prevailing a few inches away. 

The intensity of light in roosting sites of P. townsendii varies 
from the total darkness of deep caverns or tunnels, to semidarkness 
in houses, small caves, and near the entrances of deeper structures. 
With regard to Californian populations, Dalquest (1947, p. 24) re- 
ported that all retreats of Plecotus that he had examined were dark or 
but dimly lighted. In the best lighted roosts the bats could be seen 
without the use of a flashlight once the observer’s eyes had become 
accustomed to the darkness. Lights were necessary for observation 
in the other retreats. 

Likewise, humidity seems to be usually of secondary importance in 
roost-site selection. Twente (1955b, p. 716) noted an apparent indif- 
ference of the big-eared bat to varying degrees of humidity (90 to 21 
per cent in hibernating sites). Beer and Richards (1956, p. 34) re- 
ported a similar variation (100 to 38 per cent) in the relative humidity 
of E’ptesicus hibernating sites. 

Temperature appears to be the critical factor. Twente (1955b, p. 
706) observed that bats must rest in habitats whose temperatures are 
suited to their metabolic needs. Thus, there may be variation in 
choice of roost according to season, time of day, weather, sex of bat, 
etc. For example, hibernating bats must roost in cool places to keep 
metabolism low so that energy reserves will be conserved. During the 
feeding season warm roosting sites permit digestion and assimilation 
of food to proceed. In general, P. townsendii roosts in warm places 


170 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


after having eaten and in cool places when food is scarce. In addition, 
it has been postulated that warm sites may be required for the satis- 
factory progress of some reproductive processes (pp. 171, 182). 

The bat’s perception of the microclimate of its roosting site does 
not seem to be especially good. Twente (1955b, p. 714) determined 
that P. townsendii apparently locates a suitable roosting spot by a 
trial and error system. If the spot first selected is unsuitable the 
bat awakens and moves to another place. Eventually optimum con- 
ditions may be located. The social tendency of the bats to seek their 
Own species increases the probability of most of the individuals 
finding suitable sites. 

Sociapiuity: P. townsendii seems as a rule not to associate in its 
daytime and hibernation roosts with other species of bats, although 
scattered individuals of other normally colonial species occasionally 
may be present. Often such joint habitation does not actually bring 
the bats into close contact, for they may occupy different parts of the 
roosting structure or may occupy the same part at different times. 

Goldman (field notes) found P. townsendiit hibernating with P. 
mexicanus in a cave and in an abandoned mine shaft. At one time or 
another Dalquest (1947, pp. 21-23) noted P. townsendii in the same 
retreats as Antrozous pallidus, Myotis californicus, M. evotis, M. 
thysanodes, M. volans, M. yumanensis, and Tadarida mexicana. Stager 
(1939, p. 226) observed large numbers of P. townsendii, Myotis velifer, 
and Macrotus californicus, together with smaller numbers of four other 
species of bats, in an abandoned mine shaft. In addition to the species 
already noted, Pearson, et al. (1952, p. 275) found AZyotis lucifugus, 
M. subulatus, and E’ptesicus fuseus roosting with P. townsendit. Two 
M. subulatus were actually in a cluster of hibernating Plecotus. Ihave 
found numerous scattered dormant individuals of Myotis lucifugus, 
M. sodalis, and Pipistrellus subflavus only a few inches from hibernating 
clusters of Plecotus townsendii in a West Virginia cave. 

Intraspecific sociability in P. townsendii is seasonally variable. 
Nursery colonies of females and young are common in summer, but 
solitary pregnant females are frequently encountered. Males at this 
season are usually, but not always, solitary. Both sexes have been 
found together in summer roosts, though usually not clustering to- 
gether. Winter aggregations may include a random assortment of 
both sexes, or, according to the physical aspect and atmospheric con- 
dition of the roost, males and females alike may tend to be solitary. 
If there is an average seasonal divergence in roost-site selection by the 
sexes, probably the female prefers cooler winter roosts and warmer 
spring and summer sites than the male. 

Dalquest (1947, p. 22) noted females with newborn young gathered 
so closely together that the edges of the cluster appeared to be a solid 


AMERICAN BATS—HANDLEY 171 


line. The bats normally pack so tightly in the cluster that their 
number is usually considerably underestimated in a casual observa- 
tion. As shown by Twente (1955b, pp. 707, 714, 723) the function of 
the cluster is seasonally variable. 

Contrary to the opinion of many authors, the winter cluster func- 
tions to protect the bats from heat rather than from cold. The body 
temperatures of the clustered bats approach the temperature of the 
substrate upon which they are resting, whereas the body temperature 
of the solitary bat is more affected by the surrounding air temperature. 
Low body temperatures and consequent low metabolic rates are im- 
perative to insure conservation of fat reserves through the hibernating 
season. Beer and Richards (1956, p. 39) have demonstrated for 
Eptesicus that in some circumstances there may be a delicate balance 
between length of fasting (hibernating) season and quantity of stored 
fat in the bat. Unusually high metabolism in the bat or abnormally 
protracted cold weather result in starvation of the bat. 

Since temperatures within spring and summer clusters may exceed 
the environmental temperatures, heat conservation seems to be the 
primary function of the cluster at these seasons. The higher tempera- 
tures presumably would aid digestion and assimilation of food. 
Twente found that Myotis moved to the cooler parts of caves and 
assumed deeper torpidity, as in winter hibernation, when stormy 
weather prevented their normal summer foraging flights. Warmer 
sites were chosen and higher cluster temperatures prevailed after the 
bats had eaten. Pearson, et al. (1952, pp. 303, 311) suggested that 
gestation, lactation, and other reproductive processes also might be 
facilitated by the elevated temperatures maintained in clusters. 
Clustered young, their metabolism possibly accelerated by the higher 
temperatures, may grow more rapidly than solitary young. 

According to the observations of Pearson, et al. (1952, p. 275) in 
central California and Twente (1955b, p. 724) in Oklahoma and 
Kansas, nursery colonies of pregnant females begin to form early in 
April and increase gradually well into May. The colonies may consist 
of individuals from diverse hibernating sites. Varying from a few to 
200 individuals, these colonies remain more or less intact until after 
the young are weaned in late summer. The bats ordinarily return to 
the same sites—buildings, mine tunnels, or caves—year after year. 

Porutation: By studying and banding nursery colonies, Pearson, 
et al. (1952, p. 314) were able to calculate probable survival rates in 
P. townsendii. Although differences in habits of males and females 
render it difficult or impossible to determine the sex ratio in the adult 
population, the ratio is about 50 : 50 in the young. Only 5.5 per cent 
of 470 females examined in nursery colonies were neither pregnant nor 
lactating (reproductive failure is more common in young females than 


172 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


in adults). Basing their figures on these facts and on recovery of 
banded individuals, Pearson, et al., reasoned that a colony of 100 fe- 
males should produce about 45 young females in a season. Of this 
number, 20 would survive to join the colony as breeding adults the 
following season. Assuming an average survival of 80 per cent of these 
bats each year after their first and an equal rate of mortality for adults 
of all ages, the colony would, as they found through three years of 
observations, remain static at about 100 breeding females each season. 
Age composition of such a colony would be as in table 6, with the me- 
dian age 3.1 and the average age 5.0 years. Figures beyond the 3-year- 
old age class are theoretical. 


TaBLE 6.—Theoretical age composition of a nursery colony of 100 female Plecotus 
townsendii. (Modified from Pearson, et al., 1952, p. 315.) 


ACOH NYyCArSe.. wen = sone se 1)2/3,4/]5] 6) 7) 8] 9 j10 {11 12 J13 |14 |15 |16 |17 {18 | 19 


Number of individuals-_--__-- ZONLEH1S OWS WS 40 SS 2 [2 ty Py ee Pole a 


Pearson, et al. (1952, p. 316) estimated a population density of one 
P. townsendii per 310 acres on 62,000-acre Santa Cruz Island off the 
coast of California. The island was shared with at least an equal 
number of Antrozous pallidus and an undetermined number of Myotis 
californicus. Basing their estimate on the spacing of known colonies, 
these authors supposed that on the mainland of California the popula- 
tion density might be as small as one Plecotus per 419 acres, but 
supposed that in actuality each Plecotus probably had available con- 
siderably less than 419 acres. This would be especially true if un- 
known food habits restrict the hunting of P. townsendit. By doubling 
the estimated total of individuals in nursery colonies, Twente (1955a, 
p. 387) determined that there might be 300 to 500 P. townsendii in 
the gypsum caves of extreme northern Oklahoma and adjacent parts 
of Kansas. For the same area he estimated 25,000 to 50,000 Tadarida 
brasiliensis, 15,000 to 20,000 Myotis velifer, and 200 to 400 Antrozous 
pallidus. No estimate was made for H’ptesicus fuscus and Pipistrellus 
subflavus which were present in smaller numbers. 

How populations of P. townsendii are limited is not clear. Probably 
the same factors do not operate throughout the bat’s range. Twente 
(1955a, p. 388) calculated that the yearly mortality could not exceed 
33.3 per cent in animals with a reproductive potential of only one 
young per year if the population was to maintain a constant level. 
On this basis the daily population decrease could not be more than 
1.1 per thousand. Twente suggested that in Kansas and Oklahoma 
the observed predation on bats by rat snakes, raccoons, hawks, owls, 


AMERICAN BATS—HANDLEY 173 


and other animals might account for such a loss and thus would be 
the primary limiting factor. Rice (1957, p. 30) postulated that 
predation by rat and corn snakes, opossums, owls, and cockroaches 
was the most important mortality factor among populations of 
Myotis austroriparius in Florida. 

The possibility of antisocial tendencies and consequent need of 
some degree of privacy from other species of bats might be a limiting 
factor for P. townsendii where roosting places are not abundant. 
Pearson, et al. (1952, p. 317) believed it most likely that the popula- 
tion level of this species in California depended upon the number 
of suitable winter roosting sites and the number of summer roosting 
sites surrounded by adequate feeding territory. Their data indicated 
that disease, predation, and amount of food and water probably did 
not limit the Californian populations. They observed that colonies 
remained static in size, year after year, and believed that a species 
population increase might depend on establishment of new nursery 
colonies. 

How new nursery colonies become established is not known. 
Pearson, et al. (1952, p. 317) determined that when a satisfactory 
nursery site becomes available near an already-established colony, 
the colony does not divide to make use of both sites, but uses either 
one or the other, or alternates between the two. Consequently, they 
thought, for a new nursery colony to become established, some 
females either through chance, faulty navigation, or some inner 
drive must find a suitable roosting place outside of the range of an 
existing colony. Furthermore, they believed it necessary for several 
females to arrive at this place almost simultaneously if gestation 
were to proceed normally, since females are gregarious and may 
depend upon clustering to keep their body temperatures high. 

These hypotheses might serve to explain the evident slowness of 
Plecotus in reoccupying apparently suitable territory to the north 
of their present geographic range since the retreat of the Wisconsin 
glaciers. 

PARASITES AND DISEASE: No ticks, lice, fleas, or bedbugs have 
been reported on P. townsendit, and this bat is not particularly noted 
for heavy infestations of other ectoparasites. 

Winged parasitic flies of the family Streblidae (Zrichobius cory- 
norhini Cockerell and Trichobius quadrisetosus Kessel) appear to be 
encountered most often. I have noted flies of this family on specimens 
of Plecotus from Veracruz and West Virginia. Others have reported 
them as follows: Sprague (1938, p. 500) an infestation in Oklahoma, 
Wilson (1946, p. 86) in West Virginia, Dalquest (1947, p. 30) no 
more than one per bat in California, Krutzsch (1955, p. 458) up to 
three or four per bat on fur and membranes of specimens from “‘western 


7A: PROCEEDINGS OF THE NATIONAL MUSEUM You. 110 


North America,” and Smith (1934, p. 63) up to eight or ten per bat, 
mostly in the ears, on specimens from Kansas and Oklahoma. 

Wingless parasitic flies of the family Nycteribiidae (Baszlia antrozot 
Townsend and Basilia corynorhini Ferris) (Stiles and Nolan, 1931, 
p. 715) seem to be generally less common on P. townsendiz, although 
Dalquest (1947, p. 30) found up to five per bat in California and 
Krutzsch (1955, p. 458) up to two or three per bat in ‘‘western North 
America.”’ 

Sprague (1938, p. 500) reported mites on specimens from Oklahoma, 
and Krutzsch (1955, p. 458) found up to eight or nine mites of the 
genus Ichoronyssus per bat on the membranes of specimens from 
‘western North America.” 

One of four California specimens of P. townsendii examined by 
Mitchell (1956, p. 444) was infected with Trypanosoma vespertilionis. 

Twente (1955a, p. 387) discovered a possibly diseased specimen 
of P. townsendii in a cavern in Kansas on 31 October. It was a fat 
male, and although it had a rectal temperature of 93.9° I. it was 
very feeble and was unable to fly. A week later it was found dead. 

Foop: Apparently no one has observed P. townsendii engaged in 
its normal feeding activity. Little is known of its food habits. 
Hamilton (1943, p. 104) reported that all specimens that he had 
examined contained only the remains of Lepidoptera. Pearson, 
et al. (1952, p. 317) believed that P. townsendii eats moths and other 
insects (probably mostly flying insects). In stomachs of P. townsendit 
and Myotis velifer hibernating together in an Oklahoma cave, Sprague 
(1988, p. 500) found remains of insects and hairs of both species 
of bats. Inasmuch as the bats were heavily parasitized with bat 
flies (Streblidae), Sprague supposed that the bats might be feeding 
on the parasites. Twente (1955b, p. 713) observed no evidence 
of feeding among hibernating populations that he studied in Oklahoma 
and Kansas. 

Pearson, et al. (1952, p. 274) found it difficult to induce captive big- 
eared bats to feed themselves. The bats were offered mealworms, 
supplemented occasionally by percomorph oil. Most could be taught 
in a day or so to accept decapitated mealworms forced into their 
mouths. Self-feeding was encouraged by stringing mealworms on 
wires along the sides of the cage. Some individuals eventually learned 
to eat worms from a dish, although many had to be hand fed through- 
out their captivity. As many as 40 mealworms might be taken daily. 
The most successful captives were ones that had learned to feed them- 
selves and were kept in a cage large enough to permit flight. However, 
pregnant individuals, even though eating well, aborted or resorbed 
their embryos. The maximum period of survival in captivity was 
about six months. Some individuals survived without food when kept 


AMERICAN BATS—HANDLEY 5 


at a temperature of 45° F. in almost-covered aquaria stocked with 
drinking water. 

Movements: Cockrum (1956a, p. 51) classified the movements of 
bats in three major categories: Diurnal movements (routine daily 
movements in quest of food and water), local migrations (irregular 
movements from one roost to another without regard for season), and 
seasonal migrations (seasonal movement from one roost to another, 
perhaps as little as ten miles or less apart). These categories overlap 
and are not clearcut. Consequently, some movements are difficult to 
classify. 

DiurRNAL MOVEMENTS: H. H. T. Jackson (field notes) encountered 
P. townsendii in the month of August in a cave at the east side of 
Montezuma Well, Arizona. During the day the bats occupied the 
dark interior parts of the cave, but late in the afternoon they were to be 
found in the partially lighted section near the entrance. At this time 
they were unusually wild and the slightest noise or shadow caused 
them to retire to the deeper parts of the cave. Twente (1955b, p. 727) 
described a similar light-sampling behavior of P. townsendii in a 
cavern in Kansas. ‘Toward evening the bats moved from the depths of 
the cavern to lateral depressions near the entrance. From these 
stations they made periodic sorties to the entrance until nightfall 
allowed their departure from the cavern. 

Many authors have noted that P. townsendii leaves its daytime 
retreat only after darkness has fallen. Dalquest (1947, p. 24) believed 
that the species had never been observed in its hunting flight. Pear- 
son, et al. (1952, p. 274) stated that it is a relatively late-flying species, 
emerging from its roosting structure after dark and not returning 
until almost dawn. 

On the other hand, Vernon Bailey (field notes) reported that in the 
Sundance region of Wyoming: 

These are the earliest bats seen flying in the evening, and they seem to come from 
various parts or openings in the canyon walls. When they first come out it is 
light enough to see their long ears as they fly about. Their flight is rather slow 
and they would be easy to shoot... . 

Bailey (1930, p. 181) again saw big-eared bats in early evening flight at 
Yellowstone National Park, and shot two (1936, p. 387) one evening at 
McKenzie Bridge, Oregon, when they circled about at the edge of the 
forest. Krutzsch and Heppenstall (1955, p. 127) shot a P. townsendii 
in Utah as the bat foraged in the early twilight. Hamilton (1943, 
p. 102) watched big-eared bats leave a West Virginia cave during the 
late dusk. The cave opening was well above the valley floor, and the 
emerging bats first flew yet higher, soaring and circling at a height of 
several hundred feet until it was too dark for successful shooting. 


Then they descended to within a few feet of the ground. 
497256—59 6 


176 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


During the hibernating season there is considerable movement of 
bats within the roost. Apparently the selection of a suitable spot for 
hibernation is a hit or miss proposition. Thus the bats repeatedly 
awaken from dormancy and move to other spots until a favorable site 
is located. Consequently, a few flying bats are likely to be found in 
the roost at any time, although the majority of individuals may be 
lethargic (p. 179). To illustrate this, Pearson, et al. (1952, p. 278) 
in midwinter marked with chalk the locations of 56 torpid bats in a 
California cave. Temperature within the cave was 34°F. Particu- 
lar effort was made to avoid disturbing the bats. The following day 
there were as before 56 torpid bats in the chamber, but at least 17 
had moved to new locations. Records indicate that males are more 
active in winter than females. 

Loca MIGRATIONS: At all seasons individuals or populations of 
P. townsendit may normally alternate between two or more roosts, or 
may irregularly seek new roosts. These facts have been emphasized 
by the banding of hibernating populations and by the observation of 
nursery colonies. Banding operations in West Virginia (504 Plecotus, 
Mohr, 1952, p. 7), Kansas (155 Plecotus, Twente, 1955a, p. 380), 
and California (1500 Plecotus, Pearson, et al., 1952, p. 274) all indicate 
movement of P. townsendii from cave to cave during even the coldest 
weather. Much of this movement undoubtedly was precipitated by 
the disturbance to the bats when they were banded, or when the band 
numbers were read. Twente (1955a, p. 380) found that when thus 
disturbed the bats invariably awakened and left the cavern, not to be 
seen there again for several days or months, if ever. The banding 
data indicated in addition that there was considerable movement 
from cave to cave by bats that had not been disturbed by banding. 
Apparently these individuals were seeking circumstances more favor- 
able for hibernation. 

Pearson, et al. (1952, p. 276) described the effect of disturbance on 
a nursery colony. One evening, after the adult bats had left the 
roost, their 75 young were banded. ‘The adults returned, picked up 
their young, and by morning all were settled in an alternate roost 1.3 
miles away. How the colony quickly agreed to move and arrived 
intact at a distant new roosting place is an intriguing mystery. 

SEASONAL MIGRATIONS: There is no evidence of long migrations by 
P. townsendu. Among the 1500 banded in California by Pearson, 
et al. (1952, p. 275) most that were recaptured were recovered at the 
original banding site or not more than 1.5 miles from it. A young 
male found dead 20 miles from his birthplace was the most distant 
recovery. ‘Twenty-four percent of 504 individuals banded in West 
Virginia were recovered in a subsequent season, all near their banding 


AMERICAN BATS—HANDLEY Lhd 


sites. For information on dates of movements to and from nursery 
and hibernating roosts, see pp. 171, 180, and 183. 

Homrine: Only Cockrum (1956a, p. 49) has reported homing experi- 
ments with P. townsendii. He secured 54 individuals from a cave in 
Arizona in May. These were released at a point 28 miles away. 
Three of the bats were recovered at the home cave two days later and 
a fourth two months later. 

Fuicut: After spending several hours swinging a butterfly net in 
a nearly futile attempt to capture P. townsendw confined in a room, 
Dalquest (1947, p. 24) concluded that this bat sees well, is a swift 
flier, an agile dodger, and possesses an excellent sense for danger. 
Hundreds, if not thousands, of sweeps netted but one bat in free 
flight. 

Not only is P. townsendii adroit at avoiding hand-operated nets, 
but some characteristic keeps it from becoming entangled in mist nets 
as well. Although these nets are very effective for capturing many 
species of bats, those that do not forage near the ground and those 
whose flight is slow, cautious, and butterfly-like are seldom taken. 
Also, perhaps those with an unusually well developed echo location 
apparatus can detect and avoid the nets more readily. Only Baker 
(1956, p. 187) has reported taking P. townsendv in a mist net. 

The flight of captive big-eared bats has been described in detail by 
Grinnell (1918, p. 344) and Dalquest (1947, p.25). In the confinement 
of a room, the flight pattern most often was a figure eight, varied with 
ovals and circles, extending from floor level to the beamed recesses 
of the ceiling. Flight character varied from hovering like a humming- 
bird to slow and butterfly-like to swift and swallow-like. Wing-beats, 
deep and smooth, often alternated with short glides. In average 
flight the head was bent ventrally so that the body formed a smooth 
curve from chin to tip of tail. The ears, pointed forward and held 
almost parallel to the horizontal plane of the body, were, surprisingly, 
scarcely noticeable. The legs were spread apart so that the inter- 
femoral membrane was expanded. I have noted that in slow flight 
the head is not so depressed and the ears are more conspicuous, as 
they are held more nearly perpendicular to the horizontal plane of the 
body. On some occasions the flight of P. townsendii may be silent, 
with no squeaking, swishing, or fluttering. At other times it is 
relatively noisy. 

When it alights on a wall or ceiling, P. townsendii swoops up from 
below, its wings close, and with great agility it flips over so that its 
feet become foremost and almost simultaneously grasp a foothold. 

Restine posture: In ordinary sleep the big-eared bat normally 
hangs free, suspended only by the feet (see Dalquest, 1947, p. 25, for 
a detailed description). Many observers have noted its peculiarity 


178 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


of frequently hanging suspended*by only one foot, and Dalquest 
(1947, p. 29) has pointed out that the truly lethargic bat clings with 
both feet and thumbs and presses its body tightly to the surface of 
its roost-site. Among big-eared bats hibernating in the gypsum cav- 
erns of Oklahoma and Kansas, Twente (1955b, p. 721) observed a 
different posture. The ears were folded back along the neck, the 
fingers were spread so that the wings effectively covered the ventral 
surface of the body, the tail and interfemoral membrane were bent 
down to cover the venter and hind parts of the wings, and the long 
body hairs stood out perpendicularly. In this position the bat was 
practically enveloped with a layer of dead air, effectively insulating it 
from the surrounding warm air of the cavern, and helping it to assume 
a body temperature similar to that of the substrate from which it 
was suspended. Grinnell and Swarth (1913, p. 380) noted a similar 
posture in lethargic big-eared bats in California. 

The normal position of the ear in the undisturbed resting bat ap- 
pears to be coiled back and down tightly against the side of the neck 
like aram’s horn. The posterior portion of the ear is wrinkled, rather 
accordion-like, and there is lateral contraction in accomplishing the 
spiral shape. The ears can be coiled independently, and sometimes 
an individual is seen with one ear erect while the other is coiled. Ap- 
parently the tragus always remains erect, regardless of the position 
of the ear. Upon superficial examination, when the ear is coiled, the 
erect tragus may be mistaken for the ear. 

Possibly P. townsendii sometimes adopts the common ear posture 
of the resting P. auritus, that is, with the ear folded back against the 
side of the neck and chest and partially concealed beneath the folded 
wing, rather than coiled against the side of the head and neck (Grin- 
nell and Swarth, 1913, p. 380; Howell, 1920, p. 174; Alcorn, 1944, p. 
309). 

When the bat is not lethargic, the ears are unfurled at the slightest 
alarm, and move with increasing vigor as the observer approaches. 
The sight of a tightly packed cluster of bats with long ears quivering, 
waving, and in constant motion is a memorable one. 

Hamilton (1943, p. 102) and Mohr (1933, p. 50) noted that big- 
eared bats lapsed into disoriented flight or fell stunned when guns 
were discharged near them in caves. These authors presumed that 
the auditory apparatus of P. townsendii is more sensitive than that 
of bats with smaller ears. 

Voice: The voice of P. townsendir is difficult to describe. A high 
pitched twittering is commonly heard both in roosts and in captivity. 
H. W. Grinnell (1918, p. 344) noted that an individual “protested in 
shrill cicada-like notes” when handled. Grinnell and Swarth (1913, 
p. 380) described the voice as a “‘grating squeak,”’ and Dalquest (1947, 


AMERICAN BATS—HANDLEY 179 


p. 29) stated that semi-lethargic individuals were able to “‘hiss’’ when 
disturbed. Mohr (1933, p. 50) reported that E. L. Poole made the 
following observations on the call of an individual less than a week old: 
“Tt has a ventriloquial hollow cuc-cuc-cuc, like a yellow-billed cuckoo; 
close at hand a high thin tse-tse-tse, like the song of a blackpoll warbler, 
which could be heard synchronized with the other sound.” 

HisernaTion: Hibernation in an animal is a state of lethargy in 
which body temperature is depressed in correspondence with a low 
environmental temperature and in which physiological activity is 
proportionally curtailed. Hibernation may be of long or short dura- 
tion. Among bats there appears to be no clear distinction between 
sleep and hibernation. When the bat rests its temperature approaches 
that of its environment. Thus, without regard for season, if the air 
temperature of its roosting place is low enough, the bat assumes a 
lethargic condition. 

However, Twente (1955b, p. 713) found that the body temperature 
of bats difficult to awaken and bats easy to awaken may be the same. 
He postulated that the degree of lethargy is dependent upon how high 
or how low the bat’s body temperature has been during the period of 
lethargy. This factor would determine the amount of stored food 
that had been catabolized during the period of inactivity. The pri- 
mary wastes of this process are carbon dioxide, nitrogenous byprod- 
ucts, and water. The observation that Myotis nearly always urinates 
when awakened suggested to Twente that a possible factor causing 
awakening is the amount of urine in the bladder and the nervous con- 
sequences of bladder distension. He proposed that it would be more 
appropriate to refer to the bat’s lethargic state in degrees of irritabil- 
ity rather than in degrees of lethargy. Thus, bats hibernating in 
places that are warm or have fluctuating low temperatures are usually 
more irritable than those that have been hibernating in places that 
have relatively constant low temperatures. Similarly, bats at the 
edges of clusters tend to be more irritable than those in the center, 
for they have been members of the cluster the shortest time and have 
higher temperatures and a higher rate of metabolism. If the place 
that the bat has selected for hibernation is unsuitable, the animal 
awakens, probably because a high metabolic rate has made it irritable, 
and moves to another place. The bat always awakens and moves 
when the temperature of its environment approaches 32° F. 

Twente (loc. cit.) found that once the awakening process is initiated, 
the bat’s body temperature rises at a constant and rapid rate (about 
1.8° F. per minute for P. townsendii) until flight becomes possible for 
the animal. Reeder and Cowles (1951, p. 395) determined the thresh- 
old of flight to be at a rectal temperature of 73.9° F. in P. townsendia. 
Twente’s specimens of this species flew at a rectal temperature of 


180 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


75.2° F. but did not fly agily until a temperature of 82° to 86° F. was 
reached. 

Pearson, et al. (1952, p. 276) found most P. townsendi lethargic at 
air temperatures below 62° F. and recorded temperature as low as 
28.5° F. near hibernating individuals. Twente (1955b, p. 715) ob- 
served an air temperature range of 37.4° to 69.8° F’. where lethargic 
big-eared bats were resting. When Dalquest (1947, p. 29) placed 
active specimens of P. townsendii in a refrigerator at a temperature of 
40° F. all became lethargic within 20 minutes. 

Pearson, et al. (1952, p. 277) observed in their studies of P. town- 
sendii in California that the bats began to arrive in their hibernating 
quarters in late October, reached peak numbers in January, and de- 
parted by April. Females arrived earlier and stayed later than males. 
Females were usually lethargic both day and night during the coldest 
periods, whereas males tended to awaken at night the year around, 
even in midwinter. During spring, summer, and fall both sexes were 
usually alert day and night and lethargic individuals were uncommon, 
especially in September when the males were coming into breeding 
condition. 

In central California, Pearson, et al. (1952, p. 280) found that some 
male P. townsendii apparently did not hibernate. Individuals came 
to night roosting places throughout the winter in outside air tempera- 
tures ranging from 37° to 57° F. Winter records of females at night 
roosts were more irregular but indicated activity on nights when the 
outside air temperature ranged between 47° and 57° F. Pearson, 
et al., did not determine what food sustained this winter activity. 
Although flying insects were probably scarce at this season, these 
authors captured adult male bats that lacked stored fat. Elsewhere, 
males that they weighed averaged 8.36 grams on 3 February and had 
dropped 0.64 grams to 7.72 grams by 2 April. This loss probably rep- 
resented the amount of stored fat required to sustain the bats for this 
period (temperatures 37° to 50° F.) without intake of additional food. 

For further data on roosts, sociability, and movement during hiber- 
nation, see pp. 168, 170, and 176. 

Repropuction: Unless otherwise noted, all of the following infor- 
mation on repreduction in P. townsendii is taken from Pearson, et al. 
(1952, pp. 281-314). 

Insemination.—The mating season extends from early October to 
late February. Copulatory activity is greatest in the winter roosts, 
but some, perhaps most, females have mated at least once before they 
arrive at the wintering places. Unmated females may be found 
throughout September, but spermatozoa are to be found in all after 
about the third week of October. Young females (as little as 3 or 4 
months old) mate as early as do adult females, but because they pro- 


AMERICAN BATS—HANDLEY 181 


duce few sperms and because their sex accessories are small, young 
males (4 to 9 months old) are almost certainly sterile in their first 
breeding season. 

At times the males, more or less active throughout the winter, ap- 
proach and probably achieve intromission with lethargic females. 
The passive state of the females and the absence of a vaginal plug 
would permit each female to be mated many times in the course of the 
winter. Since all females in the hibernating quarters contain sperma- 
tozoa by late October, subsequent copulatory activity on the part of 
the males must be directed toward females already inseminated. 

It is not known how the male circumvents the female’s interfemoral 
membrane, which during lethargy is usually curled tightly over the 
abdomen. The only copulation that Pearson, et al., observed took 
place on a cave ceiling too high to allow observation of detail. How- 
ever, the male embraced the female from behind in the usual manner 
of copulating mammals. 

Precopulatory activity was observed among captive bats. The 
female hung by her feet and was approached from the front by a male. 
Twittering, the male proceeded to embrace the female with his wings, 
then vigorously rubbed his snout, perhaps making use of his muzzle 
glands, over the foreparts and ventral surface of the female. A 
strong bat odor was noted. 

Sperm storage.—Spermatozoa of bats are viable for surprisingly long 
periods. In male P. townsendii they retain their motility for six or 
more months after they reach the epididymis. However, it is not 
known whether motile spring spermatozoa are capable of fertilizing. 
Spermatozoa remained abundant in the uteri of females kept isolated 
in cool temperatures for as long as 108 days. Probably in P. town- 
sendii, as in other vespertilionid bats, spermatozoa stored over winter 
in the female reproductive tract are capable of fertilizing in the spring. 
After ovulation, spermatozoa disappear quickly from the uterus. 

Ovulation.—In P. townsendii ovulation occurs from February to 
April. Maturation of the ovarian follicle and its subsequent ovula- 
tion may depend upon the stimulation of copulation, and this stimulus 
may be effective even though administered several months before 
ovulation. It appears that there is an autumnal and early winter 
period during which ovulation normally will not occur. Neither fe- 
males that are active in winter nor those that are lethargic in winter 
ovulate before spring, and those that are removed from hibernation 
and kept at warm temperatures do not ovulate readily. The possi- 
bilities of sperm storage in the female and insemination of unreceptive 
females while they are lethargic may have permitted the advanced 
copulatory period to evolve. 

Gestation.—The duration of gestation may depend upon the body 


182 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


temperature of the bat and upon whether or not it was lethargic during 
a portion of the period. Variation in environmental conditions in 
hibernating quarters and nursery roosts thus may have a profound 
effect on gestation. The gestation period varied from 56 to 100 days 
in the California populations of P. townsendi studied by Pearson, 
et al. 

Parturition.—The birth process was observed in a captive female. 
She hung from the wire top of her cage by widespread hind feet and 
thumbs. Her tail was curled upward so that it completed the ‘“in- 
verted umbrella” effect and made it quite unlikely that the young 
could fall. In fact, not even a drop of blood or fluid reached the cage 
floor. This position was maintained all during parturition and for 
half an hour afterward. The birth was a breech presentation. With- 
in minutes the fluids and afterbirth were consumed by the mother, and 
while she licked her membranes and ofispring, the young moved for- 
ward, secured a nipple, and attached its feet to the top of the cage. 

A single young is the rule in this species, and 49 out of 50 embryos 
were found in the right horn of the uterus. After parturition this 
horn never returns to its original small size, but remains larger than 
the left, about 1.5 to 2 mm. in diameter. This disparity in size may 
lead to presumably erroneous reports of fall pregnancies as that of 
Bailey (1936, p. 390). 

The average date of parturition of a colony varies from year to 
year, and there seems to be a considerable variation in the spread of 
birth dates in different colonies (19 to 35 days). It is probable that 
birth dates extend over a long period in a colony whose females 
roosted in diverse places with a variety of environmental temperatures 
before they congregated at the nursery colony. There is evidence 
also that young females tend to join the nursery colony later in the 
spring than do adults, and that they bear their young later in the 
season than do adults. 

Pearson, et al., estimated birth dates for several hundred western 
American P. townsendzi from specimens, field notes, and literature by 
reference to graphs relating embryo size and forearm length to age 
(figs. 20, 21). The earliest calculated birth date was 19 April in 
California, and the latest was 22 July in Arizona. Records for the 
Appalachian population indicate a similar breeding season. 

Growth of young.—Pearson, et al., described the newborn P. town- 
sendit as a grotesque creature, its large ears flopping over its unopened 
eyes and the disproportionately large size of its thumbs and hind feet 
giving it a spider-like appearance. The following chronology of 
growth was set forth: 

The average weight of newborn young was 2.4 grams, and the 
forearm length averaged 16.6 mm. (there is no significant size varia- 


AMERICAN BATS—-HANDLEY 183 


= 
= Ig 
°|. wi 
te 23 [6 
= 21 “14 
ae 12 
to me he 
= = 15 z 
a 2 er 
ww 15 ae (e! 
ost 4 
a 
u 9 
= fee 
pees 
St o 5 
x 
ES: oO 
°o 
= 3 
i 
o 
= 
Ww 
— 


AGE IN DAYS. 


Ficure 20.—Average growth-rate of embryo in Plecotus townsendit pallescens. Measure- 
ments in millimeters. (Modified from Pearson, et al., 1952, p. 302.) 


tion between sexes until late in development when smaller size of 
male becomes apparent). Within a few hours the young one is able 
to utter a characteristic sharp, metallic “chirp.”’ The dried umbilical 
cord may remain attached for a day or two. Before five days of age 
the naked pink body becomes covered with short gray hair. At 
seven days the young bat can make the ‘‘squaking” noise of the dis- 
turbed adult, and the ears assume the erect position of the adult. A 
few days later the eyes open. Up to the age of 16 days (forearm less 
than 35 mm.), while the growth curve is approximately linear, the 
forearm lengthens 1.2 mm. per day. Forearm length is a reliable 
indicator of age up to about three weeks. At four weeks the forearm 
is so close to adult length that it can no longer be relied upon. The 
young fly at about two and a half to three weeks of age, and at six 
weeks some young fly out at night as do the adults. The summer 
colonies start to break up in August when the older young are just 
over three months old. 

Lactation.—In Plecotus there is a single pair of pectoral mammae, 
lateral in position. Tiny nipples (less than 0.6 mm. long in fresh 
specimens) are characteristic of young and nonparous females, and 
once a female has borne young the nipples do not return again to the 
tiny category, although they frequently become small again. The 
nipples of the young remain tiny through the summer, fall, and 
winter of their first year, and begin to enlarge at about the time of 
implantation of the embryo. The nipples reach full size (up to about 
3 mm. in length) at the end of pregnancy or shortly thereafter. 


184 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ADULT ¢ 


42 a eeae=— os ADULT & 


| FLYING 


| ‘FLUT TERING 


leves OPEN 
| EARS UP 


t GRAY: FUZZ 


2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 
AGE IN DAYS 


Figure 21.—Average growth-rate of forearm in Plecotus townsendii pallescens. Measure- 
ments in millimeters. (Modified from Pearson, et al., 1952, p. 313.) 


The young probably nurse with their heads hanging down, and 
most nursing must occur during the day, for from the time that they 
are born or within a day or two of their birth, the young bats are left 
in the roost at night while the adults go out to feed. Banding has 
proved that each female bat unerringly selects her own young from 
amongst the many in the cluster when she returns to the nursery 
from her foraging flight. 

The young bats cling to anything that comes into their grasp, and 
it is difficult to remove them from their mother. A disturbed mother 
may sometimes be seen flying with a young-one dangling from a 
nipple, but usually the young bat clings to its mother’s fur with well- 
clawed thumbs and feet, as well as to the nipple with its mouth. 
Adults may fly with attached volant young weighing as much as six 
grams—about two-thirds of the adult’s weight. 

It is difficult to tell when lactation ceases, but it appears that 
females nurse their young for about two months. By that time the 
young are as large as their mothers and are skillful fliers. 

Mott: The young big-eared bat molts from its juvenile pelage to 
adult pelage at about the time that it reaches adult body size, be- 
tween the ages of one and two months. One collected on 7 August 
at Burkes Garden, Va. (USNM 157075), was beginning the molt on 
chest and belly. Three collected on Sand Creek, Wyo. (USNM), on 
24 August, had barely begun to molt. 

Adults have a single complete molt annually, most commonly in 
August. Possibly the molt normally occurs earlier in males than in 
females. New hairs usually appear at almost the same time on all 


AMERICAN BATS—HANDLEY 185 


parts of the body. Short, prostrate, new hairs are found on some 
individuals as early as June (MVZ 103184, o&, San Luis Obispo 
County, Calif., 24 June) or July (USNM 250084, o, Fallon, Nev., 
7 July). Specimens from Evergreen, Calif. (MVZ 71619, o, 2 Au- 
gust), Huachuca Mountains, Ariz. (UI 4674, o', 7 August, and UI 
1425, 9, 15 August), and Vidal, Calif. (LSU 1121, 9, 29 August) 
were molting when captured. Six of seven females (USNM) taken 
on 24 August along Sand Creek, Wyo., were molting. Of five speci- 
mens (USNM) collected on 24 August at Las Cruces, N. Mex., one 
male had completed the molt; one female had completed the molt on 
the dorsum, but was still molting on the belly; two females were 
molting on all parts of the body; and one male apparently had not 
begun to molt. 

Seme of the summer specimens from New Mexico and southern 
California have a peculiar, short, woolly, relatively sparse pelage, 
possibly due to early shedding of guard hairs. The same condition 
has been observed in some specimens of P. mexicanus. 


Plecotus townse ndii australis Handley 


Plecotus (Corinorhinus) townsendi J. A. Allen, 1890, p. 176. 
Corynorhinus macrotis pallescens Miller, 1897, p. 52. (Part.) 
Corynorhinus megalotis mexicanus G. M. Allen, 1916, p. 347. (Part.) 
Corynorhinus rafinesquit mexicanus Miller, 1924, p. 83. (Part.) 
Plecotus rafinesquizt mexicanus Dalquest, 1953, p. 64. (Part.) 
Corynorhinus townsendii australis Handley, 1955c, p. 147. 

Hoiotypge: USNM 297265; adult female in alcohol, skull removed; 
collected 20 December 1952 by Aurelio Maélaga Alba; 2 miles west of 
Jacala, 5,500 feet, Hidalgo, Mexico; collector’s No. 1053. 

Distrispution: Arid interior mountain ranges of central and north- 
ern Mexico (fig. 24). North to San Juan, Durango, and near Hacienda 
La Mariposa, Coahuila. East to near Hacienda La Mariposa and 
near Bella Unién, Coahuila; Presa de Guadalupe, San Luis Potosi; near 
Jacala and near Yoltepec, Hidalgo; Lago Texcoco, México; and 
Oaxaca, Oaxaca. South to Oaxaca, Oaxaca; Cuernavaca, Morelos; 
and San Andrés, Jalisco. West to San Andrés, Jalisco; Sierra de Val- 
paraiso, Zacatecas; and San Juan, Durango. Zonal distribution: 
Upper Sonoran and upper portion of Lower Sonoran. Altitudinal 
distribution: 1,800 to 9,500 feet. Most collecting stations are between 
4,000 and 7,000 feet. 

Description: Adult coloration: Hair of upperparts between Benzo 
Brown and Fuscous at base; tips brighter brown, burnished with dark 
brown; mass effect between Russet and Cinnamon-Brown; hair bases 
sharply differentiated from tips. Hair of underparts Natal Brown at 
base; tips about Light Pinkish Cinnamon on belly, somewhat darker 
on throat. 


186 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


V 
Ks 
© 


aR 


NTT 
q : 
Hin 
,  Seeenas) 


Ficure 22.—Distribution of Plecotus townsendii Cooper (western section): 1, P. t. pal- 
lescens; 2, P. t. townsendii. Solid symbols, specimens examined; open symbols, other 
records; encircled symbols, type localities. Cross-hatching indicates known areas of 
intergradation between subspecies (between pallescens and townsendit in California, 
Nevada, Idaho, Oregon, Washington, and British Columbia; between pallescens and 
ingens in northern Texas, Oklahoma, and Kansas; and between pallescens and australis 
in western Texas, Chihuahua, and Coahuila). 


Size averages medium for the subgenus; forearm averages relatively 
long. Rostrum averages relatively long, dorsolaterally inflated, and 
usually not particularly depressed; anterior nares relatively large and 
usually rounded posteriorly (dorsal view). First upper incisor nor- 
mally without secondary cusp; upper canine averages less robust than 
in northern populations; anterointernal cingular cusp of P* frequently 
present. : 

MEASUREMENTs: See tables 10, 16. 


AMERICAN BATS—-HANDLEY 187 


Wn a Se ee: 


X 


oe ne 


-—— 


Figure 23.—Distribution of Plecotus townsend1i Cooper (eastern section): 1, P. t. ingens; 
2, P. t. virginianus. Solid symbols, specimens examined; open symbols, other records; 
encircled symbols, type localities. Cross-hatching indicates known areas of intergrada- 
tion between subspecies ingens and pallescens. 


Comparisons: P. t. australis is most similar to P. t. pallescens, but 
averages darker, browner, and less cinnamon dorsally. P. t. australis 
averages larger in most cranial dimensions than Arizonan P. t. pal- 
lescens, but can be scarcely distinguished cranially from other popula- 
tions of P. t. pallescens. 

For comparison with P. mexicanus, see account of that species. 

Remarks: Mexican populations of P. townsendii are relatively uni- 
form in coloration and cranial characters despite the wide range of 
latitude and elevation inhabited. The greatest divergence noted is in 
specimens from San Andrés, Jalisco, in which the rostrum is unusually 


188 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


i 


SHTsiaaiy 


Ficure 24.—Distribution of Plecotus townsendii Cooper (southern section): 1, P. t. australis; 
2, P. t. ingens; 3, P. t. pallescens. Solid symbols, specimens examined; open symbols, 
other records; encircled symbols, type localities. Cross-hatching indicates known areas 
of intergradation between subspecies (between pallescens and ingens in northern Texas, 
Oklahoma, and Kansas; between pallescens and australis in western Texas, Chihuahua, 


and Coahuila). 


broad and inflated. Indeed, the whole skull is broad (reflected in 
zygomatic breadth, breadth of brain case, etc.) in those specimens. 
P. t. australis intergrades with P. t. pallescens. Samples from south- 
ern Coahuila have the coloration of P. ¢. australis. 'Two specimens 
from near San Buenaventura in central Coahuila resemble P. ¢. pal- 
lescens, whereas one from the Chisos Mountains, Texas, is very similar 
to P. t. australis. Other specimens from the Big Bend region of Texas 
and Chihuahua are, on the average, sufficiently similar to P. t. palles- 
cens to be identified as that subspecies, and central and southeastern 
New Mexican specimens are essentially identical with samples of 
typical P. t. pallescens (northern and central Arizona). Apparently 


AMERICAN BATS—-HANDLEY 189 


intergradation between these two races of P. townsendw occurs in 
northern Coahuila and western Texas. 

SPECIMENS EXAMINED.—A total of 58 from the following locali- 
ties: 

MEXICO: Coaunuita: 1 mile south and 4 miles west of Bella Unién, 7,000 feet, 
3s (KU); 4 miles west of Hacienda La Mariposa, 2,300 feet, 1s (KU); one-half 
mile north of Muralla, 4,500 feet, 19bs (IU); 9 miles west and 4 miles south of 
San Buenaventura, 2,000 feet, 8s (KU); Sierra Guadalupe, 7,800 feet, 10 miles 
south and 5 miles west of General Cepeda, Is (KU). Duranao: San Juan, 3,800 
feet, 12 miles west of Lerdo, 2s (UMMZ). Guanasvato: Santa Rosa [9,500 feet], 
7a (USNM). Guanasuato?: Charcas, 5a (USNM). Hrpauco: Grutas Xoxafi, 
6.6 miles southeast of Yoltepec [= Yolotepec?, 6,600 feet], 1s (KU); 2 miles west 
of Jacala, 5,500 feet, 3a (USNM); Rfo Tasquillo (Rio de Tula), 5,200 feet, 16 
miles east [=12 miles south-southeast?] of Zimapdn, ls (TCWC). Jauisco: San 
Andrés [4,900 feet], 10 miles west of Magdalena, 3s (UMMZ); San Pedro, Guadala- 
jara, ls (AMNH). Mégxico: Convento de Acolman, 9 miles north of México, 
Distrito Federal, 1s (IB); Lago Texcoco, 7,500 feet, la (USNM). Moretos: 
Cuernavaca [4,900 feet], 1a (USNM). Oaxaca: Oaxaca [Motne Albdn, 6,500 feet, 
3 miles southwest of Oaxaca], la (USNM). San Luts Porosf: Bledos [6,200 feet], 
Is (LSU)1; Presa de Guadalupe [4,000 feet] Is (LSU). Zacarncas: Sierra de Val- 
paraiso [13 miles west of Valparaiso, 8,200 feet], Is (USNM). ‘‘Merxico’’: No 
exact locality, 1b (USNM). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: ‘ 
MEXICO: Distrito FEDERAL: Desierto de los Leones [measurements taken 
by Villa in 1958, according to my instructions, agree fairly well with those of 


P. t. australis] (Villa R., 1953, p. 340). San Luis Potosf; Hacienda La Parada 
[6,000 feet] (Miller, 1897, p. 53). 


Plecotus townsendii ingens Handley 


Corynorhinus rafinesquet rafinesquet Sealander, 1951, p. 465. 
Corynorhinus townsendit ingens Handley, 1955c, p. 148. 

Houotryrpe: USNM 296767; adult male, skin and skull; collected 
4 December 1950 by John A. Sealander; Hewlitt Cave, 12 miles west 
of Fayetteville, Washington County, Ark; collector’s No. 50-14. 

DistripuTion: Ozark Highlands in northwestern Arkansas and 
southwestern Missouri (fig. 23); possibly also extreme eastern Okla- 
homa. North to Stone County, Missouri. South to Washington 
County, Ark. Zonal distribution, Upper Austral. Altitudinal dis- 
tribution, approximately 1,000 to 1,500 feet. ‘ 

Description: Adult coloration: Mass effect of upperparts be- 
tween Hazel and Mars Brown; hair bases Fuscous. Hair of under- 
parts between Light Vinaceous-Cinnamon and Light Pinkish Cinna- 
mon distally, the bases, except those bordering membranes, Fuscous; 
bases of border hairs same as tips. Distinction between bases and 
tips of hairs fairly sharp, on both dorsum and underparts. 


190 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Size averages large; forearm averages relatively long; skull of 
heavy construction; rostrum relatively long, inflated, and not de- 
pressed; anterior nares average relatively large and rounded in 
posterior outline (dorsal view). First upper incisor usually with at 
least a trace of a secondary cusp; anterointernal cingular cusp of 
P* absent; molariform teeth robust. 

MEASUREMENTS: See tables 10, 16. 

Comparisons: Bats of the race P. t. ingens are the most reddish 
and the largest of P. townsendii. From P. t. pallescens, the only 
geographically adjacent race of this species, ingens is distinguished 
by darker, more orange or reddish coloration, average larger size, 
relatively larger auditory bullae, more inflated rostrum, relatively 
more robust molariform teeth, and more frequent development of a 
secondary cusp on the first upper incisor. 

For comparison with P. ¢. virginianus see account of that sub- 
species (p. 202). 

Remarks: The Ozark form is similar to both P. t. virginianus to 
the eastward and P. t. pallescens to the westward. Specimens of 
P. t. pallescens from western Oklahoma and south-central Kansas 
approach P. ¢. ingens in size and coloration, although they are more 
like typical P. t. pallescens. 

SPECIMENS EXAMINED: A total of 18 from the following U.S. 
localities: 

ArKANsas: Washington County: Basset Cave, near Hicks, ls (UAZ); Devil’s 
Icebox, Devil’s Den State Park, 25 miles southwest of Fayetteville, 2s (CNHM), 
9s (UAZ), Is (USNM), 1s (GF); Hewlitt Cave, 12 miles west of Fayetteville, 


2s (UAZ), 1s (USNM). Missouri: Stone County: No exact locality, la 
(AMNH). 


Plecotus townsendii pallescens Miller 


Synotus macrotis H. Allen, 1864, p. 68. (Part.) 

Synotus townsendi H. Allen, 1864, p. 65. 

Clorynorhinus] macrotis H. Allen, 1865, p. 174. (Part.) 
Clorynorhinus] townsendi H. Allen, 1865, p. 175. 

Corynorhinus macrotis pallescens Miller, 1897, p. 52. 

Corynorhinus macrotis intermedius H. W. Grinnell, 1914, p. 320. (Part.) 
Corynorhinus megalotis megalotis G. M. Allen, 1916, p. 338. (Part.) 
Corynorhinus megalotis pallescens G. M. Allen, 1916, p. 341. 
Corynorhinus rafinesquii pallescens H. W. Grinnell, 1918, p. 340. 
Plecotus rafinesquii pallescens Blair, 1952, p. 96. 

Corynorhinus tlownsendii] pallescens Handley, 1955e, p. 147. 


Hototrypr: USNM (Biol. Surv. Coll.) 66534; adult female, skin 
and skull (crushed); collected 3 August 1894 by A. K. Fisher; Keam 
Canyon, Navajo County, Ariz.; collector’s No. 1715. 

DistripuTIon: Western United States, northwestern Mexico, 
and southwestern Canada, including parts of the Great Plains, Rocky 


AMERICAN BATS—HANDLEY 191 


Mountains, Great Basin, Mohave and Colorado Deserts, ete., but 
excluding the northwestern coast region (fig. 22). North to Stanley 
and Harding Counties, S. Dak.; Powder River and Yellowstone 
Counties, Mont.; Yellowstone National Park, Wyo.; Riske Creek 
and Williams River, British Columbia. West to Riske Creek and 
Williams River, British Columbia; Okanogan and Yakima Counties, 
Wash.; Harney County, Oreg.; Siskiyou, Tehama, Placer, Tulare, 
and Kern Counties, Calif., and the coast and Channel Islands of 
California south of Point Conception. South to San Diego and 
Imperial Counties, Calif.; northern Baja California, northern Sonora, 
and northern Chihuahua, Mexico; Dona Ana County, N. Mex.; 
Culberson, Presidio, Brewster, Val Verde, and Edwards Counties, 
Tex. East to Val Verde and Edwards Counties, Tex.; Eddy County, 
N. Mex.; Cimarron County, Okla.; El Paso, Boulder, and Larimer 
Counties, Colo.; Custer and Stanley Counties, S. Dak. An appar- 
ently isolated population inhabits the gypsum cave region in south- 
central Kansas, central Oklahoma, and north-central Texas (speci- 
mens from Barber and Comanche Counties, Kans., south to Garza 
County, Tex. Zonal distribution: Most numerous in Upper Sonoran 
and Transition zones, but also occurs at the upper edge of the Lower 
Sonoran and the lower edge of the Canadian. Altitudinal distribu- 
tion: Sea level (possibly below sea level in southern California) to 
9,600 feet. 

Description: Adult coloration: Mass effect of upperparts averages 
between Sayal Brown and Cinnamon; hair bases Light Cinnamon- 
Drab to Benzo Brown. Hair bases of underparts average between 
Fawn Color and Benzo Brown; tips between Light Pinkish Cinnamon 
and Pinkish Buff. Distinction between bases and tips of hairs usually 
everywhere slight. Immature coloration: Mass effect of upperparts 
between Drab and Cinnamon-Drab; hair bases slightly grayer. Hair 
tips of underparts “dirty” white; bases Benzo Brown to Light Mouse 
Gray. 

Size averages medium for the subgenus; forearm averages relatively 
long; skull relatively light in construction; rostrum averages relatively 
long, not particularly depressed, and not inflated; anterior nares 
average relatively small and angular in posterior outline (dorsal view). 
Canine relatively strong; secondary cusp on I! usually absent; 
anterointernal cingular cusp of P* frequent. 

MEASUREMENTS: See tables 11, 17. 

Comparisons: In coloration of fur, membranes, and ears P. t. 
pallescens is the palest and most yellowish known population of the 
species. 

For comparison with related subspecies see accounts of P. ft. 
australis (p. 187), P. ¢. ingens (p. 190), and P. t. townsendii (p. 198). 

497256—59 7 


192 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Remarks: There are four centers of saturate coloration in P. 
townsendii: (1) the central Appalachians; (2) the Ozark Plateau; 
(3) the Pacific Northwest; and (4) to a lesser degree, north-central 
Mexico. Between the three latter centers is a pale-colored population 
(P. t. pallescens) which exhibits maximum pallor and yellowness in the 
area extending from southeastern California and central Arizona 
northeastward to Wyoming, southeastern Montana and southwestern 
South Dakota. The extreme in pallor is found in the northeastern 
and southwestern parts of this belt. ‘The very palest specimens that 
I have seen come from southeastern California. Southward, eastward, 
and westward from the pallid belt there is a gradual and progressive 
darkening of pelage toward the saturate centers (fig. 22). 

Populations inhabiting the northern flanks of the Sierra Madre 
Occidental in northern Sonora and adjacent parts of Arizona are 
somewhat darker than populations of central and northern Arizona, 
but are much paler than P. mexicanus which is at the northwestern 
limit of its range in the same region. New Mexican specimens are 
but slightly darker and less yellow than the typical coloration of 
pallescens. Specimens from Oklahoma and Kansas are clearly 
intermediate between pallescens and the Ozark form, there being 
considerable variation in color. The series as a whole, however, is 
nearer pallescens. Most Texas (Big Bend) specimens, while not 
appreciably darker, are much less yellowish than typical pallescens. 
One specimen is darker and resembles more southerly populations 
called P. t. australis. 

Proceeding westward from the pallid belt, increasing darkness of 
coloration is encountered as the range of the dark-colored north- 
western coast population of P. townsendii is approached. The zone of 
intergradation between pale-colored and dark-colored populations is 
broad and includes the coast region of northern and central California; 
much of the remainder of California east of the Coast Range, but 
excepting the southern deserts; northwestern Nevada; Idaho; eastern 
Oregon; eastern Washington; and south-central British Columbia. 

Within this zone there is considerable individual variation in 
dorsal coloration, shades of yellowish brown and grayish brown being 
most common, but reddish brown individuals also occurring. Tone is 
very variable throughout the zone, the range in some samples almost 
bridging the gap between the pale and dark extremes of the north- 
western coast and the pallid belt. 

Various authors (among them, H. Grinnell, 1914, p. 320; Whitlow 
and Hall, 1933, p. 245; Hall, 1946, p. 160; and Dalquest, 1947, p. 20) 
have regarded the populations of this region as being subspecifically 
distinct from townsendiz and pallescens. The appropriate name, 
“intermedius” H. W. Grinnell, is available for this population. 


AMERICAN BATS—HANDLEY 193 


Dalquest (1947, p. 20) suggested that this intergrading population 
probably developed intermediate characters in response to the nature 
of its environment—intermediate between the humid, forested coast 
and the arid, treeless deserts, and further postulated that genetic 
intergradation was not necessarily implied. If this were not an 
intergrading population I would expect to find constant adaptive 
characters developed in some portion of its geographic range. How- 
ever, in examining specimens from all portions of the range assigned to 
‘“Ontermedius,” I have failed to find such a condition. 

A series of 25 specimens from Mission San Jose, Alameda County, 
Calif., when arranged in order from pale to dark, quite effectively 
bridges the color gap between pallescens and townsendii; the average 
is nearest townsendiz. Among 14 specimens from Tulare County, 
Calif., are three as pale as pallescens, two as dark as townsendii (al- 
though more reddish), and nine that are intermediate; the average is 
nearest pallescens. In a series of 16 specimens from Pyramid Lake, 
Nev., the palest individuals are indistinguishable from typical palles- 
cens, while the darkest, without reservation, could be called townsendii; 
the average is nearest pallescens. Six of seven from Malheur County, 
Oreg., are indistinguishable from typical pallescens, while one is inter- 
mediate but nearer pallescens. In a series of three specimens from 
Boulder Cave, Kittitas County, Wash., one is as dark as typical 
townsendti, while the others are somewhat paler, but nevertheless 
nearer townsendit. Whitlow and Hall (1933, p. 246) regarded a large 
series from the Pocatello Region of Idaho as intermediate. 

Similarly, there is intergradation between pale-colored and dark- 
colored populations in cranial characteristics. This is best seen in the 
relative stoutness of the rostrum (normally stout in dark-colored 
northwestern coast populations, normally less stout in pale-colored 
interior populations). Within the zone of intergradation, stoutness 
of the rostrum can not be correlated with coloration: Some pale- 
colored individuals have the rostrum very stout; some dark-colored 
individuals have the rostrum rather frail. 

From the specimens and information available to me, it does not 
appear that there is a population with constant morphological char- 
acters that could justifiably bear the name ‘‘intermedius.”” The fact 
that intergradation between P. t. townsendit and P. t. pallescens occurs 
over an extensive geographic area does not make the application of 
the name “intermedius” to the intermediate population more valid. 
The ideal in mammalian systematics is to avoid application of special 
names to geographically and genetically intermediate populations. 
The use of the name ‘‘intermedius”’ is clearly in violation of this prin- 
ciple, and the name is best relegated to synonymy. 


194 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


SPECIMENS EXAMINED: A total of 313, from the following localities: 


MEXICO: Cuinvuaunua: Tinaja de Ponce, 2,600 feet, Sierra de Ponce, 12 miles 
southwest of Santa Helena, 16as (USNM). Sonora: Santa Maria Mine, El 
Tigre Mts., 3a (UMMZ); Pilares, la (UMMZ). 

UNITED STATES: Catirornia: Imperial County: Potholes, la (USNM); 6 
miles north of Potholes, la (MCZ). Inyo County: Lone Pine, Alabama Hills, 
4as (MCZ); Owen’s Lake, la (USNM); Owen’s Valley, 1a (USNM). Riverside 
County: 2 miles west of Red Mountain Lookout, 2s (STPM), 2s (USNM); River- 
side Mts., 2s (LSU). San Bernardino County: ‘‘Manvel” or “Marvel,” la 
(USNM); Eagle Pass, 12 miles southwest of Needles, 1s (LSU); Oro Grande, 2s 
(USNM); Vidal (Alice Mine), 4a (USNM); 6 miles south of Vidal, 1s (LSU). 
San Diego County: Dulzura, la (USNM); El Monte Oaks Park, la (CNHM); 
Pine Valley, la (USNM); Vallecito Stage Station, Is (CNHM). Shasta County: 
Fort Creek, la (USNM). Siskiyou County: Lava Cave, 5 miles southwest of 
Tule Lake, ls (USNM). Tulare County: 4 miles southeast of Porterville, 550 
feet, 10s (AMNH); 10 miles east of Porterville, ls (AMNH); 4 miles east-south- 
east of White River, 1,700 feet, 38s (AMNH). Orecon: Malheur County: 15 
miles south-southwest of Carol, 17as (USNM); Malheur Cave, 20 miles east of 
Malheur Lake, 1s (USNM). Wasuineton: Okanogan County: Okanogan High- 
lands, 15 miles east of Tonasket, ls (MCZ). Ipano: Bannock County: Portneuf 
River, 3 miles northwest of Inkom, lbs (USNM); Chinks Peak, 1.5 miles southeast 
of Pocatello, lbs (USNM). Butte County: Craters of the Moon National Monu- 
ment, Is (USNM). Nevapa: Churchill County: Fallon, is (USNM). Mineral 
County: 1 mile northwest of Rawhide, 1s (UI). Washoe County: Sutcliffe, 3,700 
feet, ls (AMNH); The Needles (The Pinnacles), 3,900 feet, north end of Pyramid 
Lake, 35as (USNM). Utanw: Cache County: Logan Canyon, 15 miles above 
Logan, la (MCZ), 6a (USNM). No eract locality: la (USNM). Arizona: 
Cochise County: Chiricahua Mts., la (USNM); Fort Huachuca, 3a (USNM); 8 
miles west of Fort Huachuca, 2s (UI); 11 miles south of Fort Huachuca, Is (UI); 
Ash Canyon, 19 miles south of Fort Huachuca, 1s (UI); Carr Canyon, 7,200 feet, 
Huachuca Mts., 4a (MCZ), 3s (UI); Miller Canyon, Huachuca Mts., 3a (MCZ); 
Montezuma Canyon, 5,300 feet, Huachuca Mts., 4a (MCZ); Ramsey Canyon, 
Huachuca Mts., 2a (MCZ); Huachuca Mts., ls (MCZ). Graham County: Ash 
Creek, 4,200 feet, Graham Mts. (Pinalino Mts.), ls (USNM). Maricopa County: 
Roosevelt [Bumblebee Creek, below Roosevelt Dam], 2,300 feet, 1s (USNM); 
Christmas Mine, 3,000 feet, 5 miles southwest of Roosevelt, ls (USNM). Mohave 
County: Oatman, 6as (MCZ); Rampart Cave, 2 miles southeast of Pierce’s Ferry, 
Hoover Dam National Recreational Area, ls (USNM). Navajo County: Keam 
Canyon, ls (USNM). Pima County: 2 miles northeast of Arivaca, 4,000 feet, 
1ls (AMNH); Las Guijas Mts., north slope, 7 miles north-northwest of Arivaca, 
5s (AMNH); Las Guijas, about 10 miles north-northwest of Arivaca, la (USNM); 
Santa Catalina Mts., 3a (AMNH); Colossal Cave, 28 miles southeast of Tucson, 
2s (USNM). Pinal County: No exact locality, ls (AMNH). Santa Cruz County: 
Oro Blanco Mts., 4,600 feet, 12a (MCZ); Patagonia Mts., 3a (MCZ); McCleary’s 
Ranch, Santa Rita Mts., ls (USNM). Yavapai County: Camp Verde, 3,200 feet, 
Is (USNM), Is (AMNH); Montezuma Well, 3,500 feet, 6as (USNM); Prescott, 
1s (AMNH); Turkey Creek [east slope of Bradshaw Mts., 7 miles northeast of 
Crown King], 3,400 feet, Is (USNM). New Mexico: Dona Ana County: San 
Andreas Mts., 5,600 feet, 35 miles northeast of Las Cruces, 5s (USNM). Eddy 
County: Carlsbad, 1s (USNM); McKitterick Cave, 20 miles west of Carlsbad, 
1s (USNM); Carlsbad Caverns, 4s (CC). Socorro County: Socorro, Is (USNM). 
Texas: Brewster County: West slope of Emory Peak, 7,100 feet, Chisos Mts., 4s 


AMERICAN BATS—HANDLEY 195 


(UMMZ); Kibbe Spring, 5,700 feet, Chisos Mts., 1s (CNHM); Pine Canyon, 
4,700 feet, Chisos Mts., 1s (UMMZ); Johnson Ranch, la (USNM); Viviani 
Mines, Mariscal Mts., Is (TSDH). Garza County: Post, 1s (TT). Val Verde 
County: East Painted Cave, northwest of Del Rio, la (USNM); Langtry, 1a 
(USNM). Oxtanoma: Comanche County: Wichita Mountains National Wildlife 
Refuge, 3s (USNM). Woods County: Alva, 1a (USNM); no exact locality, 2b 
(MCZ). Woodward County: Alabaster Cave, 2s (CNHM); 6 miles southwest of 
Freedom, la (USNM). Kansas: Barber County: Sun City, 2s (USNM). Wro- 
MING: Big Horn County: 25 miles northeast of Greybull, 3a (AMNH). Crook 
County: Sand Creek, 3,750 feet, Black Hills, 10s (USNM). Yellowstone National 
Park: Devil’s Kitchen, Mammoth Hot Springs, 6as (USNM). Montana: Powder 
River County: Moorhead, 24s (USNM). Yellowstone County: 5 miles west of 
Billings, la (CNHM). Soutra Daxora: Custer County: Hell Canyon, 13 miles 
west of Custer, 7s (USN M); 18 miles southwest of Custer, 1s (USNM). Penning- 
ton County: Mouth of Spring Creek, Cheyenne River, 3s (AMNH). County un- 
certain: Crowfeet [=Crow Peak, Lawrence County?], Black Hills, 7a (AMNH). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: 


MEXICO: Basa Canirornia: No exact locality (Pearson, et al., 1952, p. 273). 
CuIHUAHUA: Casas Grandes (Mdlaga, 1953, in litt.). Sonora: Sdric (Burt, 
1938, p. 26). 

UNITED STATES: Caurrornia: El Dorado County: No exact locality (Taylor, 
1919, p. 86). Imperial County: Palo Verde (H. W. Grinnell, 1918, p. 348). Inyo 
County: Mountain Springs Canyon, 5,500 feet, Argus Mts. (Dalquest, 1947, p. 19); 
4.5 miles southwest of Bishop, 5,200 feet, and 4 miles south-southeast of Bishop, 
4,700 feet (Dalquest, 1947, p. 19). Kern County: Old Fort Tejon, near Lebec 
(Dalquest and Ramage, 1946, p. 62); Red Rock Canyon (Dalquest, 1947, p. 19); 
Weldon (Dalquest, 1947, p. 20). Lassen County: Near Wendel (Dalquest, 1947, 
p. 20). Los Angeles County: Johnson’s Harbor, Santa Catalina Island (H. W. 
Grinnell, 1914, p. 320). Placer County: Auburn, 1,300 feet (H. W. Grinnell, 1914, 
p. 320); Pioneer Cave (H. W. Grinnell, 1914, p. 320). Riverside County: Hemet 
Peak, 6,500 feet, San Jacinto Mts., 2 miles northeast of Kenworthy (Grinnell and 
Swarth, 1913, p. 379); near Whitewater (H. W. Grinnell, 1918, p. 343). San 
Bernardino County: 5 miles southwest of Ivanpah, 4,500 feet (Dalquest, 1947, p. 
19); Horse Spring, 2,700 feet, Kingston Range (Dalquest, 1947, p. 19); Mitchell’s 
(Johnson, et al., 1948, p. 343). San Diego County: One-half mile west of Banner 
(Frum, 1954, in litt.); Excondido (Dalquest, 1947, p. 19); Julian (H. W. Grinnell, 
1918, p. 343); Oak Grove (Frum, 1954, in litt.); San Diego (Stephens, 1906, p. 
265); Suncrest (Mitchell, 1956, p. 444); Vallecito (H. W. Grinnell, 1918, p. 348); 
1.5 miles northwest of Vallecito (Dalquest, 1947, p. 19). Santa Barbara County: 
Santa Cruz Island (Pearson, 1951, p. 366). Shasta County: Old Station, 4,300 
feet (Pearson, et al., 1952, p. 277). Siskiyou County: Lava Beds National Monu- 
ment, 4,800 feet (Pearson, et al., 1952, p. 278). Tehama County: Mouth of Battle 
Creek, near Bloody Island, 350 feet (Dalquest, 1947, p. 20). Ormcon: Harney 
County: Roaring Springs Cave, 80 miles south of Burns (Cressman and Reed, 1938, 
p. 248). Umatilla County: 5 miles east of Milton (Booth, 1945, p. 26). Wallowa 
County: Grande Ronde Canyon, [12 miles northeast of] Paradise (Bailey, 1936, 
p. 388). Wasurneton: Spokane County: 10 miles north of Spokane (Svihla, 1938, 
p.18). Yakima County: Selah (Dalquest, 1948, p. 164). IpaHo: Bannock County: 
3-4 miles east of Pocatello, 5,000 feet (Whitlow and Hall, 1933, p. 243); Schutt’s 
Mine, 6,300 feet, 9 miles east of Pocatello (Whitlow and Hall, 1933, p. 243); 1 
mile east of Portneuf, 4,500 feet (Whitlow and Hall, 1933, p. 248). Bingham 


196 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


County: Blackfoot, 4,500 feet (Whitlow and Hall, 1938, p. 246). Bonner County: 
Deer Island, Lake Pend Oreille, and near Lakeview (Jones, 1948, p. 416). 
Nevapa: Churchill County: 9 miles north of Eastgate (Alcorn, 1944, p. 309); 10 
miles southeast of Fallon (Hall, 1946, p. 159); 10 miles south and 5 miles west of 
Fallon, 4,300 feet (Hall, 1946, p. 162); 4.5 miles east of Stillwater, 4,000 feet (Hall, 
1946, p. 160). Clark County: 7 miles east of Las Vegas (Burt, 1934, p. 397). 
Esmeralda County: Chiatovich Ranch, 4,850-4,900 feet (Hall, 1946, p. 162); 1 
mile southeast of Dyer Ranch, 5,300 feet (Hall, 1946, p. 160). Lander County: 
one-quarter mile west-northwest of Austin (Alcorn, 1944, p. 309). Lyon County: 
2 miles southwest of Pine Grove, 7,250 feet (Hall, 1946, p. 160); near Ramsey 
(Alcorn, 1944, p. 309); Muriel Bee Mine, 5,800 feet, 4 miles west of Wichman 
(Hall, 1946, p. 159); 9 miles east and 2 miles north of Yerington, 4,700 feet (Hall, 
1946, p. 159); Bluestone Mine, 5,200 feet, 3.5 miles southwest of Yerington (Hall, 
1946, p. 159). Mineral County: Fletcher, 6,100 feet (Frum, 1954, in litt.) ; Stan- 
moore Mine, about 7,000 feet, Lapon Canyon, Mount Grant (Hall, 1946, p. 159). 
Nye County: Amargosa River, 3,400-3,500 feet, 3.5 miles northeast of Beatty 
(Hall, 1946, p. 162); Hot Creek Range, 1 mile west of Hot Creek (Hall, 1946, p. 
162); Springdale, north end of Oasis Valley (Davis, 1939, p. 124). Washoe 
County: 10 miles north-northwest of Deep Hole, 5,150 feet (Hall, 1946, p. 161); 
near Olinghouse (Alcorn, 1944, p. 309); Horse Canyon, 5,800 feet, Pahrum Peak 
(Hall, 1946, p. 161); 1 mile southwest of Pyramid Lake (Hall, 1946, p. 162); 
Smoke Creek, 9 miles east of California boundary, 3,900 feet (Hall, 1946, p. 162); 
2 miles north of Sparks (Alcorn, 1944, p. 309). White Pine County: Lehman Cave, 
7,200 feet (Hall, 1946, p. 162). Utan: Beaver County: Fort Beaver, 5,900 feet 
(Long, 1940, p. 172). Carbon County: Range Creek (Hardy, 1941, p. 293); Sun- 
nyside (Hardy, 1941, p. 293). Emery County: Horse Creek Canyon (Hardy, 1941, 
p. 293). Grand County: 4 miles north of Thompsons (Krutzsch and Heppenstall, 
1955, p. 127); Post Canyon, Book Cliffs, 8,200 feet, 75 miles south of Ouray 
(Krutzsch and Heppenstall, 1955, p. 127). Kane County: 4 miles north of Kanab 
(Hardy, 1941, p. 293). San Juan County: Bluff, 3,300 feet (Hardy, 1941, p. 293). 
Sanpete County: Pete Canyon, 5,800 feet, west of Wales (Hardy, 1941, p. 293). 
Tooele County: Goldhill (Durrant, et al., 1955, p. 72). Uintah County: 1 mile 
southeast of Ouray, 4,700 feet (Durrant, 1952, p. 59). Utah County: Rock Can- 
yon, near Provo (Biol. Surv. files); east of Springville (Hardy, 1941, p. 293). 
Washington County: Apex Mine (Hardy, 1941, p. 293); Bloomington Cavern 
(Hardy, 1941, p. 293); Mineral Mountain (Hardy, 1941, p. 293); Zion National 
Park (Hardy, 1941, p. 293). Weber County: South fork of Ogden River (Durrant, 
et al., 1955, p. 72). Anizona: Cochise County: Portal (Cahalane, 1939, p. 422). 
Coconino County: Grand Canyon National Park (Hearst Ranch, Grandview, and 
Pasture Wash Ranger Station) (Hoffmeister, 1955, p. 5). Mohave County: Virgin 
Narrows, northeast of Littlefield (Hardy, 1949, p. 434). Santa Cruz County: Peiia 
Blanca Spring (Campbell, 1934, p. 242). Yavapai County: Martinez Creek, 7 
miles southwest of Yarnell (Hatfield, 1942, p. 148). New Mexico: Grant County: 
Mule Creek (Bailey, 1931, p. 381). Santa Fe County: Santa Fe (H. Allen, 1864, 
p. 65). Texas: Brewster County: Maravillas Canyon, east of Black Gap (Tamsitt, 
1954, p. 46) ; southeast slope of Mariscal Mountain, 2,800 feet (Borell and Bryant, 
1942, p. 10). Culberson County: Bell Creek, 5,300 feet, 7 miles north of Pine 
Springs (Davis and Robertson, 1944, p. 262); Upper Sloth Cave, west slope of 
Guadalupe Mts., 7,000 feet (Davis, 1940, p. 74). Hdwards County: Devil’s Sink- 
hole, 6.5 miles northeast of Rocksprings, 2,300 feet (White, 1948, p. 6). Harde- 
man County: Acme (Dalquest, 1954, in litt.) ; 3 miles southeast of Lazare (Dalquest, 
1954, in litt.); Panther Cave, 10 miles southeast of Quanah (Blair, 1954, p. 242). 
Presidio County: Sierra Vieja (Blair and Miller, 1949, p. 76). Val Verde County: 


AMERICAN BATS—HANDLEY 197 


Mouth of Pecos River (Blair, 1952, p. 96). Oxuanoma: Blaine County: Salt 
Creek Canyon (Blair, 1939, p. 102); 3 miles southeast of Southard (Blair, 1939, 
p. 102). Cimarron County: Tesequite Canyon [1.7 miles southeast of Kenton] 
(Glass, 1951, p. 26). Greer County: 3.5 miles north of Jester (Glass, 1955, p. 128). 
Harmon County: 3 miles west and 1 mile south of Reed (Glass, 1955, p. 128). 
Woods County: 2 miles west of Edith (Blair, 1939, p. 102); 6 miles northeast of 
Freedom (Blair, 1939, p. 102); Marehew Cave, one-half mile south of Oklahoma- 
Kansas boundary (Hibbard, 1934, p. 237). Woodward County: 10 miles south 
of Freedom (Blair, 1939, p. 102). Kansas: Barber County: Several caves 3 to 5 
miles southwest of Aetna (Cockrum, 1952, p. 79; Frum, 1954, in litt.; Twente, 
1955a, p. 379); several caves 2 to 18 miles south of Sun City (Cockrum, 1952, 
p. 79; Hibbard, 1933, p. 233; Twente, 1955a, p. 379). Comanche County: Several 
caves 4 to 6 miles west and northwest of Aetna (Cockrum, 1952, p. 79; Hibbard, 
1934, p. 287; Twente, 1955a, p. 379). Conorapo: Boulder County: Boulder 
Canyon, 7,700 feet (Young, 1908, p. 407); Fourmile Canyon, 7,000 feet, near 
Crisman (Cary, 1911, p. 204); 12 miles south of Lyons (G. Allen, 1916, p. 341). 
El Paso County: 1 mile north of Glen Eyrie (near Colorado Springs) (Warren, 
1942, p. 80). Gunnison County: Gothic, 9,600 feet (Findley and Negus, 1953, 
p. 237). Larimer County: Fort Collins (Cary, 1911, p. 204); no exact locality 
(Miller, 1897, p. 53). Las Animas County: Trinidad (Warren, 1906, p. 267). 
SoutH Dakota: Custer County: Jewel Cave National Monument (National 
Park Service Collection, Washington, D.C.). Harding County: Ludlow Cave 
Hills (Biol. Surv. files). Stanley County: White River (Biol. Surv. files). 

CANADA: British CotumBia: Adam’s River, northwest of Shuswap Lake 
(Anderson, 1946, p. 32); Creston, Kootenay River (Anderson, 1946, p. 32); 
Keremeos, Similkameen Valley (Anderson, 1946, p. 32); Riske Creek and Williams 
River (Cowan and Guiguet, 1956, p. 67). 


Plecotus townsendii townsendii Cooper 


Plecotus townsendii Cooper, 1837, p. 73, pl. 3, fig. 6. 

S[ynotus] townsendit Wagner, 1855, p. 720. 

Corinorhinus townsendi Dobson, 1875, p. 356. 

Plecotus (Corinorhinus) macrotis Dobson, 1878, p. 180. (Part.) 
Corynorhinus macrotis townsendii Miller, 1897, p. 53. 
Corynorhinus macrotis intermedius H. W. Grinnell,3 1914, p. 320. 
Corynorhinus megalotis townsendi G. M. Allen, 1916, p. 344. 
Corynorhinus rafinesquit townsendi H. W. Grinnell, 1918, p. 345. 
Corynorhinus townsendit townsendit Handley, 1955c, p. 147. 


Hoxrotyrz: Not known to be in existence. Type locality: ‘‘Co- 
lumbia River,” herein restricted to Fort Vancouver, Clark County, 
Wash. 

Distrisution: Coast region of the northwestern United States 
and southwestern Canada (fig. 22). North to Comox, Vancouver 
Island, British Columbia. South to Santa Barbara County, Calif. 
Inland to eastern flanks of Coast Range in south; to eastern flanks of 
Cascade Range in north. Zonal distribution: Transition and Upper 
Sonoran. 


8 Holotype: MVZ No. 7753; adult female, skin and skull; collected 31 July 1909, by J. C. Hawver; 
Auburn, Placer County, Calif.; original No. 2387. 


198 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Description: Adult coloration: Mass effect of upperparts in 
darkest individuals between Natal Brown and Bone Brown, with 
little distinction between bases and tips of hairs; mass effect in palest 
individuals between Vandyke Brown and Verona Brown, with hair 
bases considerably paler than the tips, about Benzo Brown. Hair 
of underparts Fuscous at base; darkest individuals have hair tips of 
throat and breast between Snuff Brown and Bister, those of the 
belly Light Pinkish Cinnamon; palest individuals have hair tips of 
entire underparts Light Pinkish Cinnamon. Immature coloration: 
Similar to the darkest adults or slightly darker. 

Size averages medium for subgenus; forearm averages relatively 
short. Construction of skull heavy; rostrum relatively long, inflated, 
and not depressed; anterior nares, viewed from above, usually rounded 
in posterior outline. Canine strong; first upper incisor frequently 
with trace of secondary cusp; P* usually with anterointernal cingular 
cusp. 

MEASUREMENTS: See tables 12, 18. 

Comparisons: P. t. townsendii is the darkest colored and shortest 
winged population of the species P. townsendii. Compared with 
P. t. pallescens, into which it grades to the east and south, P. t. town- 
sendii is darker, averages slightly larger, and has a more robust skull, 
heavier rostrum, and anterior nares that average relatively larger 
and more rounded in posterior outline. 

Remarks: The type locality of this form is not “Columbia River, 
Oregon” as has been assumed by many authors (e.g., Miller, 1897, 
p. 53; G. M. Allen, 1916, p. 344; Anderson, 1946, p. 33; Miller and 
Kellogg, 1955, p. 110). Bailey (1936, p. 387) and Dalquest (1948, 
p. 163) were correct in naming “Fort Vancouver, Washington” as 
the type locality. 

Cooper’s description (1837, p. 73) of Plecotus townsendit was based 
on three specimens which he stated had been obtained by J. K. Town- 
send on the Columbia River. In the appendix to the narrative of his 
journey to the Columbia, Townsend (1839, pp. 324, 325) mentioned 
the species as follows: 

[Plecotus Townsendii is the] So-capual of the Chinook Indians .... Inhabits 
the Columbia river district, rather common. Frequents the store houses attached 
to the forts, seldom emerging from them even at night. This, and a species of 
Vespertilio, (V. subulatus), which is even more numerous, are protected by the 
gentlemen of the Hudson’s Bay Company, for their services in destroying the 
dermestes which abound in their fur establishments.—Towns. 

In these sentences from Townsend’s narrative there are many 
clues to the whereabouts of the type locality of P. townsendit. First 
is the fact that the bat ‘Inhabits the Columbia river district.”” Next, 
“Frequents the store houses attached to the forts.” According to 


AMERICAN BATS—HANDLEY 199 


his narrative, Townsend visited four Columbia River forts: Fort 
Walla-Walla, near the present site of Wallula, Washington, half a mile 
above the mouth of the Walla Walla River; Fort Vancouver, 6 miles 
above the mouth of the Willamette River, Clark County, Wash.; 
Fort William, on the southwest side of Sauvie Island, Multnomah 
County, Oreg.; and Fort George at Astoria, Clatsop County, Oreg. 
Fort William can be eliminated immediately, for it was an American 
establishment, built during Townsend’s visit. There the bats could 
not have been “protected by the gentlemen of the Hudson’s Bay 
Company.” ‘Townsend’s citation of a Chinook Indian name for the 
bat, ‘‘So-capual of the Chinook Indians,” eliminates Fort Walla-Walla, 
because the Chinooks normally did not occur above the cascades of 
the Columbia (Townsend, 1839, p. 220), although Townsend’s party 
did encounter a band at The Dalles (1839, p. 159). Furthermore, 
Cooper’s description agrees with the dark, coastal race of the bat, 
whereas a pallid form occurs in the vicinity of Fort Walla-Walla. 
Finally, Townsend’s description of Fort George (1839, p. 182) elimi- 
nates it as a possibility: 

We anchored off Fort George, as it is called, although perhaps it scarcely deserves 
the name of a fort, being composed of but one principal house of hewn boards, 
and a number of small Indian huts surrounding it, presenting the appearance, 
from a distance, of an ordinary small farm house with its appropriate outbuildings. 
Compare with this the statement, ‘frequents the store houses at- 
tached to the forts.’”’ Townsend described Fort Vancouver, where 
he had residence during his months on the Columbia, as follows 
(1839, pp. 170-172): 

The space comprised within the stoccade is an oblong square, of about one 
hundred, by two hundred and fifty feet. The houses built of logs and frame-work, 
to the number of ten or twelve, are ranged around in a quadrangular form . 
in the vicinity of the fort, are thirty or forty log huts . . . placed in rows, with 
broad lanes or streets between them, and the whole looks like a very neat and 
beautiful village. 

Fort Vancouver appears to be the logical choice for the restricted 
type locality of Plecotus townsendii Cooper. 

P. t. townsendii intergrades with the very differently colored 
P. t. pallescens over a wide area in central and northern California 
and northward between the Cascades and the Rockies. Allocation of 
specimens, especially in inadequate series, from much of this area 
to one race or the other is largely a matter of personal opinion. As a 
result, various authors have disagreed on just where the artificial 
boundary between the ranges of townsendii and pallescens should be 
set. Other authors have chosen to solve the problem by applying a 
subspecific name, intermedius, to populations of the intergrading area. 
For a discussion of this problem, see the remarks under P. t. pallescens 


(p. 192). 


200 PROCEEDINGS OF THE NATIONAL MUSEUM You. 110 


Dark-colored populations, resembling those at the type locality 
probably occur only in the Coast Ranges and humid lowlands west of 
the Cascade Range from northern California northward to Vancouver 
Island. Progressive increase in pallor apparently takes place even 
as the western slopes of the Cascades are approached. ‘Thus, in the 
Willamette Valley in Oregon, specimens from Carver and Vida, at 
lower elevations, are darker than those from McKenzie Bridge a little 
further up-stream. On the eastern slopes of the Cascades, as at 
Boulder Cave, Wash., specimens average even paler, but are, never- 
theless, nearer townsendit than pallescens. A single specimen from 
Happy Camp, Siskiyou County, Calif., is paler than most other 
specimens identified as townsendii, but it is from an area where 
townsendii theoretically should occur, and on the supposition that a 
larger series would show the average to be nearer townsendit, it has 
been identified as that form. 

Samples from collecting stations in the lowlands and Coast Range 
south of San Francisco Bay are variable, showing intergradation with 
P. t. pallescens. From Santa Barbara County northward the samples 
average nearer P. t. townsendii in morphological characters. 

Plecotus appears to be absent from most parts of the Great Valley 
of California. 

SPECIMENS EXAMINED: A total of 83, from the following localities: 

UNITED STATES: Caurrornia: Alameda County: Mission San Jose, 25s 
(MVZ). Fresno County: New Jaria [-New Idria?], 1b (USNM). Marin County: 
Nicasio, ls (USNM). Monterey County: Stonewall Creek, 1,300 feet, 6.3 miles 
northeast of Soledad, 2s (MVZ). Napa County: Mount Veeder [1,950 feet], is 
(USNM). San Benito County: Bear Valley [The Pinnacles, near Cook, 2,000 
feet], 2s (USNM); 4 miles west-northwest of Panoche, Is (MVZ). San Luis 
Obispo County: 5 miles south-southwest of Adelaida, 2,200 feet, 1s (MVZ); 
San Simeon, Ils (MVZ); 6.5 miles southeast of Shandon, 1s (MVZ). San Mateo 
County: 2.0 to 2.5 miles east of Pescadero, 2s (MVZ). Santa Clara County: 
Evergreen, 2s (MVZ); 2.25 miles northwest of San Felipe, 2s (MVZ). Santa 
Cruz County: 2 miles southwest of Brookdale, 1s (MVZ); Mouth of Meder Creek, 
2s (MVZ); 2 to 3 miles east of Santa Cruz, 2s (MVZ). Siskiyou County: Happy 
Camp, ls (USNM). Orxaon: Clackamas County: Carver, 4s (CNHM), 3s 
(USNM). Curry County: Gold Beach, 5s (CNHM), Is (MCZ), 1a (USNM). 
Lane County: Creswell, 1a (USNM); McKenzie Bridge, 3s (USNM); Vida, 1s 
(USNM). Wasuineton: Kittitas County: Boulder Cave, Naches River, 43 miles 
northwest of Yakima, l4as (USNM). Thurston County: Olympia, 1s (USNM). 

CANADA: Britis CotumBia: Vancouver Island: Comox, 1s (USNM). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: 


UNITED STATES: Caurirornia: Alameda County: 4 miles east-southeast of 
Mission San Jose (Dalquest, 1947, p. 20). Colusa County: 2.25 miles south- 
southeast of Wilbur Springs, 1,800 feet (Frum, 1954, in litt.). Humboldt County: 
Carlotta (Dickey, 1922, p. 116). Lake County: Long Valley (Dalquest,1947, 
p. 20); Lucerne (Frum, 1954, in litt.); 4 miles east of Upper Lake (Dalquest, 


AMERICAN BATS—-HANDLEY 201 


1947, p. 20). Marin County: Inverness (Pearson, et al., 1952, p. 275); 1.3 miles 
southeast of Inverness (Frum, 1954, in litt.). Mendocino County: 1.5 miles 
north of Gualala (Dalquest, 1947, p. 20). Napa County: Aetna Springs (Pearson, 
et al., 1952, p. 275); one-half mile south of Angwin, 1,600 feet, and 1 mile south of 
Angwin, 1,500 feet (Dalquest, 1947, p. 20); 4 miles south of Calistoga (Dalquest, 
1947, p. 20); near Knoxville, 2,000 feet (Pearson, et al., 1952, p. 279); Pope Creek, 
8 miles northwest of Monticello (Dalquest, 1947, p. 20). San Benito County: 
Hernandez (H. W. Grinnell, 1914, p. 320). Sonoma County: 10 miles east of 
Stewart’s Point (Dalquest, 1947, p. 20). Orscon: Clackamas County: Clacka- 
mas (V. Bailey, field notes). Josephine County: Siskiyou Mts., Oregon Caves 
National Monument (Roest, 1951, p. 346). Lane County: Eugene (Bailey, 1936, 
p. 387). Multnomah County: Portland (Bailey, 1936, p. 387). WasHINGTON: 
Clallam County: Elwha River (Johnson and Johnson, 1952, p. 33). Clark County: 
Fort Vancouver (Bailey, 1936, p. 386). King County: Seattle (Dalquest, 1948, 
p. 163). Mason County: Lake Cushman (Dice, 1932, p. 48). Pierce County: 
Puyallup (Dalquest, 1938, p. 213). San Juan County: Blakeley Island (Dal- 
quest, 1948, p. 162); Friday Harbor (Dalquest, 1948, p. 161). County uncertain: 
Hood’s Canal (Johnson and Johnson, 1952, p. 33). 

CANADA: British Cotumpra: Nanaimo Bay, Vancouver Island (Anderson, 
1946, p. 33); Newcastle Island (Anderson, 1946, p. 33). 


Plecotus townsendii virginianus Handley 


Corynorhinus macrotis A. H. Howell, 1909, p. 68. (Part.) 
Corynorhinus megalotis megalotis G. M. Allen, 1916, p. 338. (Part.) 
Corynorhinus rafinesquit Thomas, 1916, p. 127. (Part.) 
Corynorhinus townsendii virginianus Handley, 1955c, p. 148. 


Hoxtotyese: USNM 269163; adult male, skin and skull; collected 
12 November 1939, by W. J. Stephenson; Schoolhouse Cave, 4.4 
miles northeast of Riverton, 2,205 feet, Pendleton County, W. Va.; 
no collector’s number. 

Distrrisution: Central part of Appalachian Highlands in eastern 
Kentucky, western Virginia, and eastern West Virginia (fig. 23). 
North to Hardy, Grant, and Preston Counties, W. Va. West to 
Preston, Tucker, and Randolph Counties, W. Va. South to Ran- 
dolph and Pendleton Counties, W. Va. East to Pendleton and Hardy 
Counties, W. Va. Apparently isolated populations further south 
and west, in Tazewell County, Va., and Powell County, Ky. Zonal 
distribution, Transition. Altitudinal distribution, 1,700 to 3,500 
feet. 

Descrietion: Adult coloration: Hair of upperparts about Benzo 
Brown at base; subterminal band orange-brown, tip dark brown; 
mass effect between Prout’s Brown and Bister. Hair of underparts 
between Light Vinaceous-Cinnamon and Light Pinkish Cinnamon 
distally, occasionally somewhat darker on throat and chin; bases, 
except of those hairs bordering membranes, Fuscous; border hairs 
monocolored. Distinction between tip and base of hair sharp on under- 
parts, poor on dorsum. Immature coloration: Hair of upperparts 


202 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


between Natal Brown and Benzo Brown distally; bases slightly grayer, 
hardly distinguished from tips. Hair of underparts Mouse Gray at 
base, pale buff at tip; distinction between base and tip slight. 

Size averages medium for subgenus; forearm averages relatively 
long. Rostrum relatively long and not depressed; anterior nares 
(viewed from above) wide and rounded in posterior outline. First 
upper incisor usually without trace of secondary cusp. 

MEASUREMENTS: See tables 12, 18. 

Comparisons: Compared with P. ft. ingens, P. t. virginianus is 
more sooty dorsally and averages slightly smaller in all dimensions; 
the first upper incisor rarely has a trace of a secondary cusp; and the 
rostrum is less heavy and inflated. 

Remarks: The Appalachian populations (P. t. virginianus) appear 
to be isolated by no less than 600 miles from the nearest closely 
related populations (P. ¢. ingens) in the Ozark Highlands. Despite 
this separation, the two forms are not strikingly different, and although 
there is no possibility of demonstrating intergradation between them, 
they may be treated best as being conspecific. On the other hand, 
the Appalachian populations of P. townsendii are specifically distinct 
from P. rafinesquii, whose geographic range they overlap. This 
peculiar pattern of distribution, perhaps related to Pleistocene dis- 
turbances, is discussed in the section on history (p. 215). 

There is no obvious explanation for the limited and apparently 
discontinuous distribution of the populations within the bounds of 
the Appalachian Highlands. The species is relatively numerous, even 
the most abundant bat in some caves, in two adjacent drainage sys- 
tems in northeastern West Virginia. It has been found in at least 15 
caves at the head of the Potomac River and in four caves at the head 
of the Cheat River. All of these caves are in limestone, in an area 
30 miles wide and 40 miles long. Three other populations have been 
discovered away from this main body: One a few miles to the north 
in Preston County, W. Va. (Kellogg, 1937, p. 450), another 150 miles 
to the south-southwest in a limestone cavern in Tazewell County, Va. 
(Howell, 1909, p. 68; Mohr, 1933, p. 49), and the third in a cave 200 
miles to the west-southwest in Powell County, Ky. (Barbour, 1957, 
p. 141). The intervening extensive limestone caverns, mostly well 
explored, in Pocahontas, Greenbrier, and Monroe Counties, W. Va., 
and in the Valley of Virginia do not seem to be occupied by the 
species. 

Isolation such as apparently exists among the Kentucky, Virginia, 
and West Virginia populations of this species favors differentiation. 
It is not surprising then that differentiation is evident. Virginia and 
Kentucky specimens average smaller than those from West Virginia. 
Kentucky and West Virginia specimens are similar in color to Virginia 


AMERICAN BATS—-HANDLEY 203 


specimens obtained in 1908. However, specimens collected in Virginia 
in 1957 average considerably darker, more sooty, both dorsally and 
ventrally, than those collected recently in Kentucky and West Virginia 
or in the same Virginia cave fifty years earlier. They resemble typical 
P. t. townsendii in coloration. 

SPECIMENS EXAMINED: A total of 100, from the following U.S. 
localities: 


Kentucky: Powell County: Natural Bridge State Park, 2s (UK). Virerntia: 
Tazewell County: Burkes Garden, 3,200 feet, 9as (USNM). West VircInia: 
Grant County: 10 miles south of Petersburg, 3s (CM). Pendleton County: Cave 
Mountain Cave, 2,450 feet, 1.4 miles west of Brushy Run, llas (USNM); Hellhole, 
2,200 feet, 3.6 miles northeast of Riverton, 5a (USNM); Hoffman School Cave, 
2,175 feet, 4.9 miles south-southwest of Franklin, 2s (USNM); Schoolhouse Cave, 
2,205 feet, 4.4 miles northeast of Riverton, 3las (USNM); ‘“‘Smokehole” [=Smoke- 
hole Cave, 1,730 feet, 1.9 miles west-northwest of Brushy Run?], 29abs (AMNH); 
“Cave Rock Cave, Smokehole” [=Cave Mountain Cave, 2,450 feet, 1.4 miles 
west of Brushy Run?], 8s (AMNH). 


ADDITIONAL RECORDS: The following records probably refer to this 
form: 


Kentucky: Rowan County: Carter Caves (Welter and Sollberger, 1939, p. 79). 
West VirGinia: Grant County: Klines Gap Cave, 1,700 feet, 4 miles southwest 
of Maysville (Frum, 1954, in litt.); Peacock Cave, 1,755 feet, 7.6 miles southwest 
of Petersburg (Fowler, 1941a, p. 35) ; Petersburg (McKeever, 1952, p. 47). Hardy 
County: Baker’s Cave, near Durgon [apparently not the ‘‘Baker Cave” of Davies, 
1949, p. 139] (Wilson, 1946, p. 86). Pendleton County: Mill Run Cave, 1,800 
feet, one-half mile southeast of Kline (Frum, 1954, in litt.); Seneca Caverns, 
2,200 feet, 3 miles northeast of Riverton (Reese, 1934, p. 47); Sinnit Cave, 2,250 
feet, one-quarter mile northwest of Moyers (Frum, 1954, in litt.) ; Thompson Cave, 
2,350 feet, 1.5 miles east-southeast of Riverton (Frum, 1954, in litt.) [=5.6 miles 
north-northeast of Circleville, 2,350 feet (McKeever, 1952, p. 47)?]; Torys Cave, 
2,000 feet, 5 miles southwest of Franklin (Frum, 1954, in litt.) [=Tory’s Cave, 10 
miles south of Franklin (Barbour, 1951b, p. 369)?]; Trout Cave, 5 miles southwest 
of Franklin (Fowler, 1941b, p. 47). Preston County: No exact locality (Kellogg, 
1937, p. 450; Barbour, 195la, p. 50). Randolph County: 6.7 miles east-northeast 
of Glady, 3,500 feet (McKeever, 1952, p. 47); The Sinks, Cave No. 1, 3,500 feet, 
one-half mile west of Osceola (Reese, 1934, p.47). Tucker County: Arbegast Cave, 
2,600 feet, 6 miles southeast of Hendricks (Frum, 1954, in litt.); Cave Hollow 
Cave, 2,400 feet, 5 miles southeast of Hendricks (Frum, 1954, in litt.). 


INCERTAE SEDIS 
Vespertilio maugei Desmarest 


Vespertilio mauget Desmarest, 1819, p. 480. 
Plecotus Maugei Lesson, 1827, p. 96. 

Vespertilio (Plecotus) Maugei Cooper, 1837, p. 71. 
Synotus Maugei Wagner, 1855, p. 719. 

Vespertilio mauget H. E. Anthony, 1916, p. 360. 


Houotyrr: From Puerto Rico. Collected by Maugé. The pres- 
ent disposition of the holotype is unknown. Some of Desmarest’s 


204. PROCEEDINGS OF THE NATIONAL MUSEUM You. 110 


types are in the Paris Museum, but Rode (1941) did not list Vesper- 
tilio maugei in his catalog of the type specimens of Chiroptera in the 
Paris Museum. 

Distrispution: ‘Puerto Rico.” 

Description: Following is my translation of Desmarest’s (1819, 
p. 480) description: 

Vespertilio maugei, new species. 

This bat, brought back from Puerto Rico by the late Maugé, is of medium size 
and approaches, in its physiognomy, the barbastelle, notably by the distinguishing 
character of the union of the ears at their internal bases. 

Its pelage is long, silky, of a blackish brown above, lighter beneath, particularly 
in the region near the interfemoral membrane, where it becomes almost white; 
its tail is nearly as long as the body; its membranes are of a dark gray. 

Its mouth is provided with four upper incisors of which the two inner are the 
largest, separated from one another and bifurcate, the exterior being simple; 
the six lower incisors overlap one another and have three lobes, and medium-sized 
canines are very closely followed by the molars. 

The muzzle is short, slender, pointed, and forms, with the ears, a right angle; 
the nose is rather broad, that is to say, that the nostrils are separated by a cartilage 
in the form of a plate, which resembles somewhat a lyre; the ears are large, their 
extremity is rounded, their exterior border notched, with a longitudinal fold 
provided with very dense and very noticeable hairs; the tragi are pointed and do 
not reach half the height of the ears; the eyes are small and placed at the base of the 
latter. 


Desmarest (1820, p. 145) modified this description slightly the fol- 
lowing year. My translation of the most significant portions of the 
new description is as follows: 

Ears very large, united, notched exteriorly near the tip . .. Dimensions, a 
little larger than the barbastelle. . . .” 

Remarks: For the early 19th century these are surprisingly detailed 
descriptions. However, the bat described is quite unlike any bat 
known to occur in the West Indies. Eptesicus is the only vespertil- 
ionid that has subsequently been collected on Puerto Rico, and 
Myotis, Nycticeius, and Lasiurus are the only other vespertilionid 
genera recorded from the West Indies. 

Vespertilio maugei has been overlooked or ignored by most authors. 
Those who have considered it (see synonymy, p. 203) have thought 
it to be a species of Plecotus. Plecotus has not been collected subse- 
quently in the West Indies, however, and no skeletal remains of 
Plecotus have been recognized in the sub-Recent bone deposits that 
are abundant in West Indian caves. 

Anthony (1918, p. 360) suggested that Maugé’s specimen actually 
was a Plecotus which had been obtained in the United States and was 
attributed to Puerto Rico by mistake. It is true that Desmarest’s 
descriptions agree in some details with Plecotus, but there are many 
discrepancies: Coloration of the underparts is wrong, the canines 


AMERICAN BATS—HANDLEY 205 


are not “very closely followed by the molars,” the nostrils are not 
separated by a lyre-shaped plate, and the auricles are not notched on 
their exterior borders. 

There is a possibility that the description of V. mauget is a com- 
posite. The dental characters could apply to Eptesicus. Externally, 
the West Indian phyllostomid Macrotus waterhousi: agrees fairly 
well with Desmarest’s description. Although its tail is a trifle short, 
and its nose leaf can hardly be said to be “‘lyre-shaped,” it nevertheless 
has unusually large ears, connected at their internal bases, and its 
coloration exactly matches Desmarest’s description. On the other 
hand, with regard to its dentition, Macrotus bears no resemblance 
whatsoever to the description of “V. mauger.”’ 

It seems probable that the bat Desmarest described was 
the Eurasian Barbastella, with which his descriptions agree in all 
details. In all probability the specimen did not originate in Puerto 
Rico or elsewhere in the Western Hemisphere. Consequently, the 
name Vespertilio maugei should be placed in the synonymy of Barba- 
stella and should be disassociated from Plecotus and the American 
fauna. 


History 


PALEONTOLOGICAL RECORD 
CHIROPTERA 


The paleontological record is so fragmentary for bats that few 
conclusions can be drawn from it. Crania and upper dentitions are 
disappointingly rare in fossil bat material. The commonest remains 
are mandibles or long bones, neither of which, when isolated and 
fragmentary, are very satisfactory for studies of evolution and rela- 
tionships or even for taxonomic classification. Paradoxically, the 
oldest known chiropteran remains—those of Archaconycteris, Palaeo- 
chiropteryz, and Cecilionycteris of the Middle Eocene—are also 
the best preserved. Complete dentitions and skeletons are intact, 
and even the outlines of the wing membranes can be traced (G. 
Allen, 1939, p. 177). 

Most of the work on fossil Chiroptera has been done in Europe. 
A majority of the named forms and a preponderance of the literature 
on the subject are from that region. Investigations conducted else- 
where, principally in China and in the United States are insignificant 
by comparison. Preglacial fissure and cave microfaunas have been 
virtually untouched outside of Europe. 

The European record indicates that the extinct families Archaeonyc- 
teridae and Palaeochiropterygidae occurred with some variety in 
Middle Eocene time. Rhinolophidae were numerous and predominant 
in Upper Eocene and Oligocene deposits. Species of Megadermatidae 


206 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


and Emballonuridae were somewhat less numerous during the same 
period. The Vespertilionidae became abundant somewhat later. 


FAMILY VESPERTILIONIDAE 


According to Revilliod (1919, p. 94), the genus Stehlinia [= Nycter- 
obius Revilliod and Revilliodia Simpson (Handley, 1955a, p. 128)] of 
the Quercy fauna of the Upper Eocene or Lower Oligocene of Europe 
is the most ancient known vespertilionid. It has cranial characters 
of the Vespertilionidae, but is distinguished from modern genera by 
several primitive characters, including very elongated rostrum, rela- 
tively very long premolar series, and 2-rooted P!. It most closely 
resembles some of the small species of Myotis, which is generally con- 
ceded to be the most primitive living genus of the family. 

True Myotis appears for the first time in the Stampien and Aqui- 
tanien faunas of the Middle and Upper Oligocene of Europe [Myotis 
salodorensis Revilliod (1919, p. 94) and Myotis insignis von Meyer 
(Revilliod, 1922, p. 166)]. Other Myotis have been described from 
the Middle Miocene of Europe [Myotis antiquus Gaillard (1899, p. 8) 
and Myotis sancti-albani Viret (1951, p. 19)], and Myotis-like remains 
occur in Lower Miocene beds in Florida (Lawrence, 1943, p. 364). 
Some of the Recent species of Myotis (M. bechsteinn, M. mystacinus, 
and M. oxygnathus) were possibly derived by Pliocene or Lower 
Pleistocene time (e.g., see Heller, 1930, p. 154; Kormos, 1937, p. 338; 
and Kowalski, 1956, p. 331), although most late Tertiary and Quater- 
nary material has been described as now extinct species (e.g., see 
Heller, 1936, p. 113; Kormos, 1934, p. 306; Kowalski, 1956, p. 331; 
Sickenburg, 1939, p. 8; and Wettstein, 1923, p. 40). 

Another primitive vespertilionid genus, Miniopterus, is thought to 
have been recognizable as far back as the Miocene epoch (WM. fossilis, 
Zapfe, 1950, p. 60). The Lower Pliocene Mystipterus (Hall, 1930, 
p. 319) from Nevada, first supposed to be related to Miniopterus, 
subsequently has been shown to be an insectivore (Patterson and 
McGrew, 1937, p. 256). 

The oldest Eptesicus-like remains are from Lower Miocene beds 
of Florida (Lawrence, 1943, p. 367). Other Eptesicus-like genera, 
Samonycteris (Revilliod, 1919, p. 95) and Pareptesicus (Zapfe, 1950, 
p. 58), have been described from the Miocene of Europe. From the 
Upper Pliocene of Arizona comes Simonycteris (Stirton, 1931, p. 27), 
also possibly a relative of Eptesicus. The most ancient remains at- 
tributed to true H’ptescius are those discovered in the Pliocene or Lower 
Pleistocene of Hungary (Kormos, 1930a, p.41). Brown (1908, p. 174) 
regarded Middle Pleistocene material from the Conard Fissure of 
Arkansas as being only subspecifically distinct from the Recent 
Eptesicus fuscus. Gidley and Gazin (1938, p. 11), with more abundant 


207 


AMERICAN BATS—HANDLEY 


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208 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


material from the Pleistocene of Cumberland Cave, Md., agreed that 
the relationship of grandis to the Recent species is close but believed 
it to be sufficiently different to warrant specific separation. 

Records for Barbastella (Wettstein, 1931 p. 780), Lasiurus (Hibbard, 
1950, p. 134), and Vespertilio (Kormos, 1934, p. 312) extend back at 
least to the Pliocene. Numerous other Recent vespertilionid genera 
have been identified in Pleistocene deposits. 

To recapitulate: The oldest recognizable vespertilionid was of 
Upper Eocene or Lower Oligocene age. The morphologically more 
primitive Recent genera emerged in the Oligocene and Miocene. 
Recent species of these genera were in some instances perhaps derived 
as early as the Pliocene, although fossil species of the same genera 
persisted into the Pleistocene (fig. 25). 


Genus Plecotus 
Vespertilio (Plecotus) grivensis Depéret 


Remains undoubtedly referable to Plecotus have been found only 
in Upper Pliocene (possibly actually Lower Pleistocene) and Pleisto- 
cene deposits. Records of greater antiquity are erroneous or question- 
able. The controversial Vespertilio (Plecotus) grivensis of Depéret 
(1892, p. 11, pl. 2) from the Vindobonien (Middle Miocene) fauna of 
Grive Saint-Alban, France,is a case in point. Depéret’s material 
consisted of two fragmentary mandibles. Gaillard (1899, p. 7) later 
reported a total of four mandible fragments from Grive Saint-Alban 
in the Museum of Lyon. All apparenty agreed with V. grivensis, 
but did not elucidate the unknown portions of the mandible. 

Depéret (loc. cit.) believed that the remains most closely resembled 
Plecotus, and to this opinion Wettstein (1931, p. 779) agreed. On the 
other hand, Gaillard (loc. cit.) preferred to leave the generic desig- 
nation questionable. When Revilliod (1922, p. 172) reviewed all the 
Tertiary Chiroptera, his conclusion was that Vespertilio grivensis 
was in actuality a Myotis. 

More recently, Viret (1951, p. 21) redescribed and figured the 
specimens (now totaling five fragments) of Vespertilio grivensis which 
were initially reported by Depéret and Gaillard. Viret argued con- 
vincingly that the type and one additional fragment should beregarded 
as Rhinolophus grivensis, while the third fragment proved to be Myotis 
sancti-albani, the fourth Sorex pusillus, and the fifth indeterminate. 


Plecotus crassidens Kormos 


Next in geologic age is the Pliocene or Lower Pleistocene Plecotus 
crassidens of Kormos (1930b, p. 238) from Hungary and Poland. 
Kormos did not figure the fragmentary mandible upon which he based 
the name, and there was nothing positive about his description. 


AMERICAN BATS—-HANDLEY 209 


Subsequently, however, Kowalski (1956, p. 359, pl. 3) figured and 
described in detail ten maxillary and cranial frgaments which he had 
compared directly with Kormos’ type of P. crassidens. He character- 
ized P. crassidens as follows: RostRUM BROAD (much more so than in 
P. auritus), FLATTENED, AND WITH A MEDIAN DEPRESSION; intermaxil- 
lary notch broad and cordate; palate extends one molar breadth poster- 
ior to toothrows;LACHRYMAL [AND SUPRAORBITAL] REGIONS NOT RIDGED; 
P‘ labially broader and less obliquely placed than in P. auritus; molars 
larger and more massive than in P. auritus; coronoid process of man- 
dible distinctly pointed; angular process broad and blunt and deflected 
outward; P; smaller than P,; P, 2-rooted, subquadrate in basal cross- 
section, provided with strong cingulum, and not so high as Mi; Ms; 
talonid narrower than trigonid. Also apparent from Kowalski’s 
figures, although not stated in his description are the facts that the 
braincase bulges dorsally in the parietal region, and THE ZYGOMATIC 
ARCH IS NOT EXPANDED IN ITS MIDDLE THIRD (posterior third missing). 

The characters in larger type relate P. crassidens to the subgenus 
Corynorhinus, while the shape and 2-rooted condition of P, show an 
affinity to the subgenus Plecotus. 

On the basis of these specimens Kowalski (1956, p. 340) supposed 
that the North American Corynorhinus might be a relict of the Tegelien 
Interglacial of Europe (possibly equivalent to the Aftonian Inter- 
glacial of North America), analogous to the several species of the 
Tegelien flora, now extinct in Europe but persisting in North America. 


Plecotus abeli Wettstein 


Wettstein (1923, p. 39; 1931, p. 779) described abundant Pleistocene 
remains from the Drachenhéhle in Austria as belonging to a distinct 
species, Plecotus abeli, averaging slightly smaller than the Recent 
form and having the cusps of the lower molars not so high and sharp. 
Kowalski (1956, p. 361), judging from Wettstein’s descriptions, ven- 
tured the opinion that P. abeli was no more than subspecifically 
distinct from P. auritus. 


Plecotus auritus Linnaeus 


Most of the undoubted remains of Plecotus that have turned up in 
the Pleistocene and dubiously Upper Pliocene fissure and cave de- 
posits of Europe have been identified as Plecotus auritus, the Recent 
species now inhabiting Europe (e.g., see Heller, 1930, p. 154; 1936, 
p. 112; Kowalski, 1956, p. 361; and Sickenburg, 1939, p. 9). 


Corynorhinus alleganiensis Gidley and Gazin 


Corynorhinus alleganiensis from Cumberland Cave, Md. (Gidley, 
1913, p. 96; Gidley and Gazin, 1933, p. 345; 1938, p. 12), is the best 


210 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


known North American fossil Plecotus. The geologic age of this 
species cannot be determined stratigraphically, since the material 
upon which it is based is from a fissure type deposit. However, the, 
fauna associated with the bat remains is certainly Pleistocene in age 
and Gazin (in Gidley and Gazin, 1938, p. 7) concluded from the number 
of extinct forms present and from the difficulty encountered in identi- 
fying others as Recent that the Cumberland Cave fauna can hardly 
be considered Upper Pleistocene, and is more likely Middle Pleistocene 
in age. There is no reason to suppose that the bat remains were not 
deposited at the same time as the remainder of the Cumberland Cave 
fauna. 

P. alleganiensis was similar in many respects to (and possibly 
directly ancestral to) P. townsendii, the Plecotus occurring nearer 
than any other to Cumberland Cave today. It possessed the primitive 
characteristics of supraorbital ridge and lack of a sagittal crest. See 
pages 137-139 for a full description of P. alleganiensis. 


Corynorhinus tetralophodon Handley 


The other American Pleistocene species, C. tetralophodon, from San 
Josecito Cave, Nuevo Leén, Mexico, is known from a single skull. It 
was found in a cave deposit of indeterminable age, but judging from 
its similarity to Recent forms and from the associated fauna, it is 
probably Upper Pleistocene (Wisconsin) in age and younger than 
Plecotus alleganiensis. It differs from Recent forms in possession of 
a fourth commissure on M? and in failure of the temporal ridges to 
coalesce completely. Probably it was related most closely to P. 
townsendii, but possibly was not a direct ancestor of that species. 
See page 140 for a full description of P. tetralophodon. 


ORIGIN AND DISPERSAL OF EUDERMA AND PLECOTUS 


The paleontological record is so incomplete that a discussion of 
the origin and dispersal of Huderma and Plecotus necessarily must rely 
heavily upon speculation. However, with the historical record of 
other vespertilionid groups in mind, and with the evidence of present 
day geographic distribution of taxonomic characters in Huderma and 
Plecotus as a factual foundation, there is ground for speculation on the 
evolution of these genera. In the systematic section and in the pre- 
ceding paragraphs of this section the facts have been presented. 
The remainder of this section is devoted to an interpretation of them. 

Several factors govern the dispersal of bats. Of paramount im- 
portance are food supply and shelter. For insectivorous forms, such 
as Huderma and Plecotus, the vegetation controlling the distribution 
of the bat’s insect prey and the climate controlling the distribution 
of the vegetation are the ultimate critical factors in the bat’s dispersal 


AMERICAN BATS—HANDLEY 211 


as far as food supply is concerned. For antisocial, nonmigratory, 
cool climate inhabitants, such as some Plecotus are, not only is the 
presence of caves suitable for hibernation essential, but the caves 
must be sufficiently abundant to allow a degree of privacy and isola- 
tion from other species of bats. Unfortunately, it is not possible to 
index all of these factors through time. Otherwise, a more satisfactory 
picture of the differentiation and dispersal of these bats might be 
drawn with less recourse to speculation. 

The present distribution of Huderma and Plecotus and their relative, 
Barbastella, coincides rather closely with the North Temperate zone. 
Thus, it may be assumed that tropical and boreal climates have 
served as barriers to their dispersal. The center of origin for the 
group may have been either Nearctic or Palearctic. Since distantly 
related primitive forms and closely related progressive forms occur in 
both regions, a hypothesis of multiple invasion is required to explain 
present distributions regardless of which region may have served as 
the center of origin. No paleontological record is available for the 
more primitive forms, so paleontology gives no hint of origin or an- 
cestry. If it be assumed that the area of most active differentiation 
and greatest elaboration of forms indicates the center of origin of a 
group of animals, and that the most advanced, most progressive, and 
most specialized form is to be found about the center of origin, while 
the most primitive, most conservative, and least progressive is to be 
found on the peripheries (Matthew, 1939, p. 32), then it must be 
reckoned that this group of bats originated in the Nearctic. 


ANCESTRAL FORMS 


It is probable that the ancestral forms were derived early in the 
Cenozoic, since the related living genus Myotis is recognizable at least 
as far back as Middle Oligocene. During the Eocene and Oligocene 
epochs, low relief of the northern continents and lack of climatic 
barriers permitted a moist temperate forest to extend from Greenland 
across northern America to Alaska, Siberia, northern Europe, and 
Spitzbergen. A land bridge at Bering Strait intermittently connected 
North America and Eurasia [indicated by strong Lower Eocene, 
Upper Eocene, and Lower Oligocene migrations of mammals (Simp- 
son, 1947, p. 625)]. The temperate forest of this region was char- 
acterized by redwood (Sequoia), and included such other trees and 
shrubs as alder (Alnus), dogwood (Cornus), maple (Acer), oak (Quer- 
cus), ash (Fraximus), madrone (Arbutus), beech (Fragus), and elm 
(Ulmus) (Cain, 1944, p. 107). During this period the ancestors of 
Euderma and Plecotus possibly could have had a Holarctic distribution. 

Continental uplift, beginning in the Oligocene epoch and continuing 
to the present time, led to a gradual southward spread of the temperate 


212 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


forests, so that in the Miocene they became discontinuous, with seg- 
ments in North America and Eurasia (Cain, 1944, p. 107). Presum- 
ably, the distribution of the ancestors of Huderma and Plecotus may 
have become discontinuous at the same time, providing isolation for 
major evolution within the group. 

The Eurasian segment developed into an evolutionary dead end, 
Barbastella, which, though greatly modified in form, represents the 
ancestral stock. Barbasiella has a geographic range most remote from 
the postulated Nearctic center of origin of the group. 


Genus Fuderma 


Meanwhile, in the North American segment, evolution was rapid 
and spectacular, directed toward auditory specialization. Huderma is 
a derivative of the early stages of this evolution. It is the most 
specialized, and, in many respects, the most primitive of the big-eared 
forms. It must have had a long period of evolution along its own 
line, distinct from that of the other members of the group. Whereas 
Euderma has developed characters of its own, such as the connection 
of the posterior basal lobe of the auricle with the tragus, unique colora- 
tion, reduction of dentition, and specialization of the canines, among 
the other big-eared forms (genus Plecotus) there have been only minor 
variations of features common to all. The nature of the isolating 
mechanism that allowed the divergence of the Plecotus and Huderma 
stems is not apparent. Possibly Huderma was southern and wide- 
spread during Pliocene and had its range restricted and displaced by 
the more northern Plecotus during the Pleistocene glaciations. 


Genus Plecotus 
Subgenus Idionycteris 


The species of Plecotus seem to represent evolutionary stages. 
P. phyllotis is most primitive and conservative, and most like Huderma; 
P. auritus is somewhat more progressive; and P. mexicanus, P. rafines- 
qu, and P. townsendui are most progressive. P. phyllotis presumably 
diverged from the main stem while it still bore much resemblance to 
the earlier derived Huderma. P. phyllotis failed to match the special- 
izations of the later derived forms, and if abundance and area of 
distribution be taken as measures of success, it definitely occupies a 
position inferior to them. It seems to be extremely rare, and inhabits 
a restricted area on the fringe of the ranges of P. mexicanus and 
P. townsend. On the other hand, the more progressive P. auritus 
and P. townsendw are widely distributed, with ranges trans-continental 
in scope, and are relatively abundant. P. auritus is even the most 
abundant bat in some parts of its range. 


AMERICAN BATS—HANDLEY 213 


Subgenus Plecotus 


Invasion of Eurasia by way of the Bering Strait area by the P. 
auritus stock probably occurred during the Pliocene [much migration 
in Middle and Upper Pliocene (Simpson, 1947, p. 625)]. This would 
have required @ more northern distribution than any species of Plecotus 
has today. Changes in habits or habitat could have allowed a more 
northern range. 

Species of Plecotus now range into the warmer portions of the Cana- 
dian life zone (and its Eurasian equivalent). The cooler portions of this 
zone lie within 1,000 miles of Bering Strait. Only aslight amelioration 
of today’s climate would be required to provide similar floral conditions 
to Bering Strait. The present wide distribution of cool climates seems 
to be an exception in geological history. Even during the Pleistocene 
“Tce Age” warm climates and their associated faunas extended farther 
north during the interglacial periods than they do today. Presumably, 
during the Pliocene, warmer climates may have extended even farther 
north. 

If the pre-Pliocene Plecotus-like bats were not cave inhabitants, then 
a change in habit to include cave roosting would have allowed a more 
northern dispersal, provided suitable food supplies and adequate caves 
were available. Resort to caves for hibernation permits habitation of 
areas with colder winters. It also provides refuge in eras of changing 
climate, for modification of the climate within the cave may be negli- 
gible in comparison with the change on the surface. Barbastella and 
Plecotus auritus frequently roost during the summer under loose bark 
on tree trunks, in cavities in trees and under the eaves of houses, but 
usually hibernate in caves or in attics of houses. The most primitive 
living species of the subgenus Corynorhinus, P. rafinesquii, appears 
to be primarily a tree rooster, whereas the progressive species P. 
townsendiz seems to roost only in caves and houses. 

The ancestors of Plecotus auritus spread through most of the tem- 
perate portions of Eurasia, but were blocked from southward expansion 
into Maylasia and Africa by the hot climates of the equatorial regions. 
They came eventually to overlap broadly and perhaps partially dis- 
place the range of their relative, Barbastella, a rather rare bat whose 
continent-spanning range averages somewhat more southern than the 
range of P. auritus. 

The Plecotus that now inhabits Eurasia and northern Africa is 
probably not very different in appearance from the pre-Pleistocene 
migrant. This may be inferred from its similarity to the primitive 
New World species P. phyllotis and Pleistocene P. alleganiensis and 
P. crassidens. If a common origin for these forms and P. auritus be 
envisioned, and if a pre-Pleistocene dispersal of Plecotus into Eurasia 


VOL. 110 


PROCEEDINGS OF THE NATIONAL MUSEUM 


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AMERICAN BATS—-HANDLEY 215 


be accepted, then the morphological similarities between Old and New 
World Plecotus must be characters that were shared before dispersal 
occurred. 

The primitive Corynorhinus-like Plecotus crassidens may have 
reached Eurasia in a secondary Pliocene invasion, bringing it into 
contact with the already widespread and firmly established P. auritus. 
Apparently it did not persist beyond the early Pleistocene and left 
no derivatives. 


Subgenus Corynorhinus 


The group of species that form the American subgenus Corynorhinus 
seem to represent the stem of the evolutionary line that has developed 
Barbastella, Euderma, and Plecotus. The present center of differentia- 
tion of this division appears to be in the arid southwestern portion of 
the United States. This area is presently inhabited by Plecotus town- 
sendit, a form that is actively differentiating today. It is the most 
abundant and widespread member of the subgenus, there is much 
regional variation within its large geographic range, and its range is 
fringed with derivatives from it or from its immediate ancestors. 

The evolution and present distribution of the Recent species of the 
subgenus Corynorhinus can possibly be related to glacial disturbances 
during the Pleistocene (fig. 26). The picture presented in this figure 
reduces the problem to its barest essentials and is almost certainly 
oversimplified. Forms that occurred earlier in the Pleistocene 
(Nebraskan, Aftonian, and Kansan ages) cannot be accounted for, 
and it is very likely that other, now extinct, species existed within 
the time represented in this diagram. For example, Plecotus tetra- 
lophodon lived perhaps as recently as late Wisconsin glacial age. It 
was closely related to P. townsendii but probably was not ancestral 
to it. P. alleganiensis probably lived as recently as the Sangamon 
interglacial age. It was closely related to P. townsendi and P. 
rafinesquit and may have been a direct ancestor of either of these 
Recent species. 

The Pleistocene epoch was characterized by an alternation of 
periods of warmth and cold. During the warm interglacial ages, 
forms such as the species of Plecotus conceivably could have been 
distributed across the continent from coast to coast, provided trees 
or caves were present. During glacial times distribution patterns 
were shifted southward, contracted, and may have been split into 
isolated segments as icecaps moved southward toward the Gulf of 
Mexico and through the Rocky Mountain range. ‘The most southerly 
advance of the ice brought it within 500 miles of the present Gulf 
Coast. Adjacent to the ice front was a belt of treeless tundra of 
unknown width. Faunas and floras were compressed southward 


216 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


——— 
—_—- 
—\_—_- 
——— 


il 


I 


P _—__—_—_—_—_— 


Figure 27.—Relationship of the present distribution of the subgenus Corynorhinus and the 
maximum southward extension of Wisconsin glaciation. Plecotus townsendti, horizontal 


shading; P. mexicanus and P. rafinesquit, vertical shading; southern limit of glaciers, 
heavy line. 


before the advance of the ice. Probably temperate climate inhabit- 
ants found conditions least severe in the southwest and in the extreme 
southeast, in the region of the Carolinas and Florida which was pro- 
tected by the Appalachian Highlands from the winds that blew off 
the icecaps during some phases of the glacial cycle. The Gulf Coastal 
pathway south of the ice may have been less hospitable for these forms 
inasmuch as it was unprotected from the ice chilled winds and had 
rivers of meltwater flowing across it. Furthermore, competition for 
its limited niches may have been acute. 

Under these conditions, a population of Plecotus that may have 
had a transcontinental distribution in the Yarmouth interglacial age, 
might have been isolated into southeastern and southwestern seg- 
ments during the ensuing Illinoian glacial age. The southwestern 


segment included the mountains of northern and central Mexico in 
its range. 


AMERICAN BATS—HANDLEY 217 


With the retreat of the Illinoian icecap and the warming during 
Sangamon time, Plecotus could have expanded its range again from 
coast to coast and perhaps northward into Canada and Alaska if 
caves were available. By this time the southwestern and south- 
eastern segments had differentiated to the species level and remained 
genetically distinct when their expanding ranges came into contact. 
In the meantime, northward advance of warm climate floras and 
faunas isolated the Plecotus of the Mexican mountains from the 
remainder of the southwestern segment. 

Southward advance of the Wisconsin icecaps and cooling climates 
once again contracted the range of Plecotus and broke its wide, con- 
tinuous distribution into several isolated segments, remnants of which 
now occur in the southeastern lowlands, in the Appalachians, in the 
Ozarks, and in southwestern North America. Contact was reestab- 
lished between the southwestern segment and the Mexican segment 
which had by this time diverged to the species level. 

Withdrawal of the Wisconsin icesheets and amelioration of climate, 
still proceeding, make it possible once more for the ranges of Plecotus 
to expand. Plotting the present distribution of American Plecotus 
on a map (fig. 27) showing the maximum southward extension of 
Wisconsin glaciation lends emphasis to the probability that range 
expansion has progressed slowly. The present distribution of Plecotus 
probably bears considerable resemblance to its distribution in the 
Wisconsin age. It is significant that, in the several thousand years 
since the last glacial retreat, bats of several genera have moved north- 
ward, far into the glaciated area, but Plecotus has barely penetrated 
it. No sharp biotic division marks the drift border, and there is no 
lack of caves north of the present range of Piecotus (caves in Wisconsin, 
Pennsylvania, New England, and Ontario, for example) to prevent 
its northward spread. To assume a relatively slow rate of dispersal 
for Plecotus could explain its present pattern of distribution, and 
furnish an important mechanism for speciation. 


110 


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AMERICAN BATS—HANDLEY 239 


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240 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Harpy, R. 

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Harper, F. 

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AMERICAN BATS—HANDLEY 241 


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244 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


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U. $. GOVERNMENT PRINTING OFFICE: 1959 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 110 Washington : 1959 No. 3418 


A REVISION OF THE APION SUBGENUS TRICHAPION 
WAGNER IN THE NEW WORLD (COLEOPTERA : CURCU- 
LIONIDAE) 


By Davin G. KissiIncER 


This paper ' is one of a series in which the species of Apion occurring 
in the New World will be revised. Acknowledgments, explanation of 
measurements, and explanation of abbreviations used to designate 
deposition of specimens are given in the first paper in the series 
(Kissinger, 1957a). 

Ninety-two species are treated; of these 17 are new, 11 were not 
seen by the author, and four are known to the author only from the 
female. Of the 11 species not seen, eight were originally included in 
the subgenus by Wagner, the other three species, Jaminatum Sharp, 
peninsulare Fall, and tabogense Sharp, were adequately described or 
figured so that it has been possible to place them in this subgenus. 

A key to the males of the New World species of subgenus Trichapion 
is presented. The species either unknown to the author or repre- 
sented by females only are not included in the key. The species 
treated are arranged in groups as shown in table 1. The structural 
characters of these groups are summarized in tables 2 and 3 and are 
discussed after the definition of the subgenus. 

1 Based on a thesis submitted in 1957 to the Graduate School of the University of Maryland in partial 


fulfillment of the requirements for the degree of Doctor of Philosophy. The research was aided by a grant 
from the Sigma Xi-RESA Research Fund. 


247 


248 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Because the majority of the material seen was from Canada, the 
United States, and Mexico, the treatment of the species occurring in 
these regions is more complete than that for Central America and 
South America. In many cases only one or a few determined speci- 
mens of a species from the latter regions were seen. No doubt more 
described species of South American Apion will be found to belong to 
this subgenus when determined material is available for study. 

Trichapion Wagner (1912) was proposed as a subgenus for 16 new 
species described from Mexico and Central America. No type species 
was designated. I hereby designate Apion (Trichapion) aurichaleeum 
Wagner as the type species of subgenus Trichapion Wagner. This 
species is selected because it is relatively abundant. In addition I 
was able to study a pair of specimens determined by Wagner and 
I am informed by Mr. J. Balfour-Browne that the types of this species 
are in good order. 

No previously described species were included by Wagner in the 
subgenus. With the exception of Voss (1955), whose paper is dis- 
cussed below, the name 7richapion has not been used in the literature 
since it was first proposed. At the present time 92 of the approxi- 
mately 425 New World species of Apion are known to belong to this 
subgenus. Fall (1898) divided the North American Apion into four 
groups. His group No. 3 corresponds quite closely to subgenus 
Trichapion Wagner. 

Voss (1955) uses Trichapion Wagner as a subgenus for 18 species 
of Apion from Ruanda-Urundi, Belgian Congo. Because Voss does 
not mention the chief diagnostic character of the subgenus—the mu- 
cronate tibiae of the males—in the descriptions of the species he in- 
cluded in subgenus Trichapion, it may be that the African species are 
incorrectly assigned. 

At the present time it is difficult to define a subgenus of Apion. 
The chief reason is the large number of species not assigned to sub- 
genera. In the New World six subgenera have been erected, viz., 
Bothryopteron Wagner with four species, Coelocephalapion Wagner 
with six species, Coleopterapion Wagner with five species, Heterapion 
Sharp with two species, Stenapion Wagner with 18 species, and Tricha- 
pion Wagner with 92 species. In this region 127 out of about 425 
species of Apion have been placed in subgenera. Previous to the 
present study no species of Apion from the United States were placed 
in a subgenus. The majority of Apion subgeneric names have been 
founded on European species, that is, 30 out of 42 subgenera founded 
on approximately 250 species of Apion from Europe. From a world- 
wide standpoint well over a thousand species of Apion have not been 
assigned to subgenera. 


SUBGENUS TRICHAPION—KISSINGER 249 


The mucronate tibiae of the male are the chief diagnostic character 
of the subgenus Trichapion. The middle and hind tibiae of the males 
of most species are mucronate; a few have all of the tibiae mucronate, 
while others have only the middle tibia mucronate. In the type 
species, A. aurichalceum Wagner, and closely allied species the front 
tibiae bear a flat, polished, striate area on the inner anterior surface. 
Other male secondary sexual modifications of the legs are swollen 
femora, spined tarsi, or enlarged tarsal segments. 

The males of a few other species groups of Apion not belonging to 
the subgenus Trichapion have mucronate tibiae but lack one or more 
of the other characters that will be listed in the definition. The 
A. corale group differs in that the front coxae of the male are tuber- 
culate. Thespecies of Fall’s Group I differ, in addition to many other 
characters, by the smooth, polished area limited by a raised ridge on 
the apical ventral surface of the front femur of the male, the nearly 
simple claws, and the metasternal spicules. Apion reclusum Fall 
and A. acrophilum Fall also differ by their simple claws. The male 
of A. opacicolle Smith differs by having metasternal spicules similar 
to Fall’s Group I. A. tenuirostrum Smith, A. smitht Wagner, A. 
impecum Fall, A. metallicum Gerstaecker, and A. troglodytes Manner- 
heim differ in that the prothorax is widest before the base, the sides 
not expanded laterally at the base. 

The definition of the subgenus Trichapion Wagner is as follows: 

Legs, coxae, and beak biack (evustum, rufipenne, subrufum, 
subtinctum, and vinosum with rufescent legs); body not gibbose in 
outline, without transverse pattern of scales, not clothed with fine, 
suberect pubescence; beak curved, apex more slender than base, at 
most expanded laterally at tip, apical region more finely sculptured 
than basal region; anterior portion of the dorsal margin of the antennal 
scrobe may or may not be oblique; frons generally wider than dorsal 
top of beak (a few species with frons equal to or narrower than dorsal 
tip of beak especially of male), not depressed adjacent to eye, with a 
rather broad median area which may be flat, concave, or with an 
obvious, deep, longitudinal sulcus; prothorax expanded laterally at 
base, constricted apically (perforicolle not constricted), apex obviously 
narrower than base; middle coxae separated by a complete lamina; 
first segment of fore tarsus generally longer than wide (with exception 
of rostrum); third tarsal segment strongly bilobed; claws toothed. 

Although some intermediate forms occur it is convenient to consider 
the species of Trichapion as falling into two sections, one related to 
the holarctic Apion simile Kirby and the other to such species as 
Apion patruele Smith, A. reconditum Gyllenhal, and A. rostrum Say. 
The first section related to A. simile is characterized by having two 


250 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


TABLE 1.—Species groups of subgenus Trichapion Wagner 


I. Species of uncertain position 


1. aequabile Fall 
2. aestimabile Wagner 

3. albidulum Fall 

4. alticola Wagner 

5. asellum Wagner 

6. auronitidum Wagner 
7. bettyae, new species 
8. chalybaeum Wagner 
9. grossulum Fall 

10. guatemalense Wagner 
11. laminatum Sharp 

12. peninsulare Fall 

13. persuleatum Wagner 
14. spiculiferum Wagner 
15. tabogense Sharp 


II. Species groups related to Apion 
simile Kirby. 


A. 


SIMILE GROUP 
1. meorrhynchum Philippi 
2. modestum Smith 

3. propinquicorne Fall 

4. simile Kirby 

PROCLIVE GROUP 
acanonicum, new species 
adaetum, new species 
coryi, new species 
imitator Wagner 
plectrocolum, new species 
proclive LeConte 
GLYPHICUM GROUP 

1. chuparosae Fall 

2. glyphicum Sharp 

3. vinosum Sharp 

GRISEUM GROUP 
aurichalceum Wagner 
godmani Wagner 
griseum Smith 
nebraskense Fall 
novellum Fall 

oriotes, new species 
parcum, new species 
sayi Gyllenhal 
SPINITARSE GROUP 
brachycephalum Wagner 
caenum, new species 
calcaratipes Sharp 
enoplus, new species 
innocens, new species 
innocuum, new species 
latitator, new species 


Cah ha tere ast 


Co RSS Loe SI 


Se ees 


8. mirandum, new species 
9. mirificum, new species 
10. sancti-felicis Sharp 

11. spinitarse Wagner 
PUNCTULIROSTRE GROUP 

1. brunnicornis Fall 

2. eccentricum Fall 

3. punctulirostre Sharp 
SUBMETALLICUM GROUP 

1. auriferum Wagner 

2. hadromerum Wagner 
3. perpilosum Wagner 

4, submetallicum Boheman 


III. Species groups related to Apion 


patruele Smith 
PATRUELE GROUP 
abdominale Smith 
centrale Fall 
evustum, new species 
fusconitidum Wagner 
gulare Fall 
nitidum Kirsch 
patruele Smith 
perforicolle Fall 
porcatum Boheman 
10. rufipenne Gyllenhal 
RECONDITUM GROUP 
acupunctatum Sharp 
bicolor Gerstaecker 
davidis Sharp 
gracilirostre Sharp 
importunum [all 
managuense Wagner 
minor Smith 
nanulum, new species 
nitidirostre Sharp 
10. oscillator Sharp 
11. pervicax Fall 
12. reconditum Gyllenhal 
13. subrufum Sharp 
14. subsequens, new species 
15. subtinctum Fall 
OBLITUM GROUP 
1. mediocre Sharp 
2. oblitum Smith 
NIGRUM GROUP 
cordatum Smith 
dolosum Fall 
heterogeneum Sharp 
lativentre Béguin-Billecocq 
nigrum Herbst 


CHINA WHE 


See Oa ras Cte ie 12 ROT 


BAL Goa eed oe 


SUBGENUS TRICHAPION—KISSINGER 251 


TABLE 1.—Species groups of subgenus Trichapion Wagner—Continued 


III. Species groups related to Apion 3. confertum Smith 
patruele Smith—Con. 4. furtivum Fall 
EK. ROSTRUM GROUP 5. mexicanum Wagner 
1. coloradense Fall 6. rostrum Say 


2. commodum Fall 


rows of scales on the elytral intervals, the beak of the male in dorsal 
view rather evenly attenuate slightly distad of the antennal insertion 
to the apical third, and the beak of the male noticeably pubescent in 
the basal two-thirds. The other section related to A. patruele is 
characterized by having a single row of generally inconspicuous 
scales on the elytral intervals, the beak of the male in dorsal view 
nearly parallel-sided in the apical half, and the beak of the male 
nearly glabrous distad of the insertion of the antennae. The uniserial 
or biserial pubescence is subject to some variation but the pattern 
in general falls into one of the two groups. The dorsal pubescence 
of the uniserial section is generally sparse and more inconspicuous 
than that of the other section. 

The more notable intermediate forms are among the species whose 
males have all three tibiae mucronate. Apion furtivum Fall and A. 
confertum Smith have uniserially pubescent elytral intervals but the 
beak is attenuate to the apical third. Apion commodum Fall has 
biserially or triserially pubescent intervals and the beak is attenuate. 
These three species were considered by Fall (1898) to be allies of A. 
rostrum Say, @ species with one row of inconspicuous scales on the 
elytral intervals, beak attenuate to apical third, and the male with 
two pairs of tibiae mucronate. The A. rostrum group may link the 
A. patruele section to the A. simile section through the A. proclive 
group, a member of which also has all three tibiae mucronate. 

The key to species is based entirely upon the male. Generally the 
species of subgenus Trichapion are captured in a series and males are 
usually present. In most cases it is possible to match males and 
females of a species without too much difficulty by comparing the 
sculpture and vestiture of the dorsal surface. 

The key is not based on the species groups as outlined in tables 
2 and 3 because this would be too indefinite since these groups are based 
on rather tenuous differences of degree rather than upon concrete 
structural characters. The general plan of the key is as follows. 
First the species with one or three pairs of mucronate tibiae are split 
off. Then the A. griseum group with its modified front tibiae is 
eliminated. The position of the antennal insertion is then utilized to 


252 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


remove those species with the antennae inserted rather far from the 
base of the beak. The remaining species are split into two groups 
based on uniserially or biserially pubescent elytral intervals. 

Generally the beak and secondary sexual characters of the male 
are the best diagnostic specific characters. Other characters include 
the proportions of the prothorax and elytra, the sculpturing of these 
parts, and the flatness or convexity of the elytral intervals. 

The following explains metheds not included in Kissinger (1957b). 
In giving the measurements of the scutellum the length is always listed 
first; the same applies to measurements of the antennal club. In the 
description of the prothorax the approximate diameter of the punctures 
is given. This may vary somewhat on the same specimen. If the 
variance is great the range of variation is approximated; in general it 
is small so that the measurement given represents an average. To 
indicate approximate robustness of the legs the relation of width to 
length of the front femur is indicated. If the legs are slender the 
number of times the front femur is longer than wide will be about 
four, if the legs are stout it will be about three. The male of A. brun- 
nicornis Fall is an example of very robust legs, in which case the middle 
femur is only 2.25 times as long as wide. 

As an aid to identification, in the key are included the states or 
provinces of Canada, the United States, and Mexico from which the 
species are known. In the case of extensive distribution an attempt 
was made to indicate the extremes of the range. 


Key to males of New World species of subgenus Trichapion 


1. Integument nearly concealed by dense white scales; elytral intervals with 
three or four rows of scales; antennae inserted at basal fourth of beak 
(Southern California, Arivonay ake . . . . albidulum Fall (p. 260) 

Integument not concealed by eect dorsal surface of prothorax and 
elytra with fine pubescence; elytral eae generally with one or two rows 


of fine scales or antennae inserted near middle of beak. . ..... 2 

2. Three pairs of tibiae mucronate, anterior pair sometimes minutely so; pro- 
thorax somewhat wider at middle than base ........... 3 

Not more than two pairs of tibiae mucronate. . .. . fie Fees 9 

3. Dorsal margin of antennal scrobe oblique, angulate Bbes antennal inser- 
ION” sare pet Pe x hone 4 


Dorsal margin of anienial aCrobe ev venly deseondine to belo eve 3- oh x 5 

4. Mucrones denticulate; second elytral interval with two or more rows of fine 

scales; in dorsal ios beak not strongly expanded at antennal insertion and 
slightly attenuate apically (Montana, Manitoba). 

commodum Fall (p. 380) 

Mucrones simple; second elytral interval with one row of fine scales; in dorsal 

view beak strongly expanded at antennal insertion, strongly attenuate to 

apex (Florida) ... . . . . confertum Smith (p. 381) 

5. Beak as long as head and protons pombe deflexed at apical fourth; 

mucrones 2 and 3 dentate (San Luis Potosf) . coryi, new species (p. 285) 

Beak shorter than head and prothorax combined .... . «ae Seen 6 


10. 


12. 


13. 


14. 


15. 


‘16. 


1%. 


18. 


SUBGENUS TRICHAPION—KISSINGER 253 


. Beak in lateral view noticeably narrowed immediately beyond antennal 


insertion, apical half subcylindrical, polished, and subimpunctate . . 7 
Beak in lateral view not abruptly attenuate, eaib ae to near apex; tip 
only shining ... Aas ers 8 


Prothorax deeply Demonic pete 2 An 3 inh ehoet mucrones; first segment 
of tarsus 1 shorter than segments 2 and 3 combined (Illinois, Nebraska, 
Colorado, Kansas, Texas, Arizona) ... . . . oblitum Smith (p. 368) 

Prothorax shallowly punctured; tibiae 2 and 3 armed with long mucrones; 
first segment of tarsus 1 about as long as 2 and 3 combined (Guerrero, 
Puebla). cst os . . . mediocre Sharp (p. 366) 


. Antennae inserted a Bacal fourth of jive oh} distance from eye equal to width 


of frons; elytra at base one-third wider than prothorax at base (Washington 
to Southern California). .. . . . . cordatum Smith (p. 369) 
Antennae inserted at basal third of heals at distance from eye one-half greater 
than width of frons; elytra at base one-half wider than prothorax at base 


(Georgia, Arkansas). 20. =. + < = « ss «,+, . furtivum Fall i 382) 

7 One pair of-tibiae-mucromate. . as esc se ws 10 
Two pairs of tibiae mucronate. . .. . ce ot 
First segment of tarsus 1 with outer apical atiple produped into'e a epi. ee | 
First segment of tarsus 1 not so modified. . .... malay >) 

. Beak in dorsal view attenuate from antennal insertion to epicel finde Feo 


Beak in dorsal view nearly parallel-sided from antennal insertion to apex 
(Mexico). . . . . . . . latitator, new species (p. 317) 
Antennae yellowish: inserted a distance from eye slightly less than width of 
frons; dorsal surface of beak clothed with scales coarser, shorter, and whiter 
than pies on dorsal surface of prothorax, intervals in part with two rows 
of punctures (San Luis Potosf and Puebla). innocens, new species (p. 314) 
Antennae dark, inserted at distance from eye slightly greater than width of 
frons; dorsal surface of beak clothed with scales similar to those on dorsal 
surface of prothorax; intervals with one row of punctures (Guatemala, 


British Honduras). . . ... . . brachycephalum Wagner (p. 310) 
First segment of tarsus 1 elonzate. four times as long as wide (Michoacd4n, 
Ruebla) a ee . . . mirandum, new species (p. 318) 
First segment phe ae ploneata: 6k’ nc MFS igsle BPs al. ceetspass hiethctastga, A 


Tarsus 1 with first segment with inner margin strongly rounded from base to 
apex, forming a broad, rounded lobe projecting distally beyond apex of 
segment, from above inner pers of segment much broader than outer 
half (Veracruz)... . . . mirificum, new species (p. 319) 

Tarsus | with first eeement. not so iredsaed. Sh sy We cee ee ie mead epee 

Front femur distinctly swollen, antennae inserted at basal fourth of beak, at 
distance from eye equal to width of frons (Michoac4én). 

caenum, new species (p. 312) 


Hrontwemurmot swolleni=.cepee es oo us Gee) eee ea Gees css ee HG 
First segment of tarsus 3 with inner angle produced into a short spine (Vera- 

cruz, Panama) .......... .. . sancti-felicis Sharp (p. 320) 
PP ATSUS{O SIMD lCysyci iss) ha las ial Je ge, ve eva ame iat cae et es, ees 
Antennae inserted at not more than basal third of he aoe : iene 8 
Antennae inserted just behind middle of beak, yellowish; eal strongly 

deflexed at middle (Arizona)... . . . . brunnicornis Fall (p. 3238) 


Beak shorter than prothorax, stout; size greater than 3.5 mm. (Puebla). 
bettyae, new species (p. 263) 
Beak longer than prothorax, slender; sizelessthan2.8mm....... 19 


254 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


19. 


20. 


21. 


22. 


23. 


24. 


25. 


26. 


27. 


28. 


29. 


30. 


Antennae inserted at basal fifth of beak, at distance from eye equal to width 


of frons (San Luis Potos{, Morelos). . . innocuum, new species (p. 315) 
Antennae inserted at basal third of beak, at distance from eye twice width 
of frons ...°. 20 


Beak in lateral view attenuate Age cence mention to oer (Arizona). 
eccentricum Fall (p. 324) 
Beak in lateral view nearly parallel beyond antennal insertion (Puebla, 
México, Morelos; Guatemala) . . . . punctulirostre Sharp (p. 325) 
Fifth ventral segment with lateral apical angles produced into a pair of trian- 
gular processes projecting posteriorly slightly beyond tips of elytra (Tres 
Marfas Island, Mexico) . . . . . . acanenicum, new species (p. 282) 
Fifth ventral segment not so medinedl: Sefer ay oe ; F a 77° 
Front tibia more or less flattened on inner surface, this area Fee of scales, 
generally polished and noticeably striate; ne generally inserted at 


distance from eye less than width of frons. . ........... 32 
Front:tibia not so modified... 4. Ye sae eth ed ice ea 
First segment of tarsus 1 with outer apical ek produced intoaspine. . 24 
First segment of tarsus 1 simple. . .. . Sige duh fd sb tayf ev col has eles ee 


Size about 2.0 mm.; elytral intervals with two rows of scales which are coarse 
and pearly in solon on basal margin of elytra and along basal third of suture; 
laterally and posteriorly coarse scales are bounded by a region of very fine, 
yellowish scales; elytra in posterior half and area lateral of interval 5 
clothed with white, hairlike scales; tibiae yellowish (Guatemala). 

calcaratipes Sharp (p. 312) 

Size less than 1.5 mm., elytral intervals with one row of uniform, fine scales; 
tibiae nearly: blackat.ius 2 RU a) Be ats, 02S es 

Spine on outer angle of first segment of tarsus 1 large, triangular in shape, as 
long as tarsal segment is wide at apex (Brazil, Nicaragua). 

spinitarse Wagner (p. 321) 

Spine on outer angle of first segment of tarsus 1 small, shorter than width of 
tarsal segment (Veracruz, Puebla). . . . emoplus, new species (p. 314) 

Tibia 1 with a short, flat area less than one-third its length, not or very 
slightly striate (New York-Florida-Texas-W yoming). 

sayi Gyllenhal (p. 306) 

Tibia 1 with flat or concave area extending about one-half its length. . 27 

Flat area on tibia 1 extremely densely and finely striate so that the area 


appears irridescent, individual striae not discernible. ...... 28 
Flat area on tibia 1 with coarser, distinct, well separated striae, not irri- 
descents US tate vie wy Hs ee Omi teem Ee ches Ashe, US mae 


Tibial mucrones projecting in line with feild axis of tibiae; beak strongly 
attenuate toward apex (Nebraska, Colorado, Oklahoma). 
nebraskense Fall (p. 301). 
Tibial mucrones projecting at an angle with long axis of tibiae; beak slightly 
attenuate toward apex (District of Columbia, Tennessee). 
novellum Fall (p. 303) 
Tibia 1 straight in anterior view... . 30 
Tibia 1 deflexed at apical two-fifths in arteien view (Mexico: Guatemala 
godmani Wagner (p. 297) 
Beak in dorsal view distinctly expanded at apex (Guatemala; Puebla, México, 
Distrito Federal)... .. . . .aurichalceum Wagner (p. 296) 
Beak in dorsal view cylindrical a peer Ie eae 2 6 US ee eT 


31. 


32. 


33. 


34, 


35. 


36. 


37. 


38. 


39. 


40. 


41. 


42. 


SUBGENUS TRICHAPION—KISSINGER 255 


Size about 1.6 mm.; metasternum comparatively coarsely, sparsely punc- 
tured; tibial mucrones fine, curved (Pueblo, México, Distrito Federal). 
parcum, new species (p. 305) 
Size about 2.0 mm.; metasternum comparatively finely, densely punctured; 
tibial mucrones Bante eras shanliet = Oe 
Elytra not aeneous, pubescence Bonny ee Antennal ep pearly three times 
as long as wide, covered with very sparse, long, suberect setae and shorter, 
more abundant, decumbent setae (Arizona, New Mexico). 
oriotes, new species (p. 304) 
Elytra moderately aeneous, pubescence on dorsal surface with faint yellowish 
cast, white laterally; antennal club slightly more than twice as long as 
wide, covered with nearly uniform, long, suberect setae (New York- 


Florida, Alabama). ..... . . .griseum Smith (p. 300) 
Antennae inserted at distance han ee eee than width of frons; dorsal 
pubescence squamiform (Texas, México) . . propinquicorne Fall (p. 277) 
Antennae inserted at distance from eye not less than width of frons. . 34 
Prothorax much wider at middle than base, sides greatly rounded. . . 35 
Prothorax at middle at most ime wider than ee sides not greatly 
TOUNGEG! 57 ouis Henne Saher ade sae! VO 


Elytral intervals SreouehaT convex; tibia 1 dilated: curved, bearing long, 
yellowish pubescence on inner surface (Puebla, Distrito Federal, Micho- 
acén, Guerrero). . . . . . heterogeneum Sharp (p. 372) 

Elytral intervals Slichtlyic convex; TonRe 1 dilated (New Hampshire—Georgia, 
Louisiana, Iowa, Wisconsin) . . . .nigrum Herbst (p. 376) 

Antennae inserted at about the ails of tis beak, scape longer than next 
five antennal segments (Chile) . . .meorrhynchum Philippi (p. 275) 

Antennae inserted distinctly behind middle of: beak’... i.) PA ae FO 

Antennae inserted at distance from eye equal to or Slightly renee than 
widthoffrons. .. . ie Taaaywi os 

Antennae inserted at dictanes from eye ‘distinctly Grentar nena or more) 
than width of frons. ... SF fice OL 

Pubescence of dorsal surface penerallsy pont icistral paieeale with one row 
of fine scales... . a aCe ae, 

Pubescence of dorsal pte Conan icnone! eivinel iteavale with two rows of 
fiNGu BOALCRU corals cualer toques eaten te berieriae bit OME od CE CNN Uae 

Mucrones dentate... . NORGE cee anvil on cbarhions 140 

Mucrones simple, at most pibenecinte Seta Boca ee es sepettcaase (40 

Frons about as wide as dorsal tip of beak, appearing fagaloete: prothorax 
basal margin strongly expanded (?Arizona) . . . grossulum Fall (p. 269) 

Frons wider than dorsal tip of beak (or elytra red), not trisuleate; prothorax 
basal margin at most) slightly expandeds:..5.....:) ....0. #2... Al 

EB lytEa red sw OrotlOrax DIACK acs qo. so -e18 1 eo ch pay ee tysy. formate Gdiieeebe legs Ee 

HKlytra and prothorax black. ... . Mine eet 4a 

Beak as long as head and prothorax pombined= Pinheras anely punctured, 
nearly glabrous; dorsal surface of elytra not at all striate; prothorax not 
constricted apically (Veracruz, Tabasco, Guerrero; British Honduras; 
Guatemala; Nicaragua; Panama; Colombia; Venezuela; Brazil). 

rufipenne Gyllenhal (p. 346) 

Beak shorter than head and prothorax combined; prothorax with rather 
coarse, deep punctures which bear hairlike setae; striae adjacent to suture 
distinctly impressed throughout; prothorax distinctly constricted at apex 
(Tamaulipas, San Luis Potosf). . . . . . evustum, new species (p. 337) 


256 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


43. 


44. 


45. 


46. 


47. 


48. 


49. 


50. 


52. 


53. 


or 
i 


Abdomen and metasternum finely, sparsely punctate. ....... 44 
Abdomen and metasternum coarsely, closely, deeply punctate. . ... 45 
In dorsal view beak not expanded over antennal insertion; beak distinctly 


shorter than head and prothorax combined (New England—Florida—Texas— 
Wisconsin) ... . . . patruele Smith (p. 341) 
In dorsal view beak cintinety eeaenied over antennal insertion; beak about 
as long as head and prothorax combined (Peru; Amazonas). 
nitidum Kirsch (p. 341) 
Mucrones moderately long, slender; elytral intervals not much wider than 
striae (California, Arizona). . .. .. . . abdominale Smith (p. 333) 
Mucrones short, blunt; elytral intervals nearly twice as wide as striae (Colo- 
rado, Idaho, Montana; British Columbia, Alberta, Hudson Bay Territory). 
centrale Fall (p. 335) 
Beak about as long as prothorax (Illinois, Nebraska, Colorado, Arizona). 
modestum Smith (p. 276) 
Beak distinctly longer than prothorax. .. . Faerie Ch 
Prothorax slightly wider at middle than base; civtsal iter ae eironale convex, 
slightly wider than striae (Massachusetts—Tennessee—Texas—Iowa). 
porcatum Boheman (p. 344) 
Prothorax at middle not wider than base; elytral intervals at most slightly 


convex, at least one-half wider than striae. .. . pieces), eis) 
Elytra at humeri at least two-fifths wider than Sratnorae a base; elytra 
more than two and three-fourths times as long as prothorax. . .. 49 
Elytra at humeri one-third wider than prothorax at base; elytra less than two 
and three-fourths times as long as prothorax. . . . Bares) te, | OH) 
In lateral view dorsal surface of head above eyes distinetly declivitous 
(Tamaulipas). . . ; . . . nanulum, new species (p. 357) 
In lateral view dorsal purines of endl is flat from eyesto base. .... 50 


Elytral intervals convex (Tabasco; British Honduras; Guatemala). 
oscillator Sharp (p. 359) 
Elytral intervals flat. . ... ee ROL 


. Pubescence on side of aaiLOrae aii as coarse as a that c on ‘dorsal surface or 


midline wan eae pao ha 52 
Pubescence on side lof prathoras BRAG twice as coarse as that on dorsal sur- 
face; in dorsal view beak strongly expanded over antennal insertion (Ta- 
pacco: Guatemala; Nicaragua). . . . .. mamaguense Wagner (p. 354) 
Prothorax with uniform, sparse pubescence on dorsal surface . .... 53 
Prothorax with scales along median line coarser than those on lateral portion 
of dorsal surface... . se Oe 
Prothorax sparsely, finely, shallowly, manotured: dows ay aad daterater eyes 
acutely prominent; scales in elytral striae minute, not larger than those 
on intervals (Tamaulipas). . . . . . subsequens, new species (p. 364) 
Prothorax moderately densely, finely, deeply punctured; eyes more rounded; 
scales in elytral striae distinctly larger than those on intervals (Ontario; 
New York, Virginia, Tennessee, Texas, Kansas, Iowa, Michigan). 
reconditum Gyllenhal (p. 361) 
Beak slightly shorter than head and prothorax combined; prothorax more 
deeply, uniformly punctured (Texas, Tamaulipas). 
subtinctum Fall (p. 365) 
Beak slightly longer than head and prothorax combined; prothorax more 
shallowly, irregularly punctured (Guatemala). subrufum Sharp (p. 363) 


55. 


56. 


57. 


58. 


60. 


61. 


62. 


64. 


SUBGENUS TRICHAPION—KISSINGER 257 


Mucrones minute; dorsal margin of antennal scrobe evenly descending to 


below eye (Florida)... .. . « .pervicax Fall (p. 360) 
Mucrones larger; dorsal margin of lantennan sorebe somewhat angulate above 
antennal insertion .. . {ieyia., 56 


Mucrones short, subangulate (New. Teer Florida, Mieseeiap? Indiana). 
perforicolJe Fall (p. 343) 
Mucrones long, simple (Georgia, Florida, Mississippi). 
importunum Fail (p. 353) 
Pubescence noticeably denser at base of elytral interval 3; size generally less 


thanl.6@mm.... Anchen meen earn n eae Uha te: 
Pubescence on elytra nado size Core ant le a MMe Psa es eS 4D 
Mucrones projecting at an angle with tibiae; elytral intervals nearly flat 
(Guerrero; Guatemala) .... . . . perpilosum Wagner (p. 329) 


Mucrones in basal half projecting in ane with long axis of tibiae; apical half 
curved out laterally; elytral intervals moderately convex (@abaees: Guate- 
mala; Panama). . . . . . auriferum Wagner (p. 327) 

Frovs wider than dorsal ip on Bente nor sulcate; prothorax at base one-fifth 
wider than long; elytral intervals moderately convex; size 1.6 to 2.0 mm. 
(Arizona; Baja California, Chihuahua, Guerrero). aequabile Fall (p. 259) 

Frons as wide as dorsal tip of beak, with shallow median depression; pro- 
thorax at base one-third wider than ss elytral intervals nearly flat; size 


greater than 2.1 mm... . SH eeart 260 
Middle and hind tibiae with siunjlar’ mucrones, ‘both soutely, angulate ven- 
trally (Guerrero; Guatemala). . . . . . . . imitator Wagner (p. 286) 


Middle tibia with a short, acutely angulate mucro, hind tibia with long, 
slender, straight mucro projecting at about 120° from tibia (Distrito 


Hederal)s ss 4: . . . . plectrocolum, new species (p. 288) 
Beak as long or longen tana head and prothorax combined. ..... 62 
Beak distinctly shorter than head and prothorax combined. . . . . . 67 


Antennae inserted at distance from eye four times as great as width of frons; 
beak one-fourth longer than head and prothorax; mucro 2 short, subangu- 
late; mucro 3 long, curved; size 2.25 to 2.50 mm. (Puebla, Michoacdn, 
Guerrero; Guatemala; Honduras) . . . . . gracilirostre Sharp (p. 352) 

Antennae inserted at distance from eye not more than twice as great as 
width of frons; beak at most slightly longer than head and prothorax 
Combimnedaar cise Sh 2auew eae COS 


. Size 2.5 to 3.0 mm.; “egal terse ecient as Stone as Eide) not longer than 


lobes of third segment (New Hampshire, Florida, Texas, Wisconsin). 
rostrum Say (p. 384) 
Size generally less than 2.5 mm.; second tarsal segment longer than wide, 
longer than lobes of third segment. . ... . gan SER A 
Head excavated and polished ventrally; elytral airine ‘shallow; mucrones 
minute (Florida; Tamaulipas). . . . . . . gulare Fall (p. 338) 
Head not excavated ventrally; elytral striae dean: mucrones larger. . . 65 


5. Elytra strongly bronzed, intervals polished, nearly impunctate, elytral pubes- 


cence nearly lacking, scales in striae larger than those on intervals (Guate- 
mala; Honduras; Colombia; Venezuela; Bolivia). 

bicolor Gerstaecker (p. 350) 

Elytra black, not bronzed, intervals with more or less distinct transverse 

minute rugae, intervals with distinct punctures bearing conspicuous scales 

as largeras thosedmesttides, . .. . 4 < ).< » # «6 2 2 2 « @ 9 OG 


258 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


66. 


67. 


68. 


69. 


70. 


71. 


72. 


73. 


74. 


75. 


76. 


Apex of prothorax distinctly constricted; beak slightly punctured beyond 


middle (México, Distrito Federal) . . . . . mexicanum Wagner (p. 383) 
Apex of prothorax slightly constricted; beak strongly punctured in rows, tip 
smoother (Illinois, Kansas, Colorado) . . . . .coloradense Fall (p. 378) 
Mucrones simple. . . . hat eR a arr LRT ASU eee 
Mucrones dentate or sarees Seyitaniie ee er ere ee me fC) 
Size 1.5 mm.; dorsal surface of prothorax deeply, closely, coarsely punctured 
(Iowa, ne iceratig Kansas) . . . . . .minor Smith (p. 355) 
Size generally larger than 1.8 mm.; Prioreal aie of prothorax more shallowly 
muncbtired sc. '0 i ee oe a 


Antennae inserted at basal two-fifths of beak; beak in dorsal view nearly 
parallel-sided; prothorax little wider at base than apex (holarctic; North 
Carolina to New England, west to Washington, south to Utah). 

simile Kirby (p. 278) 

Antennae inserted at basal third of beak; in dorsal view beak attenuate 
toward apex; apex of prothorax seven-tenths as wide as base (Guerrero; 
Guatemala). .......... .. .hadromerum Wagner (p. 328) 


Beak shorter and stout, at most slightly longer than prothorax, in lateral 
view attenuating to apex, more or less suleate laterally; frons not wider 
than dorsal tip of beak. . ... oc eneeel fe a ee 

Beak more slender and subparallel apically: Pieuincie fonees nee prothorax, 
not sulcate laterally; frons generally wider than dorsal tip of beak. . 73 


Elytra reddish bronze, legs in part rufous (Veracruz; Guatemala). 
vinosum Sharp (p. 294) 
Elytra black, aeneous; legs black. . . ...... : Mer ALi) P70 


Beak shorter than prothorax; frons slightly narrower fee ee tip of beak 
(Veracruz, México, Morelos, Michoacan, Guerrero). 

glyphicum Sharp (p. 292) 

Beak slightly longer than prothorax; frons equal to dorsal tip of beak (Baja 

California)... .....4.2.+.. =... .. .chuparosae Fall (p. 291) 


Prothorax black, elytra blue; prothorax with noticeable prominence on lateral 
basal region (Argentina; Brazil) . . .lativentre Béguin-Billecocq (p. 374) 
Prothorax and elytra of same color, black; lateral basal region of prothorax 
NOL: Prominent +... oF ek see See : aid Yak: 5, Ao et meme 


Antennae inserted at distance from eye eerie greater than width of frons; 
beak in side view nearly parallel-sided in apical third’ (Washington, Wyo- 
ming, Colorado, Utah, California, Durango, México). 

proclive LeConte (p. 289) 

Antennae inserted at distance from eye more than one-half greater than 
width of frons; beak in side view attenuate to apex ........ 75 

Antennae inserted in front of basal third of beak; elytral pubescence uniform; 
prothorax at middle not narrower than base (New Mexico, Arizona). 

dolosum Fall (p. 370) 

Antennae inserted behind basal third of beak; pubescence forming a more or 
less distinct spot at base of interval 3; prothorax widest at base. . . 76 

Size larger than 2.50 mm.; mucrones dentate apically (Puebla, México, Dis- 
trito Federal, Guerrero). . . . . . .submetallicum Boheman (p. 330) 

Size about 1.75 mm.; mucrones angulate apically (Tabasco). 

adaetum, new species (p. 284) 


SUBGENUS TRICHAPION—KISSINGER 259 


Species of Uncertain Position 
Apion (Trichapion) aequabile Fall 
FicureE 1,),c,g,7. 
Apion aequabile Fall, Trans. Amer. Ent. Soc., vol. 25, p. 148, 1898. 


Description: Length, 1.6 to 2.0mm. Moderately robust. Black; 
pubescence conspicuous, coarse, white, sparse, coarser and denser 
laterally. Male beak one-eighth longer than prothorax; deflexed in 
apical third; in lateral view basal one-third parallel, attenuate through 
middle third, apical third nearly parallel, five-sixths as thick as basal 
region; from dorsal view widest in basal third at antennal insertion 
where it is one-fourth wider than base, attenuate to apex, there three- 
fourths as wide as base; sparsely pubescent in basal two-thirds, 
punctures there coarse, apical one-third with fine punctures, shining. 
Female beak as long as head and prothorax combined, one-half 
longer than prothorax, slightly curved, apical half nearly cylindrical, 
somewhat depressed near tip; strongly punctured laterally in basal 
half, fine scales in two short sulci above antennal insertion, shining 
and glabrous beyond antennal insertion. Antennae inserted at dis- 
tance from eye equal to width of frons, of male in basal fourth of 
beak, of female at basal fifth of beak; first segment slightly shorter 
than next two, second segment a little longer than third, club 0.18 
by 0.06 mm. Eyes moderately prominent; frons wider than dorsal 
tip of beak, with a wide, flat median area and one lateral row of punc- 
tures. Prothorax at base one-fifth wider than long, middle as wide 
as base, apex two-thirds to three-fourths as wide as base; sides with 
moderate lateral expansion at base, slightly diverging to middle, 
rounding to constricted apex; in profile dorsal surface very slightly 
arcuate from base to apex; punctation shallow, sparse, 0.03 mm. in 
diameter, interspaces about one-half as great as diameter of punc- 
tures; basal fovea nearly obsolete, elongate. Elytra at humeri one- 
third wider than prothorax at base, 2.5 times as long as prothorax, 
length to width as 10:7.5; intervals twice as wide as striae, moderately 
convex, with two rows of fine punctures bearing fine scales; striae 
deep. Scutellum narrow, elongate, 0.08 by 0.04 mm. Claws with 
strong, acute basal tooth. Front femora 3.3 times as long as wide. 

Special male characters: Tibiae 2 and 3 armed with subdentate 
mucrones. 

Typss: I hereby designate the lectotype of this species as the male 
specimen (MCZ 25071) in the Fall Collection labeled La Chuparosa. 
Cotypes are in the J. L. LeConte Collection and in the California 
Academy of Sciences. 


260 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


MATERIAL EXAMINED: Lectotype and 30 specimens. 

KNOWN DISTRIBUTION: 

Unrrep Strarss: Arizona: Chiricahua Mts., July 4, 1940, D. E. Hardy (UK); 
Globe (CU); Huachuca Mts., Ramsay Canyon, Cochise Co., July 16, 1948, C. 
and P. Vaurie (AMNH); Santa Rita Mts., Madera Canyon, beating oak (CU); 
Santa Rita Mts. (USNM); Whiteriver, July 4, 1950, D. E. Hardy (UK). 

Mexico: Baja California: La Chuparosa (MCZ, CAS). Chihuahua: Chihua- 
hua, H. F. Wickham (MCZ). Guerrero: Rio Balsas, H. F. Wickham (TLCC). 


Apion (Trichapion) aestimabile Waguer 


Apion (Trichapion) aestimabile Wagner, Arch. Naturg. Berlin, vol. 78, p. 115, 
1912. 

Description: This species was described from a single male in the 
Wagner Collection which was not seen by the author. The following 
notes are taken from the original description. 

Length, 1.8 mm. 

Very near A. mexicanum Wagner, head much smaller and narrower, 
frons flat with two rows of fine, scale-bearing punctures. Male beak 
a little longer than head and prothorax (A. mexicanum beak of male 
equal to head and prothorax combined), distinctly less curved than 
A. mexicanum, stouter, distinctly swollen at antennal insertion, thence 
slightly attenuate. Antennae of male inserted somewhat in front of 
middle of beak. Prothorax at base one fifth wider than long, sides 
nearly parallel to middle, strongly constricted at apex, there one- 
fifth narrower than base; punctation sparser and deeper than A. 
mexicanum; basal fovea continues on to near apex, but deep only at 
base. 

Type Locauity: Veracruz, Mexico. 

Remarks: The insertion of the antennae somewhat in front of the 
middle of the beak will distinguish this species from all known mem- 
bers of the subgenus Trichapion. 


Apion (Trichapion) albidulum Fall 
Fiaure 1,d,f,h. 
Apion albidulum Fall, Journ. New York Ent. Soc., vol. 26, p. 219, 1918. 


Description: Length, 2.0 to 2.2 mm.; width, 1.00 to 1.12 mm. 

Robust. Black; densely clothed with appressed white scales, 
scutellum and beak beyond basal one-fifth glabrous. Male beak 
slightly longer than prothorax, female beak shorter than head 
and prothorax combined, one-fourth longer than prothorax; mod- 
erately evenly curved; dilated at antennal insertion, attenuate to 
middle, cylindrical beyond middle; coarsely punctured in basal one- 
half, sparsely punctured apically. Antennae inserted at basal one- 
fourth, male, at basal one-fifth, female, at distance from eye equal 


SUBGENUS TRICHAPION—KISSINGER 261 


to width of frons; first segment a little shorter than next two, second 
equal to next three combined, club 0.18 by 0.09 mm. Eyes prominent; 
frons moderately wide, densely clothed with white scales on either 
side of bare median sulcus. Prothorax slightly wider at base than 
long, middle as wide as base, apex three-fourths as wide as base; 
sides with slight basal lateral expansion, slightly sinuate before base, 
apex moderately constricted; in profile dorsal surface slightly arcuate; 
punctation moderately deep, 0.03 mm. in diameter, interspaces about 
one-half as great as diameter of punctures; basal fovea deep, elongate, 
a slightly raised longitudinal median carina occurs in apical one-half. 
Elytra at humeri two-fifths wider than prothorax at base, about 2.5 
times as long as prothorax, length to width as 11:8.5; intervals flat, 
more than twice as wide as striae, with three or four rows of punctures 
bearing scales; striae moderately deep, with a single row of scales. 
Scutellum elongate, twice as long as wide, 0.12 by 0.05 mm., with 
deep, median basal foven. Front femora three times as long as wide. 
Claws with a large, acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, straight, denticulate mucrones. 

MATERIAL EXAMINED: Type, male (MCZ 25073), labeled Coachella, 
Calif., in Fall Collection; and 25 specimens. 

KNOWN DISTRIBUTION: 


UnitEep States: California: Riverside Co., Coachella (MCZ); Palm Springs 
(CU, UK, USNM). Arizona: Fort Yuma (USNM). 


Apion (Trichapion) alticola Wagner 
Apion (Trichapion) alticola Wagner, Arch. Naturg. Berlin, vol. 78, p. 107, 1912 


Description: This species was described from a pair of specimens 
in the British Museum (Natural History) not seen by the author, 
The following notes are taken from the original description. 

Length, 2.2 to 2.4 mm. 

Very near A. imitator Wagner, with different beak and legs. Male 
beak equal to head and prothorax combined, stouter than A. imitator, 
more strongly curved, more distinctly widened at antennal insertion, 
more distinctly attenuate toward apex; female beak much stouter 
than in A. imitator, only a little longer than head and prothorax, 
more strongly curved, punctation deeper, denser; at antennal insertion 
beak of female of A. imitator more distinctly swollen than A. altzcola. 
Frons striate, middle stria deeper. Legs, especially femora, stouter, 
tarsus longer as in A. mediocre Sharp. Male mucrones sharper than 
those of A. imitator, somewhat shorter than those of A. mediocre. 
Scutellum a little larger than that of A. mediocre, but distinctly 
smaller than that of A. mitator, deeply grooved, enclosed in a groove. 
Prothorax and elytra the same as A. tmitator. 


262 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


TYPE LOCALITY: Omilteme, Guerrero, Mexico, 8,000 feet (BMNH). 

Remarks: Wagner’s statement that the elytra of A. alticola are 
the same as those of A. imitator is accepted literally. The elytral 
intervals of A. imitator are biserially pubescent. The apparently 
long beak of this species should distinguish it from related forms. 


Apion (Trichapion) asellum Wagner 
Apion (Trichapion) asellum Wagner, Arch. Naturg. Berlin, vol. 78, p. 102, 1912. 


Description: This species was described from three males from 
Managua, Nicaragua, one in the Wagner Collection and two in the 
Solari Collection; this material was not seen by the author. The 
following notes are a translation of the original description. 

Length, 1.7 to 1.9 mm. 

Near preceding species [godmanz], of same body size and general 
habitus, also same color; but has much larger and denser gray-white 
pubescence on head, prothorax, and elytra; and denser pubescence on 
ventral surface and following characters: eye of male distinctly larger; 
frons somewhat smaller, nearly smooth, indistinctly striate; eyes 
beneath with snow white cilia. Beak slightly shorter than head and 
prothorax combined, hardly curved; from above, base to antennal 
insertion with wrinkled sculpture and single punctures, rather dull; 
at antennal insertion slightly wider, from there to apex rather strongly 
attenuate; with rows of punctures on both sides of smooth middle 
line, gradually becoming finer toward apex; more shining and glabrous 
toward apex; from side angulately swollen under the antennal 
insertion, on dorsal line slightly angulate over antennal insertion. 
Antennae inserted at distance equal to long diameter of eye, otherwise 
much like godmani. The prothorax is larger in comparison to the 
elytra, at the bisinuate base only one-sixth wider than long, in front 
with a slightly uniform constriction, the constriction before the hind 
angle only a little stronger than that before the anterior margin and 
rather less than the same in godmani; the punctation is distinctly 
denser but finer and deeper, the interspaces smaller than the punctures; 
the basal stria is rather short but deep. The elytra hardly differ, 
a little wider and from the apex with less sinuation. The scutellum 
somewhat larger, triangular, deeply grooved. The legs—especially 
the femur—are similar in length but somewhat stouter; the first 
tarsal segment distinctly longer and narrower than the second, which is 
as long as broad; the claws are longer, widely spread, strongly toothed. 
The male with the second and third pair of tibiae mucronate. 


SUBGENUS TRICHAPION—KISSINGER 263 


Apion (Trichapion) auronitidum Wagner 


Apion (Trichapion) auronitidum Wagner, Arch. Naturg. Berlin, vol. 78, p. 113, 
1912. 

Description: This species was described from a single female 
from Capetillo, Guatemala, in the British Museum (Natural History) 
that was not seen by the author. The following notes are a transla- 
tion of the original description. 

Length, 1.8 mm. 

Closely related to A. subrufum Sharp, somewhat smaller and more 
compact in form, can be recognized by the following characters: 
beak of female somewhat shorter, at antennal insertion somewhat 
more widened and a little more strongly curved. The head is a 
little wider, the frons with a wide, shallow middle striae and finely, 
sparsely punctured; the prothorax is of the same general form, but 
the disc is nearly impunctate, the sides with large, shallow, distant 
punctures; in front of the scutellum is an indistinct, shallow impres- 
sion. The elytra in comparison with the prothorax is distinctly shorter 
than in subrufum, widest point nearly at middle, with strong humeri; 
striae with finer, shallower punctures, intervals slightly convex, with 
fine hair-bearing punctures. Legs same as subrufum, only the 
tibia blackish, while in subrufum the tibia are rusty red. With 
yellowish pubescence especially on head and prothorax, where it is 
distinctly longer and somewhat denser than subrufum. Head, 
prothorax, and elytra with a bright metallic gold luster. De- 
scribed from a specimen in Sharp’s type series of subrufum, from 
which it differed by the rust red color of the elytra and prothorax with 
its slight metallic luster. 


Apion (Trichapion) bettyae, new species 
Fiaure 1,¢,7,k 


Duscrietion: Length, 3.90 mm.; width, 1.75 mm. 

Moderately robust, depressed. Black; pubescence conspicuous, 
white, long, fine, somewhat coarser and denser on sides of mesothorax 
and metathorax. Male beak stout, shorter than prothorax, nearly 
straight; in lateral view nearly parallel-sided, upper surface abruptly 
deflexed downward toward tip; in dorsal view tapering from antennal 
insertion to apex, not expanded laterally at antennal insertion; 
coarsely, deeply, rather densely punctured, pubescence conspicuous 
to near tip, tip smoother; dorsal margin of antennal scrobe continues 
to apical third as a carina overhanging a series of deep punctures 
which appear to form a sulcus. Female beak equal in length to 


483661—59—_2 


VOL. 110 


ve 
k 


Figure 1.—a, Apion laminatum Sharp, lateral view of head (after Sharp, 1890). , c, g, 1, 
A. aequabile Fall: b, lateral view of head and prothorax of male; c, mucro of tibia 3 of 
male; g, lateral view of head and prothorax of female; i, entire dorsal view of male. d,f, h, 
A. albidulum Fall: d, mucro of tibia 3 of male; f, lateral view of head and prothorax of 
male; h, entire dorsal view of male. ¢, 7, k, A. bettyae, new species: ¢, mucro of tibia 2 
of male; 7, lateral view of head and prothorax of male; k, entire dorsal view of male. 
Line equals 0.50 mm. 


SUBGENUS TRICHAPION—KISSINGER 265 


prothorax, stout, nearly straight; in lateral view ventral surface 
slopes upward slightly from antennal insertion to apical third, dorsal 
surface slopes strongly to apex; in dorsal view attenuate strongly from 
antennal insertion to middle, apical half nearly parallel; sculpture 
similar to male. Antennae inserted near basal third of beak, male, 
at basal fourth of beak, female, at distance from eye one-third greater 
than width of frons; first segment slightly shorter than next four, 
second equal to next two, club 0.24 by 0.12 mm. Eyes moderately 
prominent; frons narrower than dorsal tip of beak, with a fine, very 
deep median sulcus and one lateral row of coarser, more or less sepa- 
rated punctures. Prothorax at base slightly wider than long, middle 
narrower than base, apex two-thirds as wide as base; sides beyond 
basal lateral expansion moderately converging to distad of middle, 
rounded in apical third to constricted apex; in profile dorsal surface 
slightly arcuate toward base, strongly arcuate in apical third; puncta- 
tion deep, 0.06 to 0.08 mm. in diameter, interspaces narrow, sub- 
cariniform, dull, strongly alutaceous; basal fovae linear, deep, ex- 
tending one-third length of prothorax. Elytra at humeri one-third 
wider than prothorax at base, about 2.25 times as long as prothorax, 
length to width as 10:7; intervals about one-half wider than striae, 
concave, with one row of rather large, shallow punctures. Front 
femora 3.7 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 with a short, nearly simple mucro 
projecting in line with long axis of tibia. 

Typns: Holotype male (CAS), Atlixco, Puebla, Mexico, May, A. 
Fenyes Collection. Allotype female, same data as holotype; both 
mounted on same pin. One paratype, same data as holotype. 

Remarks: The large size; short, stout beak; and male secondary 
sexual characters will distinguish this species. 

I am pleased to name this outstanding species in honor of my wife, 
Betty, in acknowledgment of her understanding interest during the 
course of this research. 


Apion (Trichapion) chalybaeum Wagner 


Apion (Trichapion) chalybaewm Wagner, Arch. Naturg. Berlin, vol. 78, p. 111, 
1912. 

Description: This species was described from a few specimens in 
the Solari Collection that were not seen by the author. The following 
notes are taken from the original description. 

Length, 1.8 to 1.9 mm. 

Closely related to A. oscillator Sharp, of same size, shape of pro- 
thorax and elytra, and color; differs markedly in form of rostrum, 
which in both sexes is nearly cylindrical, only at the antennal insertion 
slightly angulate, somewhat stouter especially in female, less strongly 


VOL. 110 


PROCEEDINGS OF THE NATIONAL MUSEUM 


266 


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267 


SUBGENUS TRICHAPION—KISSINGER 


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PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 116 


268 


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SUBGENUS TRICHAPION—KISSINGER 269 


curved, in both sexes more strongly punctate, duller, and especially 
in the male with punctures arranged in rows above antennal insertion. 
Head shorter and wider, eyes (especially of male) differently rounded, 
widest well behind middle; the wide frons furcate, interspace between 
middle and lateral stria moderately closely and extremely finely 
punctured. Antennae are somewhat stouter, especially the club 
noticeably larger. Prothorax same general shape but a little wider, 
punctures more scattered and on disc and base distinctly sparser 
than at foremargin (in oscillator punctures denser and less pro- 
nounced); the smooth middle line is more distinct and the basal fovea 
shallower than in oscillator. The intervals are nearly flat, extremely 
finely and sparsely punctured; scutellum is nearly twice as large, 
elongate-triangular; unfurcate. Legs as in oscillator, male with four 
hind tibiae mucronate with shorter, fine spine pointed more toward 
inner angle. Body color slightly shining black, elytra with a dark 
blue metallic luster. Pubescence of mesothorax and metathorax is 
not denser than oscillator, that of elytra and prothorax somewhat 
finer. 
Typr Locauity: Managua, Nicaragua. 


Apion (Trichapion) grossulum Fall 
Apion grossulum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 141, 1898. 


Description: Length, 2.37 mm.; width, 1.67 mm. 

Robust. Black, antennae in part piceous; pubescence of dorsal 
surface white, fine, very sparse, slightly coarser and more evident 
laterally but not especially denser. Male beak slightly shorter than 
head and prothorax, a little less than one-half longer than prothorax, 
moderately, evenly curved especially on dorsal margin; in lateral 
view stoutly parallel in basal third, attenuate to tip; in dorsal view 
expanded laterally at antennal insertion, attenuate from antennal 
insertion to near tip, tip slightly expanded; punctures rather coarse, 
close, arranged in rows, tip smoother; sparsely pubescent in basal 
two-thirds. Antennae inserted at basal fifth of beak at distance from 
eye equal to width of frons; first segment equals next two, second 
segment slightly longer than third. Eyes prominent; frons about 
equal to dorsal tip of beak, with a median, moderately deep, narrow 
sulcus and one lateral row of confluent punctures, appearing tri- 
sulcate. Prothorax one-sixth wider than long at base, middle slightly 
narrower than base, apex three-fourths as wide as base; sides beyond 
basal lateral expansion slightly converging, rounded to constricted 
apex, apex slightly flared out; punctation moderately deep, 0.04 mm. 
in diameter, interspaces generally less than diameter of punctures; 
basal fovea deep, short, nearly punctiform. Elytra at humeri one- 
third wider than prothorax at base; 2.5 times as long as prothorax; 


270 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


length to width as 13:10.5; intervals twice as wide as striae, slightly 
convex, with two rows of minute punctures bearing fine scales. Scutel- 
lum elongate-triangular, 0.12 by 0.06 mm. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, triangular mucrones which are acutely dentellate ventrally; 
mucrones project almost at right angle to long axis of tibiae. 

MATERIAL EXAMINED: Holotype male (MCZ 25099) in J. L. Le- 
Conte Collection. 

KNOWN DISTRIBUTION: Type is not labeled but is thought to be 
from Arizona according to Fall (1898). 


Apion (Trichapion) guatemalense Wagner 


Apion (Trichapion) guatemalense Wagner, Arch. Naturg. Berlin, vol. 78, p. 112, 
1912. 

Description: This species was described from two females from 
Guatemala City and Retalhuleu, Guatemala, in the Wagner Col- 
lection and in the British Museum (Natural History) that were not 
seen by the author. The following notes are a translation of the 
original description. 

Length, 1.9 to 2.0 mm. 

Near A. oscillator Sharp. Body pitchy brown, with a rather strong 
though nearly dull copper luster, especially on head, prothorax, and 
elytra; with fine sparse white pubescence, pubescence on sides of meso- 
thorax and metathorax not dense. Beak blackish, legs dark reddish 
brown. Head wider than long, of female slightly conical, with rather 
strongly convex though hardly prominent eyes, with a square depres- 
sion between, vertex with microscopic sculpture, more brassy; frons 
somewhat smaller than beak at base, with two rows of moderately fine, 
close, deep punctures; behind eyes head with stronger punctures 
less densely placed. Beak of female one-fourth longer than head and 
prothorax combined, relatively slender, slightly curved. From base 
to antennal insertion hardly widened, from there to tip slightly attenu- 
ate, nearly cylindrical at tip; at base with fine punctures, otherwise 
with very fine sculpture, nearly dull, glabrous. Antennae long and 
slender, of female inserted in basal third or fourth of beak, scape 
equals first three funicular segments; first slightly longer but dis- 
tinctly stouter than next, elliptical; second barrel-shaped; 3-7 grad- 
ually diminishing in length. Prothorax is somewhat wider than long, 
its base bisinuate, hind angles not prominent, nearly rectangular; 
at base slightly, at apex feebly sinuate, between uniformly, moder- 
ately rounded; widest at middle, only slightly narrowed toward 
front, at slightly concave foremargin about one-half narrower than 
base; in side view slightly, uniformly convex; with fine, not dense 
punctures, shallow, becoming larger and shallower towards sides, 


SUBGENUS TRICHAPION—KISSINGER Dik 


interspaces as wide or slightly narrower than punctures, finely sculp- 
tured; in front of scutellum with a short but deep basal fovea. Elytra 
rather shortly egg-shaped, base much wider than prothorax, with 
prominent humeri, sides gently widening toward rear, widest point 
somewhat behind middle, then widely rounded to rear, slightly sin- 
uate at apex; moderately large punctures in striae, flat intervals 
twice as wide as striae, in middle striae are finer, with fine sculpture 
of transverse wrinkles. Scutellum rounded, unfurcate, enclosed in 
a sharp furrow. Legs moderately long, femora rather stout; first 
tarsal segment about as long as second, as long as wide, fourth longer 
than third by one-half, claws distinctly toothed. 


Apion (Trichapion) laminatum Sharp 
Ficurs l,a 


Apion laminatum Sharp, Biologia Centrali-Americana. Coleoptera, vol. 4, pt. 3, 
p. 58, pl. 3, figs. 6, 6a, 1890. 

Duscription: This species was described from a single specimen 
in the British Museum (Natural History) that was not seen by the 
author. The following descriptive notes were taken from the original 
description. 

Length, 2.25 mm. 

Black, opaque; very sparsely pubescent. Male beak not quite as 
long as head and prothorax combined, with a large ventral lamina 
extending from antennal insertion to apex. Antennae inserted very 
near to eyes. 

Special male characters: Middle tibiae bear a minute mucro at 
apex, laminate beak is probably a male secondary sexual character. 

Typr LocaLiry: Amula, Guerrero, Mexico. 


Apion (Trichapion) peninsulare Fall 


Apion peninsulare Fall, Trans. Amer. Ent. Soc., vol. 25, p. 149, pl. 5, fig. 2, 2a, 
1898. 

Description: The type of this species is in the California Academy 
of Sciences and was not seen by the author. The following notes 
are taken from the original description. 

Length, 2.0 mm. 

Black; sparsely pubescent. Male beak shorter than head and 
prothorax combined, stout, strongly dilated a little behind the middle, 
rugosely punctate laterally, more finely and sparsely punctured 
dorsally. Antennae rather short, first segment scarcely as long as 
next two, second globose, barely reaching eye. Frons_ slightly 
sulcate; eyes not prominent. Prothorax about as long as wide; 
base one-fourth wider than apex; sides divergent to middle, subparallel 
in basal half. Elytra one-third longer than wide; humeri moderately 


212 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


prominent; sides subparallel, very slightly arcuate; intervals wide, 
moderately convex. Rather strongly uniformly punctate ventrally. 
Special male characters: Tibiae 2 and 3 armed with short mucrones. 
Female unknown. 
Typr Locatiry: La Chuparosa, Baja California, Mexico. 


Apion (Trichapion) persulcatum Wagner 


Apion persuleatum Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 22; pl. 2, figs. 
3a, 3b, 1911. 

Description: This species was described-from a male in the 
Wagner Collection and a pair in the British Museum (Natural 
History) that were not seen by the author. The following notes 
are taken from the original description. 

Length, 2.8 to 2.9 mm. 

Close to Apion subnudum Wagner [which does not seem to belong 
to the subgenus Zrichapion]. The body is black, only slightly 
shining, the elytra are bright as in subnudum but are more grayish 
blue. Beak in both sexes distinctly longer than subnudum and of 
the female more slender, about one-third longer than head and 
prothorax combined, of the male somewhat shorter; in both sexes 
somewhat more curved than swbnudum. The prothorax is somewhat 
shorter than swbnudum, sides between hind angle and anterior margin 
uniformly rounded; with large, shallow punctures rather densely 
placed, microscopically finely chagrined interspaces not wider than 
punctures; with a rather strong, short basal fovea. Elytra behind 
well developed humeri strongly widened posteriorly, widest behind 
the middle. Scutellum like swhnwdum. The tarsi are more slender; 
the first segment of the front leg is about one-half longer and narrower 
than second; the four posterior legs are shorter; the last two pairs 
of legs of the male are mucronate. 

KNOWN DISTRIBUTION: 


GUATEMALA: Capetillo; Retalhuleu. 
Apion (Trichapion) spiculiferum Wagner 
Apion (Trichapion) spiculiferum Wagner, Arch. Naturg. Berlin, vol. 78, p. 104, 1912. 


Description: This species was described from a few specimens in the 
British Museum (Natural History) that were not seen by the author. 
The following notes are taken from the original description. 

Length, 1.9 to 2.2 mm. 

Closely related to A. aurichaleewm Wagner, which has a larger body 
and bright sheen and gloss, finer, sparser pubescence especially on the 
beak. Female beak a little longer than head and prothorax, strongly 
curved, brightly shining, with single, fine punctures, a little denser 
over the antennae; male beak somewhat shorter than that of female, 


-* 


SUBGENUS TRICHAPION—KISSINGER 213 


less strongly curved, basal one-half with deep, but not dense punctures, 
finely pubescent, apical one-half nearly glabrous, bare, nearly cylindri- 
cal, feebly swollen at antennal insertion. Antennae inserted in basal 
one-third of beak of male, of female at long diameter of eye. Pro- 
thorax in form and sculpture as in A. aurichalceum, with the exception 
of the less constricted front. Elytra are only a little shorter, about 2.5 
times as long as prothorax (A. aurichalceum about three times as long 
as prothorax); striae strongly punctured; intervals somewhat convex, 
asin A. aurichalceum. Legs a little more slender, especially the tibiae. 
Male as A. aurichalceum with tibiae 2 and 3 mucronate, but spine 
somewhat longer and pointing more to the rear. 

KNOWN DISTRIBUTION: 

Mexico: Veracruz: Jalapa. Guanajuato. 

GuaTEMALA: San Gerénimo. 

Remarks: Mr. J. Balfour-Browne (personal communication) has 
studied the type and is of the opinion that this species is a synonym 
of A. godmant Wagner. 


Apion (Trichapion) tabogense Sharp 


Apion tabogense Sharp, Biologia Centrali-Americana. Coleoptera, vol. 4, pt. 3, 
p. 52, 1890. 

Description: This species was described from two males in the 
British Museum (Natural History) that were not seen by the author. 
The following notes are taken from the original description. 

Length, 2.00 mm. 

Middle and hind tibiae mucronate, mucrones are slender spurs di- 
rected straight downwards, not inwards as in most of the other species. 
Rostrum is rather thick, curvate, densely sculptured except at tip, 
antennae are inserted not very far from eyes, being separated from 
them by not quite one-fourth length of rostrum; eyes rather large and 
prominent, placed close to the prothorax, frons moderately broad. 
Thorax rather broad, much narrowed towards front, scarcely impressed 
at sides, moderately closely and coarsely but obsoletely punctured, 
with a small fovea in front of scutellum. Elytra short and broad, 
coarsely striate, though striae are scarcely deep enough to be called 
sulci, intervals rather broad, flat. White setosity is not quite so 
scanty on undersurface as hae) under orbits more coneU Caan: 

Typs Locauity: Taboga Island, Panama. 


Apion simile Group 


Three species grouped with A. simile Kirby agree in general char- 
acters as outlined above. In addition the density and pattern of the 
pubescence is similar and the femora of the male tend to be stouter 


274 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


than those of the female, except in A. meorrhynchum Philippi. The 
four species apparently do not overlap in range. It is possible that 
A. modestum Smith overlaps A. simile in Nebraska or Iowa. A. pro- 
pinquicorne Fall is more southern in distribution, occurring in southern 
Texas and Mexico. A. meorrhynchum Philippi occurs in Chile. 

A. simile differs from A. modestum and A. propinquicorne by the 
longer beak of both sexes and more distally inserted antennae. The 
latter two species are closely allied—small, densely pubescent, with 
antennae inserted near the base of the beak. The male of A. propin- 
quicorne has a beak which is slightly longer than the prothorax; the 
antennae in both sexes are inserted close to the base of the beak at a 
distance from the eye less than the width of the frons; and the elytral 
intervals are biserially pubescent. The male of A. modestum has a 
beak which is as long as the prothorax; antennae of male are inserted 
at distance from eye equal to width of frons; and the elytral intervals 
are generally uniserially pubescent, but the scales may be confused, 
appearing in part as two rows. A. meorrhynchum is well distinguished 
by the antennae which are inserted at about the middle of the beak, 
the antennal scape is longer than the next five segments, and the 
elytral intervals are clothed with four to six rows of scales. 

Two lines of species groups seem to be related to the holarctic species, 
Apion simile. One line is composed of the A. proclive and A. glyphi- 
cum groups. They have been differentiated from A. simile along the 
following lines: the male mucrones are generally strongly dentate, 
the frons is narrow, in some cases narrower than the dorsal tip of the 
beak, head is much more strongly constricted dorsally behind the eyes, 
and the female beak, while elongate, is not cylindrical but gradually 
attenuate toward the apical region and the apex is generally expanded 
laterally. The A. proclive group overlaps the range of the A. simile 
group in the western portion of the United States and extends well 
into Mexico. The A. glyphicum group overlaps the range of the A. 
proclive group in Mexico and extends into Central America. 

The other line is composed of the A. griseum, A. spinitarse, A. 
punctulirostre, and A. submetallicum groups. They have become 
differentiated from A. simile along the following lines: the A. griseum 
group exhibits additional male secondary sexual modifications of the 
fore tibiae; in both sexes the beak is more attenuate toward the apical 
third; generally the female beak is expanded laterally at the tip; 
the frons is comparatively broader, and in general the antennae of the 
male are inserted closer to the eyes at a distance less than the width 
of the frons; the males of the A. spinitarse and A. punctulirostre groups 
have only the middle tibiae mucronate; the A. submetallicum group 
exhibits modification of the beak in that it is more attenuate to the 


SUBGENUS TRICHAPION—KISSINGER 275 


tip; in the male it is pubescent to the tip and in the female is pubescent 
to a point well in front of the antennal insertion. The A. griseum 
group overlaps the range of the A. stmile group particularly in Eastern 
North America and extends into South America. It is interesting 
to note that the superficial resemblance of A. sayi Gyllenhal (of the 
A. griseum group) to A. simile is great enough that in at least two 
cases in the material studied the two species were confused by com- 
petent coleopterists. 


Apion (Trichepion) meorrhynchum Philippi 


Apion meorrhynchum Philippi, Stettiner Ent. Zeit., vol. 25, p. 364, 1864.— 
Kuschel, Agr. Tecn. Chile, vol. 10, p. 16, 1950. 

Apion tenebricosum Gemminger, Coleopterologische Hefte, No. 8, p. 123, 1871.+~ 
Kuschel, Agr. Teen. Chile, vol. 10, p. 16, 1950 (new name for Apion obscurum 
Blanchard, in Gay, Historia fisica y politica de Chile, vol. 5, p. 309, 1851; 
nec Kirby, 1811). 

Description: Length, 2.75 to 3.25 mm.; width, 1.33 to 1.50 mm. 

Moderately robust. Black; pubescence white, silvery, fine, long, 
dense. Male beak equal to, to slightly longer than, head and pro- 
thorax combined, three-fifths to four-fifths longer than prothorax, 
slightly curved; subcylindrical throughout, slightly expanded laterally 
at antennal insertion; rather densely, finely punctured, with minute 
scales to near apex, tip slightly smoother. Female beak one-ninth to 
two-fifths longer than head and prothorax combined, four-fifths to 

2.25 times as long as prothorax, slightly curved, subcylindrical, 

slightly expanded laterally at antennal insertion and at apex; punc- 

tation and pubescence similar to male, tip polished, bare. Antennae 
of male inserted at distance from eye 2.5 times width of frons, at 
middle to slightly basad of middle; of female at distance from eye 
three times width of frons, slightly basad of middle; first segment 
equals next five, second segment slightly longer than third; club 0.27 
by 0.10 mm. Eyes prominent; frons of male slightly narrower than 
dorsal tip of beak, of female equal to dorsal tip of beak, with slightly 
concave area medially. Prothorax at base about one-fifth wider than 
long, middle slightly narrower than base, apex three-fourths as wide 
as base; sides beyond acute basal lateral expansion subparallel to 
middle, rounded to slightly constricted apex; in profile dorsal surface 
nearly flat (slightly convex in some females); punctation fine, moder- 
ately deep, dense, uniform; basal fovea shallow, broad, short. Elytra 
at humeri one-third wider than prothorax at base, about three times 
as long as prothorax, length to width as 15:11 to 3:2; intervals 
convex, about twice as wide as striae, with four to six rows of fine 
scales; striae deep, coarse, scales in striae similar to those on interval. 
Scutellum 0.06 by 0.06 mm., subquadrate, with slight median longi- 


276 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


tudinal depression basally, with a more or less distinct apical trans- 
verse pit. Front femora about four times as long as wide. Claws 
with large, blunt basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, nearly simple mucrones. 

MATERIAL EXAMINED: 14 examples including material determined 
by Wagner. 

KNOWN DISTRIBUTION: 

Cuivre: Caramavida, Feb. 5-10, 1953, Luis E. Pefia (DGK); Pichinahuel, 


February 1953, Luis E. Petia (DGK); Recinto, Jan. 16, 1953, Luis E. Pefia 
(DGk). 


Apion (Trichapion) modestum Smith 
FIGURE 2,a 
Apion modestum Smith, Trans. Amer. Ent. Soe., vol. 11, p. 58, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 138, pl. 4, figs. 8, 8a, 1898.—Blatchley 
and Leng, Rhynchophora or weevils of northeastern America, p. 78, 1916. 

Descriprion: Length, 1.36 to 1.62 mm.; width, 0.67 to 0.77 mm. 
Moderately robust; black, antennae piceous at base. Pubescence 
conspicuous, white, on dorsal surface rather coarse, coarser and denser 
laterally and ventrally. Male beak about as long as prothorax, 
nearly straight, nearly cylindrical, slightly expanded laterally at 
antennal insertion, punctured and pubescent in basal two-thirds, 
apical third smooth, bare, shining. Female beak as long as head and 
prothorax combined, one-half longer than prothorax, slightly curved; 
slightly attenuate beyond antennal insertion, slightly expanded later- 
ally at tip; scantily pubescent behind antennal insertion, with sparse, 
moderate punctures laterally, smoother at tip. Antennae of male 
inserted at distance from eye equal to width of frons, at basal one- 
fourth of beak, of female inserted at distance from eye slightly less 
than width of frons, at basal one-sixth of beak; first segment and 
second segment each as long as next two, dimensions of club from 
0.14 by 0.06 mm. to 0.16 by 0.07 mm. Eyes prominent; frons wider 
than dorsal tip of beak. Prothorax one-fourth wider at base than 
long, middle slightly narrower than base, apex four-fifths as wide as 


Ficure 2.—a, Apion modestum Smith, lateral view of head and prothorax of male. 5, 4, 
propingutcorne Fall, lateral view of head and prothorax of male. c, 4. simile Kirby 
lateral view of head and prothorax of male. Line equals 0.50 mm. 


SUBGENUS TRICHAPION—KISSINGER DAG | 


base; sides beyond basal lateral expansion nearly parallel to middle, 
rounded to moderately constricted apex; in profile dorsal surface 
nearly flat; punctures deep, 0.03 mm. in diameter, interspaces less 
than diameter of punctures, generally one-half as wide, alutaceous; 
basal fovea punctiform, moderately deep. Elytra at humeri one- 
third wider than prothorax at base, 2.75 to 3.00 times as long as 
prothorax, length to width as 11:8; intervals about twice as wide as 
striae, nearly flat on disc, generally with one row of scales, these 
sometimes confused and appear in part as two rows of scales; striae 
fine, deep. Scutellum triangular, 0.06 by 0.06 mm., with slight 
median furrow. Front femora about three times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, subangulate mucrones. 

MATERIAL EXAMINED: 10 specimens including material determined 
by Fall. 

KNOWN DISTRIBUTION: 

Unirep States: Arizona: Chiricahua Mts. (USNM). Colorado (Fall, 1898). 
Illinois (MCZ). Nebraska (MCZ). 

Remarks: No type specimens were seen in either the U.S. National 
Museum Collection or in the J. L. LeConte Collection, the two 
collections containing most of Smith’s types. IJ am following Fall 
(1898) in the definition of this species. 


Apion (Trichapion) propinquicorne Fall 
FIGURE 2,b 


Apion propinquicorne Fall, Trans. Amer. Ent. Soc., vol. 25, p. 138, pl. 4, figs. 7, 
7a, 1898. 


Descriprion: Length, 1.67 mm.; width, 0.92 mm. 

Robust; black. Pubescence conspicuous, white, rather coarse, 
denser on ventral surface. Male beak shorter than head and pro- 
thorax combined, one-fifth longer than prothorax, slightly, evenly 
curved; expanded laterally at antennal insertion, apical half nearly 
cylindrical; basal half sparsely punctured and pubescent, apical half 
smoother, bare, shining. Female beak as long as head and prothorax 
combined, two-fifths longer than prothorax, moderately, evenly 
curved; apical two-thirds nearly cylindrical; shining beyond antennal 
insertion, basal half with strong, lateral punctures. Antennae in- 
serted at distance from eye less than width of frons, of male inserted 
at basal sixth of beak, of female at basal one-seventh; first segment 
shorter than next two, second segment equal to next two, club 0.17 
by 0.06 mm. Eyes prominent; frons wider than dorsal tip of beak. 
Prothorax at base one-fifth wider than long, middle slightly narrower 


278 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


than base, apex three-fourths as wide as base; sides beyond basal 
lateral expansion nearly parallel to middle, rounded to constricted 
apex; in profile dorsal surface nearly flat; punctures moderately deep, 
0.03 mm. in diameter, interspaces less than diameter of punctures; 
basal fovea punctiform, moderately deep. Elytra at humeri one-half 
wider than prothorax at base, 2.8 times as long as prothorax, length to 
width as 7:5; intervals twice as wide as striae, nearly flat, with two 
rows of scales; striae moderately deep, fine. Scutellum triangular, 
0.05 by 0.04 mm., with a slight median furrow. Front femora 3.4 
times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 with moderately long, 
nearly simple mucrones. 

Typrs: I hereby designate the lectotype of this species as the 
female specimen (USNM 4233) labeled San Diego, Tex. A male 
cotype (MCZ 25117) in the Fall Collection has the same data as 
lectotype. Fall (1898) states that the material on which the species 
was based was collected by Mr. Schwarz and was in the collection of 
the U. S. Department of Agriculture, hence the selection of the 
National Museum specimen as lectotype. 

MATERIAL EXAMINED: Lectotype, cotype, and five specimens. 

KNOWN DISTRIBUTION: 

Unitep States: Texas: Del Rio (MCZ); San Diego (MCZ, USNM). 

Mexico: Mexico: 30 miles southeast of Mexico City, July 1954, D. G. Kis- 
singer. 

Remarks: The material from Mexico was taken while beating 
scrubby oak and acacia bushes. 


Apion (Trichapion) simile Kirby 


FIGURE 2,c 


Apion simile Kirby, Trans. Linn. Soc. London, vol. 10, p. 351, 1811.—Germar, 
Mag. Ent., vol. 2, p. 208, 1817.—Gyllenhal, in Schoenherr, Genera et species 
curculionidum, vol. 1, p. 424, 1833.—Wencker, L’Abeille, vol. 1, p. 181, 
1864.—Bedel, Faune des Coléoptéres du bassin de la Seine, vol. 6, p. 377, 
1885.—Desbrochers, Le Frelon, vol. 5, p. 244, 1895-1896.—Schilsky, in 
Kiister and Kraatz, Die Kafer Europa’s, vol. 39, p. 89, 1902.—Wagner, 
Genera insectorum, Coleoptera, fam. Curculionidae: subfam. Apioninae, 
fasc. 130, p. 84, 1912.—Hustache, Famille Lxx1x Curculionidae, Tribu: 
Apioninae. Tableaux analytiques des Coléoptéres de Ja Faune Franco- 
Rhenane, p. 268, 1931.—Bartoszynski, Polaski Pismo Ent., vol. 14-15, p. 
120, 1935.—Donisthorpe, Ent. Monthly Mag., vol. 74, p. 171, 1938.— 
Walhgren, Ent. Tidskr., vol. 72, p. 140, 1951.—Kissinger, Proc. Ent. Soc. 
Washington, vol. 59, p. 40, 1957. 

Apion superciliosum Gyllenhal, Fauna Suecica, vol. 3, p. 58, 1813.—Wagner, 
Genera insectorum, Coleoptera, fam. Curculionidae: subfam. Apioninae, 
fasc. 130, p. 84, 1912.—Kissinger, Proc. Ent. Soc. Washington, vol. 59 
p. 40, 1957. 


SUBGENUS TRICHAPION—KISSINGER 279 


Apion triste Germar, Mag. Ent., vol. 2, p. 233, pl. 3, fig. 2, 1817.—-Wagner, Genera 
insectorum, Coleoptera, fam. Curculionidae: subfam. Apioninae, fase. 130, 
p. 84, 1912.—Kissinger, Proc. Ent. Soc. Washington, vol. 59, p. 40, 1957. 

Apion walshiit Smith, Trans. Amer. Ent. Soc., vol. 11, p. 57, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 142, pl. 4, figs. 12, 12a, 18, 1898.—Blatch- 
ley and Leng, Rhynchophora or weevils of northeastern America, p. 79, fig. 
37r, 1916.—Wolcott and Montgomery, Amer. Midl. Nat., vol. 14, p. 161, 
1933.—Kissinger, Proc. Ent. Soc. Washington, vol. 59, p. 40, 1957. [New 
name for Apion lanuginosum Walsh, Proc. Ent. Soc. Philadelphia, 1866, 
vol. 6, p. 269, 1867, nec Gerstaecker, 1854.] 

Apion vicinum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 58, 1884.—Fall, Trans. 
Amer. Ent. Soc., vol. 25, p. 143, 180, 1898.—Kissinger, Proc. Ent. Soc. 
Washington, vol. 59, p. 40, 1957. 

Apion eppelsheimi Faust, Deutsche Ent. Zeitschr., 1887, p. 179, 1887.—Wagner, 
Genera insectorum, Coleoptera, fam. Curculionidae: subfam. Apioninae, fase. 
130, p. 84, 1912.—Kissinger, Proc. Ent. Soc. Washington, vol. 59, p. 40, 1957. 

Description: Length, 1.70 to 2.20 mm.; width, 0.70 to 1.00 mm. 

Elongate, moderately slender. Black; pubescence conspicuous, 
fine, long, whitish to yellowish white, sparse, uniform. Male beak 
one-fourth longer than prothorax, nearly straight, cylindrical, slightly 
dilated laterally over antennal insertion; basal two-thirds mod- 
erately punctured and with sparse pubescence, apex smooth, more 
shining. Female beak equal to, to much longer than, head and 
prothorax, one-half to twice as long as prothorax, slightly curved, 
nearly cylindrical throughout; finely punctured throughout, tip 
slightly more shining. Antennae of male inserted at distance from 
eye twice as great as width of frons, at basal two-fifths of beak; of 
female inserted at distance from eye one-half greater than width of 
frons, at basal third of beak; first segment about equal to next three; 

second segment longer than third, shorter than next two; club 0.16 

by 0.08 to 0.21 by 0.10 mm. Eyes moderately prominent; frons 

moderately wide, shallowly punctured, alutaceous. Prothorax one- 
seventh to one-fourth wider at base than long, middle a little wider 
than base, apex from three-fourths to four-fifths as wide as base; sides 
with very slight lateral expansion at base, diverging slightly to middle, 
rounding to slightly constricted apex; in profile dorsal surface slightly 
arcuate, flatter at apex and base; punctation 0.03 mm. in diameter, 
moderately deep, interspaces variable, less than, to equal to, diameter 
of punctures; basal fovea shallow, rounded. Elytra one-fourth to 

one-third wider at humeri than base of prothorax, from 3.2 to 3.5 

times as long as prothorax, length to width varies from 11:7 to 

14 : 8.5; intervals nearly twice as wide as striae, variable but convex, 

with one or two rows of fine punctures bearing fine scales; striae deep, 

with one row of fine scales. Scutellum triangular, 0.06 by 0.04 mm., 

with median furrow or not. Front femora of male 3.2, of female 3.8 

times as long as wide. Claws with acute basal tooth. 

483661593 


280 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Special male characters: Tibiae 2 and 3 armed with simple, moder- 
ately long mucrones. 

Types: I hereby designate as lectotype of A. walshii Smith the 
female specimen (USNM 1256) labeled White Mts., N. H. Cotypes 
labeled Marquette, Mich., are in the J. L. LeConte Collection (MCZ 
369). 

I hereby designate as the lectotype of A. vicinum Smith the female 
specimen (USNM 1255) labeled Garland, Colo. Cotypes with the 
same data are in the J. L. LeConte Collection (MCZ 370). 

MaTERIAL EXAMINED: Lectotypes of A. walshii and A. vicinum, 
specimens of A. simile from Europe determined by Hans Wagner in 
the Bovie Collection of the U. S. Nationa] Museum, and 150 speci- 
mens from the United States and Canada. 

Biotoey: Apion lanuginosum Walsh was described from material 
taken from galls made by a Cecidomya sp. on Saliz strobiloides. Do- 
nisthorpe (1938) records A. simile from Windsor Forest, England, 
by sweeping and under faggots in a willow swamp. Fall (1898) 
states that in New England A. walshii occurs abundantly on white 
birch, Betula alba. Hustache (1931) records A. simile as occurring 
on Betula alba, stating that the species is common in France. I have 
taken a large series from birch leaves in New Hampshire. Wahlgren 
(1951) cites A. simile as a gall maker on Betula verrucosa. Wolcott. 
and Montgomery (1933) took A. walshit by sweeping in the maple 
zone of a Tamarack swamp in Porter Co., Ind. 

KNOWN DISTRIBUTION: 

Europe, Algeria, Asia Minor, and Siberia according to Hustache (1931). 

Canapa: Newfoundland: Lomond, Bonne Bay, July 15, 1949, E. Palmen 
(CNC); Stead Bank, Humber, July 10, 1949, E. Palmen (CNC). Nova Scotia: 
Annapolis Royal, July 4, 1928, W. J. Brown (CNC). New Brunswick: Boies- 
town, July 13, 1928, W. J. Brown (CNC). Quebec: Knowlton, Aug. 1, 1929, 
L. J. Milne (CNC); Queens Park, Aytmer, July 23, 1926, C. B. Hutchings (CNC). 
Ontario: Merivale, June 16, 1931, W. J. Brown (CNC); Ottawa (CNC). Mani- 
toba: Aweme, June 9, 1930, R. M. White (CNC); Onah, June 19, 1930, R. M. 
White (CNC). 

Unirep States: North Carolina: Mt. Mitchell, 4,000-6,000 ft., June 1939, 
Quirsfeld (CAF). Maryland: Beltsville (USNM); Branchville (USNM). New 
York: Colden, May 1885, E. P. Van Duzee (CIS); Ithaca (USNM); Lancaster, 
May 9, 1888, E. P. Van Duzee (CIS); Portage, May 30, 1888, E. P. Van Duzee 
(CIS). Rhode Island (TLCC). New Hampshire: Manchester (INHS); Mt. 
Washington, Ammonoosuc Ravine, 4,000 ft., June 4, 1940, M. A. Hanson (CAF), 
Great Gulf Trail, 1,500 ft., June 1, 1940, J. F. Hanson (CAF), Pinkham Notch, 
July 3, 1951, common on birch leaves, D. G. Kissinger (DGK), Tuckerman’s 
Ravine, 2,000 ft., June 2, 1940, J. F. Hanson (CAF). Vermont: Lake Willoughby, 
1,400 ft., June 17-29, 1945, C. P. Alexander (CAF). Ohio: Champaign Co., 
Cedar Swamp, May 22, 1954, R. E. Woodruff (ELS); Hocking Co., March 2], 
1951, May 1, 1954, Sept. 10, 1953, N. J. and E. L. Sleeper (ELS); Vinton Co., 
June 5, 1953, N. J. and E. L. Sleeper (ELS). Indiana: Porter Co. Michigan: 


SUBGENUS TRICHAPION—KISSINGER 281 


Marquette (MCZ). Jowa: Iowa City, L. L. Buchanan (USNM); Lake Okoboji, 
July 22, 1916, L. L. Buchanan (USNM). South Dakota: Custer, Aug. 28, 1935, 
M. W. Sanderson (UK). Washington: Deeplake, May 5, 1949, G. H. Nelson 
(DGK). California: Siskiyou Co. (TLCC). Utah: Cedar City, Aug. 13, 1929, 
P. W. Oman (UK); Emery, Aug. 16, 1929, P. W. Oman (UK). Colorado (MCZ, 
USNM). 


Remarks: This is the first time an Apion species is recorded as 
being holarctic in distribution. J am indebted to Dr. W. J. Brown 
for informing me (personal communication) about the new record of 
a European species of Apion being found in Newfoundland by Dr. 
Palmen. 


Apion proclive Group 


The six species grouped here agree in general characters as mentioned 
above. In addition the beak of the male in dorsal view is attenuate 
distad of the antennal insertion, generally to the apical third, and there 
is marked sexual dimorphism in the structure of the beak; in the 
female this part is slender, elongate, and more strongly attenuate. 
The ranges of the Mexican species are incompletely known due to lack 
of collections. The ranges of A. coryi, new species, and A. proclive 
LeConte probably do not overlap. 

A comparison shows that the males of two species, A. proclive and 
A. coryi, have dentate mucrones, while the mucrones of the males of 
the remaining species are nearly sunple. The former two species are 
apparently closely allied. The male of A. coryi has all three pairs of 
tibiae mucronate while A. proclive has only two pairs of tibiae mucron- 
ate. However under the discussion of A. proclive an interesting series 
from Arizona is noted which is apparently identical to A. proclive but 
the front tibiae of the male are minutely mucronate. The main 
differences between the two species are as follows: A. coryi has the 
elytra comparatively short and more robust, being only 2.5 times as 
long as the prothorax, the elytral intervals are more convex, the 
prothorax is more coarsely and closely punctured, the basal fovea of 
the prothorax extends one-third its length, the beak of the male is as 
long as the head and prothorax combined, more noticeably deflexed at 
the apical fourth, the beak of the female is more strongly deflexed at 
the apical third, and the eyes are more prominent. The elytra of A. 
proclive are comparatively longer, being 2.8 times as long as the pro- 
thorax, the elytral intervals are flatter, the basal fovea of the pro- 
thorax short, the beak of the male is shorter than the head and pro- 
thorax combined, the beak of both sexes is evenly curved, the eyes are 
not prominent, and the species is generally larger in size. 

Of the other species, A. acanonicum, new species, is immediately 
distinct because of the unusual male characters. Apion imitator 


282 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Wagner has tibiae 2 and 3 of the male armed with similar, simple 
mucrones. The male of A. plectrocolum, new species, has the mucro 
on tibia 3 much longer than that on tibia 2. A. adaetum, new species, 
has the frons distinctly wider than the dorsal tip of the beak; A 
imitator and A. plectrocolum have the frons equal to the width of the 
dorsal tip of the beak. 

The affinites of A. proclive and A. cory? are close to the A. glyphicum 
group. The main difference between the A. proclive and A. glyphicum 
groups is that the characters of A. proclive which separate it from A. 
simile Kirby are more strongly developed in the A. glyphicum group, 
namely the more strongly dentate mucrores of the male; the shorter, 
stouter beak; the larger, more protruding eyes; the narrower frons; 
and the much more strongly constricted dorsal portion of the head 
above the hind margin of the eyes. 

An interesting character shared by the members of this group is a 
more or less evident row of punctures extending about one-fourth the 
length of the beak above the antennal insertion in the male. In A. 
proclive this row of punctures generally appears as a definite sulcus 
above the antennal insertion. In the A. glyphicum group the beak of 
the male has a deep lateral sulcus limited by an upper and lower 
raised carina. This sulcus is apparently a development of the row of 
punctures above the antennal insertion noted in A. proelive. 


Apion (Trichapion) acanonicum, new species 
FIGURE 3,a-e 


Description: Length, 1.94 to 2.00 mm.; width, 0.09 to 1.00 mm. 

Robust. Black, base of antennae piceous; pubescence white, on 
dorsal surface very fine, sparse, much coarser and somewhat denser on 
sides of prothorax, mesothorax, and metathorax. Male beak moder- 
ately slender, five-sixths as long as head and prothorax, one-fourth 
longer than prothorax, slightly curved; in lateral view apical third 
nearly parallel; in dorsal view slightly expanded laterally at antennal 
insertion, attenuate to middle, there somewhat compressed, expanding 
to apical third which is nearly parallel sided to apex; basal two-thirds 
dull, punctured, sparsely pubescent, a coalesced row of punctures 
forming a vague sulcus above antennal insertion in basal third, apical 
third shining, finely, more sparsely punctured. Female beak one- 
third longer than head and prothorax combined, twice as long as 
prothorax, in lateral view nearly parallel-sided to tip, tip somewhat 
depressed; in dorsal view attenuate beyond antennal insertion, nearly 
parallel-sided beyond basal third, definitely expanding toward apex; 
not pubescent beyond antennal insertion, basal three-fourths finely, 
moderately densely punctate, apical fourth smoother, more shining. 
Antennae of male inserted at basal fourth of beak, at distance from eye 


SUBGENUS TRICHAPION—KISSINGER 283 


SS) 


O 
S 


h 
1 


Ficure 3.—a-e, Apion acanonicum, new species: a, fifth ventrite of male; b, dorsal view of 
head of female; c, lateral view of head and prothorax of female; d, lateral view of head 
and prothorax of male; ¢, entire dorsal view of male. f-1, A. coryi, new species: f, lateral 
view of head and prothorax of male; g, lateral view of head and prothorax of female; 
h, mucro of tibia 3 of male; i, entire dorsal view of male. Line equals 0.50 mm. 


284 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


equal to width of frons; of female inserted at basal fifth of beak at 
distance from eye one-half greater than width of frons; first segment 
equal to next three, second segment equals next two, club 0.21 by 0.08 
mm. Eyes prominent; frons slightly wider than dorsal tip of beak, 
with a shallow linear median sulcus and two lateral rows of fine 
punctures. Prothorax at base one-fourth wider than long, middle 
narrower than base, apex 0.7 as wide as base; sides beyond basal 
lateral expansion slightly, roundly expanding to middle, rounding to 
constricted apex; in profile dorsal surface slightly arcuate; punctation 
0.03 mm. in diameter, moderately deep, interspaces generally less than 
the diameter of punctures, there is a narrow, nearly impunctate 
median area; basal fovea moderately shallow, rounded. LElytra at 
humeri one-third wider than prothorax at base, 2.5 times as long as 
prothorax, length to width as 10 : 7.5; intervals twice as wide as striae, 
somewhat convex, with one row of minute punctures bearing very fine 
scales; striae moderately deep, fine. Scutellum triangular, 0.06 by 
0.06 mm., with very slightly median furrow. Front femora 3.3 times 
as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, nearly simple mucrones, femora 2 and 3 incrassate, first segment 
of tarsus 1 clothed beneath with moderately long setae, fifth ventral 
segment modified as follows: raised and convex transversely at base; 
in median third with brush of dense, very fine long cilia; lateral apical 
angles produced into a pair of triangular processes projecting poste- 
riorly and downward slightly beyond elytral tips, processes not visible 
in dorsal view. 

Typxs: Holotype male (CAS) from village on Maria Madre Island, 
Tres Marfas Islands, Mexico, May 15, 1925, H. H. Keifer. Allotype 
female (CAS), same data as holotype. Female paratype (DGK) from 
Tapanatepec, Oaxaca, Mexico, Jan. 22, 1953, D. G. Kissinger. 

Remarks: The extraordinary characters of the male will readily 
distinguish this species. The long beak of the female and its rather 
broad head will help to recognize that sex. 


Apion (Trichapion) adaetum, new species 
Fiaure 4,h 


Description: Length, 1.87 mm.; width, 0.87 mm. 

Moderately robust. Black, legs piceous; pubescence of dorsal 
surface fine, yellowish, sparse, on sides of prothorax, mesothorax, and 
metathorax white, much coarser and somewhat denser, a few whitish 
scales at base of interval 3. Beak of male shorter than head and pro- 
thorax combined, two-fifths longer than prothorax, moderately curved; 
in lateral view more or less attenuate to tip; in dorsal view moderately 


SUBGENUS TRICHAPION—KISSINGER 285 


expanded at antennal insertion, attenuate to apical two-fifths, nearly 
parallel to tip; basal two-fifths sparsely pubescent, moderately punc- 
tured, a line of punctures appears as a slight sulcus above antennal 
insertion and extends to apical two-fifths; tip smoother, bare. An- 
tennae inserted at basal fifth of beak, at distance from eye one-third 
greater than width of frons; first segment equals next two, second 
segment shorter than next two; club 0.18 by 0.06 mm. Eyes promi- 
nent; frons wider than dorsal tip of beak, with a shallow median sulcus 
and two lateral rows of punctures. Prothorax at base one-fourth 
wider than long, middle narrower than base; apex three-fourths as 
wide as base; sides beyond basal lateral expansion nearly parallel to 
middle, rounded to constricted apex; in profile dorsal surface nearly 
flat; punctation moderately deep, 0.03 mm. in diameter, interspaces 
generally narrower than punctures; basal fovea shallow, punctiform. 
Elytra at humeri one-third wider than prothorax at base, nearly three 
times as long as prothorax, length to width as 10:7; intervals twice 
as wide as striae, slightly convex, with two rows of punctures bearing 
scales. Scutellum elongate-triangular, 0.07 by 0.04 mm. Front 
femora 3.3 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, subangulate mucrones. 

Hototyrse: Male (USNM 63419) from Frontera, Tabasco, Mexico, 
in Thomas Lincoln Casey Collection. 


Apion (Trichapion) coryi, new species 
FIGURE 3, f—2 


Description: Length, 1.75 to 1.94 mm.; width, 0.87 to 1.05 mm. 

Moderately robust. Black, elytra slightly aeneous. Pubescence 
white, fine, scant on dorsal surface, denser on mesothorax. Male beak 
equal in length to head and prothorax combined, two-fifths longer than 
prothorax; slightly curved, more noticeably deflexed at apical fourth; 
apical third is nearly cylindrical; in dorsal view moderately expanded 
laterally at antennal insertion, attenuate to apical third; somewhat 
alutaceous, punctures arranged in rows, 2 moderately strong series of 
punctures above antennal insertion, tip somewhat more shining; basal 
two-thirds sparsely pubescent. Female beak longer than head and 
prothorax combined, four-fifths longer than prothorax; more strongly 
deflexed at apical third; in lateral view subparallel throughout, in 
dorsal view moderately strongly expanded laterally at antennal inser- 
tion, attenuate to middle, tip strongly expanded; dull, alutaceous, 
shallow punctures arranged in rows similar to male, tip smoother, 
slightly more shining, glabrous distad of antennal insertion. An- 
tennae inserted at basal fourth of beak at distance from eye one-half 


286 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


greater than width of frons; first segment equal in length to next three; 
second segment of male shorter than next two, of female more elongate, 
equal to next four segments combined; club 0.18 by 0.07 mm. Eyes 
moderately prominent; frons slightly wider than dorsal tip of beak, 
with a shallow median depression and one lateral row of punctures. 
Prothorax slightly wider at base than long, middle slightly narrower 
than base, apex three-fourths as wide as base; sides beyond basal 
lateral expansion slightly diverging to middle, rounded to constricted 
apex; in profile dorsal surface slightly arcuate; punctation 0.03 mm. 
in diameter, moderately deep, interspaces appear convex, alutaceous, 
range in width from one-half to equal to diameter of punctures; basal 
fovea moderately deep, extending one-third the length of the pro- 
thorax. Elytra at humeri slightly more than one-third wider than 
prothorax at base, 2.5 times as long as prothorax, length to width as 
9:7; elytral intervals somewhat convex, equal in width to two striae, 
with one row of punctures, punctures somewhat confused at base of 
interval 3, intervals smooth; striae fine, moderately deep. Scutellum 
elongate-triangular, 0.09 by 0.05 mm., with a moderate median furrow. 
Front femora 3.2 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 1 armed with minute, blunt micro; 
tibiae 2 and 3 armed with moderate, acutely dentate mucrones. 

Tyres: Holotype male (USNM 63416) from San Luis Potosf (near 
the town of Antiguo Morelos in Tamaulipas), Mexico, July 1954, 
D. G. Kissinger. Allotype female (USNM), same data as holotype. 
Forty-six paratypes, same data as holotype (24 in the collection of the 
author and 2 in each of the following: AMNH, BMNH, CAS, CIS, 
CNC, CU, INHS, UC, UK, UM, UMD). 

Remarks: It gives me much pleasure to name this species in honor 
of Dr. Ernest N. Cory in recognition of his years of service to ento- 
mology. 


Apion (Trichapion) imitator Wagner 


FIGURE 4,a-c 
Apion (Trichapion) imitator Wagner, Arch. Naturg. Berlin, vol. 78, p. 106, 1912. 


Description: Length, 2.25 mm.; width, 1.10 mm. 

Robust, black. Pubescence fine, white, sparse on dorsal surface, 
coarser and denser on sides of prothorax, mesothorax, and metathorax. 
Male beak shorter than head and prothorax, one-third longer than 
prothorax, slightly, evenly curved; in lateral view attenuate from 
antennal insertion to apex; in dorsal view expanded laterally at 
antennal insertion, strongly attenuate to apical third, tip slightly 
expanded, somewhat compressed in apical third; punctured and 
sparsely pubescent to near apex, tip smoother, bare, a line of con- 
fluent punctures above antennal insertion appears as a sulcus, extends 


SUBGENUS TRICHAPION—KISSINGER 287 


h 1 J k 

Ficure 4.—a-c, Apion imitator Wagner: a, lateral view of head and prothorax of male; 
b, dorsal view of head of male; c, mucro of tibia 3 of male. d-g, k, A. proclive LeConte: 
d, lateral view of head and prothorax of male; ¢, lateral view of head and prothorax of 
female; f, dorsal view of head of female; g, entire dorsal view of male, k, mucro of tibia 
3 of male. Ah, A. adaetum, new species, mucro of tibia 3 of male. 1, 7, 4. plectrocolum, 
new species: i, mucro of tibia 2 of male; 7, mucro of tibia 3 of male. Line equals 0.50 
mm. 


about one-third length of beak. Male antennae inserted at basal 
fourth of beak at distance from eye one-half greater than width of 
frons; first segment longer than next three, slightly shorter than next 
four; second segment about equal to next two; club 0.21 by 0.08 mm. 
Eyes large, prominent; frons slightly narrower than dorsal tip of 
beak. Prothorax at base one-third wider than long, middle narrower 
than base, apex two-thirds as wide as base; sides beyond basal lateral 
expansion nearly parallel to middle, rounded to constricted apex; in 
profile dorsal surface flat; punctation moderately deep, 0.03 mm. 
in diameter, interspaces less than diameter of punctures, alutaceous; 


288 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


basal fovea deep, punctiform. Elytra at humeri one-half wider than 
prothorax at base, nearly three times as long as prothorax, length to 
width as 3:2; intervals twice as wide as striae, nearly flat, with two 
rows of fine scales; striae moderately deep, fine. Scutellum elongate- 
triangular, 0.12 by 0.08 mm. Front femora 3.4 times as long as 
wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with dentate 
mucrones. 

MATERIAL EXAMINED: One male determined by Hans Wagner. 

Type Locauity: Chilpancingo, Guerrero, Mexico (BMNH). This 
locality was erroneously attributed to Guatemala by Wagner in his 
description. 


Apion (Trichapion) plectrocolum, new species 
FicurReE 4,i,j 


Description: Length, 2.12 mm.; width, 1.06 mm. 

Moderately robust. Black; pubescence white, moderately fine, 
sparse, coarser and denser on sides of mesothorax and metathorax. 
Beak of male shorter than head and prothorax combined, one-third 
longer than prothorax, slightly curved; in lateral view apical three- 
fifths nearly parallel-sided; in dorsal view moderately expanded at 
antennal insertion, attenuate to basal three-fifths, here somewhat 
compressed, expanded slightly, tip subparallel; basal four-fifths 
pubescent, punctured, a distinct sulcus in basal half above antennal 
insertion, tip bare, smoother. Antennae of male inserted slightly 
distad of basal fourth, at distance from eye one-fourth greater than 
width of frons; first segment slightly longer than next three, second 
segment shorter than next two, club 0.20 by 0.08 mm. Eyes large, 
moderately prominent; frons about as wide as dorsal tip of beak, with 
a moderately broad, shallow, median sulcus and one lateral row of 
punctures. Prothorax at base one-fourth wider than long, middle 
narrower than base, apex seven-tenths as wide as base; sides beyond 
basal lateral expansion nearly parallel to middle, rounded to con- 
stricted apex; in profile dorsal surface nearly flat; punctation about 
0.03 to 0.04 mm. in diameter, moderately deep, interspaces generally 
about one-half diameter of punctures, finely alutceous; basal fovea 
moderately deep, broader at base, extends about one-third length 
of prothorax. Elytra at humeri two-fifths wider than prothorax 
at base, 2.75 times as long as prothorax, length to width as 11:8.5; 
intervals slightly convex, twice as wide as striae, on disc of elytra with 
two more or less distinct rows of scales, on declivity with one row of 
slightly coarser scales; striae moderately deep, fine. Scutellum 
elongate-triangular, 0.12 by 0.06 mm., with a moderate, median 


SUBGENUS TRICHAPION—KISSINGER 289 


furrow. Front femora 3.3 times as long as wide. Claws with acute 
basal tooth. 

Special male characters: Tibia 2 armed with a short, blunt mucro; 
tibia 3 armed with a long, slender, curved mucro. 

Houotyres: Male (USNM 63420) from Guadalupe, Distrito Federal, 
Mexico, in Thomas Lincoln Casey Collection. 


Apion (Trichapien) proclive LeConte 
FicurE 4,d-g,k 


Apion proclive, LeConte, Rep. Expl. Surv. Mississippi to Pacific, vol. 12, pt. 3, 
1860, p. 53, 1857.—Smith, Trans. Amer. Ent. Soc., vol. 11, p. 58, fig. 14, 
1884.—Fall, Trans. Amer. Ent. Soc., vol. 25, p. 140, pl. 4, figs. 9, 9a, 9b, 15, 
1898. 

Apion crassinasum LeConte, Rep. Expl. Surv. Mississippi to Pacific, vol. 12, 
pt. 3, 1860, p. 53, 1857.—Smith, Trans. Amer. Ent. Soc., vol. 11, p. 58, 1884.— 
Fall, Trans. Amer. Ent. Soc., vol. 25, p. 141, 1898. 

Apion nunenmacheri Wagner, Nov. Zool, Tring, vol. 19, p. 97, figs. 1, 2, 1912.— 
Fall, Journ. New York Ent. Soc., vol. 26, p. 223, 1918. 

Description: Length, 1.70 to 2.75 mm.; width, 0.75 to 1.30 mm. 

Moderately robust. Black; pubescence conspicuous, fine, white, 
sparse, denser on sides of mesothorax and metathorax. Male beak 
slightly shorter than head and prothorax, slightly, evenly curved, 
moderately dilated laterally and ventrally at antennal insertion, 
attenuate to apical third, there nearly cylindrical; basal two-thirds 
punctured and pubescent, apex smoother, glabrous, shining. Female 
beak one-half longer than head and prothorax combined, moderately, 
evenly curved; slightly expanded laterally and ventrally at antennal 
insertion, attenuating to apical third, slightly expanded laterally at 
apex; finely punctured and alutaceous, nearly glabrous in front of 
antennal insertion. Antennae of male inserted at distance from eye 
one-third greater than width of frons, slightly distad of basal third; 
of female inserted at distance from eye about twice as great as width 
of frons, at basal fourth of beak; first segment as long as next three; 
second segment one-half longer than third, shorter than next two; 
club 0.24 by 0.10 mm.; dorsal margin of antennal scrobe subangulate. 

Eyes not prominent; frons of male as wide as dorsal tip of beak, of 

female wider than dorsal tip of beak; with moderate median sulcus 

and two lateral rows of punctures. Prothorax at base one-tenth to 
one-fifth wider than long, middle about as wide as base, apex two- 
thirds to three-fourths as wide as base; sides beyond slightly basal 
lateral expansion nearly parallel to middle, rounded to constricted 
apex; in profile dorsal surface slightly arcuate, flatter basally and 
apically; punctation deep, 0.02 to 0.03 mm. in diameter, interspaces 
less than diameter of punctures, alutaceous; basal fovea shallow, short. 


290 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Elytra at humeri one-third to two-fifths wider than prothorax at base, 
2.8 times as long as prothorax, length to width as 5:4; intervals 
twice as wide as striae, nearly flat, with one or two rows of punctures 
bearing fine scales; striae deep. Scutellum elongate-triangular, 
0.06 by 0.02 mm. Front femora 3.75 times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, dentate 
mucrones. 

MATERIAL EXAMINED: 200 specimens including types of A. proclive 
(female, MCZ 365, with gold disc signifying California, No. 1775), 
and A. crassinasum (male, MCZ 366, with dark blue disc signifying 
Oregon, No. 1774), and material determined by Hans Wagner as 
A. nunenmacheri. 

KNOWN DISTRIBUTION: 

Unirep States: Washington: Mt. Adams (USNM). Oregon: Orenco (USNM); 
15 miles southeast of Weston (GHN). Jdaho: Cassia Co., City of Rocks (UC). 
Wyoming: Yellow Stone National Park (USNM). Colorado: Mesa Verde 
(AMNH); Raymonds (UK). Utah: Ogden (USNM); Pintura (UK); Wasatch 
(USNM); Wellsville (UK). Nevada: Elko Co., Pequops Summit, 6,900 ft., 
22 miles east of Wells (BVD, DGK). California: Many records throughout the 
State. Arizona: Coconimo Co., 6 miles north of Jacobs Lake, 7,600 ft. (AMNH); 
11 miles north of Jacobs Lake, 5,500—6,500 ft. (AMNH). Cochise Co., Chiricahua 
Mts., Rustlers Park (AMNH). Flagstaff (USNM); Grand Canyon, 7,600 ft. 
(AMNH); Phoenix (AMNH); Prescott (AMNH); Santa Rita Mts. (USNM). 

Mexico: Durango: Tepehuanes (TLCC). Mezico: 23 miles south of Toluca 
(DGK). Puebla: 10 miles east of Puebla, 7,400 ft. (CIS). 

Brotoay: Material in the U.S. National Museum from Mt. Adams, 
Wash., and Orenco, Oreg., was reared from lupine seed pods. In the 
California Insect Survey Collection there is material associated with 
Tupinus sp. 

The geographical distribution of this species may seem unusual. 
Goldman (1951) classifies the Mexican localities as Upper Austral in 
character. Fall (1898) suggests that this species may range northward 
along the coast to Alaska but no material was seen from that area. 

A small series from near Jacobs Lake, Coconimo Co., Arizona, ex- 
hibits an interesting aberration of the male secondary sexual charac- 
ters. The front tibiae bear a minute but distinct mucro. While it is 
possible that this series represents another species it does not seem to 
because the principal characters of the male and female are those of 
this species. 

Remarks: Regarding the synonymy, A. crassinasum has line pri- 
ority over A. prociwe; they were described on the same page. Fall 
(1898) noted that the types represented male and female of the same 
species and chose to use proclive instead of crassinasum because “I 
have chosen to supress the latter name as being less characteristic of 


SUBGENUS TRICHAPION—KISSINGER 291 


the two.” Since Fall (1898) is the first reviser, his choice of A. pro- 
clive will stand. 

According to Fall (1918), A. nuwnenmacheri Wagner is only a minor 
variation of the typical A. proclive. Specimens determined by Wag- 
ner in the Nunenmacher Collection in the Chicago Museum of Natural 
History are typical A. proclive. 


Apion glyphicum Group 


Three very closely allied species comprise this group. Relation- 
ship of this group to the A. simile and A. proclive groups has already 
been discussed. The range of two species, A. glyphicum Sharp and 
A. vinosum Sharp, overlaps in southern Mexico. The third species, 
A. chuparosae Fall, is apparently confined to Baja California. 

As mentioned above the species are very close to each other and are 
hard to separate, however there is insuflicient evidence to warrant 
synonymizing the names. A. vinosum is distinct because of the light 
reddish brown elytra and the yellowish pubescence on the dorsal sur- 
face. The other two species are black, the elytra have a more or less 
distinct aeneous luster, and the pubescence of the dorsal surface is 
white. The beak of the male of A. chuparosae is a little more slender 
and elongate and is slightly longer than the prothorax. The beak of 
the male of A. glyphicum is shorter than the prothorax. 


Apion (Trichapion) chuparosae Fali 


Figure 5,e 
Apion chuparosae Fall, Trans. Amer. Ent. Soc., vol. 25, p. 141, pl. 4, fig. 16, 1898. 


Description: Length, 2.0 to 2.4 mm. 

Robust; black, elytra with slight aeneous luster. Pubescence fine, 
white, sparse, somewhat coarser and denser ventrally. Male beak 
slightly longer than prothorax, slightly, evenly curved, stout, attenu- 
ate from antennal insertion to tip; punctured and sparsely pubescent 
to apical fifth, with deep sulcus above antennal insertion in basal half. 
Antennae of male inserted at distance from eye one-half greater than 
width of frons, between basal fourth and third of beak; first and second 
segments each shorter than next two segments, club 0.21 by 0.09 mm. 
Kyes large, prominent; frons narrow, about as wide as dorsal tip of 
beak. Prothorax at base one-third wider than long, middle narrower 
than base, apex two-thirds as wide as base; sides beyond marked basal 
lateral expansion slightly converging to middle, rounded to constricted 
apex; in profile dorsal surface slightly arcuate in front of middle; 
punctation shallow, 0.03 mm. in diameter, interspaces generally less 
than diameter of punctures, alutaceous; basal fovea extending one- 
third length of prothorax, narrow, deep at middle. Elytra at humeri 


292 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


two-fifths wider than prothorax at base, 2.75 times as long as pro- 
thorax, length to width as 4:3; intervals twice as wide as striae, mod- 
erately convex, without transverse rugae, with two rows of fine punc- 
tures; striae deep, fine. Scutellum triangular, 0.08 by 0.06 mm. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with dentate mu- 
crones similar to those of A. glyphicum. 

Types: I hereby designate the lectotype of this species as the male 
specimen labeled La Chuparosa in the Fall Collection (MCZ 25081). 
Cotypes are in the California Academy of Sciences. 

MATERIAL EXAMINED: Lectotype and one male. 

KNOWN DISTRIBUTION: 

Mexico: Baja California: La Chuparosa; La Laguna, Sierra Laguna, Oct. 14, 
1941, Ross and Bohart (CAS). 


Apion (Trichapion) glyphicum Sharp 
Ficure 5,a—d 


Apion glyphicum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 52, 1890. 

Description: Length, 2.5 to 2.75 mm, 

Robust. Black, elytra and prothorax slightly aeneous; pubescence 
fine, white, sparse, on ventral surface and on sides of prothorax, 
mesothorax, and metathorax pubescence much coarser and denser, 
especially on the male. Male beak shorter than prothorax, slightly, 
evenly curved, moderately expanded laterally and ventrally at 
antennal insertion; in dorsal view attenuate to apical third, sides 
slightly diverging at apex; in lateral view attenuating from antennal 
insertion to apex; in basal two-thirds beak with a deep, lateral, longi- 
tudinal channel with finer punctures bearing fine scales, median dorsal 
line impunctate, tip smooth, slightly shining. Female beak about as 
long as head and prothorax, slightly, evenly curved, dorsally nearly 
parallel-sided throughout; in lateral view ventrally with slight expan- 
sion at antennal insertion, thence slightly attenuate to apex; supra- 
antennal groove evident in basal two-thirds, apical third smooth, 
impunctate, hardly shining. Antennae inserted at distance from eye 
one-half greater than width of frons, of male at basal third, of female 
at basal fourth of beak; first segment of male shorter than next two, 
of female equal to next two segments; second segment shorter than 
next two; club 0.24 by 0.10 mm. Eyes moderately prominent, of male 
slightly larger and more prominent; frons narrow, of female equal to 
width of dorsal tip of beak, of male about five-sixths as wide as tip of 
beak; trisulcate, with a deep median impunctate sulcus and a deep 
intermediate lateral sulcus with shallow punctures bearing scales, 
separated from median sulcus by a narrow, convex interval, and with a 


SUBGENUS TRICHAPION—KISSINGER 293 


b 
rangi, 


PAD J 
a) 


Figure 5.—a-d, Apion glyphicum Sharp: a, entire dorsal view of male; }, lateral view of 
head and prothorax of female; ¢, lateral view of head and prothorax of male; d, mucro of 
tibia 3 of male. ¢, A. chuparosae Fall, lateral view of head and prothorax of male. Line 
equals 0.50 mm. 


lateral raised convex interval bearing scales; frons abruptly declivitous 
behind eyes. Prothorax at base about one-fifth wider than long, 
middle slightly narrower than base, apex three-fourths as wide as base; 
base expanded laterally, sides from there to middle nearly parallel, 
then rounded to constricted apex; in profile dorsal surface nearly flat; 
punctation moderately deep, 0.03 mm. in diameter, interspaces 
irregular, usually less than diameter of punctures, alutaceous; basal 
fovea moderately deep, wider toward middle, extending one-third 
length of prothorax. Elytra at humeri one-third to two-fifths wider 
than prothorax at base, 2.3 times as long as prothorax, length to width 
as 13 : 10; intervals about 2.5 to three times as wide as striae, smooth, 
with two irregular rows of punctures bearing fine scales; striae moder- 
ately deep. Scutellum triangular, about as long as wide, 0.08 by 0.08 
mm., with a median, longitudinal sulcus. Front femora from 3.0 to 
3.3 times as long as wide. Claws with moderate, acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately long, 
broad mucrones which have an apical angulation. 


294 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


MATERIAL EXAMINED: One female determined by Sharp and 25 
other specimens. 


KNOWN DISTRIBUTION: 


Mexico: Veracruz: Jalapa (type BMNH); Cérdoba, Feb. 3, 1953, D. G. 
Kissinger (DGK). Morelos: Cuernavaca, June, Fenyes Collection (CAS). 
México: Temescaltepec, Bejucos, July 2, 1932, Hinton and Usinger (CAS). 
Michoacén: 20 miles east of Morelia, Mar. 7, 1953, D. G. Kissinger (DGK). 
Guerrero: Rio Balsas, Wickham (TLCC). 


Apion (Trichapion) vinosum Sharp 


Apion vinosum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, Cur- 
culionidae: Apioninae, vol. 4, pt. 3, p. 70, 1890. 

Description: Length, 2.17 to 2.50 mm. 

Moderately robust. Head, prothorax, scutellum, and antennae 
dark reddish brown; elytra, femora, tibiae, and tarsi lighter reddish 
brown; pubescence somewhat conspicuous, fine, yellow on dorsal 
surface, white laterally, on disc of prothorax and elytra sparse, slightly 
denser at base of elytral interval 3, moderately dense on sides of 
mesothorax and metathorax. Male beak short, stout, much shorter 
than head and prothorax, one-third longer than prothorax, moderately, 
evenly curved; in lateral view expanded ventrally at antennal inser- 
tion, more or less attentuate to tip; in dorsal view expanded laterally 
over antennal insertion, attenuate to apical third, slightly expanding 
to tip; punctures arranged in rows, deeper laterally, basal three-fourths 
sparsely pubescent, apical fourth bare, smoother, moderately shining. 
Female beak one-fifth longer than head and prothorax, seven-tenths 
longer than prothorax, moderately, evenly curved; in lateral view 
stouter in basal sixth, attenuate to apical two-thirds, there nearly 
parallel-sided; in dorsal view moderately stout to antennal insertion, 
attenuate to apical third, expanding to apex; finely punctured in rows, 
deeper laterally, very sparsely pubescent in basal third, apical two- 
thirds bare, apical third smoother, subimpunctate; beak moderately 
shining throughout. Antennae inserted at distance from eye equal to 
width of frons, of male at basal fourth of beak, of female at basal 
sixth; first segment slightly shorter than next two, second segment 
slightly shorter than next two, club 0.28 by 0.10 mm. Eyes moderately 
prominent; frons narrow, slightly wider than dorsal tip of beak, 
abruptly declivitous behind eyes. Prothorax at base three-tenths 
wider than long, middle narrower than base, apex three-fourths as 
wide as base, slightly converging to middle, roundly narrowing to 
constricted apex; in profile dorsal surface slightly arcuate; punctation 
0.03 mm. in diameter, moderately shallow, interspaces rather convex, 
slightly less than, to equal to, diameter of punctures; basal fovea 
moderately deep at middle, narrow, one-third length of prothorax. 


SUBGENUS TRICHAPION—KISSINGER 295 


Elytra two-fifths wider at humeri than prothorax at base, 2.4 to 2.8 
times as long as prothorax, length to width as 5 : 4; intervals twice as 
wide as striae, nearly flat, with one or two rows of fine punctures bear- 
ing fine scales; striae moderately coarse, deep. Scutellum triangular, 
0.10 by 0.10 mm., with slight median depression. Fore femora four 
times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with broad, short, 
angulate mucrones. 

MATERIAL EXAMINED: Male and female determined by Sharp, male 
and female determined by Hans Wagner, and three other specimens. 


KNOWN DISTRIBUTION: 


Mexico: Veracruz: Cérdoba; Orizaba (CAS). Guatemala: Chiacam; Zapote 
(BMNH); Guatemala City; Duenas (BMNH). 


Apion griseum Group 


The males of eight species grouped here have the front tibiae 
modified with a smooth, more or less polished area. The range of 
three species overlaps on the Atlantic Coast of the United States, they 
are A. sayi Gyllenhal, A. griseum Smith, and A. novellum Fall. A. 
sayi and A. nebraskense Fall overlap in the midwestern portion of the 
United States. In Mexico A. aurichalceum Wagner, A. godmant Wag- 
ner, and A. parcum, new species, have been collected in large series 
in the same area at the same time by the author. 

To aid in identifying these important weevils, a key to females is 
appended at the end of the discussion of the group. The male of A. 
godmani is distinctive because the front tibiae are abruptly deflexed 
near the apical third. The front tibiae of the males of the other 
species are straight. The males, with the exception of A. sayz, have 
a smooth polished area extending at least one-half the length of the 
front tibiae. The smooth area of A. sayi is about one-third as long 
as the tibia and bears a few, irregular striae. The modified area of 
the males of two species, A. nebraskense and A. novellum, is extremely 
finely and densely striate, the striations are so close together that the 
area appears opaque or irridescent. The modified area of the males 
of the other species exhibits distinct, separated striations. The tibial 
mucrones of A. nebraskense project in line with the long axis of the 
tibiae and the beak is strongly attenuate toward the apex; the tibial 
mucrones of A. novellum project at an angle with the tibiae and the 
beak is less strongly attenuate. Apion parcum, new species, is dis- 
tinct because of its small size and the relatively coarse, sparse punc- 
tures on the metasternum. The beak of A. aurichalceum is distinctly 
expanded laterally at the tip and is more closely punctured laterally 
than the remaining two species. The dorsal pubescence of A. grisewm 


483661—59——4, 


296 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Smith is slightly yellowish; laterally it is white, also the antennal club 
of the male is about twice as long as wide and is clothed with nearly 
uniform, long, erect pubescence. The pubescence of A. oriotes, new 
species, is unicolorous, nearly white throughout, and the antennal 
club is about three times as long as wide and is clothed with short and 
long pubescence intermixed. 

The A. griseum group is closely related to the A. simile and A. 
proclive groups but is very distinct because of the male secondary 
sexual modifications of the front tibiae. The A. spinitarse, A. punc- 
tulirostre, and A. submetallicum groups may have been derived from 
this group. 

The females of A. nebraskense and A. novellum are unknown. 


Key to females of Apion griseum group 


i. Beak distinctly pubescent distad of antennal insertion. ........ 2 
Beak nearly glabrous beyond antennal insertion. . . . . godmani Wagner 
2. Beak in dorsal view distinctly expanded at tip. ............ 838 
Beak in dorsal view not expanded at tip... . ts Orel 


Beak moderately, densely pubescent to near middle; dorsal Sans of pro- 
thorax and elytra clothed with yellowish pubescence becoming white 


co 


laterally... 4: . . . .aurichaleeum Wagner 

Beak sparsely BubeseenG! in pass Grind: ubeseence unicolorous, whitish to 
grayish “iy 31). 4 .. . . .sayi Gyllenhal 

4. Beak moderately derecly pnb eseann io, near peaiddlee . . oriotes, new species 
Beak sparsely pubescent in basal third. . . .. . zie onteae 


5. Size 1.60 mm.; metasternum comparatively coarsely, Avene panei 
parcum, new species 
Size 2.00 mm. or more; metasternum comparatively finely, densely punc- 
euTed!)-<(Toiets So lela. ee OAD SOUR wo 2) COL GS griseus enibn 


Apion (Trichapion) aurichalceum Waguer 
FicureE 6,a—d,l 


Apion (Trichapion) aurichaleeum Wagner, Arch. Naturg. Berlin, vol. 78, p. 103, 
1912.—McKelvy, et al., Mexico Sec. Agr. Ganad. Foll. Tech., vol. 8, p. 37, 
1951. 

Description: Length, 2.0 to 2.4 mm. 

Moderately robust; black, elytra strongly aeneous. Pubescence 
long, fine, yellow on dorsal surface, coarser and whitish laterally, some- 
what denser at base of elytral interval 3, much denser on sides of 
mesothorax and metathorax. Male beak slightly shorter than head 
and prothorax, one-third longer than prothorax; moderately curved; 
in lateral view apical two-fifths nearly parallel; in dorsal view mod- 
erately expanded at antennal insertion, attenuate to apical third where 
it is somewhat compressed, slightly expanded toward tip; sculpture 
laterally in basal two-thirds deeply, coarsely punctured, finely punc- 
tured dorsally, pubescent, moderately dull, slightly alutaceous, apical 


SUBGENUS TRICHAPION—KISSINGER 297 


third shining, finely punctured, glabrous. Female beak one-fourth 
longer than head and prothorax, slightly curved, subcylindrical, ex- 
panded laterally at tip, in basal three-fifths sparsely punctate in 
rows, sparsely pubescent, apex smoother. Antennae of male inserted 
at basal sixth of beak at distance from eye slightly less than width of 
frons, of female at distance from eye slightly less than width of frons 
at basal sixth of beak; first segment of male equals next two, of female 
slightly longer than next two; second segment equals next two; club 
0.21 by 0.09 mm. Hyes moderately prominent; frons wider than dorsal 
tip of beak, with two lateral rows of punctures on either side of 
median sulcus. Prothorax one-fourth to one-third wider at base than 
Jong, middle not as wide as base, apex two-thirds as wide as base; sides 
beyond basal lateral expansion nearly parallel to middle, rounded to 
constricted apex; in profile dorsal surface nearly flat; punctation 
moderately deep, 0.04 mm. in diameter, interspaces less than diameter 
of punctures; basal fovea shallow, broad, about one-third length of 
prothorax. Elytra at humeri one-third wider than prothorax at base, 
between 2.75 and 3.00 times as long as prothorax, length to width as 
4:3; intervals twice as wide as striae, nearly flat, smooth, with two 
rows of fine punctures; striae deep, fine. Scutellum triangular, 0.08 
by 0.06 mm. Front femora 3.5 times as long as wide. Claws with 
acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately long, 
slender, simple mucrones; tibia 1 with anterior inner face flattened, 
polished, longitudinally striate area along two-thirds of its length; 
tibia 1 not deflexed apically. 

MATERIAL EXAMINED: Male and female determined by Hans 
Wagner and 125 other specimens. 

KNOWN DISTRIBUTION: 

Mexico: In beans intercepted at Phoenix, Ariz., Brownsville, Tex., and Laredo, 
Tex. (USNM). Guerrero: Chilpancingo; Omilteme. Puebla: Huauchinango, 
June 1954, D. G. Kissinger (DGK); Puebla, Sept. 2, 1949, A. C. Smith (USNM). 
México: Tenancingo, Sept. 11, 1948, A. C. Smith (BMNH). Distrito Federal: 
Atzacapatalco, August 1921, H. F. Wickham, on bean (USNM); Contreras, May 
30, 1946, J. and D. Pallister (AMNH); San Jerénimo, June 11, 1946, J. and D. 
Pallister (AMNH); Tacuba, September 1921, H. F. Wickham, on bean (USNM); 
30 miles southeast of Mexico City, June 1954, D. G. Kissinger (DGK). 

GuaTEMALA: In black beans intercepted at Brownsville, Tex. (USNM); Acei- 
tuno; Capetillo; Chiacam. 

Remarks: This species apparently is a serious pest of the bean crop 
in Mexico. 

Apion (Trichapion) godmani Wagner 


FiGureE 6,9-7 


Apion (Trichapion) godmani Wagner, Arch. Naturg. Berlin, vol. 78, p. 101, 1912.— 
Estados Unidos Mexicanos, Secretaria de Agricultura y Fomento, Principales 


298 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


plagas y enfermedades de los cultivos de la Republica, Mexicana, pp. 84-85, 
1930.— Anon., Fitofilo, vol. 1, no. 2, p. 54, 1942; Fitofilo, vol. 2, no. 3, p. 111, 
1943.—McKelvey, et al., Mexico, Secr. Agri. Ganad. Foll. Tec., vol. 8, p. 11, 
figs. 1-5, 1951.—Smith, Coleopt. Bull., vol. 5; pp. 61-62, 1951. 

Description: Length, 1.75 to 2.21 mm.; width, 0.87 to 1.00 mm. 

Moderately robust. Black, elytra and prothorax with strong golden 
bronze luster; pubescence conspicuous, fine, yellow on dorsal surface, 
becoming white laterally and ventrally, sparse, coarser and denser 
laterally and ventrally. Male beak as long as head and prothorax 
combined, slightly, evenly curved; slightly expanded ventrally at 
antennal insertion, attenuate from there to apical third where it is 
nearly cylindrical; moderately punctate and pubescent in basal two- 
thirds, apical third smooth, shining. Female beak moderately 
strongly, evenly curved, about two-fifths longer than head and pro- 
thorax, nearly cylindrical throughout, with ventromedian angulation 
behind antennal insertion; moderately punctured and pubescent in 
basal fourth, smooth and dull to apex. Antennae of male inserted at 
distance from eye slightly less than width of frons at basal fifth of 
beak, of female at distance from eye equal to width of frons at basal 
fourth; first segment of male longer than next two, of female a little 
longer than next three combined; second segment a little longer than 
third; club 0.21 by 0.09 mm. Eyes moderately prominent; frons 
moderately wide, median sulcus lacking, with about six faint longi- 
tudinal rows of fine, nearly contiguous punctures and appearing faintly 
striate. Prothorax at base one-fourth wider than long, middle 
slightly narrower than base, apex two-thirds to three-fourths as wide 
as base; sides expanded laterally at base, nearly parallel to middle 
then rounded to constricted apex; in profile dorsal surface slightly, 
evenly convex; punctation moderate, 0.03 mm. in diameter, shallow, 
interspaces about equal to diameter of punctures, flat, alutaceous; 
basal fovea short, shallow, rounded. Elytra at humeri two-fifths 
wider than prothorax at base, 2.75 times as long as prothorax, length 
to width as 8.5: 11; intervals about twice as wide as striae, flet or 
slightly convex, finely alutaceous, with one or two irregular rows of 
fine punctures; striae deep, fine. Scutellum elongate, linear, about 
twice as long as wide, 0.15 by 0.07 mm., with median furrow. Front 
femora about 3.7 times as long as wide. Claws with mo‘lerate basal 
tooth. 

Special male characters: Tibiae 2 and 3 armed with moderate, 
simple mucrones; tibia 1 on inner side with a flat, smooth, longi- 
tudinally striate area extending from near base to apical one-fourth, 
from anterior view front tibiae appear to be compressed especially 
on inner side and are deflexed distad of apical two-fifths. 


SUBGENUS TRICHAPION—KISSINGER 299 


k 


Ficure 6.—a-d, 1, Apion aurichalceum Wagner; a, lateral view of head and prothorax of 
male; b, dorsal view of head of male; c, mucro of tibia 3 of male; d, dorsal view of head of 
female; J, tibia 1 of male. ¢, A. oriotes, new species, dorsal view of head of male. f, A. 
griseum Smith, dorsal view of head of female. g,-j, 4. godmani Wagner: g, dorsal view of 
head of female; h, lateral view of head and prothorax of male; i, lateral view of head and 
thorax of male; j, front leg of male. k, 4. sayi Gyllenhal, dorsal view of head of female. 
Line equals 0.50 mm. 


300 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


MATERIAL EXAMINED: Material determined by J. Balfour-Browne 
and 150 other specimens. 

KNOWN DISTRIBUTION: 

Mexico: On lima beans taken at Phoenix, Ariz., June 9, 1922, A. H. Caldwell, 
Jr. (USNM). Guerrero: Amula, 6,000 ft. (BMNH). Puebla: Near Huauchi- 
nango, June 1954, D. G. Kissinger (DGK); 14 miles west of Texmelucan, July 14, 
1953, 8,600 ft., University of Kansas expedition (UK). Hidalgo: 24 miles north- 
east of Jacala, June 22, 1953, University of Kansas expedition (UK). Veracruz: 
38 miles northeast of Cérdoba, June 30, 1953, University of Kansas expedition 
(UK). México: 7 miles south of Manzamitla, July 1, 1948, E. S. Ross (CAS); 
Real de Arriba, Temescaltepec, June 11, 1933, 8,000 ft., E. Hinton and R. L. 
Usinger (CAS). Distrito Federal: Guadalupe, H. F. Wickham (TLCC); in 
string beans from Mexico City taken at Brownsville, Tex., Sept. 14, 1946 (USNM); 
Atzcapotxalco, Aug. 16, 1921, on bean, H. F. Wickham (USNM); Tacuba, on 
bean (USNM). 

GuatTEMALA: Aceituno; Calderas; Quezaltenango, 7,800 ft., Voleén de Agua; 
Zapote (BMNH). 

Remarks: This species is a serious pest on bean crops near Mexico 
City. 


Apion (Trichapion) griseum Smith 
FiGurRE 6,f 
Apion griseum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 59, 1884.—Fall, Trans. 
Amer. Ent. Soe., vol. 25, p. 147, 1898.—Kissinger, Proc. Ent. Soc. Wash- 
ington, vol. 59, p. 40, 1957. 

Many references in the literature to this species actually refer to 
Apion sayi Gyllenhal. These will be listed and reviewed under the 
biology of the appropriate species. 

Description: Length, 2.00 to 2.35 mm.; width, 0.93 to 1.12 mm. 

Moderately robust. Black, aeneous; pubescence conspicuous, fine, 
white, with yellowish tinge on dorsal surface, sparse, denser on sides 
of mesothorax and metepisternum. Male beak equal to head and 
prothorax in length, slightly, evenly curved, moderately expanded 
laterally and slightly expanded ventrally at antennal insertion, 
attenuate to apical third which is nearly cylindrical; coarsely, sparsely, 
punctured, basal two-thirds with bristling pubescence, apex glabrous, 
shining. Female beak about one-sixth longer than head and pro- 
thorax, moderately, evenly curved, slightly angulate laterally at 
antennal insertion, attenuate to middle, apical half nearly cylindrical; 
basal third with coarse sparse punctures and short pubescence, apical 
two-thirds smoother, dull, glabrous. Antennae inserted at distance 
from eye slightly less than width of frons, of male at basal fourth, of 
female at basal fifth; first segment as long as next two, second segment 
slightly (female) to much (male) shorter than next two, club 0.21 by 
0.10 mm. Eyes moderately prominent; frons moderately wide, with 
deep, median sulcus and two lateral rows of fine, nearly or quite 


SUBGENUS TRICHAPION—KISSINGER 301 


contiguous punctures. Prothorax one-third wider at base than 
long, middle slightly narrower than base, apex two-thirds to three- 
fourths as wide as base, sides expanded laterally at base, slightly 
converging to middle, rounded to constricted apex; in profile dorsal 
surface slightly convex, flattened basally and apically; punctures 
moderate, 0.03 mm. in diameter, deep, interspaces about two-thirds 
as great as diameter of punctures, flat, alutaceous, wider and flatter 
in lateral basal third, basal fovea deep, short, rounded. Elytra at 
humeri one-third wider than base of prothorax, 2.75 to 3.00 times as 
long as prothorax, length to width as 12:9; intervals twice as wide 
as striae, flat, with two rows of fine punctures; striae deep, fine. 
Scutellum elongate-triangular, about twice as long as wide, 0.12 by 
0.06 mm., with a shallow, median furrow. Front femora about 3.75 
times as long as wide. Claws with moderate basal tooth. 

Special male characters: Tibiae 2 and 3 armed with small, simple 
mucrones; tibia 1 slightly flattened on the anterior inside surface, 
area polished with moderately close striations. 

Lectotype: Kissinger (1957) designated as lectotype of this species 
a male specimen (USNM 1253) labeled New Jersey. 

MATERIAL EXAMINED: Lectotype and 50 other specimens. 

KNOWN DISTRIBUTION: 

Unitep States: Alabama: Selma, Dec. 18, 1895, H. Soltau (USNM). District 
of Columbia (USNM). Florida: Crescent City, Hubbard and Schwarz (USNM). 
Georgia: Atlanta, Nov. 3, 1937, P. W. Fattig (USNM); Experiment, Nov. 12, 
1936, T. L. Bissell (USNM). Maryland: Piummers Island, June 17, 1906, W. L. 
McAtee (USNM). New Jersey (USNM). New York: Ithaca (USNM), Peekskill, 
on rhododendron flowers in July, Sherman (USNM). North Carolina: Round 
Knob, Hubbard and Schwarz (USNM). Virginia: Alexandria, Oct. 30, 1955, 
A. W. Vazquez; Barcroft, September 1931, J. C. Bridwell, ex seeds of Phaseolus 
polystachys (USNM); Bull Run, ex Phaseolus polystachys (USNM); Pennington 
Gap, Hubbard and Schwarz (USNM). 

Remarks: This species has been reared several times from the seed 
pods of Phaseolus polystachys. Pierce (1908), Chittenden (1908), 
Blatchley and Leng (1916), Wickham (1922), Bissell (1938), and 
Tuttle (1954) cite various species of Phaseolus as the host plants of this 
species. All except western records probably refer to A. griseum. 
The records of A. grisewm developing in beans of the genus Stropho- 
styles doubtless refer to A. say Gyllenhal. 


Apion (Trichapion) nebraskense Fall 


FiGuRreE 7,d,e 
Apion nebraskense Fall, Trans. Amer. Ent. Soc., vol. 25, p. 145, pl. 4, fig. 20, 1898. 
Description: Length, 1.75 mm.; width, 0.89 mm. 


Moderately robust. Black; pubescence fine, white, sparse, a little 
more conspicuous ventrally. Male beak shorter than head and pro- 


302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


thorax combined, two-fifths longer than prothorax, deflexed at basal 
third, in dorsal view attenuate from laterally expanded antennal in- 
sertion to near apex; basal two-thirds dull, alutaceous, sparsely punc- 
tured, apical third smooth, shining. Antennae of male inserted at 
basal third of beak at distance from eye one-third greater than width 
of frons; first and second segments each shorter than next two, club 
0.17 by 0.07 mm. Eyes moderately prominent; frons wider than dorsal 
tip of beak. Prothorax at base one-fifth wider than long, middle 
narrower than base, apex three-fourths as wide as base; sides beyond 
slight basal lateral expansion nearly parallel to middle, rounded to 
constricted apex; in profile dorsal surface nearly flat; punctation 
moderately deep, 0.02 mm. in diameter, interspaces about equal to 
diameter of punctures; basal fovea shallow, punctiform. Elytra at 


Ap ila 


Ficure 7.—a-c, Apion parcum, new species: a, lateral view of head and prothorax of male; 
b, lateral view of head and prothorax of female; c, mucro of tibia 3 of male. d, e, A. 
nebraskense Fall; d, mucro of tibia 3 of male; e, lateral view of head and prothorax of 
male. Line equals 0.50 mm. 


humeri one-third wider than prothorax at base; nearly three times as 
long as prothorax, length to width as 15: 12.5; intervals about twice 
as wide as striae, nearly flat, with one row of fine scales; striae moder- 
ately deep, fine. Scutellum triangular, 0.07 by 0.06 mm., with a 
shallow median furrow. Front femora about 3.3 times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, slender, 
simple mucrones that project in line with the long axis of the tibiae; 
tibia 1 with a smooth area extending two-thirds its length on front 
inner surface, area iridiscent in certain light because of extremely fine, 
densely placed striations. 

Female is unknown. 


MATERIAL EXAMINED: Type (MCZ 25109, in Fall Collection labeled 
Nebraska) and four specimens. 


SUBGENUS TRICHAPION—KISSINGER 303 


KNOWN DISTRIBUTION: 


Unirep States: Nebraska: No exact locality (MCZ), Platte River southeast of 
Grand Isle, June 21, 1953, B. and B. Valentine (BDV). Colorado: Boulder, 
T. D. A. Cockerell (MCZ). Oklahoma (AMNH4). 


Apion (Trichapion) novellum Fall 
Apion novellum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 144, pl. 4, fig. 19, 1898. 


Description: Length, 1.50 mm.; width, 0.75 mm. 

Moderately robust. Black; pubescence white, moderately fine, con- 
spicuous, much coarser and scale-like on sides of prothorax, meso- 
thorax and metathorax. Male beak slightly shorter than head and 
prothorax combined, one-third longer than prothorax, moderately, 
evenly curved; slightly expanded laterally and ventrally at antennal 
insertion, attenuate to apical third which is nearly cylindrical; basal 
two-thirds sparsely punctured and pubescent; apical third smoother, 
shining. Antennae of male inserted at distance from eye equal to 
width of frons, at basal one-fourth of beak; first segment equals next 
two, second segment slightly longer than third, club 0.15 by 0.06 mm. 
Eyes moderately prominent; frons wider than dorsal tip of beak. 
Prothorax at base one-third wider than long, middle narrower than 
base, apex three-fourths as wide as base, sides beyond basal lateral 
expansion nearly parallel to middle, rounded to constricted apex; in 
profile dorsal surface nearly flat; punctation moderately deep, 0.03 
mm. in diameter, interspaces less than diameter of punctures; basal 
fovea shallow, short. Elytra at base one-third wider than prothorax 
at base; 2.5 times as long as prothorax, length to width as 7.5 : 6; inter- 
vals moderately convex, twice as wide as striae, with one row of scales 
and with minute transverse rugae; striae deep, fine. Scutellum trian- 
gular, 0.06 by 0.04 mm., with slight median furrow. Front femora 3.5 
times long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with a long, fine mucro 
which is turned inward in apical half, tibia 3 armed with a long, 
straight, nearly simple mucro, tibia 1 with long, narrow, flattened 
polished area on anterior inner surface; area shines with silky luster 
due to extremely fine, densely placed striations. 

Female not available for study. 

Typxrs: I hereby designate the iectotype of this species as the male 
specimen in the Fall Collection labeled Washington, D. C. (MCZ 
25110). A cotype with the same data is in the U.S. National Museum 
(USNM 4236). 

MatTERIAL EXAMINED: Lectotype, cotype, and one specimen. 


KNOWN DISTRIBUTION: 


Unirep States: District of Columbia. Tennessee: East Ridge, Chapin 
Sanctuary, May 8, 1952, C. Peck (CNC). 


304 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Remarks: The mucro of the second pair of tibiae is similar to that 
of A. innocens, new species, from Mexico. 


Apion (Trichapion) criotes, new species 
FIGURE 6,¢ 


Description: Length, 1.75 to 2.37 mm.; width, 0.75 to 1.12 mm 

Moderately robust. Black, not aeneous; pubescence conspicuous 
unicolorous, white, fine, somewhat coarser laterally and ventrally 
denser on mesothorax and metathorax. Male beak as long as head 
and prothorax combined, one-half longer than prothorax, slightly, 
evenly curved; apical one-third nearly cylindrical, in dorsal view 
attenuate from moderately expanded antennal insertion to near 
apical third; basal two-thirds dull, alutaceous, punctured, pubescent, 
apical third shining, subimpunctate, glabrous. Female beak one- 
third longer than head and prothorax combined, about twice as 
Jong as prothorax, moderately, evenly curved; apical half nearly 
cylindrical, in dorsal view attenuate in front of antennal insertion; 
dull, alutaceous, basal half pubescent, more strongly punctured, 
apical half shallowly, sparsely punctured. Antennae of male inserted 
at basal sixth of beak at distance from eye slightly less than width 
of frons, of female at basal fifth at distance from eye slightly greater 
than width of frons; first segment of male equals next two, of female 
next three; second segment shorter than next two; club 0.21 by 0.08 
mm. Hyes prominent; frons wider than dorsal tip of beak, with slight 
median furrow and three uregular lateral rows of fine punctures. 
Prothorax at base one-third wider than long, middle narrower than 
base, apex about two-thirds as wide as base; sides beyond basal 
lateral expansion nearly parallel to middle, rounded to constricted 
apex; in profile dorsal surface slightly arcuate; punctation 0.03 mm. in 
diameter, moderately deep, interspaces generally narrower than diam- 
eter of punctures, alutaceous; basal fovea moderately deep, about 
one-third length of prothorax. -Elytra at humeri one-third wider 
than prothorax at base, three times as long as prothorax, length to 
width as 12:8.5; intervals twice as wide as striae, nearly flat, with 
two rows of fine punctures, surface rather dull, with fine transverse 
rugae; striae moderately deep, fine. Scutellum triangular, 0.09 by 
0.06 mm., with slight median furrow. Front femora 3.3 times as 
long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, nearly simple mucrones, tibia 1 with inner anterior face bearing 
a flattened, polished, striate area extending two-thirds of its length. 

Types: Holotype male (USNM 63417), Silver City, N. Mex., 
Sept. 25, 1899, Metcalfe, in seeds of Phaseolus retusa. Allotype 
female (USNM), same data as holotype. Sixty paratypes: 25, same 


SUBGENUS TRICHAPION—KISSINGER 305 


data as holotype (USNM); 4, Chiricahua Mts., Ariz., Hubbard 
and Schwarz (USNM); 1, Fort Grant, Ariz. (USNM); 1, Huachuca 
Mts., Ariz. (USNM); 9, Arizona (USNM); 20, Sunnyside Canyon, 
Huachuca Mts., Ariz., July 9, 1940, D. E. Hardy (UK). 

Remarks: The host plant of this species is evidently Phaseolus 
retusa. 


Apion (Trichapion) parcum, new species 


Figure 7,a-c 


Description: Length, 1.44 to 1.75 mm.; width, 0.68 to 0.87 mm. 

Moderately robust. Black, elytra somewhat aeneous; pubescence 
on dersal surface yellowish, fine, sparse, somewhat coarser and white 
at base of interval 3, laterally and ventrally white, slightly coarser 
there, not noticeably denser than on dorsal surface. Male beak 
slightly shorter than head and prothorax combined, one half longer 
than prothorax, moderately, evenly curved; in lateral view apical 
half nearly parallel-sided; dorsal view with moderate lateral expansion 
over antennal insertion, attenuate to beyond middle, slightly com- 
pressed, then expanding slightly, apical fourth parallel-sided; basal 
half dull, alutaceous, sparsely pubescent, sparsely punctured, with 
slight furrow above antennal insertion, apical half shining, glabrous, 
very sparsely punctured. Female beak nearly one-half longer than 
head and prothorax combined, twice as long as prothorax; moderately, 
evenly curved; nearly cylindrical throughout, slightly expanded 
laterally at antennal insertion, dull, alutaceous, with a row of 
punctures above antennal insertion, with fine, sparse scales in front 
of antennal insertion. Antennae inserted at basal fifth of beak at 
distance from eye slightly less than width of frons of male, of female 
inserted at basal sixth of beak at distance from eye equal to width 
of frons; first segment of male equals next two, of female equals next 
three; second segment of male shorter than next two, of female equal 
to next two; club 0.15 by 0.06 mm. Eyes moderately prominent; 
frons wider than dorsal tip of beak, with slight median furrow and 
two irregular lateral rows of fine punctures. Prothorax at base one- 
fourth wider than long, middle narrower than base, apex five-sevenths 
as wide as base; sides beyond basal lateral expansion nearly parallel 
to middle, rounding to constricted apex; in profile dorsal surface nearly 
flat; punctation 0.02 to 0.03 mm. in diameter, moderately deep, 
interspaces generally narrower than diameter of punctures, alutaceous; 
basal fovea moderately deep, short. Elytra at humeri one-third wider 
than prothorax at base, 2.75 times as long as prothorax, length to 
width as 15 : 11; intervals nearly flat, somewhat convex toward base, 
in part with two rows of fine punctures bearing scales, smooth, trans- 
verse rugae fine, sparse, not conspicuous; striae fine, moderately deep. 


306 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Scutellum triangular, 0.06 by 0.04 mm., with slight median furrow. 
Front femora 3.5 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with slender, 
moderately long, nearly simple mucrones; tibia 1 with smooth, 
polished, striate area along five-eighths of its length, anterior margin 
of front tibia somewhat convex. 

Typxs: Holotype male (USNM_ 63418), near Huauchinango, 
Puebla, Mexico, June 1954, D. G. Kissinger. Allotype female 
(USNM), same data as holotype. Paratypes (405): 100, same 
data as holotype (USNM); 287, same data as holotype (DGK); 2, 
30 miles southeast of Mexico City, in México, July 1954, D. G. Kissin- 
ger (DGK); 7, Tacuba, México, on bean, H. F. Wickham (USNM); 
2, Tlalpam, México, July 14, 1923, on bean, E. G. Smyth (USNM); 3, 
Atzcapotzalco, México, Aug. 17, 1921, on bean,*H. F. Wickham 
(USNM); 1, Mexico City, Distrito Federal, June 12, 1922, E. G. 
Smyth (USNM). 

Remarks: This is Apion sp. No. 3 of the series determined by 
H. C. Fall referred to by Wickham (1922). 

This species was abundant on the foliage of pine and a scrubby oak 
near Huauchinango, Puebla, Mexico, in June, at an altitude of about 
5,000 feet. 


Apion (Trichapion) sayi Gyllenha! 
Figure 6,k 


Apion sayi Gy!lenhal, in Schoenherr, Genera et species curculionidum, vol. 1, 
p. 252, 1833. 

Apion fraternum Smith, Trans. Amer. Ent. Soc. vol. 11, p. 60, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 147, 1898.—Kissinger, Proc. Ent. 
Soc. Washington, vol. 59, p. 40, 1957. New synonymy. 

Many references in the literature to Apion griseum Smith actually 
should refer to this species. These will be mentioned below in the 
discussion of the biology of this species. 

Description: Length, 1.88 to 2.44 mm.; width, 0.94 to 1.25 mm. 

Moderately robust. Black, elytra with more or less aeneous luster; 
pubescence white, fine, evident, coarser and more apparent on sides 
of prothorax, mesothorax, and metathorax. Male beak shorter than 
head and prothorax combined, one-third longer than prothorax, moder- 
ately, evenly curved; in lateral view stout and parallel at base, at- 
tenuate to apical third, thence parallel-sided to apex; in dorsal view 
stout and parallel at base, not expanded laterally at antennal inser- 
tion, attenuate to apical third, slightly expanded toward tip; punc- 
tured and pubescent in basal two-thirds, apical third glabrous, smooth, 
shining. Female beak generally longer than head and prothcrax com- 
bined, ranges from one-half to two-thirds longer than prothorax, 
moderately strongly, evenly curved; in lateral view apical half nearly 


SUBGENUS TRICHAPION—KISSINGER 307 


parallel-sided; in dorsal view stout and parallel at base, attenuate to 
apical third, distinctly expanded to tip; dull and alutaceous throughout, 
basal half punctured, more or less sparsely pubescent, apical third 
smoother, more shining. Antennae inserted at distance from eye less 
than width of frons, of male at basal fifth of beak, of female at basal 
seventh; first segment equals next two, second segment shorter than 
next two; club of male 0.18 by 0.09 mm.; of female 0.20 by 0.08 mm. 
Eyes prominent; frons much wider than dorsal tip of beak, with 
shallow median furrow and two lateral rows of punctures. Prothorax 
at base one-fourth wider than long, middle narrower than base, apex 
ranges from 0.67 to 0.70 as wide as base; sides beyond basal lateral 
expansion nearly parallel to middle, rounded to strongly constricted 
apex; in profile dorsal surface slightly arcuate; punctation moderately 
deep, 0.03 mm. in diameter, interspaces about one-half diameter of 
punctures; basal fovea moderately deep, short. Elytra at humeri 
two-fifths wider than prothorax at base, 2.8 times as long as prothorax, 
length to width ranges from 10:7 to 11 :8.5; intervals nearly flat, 
twice as wide as striae, rather smooth and polished, with two rows of 
fine punctures; striae moderately deep, fine. Scutellum triangular, 
0.08 by 0.06 mm. Front femora about 3.5 times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with small, simple 
mucrones, tibia 1 slightly flattened for about one-third its length on 
anterior inner surface, area devoid of scales and at most slightly striate. 

Typss: Kissinger (1957) designated as lectotype of A. fraternum 
the female specimen (USNM 1252) labeled Columbus, Tex. Co- 
types (MCZ 375) are in the J. L. LeConte Collection with the same 
data. 

MATERIAL EXAMINED: Lectotype of A. fraternum; type of A. sayz in 
Riksmuseum labeled “male, Ap. rostrum Say, Am. bor. Say; Typus’’; 
the ‘“allotype” of A. sayi, which is also a male labeled “female, Ap. 
rostrum?, Say, N. Carolina, Harris; Allotypus”’; and more than 500 
specimens. 

KNOWN DISTRIBUTION: 

Unitep States: Alabama: Chambers Co., Langdale, H. H. Smith (USNM). 
Arkansas: Dardanelle, Apr. 8, 1905, on wheat, G. I. Reeves, Webster No. 2907 
(USNM); Fayetteville, Aug. 16, 1901 (USNM). District of Columbia, J. B. 
Smith Collection, cotype of A. griseum (USNM). Florida: Chittenden Collec- 
tion (USNM). JTllinots: Many records throughout the State: Indiana: Tip- 
pecanoe Co., July 4, 1952, N. M. Downie (DGK). Jowa: Independence, May 
15, H. F. Wickham (USNM); Iowa City, May 19, 1917, L. L. Buchanan 
(USNM); Lake Okobija, July 1916, L. L. Buchanan (USNM). Kansas: Coffey- 
ville, June 22, 1939, L. W. Hepnor (UK); Douglas Co., 900 ft., F. H. Snow (UK); 
Hutchinson, May 2, 1948, R. H. Beamer (UK); Lawrence, June, twilight and 
night, J. S. Tucker (USNM); Reno Co., Aug. 13-20, 1917 (UK); Sedgwick Co., 


308 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


1,291 ft.,R. H. Beamer (UK); Topeka, Popenoe (USNM). Kentucky: Fulton, G. 
I. Reeves (USNM). Louisiana: Shreveport, Mar. 25, 1908, R. A. Cushman 
(USNM). Maryland: Beltsville, June 6, 1920, L. L. Buchanan (USNM); Dor- 
chester Co., near Lloyds, July 10, 1907, H. S. Barber (USNM); Glen Echo, sum- 
mer 1922, J. C. Bridwell (USNM); southern Anne Arundel Co., October 1931, 
J. C. Bridwell, ex Strophostyles helvola seeds (USNM). Missouri: Kansas City, 
June 16, 1920, Warwick Benedict (UK). Nebraska: Aurora, Hamilton Co., June 
21, 1953, B. D. Valentine, roadside sweeping (BDV); Lincoln Co., Brady, June 
21, 1953, B. D. Valentine, on Helianthus (BDV); Grand Island, June 21, 1953, 
B. D. Valentine, sweeping Helianthus (BDV). New Jersey: Many records 
throughout the State. New York: Oakwood, Sept. 8, 1944, Tuthill and Mills, 
ex Strophostyles helvola seeds (USNM). Ohio: Athens, May 13, 1935, W. Stehr 
(ELS); Hocking Co., May 27, 1950, N. J. and E. L. Sleeper (ELS). Oklahoma: 
Harrah, May 23, 1916, W. D. Pierce, on oak (USNM); Lake Texoma, July 15, 
1954, University of Kansas expedition (UK); Randlett, May 25, 1916, W. D. 
Pierce, on Hicoria (USNM). Pennsylvania: Angora, June 15, G. M. Greene 
(USNM); Glenolden, June 16, G. M. Greene (USNM). South Carolina: Isle of 
Palms, Sept. 14, 1944, in seed pods of Strophostyles umbellata (USNM). Tennessee: 
Memphis, May 20, 1916, on Ostrya, W. D. Pierce (USNM). Texas: Columbus, 
Schwarz collector (USNM). Wyoming: 8 miles east of Cheyenne, 6,100 ft., 
June 22, 1953, B. D. Valentine, sweeping roadside (BDV). 


Remarks: There are several series in the U. S. National Museum 
reared from the seed pods of Strophostyles helvola; also two smali series 
from Strophostyles umbellata. All Apion material seen from the 
United States that was reared from seed pods of beans of the genus 
Strophostyles belonged to this species. 

Tuttle (1954) cites A. grisewm Smith in association with Stropho- 
styles leiosperma, in Illinois, but this should probably be A. sayi. 
None of Tuttle’s material was seen. Several hundred specimens of 
A. sayt were seen from Illinois; no specimens of A. griseum were seen 
from that State. Chittenden (1908) cites Strophostyles pauciflora as 
a host of A. griseum but this certainly refers to A. sayi. Material 
has been seen from Nebraska and Wyoming that was collected while 
sweeping Helianthus sp. 

The name A. sayi has erroneously been considered a synonym of 
A. rostrum Say and is listed as such in the Leng catalog. I have not 
been able to locate the authority for this synonymy but it is probably 
LeConte (1876). I have been privileged to see the type of this 
species through the kindness of Dr. René Malaise. 


Apion spinitarse Group 


This group is based on the unusual secondary male sexual modifica- 
tions of the front legs of its members. Generally only the middle 
tibia of the male is mucronate. The geographical range of the species 
is very incompletely known. It is thought that most of the species 


SUBGENUS TRICHAPION—KISSINGER 309 


overlap in range or occur in the same area. The group ranges from 
central Mexico into Brazil. 

The male of Apion sancti-felicis Sharp has the inner margin of the 
first segment of tarsus 3 produced into a short spine; the front legs are 
simple. The front tarsi of the males of two species, A. caenum, new 
species, and A. innocuum, new species, are not modified. The front 
femur of the former is distinctly swollen. The front legs of the latter 
are not modified; the species is grouped here because its general facies 
are similar to those of the other species and because only tibia 2 is 
mucronate. 

The male of A. mirandum, new species, is distinct because the first 
segment of tarsus 1 is dilated and greatly elongate being about four 
times as long as wide. The male of A. mirificum, new species, has 
the inner margin of the first segment of tarsus 1 produced into a large, 
rounded lobe. 

The first segment of tarsus 1 of the males of the following species 
has the inner margin produced into a spine: A. brachycephalum 
Wagner, A. calearatipes Sharp, A. enoplus, new species, A. innocens, 
new species, A. latitator, new species, and A. spinitarse Waener. 

Together with A. spinitarse, the males of A. enoplus and A. calcara- 
tipes have tibia 3 mucronate. The spine on the first segment of tarsus 
1 of the latter two species is much smaller and not flat as in A. spini- 
tarse. -A. enoplus is smaller and the elytral intervals are clothed with a 
single row of uniform scales. A. calcaratipes is larger than A. enoplus 
and the elytral intervals are clothed with two rows of seales which 
vary in coarseness and color along the length of the elytra. 

The males of the remaining three species have tibia 3 unarmed and 
the spine on the first segment of tarsus 1 much smaller than A. 
spinitarse. A. latitator is distinct with its parallel-sided beak. The 
beak of A. innocens and A. brachycephalum is attenuate toward the 
apical third. The dorsal surface of the beak of A. innocens is clothed 
with scales that are shorter and coarser than those on the dorsal 
surface of the prothorax. The dorsal surface of the beak of A. 
brachycephalum is clothed with scales similar to those on the dorsal 
surface of the prothorax. 

This group is very closely allied to the A. griseum group. The 
species A. enoplus supports this veiw because of its intermediate 
male characters. One difference is that the frons of this group tends 
to be comparatively narrower than that of the A. grisewm group, but 
it is never narrower than the dorsal tip of the beak. Also the size 
is smaller, always under 2.0 mm. and the general facies are somewhat 
more compact. 


310 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Apion (Trichapion) brachycephalum Wagner 
Ficurs 8,a,b 


Apion (Trichapion) brachycephalum Wagner, Arch. Naturg. Berlin, vol. 78, 
p. 109, 1912. 

Description: Length, 1.94 mm. 

Moderately robust. Black, with slight brassy luster, antennae 
piceous; pubescence white, on dorsal surface of prothorax and elytra 
very fine and sparse, coarse and dense on sides of mesothorax and 
metathorax. Male beak stout, shorter than head and prothorax 
combined, one-fifth longer than prothorax, moderately strongly, 
evenly curved; in lateral view very slightly attenuate to tip; in dorsal 
view slightly expanded at antennal insertion, attenuate to apical two- 
fifths, tip slightly expanded; with dorsal median, narrow, impunctate 
line extending from base to apical two-fifths, at basal two-fifths dilated 
into a diamond-shaped ridge thence attenuating to a fine, raised line; 
slightly shining, laterally with several rows of coarse punctures, tip 
smoother, pubescent to near tip. Antennae inserted at distance from 
eye equal to width of frons; first segment equals next two, second 
segment shorter than next two, club 0.24 by 0.09 mm. Eyes moder- 
ately prominent; frons narrow, as wide as dorsal tip of beak, with a 
shallow, wide, median sulcus, top of frons with several coarse punc- 
tures. Prothorax at base one-fourth wider than long, middle narrower 
than base, apex three-fourths as wide as base; sides beyond basal 
lateral expansion rounding to constricted apex; in profile dorsal surface 
is nearly flat; punctation on dorsal surface moderately deep, 0.03 to 
0.04 mm. in diameter; interspaces irregular, narrower than diameter 
of punctures; basal fovea moderately deep, punctiform. Elytra at 
humeri one-third wider than prothorax at base, 2.5 times as long as 
prothorax, length to width as 10:7.5; intervals nearly flat, twice as 
wide as striae, with a single row of fine punctures bearing fine scales; 
striae deep. Scutellum triangular, 0.12 by 0.06 mm. Front femora 
three times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with short, blunt mucro; 
tibia 1 with anterior inside surface bearing polished, nonstriate 
area; first segment of tarsus 1 with inner margin produced into a 
short spine. 

MATERIAL EXAMINED: One male determined as a female by Hans 
Wagener. 

Known distribution: 

British Honpuras: Belize and Blancaneaux (BMNH). 

GuaTEMALA: Zapote (BMNH). 

Remarks: There is some question regarding the correctness of 
Wagner’s determination of the specimen seen. The species was 


SUBGENUS TRICHAPION—KISSINGER 311 


m nN 


Ficure 8.—a, b, Apion brachycephalum Wagner: a, entire dorsal view of male; }, lateral 
view of head and prothorax of male. c, d, 4. caenum, new species: c, front femur of male; 
d, mucro of tibia 2 of male. ¢, f, 7, A. enoplus, new species: ¢, mucro of tibia 3 of male; 
f, mucro of tibia 2 of male; 7, lateral view of head and prothorax of male. g-i, A. cal- 
caratipes Sharp: g, entire dorsal view of male; A, lateral view of head and prothorax of 
male; 2, lateral view of head and prothorax of female. k-n, 4. innocens, new species: 
k, dorsal view of head of male; /, lateral view of head and prothorax of male; m, mucro of 
tibia 2 of male; , tarsus 1 of male. Line equals 0.50 mm. 

483661—59—_5 


al2 PROCEEDINGS OF THE NATIONAL MUSEUM . VOL, 110 


described as having tibiae 2 and 3 mucronate but the specimen seen 
did not appear to have tibia 3 mucronate. 


Apion (Trichapion) caenum, new species 
FIGuRE 8,c,d 


Description: Length, 1.92 mm.; width, 0.92 mm. 

Moderately robust. Black, antennae piceous, especially at base; 
pubescence white, on dorsal surface fine, moderately sparse, under eye, 
on sides of prothorax, mesothorax, and metathorax coarse, somewhat 
denser. Beak of male shorter than head and prothorax combined, 
slightly less than one-third longer than prothorax, moderately, 
evenly curved; in lateral view attenuate from antennal insertion to 
middle, apical half nearly parallel, in dorsal view slightly expanded 
laterally at antennal insertion, attenuate to middle, slightly expanded 
at tip; rather dull, alutaceous, scantily pubescent and sparsely, 
shallowly punctate in basal two-thirds. Antennae inserted at basal 
fourth of beak, at distance from eye equal to width of frons; first 
segment equals next two, second segment equals next two, club 0.22 
by 0.08 mm. Eyes prominent; frons slightly wider than dorsal tip 
of beak. Prothorax at base one-sixth wider than long, middle nar- 
rower than base, apex five-sevenths as wide as base; sides beyond 
basal lateral expansion nearly parallel to middle, rounded to con- 
stricted apex; in profile dorsal surface nearly flat; punctation moder- 
ately deep, 0.03 mm. in diameter, interspaces slightly less than 
diameter of punctures; basal fovea shallow, punctiform. Elytra at 
base three-sevenths wider than prothorax at base, 2.6 times as long as 
prothorax, length to width as 14 : 11.5; intervals nearly flat, eque! to 
two striae, generally with one row of scales, in part with two rows; 
striae deep, fine. Scutellum elongate, triangular, 0.10 by 0.07 mm. 
Front femora 2.9 times as long as wide. Claws armed with acute 
basal tooth. 

Special male characters: Tibia 2 armed with fine mucrones, front 
femora swollen. 

Female is unknown. 

Tyrsrs: Holotype male (USNM 63421), 20 miles east of Morelia, 
Michoacdin, Mexico, Mar. 7, 1953, D. G. Kissinger. One male 
paratype, same data as holotype. 

Remarks: This species was taken on oak leaves. 


Apion (Trichapion) calcaratipes Sharp 


FIGuRE 8,g-7 


Apion calcaratipes Sharp, Biologia Centrali-Americana, Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 67. 


Description: Length, 2.06 mm. 


SUBGENUS TRICHAPION—KISSINGER 313 


Moderately robust. Black; tibiae, tarsi, and antennae dark 
reddish yellow; pubescence conspicuous, elytral suture in basal third 
behind scutellum clothed with broad, dense, pearly scales, these finer 
at base of intervals; circular area from humeri to apical third clothed 
with fine, yellow scales, giving area darker appearance, remainder of 
elytra clothed with moderately coarse, sparse, white scales; head and 
base of beak with sparse, coarse, pearly scales. Male beak one-fourth 
longer than prothorax, nearly straight; in dorsal view moderately 
expanded at antennal insertion, attenuate strongly to middle, thence 
cylindrical to tip, in lateral view expanded ventrally at antennal 
insertion, attenuate to middle; dull, punctured and scaly in basal half, 
apical half bare, polished, and subimpunctate. Female beak as long 
as head and prothorax combined, one-half longer than prothorax, 
slightly curved; apical two-thirds cylindrical, in dorsal view strongly 
dilated at antennal insertion, in lateral view basal sixth stout, slightly 
humped dorsally at antennal insertion; basal sixth dull, punctured, 
and scaly, remainder bare, polished and subimpunctate. Antennae 
inserted at distance from eye slightly less than width of frons, of male 
at basal fifth, of female at basal sixth; first segment equals next two; 
second segment nearly equals next two; club 0.21 by 0.08 mm. Eyes 
large, prominent; frons wide, with two irregular rows of punctures 
separated by a wide area with an indistinct median sulcus, base of 
frons with moderate punctures. Prothorax at base one-third wider 
than long, middle narrower than base, apex two-thirds as wide as 
base; sides beyond basal lateral expansion slightly converging to 
middle, then rounding to constricted apex; in profile dorsal surface 
slightly arcuate; punctation 0.03 mm. in diameter, deep, interspaces 
generally one-half as wide as punctures; basal fovea short, shallow, 
wide. Elytra at humeri one-third wider than prothorax at base; 2.5 
times as long as prothorax, length to width as 5:4; intervals convex, 
twice as wide as striae, with two or three irregular rows of fine punctures 
bearing scales; striae deep, with a row of scales similar to those on 
adjacent portion of interval. Scutellum triangular, 0.06 by 0.06 
mm., with slight median sulcus in basal half. Front femora 3.3 
times as long as wide. Claws with broad basal tooth. 

Special male characters: Tibiae 2 and 3 armed with small, curved 
mucrones which are slightly angulate ventrally at middle; tibia 1 
with inner anterior surface with a flattened, striate area; tarsus 1 
with outer apical angle of first segment produced into a short, blunt 
spine. 

MATERIAL EXAMINED: Male and female determined by Sharp. 

TYPE LOCALITY: Panajachel, Guatemala. 


314 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Apion (Trichapion) enoplus, new species 
FiaureE 8,e, f, 7 


Description: Length, 1.42 mm.; width, 0.71 mm. 

Robust. Black; pubescence on dorsal surface white, moderately 
conspicuous, coarser and denser on sides of prothorax, mesothorax, 
and metathorax. Male beak shorter than head and prothorax com- 
bined, three-sevenths longer than prothorax; slightly, evenly curved; 
apical third nearly cylindrical; in dorsal view moderately expanded 
laterally over antennal insertion, attenuate to apical third, slightly 
compressed in front of apical third; basal two-thirds dull, alutaceous, 
moderately punctured, sparsely pubescent; apical third shining, finely 
punctured. Antennae inserted at basal fourth of beak at distance 
from eye about equal to width of frons; first segment and second 
segment each shorter than next two segments; club 0.12 by 0.05 mm. 
Eyes prominent; frons wider than dorsal tip of beak. Prothorax at 
base three-eighths wider than long, middle slightly narrower than 
base, apex eight-elevenths as wide as base; sides beyond basal lateral 
expansion nearly parallel to middle, rounded to constricted apex; in 
profile dorsal surface nearly flat; punctation moderately shallow, 
0.03 mm. in diameter, interspaces about equal to diameter of punc- 
tures; basal fovea shailow, broadly punctiform. Elytra at humeri 
about one-half wider than prothorax at base; 2.8 times as wide as 
prothorax; length to width as 11:8.5; intervals nearly flat, twice as 
wide as striae, generally with one row of scales; striae moderately 
deep, fine. Scutellum triangular, 0.06 by 0.05 mm., with slight 
median furrow. Front femora 3.7 times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, slender, 
curved mucrones; front tibiae with flattened, finely striate area 
extending two-thirds its length on inner anterior face; first segment 
of fore tarsi with outer apical angle produced into a short spine. 

Female is not known. 

Typres: Holotype male (CAS), Jalapa, Veracruz, Mexico, July, 
Barrett collector, A. Fenyes Collection. One male paratype (DGK), 
near Huauchinango, Puebla, Mexico, June 1954, D. G. Kissinger. 

Remarks: The specimen from Huauchinango, Puebla, was taken 
while beating in a pine and scrubby oak forest at about 5,000 feet 
elevation. 

Apion (Trichapion) innocens, new species 


FIGuReE 8,k—n 


Description: Length, 1.59 mm.; width, 0.79 mm. 
Moderately robust. Black; antennae, tibiae in part, and tarsi 
piceous; pubescence dorsally very fine, long, yellowish, moderately 


SUBGENUS TRICHAPION—KISSINGER 315 


sparse, white and slightly coarser on sides and apex of elytra, on sides 
of prothorax, mesothorax, and metathorax still coarser and somewhat 
more conspicuous. Beak shorter than head and prothorax combined, 
one-third longer than prothorax, moderately, evenly curved; in 
lateral view attenuate from antennal insertion, apical{third nearly 
parallel-sided, in dorsal view attenuate from antennal‘insertion to 
apical third, there slightly compressed then expanded, apical third 
nearly parallel; basal three-fourths pubescent, basal two-thirds dull, 
alutaceous, punctured, apical third shining, sparsely punctured. 
Antennae inserted at basal fifth of beak at distance from eye about 
equal to width of frons; first segment and second segment each equal 
to next two segments combined, club 0.20 by 0.07 mm. Eyes promi- 
nent; frons slightly wider than dorsal tip of beak. Prothorax at base 
one-third wider than long, middle narrower than base, apex three- 
fourths as wide as base; sides beyond basal lateral expansion nearly 
parallel to middle, rounded to constricted apex; in profile dorsal 
surface nearly flat; punctation 0.03 mm. in diameter, moderately 
deep, interspaces narrower than diameter of punctures; basal fovea 
shallow, punctiform. Elytra at humeri one-third wider than pro- 
thorax at base, 2.65 times as long as prothorax, length to width as 
4:3; intervals nearly flat, twice as wide as striae, generally with two 
rows of fine scales; striae deep, fine. Scutellum elongate-triangular, 
0.08 by 0.05 mm. Front femora 3.3 times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibia 2 armed sath long, thin, curved 
mucro; first segment of fore tarsi with outer apical angle peda 
into a short spine. 

Female is not known. 

Types: Holotype male (USNM 63422), Huauchinango, Puebla, 
Mexico, June 1954, D. G. Kissinger. Four male paratypes: 3 
(BMNH, CAS, DGK) with same data as holotype and 1 (DGK) 
from El Salto, San Luis Potosf, Mexico, Jan. 22, 1953, D. G. Kissinger. 

ReMARKS: Specimens from Huauchinango, Puebla, were taken 
while beating in a scrub oak and pine forest at about 5,000 feet 
elevation. 


Apion (Trichapion) innocuum, new species 
Ficure 9,d,f,g 


Description: Length, 1.56 mm.; width, 0.77 mm. 

Moderately robust. Black; base of antennae yellow; pubescence 
white, on dorsal surface of prothorax and elytra fine, moderately 
sparse, on head, basal two-thirds of beak, sides of prothorax, meso- 
thorax, and metathorax very coarse, more conspicuous. Beak 
shorter than head and prothorax combined, one-sixth longer than pro- 


316 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ry : 


ci 


l 


l J 

Ficure 9.—a-c, Apion latitator, new species: a, entire dorsal view of male; b, lateral view 
of head and prothorax of male; c, mucro of tibia 2 of male. d, f, g, d. innocuum, new 
species: d, mucro of tibia 2 of male; f, dorsal view of head of male; g, lateral view of head 
and prothorax of male. ¢, h-j, 4. mirandum, new species: ¢, mucro of tibia 2 of male; 
h, lateral view of head and prothorax of male; 7, tarsus 1 of male of variant form; J, 
tarsus 1 of male of typical form. k, 4. mirificum, new species, tarsus 1 of male. 1, 
A. sanctifelicts Sharp, tarsus 3 of male. Line equals 0.50 mm. 


SUBGENUS TRICHAPION—KISSINGER S7 


thorax, slightly curved; in lateral view apical third nearly parallel, 
in dorsal view slightly expanded laterally at antennal insertion, 
attenuate to slightly past middle, beak not compressed or expanded 
toward tip; basal two-thirds punctured and densely covered with 
squamose pubescence, apical third shining, glabrous, nearly impunc- 
tate. Antennae inserted at basal fifth of beak, at distance from eye 
equal to width of frons; first segment equal in length to next two, 
second segment shorter than next two, club 0.15 by 0.07 mm. Eyes 
prominent; frons wider than dorsal tip of beak, with slight median 
furrow, and two irregular, lateral rows of fine punctures. Prothorax 
at base one-third wider than long, middle narrower than base, apex 
three-fourths as wide as base; sides beyond basal lateral expansion 
nearly parallel to middle, rounded to constricted apex; in profile 
dorsal surface slightly arcuate; punctation 0.03 mm. in diameter, 
moderately deep, interspaces somewhat narrower than diameter of 
punctures; basal fovea shallow, punctiform. Elytra at humeri 
one-third wider than base of prothorax, 2.5 times as long as pro- 
thorax, length to width as 15:12.5; intervals twice as wide as striae, 
somewhat convex, with one row of fine punctures bearing fine scales; 
striae deep, fine. Scutellum elongate-triangular, 0.08 by 0.06 mm., 
with obscure median furrow. Front femora 3.75 times as long as 
wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with moderately long, 
slender mucro. 

Types: Holotype male (CAS), Cuernavaca, Morelos, Mexico, June, 
A. Fenyes Collection. One male paratype (DGK), near Antiguo 
Morelos (in Tamaulipas), San Luis Potosi, Mexico, July 1954, D. G. 
Kissinger. 


Apion (Trichapion) latitator, new species 
FiaguRE 9,a-c 


Description: Described from a unique specimen. 

Length, 1.50 mm.; width, 0.75 mm. 

Moderately robust. Specimen teneral, antennae and apices of 
tibiae and tarsi paler; pubescence white, fine, inconspicuous on dorsal 
surface of elytra, somewhat longer and very sparse on dorsal surface 
of prothorax, somewhat denser on sides of mesothorax and metepister- 
num. Beak slightly shorter than head and prothorax combined, 
three-fifths longer than prothorax, moderately, evenly curved; apical 
half nearly cylindrical, in dorsal view not attenuate beyond slight 
lateral expansion over antennal insertion; basal two-thirds dull, very 
sparsely pubescent, finely, shallowly punctured in rows, apical third 
more shining. Antennae inserted at basal fifth of beak, at distance 
from eye about equal to width of frons; first segment equal to next 


318 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


two; second shorter than next two; club 0.18 by 0.08 mm. Eyes 
prominent; frons slightly wider than dorsal tip of beak. Prothorax 
at base two-fifths wider than long, middle narrower than base, apex 
three-fourths as wide as base; sides beyond basal lateral expansion 
slightly expanded to middle, rounded to apex; in profile dorsal surface 
slightly arcuate; punctation 0.03 mm. in diameter, moderately 
shallow, interspaces less than diameter of punctures; basal fovea 
lacking. Elytra at humeri one-third wider than prothorax at base, 
nearly three times as long as prothorax, length to width as 4 :3; 
intervals moderately convex, twice as wide as striae, with one row 
of minute scales; striae moderately deep, fine. Scutellum triangular, 
0.05 by 0.05 mm., nearly smooth. Front femora 3.6 times as long as 
wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with moderately short, 
subangulate mucro projecting in line with long axis of tibia, first 
segment of fore tarsus with outer apical angle produced into a spine. 

Female is not known. 

Houoryre: Male (USNM 63423), on banana leaf from Mexico, 
intercepted at Brownsville, Tex., Feb. 22, 1938, No. 38-2705. 


Apion (Trichapion) mirandum, new species 
FIGURE 9,e,h-j 


Description: Length, 1.75 mm.; width, 0.84 mm. 

Moderately robust. Black; antennae yellow, apices of tibiae and 
tarsi dark yellow; pubescence white, on dorsal surface fine, moderately 
sparse, on sides of prothorax, mesothorax and metathorax, under 
eyes, and base of beak denser, coarser, conspicuous. Beak of male 
shorter than head and prothorax combined, one-fourth longer than 
prothorax, slightly curved; in dorsal view slightly expanded laterally 
at antennal insertion, strongly attenuate to apical two-thirds, apical 
third strongly expanded; basal two-thirds dull, alutaceous, punctured, 
pubescent, apical third shining, sparsely punctured. Antennae 
inserted at basal fourth of beak, at distance from eye equal to width 
of frons; first segment nearly equal to next three, second segment 
equal to next two, club 0.21 by 0.08 mm. Eyes prominent; frons 
slightly wider than dorsal tip of beak. Prothorax at base one-fifth 
wider than long, middle slightly narrower than base, apex three- 
fourths as wide as base; sides beyond basal lateral expansion nearly 
parallel to middle, rounded to constricted apex; in profile dorsal 
surface nearly flat; punctation shallow, 0.03 mm. in diameter, inter- 
spaces slightly less than diameter of punctures; basal fovea shallow, 
punctiform. Elytra at humeri one-half wider than prothorax at base, 
2.6 times as long as prothorax, length to width as 13 : 10; intervals 
nearly flat, equal in width to two striae, with one row of scales; 


SUBGENUS TRICHAPION—KISSINGER 319 


striae moderately deep, fine. Scutellum elongate-triangular, 0.08 
by 0.04 mm. Front femora 4.5 times as long as wide, claws with 
acute basal tooth. 

Special male characters: Tibia 2 armed with fine mucro; first 
segment of fore tarsus elongate, nearly four times as long as wide, 
ventral surface clothed with long cilia. 

Female is not known. 

Typxrs: Holotype, male (USNM 63424), 20 miles east of Morelia, 
Michoacan, Mexico, Mar. 7, 1953, D. G. Kissinger. Five male para- 
types: 4 (BMNH, CAS, AMNH, DGK) with same data as holotype 
and 1 (DGK) from near Huauchinango, Puebla, Mexico, June 1954, 
D. G. Kissinger. 

Remarks: Material from 20 miles east of Morelia, Michoacan, 
was taken on oak leaves. The specimen from Huauchinango, Puebla, 
was taken while beating in a scrubby oak and pine forest at about 
5,000 feet elevation. 

A specimen taken with the type series from Morelia shows (fig. 9,2) 
even stronger development of the male secondary sexual modification 
of the fore tarsus. The first segment is slightly longer and about 
twice as broad as that of the typical specimens and the second segment 
is much longer and broader. This specimen is thought to be an indi- 
vidual variant and is not included in the paratype series. 


Apion (Trichapion) mirificum, new species 
Figure 9,k 


Description: Length, 2.00 mm.; width, 0.87 mm. 

Moderately robust. Black; pubescence white, on dorsal surface 
fine, inconspicuous, under eyes, on sides of prothorax, mesothorax 
and metathorax coarser and more conspicuous. Beak of male shorter 
than head and prothorax combined, one-seventh longer than pro- 
thorax, slightly curved; in lateral view attenuate from antennal 
insertion to apex; in dorsal view apical half nearly parallel-sided; 
with sparse, fine punctures and scales in basal two-thirds. Antennae 
inserted at basal fourth, at distance from eyes equal to width of 
frons; first segment shorter than next two, second segment slightly 
longer than third; club lacking on specimen. Eyes prominent; frons 
wider than dorsal tip of beak, with rather deep, narrow median 
sulcus and one lateral row of subconfiuent punctures. Prothorax 
at base one-fourth wider than long, middle slightly wider than base, 
apex three-fourths as wide as base; sides beyond acute basal lateral 
expansion rather strongly diverging to middle, rather strongly rounded 
from base to constricted apex; in profile dorsal surface slightly arcuate; 
punctation 0.03 mm. in diameter, moderately deep, interspaces 
narrower than punctures; basal fovea moderately deep, wider at 


320 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


base, extends one-third length of prothorax. Elytra at humeri one- 
third wider than prothorax at base, 2.6 times as long as prothorax, 
length to width as 10:7; intervals flat, more convex toward base, 
twice as wide as striae, with a single row of punctures; striae fine, 
deep. Scutellum 0.06 by 0.06 mm., with slight median furrow. 
Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with long, fine, curved 
mucro; tarsus 1 with inner margin of first segment produced into a 
flat lobe projecting anteriorly slightly beyond tip of segment; tibia 1 
a little stouter than other tibiae. 

Hoxtoryre: Male (USNM 63428), Coyame, on Lago de Catemaco, 
Veracruz, Mexico, June 1954, D. G. Kissinger. 


Apion (Trichapion) sancti-felicis Sharp 


FIGuRE 9,l 


Apion sancti-felicis Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 77, 1890. 

Description: Length; 1.50 to 1.83 mm. 

Robust. Black; base of antennae and fore femora and tibiae 
reddish brown; middle and hind femora and tibiae piceous; pubescence 
on dorsal surface of prothorax and elytra fine, yellowish, very sparse; 
white, coarser and denser laterally. Beak of male much shorter than 
head and prothorax combined, slightly longer than prothorax, slightly 
curved; in side view attenuate to apex, ventral margin nearly straight; 
in dorsal view slightly expanded at basal fourth, apical two-thirds 
nearly parallel-sided; moderately shining, with fine punctures arranged 
in rows, becoming sulciform laterally; basal three-fourths sparsely 
pubescent. Beak of female as long as head and prothorax combined, 
one-half longer than prothorax, moderately, evenly curved; in side 
view attenuating to apex; in dorsal view nearly parallel-sided to apex, 
antennal insertion minutely prominent; with sparse, fine punctures 
throughout, moderately shining. Antennae inserted at basal fourth of 
beak at distance from eye slightly greater than width of frons; first 
segment equals next three, second slightly shorter than next two; club 
0.21 by 0.07 mm. Eyes prominent; frons narrow, not as wide as dorsal 
tip of beak, with one lateral row of moderately coarse punctures and a 
moderately wide, flat median interval. Prothorax at base one-third 
wider than long, middle slightly narrower than base, apex two-thirds to 
three-sevenths as wide as base; sides beyond basal lateral expansion 
slightly converging to middle, rounding to constricted apex; in profile 
dorsal surface slightly arcuate, apical margin slightly raised; puncta- 
tion 0.03 mm. in diameter, moderately deep, interspaces narrower 
than diameter of punctures; basal fovea short, moderately deep, 
narrow. Elytra at humeri one-third to two-fifths wider than pro- 


SUBGENUS TRICHAPION—KISSINGER ool 


thorax at base, from 2.25 to 2.50 times as long as prothorax, length 
to width as 10:9; intervals nearly flat, twice as wide as striae, with 
one occasionally irregular row of fine punctures with fine scales, 
interval 2 with two rows of punctures, smooth, without transverse 
rugae; striae moderately deep, coarse. Scutellum rounded, 0.06 by 
0.06 mm., with broad, shallow median sulcus. Front femora 3.8 
times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with short mucro, first 
segment of tarsus 3 with inner apical margin produced into a short 
spine. 

MATERIAL EXAMINED: 50 specimens including material determined 
by Sharp. 

KNOWN DISTRIBUTION: 


Mexico: Veracruz: Coyame, Lago de Catemaco, June 1954, D. G. Kissinger; 
18 miles north of San Andrés Tuxtla, Feb. 6-12, 1953, and June 1954, D. G. 
Kissinger. 

Panama: ‘San Feliz.” 


Apion (Trichapion) spinitarse Wagner 
Apion spinittarse Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 18, 1911. 


Description: Length, 1.42 to 1.50 mm. 

Moderately robust. Black; base of antennae piceous or yellow; 
pubescence on dorsal surface of prothorax and elytra fine, white, 
very sparse, denser and coarser on sides of mesothorax and meta- 
thorax; male with ventral surface covered with scales much coarser 
than those of female. Beak of male moderately stout, strongly 
attenuate, much shorter than head and prothorax combined, one- 
third longer than prothorax, moderately curved; moderately expanded 
at antennal insertion, attenuate in middle third, parallel-sided in 
apical third, apical third slightly compressed, tip slightly expanded; 
sparse pubescence in basal two-thirds, punctures slight, sparse, 
mainly in basal two-thirds, apical region smoother; moderately 
shining beyond antennal insertion. Beak of female shghtly longer 
than head and prothorax combined, three-fourths longer than pro- 
thorax, moderately curved, more strongly deflexed in apical third; 
in lateral view attenuate beyond antennal insertion to apical third 
which is nearly parallel-sided; in dorsal view stout at base, slightly 
expanded over antennal insertion, slightly attenuate to tip; dull, 
alutaceous throughout, with shallow, very fine punctures; with a few 
scales behind antennal insertion. Antennae of male inserted at 
basal fourth of beak at distance from eye slightly greater than width 
of frons, of female inserted at basal fifth of beak at distance from 
eye equal to width of frons; first segment of male equals next two, 
of female somewhat longer than next two; second segment somewhat 


322 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


longer than third; club 0.17 by 0.05 mm. Eyes prominent; frons 
of male narrow, equals dorsal tip of beak; of female wide, much 
wider than dorsal tip of beak, with a moderately wide, flat, median 
area with at most a very slight sulcus. Prothorax at base one-third 
wider than long, middle slightly narrower than base, apex three- 
fourths as wide as base; sides beyond basal lateral expansion nearly 
parallel to middle, rounded to constricted apex; in profile dorsal 
surface slightly arcuate; punctation 0.03 mm. in diameter, moderately 
deep, interspaces equal to or slightly less than diameter of punctures; 
basal fovea shallow, broad, short. Elytra at humeri one-third wider 
than prothorax at base, 2.75 times as long as prothorax, length to 
width as 11:8; intervals twice as wide as striae, nearly flat to slightly 
convex, with one row of fine punctures bearing fine scales; striae 
moderately fine, deep. Scutellum 0.08 by 0.05 mm., elongate-tri- 
angular, with slight median depression. Front femora three times 
as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with small, short, thin, 
curved mucro; tibia 2 armed with larger curved mucro; tibia 1 with 
anterior inner surface bearing a smooth, extremely finely and closely 
striate area on about one-half of its length; tarsus 1 with inner posterior 
margin of the first segment produced into a large, triangular, flat 
projection. 

MATERIAL EXAMINED: Two males and one female determined by 
Hans Wagner. 

KNOWN DISTRIBUTION: 


Nicaraaua: Managua (Solari Collection). 
Braziut: Rio de Janeiro (BMNH). 


Apion punctulirostre Group 


Three species comprise this distinct group. Two species, A. 
brunnicornis Fall and A. eccentricum Fall, occur together in Arizona. 
A. punctulirostre Sharp occurs farther south in the mountains of 
Mexico and Guatemala. 

A. brunnicornis Fall is distinct because of the pale antennae which 
in the case of the male are inserted slightly behind the middle of the 
beak. The antennae of the other two species are black and are 
inserted at about the basal third of the beak. A. punctulirostre 
Sharp is very close to A. eccentricum Fall. The principal difference is 
that the beak of the former, in lateral view, is nearly parallel beyond 
the antennal insertion to the apex; in the latter species the beak 
attenuates, in lateral view from the antennal insertion to the apex. 


SUBGENUS TRICHAPION—KISSINGER 323 


Apion (Trichapion) brunnicornis Fall 
Ficure 10,a—d 
Apion brunnicornis Fall, Journ. New York Ent. Soc., vol. 26, p. 221, 1918. 


Description: Length, 1.6 to 1.9 mm. 

Moderately robust. Black; antennae brownish testaceous; pu- 
bescence fine, white, sparse, male with pubescence slightly more 
conspicuous on sides of mesothorax, under side of head and on anterior 
surface of front coxae, female with pubescence more uniform. Male 
beak very thick, one-third shorter than head and prothorax combined, 
strongly deflexed at middle; in lateral view dorsal margin strongly 
curved downward, apical third thickened ventromedially; in dorsal 
view not expanded laterally at antennal insertion, strongly compressed 
in apical third; sparsely, shallowly punctate to tip, pubescent to 
near tip, tip slightly shinmg. Female beak slender, equal to head 
and prothorax in length, very strongly, evenly curved, cylindrical, 
sparsely, shallowly punctate to near apex, basal two-thirds with 
sparse pubescence. Antennae inserted at distance from eye equal to 
twice width of frons, of male inserted slightly behind middle of beak, 
of female at basal one-third; first segment of male a little longer 
than next two, of female equal to next three segments; second segment 
about one-half longer than third, shorter than next two; club 0.20 
by 0.08 mm. Eyes prominent; frons moderately wide, with deep, 
elongate median sulcus, with a row of large, deep punctures next to 
sulcus and a row of finer, shallower punctures next to eye. Pro- 


LAICA! A 
14 OE 


C € 
Ficure 10.—a-d, Apion brunnicornis Fall: a, lateral view of head and prothorax of male; 
b, lateral view of head and prothorax of female; c, dorsal view of head of male; d, mucro of 
tibia 2 of male. e¢-h, A. punctulirostre Sharp: e, dorsal view of head of male; f, mucro 
of tibia 2 of male; g, lateral view of head and prothorax of male; h, lateral view of head and 
prothorax of female. Line equals 0.50 mm. 


324 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


thorax at base slightly to one-fourth wider than long, middle slightly 
narrower than base, nearly parallel to middle then rounded to con- 
stricted apex; in profile dorsal surface nearly flat; punctation 0.03 mm. 
in diameter, deep, interspaces less than diameter of punctures; basal 
fovea deep, elongate, extending to middle of prothorax. Elytra at 
humeri one-third wider than prothorax at base, 2.75 times as long 
as prothorax, length to width as 10:7.5; intervals flat, twice as wide 
as striae, not shining, with a single row of fine punctures bearing fine 
scales; striae deep with a single row of fine scales. Scutellum tri- 
angular, 0.07 by 0.05 mm., with a shallow median furrow. Front 
femora of female 3.5 times as long as wide, of male 2.5 times as 
long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 with a fine mucro; femora stouter 
than those of female, femora 1 and 3 are 2.5 times as long as wide, 
femur 2 is 2.25 times as long as wide. 

MATERIAL EXAMINED: Type, male (MCZ 25078), Chiricahua Mts., 
Ariz., in Fall Collection, and 25 specimens. 

KNOWN DISTRIBUTION: 


Unitep Status: Arizona: Chiricahua Mts. (MCZ, USNM); Santa Rita Mts., 
Madera Canyon, June 21, 1953, A. and H. Dietrich (CU). 


Apion (Trichapion) eccentricum Fall 
Apion eccentricum Fall, Journ. New York Ent. Soc., vol. 26, p. 220, 1918. 


Description: Length, 2.0 to 2.2 mm. 

Moderately robust. Black; elytra with faint bluish luster; pubes- 
cence white, fine, very sparse, uniform on dorsal surface, denser and 
coarser on sides of mesothorax, denser on sides of prothorax and meta- 
thorax. Male beak slightly shorter than head and prothorax com- 
bined, shghtly more than one-half longer than prothorax, moderately, 
evenly curved; in lateral view expanded ventrally at antennal inser- 
tion, attenuate to tip; in dorsal view slightly expanded laterally at 
antennal insertion, attenuating toward apex, apex slightly expanded, 
sides compressed at apical third; strongly punctate in rows to near 
apex, punctures bear scales to near apex, apex smooth, bare. Female 
beak two-fifths longer than head and prothorax combined, twice as 
long as prothorax, moderately, evenly curved; in lateral view stout 
in basal third, attenuating to apical third and nearly parallel to tip; 
in dorsal view stouter at base and nearly parallel-sided, attenuating 
to beyond middle, slightly compressed before tip; punctures strong, 
in rows, pubescent in basal two-thirds, apex smoother, bare. Antennae 
of male inserted at basal third of beak at distance from eye twice as 
great as width of frons, of female inserted slightly distad of the basal 
third at distance from eye 2.5 times as great as width of frons; first 
segment of male equals next three, of female slightly shorter than 


SUBGENUS TRICHAPION—KISSINGER 325 


next four; second segment slightly longer than third; club 0.24 by 
0.09 mm. Eyes moderately prominent; frons slightly wider than 
dorsal tip of beak, with shallow, broad, median sulcus and one lateral 
row of coalesced punctures. Prothorax at base one-third wider than 
long, middle narrower than base, apex three-fourths as wide as base; 
sides beyond basal lateral expansion slightly converging to middle, 
rounded to constricted apex; in profile dorsal surface flat; punctures 
deep, 0.04 mm. in diameter, interspaces alutaceous, irregular, less 
than diameter of punctures; basal fovea shallow, extending one-third 
length of prothorax. Elytra at humeri one-fourth wider than pro- 
thorax at base, three times as long as prothorax, length to width as 
12:8.5; intervals convex, twice as wide as striae, with one or two 
rows of fine punctures; striae deep. Front femora of male three times 
as long as wide, of female 3.3 times as long as wide. Claws with 
acute basal teoth. 

Special male characters: Tibia 2 armed with mucro, all femora 
swollen. 

MatTerRIAL EXAMINED: Type, male (MCZ 25090), Santa Rita Mts., 
Ariz., in Fall Collection, and three specimens in Fall Collection. 

Known DIstTriBurion: 

Unirep States: Arizona: Santa Rita Mts.; Huachuca Mts., Ramsay Canyon 
(MCZ). 

Apion (Trichapion) punciulirostre Sharp 


FiaurEe 10,e-h 


Apion punctulirostre Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 57, 1890—Kissinger, Proc. Ent. 
Soc. Washington, vol. 59, p. 40, 1957. 

Apton spectator Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, Curcu- 
lionidae: Apioninae, vol. 4, pt. 3, p. 61, 1890.—Kissinger, Proc. Ent. Soe. 
Washington, vol. 59, p. 40, 1957. 

Description: Length, 2.0 to 2.5 mm. 

Moderately robust; black. Pubescence fine, white, sparse, denser 
on sides of mesothorax. Male beak as long as head and prothorax 
combined, slightly, evenly curved, middle (in dorsal view) slightly 
thicker than base, apical third compressed; with slight, dorsal, median 
carina, and a lateral line of punctures in a slight groove extending to 
apical third; pubescence conspicuous, apex smoother, glabrous. Fe- 
male beak nearly twice as long as head and prothorax combined, 
slightly, evenly curved, nearly cylindrical throughout; apex slightly 
expanded; finely, deeply punctured, apex not as shining as base, basal 
third with scant pubescence. Antennae inserted in basal third of 
beak, of male at distance from eye 2.25 times as great as width of 
frons, of female at distance from eye three times as great as width of 
frons; first segment longer than next three, shorter than next four; 


326 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


second segment shorter than next two; club 0.21 by 0.09 (male) to 
0.28 by 0.10 mm. (female). Eyes moderately prominent; frons nar- 
row, equal to width of dorsal tip of beak, with slight median impres- 
sion and a lateral row of confluent punctures. Prothorax at base one- 
fourth to one-third wider than long, middle narrower than base; sides 
strongly expanded laterally at base, nearly parallel to middle, rounded 
to constricted apex; in profile dorsal surface nearly flat, slightly sin- 
uate before base and apex; punctation 0.04 to 0.05 mm. in diameter, 
deep, interspaces less than diameter of punctures, slightly convex; 
basal fovea deep, linear, extending one-third length of prothorax. 
Elytra at humeri one-third wider than prothorax at base, 2.75 times 
as long as prothorax, length to width as 4:3; intervals twice as wide 
as striae, moderately convex, with one irregular row of fine punctures 
bearing fine scales; striae deep. Scutellum triangular, 0.12 by 0.08 
mm., with median furrow. Front femora of male 3.4 times as long 
as wide, of female 4.2 times as long as wide. Claws with acute basal 
tooth. 

Special male characters: Tibia 2 with moderately long, simple 
mucro; all femora much stouter than those of female. 

MatTERIAL EXAMINED: Male and female compared with type of 
punctulirostre by G. A. K. Marshall, two males and two females 
determined as A. spectator by Sharp, and six other specimens. 

KNOWN DISTRIBUTION: 

Mexico: Morelos: Cuernavaca (BMNH). Meézico: Temescaltepec, Real de 
Arriba, July 1932, H. Hinton (CAS). Puebla: Near Huauchinango, June 


1954, D. G. Kissinger (DGK). 
GUATEMALA: Aceituno; San Gerénimo (BMNB#). 


Remarks: Specimens from Huauchinango, Puebla, were taken 
while beating in a scrubby oak and pine forest at about 5,000 feet 
elevation. 

A. punctulirostre was described from two females. The original 
description is rather brief and is more or less a comparison with A. 
gracilirostre Sharp. A. spectator was described in more detail with 
the statement that only the middle tibiae of the male are mucronate. 
A comparison of specimens reveals no significant differences. 


Apion submetallicum Group 


The four species comprising this group apparently overlap in geo- 
graphical range. Apion auriferum Wagner and A. perpilosum Wagner 
occur in Guatemala; the latter, A. submetallicum Boheman, and A. 
hadromerum Wagner occur in Guerrero, Mexico. 

A. submetallicum is about 1 mm. longer than the other species and 
the antennae are inserted more distad on the beak. The beak of the 
male of A. hadromerum is clothed with long, bristly pubescence. The 


SUBGENUS TRICHAPION——KISSINGER 327 


beak of the males of the other species is clothed with shorter pubes- 
cence. The mucrones of the males of A. perpilosum project at an 
angle with the tibiae and the elytral intervals are flat. The mucrones 
of the male of A. auriferum project in line with the tibiae at the base 
and curve out at an angle in the apical half and the intervals are 
convex. 


Apion (Trichapion) auriferum Wagner 
Figure 11,a,b,k 
Apion (Trichapion) auriferum Wagner, Arch. Naturg. Berlin, vol. 78, p. 100, 1912. 
Description: Length, 1.4 to 1.6 mm. 
Moderately robust. Black; pubescence conspicuous, fine, white, 
sparse, coarser and denser at base of elytral interval 3 and sides of 
prothorax, mesothorax, and metathorax. Male beak one-seventh 


longer than prothorax, slightly curved, slightly expanded laterally over 
antennal insertion, evenly attenuate toward apex; in basal three- 


a eee 


Ficure 1l1.—a, b, k, Apion auriferum Wagner: a, entire dorsal view of male; b, lateral view 
of head and prothorax of male; k, mucro of tibia 3 of male. c, f, g, j, 4. perpilosum 
Wagner: c, dorsal view of head of male; f, lateral view of head and prothorax of male; g, 
lateral view of head and prothorax of female; 7, mucro of tibia 3 of male. d, A, l, A. 
hadromerum Wagner: d, dorsal view of head of male; h, lateral view of head and prothorax 
of male; J, mucro of tibia 3 of male. ¢,i,m, A. submetallicum Boheman: ¢, dorsal view of 
head of male; i, lateral view of head and prothorax of male; m, mucro of tibia 3 of male. 
Line equals 0.50 mm. 

483661—59 6 


328 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


fourths sparsely punctate and pubescent, apical fourth smoother and 
shining. Antennae inserted at distance from eye equal to width of 
frons, at basal fifth of beak; first segment one-half longer than second, 
shorter than next two; second segment slightly longer than third; 
club 0.17 by 0.09 mm. Eyes moderately prominent; frons moderately 
wide, with two rows of punctures on either side of wide, slightly con- 
cave median area. Prothorax at base one-third wider than long, mid- 
dle slightly narrower than base, apex two-thirds as wide as base; sides 
expanded laterally at base, nearly parallel to middle, rounded to con- 
stricted apex; in profile dorsal surface slightly, evenly convex; puncta- 
tion 0.03 mm. in diameter, shallow, interspaces less than diameter of 
punctures; basal fovea round, moderately deep. Elytra at humeri 
one-fourth wider than prothorax at base, about 2.5 times as long as 
prothorax, length to width as 9 : 7; intervals more than twice as wide 
as striae, moderately convex, smooth, with two rows of punctures 
bearing fine scales; striae deep, fine. Scutellum triangular, as wide as 
long, about 0.06 by 0.06 mm., with slight median furrow. Front 
femora about three times as long as wide. Claws with large, acute 
basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long mucrones which at base are in line with the long axis of the tibiae 
and curve out laterally in apical half, mucrones angulate ventrally. 

MATERIAL EXAMINED: One male determined by Hans Wagner. 

KNOWN DISTRIBUTION: 

Mexico: Tabasco: Teapa. 

GUATEMALA: San Gerénimo (BMNH). 


Nicaraacua: Managua (Solari Collection). 
PanaMa: ‘‘San Felice” (BMNH). 


Apion (Trichapicn) hadromerum Wagner 


FicureE 11,d,h,l 


Apion (Trichapiton) hadromerum Wagner, Arch. Naturg. Berlin, vol. 78, p. 105, 
1912. 

Description: Length, 1.8 to 2.0 mm. 

Moderately robust. Black; pubescence conspicuous, fine, on dor- 
sal surface in part yellowish, sparse; coarser, white laterally, on head 
and base of beak, and at small spot at base of elytral interval 3. Male 
beak four-fifths as long as head and prothorax, nearly straight ; moder- 
ately expanded ventrally at antennal insertion, attenuating to apex; 
punctured and pubescent in basal three-fourths, apex, smooth, shining; 
with dorsomedian narrow carina in basal three-fourths. Antennae in- 
serted at distance from eye one-fourth greater than width of frons, 
shghtly distad of basal third; first segment shorter than next two, 
second segment equal to third, club 0.18 by 0.08 mm. Eyes promi- 


SUBGENUS TRICHAPION—KISSINGER 329 


nent; frons moderately wide, with median sulcus and two or three 
irregular rows of moderate punctures. Prothorax at base one-fourth 
wider than long, middle slightly narrower than base, apex seven-tenths 
as wide as base; sides expanded laterally at base, nearly parallel to 
middle, slightly rounded to constricted apex; in profile dorsal surface 
nearly flat; punctation 0.03 mm. in diameter, moderately deep, inter- 
spaces about equal to diameter of punctures, flat, alutaceous; basal 
fovea shallow, moderately broad, short. Elytra at humeri two-fifths 
wider than prothorax at base, three times as long as prothorax, length 
to width as 3 : 2; intervals less than twice as wide as striae, third in- 
terval twice as wide as striae, moderately convex, smooth with two 
irregular rows of fine punctures bearing fine scales. Scutellum tri- 
angular, about as long as wide, 0.06 by 0.06 mm., smooth, alutaceous. 
Front femora 3.5 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, curved, 
simple mucrones. 

MatTERIAL EXAMINED: One male determined by Hans Wagner and 
seven other specimens. 

KNOWN DISTRIBUTION: 

GuATEMALA: Capetillo; Duefias (BMNH). 

Mexico: Distrito Federal: Mexico City, July 29, 1944, N. L. H. Kraus (USNM); 
Contreras, May 30, 1946, J. and D. Pallister (AMNH). Guerrero: Omilteme; 
Chilpancingo (BMNH). Michoacén: South side of Lago de Cuitzeo, 1,950 
meters, July 8, 1947, No. 82, T. H. Hubbell (UM). Morelos: Coajomulco, June 7, 
1946, J. and D. Pallister (AMNH); near Cuernavaca, Aug. 14, 1953, about 
7,000 feet, R. B. and J. M. Selander (DGK). Puebla: Near Huauchinango, 
June 1954, D. G. Kissinger. 

Remarks: The first two Mexican localities were erroneously attrib- 
uted to Guatemala by Wagner in his description. 


Apion (Trichapion) perpilosum Wagner 


FIGURE 11,¢,f,9,9 


Apion (Trichapion) perpilosum Wagner, Arch. Naturg. Berlin, vol. 78, p. 107, 
fig. b, 1912—McKelvey, et al., Mex. Sec. Agr. Ganad. Fol. Tec., vol. 8, 
pp. 8-42, 1951. 

Description: Length, 1.45 to 1.60 mm. 

Moderately robust. Black; aeneous; pubescence conspicuous, 
whitish and in part yellowish on dorsal surface, fine sparse, denser 
and coarser at base of third elytral interval and sides of prothorax, 
mesothorax, and metathorax. Male beak three-fourths as long as 
head and prothorax combined, slightly, uniformly curved; moderately 
expanded laterally and ventrally at antennal insertion, attenuate to 
apex; moderately densely punctured and pubescent to near apex, 
pubescence long and bristling, apex smooth and shining. Female 
beak slightly longer than head and prothorax combined, moderately, 


330 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


uniformly curved; slightly expanded laterally and ventrally at an- 
tennal insertion, apical two-thirds nearly cylindrical; basal fourth 
moderately punctured and pubescent, remainder smooth, glabrous, 
and shining. Antennae inserted at distance from eye equal to width 
of frons, of male slightly distad of basal fourth of beak, of female at 
basal fifth; first segment shorter than next two; second segment 
shorter than next two; club 0.18 by 0.08 to 0.21 by 0.09 mm. Eyes 
moderately prominent; frons moderately wide, with slight, narrow 
median sulcus and two lateral rows of punctures. Prothorax at base 
one-eighth wider than long, middle slightly narrower than base, apex 
three-fourths as wide as base; sides with slight lateral expansion at 
base, slightly expanded to middle, rounded to constricted apex; in 
profile dorsal surface slightly arcuate, flattened basally and apically; 
punctation 0.02 to 0.03 mm. in diameter, deep, interspaces variable, 
usually less than diameter of punctures; basal fovea moderately deep, 
about one-fourth length of prothorax. Elytra at humeri two-fifths 
wider than prothorax at base, 2.5 times as long as prothorax, length 
to width as 10:7.5; intervals twice as wide as striae, nearly flat, with 
one or two rows of punctures; striae deep, fine. Scutellum roundly 
triangular, about 0.08 by 0.06 mm., with moderate, median furrow. 
Front femora 3.75 times as long as wide. Claws with acute basal 
tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, subangulate mucrones. 

MatTERIAL EXAMINED: Male and female determined by Hans 
Wagner. 

KNOWN DISTRIBUTION: 


Mexico: Guanajuato. Guerrero: Amula; Chilpancingo (BMNH). 
GUATEMALA: Guatemala City (BMNH). 


Apion (Trichapion) submetallicum Boheman 
Figure 11,e,i,m 

Apion submetallicum Boheman, in Schoenherr, Genera et species curculionidum 
vol. 5, p. 376, 1839.—Sharp, Biologia Centrali-Americana. Insecta, Coleop 
tera, Curculionidae: Apioninae, vol. 4, pt. 3, p. 48, 1890. 

Apion hystriculum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 56, 1890.— Wagner, Arch. Naturg. 
Berlin, vol. 78, p. 108, fig. a, 1912. New synonymy. 

Description: Length, 2.5 to 3.0 mm. 

Moderately robust. Black; pubescence conspicuous, fine, white, 
sparse, denser and coarser at base of elytra interval 3, laterally and 
ventrally. Male beak three-fourths as long as head and prothorax, 
slightly curved, moderately dilated laterally and ventrally at antennal 
insertion, attenuate to apical third which is nearly cylindrical; moder- 
ately punctured and pubescent in basal two-thirds, apex smooth, 


SUBGENUS TRICHAPION—KISSINGER 301 


shining. Antennae inserted at distance from eye four-fifths greater 
than width of frons at basal one-third of beak; first segment longer 
than second, shorter than next two; second segment shorter than 
next two; club 0.24 by 0.08 mm. Eyes moderately prominent; frons 
moderately wide, with median sulcus and two or three lateral rows of 
moderate punctures. Prothorax slightly wider at base then long, 
middle slightly narrower than base, apex four-fifths as wide as base; 
sides slightly expanded laterally at base, nearly parallel to middle, 
slightly rounded to broadly constricted apex; in profile dorsal surface 
flat; punctures deep, 0.03 mm. in diameter, interspaces narrow, about 
one-half as wide as diameter of punctures, convex, shining, alutaceous; 
basal fovea deep, linear, extending one-fifth length of prothorax. 
Elytra at humeri three-fifths wider than prothorax at base, 2.6 times 
as long as prothorax, length to width as 13:9; intervals less than 
twice as wide as striae, flat, with two rows of moderately fine punctures ; 
striae deep. Scutellum elongate, triangular, 0.09 by 0.06 mm., with 
a median furrow. Front femora four times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long blunt 
mucrones. 

MATERIAL EXAMINED: 10 males, including Boheman’s type and a 
male compared with the type of A. hystriculum by G. A. K. Marshall. 

KNOWN DISTRIBUTION: 

Mexico: Puebla: Near Huauchinango, June 1954, D. G. Kissinger (DGK). 
Distrito Federal: Mexico City, A. W. Barret (USNM). Mezico: Amecameca, 
June 8, 1897, Koebel Collection (CAS). Guerrero: Xucumanatlan, Omilteme, 
H. H. Smith. 

Remarks: Material from Huauchinango was taken while beating 
in a scrubby oak and pine forest at about 5,000 feet elevation. 

Apion submetallicum has not been recognized since its description 
in 1839. Sharp (1890) simply listed the name, stating that he could 
not recognize the species in any material before him. An examine- 
tion of Boheman’s type, through the kindness of Dr. René Malaise, 
has revealed the above synonymy. 


Apion patruele Group 


The 10 species comprising this group are characterized by their 
robust body; generally deep, elongate prothoracic basal fovea (lacking 
in rufipenne and evustum); and the strongly expanded beak over the 
antennal insertion. Also the second and third pairs of tibiae of the 
male are mucronate, tibia 1 of the male is simple, prothorax is widest 
at base, and the beak is subcylindrical in the apical half. The range 
of three species, A. patruele Smith, A. perforicolle Fall, and A. porcatum 
Boheman overlaps in the eastern United States. Three species, A. 


3a2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


abdominale Smith, A. centrale Fall, and A. perforicolle, are very 
similar but occur in widely separated localities. 

A. rufipenne Gyllenhal and A. evustum, new species, are distinct 
with their red elytra. The other species are black. A. gulare Fali 
is distinct in that the ventral surface of the head is deeply excavated 
and polished and the antennae are inserted far front on the beak. 
Apion porcatum is a large species with narrow, strongly convex 
elytral intervals, and the antennal club is rather small, being shorter 
than segments 5 through 8 of the antennal funicle. The elytral inter- 
vals of the other species are nearly twice as wide as the striae (except 
in abdominale) and are flatter and the antennal club is about as long as 
seements 3 through 8 of the antennal funicle. The mucrones of 
A. perforicolle are rather short and subangulate, the elytral intervals 
are twice as wide as the striae, the beak is strongly dilated laterally 
over the antennal insertion and in comparison with A. centrale the 
prothorax is nearly parallel in the basal half and not constricted 
apically, and the interspaces on the dorsal surface of the prothorax 
are about equal to the diameter of the punctures and are slightly 
shining and strongly alutaceous. The mucrones of A. patruele, A. 
abdominale, and A. centrale are dentate and the sides of the pro- 
thorax of the latter are sinuate before the base (the basal lateral ex- 
pansion is more pronounced) and constricted apically; the dorsal 
surface of the prothorax is closely punctured; and the interspaces are 
about one-half the diameter of the punctures, polished and very 
finely alutaceous. The beak of A. patruele is not expanded laterally 
at the antenna insertion; the abdomen and metasternum are rather 
finely, sparsely punctured; and the elytral intervals are somewhat 
convex. The beak of A. centrale and A. abdominale is strongly 
expanded laterally at the antennal insertion; the abdomen and meta- 
sternum. are coarsely, closely, and deeply punctured; and the elytral 
intervals are flatter. The intervals of A. centrale are nearly twice 
as wide as the striae and slightly wider at the middle than at the base 
and the mucrones of the male are short and strongly angulate. The 
intervals of A. abdominale are narrower and are slightly wider than 
the striae and are especially narrow basally and the mucrones are 
moderately long and slender. A. fusconitidum Wagner is distinct 
with its large, robust form; narrow, convex intervals; and irregularly 
punctured dorsal surface of the prothorax. 

The name Apion patruele Smith has been selected arbitrarily as 
the one to be used to designate the section of Trichapion Wagner 
that is coordinate with the Apion simile section. As yet no member 
of the A. patruele section is known to be holarctic in distribution. 

The differentiation of the groups of the A. patruele section with 
respect to the A. patruele group is as follows. The A. reconditum 


SUBGENUS TRICHAPION—KISSINGER 333 


group is very closely allied, most of its members exhibit characters 
common to both groups, i. e., parallel-sided beak in dorsal view in 
apical half, male tibiae 2 and 3 mucronate, and generally little sexual 
dimorphism in length of beak. The members of the A. reconditum 
group are more slender, the elytra are longer in proportion to the 
prothorax, the basal lateral expansion of the prothorax is more pro- 
nounced, and the beak tends to be depressed apically. 

The A. oblitum group is similar to the A. patruele group with its 
robust body and cylindrical beak in apical half, but the base and 
middle of the prothorax are nearly equal, the beak is abruptly nar- 
rowed beyond the antennal insertion and cylindrical and polished 
to the tip, and the male has all three tibiae mucronate. 

The A. nigrum group is similarly stout and has the beak cylindrical 
in its apical half, but the prothorax is modified so that it is much 
wider at the middle than at the base, also the front legs of the male 
are modified either with mucronate tibiae (cordatum), or fore tibia 
dilated, curved, and set with long ciliae within (heterogeneum), or 
with dilated or densely pubescent first tarsal segment (nigrum and 
dolosum). 

The prothorax of the A. rostrum group is generally about as wide 
at the base as at the middle. The beak is distinctly attenuate to 
near the apex, also there is sometimes marked sexual dimorphism in 
the length of the beak. Some members of this group exhibit further 
affinity with the A. simile section because they have multiserially 
pubescent elytral intervals. 


Apion (Trichapion) abdominale Smith 


FIGuReE 12,a,e-g 


Apion abdominale Smith, Trans. Amer. Ent. Soc., vol. 11, p. 53, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 143, pl. 4, fig. 13, 1898. 

DescrirTion: Length, 2.25 to 2.40 mm.; width, 1.12 to 1.18 mm. 

Robust. Black; pubescence scant, white, fine, not much more 
conspicuous on ventral surface. Male beak slightly shorter than 
head and prothorax combined, one-third longer than prothorax; 
slightly curved; in lateral view apical half is nearly parallel-sided; in 
dorsal view strongly expanded laterally at antennal insertion, from 
middle to apex nearly parallel-sided; coarsely punctured laterally, 
more finely dorsally, with short, deep sulcus above antennal insertion. 
Female beak as long as head and prothorax combined, one-half longer 
than prothorax; moderately, evenly curved, in lateral view sides 
nearly parallel in apical half; dull, smoother and finely, sparsely punc- 
tured, tip slightly shining. Antennae inserted at basal fifth of beak 
at distance from eye equal to width of frons; first segment equals 
next two, second segment slightly shorter than next two, club 0.24 by 


334 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 12.—a, e-g, Apion abdominale Smith: a, entire dorsal view of male; ¢, mucro of 
tibia 2 of male; f, mucro of tibia 3 of male; g, lateral view of head and prothorax of male. 
b, h, i, A. centrale Fall: b, entire dorsal view of male; h, lateral view of head and prothorax of 
male; 7, lateral view of head and prothorax of female. c, d, A. fusconitidum Wagner: ¢, 
entire dorsal view of female; d, lateral view of head and prothorax of female. 4, k, A. 
evustum, new species: j, lateral view of head and prothorax of male; k, dorsal view of 


prothorax. Line equals 0.50 mm. 


SUBGENUS TRICHAPION—KISSINGER 335 


0.09 mm. Eyes moderately prominent; frons slightly wider than 
dorsal tip of beak. Prothorax at base slightly wider than long, 
middle and base about equal, apex four-fifths as wide as base; sides 
beyond slight basal lateral expansion expanding slightly to middle, 
rounded to slight apical constriction; in profile dorsal surface slightly 
arcuate; punctation deep, rounded, 0.04 mm. in diameter, inter- 
spaces less than diameter of punctures, generally one-half as wide; 
basal fovea deep, narrow, extending beyond middle. Elytra at 
humeri one-third wider than prothorax at base, about three times as 
long as prothorax, length to width as 4 : 3; intervals narrower at base, 
not more than two-third wider than striae, flat, with one row of fine 
punctures; striae deep, coarse. Scutellum triangular, 0.06 by 0.06 
mm. Front femora 3.5 times as long as wide. Claws with acute 
basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, dentate mucrones. 

Lecrotyprr: J hereby designate as lectotype of this species the 
male specimen (USNM 1263) labeled California, J. B. Smith Collection. 

MATERIAL EXAMINED: Lectotype and five specimens. 

KNOWN DISTRIBUTION: 


Unitep States: California: No exact locality (USNM). Arizona: No exact 
locality (CU); Santa Rita Mountains, July 18, 1938, R. I. Sailer (UK). 


Apion (Trichapien) centrale Fall 
Figure 12,b,h,i 
Apion centrale Fall, Trans. Amer. Ent. Soc., vol. 25, p. 151, 1898—Bleasdell, 
Journ. Sci. lowa State Coll., vol. 11, p. 411, 1937. 

Description: Length, 2.1 to 2.4 mm. 

Robust. Black; pubescence white, fine, sparse, a little coarser 
laterally and ventrally. Male beak as long as head and prothorax 
combined, slightly, evenly curved, expanded laterally and ventrally at 
antennal insertion, rather abruptly attenuate beyond tbat point, from 
middle to apex nearly cylindrical, apex about two-thirds as wide as 
base of beak; punctation coarse, sparse to apical third, pubescence 
inconspicuous. Female beak a little longer than head and prothorax; 
slightly, evenly curved, antennal insertion as in male; punctation 
beyond antennal insertion sparse, fine. Antennae inserted at distance 
from eye equal to width of frons, at basal fifth of beak; first segment 
of male equal to next two, of female longer than next two; second 
segment equals next two; club ranges from 0.24 by 0.10 to 0.27 by 0.12 
mm. Eyes prominent; frons wide with a slight central depression, 
laterally with two rows of punctures. Prothorax at base slightly 
wider than long, at middle a little wider than base, apex three-fourths 
as wide as base; sides in basal half slightly diverging to middle, 


336 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


rounded to slightly constricted apex; in profile dorsal surface slightly, 
evenly convex; punctation coarse, 0.04 mm. in diameter, deep, 
interspaces narrower than diameter of punctures, usually one-half 
as wide; basal fovea deep, elongate, extending about one-third length 
of prothorax. Elytra at humeri two-fifths wider than prothorax at 
base, 2.75 times as long as prothorax, length to width as 13 : 10; in- 
tervals twice as wide as striae, nearly flat to slightly convex, with an 
irregular row of fine punctures, ground sculpture fine, transverse; 
striae deep. Scutellum triangular, about as long as wide, 0.06 by 
0.06 mm., smooth. Front femora 3.5 times as long as wide. Claws 
with moderate basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, subangu- 
ate mucrones. 

Typss: I hereby designate as lectotype of this species the male 
specimen in the Fall Collection labeled Garland, Colo. (MCZ 25080). 
Cotypes are in the J. L. LeConte Collection labeled Hudson Bay 
Territory and Colorado (MCZ 237) and in the U.S. National Mu- 
seum labeled Hudson Bay Territory (USNM 4224). 

Material examined: Lectotype and 50 specimens. 

KNOWN DISTRIBUTION: 


Unitep States: Colorado: Garland, Hubbard and Schwarz (USNM); 
Gunnison, Baker (AMNH); Pagosa Springs, July 22, C. F. Baker (USNM); 
Beta Pass, May 27, Hubbard and Schwarz (USNM). Idaho: Bonner Co., Sagle, 
July 16, 1950, N. M. Downie (DGK). Jowa: Emmitsburg, July 6, 1916, D. 
Stoner (USNM); Iowa City, May 30, 1916, L. L. Buchanan (USNM); Lake 
Okoboji, July 2, 1917, L. L. Buchanan (USNM); Dickinson Co.; Johnson Co.; 
Palo Alto Co.; and Story Co. Montana: Bear Paw Mountain, Sept. 17, Hubbard 
and Schwarz (USNM); Kalispell, June 13-20, H. F. Wickham (USNM). North 
Dakota: Wahpeton, July 7 (USNM). 

Canapva: Alberta: Cypress Hills, July 21, 1930, J. H. Pepper (CNC); Edmonton, 
June 8, 18, 28, July 8-27, F. 8. Carr (AMNH, DGK, UC, USNM); Medicine 
Hat, July 23, 1930, J. H. Pepper (CNC); Morrin, June 17, 1929, G. F. Manson 
(CNC); Lethbridge, June 2, 1930, J. H. Pepper (CNC); Little Smokey River, 
June 23, 1954, C. P. Alexander (CAF). British Columbia: Clinton, June 14, 
1938, J. K. Jacobs (CNC); Salmon Arm, May 29, 1932, sweeping bushes, H. B. 
Leech (CNC, USNM); Vernon, July 2, 1929, H. B. Leech (CNC); Oliver, Aug. 6, 
1930, J. Nottingham (UK); Robson, May 22, 1948, H. R. Foxlie (CNC); Terrace, 
Mrs. M. E. Hippisley (CAF). Hudson Bay Territory (MCZ, USNM). Manitoba: 
Aweme, July 3, 1909, E. Criddle (CNC); Winnipeg (USNM). Ontario: Leaming- 
ton, June 10, 1929, G. 8S. Walley (CNC); Moosonee, July 17, 1934, G. S. Walley 
(CNC); Ogoki, July 6, 1932, J. B. Wallis (CNC); Smoky Falls, Mattagami 
River, June 11, 1939, G. 8. Wallace (CNC). Saskatchewan: Fort a la Corne, 
July 17, 1925, K. M. King (CNC); Waskensin Lake, June 2, 1939, A. R. Brooks 
(CNC). 


SUBGENUS TRICHAPION—KISSINGER 337 


Apion (Trichapion) evustum, new species 
FigurE 12,j,k 


Description: Length, 2.12 to 2.25 mm.; width, 1.12 mm. 

Robust. Beak, antennae, head, and prothorax black; elytra, 
abdomen, mesothorax and metathorax pale reddish; coxae and legs 
darker red. Pubescence white, fine, prothorax with a number of 
scales on basal median region and a few along median line; elytra 
glabrous except at apex; denser on sides of mesothorax and mete- 
pisternum. Beak of male shorter than head and prothorax combined, 
one-half longer than prothorax, slightly curved; in lateral view rather 
stout and parallel-sided in basal third, thence rather suddenly nar- 
rowed, apical half subparallel-sided; in dorsal view subparallel in 
apical half, with moderate expansion over antennal insertion; moder- 
ately punctured in basal half, with sparse pubescence in basal third, 
apical half finely, sparsely punctured; dorsal surface sparsely, finely 
punctured. Beak of female as long as head and prothorax combined, 
three-fourths longer than prothorax, slightly curved; apical half 
subcylindrical, with very sparse, fine punctures; with minute pube- 
scence slightly distad of antennal insertion. Antennae inserted at 
distance from eye equal to width of frons, of male at basal sixth, of 
female at basal seventh; first segment equals next two, second seg- 
ment shorter than next two, club 0.26 by 0.10 mm. Eyes moderately 
prominent; frons equals dorsal tip of beak, flat medially. Prothorax 
at base one-third wider than long, middle slightly narrower than base, 
apex three-fourths as wide as base; sides beyond moderate basal 
lateral expansion subparallel in basal half, thence rounded to dis- 
tinctly constricted apex; in profile dorsal surface slightly arcuate; 
punctation shallow, coarse, sparse but irregularly placed; basal fovea 
lacking. Elytra at humeri two-fifths wider than prothorax at base; 
2.75 times as long as prothorax, length to width as 11:9; intervals 
flat, about three times as wide as striae, glabrous, nearly impunctate; 
stria 1 complete, uniformly impressed throughout, 2 and 9 nearly 
complete, other striae more or less distinct at base and appear as a 
row of separated punctures in middle of elytra. Scutellum triangular, 
0.09 by 0.09 mm. Front femora four times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with large, blunt 
strongly dentate mucrones. 

Typrs: Holotype male (USNM 63133), Tampico, Tamaulipas, 
Mexico, December 21, E. A. Schwarz. Allotype female, same data 
as holotype (USNM). Eight paratypes; five, same data as holotype 


338 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


(USNM); three, Tamazunchale, San Luis Potosi, Mexico, Jan. 28, 
1953, D. G. Kissinger (DGK). 


Apion (Trichapion) fusconitidum Wagner 
Figure 12,c,d 


Apion fusconitidum Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 23, pl. 2, fig. 4, 
1911. 

Description: Length, 2.62 mm.; width, 1.50 mm. 

Robust. Black, slightly aeneous; tibiae and tarsi castaneous; 
pubescence white, very fine, on dorsal surface of prothorax and elytra 
minute, very sparse, slightly more conspicuous on sides of mesothorax 
and metathorax. Female beak about as long as head and prothorax 
combined; three-fourths longer than prothorax, moderately curved; 
basal third stoutly cylindrical, then attenuate, apical three-sevenths 
subcylindrical; from above not expanded laterally at antennal inser- 
tion; with fine sparse punctures evident to tip, with minute scales 
somewhat distad of antennal insertion. Antennae inserted at basal 
fourth, at distance from eye slightly greater than width of frons; 
first segment slightly shorter than next three, second segment shorter 
than next two, club 0.30 by 0.10 mm. Eyes prominent; frons slightly 
wider than dorsal tip of beak, flat medially. Prothorax at base one- 
third wider than long, middle narrower than base, apex seven-tenths 
as wide as base; sides beyond slight basal lateral expansion converging 
to apical two-fifths, thence strongly rounded to strongly constricted 
apex, extreme apex flared out laterally; in profile dorsal surface nearly 
flat; punctation about 0.03 mm. in diameter, shallow, very irregularly 
placed; with an impunctate region in lateral basal half; basal fovea 
deep, punctiform, with a short, shallow anterior furrow. Elytra at 
humeri one-half wider than prothorax at base, nearly three times as 
long as prothorax, length to width as 5:4; intervals convex, about 
twice as wide as striae, with several rows of minute punctures bearing 
extremely small scales; striae deep, fine. Scutellum triangular, 0.09 
by 0.09 mm., with slight median impression. Front femora about 
four times as long as wide. Claws with acute basal tooth. 

Material examined: One female determined by Hans Wagner. 

KNOWN DISTRIBUTION: 


Braziu: Rio de Janeiro and Petrépolis (BMNH). 
Apion (Trichapion) gulare Fall 
FiaureE 13,a-c 


Apion gulare Fall, Trans. Amer. Ent. Soc., vol. 25, p. 140, 1898.—Blatchley and 
Leng, Rhynchophora or weevils of northeastern America, p. 78, 1916. 


Description: Robust. Black, with reddish aeneous luster; pubes- 
cence moderate, fine, white, sparse, slightly denser on sides of meso- 


SUBGENUS TRICHAPION—-KISSINGER 339 


Figure 13.—a-c, Apion gulare Fall: a, entire dorsal view of male; b, lateral view of head and 
prothorax of male c, mucro of tibia 3 of male. d, e, 4. nitidum Kirsch: d, mucro of tibiae 
3 of male; e, lateral view of head and prothorax of male. f, m, n, 4. porcatum Boheman: 
f, mucro of tibiae 3 of male; m, entire dorsal view of male; 2, lateral view of head and 
prothorax of male. g-j, 4. patruele Smith: g, mucro of tibia 3 of male; A, lateral view of 
head and prothorax of male; 1, lateral view of head and prothorax of female; 7, dorsal 
view of head of male. &, 1, 4. perforicolle Fall: k, lateral view of head and prothorax of 
male; J, entire dorsal view of male. 0, p, 4. rufipenne Gyllenhal: 0, dorsal view of pro- 
thorax; p, lateral view of head and prothorax of male. Line equals 0.50 mm. 


340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


thorax. Male beak as long as head and prothorax combined, slightly 
curved, moderately dilated laterally and ventrally at antennal inser- 
tion, nearly cylindrical beyond middle, tip slightly expanded; punc- 
tures sparse, very fine, with a lateral row of coarse punctures above 
antennal insertion, apical half shining, pubescence inconspicuous. 
Female beak slightly longer than head and prothorax, slightly curved, 
nearly cylindrical throughout, expanded slightly ventrally in basal 
fourth, apex expanded; in basal half with a lateral row of coarse punc- 
tures above antennal insertion, apical half dull, alutaceous, pubescence 
inconspicuous. Antennae inserted at distance from eye one-half 
greater than width of frons, of male inserted slightly distad of basal 
third, of female slightly distad of basal fourth; first segment equal to 
next two segments, second segment equal to next two, club 0.18 by 
0.08 mm. Eyes moderately prominent; frons narrow, wider than 
dorsal tip of beak, with a wide shallow median impression and a row 
of four or five coarse punctures tateral to this; underside of head 
deeply excavated and strongly polished. Prothorax one-third wider 
at base than long, middle slightly narrower than base, apex three- 
fourths as wide as base; sides moderately expanded laterally at base, 
nearly parallel to middle, slightly rounded to slightly constricted 
apex; in profile dorsal surface nearly flat; punctation shallow, 0.03 mm. 
in diameter, interspaces narrower than diameter of punctures; basal 
fovea rounded, shallow. Elytra at humeri one-third wider than 
prothorax at base, three times as long as prothorax, length to width 
as 9:7; intervals twice as wide as striae, nearly flat, alutaceous, with 
one row of fine, distant punctures bearing very fine scales; striae not 
deep, with a row of scales slightly more conspicuous than those on 
intervals. Scutellum triangular, about 0.06 by 0.04 mm., smooth. 
Front femora 2.75 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with minute, simple 
mucrones. 

Typrs: I hereby designate as lectotype of this species the male 
specimen in the Fall Collection labeled Key West, Fla. (MCZ 25100). 
Cotypes with the same data are in the J. L. LeConte Collection 
(MCZ 364) and in the U.S. National Museum (USNM 4235). 

MATERIAL EXAMINED: Lectotype and 20 specimens. 

KNOWN DISTRIBUTION: 

UniTep States: Florida: Biscayne; Key West (MCZ, USNM). 

Mexico: Tamaulipas: Tampico, December 8, E. A. Schwarz (USNM). 

Remarks: Material in the U. S. National Museum was reared by 
HK. A. Schwarz from Piscidea erythrina. 

The specimen from Mexico undoubtedly belongs to this species. It 
differs in that the beak is somewhat more distinctly punctured toward 
the apex. 


SUBGENUS TRICHAPION—KISSINGER 341 


Apion (Trichapion) nitidum Kirsch 
Figure 13,d,e 
Apion nitidum Kirsch, Berliner Ent. Zeitschr., vol. 18, p. 416, 1874 


Description: Length, 2.12 mm.; width, 1.06 mm. 

Robust. Black; pubescence white, fine inconspicuous on dorsal 
surface of elytra, prothorax dorsally with scattered, sparse scales; on 
sides of mesothorax and metathorax moderately dense. Beak of male 
about equal to head and prothorax combined, three-fifths longer than 
prothorax, slightly curved; basal fifth rather stout, subcylindrical, not 
expanded laterally at antennal insertion; basal half with sparse punc- 
tures and scales; apical half smoother, bare. Antennae inserted at 
basal sixth of beak at distance from eye equal to width of frons; first 
segment equals next two, second segment shorter than next two, club 
0.24 by 0.10 mm. Eyes moderately prominent; frons slightly wider 
than dorsal tip of beak, flat medially. Prothorax at base one-fourth 
wider than long, middle slightly narrower than base, apex three- 
fourths as wide as base; sides beyond basal lateral expansion nearly 
parallel-sided to middle, rounded to constricted apex; in profile dorsal 
surface nearly flat; punctation 0.03 mm. in diameter, rather shallow, 
irregularly placed, a large impunctate area laterally at base, inter- 
spaces from one-fourth as wide to wider than punctures; basal fovea 
deep, linear, extending about one-half length of prothorax. Elytra 
at humeri one-half wider than prothorax at base, 2.5 times as long as 
prothorax, length to width as 10:9; intervals nearly flat, about one- 
half wider than striae, with a single irregular row of minute punctures 
bearing minute scales; striae deep, coarse, scales in striae somewhat 
larger than those on intervals. Scutellum triangular, 0.09 by 0.09 
mm. Front femora 3.7 times as long as wide. Claws with acute 
basal tooth. 

Special male characters: Tibiae 2 and 3 armed with angulate 
mucrones. 

MatTERIAL EXAMINED: One male (BMNH, labeled “‘Amazonas’’) 
compared with type by Hans Wagner. 

KNOWN DISTRIRUTION: Peru (type locality). 


Apion (Trichapion) patruele Smith 


Figure 13,g-7 


Apion patruele Smith, Trans. Amer. Ent. Soc., vol. 11, p. 64, fig. 11, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 142, pl. 4, figs. 11, 1la, 17, 1898.—Blatch- 
ley and Leng, Rhynchophora or weevils of northeastern America, p. 78, fig. 
37q, 1916. 

Description: Length, 1.68 to 2.12 mm.; width, 0.85 to 1.12 mm. 
Robust. Black; pubescence fine, white, sparse, coarser and denser 
on mesothorax and metathorax. Male beak stout, shorter than head 


342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


and prothorax combined, two-fifths longer than prothorax, deflexed 
at basal third; not expanded laterally at antennal insertion, apical 
third nearly cylindrical; more coarsely punctate laterally, with a row 
of confluent punctures above antennal insertion bearing scales, punc- 
tures to near tip bear minute scales. Female beak as long as head 
and prothorax combined, three-fifths longer than prothorax, evenly, 
strongly curved; nearly cylindrical beyond antennal insertion; finely 
sparsely punctured to tip, punctures bearing minute scales. Antennae 
inserted at distance from eye equal to width of frons, of male at basal 
fourth of beak, of female at basal fifth; first sezment equals next two, 
second segment shorter than next two, club 0.21 by 0.08 mm. Eyes 
moderately prominent; frons slightly wider than dorsal tip of beak, 
with deep median sulcus and one lateral row of somewhat confluent 
punctures. Prothorax at base two-fifths wider than long, middle 
narrower than base, apex five-sevenths as wide as base; sides beyond 
moderately basal lateral expansion nearly parallel to middle, rounded 
to constricted apex; in profile dorsal surface slightly arcuate; puncta- 
tion moderately deep, 0.03 mm. in diameter, interspaces about equal 
to diameter of punctures, somewhat alutaceous. Elytra at humeri 
two-fifths wider than prothorax at base, three times as long as pro- 
thorax, length to width as 16:13; intervals twice as wide as striae, 
rather convex, generally with one row of fine scales; striae deep, fine. 
Scutellum triangular, 0.06 by 0.06 mm., nearly smooth. Front femora 
about 3.25 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, dentate 
mucrones. 

Typxs: I hereby designate the lectotype of this species as the male 
specimen (USNM 1246) labeled Massachusetts. Cotype in the J. L. 
LeConte Collection labeled with an orange disc signifying the Southern 
States (MCZ 368). 

MATERIAL EXAMINED: Lectotype and 100 specimens. 

KNOWN DISTRIBUTION: 

Unitep Srares: Alabama: Mobile, Apr. 18, 1910, W. D. Pierce (USNM); 
Tuscaloosa, May 21, 1955, at light, B. D. Valentine; 5 miles north of York, July 
1954, D. G. Kissinger (DGK). Connecticut: Shelton, June 15, 1934, M. P. Zoppe 
(USNM); West Haven, Sept. 15, 1944 (USNM). District of Columbia: Wood- 
ridge, July 3, 1919, L. L. Buchanan (USNM). Florida: Dunedin, Feb. 30, 1926, 
W.S. Blatchley (CU, USNM); Lake Harney, May 4, E. A. Schwarz (USNM). 
Georgia: Griffin (UK); Okefenokee Swamp, May 7, 1953, N. J. and E. L. Sleeper 
(ELS). Illinois: Funkhouser, May 21, 1950 (INHS); Herod, May 29, 1936 
(INHS); McClure, June 25, 1931, on locust (INHS); Muncie, Aug. 31, 1907 
(USNM); Palisades State Park, July 4, 1946 (INHS); Pittsfield, Pike Co. (UC); 
Starved Rock State Park, July 4, 1941 (INHS); Valmeyer, Aug. 25, 1949, on red 
clover (INHS). Jndiana: (Blatchley and Leng, 1916) common throughout the 
State. Maryland: Beltsville, May 21, 1922, L. L. Buchanan (USNM); Glen 
Kcho, Aug. 3, 1922, L. L. Buchanan (USNM); Sparrow Point, July 4, 1932 (UC); 


SUBGENUS TRICHAPION—KISSINGER 343 


2 miles east of Silver Spring, Northwest Branch, June 28, 1952, D. G. Kissinger 
(DGK). Massachusetts: Canton, August 1924, D. H. Blake (USNM); Chicopee, 
July 23, 1903, F. Knab (USNM); Framingham, July 2, 1950, C. A. Frost (CAF); 
Marion, Wickham (USNM); Sherborn, July 19, 1947, C. A. Frost (CAF). Michi- 
gan: Clarkston (USNM); Detroit (USNM); Washtenaw Co., Aug. 1, 1922, 
M. H. Hatch (USNM). Missouri: No exact locality, J. B. Smith (USNM). 
New Jersey: Anglesea (USNM); Arlington (USNM); Avon, Aug. 30, 1923 
(USNM); Bayhead (USNM); Burlington Co. (USNM); Camden, December 1, 
G. M. Greene (USNM); Cape May (UC); Clifton (USNM); Montclair (USNM); 
Newark, E. A. Bischoff (USNM); Orange Mt. (USNM); Point Pleasant, July 3, 
1926, A. Nicolay (USNM); Roselle Park, Nov. 9, 1924, A. Nicolay (USNM); 
Woodside (USNM). New York: Ithaca (USNM); Riverhead, Long Island, 
June 23, 1924 (USNM); Rosedale, Long Island, June 3, 1923 (USNM); Schenec- 
tady, June 27, 1935, N. M. Downie; South Jamesport, Aug. 25, 1944, Tuthill, 
on Apios tuberosa leaves (USNM). North Carolina: Andrews, July 11, 1954 
(DGK); Bell Island, May 10, 1952, D. G. Kissinger (DGK); Nelson, August 31, 
L. D. Beamer (UK). Ohio: Champaign Co., July 4, 1950, E. L. Sleeper (ELS); 
Hocking Co., May 27, July 18, N. J. and E. L. Sleeper (ELS); Licking Co., 
Aug. 2, 1950, E. L. Sleeper (ELS); Scioto Co., Aug. 1, 1950, E. L. Sleeper (ELS) ; 
Summit Co. (UK). Pennsylvania: No exact locality (TLCC); Angora, June 15 
(USNM); Philadelphia (USNM); Wind Gap, June 18, 1931 (UC). Rhode Island: 
No exact locality (TLCC). Tennessee: Gatlinburg (Univ. Illinois). Texas: 
Dallas, May 22, 1910, H. Pinkus (USNM); San Diego, Chittenden (USNM). 
Virginia: Falls Church, July 4, 1919, L. L. Buchanan (USNM); Fredricksburg, 
May 27, 1900, W. N. Richardson (USNM); Lake Drummond, June 8, 1905, 
H. S. Barber (USNM); Pennington Gap, Hubbard and Schwarz (USNM). 
West Virginia: White Sulphur Springs (TLCC). Wisconsin: Green Lake, 
June 12, 1928, Ralf Martin (UK). 


Remarks: The host plant is not known definitely; Tuthill took 
specimens on the foliage of Apios tuberosa in New York. 


Apion (Trichapien) perforicolle Fall 
Figure 13,k,l 


Apion perforicolle Fall, Trans. Amer. Ent. Soc., vol. 25, p. 144, pl. 5, fig. 1, 1898.— 
Blatchley and Leng, Rhynchophora or weevils of northeastern America, p. 
79, 1916. 
Description: Length, 1.80 to 2.40 mm.; width, 0.87 to 1.12 mm. 
Moderately robust. Black; elytra sometimes faintly aeneous; 
pubescence white, rather coarse, sparse, denser on sides of mesothorax 
and metepisternum. Male beak shorter than head and prothorax 
combined, one-half longer than’ prothorax, slightly, evenly curved; 
expanded laterally at antennal insertion, apical half nearly cylindrical; 
more coarsely punctured above antennal insertion, more finely punc- 
tured dorsally, apical third smoother. Female beak slightly longer 
than head and prothorax, two-thirds longer than prothorax, slightly 
curved; otherwise similar to male. Antennae inserted at distance 
from eye equal to width of frons, of male inserted at basal fourth, of 
female at basal fifth of beak; first segment equals next two, second 
483661—59-—7 


344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


segment shorter than next two, club 0.25 by 0.10 mm. Eyes moder- 
ately prominent; frons much wider than dorsal tip of beak. Prothorax 
at base one-third wider than long, middle about equal to base, apex 
three-fourths as wide as base; sides beyond minute basal lateral ex- 
pansion slightly expanded to middle, thence rounded to apex, not con- 
stricted at apex; in profile dorsal surface slightly arcuate; punctation 
deep, 0.04 mm. in diameter, interspaces less than diameter of punc- 
tures, alutaceous; basal fovea deep, narrow, extending about one- 
third length of prothorax. Elytra three-eighths wider at humeri than 
base of prothorax, 2.8 times as long as prothorax, length to width as 
9 :7; intervals twice as wide as striae, nearly flat, with one row of fine 
scales; striae moderately deep, fine. Scutellum triangular, 0.06 by 
0.06 mm., with deep, median groove. Front femora about 3.5 times as 
long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
short, subdentate mucrones. 

I hereby designate the lectotype of this species as the male specimen 
(MCZ 25113) in the Fall Collection labeled Georgia. Cotypes are in 
the J. L. LeConte Collection (MCZ 371) and in the U. S. National 
Museum (USNM 4218). 

MatTERIAL EXAMINED: Lectotype and 20 specimens. 

Known DistrIBUTION: 

Unirep States: Alabama: Tuscaloosa, May 24, 1953, on Tephrosia spicata, 
B. D. Valentine (BDV). Florida: Mount Pleasant, April 1952, O. Peck (CNC). 
Georgia: Augusta, May 4, 1944, P. W. Fattig (USNM). Indiana: Pulaski Co., 
June 19, July 16, determined by Fall. Louisiana: Longville, June 17, 1948, H. W. 
Crowder (UK). Mississippi: Fulton, July 14, 1930, R. H. Beamer (UK). North 
Carolina: Southern Pines (TLCC). New Jersey: Lakehurst, July 4, E. A. Bischoff 
(USNM). 

Remarks: Blatchley and Leng (1916) record this species as swept 
from the lead plant, Amorpha canescens Pursh. Specimens were 
taken by B. D. Valentine on Tephrosia spicata. 


Apion (Trichapion) porcatum Boheman 


FIGuRE 13,f,m,n 


Apion porcatum Boheman, in Schoenherr, Genera et species curculionidum, vol. 
5, p. 374; 1839.—Smith, Trans. Amer. Ent. Soc., vol. 11, p. 64, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 150, 1898.—Blatchley and Leng, Rhyn- 
chophora or weevils of northeastern America, p. 81, 1916.—Bleasdell, Journ. 
Sci. Iowa State Coll., vol. 11, p. 411, 1937. 

Description: Length, 2.2 to 2.6 mm. 

Robust. Black; pubescence inconspicuous, very fine, white, uni- 
form. Male beak slightly shorter than head and prothorax combined; 
deflexed at basal two-fifths; moderately expanded laterally and ven- 
trally at antennal insertion, attenuate to middle, apical half nearly 


SUBGENUS TRICHAPION—KISSINGER 345 


cylindrical; moderately punctate throughout, with a confluent line 
of punctures over antennal insertion, pubescence inconspicuous. Fe- 
male beak as long as head and prothorax combined, slightly, evenly 
curved, otherwise similar to male with exception that beak is more 
slender and punctation is sparse. Antennae inserted at distance from 
eye slightly greater than width of frons, at basal fourth of beak; first 
segment longer than next two, shorter than next three; second segment 
equal to next two, club 0.21 by 0.10 mm. Eyes prominent; frons 
moderately wide, with a wide median area with a faint median depres- 
sion and one lateral line of coalescent punctures. Prothorax at base 
one-third wider than long, middle slightly wider than base, apex three- 
fourths as wide as base; sides moderately expanded laterally at base, 
slightly expanding to middle, rounded to constricted apex; in profile 
dorsal surface slightly, evenly arcuate; punctation 0.04 mm. in diame- 
ter, deep, interspaces irregular, from one-half as great to equal to 
diameter of punctures; basal fovea deep, elongate, extending one-half 
length of prothorax. Elytra at humeri two-fifths wider than pro- 
thorax at base, 2.8 times as long as prothorax, length to width as 
13 : 10; intervals strongly convex, slightly wider than striae, with one 
row of moderate punctures bearing inconspicuous scales; striae deep. 
Scutellum triangular, 0.06 by 0.06 mm., with slight median impression. 
Front femora four times as long as wide. Claws with acute basal 
tooth. 

Special male characters: Tibiae 2 and 3 armed with long, slender 
denticulate mucrones. 

MATERIAL EXAMINED: Type labeled Massachusetts in the Riks- 
museum and 50 specimens. 

KNOWN DISTRIBUTION: 

Unitep States: District of Columbia: Collected by Hubbard and Schwarz 
(USNM). Georgia: Neil Gap, Apr. 27, 1946 (USNM). Illinois: East St. Louis, 
May 9, 1924 (USNM). Jndiana: Clark, Floyd, Perry, and Posey Counties. 
Iowa: Des Moines and Lee Counties. Maryland: Beltsville, June 8, 1919, L. L. 
Buchanan (USNM); Bladensburg, May 4, 1915, L. L. Buchanan (USN M); Cabin 
John, Potomac River, May 24, 1952, G. H. Nelson, beating foliage; Glen Echo, 
May 27, 1923, J. R. Malloch (USNM); Plum Point, May 28, 1922, L. L. Buchanan 
(USNM); Plummers Island, June 29, 1920, H. S. Barber (USNM); South Moun- 
tain near Pine Knob, May 14, 1918, R. E. Van Zwaluwonburg (USNM). Massa- 
chusetts: Chicopee (USNM). New Hampshire: Pike, July 16, 1931, on locust, 
Mrs. A. W. Jones (USNM). New Jersey: Nutley, E. A. Bischoff (USNM); 
Point Pleasant, Aug. 8, 1926, A. Nicolay (USNM). New York: Ithaca, Chit- 
tenden (USNM). Ohio: Hocking Co., Cantwell Cliffs, June 1, 1941, Clem Wolfe 
(USNM); Marietta, Hubbard and Schwarz (USNM). Pennsylvania: Berks Co., 
Stony Creek Mills, June 11, July 5, D. G. Kissinger (DGK); Hummelstown, 
June 11, 1925, J. N. Knull (USNM); Lehigh Gap, July 10, 1897, G. M. Greene 
(USNM); Pen Mar, Hubbard and Schwarz (USNM). Tennessee: Clarksville, 
June 23, 1919, E. R. Jones (USNM); near Nashville, Oct. 30, 1937, R. H. Adams 
(USNM). Vermont: Brattleboro, spring 1894 (USNM). Virginia: Afton, Hub- 


346 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


bard and Schwarz (USNM); Dead Run to Turkey River, Apr. 30, 1922, wW. 1G) 
McAtee (USNM); Falls Church, May 22, 1921, E. A. Chapin (USNM); Great 
Falls, June 12, 1918, L. L. Buchanan (USNM); Petersburg, June (USNM); 
Rosslyn, Feb. 6, Chittenden (USNM); St. Elmo, Chittenden (USNM). West 
Virginia: Berkley, Hubbard and Schwarz (USNM); Harper’s Ferry, May 19, 
Hubbard and Schwarz (USNM). 

Remarks: Blatchley and Leng (1916) record this species as occur- 
ring on black locust with Apion nigrum Herbst. 


Apion (Trichapion) rufipenne Gyllenhal 


FIGuReE 13,0,p 


Apion rufipenne Gyllenhal, in Schoenherr, Genera et species curculionidum, vol. 
5, p. 397, 1839.— Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 31, 1911. 
Apion rufinulum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, Cur- 

culionidae: Apioninae, vol. 4, pt. 3, p. 69, pl. 2, fig. 24, 1890—Wagner, Mém. 
Soc. Ent. Belgique, vol. 19, p. 31, 1911. 
Apion semicastaneum Faust, Stettiner Ent. Zeitschr., vol. 54, p. 320, 1893.— 
Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 31, 1911. 
Description: Length, 2.38 to 2.62 mm.; width, 1.12 to 1.18 mm. 
Robust. Beak, antennae, head and prothorax black; elytra, abdo- 
men, mesothorax and metathorax, coxae, and legs reddish. Pubes- 
cence white, fine, dorsal surface nearly glabrous; apex of elytra, head 
and beak sparsely clothed; denser on anterior surface of front coxae, 
sides of mesothorax, and metepisternum. Beak of male as long as 
head and prothorax combined, one-half longer than prothorax, shghtly 
curved; apical four-sevenths subcylindrical, not expanded laterally at 
antennal insertion; basal five-sevenths with coarse, rather close punc- 
tures bearing scales; apical region smoother, bare; dorsal surface with 
sparse, shallower punctures. Beak of female longer than head and 
prothorax combined, four-fifths longer than prothorax, slightly, evenly 
curved; subcylindrical, slightly attenuate distad of antennal insertion; 
basal fifth with sparse pubescence, moderate punctures; apical region 
finely, sparsely punctured. Antennae inserted at basal fifth of beak, 
of male at distance from eye equal to width of frons; first segment 
slightly shorter than next three; second segment shorter than next 
two; club 0.30 by 0.11 mm. Eyes moderately prominent; frons wider 
than dorsal tip of beak, flat medially. Prothorax at base one-third 
wider than long, middle narrower than base, apex three-fourths as 
wide as base; sides beyond basal lateral expansion nearly parallel to 
slightly distad of middle, thence rounded to apex which is not con- 
stricted; in profile dorsal surface moderately convex; punctation 
shallow, moderately sparse but irregularly placed; basal fovea lacking. 
Elytra at humeri two-fifths wider than prothorax at base, 2.8 times 
as long as prothorax, length to width as 13:10; intervals flat, three 
to four times wider than punctures marking striae, nearly impunctate; 


SUBGENUS TRICHAPION—KISSINGER 347 


striae are not complete, 9 indicated beyond middle, striae are indicated 
by a row of well separated punctures, somewhat impressed on apex 
of elytra. Scutellum triangular, 0.09 by 0.09 mm. Front femora 
four times as long as wide. Claws with acute basal tooth. 

Special male characters: Mucro of tibia 2 smaller than that on 
tibia 3, both strongly dentate. 

MATERIAL EXAMINED: 10 specimens including some determined by 
Hans Wagner. 

KNOWN DISTRIBUTION: 

Mexico: Guerrero: Chilpancingo. Tabasco: Teapa. Veracruz: 20 kilometers 
south of Catemaco. 

British Honpuras: Belize. 

GUATEMALA: Panajachel; Mirandilla; Zapote; Aceituno; San Gerénimo; San Juan 
in Vera Paz; Cahabén; Chiacam. 

NicaraGcua: Chontales. 

PanaMa: David. 

Cotomsia: No exact locality. 

VENEZUELA: San Esteban. 

Remarks: I have followed the synonymy of Wagner (1911). 
Characters separating this species from A. evustum Kissinger are 
summarized in the key. 


Apion reconditum Group 


The 15 species assigned to this group have characters in common as 
discussed above. In addition the frons is narrow, being slightly wider 
than the dorsal tip of the beak, eyes are prominent, elytral humeri are 
prominent, elytral intervals are generally smooth and_ uniserially 
pubescent, and the scales in the striae tend to be more conspicuous 
than those on the intervals. 

Three species assigned to this group, A. acupunctatum Sharp, A. 
davidis Sharp, and A. nitidirostre Sharp, are known to the author 
only from females. There is naturally some question as to whether 
these three species actually belong to Trichapion, but this can be 
settled only when males are available. A. davidis and A. nitidirostre 
lack the basal lateral expansion and basal fovea of the prothorax. 
The beak of the former is very strongly, evenly curved. 

In general the antennae are inserted close to the base of the beak 
at a distance from the eyes equal to the width of the frons, but in A. 
minor Smith, A. gracilirostre Sharp, and A. bicolor Gerstaecker the 
antennae are inserted more distad on the beak at a distance from the 
eyes distinctly greater than the width of the frons. .A. minor is a small 
species, 1.5 mm.; the male has an attenuate beak about as long as head 
and prothorax combined; the dorsal surface of the prothorax is deeply, 
closely, and coarsely punctured; and the mucrones of the male are 
long, straight, and slender. A. gracilirostre is larger, 2.25 to 2.50 mm., 


348 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


the beak of the male is one-fourth longer than head and prothorax, 
and the antennae are inserted at a distance from the eye four times as 
great as width of frons, and the prothorax is finely punctured. A. bi- 
color has the prothorax and elytra strongly bronzed, the beak is slightly 
longer than head and prothorax combined, the antennae are inserted 
at a distance from the eye one-half greater than width of frons, and 
the prothorax is finely punctured. 

A. nanulum, new species, has the head strongly humped above the 
eyes. The heads of the other species are nearly flat. A. subrufum 
Sharp and A. subtinctum Fall are distinct because of the reddish elytra 
and yellowish legs. The beak of the male of A. subrufum is a little 
longer than the head and prothorax combined while the beak of the 
male of A. subtinctum is about as long as the head and prothorax. 

A. pervicaz Fall and A. importunum Fall are comparatively shorter 
and more compact than the other species. The mucrones of the male 
of the former are very small while those of the latter are long and 
prominent. The elytral intervals of A. oscillator Sharp are convex 
while those of A. reconditum Gyllenhal and A. managuense are flat. 
A. reconditum differs from A. managuense in that the pubescence of the 
sides of the prothorax is about as coarse as that on the dorsal surface 
and the beak is slightly expanded at the antennal insertion. The 
pubescence on the sides of the prothorax of the latter species is dis- 
tinctly coarser than that on the dorsal surface and the beak is strongly 
expanded at the antennal insertion. 

A. subsequens, new species, differs from A. reconditum by its more 
acutely prominent eyes, more shallowly punctured prothorax, and 
scales in striae being not larger than those on elytra intervals. 

This group appears to be closely related to the A. patruele group 
but its members differ in the more slender shape, the shallower basal 
fovea of the prothorax, the narrower frons, and the slight expansion of 
the beak over the antennal insertion. A. pervicax and A. importunum 
make a close approach to the A. patruele group. A. minor, with its 
attenuate beak, seems to make an approach to the A. rostrum group. 


Apion (Trichapion) acupunctatum Sharp 


Ficurs 14,a,b 


Apion acupunctatum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 51, 1890. 

Description: Length, 1.75 mm. 

Moderately robust. Black; pubescence inconspicuous, very fine, 
white, uniform. Female beak one-fifth longer than head and pro- 
thorax combined, moderately, evenly curved; basal one-third cylindri- 
cal, very gradually attenuate to apex; fine, sparsely punctured through- 
out. Antennae of female inserted at basal one-fourth of beak, at dis- 


SUBGENUS TRICHAPION—KISSINGER 349 


Ficure 14.—a, b, Apion acupunctatum Sharp: a, entire dorsal view of female; }, lateral 
view of head and prothorax of female. c, d, A. davidis Sharp: c, lateral view of head and 
prothorax of female; d, entire dorsal view of female. e¢, h, 0, p, A. gracilirostre Sharp: 
e, lateral view of head and prothorax of male; h, dorsal view of head of male; 0, mucro 
of tibia 2 of male; p, mucro of tibia 3 of male. f, g, 4. bicolor Gerstaecker: f, lateral view 
of head and prothorax of female; g, lateral view of head and prothorax of male. 1-k, q, 
Apion importunum Fall: i, lateral view of head and prothorax of male; 7, lateral view of 
head and prothorax of female; &, dorsal view of head of male; g, mucro of tibia 3 of male. 
l-n, A. managuense Wagner: /, dorsal view of head of male; m, lateral view of head and 
prothorax of female; 7, lateral view of head and prothorax of male. Line equals 0.50 mm. 


350 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


tance from eye one-half greater than width of frons; first segment as 
long as next three, second segment slightly longer than third, club 0.18 
by 0.08 mm. Eyes very prominent; frons wider than dorsal tip of 
beak, with a smooth, flat median area and one or two irregular lateral 
rows of punctures. Prothorax at base one-third wider than long, 
middle slightly narrower than base, apex three-fourths as wide as base; 
sides beyond slight basal lateral expansion parallel to middle, rounded 
to constricted apex; in profile dorsal surface slightly, evenly arcuate; 
punctation deep, 0.03 mm. in diameter, interspaces irregular, from 
one-half as wide to equal to diameter of punctures; basal fovea shallow, 
short. Elytra at humeri one-third wider than prothorax at base, 
2.75 times as long as prothorax, length to width as 9 : 7; intervals. 
somewhat more than twice as wide as striae, flat, with one row of in- 
conspicuous scales; striae fine, shallow. Scutellum triangular, 0.05 by 
0.05 mm., not furcate. Front femora about 3.3 times as long as wide. 
Claws with acute basal tooth. 

MATERIAL EXAMINED: Two females determined by Sharp. 

KNOWN DISTRIBUTION: 


Panama: David (type, BMNH). 


Apion (Trichapion) bicolor Gerstaecker 


Ficure 14,f,g 


Apion bicolor Gerstaecker, Stettiner Ent. Zeitschr., vol. 15, p. 244, 1854.— Wagner, 
Mém. Soc. Ent. Belgique, vol. 19, p. 31, 1911. 

Apion mandoni Wencker, Ann. Soc. Ent. France, vol. 3, p. 178, 1863.—Wagner, 
Mém. Soc. Ent. Belgique, vol. 19, p. 31, 1911. 

Apion guatemalenum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 50, 1890.— Wagner, Mém. Soc. 
Ent. Belgique, vol. 19, p. 31, 1911. 

Apion sagax Faust, Stettiner Ent. Zeitschr., vol. 54, p. 321, 1893—Wagner, Mém. 
Soe. Ent. Belgique, vol. 19, p. 31, 1911. 

Description: Length, 1.88 to 2.06 mm. 

Moderately robust. Black; elytra and prothorax with strong 
bronzy luster, base of antennae piceous; pubescence on dorsal surface 
of prothorax and elytra inconspicuous, very fine, white, coarser and 
denser on sides of mesothorax and metathorax. Beak of male slender, 
a little longer than head and prothorax combined, nine-tenths longer 
than prothorax, moderately, evenly curved; not expanded at antennal 
insertion, in lateral view basal fourth stout, parallel-sided, attenuate 
to apical third which is nearly parallel-sided; in dorsal view expanded 
slightly at tip; punctures arranged in rows, fine dorsally, coarser later- 
ally, surface nearly polished, shining, sparsely pubescent in basal 
third. Female beak long and slender, one-third longer than head and 
prothorax combined, about twice as long as prothorax, strongly, 
evenly curved; in lateral view slightly dilated beyond basal fourth, 


SUBGENUS TRICHAPION—KISSINGER 351 


slightly attenuating to apex; in dorsal view nearly parallel-sided, tip 
slightly expanded; finely, sparsely punctured, polished, with minute 
pubescence basally. Antennae inserted at distance from eye one-half 
greater than width of frons, of male at basal fifth, of female slightly 
distad of basal sixth; first segment equals next three, second segment 
shorter than next two, club 0.24 by 0.08 mm. Eyes moderately promi- 
nent; frons narrow, of male slightly narrower than, of female about as 
wide as dorsal tip of beak, median sulcus variable, usually obsolete, 
with one lateral row of punctures. Prothorax at base one-third 
wider than long, middle slightly narrower than base, apex three- 
fourths as wide as base; sides beyond basal lateral expansion nearly 
parallel to middle, rounded to constricted apex; in profile dorsal 
surface slightly convex; punctation moderately deep, 0.02 to 0.03 mm. 
in diameter, interspaces less than diameter of punctures; basal fovea 
narrow, elongate, shallow. Elytra at humeri two-fifths wider than 
prothorax at base, about 2.8 times as long as prothorax, length to 
width as 9 : 7; intervals flat, twice as wide as striae, smooth, with one 
row of minute, distant punctures bearing minute scales; striae deep, 
fine. Scutellum rounded, 0.06 by 0.06 mm., with slight median 
sulcus. Front femora about 3.3 times as long as wide. Claws with 
acute basal tooth. 

Special male characters: Tibiac | 2 and 38 armed with mucrones; 
mucro 2 shorter than mucro 3, both denticulate. 

MATERIAL EXAMINED: One nelle and three females determined as 
A. guatemalenum by Sharp and five other specimens. 

KNOWN DISTRIBUTION: 

Honpuras: Morazan: Cerro Uyuca. 

GuateMa.a: Cerro Zunil; San Gerénimo; El Tumbador; Senaht. 

VENEZUELA: Colonia Tovar. 


Boutvia: Sorata. 
CotomsiA: No exact locality. 


Apion (Trichapion) davidis Sharp 


Ficure 14,c¢,d 


Apion davidis Sharp, Biologia Centrali-Americana, Insecta, Coleoptera, Cur- 
culionidae: Apioninae, vol. 4, pt. 3, p. 53, 1890. 


Description: Length, 2.0 mm.; width, 1.0 mm. 

Robust. Black; nearly glabrous on dorsal surface; sides of meso- 
thorax and metathorax with noticeable, white pubescence. Beak of 
female slightly longer than head and prothorax combined, four-fifths 
longer than prothorax, strongly, evenly curved; cylindrical beyond 
antennal insertion; with a few punctures arranged in rows in basal 
third, beyond antennal insertion polished and nearly impunctate. 
Antennae inserted a little behind basal sixth, at distance from eyes 


S02 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


slightly less than width of frons; first segment equals next three, 
second segment equals next two, club 0.21 by 0.07 mm. Eyes promi- 
nent; frons wider than dorsal tip of beak, flat medially. Prothorax 
at base one-fourth wider than long, middle narrower than base, apex 
three-fourths as wide as base; basal lateral expansion lacking, sides 
evenly converging to middle, then slightly rounded to strongly con- 
stricted apex, apical lateral margin flared out; in profile dorsal surface 
is shghtly arcuate; punctation about 0.02 mm. in diameter, rather 
deep, irregularly placed, interspaces wider basally and laterally; basal 
fovea lacking. Elytra at humeri one-third wider than prothorax at 
base, three times as long as prothorax, length to width as 5:4; 
intervals about three times as wide as striae, flat, smooth, nearly 
impunctate; striae fine, inner margin more pronounced than outer 
margin. Scutellum triangular, 0.06 by 0.06 mm., with shallow median 
furrow. Front femora four times as long as wide. Claws with acute 
basal tooth. 

MATERIAL EXAMINED: One female compared with type by G. A. K. 
Marshall and five other specimens. 

KNOWN DISTRIBUTION: 

Panama: David (type BMNH); Porto Bello, Feb. 28, 1901, E. A. Schwarz 
(USNM). 

Apion (Trichapion) gracilirostre Sharp 


FicurE 14,e,h,0,p 


Apion gracilirostre Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 57, pl. 2, fig. 22, 1890. 

Description: Length, 2.25 to 2.50 mm. 

Moderately robust. Black; pubescence fine, white, sparse, coarser 
and denser on sides of mesothorax and metathorax. Male beak one- 
fourth longer than head and prothorax, distinctly curved, slender, 
basal third cylindrical, apical two-thirds depressed; dorsally polished 
and with fine, sparse punctures, laterally with irregular rows of coarser, 
denser punctures. Antennae inserted at basal fourth at distance from 
eye four times as great as width of frons; first segment slightly shorter 
than next five segments, second segment shorter than next two, club 
0.23 by 0.07 mm. Eyes prominent; frons narrow, equal to dorsal tip 
of beak, with a shallow median depression and a lateral row of con- 
fluent punctures. Prothorax at base one-fourth wider than long, 
middle about equal to base, apex seven-tenths as wide as base; with 
moderate basal lateral expansion, sides expanded slightly from base to 
middle, rounded to constricted apex; in profile dorsal surface nearly 
flat, apex slightly depressed; punctation moderate, 0.03 mm. in 
diameter, deep, interspaces slightly narrower than diameter of punc- 
tures, alutaceous, flat; basal fovea deep, punctiform at base, with a 


SUBGENUS TRICHAPION—KISSINGER 353 


fine impressed line extending one-fourth length of prothorax. Elytra 
at base two-fifths wider than prothorax at base, three times as long 
as prothorax, length to width as 3 : 2; intervals one-half wider than 
striae, nearly flat, dull, alutaceous, with one row of punctures with 
very fine scales; striae deep, with one row of scales more conspicuous 
than those on intervals. Scutellum rounded, 0.06 by 0.08 mm., with 
slight median depression. Front femora four times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with mucrones; 
second mucro short, subangulate; third mucro long, curved, simple. 

Female not available for study. 

MATERIAL EXAMINED: Five males, one compared with type by 
G. A. K. Marshall. 

KNOWN DISTRIBUTION: 

Mexico: Guerrero (type BMNH). Puebla: Huauchinango, June 1954, D. G. 
Kissinger (DGK). Michoacén: 20 miles east of Morelia, Mar. 7, 1958, D. G. 
Kissinger (DGK). 

Honpvuras: Tegucigalpa, Dec. 8, 1917, F. J. Dyer (USNM). Morazan: Cerro 
Uyuca, 5,500—5,700 feet, July 20, 1948, T. H. Hubbell (UM). 

Remarks: Specimens from Huauchinango, Puebla were taken 
while beating in a scrubby oak and pine forest at about 5,000 feet 
elevation. 


Apion (Trichapion) importunum Fall 
Figure 14,7-k,q 
Apion importunum Fall, Trans. Amer, Ent. Soc., vol. 25, p. 146, 1890. 


Description: Length, 1.75 mm.; width, 0.85 mm. 

Moderately robust. Black; pubescence sparse, white, rather 
coarse, coarser and more conspicuous laterally and ventrally. Male 
beak shorter than head and prothorax combined, one-third longer 
than prothorax; more strongly deflexed in apical third; in lateral 
view nearly parallel-sided ; in dorsal view expanded laterally at antennal 
insertion, attenuate to apical third, slightly compressed there, apex 
nearly parallel; basal half dull, alutaceous, with scant pubescence, a 
row of confluent punctures above antennal insertion, apical half 
shining, sparsely, finely punctured. Female beak slightly shorter 
than head and prothorax, one-half longer than prothorax; slightly, 
evenly curved; apical two-thirds nearly cylindrical, slightly expanded 
laterally at antennal insertion; nearly smooth, shining beyond antennal 
insertion, finely sparsely punctate. Antennae inserted at distance 
from eye equal to width of frons; of male at basal fourth of beak, of 
female at basal fifth; first segment as long as next two, second shorter 
than next two, club 0.17 by 0.07 mm. Eyes moderately prominent; 
frons wider than dorsal tip of beak. Prothorax at base about one-fifth 


354 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


wider than long, middle about equal to base, apex three-fourths as 
wide as base; sides beyond basal lateral expansion slightly expanding 
to middle, rounded to constricted apex; in profile dorsal surface nearly 
flat; punctation shallow, 0.03 mm. in diameter, interspaces about 
equal to diameter of punctures, alutaceous; basal fovea moderately 
deep, punctiform. Elytra at humeri one-third wider than prothorax 
at base, 2.6 times as long as prothorax, length to width as 13 : 10; 
intervals twice as wide as striae, somewhat convex, with one row of 
rather coarse scales, with irregular, fine, transverse rugae; striae fine, 
with deep, coarse, well separated punctures; striae rather shallow. 
Scutellum triangular, 0.06 by 0.06 mm., with slight median furrow. 
Front femora about 3.5 times as long as wide. Claws with acute 
basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, subangulate mucrones. 

Types: I hereby designate as lectotype of this species the male 
specimen (MCZ 25106) in the Fall Collection labeled Georgia. Co- 
types are in the J. L. LeConte Collection labeled Lake Harney, Fla., 
May (MCZ 373) and in the U. S. National Museum labeled Lake 
Harney, Fla., May, Hubbard and Schwarz (USNM 4217). 

MATERIAL EXAMINED: Lectotype and 7 specimens. 

KNOWN DISTRIBUTION: 


UniTEpD States: Georgia: No exact locality (MCZ). Florida: Hilliard, June 8, 
1930, J. O. Nottingham (UK); Jacksonville, Hubbard and Schwarz (USNM); 


’ 


Lake Harney, May 4, Hubbard and Schwarz (MCZ, USNM). Mississippi: 
Fulton, July 14, 1930, R. H. Beamer (UK). 


Apion (Trichapion) managuense Wagner 


Ficure 14,l-n 


Apion (Trichapion) managuense Wagner, Arch. Naturg. Berlin, vol. 78, 
p. 110, 1912. 

Description: Length, 1.6 to 1.9 mm. 

Moderately robust. Black; elytra slightly aeneous; pubescence 
moderate, fine, whitish, sparse, coarser and denser on sides of meso- 
thorax and metathorax. Male beak as long as head and _ pro- 
thorax combined, deflexed in middle, strongly expanded laterally and 
ventrally at antennal insertion, attenuate to middle, nearly cylindrical 
to apex; in basal two-thirds moderately punctured and sparsely 
pubescent, apex smooth, shining. Female beak slightly longer than 
head and prothorax, moderately strongly curved, basal fifth stout, 
cylindrical, in dorsal view beak at middle two-thirds as wide as at base, 
apical four-fifths nearly cylindrical; basal fifth with scant pubescence, 
moderately punctured, apical four-fifths smooth, glabrous, shining. 
Antennae of male inserted at distance from eye equal to width of 


SUBGENUS TRICHAPION—KISSINGER 355 


frons, at basal fifth of beak, of female at distance from eye about 
two-thirds as great as width of frons, at basal one-seventh; first 
segment as long as next two; second segment shorter than next two; 
club 0.22 by 0.10 mm. Eyes moderately prominent; frons narrow, 
wider than dorsal tip of beak, with a shallow, median depression and 
an irregular lateral row of moderate punctures. Prothorax at base 
one-fourth to one-third wider than long, middle slightly narrower 
than base, apex three-fourths as wide as base; sides slightly expanded 
laterally at. base, nearly parallel to middle, rounded to constricted 
apex; in profile dorsal surface slightly, evenly arcuate; punctation 
moderate, deep, 0.03 mm. in diameter, interspaces narrower than 
diameter of punctures, flat, alutaceous; basal fovea deep, short. 
Elytra at humeri one-half wider than prothorax at base, three times 
as long as prothorax, length to width as 9 : 7; intervals twice as wide 
as striae, flat, alutaceous, with one row of punctures bearing minute 
scales; striae deep, with one row of fine scales coarser than those on 
intervals. Scutellum roundly triangular, 0.06 by 0.06 mm., with 
median furrow. Front femora 3.5 times as long as wide. Claws with 
acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, blunt 
mucrones. 

MaTERIAL EXAMINED: Male and female determined by Hans 
Wagener. 

IKNOWN DISTRIBUTION: 

Nicaracua: Managua (type, Solari Collection). 

Mexico: Tabasco: Teapa (BMNH). 

GuaATEMALA: Capetillo; San Gerénimo (BMNH). 

Panama: ‘‘San Felice’? (BMNH). 


Apion (Trichapion) minor Smith 


Figure 15,a—c 


Apion minor Smith, Trans. Amer. Ent. Soc., vol. 11, p. 56, 1884.—Fall, Trans. 
Amer. Ent. Soc., vol. 25, p. 145, pl. 4, fig. 10, 10a, 1898. 

Description: Length, 1.50 to 1.67 mm.; width, 0.67 to 0.71 mm. 

Moderately slender. Black; pubescence conspicuous, white, mod- 
erately fine, not much denser laterally. Male beak shorter than 
head and prothorax combined, one-fifth longer than prothorax, 
slightly curved; attenuate from laterally expanded antennal insertion 
to tip; basal two-thirds dull, alutaceous, sparsely pubescent behind 
antennal insertion; apical third smoother, shining, glabrous. Female 
beak slightly longer than head and prothorax combined, three-fourths 
longer than prothorax, slightly curved; slightly attenuate beyond 
slight lateral expansion over antennal insertion to apical third, thence 
nearly cylindrical to apex; basal three-fourths dull, alutaceous, apical 


356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


nN 


Ficure 15.—a-c, Apion minor Smith: a, entire dorsal view of male; b, lateral view of head 
and prothorax of male; c, mucro of tibia 3 of male. d, 0, 4. nanulum, new species: d, 
lateral view of head and prothorax of male; 0, mucro of tibia 3 of male. e, f, 4. niti- 
dirostre Sharp: e, lateral view of head and prothorax of female; f, entire dorsal view of 
female. g—1, A. oscillator Sharp: g, lateral view of head and prothorax of male; h, mucro 
of tibia 2 of male; 7, mucro of tibia 3 of male. j-l, A. reconditum Gyllenhal: j, lateral 
view of head and prothorax of male; &, lateral view of head and prothorax of female; J, 
entire dorsal view of male. m, n, A. subrufum Sharp: m, lateral view of head and pro- 
thorax of male; 1, entire dorsal view of male. 9, g, 4. subsequens, new species: , lateral 
view of head and prothorax of male; q, dorsal view of head of male. Line equals 0.50 mm. 


SUBGENUS TRICHAPION—KISSINGER 357 


fourth smooth, shining. Antennae inserted at distance from eye 
one-half greater than width of frons, of male at basal third, of female 
at basal fourth of beak; first and second segments each equal to next 
two, club 0.17 by 0.07 mm. Eyes moderately prominent; frons wider 
than dorsal tip of beak. Prothorax at base slightly wider than long, 
middle about equal to base, apex three-fourths as wide as base; sides 
beyond slight basal lateral expansion slightly expanding to middle 
then rounded to constricted apex; in profile dorsal surface nearly 
flat; punctures deep, 0.03 mm. in diameter, interspaces about one-half 
diameter of punctures, slightly alutaceous; basal fovea deep, puncti- 
form. Elytra at humeri one-third wider than prothorax at base, 2.6 
times as long as prothorax, length to width as 13: 8.5; intervals 
about twice as wide as striae, rather convex, with one row of punc- 
tures; striae fine, moderately deep. Scutellum triangular, 0.06 by 
0.06 mm., with moderate median furrow. Front femora slightly 
more than three times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, simple 
mucrones. 

MATERIAL EXAMINED: Five specimens including material in the 
Fall Collection. 

KNOWN DISTRIBUTION: 

Unitep States: Jowa: Dickinson, Johnson, and Story Counties; Iowa City, 
H. F. Wickham (USNM); Lake Okoboji, July 24, Aug. 26, 1916, L. L. Buchanan 
(USNM). Missouri (USNM). Nebraska (USNM). Kansas: Topeka (USNM). 

Remarks: No types of this species were found in either the J. L. 
LeConte Collection or the U.S. National Museum, the two collections 
containing most of Smith’s types. It is necessary therefore to follow 
Fall (1898) in his interpretation of this species. 

Fall (1898) did not list distribution records for this species, evidently 
through an oversight. Leng (1920) gives the locality of this species 
as Lower California, which is certainly an error. 


Apion (Trichepion) nanulum, new species 
Figure 15,d,o 


Description: Length, 1.75 to 1.80 mm.; width, 0.80 to 0.87 mm. 
Moderately robust. Black, prothorax and elytra aeneous; pubes- 
cence inconpsicuous, white, very fine, slightly coarser and more 
conspicuous on sides of prothorax, mesothorax, and -metathorax. 
Beak of male shorter than head and prothorax combined, one-third 
longer than prothorax, moderately curved; in side view nearly parallel- 


358 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


sided throughout; in dorsal view slightly expanded at antennal inser- 
tion, attenuate to apical third, there slightly compressed, tip parallel; 
alutaceous, basal two-thirds very sparsely pubescent, sparsely 
punctured, tip smooth. Beak of female shorter than head and 
prothorax combined, slightly longer than beak of male and more 
slender, moderately curved; nearly cylindrical, in side view very 
slightly attenuate toward tip; nearly glabrous, smoother and more 
shining than that of male. Antennae of male inserted at basal fourth 
of beak at distance from eye one-third greater than width of frons, of 
female inserted at basal fifth at distance from eye equal to width of 
frons; first segment equals next two; second segment shorter than 
next two; club 0.16 by 0.06 mm., eyes prominent; frons equals dorsal 
tip of beak, with indistinct median furrow. Prothorax slightly wider 
than long, middle slightly wider than base, apex 0.85 as wide as 
base; sides beyond slight basal lateral expansion expanding to widest 
point at middle, rounded to slightly constricted apex; in profile 
dorsal surface shghtly arcuate; punctation 0.02 mm. in diameter, 
shallow, interspaces much greater than diameter of punctures, 
minutely alutaceous; basal fovea shallow, punctiform. Elytra at 
humeri two-fifths wider than prothorax at base, 2.75 times as long 
as prothorax, length to width as 9 : 6.5; intervals flat, intervals 2 and 
4 more than three times as wide as striae, other intervals somewhat 
narrower, with one row of indistinct punctures, surface smooth; 
striae shallow, very fine. Scutellum narrowly elongate-triangular, 
0.08 by 0.04 mm., convex. Front femora 3.6 times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short mucrones. 

Tyres: Holotype male (USNM 64129), Tampico, Tamaulipas, 
Mexico, December 12, E. A. Schwarz. Allotype female, same data 
as holotype (USNM). 


Apion (Trichapion) nitidirostre Sharp 
Ficure 15,e,f 


Apion nitidirostre Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 49, 1890. 

Description: Length, 1.88 mm. 

Robust. Black, elytra faintly bluish; dorsal surface of prothorax 
and elytra nearly glabrous, sides of mesothorax and metathorax with 
dense, white, fine scales. Beak of female longer than head and 
prothorax combined, nearly two times as long as prothorax, moder- 
ately, evenly curved; stout and nearly parallel-sided in basal one-fifth, 
then attenuate, apical three-fourths nearly cylindrical; polished and 
shining throughout, slightly punctured and minutely pubescent in 
basal fourth; apical three-fourths impunctate, bare. Antennae 


SUBGENUS TRICHAPION—KISSINGER 359 


inserted at basal sixth, at distance from eye equal to width of frons; 
first segment slightly shorter than next two, second segment slightly 
shorter than next two, club 0.22 by 0.08 mm. Eyes prominent; 
frons narrow, about equal to dorsal tip of beak, with one row of coar se 
punctures on either side of a rather broad, concave, median area, 
Prothorax at base one-third wider than long, middle three-fourths 
as wide as base, apex seven-tenths as wide as base; basal lateral 
expansion lacking, sides converging from base to middle, then stron gly 
rounded and constricted, lateral apical margin expanded, dorsal 
apical margin emarginate medially; in profile dorsal surface slightly 
arcuate; punctation minute (0.02 mm. in diameter), shallow, sparse, 
interspaces one to three times as wide as punctures; basal fovea 
lacking. Elytra at humeri two-fifths wider than prothorax at base, 
three times as long as prothorax, length to width as 5 :4; intervals 
flat, four to five times as wide as striae, smooth, with minute ground 
sculpture, with two rows of minute punctures, apparently glabrous; 
striae very fine, moderately deep, punctures vague, widely separated, 
apparently glabrous. Scutellum triangular, 0.06 by 0.06 mm., not 
furcate. Front femora 3.8 times as long as wide. Claws with acute 
basal tooth. 


MATERIAL EXAMINED: One female determined by Sharp. 
KNOWN DISTRIBUTION: 


Panama: Bugaba (type, BMNH). 


Apien (Trichapion) oscillator Sharp 


Figure 15,g-7 


Apion oscillator Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 64, 1890. 

Description: Length 1.6 to 2.0 mm. 

Moderately robust. Black, slightly aeneous; pubescence scant, 
fine, slightly denser on sides of mesothorax and metepisternum. 
Male beak equal to head and prothorax combined, moderately, evenly 
curved; at basal fourth moderately stout, cylindrical, attenuate to 
middle, apical half nearly cylindrical; finely, sparsely, deeply punctate, 
a coarse lateral row of punctures above antennal insertion. Antennae 
inserted at basal sixth of beak at distance from eye equal to width of 
frons; first segment slightly shorter than next two, second segment 
slightly shorter than next two, club 0.18 by 0.08 mm. Eyes promi- 
nent; frons narrow, equal to dorsal tip of beak, with a shallow median 
depression and a lateral row of coarse punctures. Prothorax at base 
more than two-fifths wider than long, at middle slightly narrower than 
base, apex seven-tenths as wide as base; sides expanded laterally at 
base, nearly parallel to middle, rounded to constricted apex; in profile 

483661—59—8 


360 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


dorsal surface moderately convex, flattened basally and apically; 
punctation deep, 0.03 to 0.04 mm. in diameter, interspaces usually 
about one-half as great as diameter of punctures, with a narrow 
impunctate median area; basal fovea moderately deep, elongate, oval. 
Elytra at humeri two-fifths wider than prothorax at base, more than 
2.8 times as long as prothorax, length to width as 7.5 : 6; intervals 
one-half wider than striae, moderately convex, smooth, alutaceous, 
with one row of very fine, distant punctures, apparently glabrous; 
striae deep, with one row of conspicuous scales. Scutellum rounded, 
triangular, 0.06 by 0.08 mm., smooth. Front femora four times as 
long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 2 armed with a moderately long, 
dentate mucro; tibia 3 armed with a moderately long, subangulate 
mucro. 

MATERIAL EXAMINED: One male determined by Hans Wagner, 
two broken specimens determined by Sharp, and one other male. 

KNOWN DISTRIBUTION: 

Mexico: Tabasco: Frontera (type, BMNH). 


GuatremaLa: Guatemala City (BMNH). British Honpvuras: “M-tee” dis- 
trict, May 25, 1906, Bowditch Collection (MCZ). 


Apion (Trichapion) pervicax Fall 
Apion pervicaz Fall, Trans. Amer. Ent. Soc., vol. 25, p. 139, pl. 4, fig. 2, 14, 1898. 


Description: Length 1.4 to 1.6 mm. 

Moderately robust. Black, elytra slightly aeneous; pubescence 
scant, fine, white, nearly uniform. Male beak four-fifths as long as 
head and prothorax combined, moderately deflexed at middle, mod- 
erately expanded laterally and ventrally at antennal insertion, 
attenuate to apex; basal two-thirds finely punctured, with scant 
pubescence, finely densely alutaceous, apical third smoother, more 
shining. Female beak a little longer than head and prothorax com- 
bined, moderately, evenly curved; basal fifth moderately stout, 
attenuate to middle, apical half nearly cylindrical; basal fifth with 
scant pubescence, densely, finely alutaceous, apical third polished, 
finely punctured. Antennae inserted at distance from eye equal to 
width of frons, of male at basal fourth of beak, of female at basal fifth; 
first segment nearly as long as next two, second segment nearly as 
long as next two, club 0.15 by 0.08 mm. Eyes slightly prominent; 
frons moderately wide, with slight median sulcus and two lateral 
rows of punctures. Prothorax one-fifth wider at base than long, 
middle about equal to base, apex three-fourths as wide as base; sides 
expanded laterally at base, slightly expanded to middle, rounded to 
constricted apex; in profile dorsal surface nearly flat, apical third 
slightly deflexed; punctation deep, 0.03 mm. in diameter, interspaces 


SUBGENUS TRICHAPION—KISSINGER 361 


narrower than diameter of punctures; basal fovea deep, elongate, 
one-third length of prothorax. Elytra at humeri one-third wider 
than prothorax at base, 2.6 times as long as prothorax, length to width 
as 4:3; intervals nearly flat, twice as wide as striae, finely, trans- 
versely rugulose, with one row of fine punctures bearing fine scales; 
striae deep, with one row of scales. Scutellum triangular, 0.06 by 0.06 
mm., smooth. Front femora slightly less than three times as long as 
wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with very small 
mucrones. 

Tyres: I hereby designate the lectotype of this species as the male 
specimen (MCZ 25116) in the Fall Collection labeled Tampa, Fla. 
Cotype with same data in U. S. National Museum (USNM 4234). 

MATERIAL EXAMINED: Lectotype and two other specimens. 

KNOWN DISTRIBUTION: 


Unitep States: Florida: Hillsborough Co. (USNM); Tampa (MCZ, USNM). 


Apion (Trichapion) reconditum Gyllenhal 
FiaurE 15,j-l 


Apion reconditum Gyllenhal, zn Schoenherr, Genera et species curculionidum, vol. 
5, p. 432, 1839. 

Apion turbulentum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 56, 1884.—Fall, : 
Trans. Amer. Ent. Soc., vol. 25, p. 146, 1898.—Chittenden, Bur. Ent. Bull., 
vol. 64, p. 31, 1908.—Blatchley and Leng, Rhynchophora or weevils of 
northeastern America, p. 80, 1916.—Buchanan, Proc. Ent. Soc. Washington, 
vol. 24, p. 84, 1922—Bruhn, Great Basin Nat., vol. 8, p. 12, 1947 (mew 
synonymy). 

Description: Length, 1.54 to 2.00 mm.; width, 0.81 to 0.93 mm. 

Moderately robust. Black, slightly aeneous; pubescence scant, 
fine, white, denser on sides of mesothorax and metepisternum. Male 
beak slightly shorter than head and prothorax, moderately, evenly 
curved, slightly dilated laterally and ventrally at antennal insertion, 
attenuate to middle, apical half nearly cylindrical; laterally with rows 
of strong punctures, dorsally punctures sparse, fine, tip smoother, 
more shining; pubescence scant. Female beak a little longer than 
head and prothorax combined, moderately, evenly curved, lateral 
antennal dilation very slight, apical two-thirds nearly cylindrical; 
shining beyond antennal insertion, finely punctured. Antennae 
inserted at distance from eye equal to width of frons at basal fifth of 
beak; first segment nearly equal to next three; second segment 
shorter than next two; club 0.21 by 0.09 mm. Eyes moderately 

prominent; frons narrow, about equal to dorsal tip of beak, with a 

rather broad median sulcus, deep in middle, rather broadly V-shaped, 

with a lateral line of deep, confluent punctures. Prothorax at base 
one-fourth wider than long, middle slightly narrower than base, apex 


362 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


three-fourths as wide as base; sides moderately expanded laterally at 
base, nearly parallel to middle, rounded to constricted apex; in profile 
dorsal surface nearly flat, more convex in some females; punctation 
deep, irregular, about 0.03 mm. in diameter, interspaces irregular, 
flat, shining, alutaceous, from one-third to twice as great as diameter 
of punctures; basal fovea deep, linear, about one-third length of pro- 
thorax or shorter. Elytra at humeri two-fifths wider than prothorax 
at base, more than 2.8 times as long as prothorax, length to width as 
10 : 7; intervals at least one-half wider than striae, nearly flat, smooth, 
alutaceous, with one row of fine punctures bearing inconspicuous 
scales; striae deep, with one row of conspicuous scales. Scutellum 
triangular, 0.06 by 0.06 mm., with slight median furrow. Front 
femora 3.6 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderate, nearly 
simple mucrones. 

Typxs: I hereby designate the lectotype of A. turbulentum Smith 
as the male specimen (USNM 1260) labeled District of Columbia. 
A cotype (MCZ 372) in the J. L. LeConte Collection is labeled 
Pennsylvania. 

MATERIAL EXAMINED: Type of A. reconditum, female, labeled 
Pennsylvania in the Riksmuseum, through the kindness of. Dr. 
‘René Malaise; lectotype of A. turbulentum; and 150 other specimens. 

KNOWN DISTRIBUTION: 

Canapa: Ontario: Fisher Glen, June 12, 1931, W. J. Browne (CNC); Ottawa, 
July 4, August 25, R. De Ruette (CNC). Quebec: Montreal, July 2, 1902, C. J. 
Quellet (USNM). 

Unirep States: Alabama: “Coleta,” H. H. Smith (USNM); Corona, Walker 
Co., October 12, 1950, A. F. Archer (BDV); Langsdale, Chambers Co., H. H. 
Smith (USNM); Mobile, Apr. 18, 1910, W. D. Pieree (USNM); Pyziton, Clay 
Co., H. H. Smith (USNM); Wadley, H. H. Smith (USNM). Arkansas: Little 
Rock, Wickham (USNM); McNeil, Oct. 1, 1906, in cotton field, J. D. Mitchell 
(USNM). Connecticut: East Haddam, June 19, 1936, M. P. Zoppe (USNM). 
District of Columbia: Apr. 24, May 15 (USNM). Georgia: Okefenokee Swamp, 
Aug. 8, 1902, P. L. Richer (USNM); Rabun Co., July, Leng (USNM)._ IIlinois: 
Muncie, Aug. 31, 1907 (USNM). Indiana: Blatchley and Leng (1916) state this 
species is common throughout the State. Jowa: Iowa City, H. F. Wickham 
(USNM); Lake Okoboji, June 21, 1916, L. L. Buchanan (USNM). Kansas: 
Leavenworth Co., June 30, 1924, R. H. Beamer (UK). Kentucky: Brooklyn 
Bridge, June 29, 1925 (USNM). Louisiana: Forbing, Mar. 24, 1908, R. A. 
Cushman (USNM); Mansura, Apr. 3, 1909, W. D. Pieree (USNM). Maryland: 
Many records throughout the State. Massachusetts: Chicopee, Aug. 16, 1902, 
F. Knab (USNM); Springfield (USNM). Michigan: Detroit, May, Hubbard and 
Schwarz (USNM); Washtenaw Co., Aug. 1, 1922, M. H. Hatch (USNM). 
Mississippi: Utica, August (USNM). Nebraska: Nebraska City, June (USNM). 
New Jersey: Greenwood Lake (USN M); Orange Mt. (USNM); Pompton (USNM); 
Raritan (USNM); Springfield (USNM); Warren Co. (USNM); Woodside 
(USNM). New York: Ithaca, July 10, 1951, B. D. Valentine (BDV). North 
Carolina: Round Knob (USNM). Ohio: Adams Co., Aug. 1, 1950 (ELS); Ash- 


SUBGENUS TRICHAPION—KISSINGER 363 


land Co., Apr. 9, 1953 (ELS); Cincinnati, H. Soltau (USNM); Franklin Co., 
June 21, 1950 (ELS); Westerville, June 26, 1949 (ELS). Pennsylvania: Ash- 
bourne, May 24, 1900 (USNM); Frankford (USNM); Glenolden, June 16 
(USNM); Glenside, June 16 (USNM); Hanover, May 15, 1930 (USNM); Lehigh 
Gap, Aug. 8, 1904 (USNM). Tennessee: Gatlinburg, Sept. 20, 1941 (USNM); 
Memphis, May 20, 1916 (USNM); Nashville, Aug. 4-15, 1897 (USNM). Tezas: 
Columbus, Aug. 14, E. A. Schwarz (USNM). Virginia: Chain Bridge, May 7, 
1922 (USNM); Covington, May 6, 1950 (DGK); Fairfax Co., Dyke, May 28 
(USNM); Falls Church, July 2, 1922 (USNM); Fredericksburg, July 15, 1900 
(USNM); Nelson Co., Aug. 4, 1910 (USNM); Stone Creek, Lee Co. (USNM). 

Remarks: Chittenden (1908) records this species as developing in 
the seeds of tick-trefoil, Meibomia marylandica L. Blatchley and 
Leng (1916) state that it was beaten from flowers of panicled dogwood, 
Cornus candidissma Marsh., and swept from huckleberry. 

Apion reconditum Gyllenhal is a name that has not been recognized 
since its description in 1839. Both Smith (1884) and Fall (1898) 
treat the name as an unrecognized species. An examination of the 
type has revealed the above synonymy. 


Apion (Trichapion) subrufum Sharp 


Ficure 15,m,n 


Apion subrufum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, Cur- 
culionidae: Apioninae, vol. 4, pt. 3, p. 68, 18990. 


Description: Length, 1.67 mm. 

Moderately robust. Dark reddish; beak, head and prothorax 
darker than elytra; tibiae and tarsi light reddish yellow; pubescence 
moderately fine, white, on dorsal surface of prothorax and elytra 
inconspicuous, denser on median basal spot of prothorax; denser on 
sides of mesothorax and metathorax. Beak of male slender, slightly 
longer than head and prothorax combined, three-fifths longer than 
prothorax, slightly curved; more strongly deflexed in apical third; in 
lateral view stouter in basal fourth, attenuate, apical one-half nearly 
parallel-sided; in dorsal view moderately expanded over antennal 
insertion, apical two-thirds nearly parallel-sided; polished, with 
irregularly placed, sparse, moderately deep punctures, more strongly 
punctured toward base and laterally. Antennae of male inserted at 
basal one-eighth at distance from eye equal to width of frons; first 
segment equals next two, second segment shorter than next two, 
club 0.25 by 0.09 mm. Eyes moderately prominent; frons narrow, 
about equal to dorsal tip of beak, with a moderately wide median 
area and a shallow, wide sulcus. Prothorax at base one-fifth wider 
than long, middle about as wide as base, apex seven-tenths as wide 
as base; sides beyond minute basal lateral expansion slightly expand- 
ing to middle, rounding to constricted apex; in profile dorsal sur- 
face nearly flat; punctation 0.03 mm. in diameter, shallow, margin 


364 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


of punctures more distinct basally, interspaces irregular, from one to 
three times as wide as punctures; basal fovea absent. Elytra at 
humeri one-third wider than prothorax at base, 2.6 times as long as 
prothorax, length to width as 6:5; intervals flat, twice as wide as 
striae, with one row of distant, minute punctures bearing minute 
scales; striae moderately fine, deep, with one row of scales larger than 
those on intervals. Scutellum triangular, 0.08 by 0.08 mm., with 
no median depression. Front femora four times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, trian- 
cular-acute mucrones. 

MATERIAL EXAMINED: One male determined by Sharp. 

KNOWN DISTRIBUTION: 


GuATEMALA: La Tinta in Vera Paz (type). 
Apion (Trichapion) subsequens, new species 
FiaureE 15,p,q 


Description: Length, 1.67 to 1.91 mm.; width, 0.78 to 0.94 mm. 

Moderately robust. Black, elytra and prothorax somewhat aeneous, 
femora and tibiae piceous; pubescence very fine, white, inconspicuous 
on dorsal surface, somewhat coarser and denser on mesothorax. Beak 
of male slightly shorter than head and prothorax combined, four- 
sevenths longer than prothorax, moderately, evenly curved; apical 
third nearly cylindrical, in dorsal view attenuate from antennal 
insertion to apical third; basal two-thirds moderately dull, alutaceous, 
sparsely punctured and pubescent, apical third shining, minutely 
punctured. Beak of female longer than head and prothorax combined, 
nearly twice as long as prothorax, moderately, evenly curved; apical 
half nearly cylindrical, somewhat depressed at tip; in dorsal view 
attenuate from antennal insertion to middle; basal three-fourths 
rather strongly punctured, a distinct row of punctures above antennal 
insertion extends to middle; pubescence sparse at base. Antennae 
inserted at distance from eye equal to width of frons, of male at basal 
fourth, of female at basal sixth of beak; first segment equals next 
two, second segment shorter than next two; club 0.20 by 0.07 mm. 
Eyes acutely prominent; frons wider than dorsal tip of beak, with a 
slight, broad median depression and one lateral row of fine punctures. 
Prothorax at base one-fourth longer than wide, middle about as wide 
as base, apex three-fourths as wide as base; sides beyond obsolete 
basal lateral expansion subparallel in basal half, rounded to constricted 
apex; in profile dorsal surface is nearly flat; punctation very shallow, 
about 0.03 mm. in diameter, interspaces equal to or wider than 
punctures; basal fovea shallow, short, Elytra at humeri one-half 


SUBGENUS TRICHAPION—KISSINGER 365 


wider than prothorax at base, three times as long as prothorax, length 
to width as 9 : 6.5; intervals more than twice as wide as striae, flat, 
with one row of minute punctures bearing minute scales; striae 
fine, deep. Scutellum elongate-triangular, 0.06 by 0.04 mm., convex, 
no median furrow. Front femora four times as long as wide. Claws 
with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately long, 
subdentate mucrones, mucro of tibia 3 longer than that of tibia 2. 

Types: Holotype male (USNM 64130), Tampico, Tamaulipas, 
Mexico, Dec. 16, E. A. Schwarz. Allotype female, same locality as 
holotype, December 7 (USNM). One paratype, male, same locality 
as holotype, December 6 (DGK). 


Apion (Trichapion) subtinctum Fall 


Apion subtinctum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 138, 1898. 


Description: Length, 1.37 to 1.70 mm.; width, 0.68 to 0.82 mm. 

Moderately robust. Black with reddish bronze luster; pubescence 
scant, fine, white, denser mediobasally on prothorax and on sides of 
mesothorax and metathorax. Male beak as long as head and prothorax 
combined, slightly, evenly curved; with a moderate, lateral dilation 
over antennal insertion, attenuate to middle, apical half nearly 
cylindrical; polished, laterally with a few strong punctures at antennal 
insertion; glabrous beyond base. Female beak similar. Antennae 
inserted at distance from eye equal to width of frons, at basal one- 
sixth of beak; first segment longer than next two, shorter than next 
three; second segment shorter than next two; club 0.18 by 0.06 mm. 
Eyes prominent; frons narrow, about equal to dorsal tip of beak, with 
slight median sulcus and a lateral row of punctures. Prothorax at 
base one-sixth wider than long, middle slightly narrower than base, 
apex three-fourths as wide as base; sides moderately expanded 
laterally at base, nearly parallel to middle, rounded to moderately 
constricted apex; in profile dorsal surface nearly flat; punctation 0.01 
to 0.02 mm. in diameter, shallow, posterior margin of punctures more 
abrupt, interspaces flat, alutaceous, variable, greater than diameter of 
punctures, usually twice as wide; basal fovea shallow, linear, reduced. 
Elytra at humeri two-fifths wider than prothorax at base, 2.8 times as 
long as prothorax, length to width as 4:3; intervals about twice as 
wide as striae, flat, with one row of fine punctures bearing inconspicuous 
scales, smooth; striae deep, with one row of rather conspicuous scales. 
Scutellum triangular, 0.04 by 0.04 mm., with median furrow. Front 
femora 3.75 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately long, 
simple mucrones. 


366 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Typus: I hereby designate the lectotype of this species as the male 
specimen (MCZ 25126) in the Fall Collection labeled San Antonio, 
Tex. A cotype (USNM 4221) is labeled Columbus, Tex. 

MatTERIAL EXAMINED: Lectotype and 20 specimens. 

KNOWN DISTRIBUTION: 

UnitTep States: Texas: Brownsville (USN M); Columbus (USN M); La Paloma, 
Oct. 14, 1944 (ELS); San Antonio (MCZ); Victoria (USNM); 

Mexico: Tamaulipas: Matamoros, Mar. 23, 1942, Libby and Moreno (USNM), 
May, Fenyes Collection (CAS). Tampico, December 21, E. A. Schwarz (USNM), 

Remarks: Material in the U. S. National Museum was reared from 
blooms of Malvaviseus drummondi from Matamoros, Mexico. Also 
in the same collection is material collected at Victoria, Tex. on 
Bumelia sp. 


Apion oblitum Group 


The males of the two species placed in this group have all three 
tibiae mucronate, in addition the beak of both sexes is abruptly 
narrowed slightly distad of the antennal insertion and is polished and 
nearly cylindrical to the tip. The two species do not overlap in range. 
A. oblitum Smith ranges from Nebraska to Texas and Arizona: A. 
mediocre Sharp occurs in Mexico in the States of Guerrero and Puebla. 

The two species are rather similar in appearance. The dorsal surface 
of the prothorax of A. oblitum is deeply punctured and mucrones 2 and 
3 of the male are rather short. The dorsal surface of the prothorax of 
A. mediocre is shallowly punctured and mucrones 2 and 3 of the male 
are long. 


Apion (Trichapion) mediocre Sharp 
Figure 16,a—d 


Apion mediocre Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 

Curculionidae: Apioninae, vol. 4, pt. 3, p. 56, 1890. 

Description: Length, 1.95 to 2.50 mm. 

Moderately robust. Black; elytra dull with bluish luster, prothorax 
more shining; pubescence conspicuous, fine, white, sparse, coarser and 
denser on sides of mesothorax and metathorax. Male beak slightly 
shorter than head and prothorax, slightly curved, basal third stoutly 
cylindrical, abruptly attenuate to basal two-fifths, nearly cylindrical 
to apex; in dorsal view middle about two-thirds as wide as base; 
sparsely, finely punctate, basal third very sparsely pubescent. Female 
beak a little longer than head and prothorax, slightly deflexed at apical 
two-thirds; in dorsal view stoutly cylindrical at basal fifth, abruptly 
attenuate to basal two-fifths, cylindrical to apex; in lateral view atten- 
uating from base to slightly distad of antennal insertion, at middle 
two-thirds as wide as base; basal fifth sparsely pubescent, dull, apical 


SUBGENUS TRICHAPION—KISSINGER 367 


a ee 


a 


Ficure 16.—a-d, Apion mediocre Sharp: a, entire dorsal view of male; 6, lateral view of 
head and prothorax of male; c, lateral view of head and prothorax of female; d, mucro of 
tibia 3 of male. e¢, f, A. oblitum Smith: e, lateral view of head and prothorax of male; 
f, mucro of tibia 3 of male. Line equals 0.50 mm. 


four-fifths shining, glabrous, very finely, sparsely punctate. Antennae 
inserted at distance from eye slightly less than width of frons, at 
basal fifth of beak; first segment slightly shorter than next three; 
second segment shorter than next two; club 0.27 by 0.09 mm. Eyes 
moderately prominent; frons narrow, about as wide as dorsal tip 
of beak, with slight median depression and a lateral row of coalesced 
punctures. Prothorax at base one-fourth wider than long, middle 
slightly narrower than base, apex three-fourths as wide as base; sides 
slightly expanded laterally at base, slightly diverging to middle, 
rounded to constricted apex; in profile dorsal surface slightly, evenly 
arcuate; punctation moderately deep, 0.03 mm. in diameter, inter- 
spaces narrower than diameter of punctures; basal fovea shallow, 
linear, extending one-third length of prothorax. Elytra at humeri 
two-fifths wider than prothorax at base, 2.7 times as long as prothorax, 
length to width as 4:3; intervals twice as wide as striae, convex, 
with one row of fine scales. Scutellum rounded, 0.06 by 0.06 mm., 
with slight median furrow. Front femora slightly more than four 
times as long as wide. Claws with acute basal tooth. 

Special male characters: Three pairs of tibiae mucronate, tibia 1 
with minute mucro, tibiae 2 and 3 with long, dentate mucrones. 


368 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


MATERIAL EXAMINED: One male compared with type by G. A. K. 
Marshall and 70 specimens. 
KNOWN DISTRIBUTION: 
Mexico: Guerrero (type, BMNH). Puebla: Huauchinango, June 1954, D. G. 
Kissinger. 
Apion (Trichapion) oblitum Smith 


Ficure 16,e,f 


Apion oblitum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 54, 1884.—Fall, Trans. 
Amer. Ent. Soc., vol. 25, p. 153, 1898.—Schwitzgebel and Wilbur, Journ. 
Kansas Ent. Soc., vol. 15, p. 44, 1942. 

Apion capitatum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 54, 1884.—Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 154, 1898. 

Description: Length, 2.0 to 2.4 mm. 

Moderately robust. Black; pubescence fine, white, sparse, denser 
on mesothorax and metepisternum. Male beak shorter than head 
and prothorax combined, one-fifth longer than prothorax; deflexed at 
basal third; in lateral view abruptly narrower in front of antennal 
insertion; in dorsal view expanded laterally over antennal insertion, 
attenuate beyond dilated portion, apical third nearly cylindrical; 
basal three-sevenths dull, alutaceous, punctured, sparsely pubescent, 
apical four-sevenths dull, shining. Female beak similar to male except 
that in lateral view it is not as abruptly narrowed beyond antennal 
insertion. Antennae inserted at distance from eye equal to width of 
frons; nearly at basal fourth of beak; first segment equals next two, 
second segment shorter than next two, club 0.21 by 0.08 mm. Eyes 
moderately prominent; frons wider than dorsal tip of beak. Prothorax 
at base slightly wider than long, middle and base about equal, apex 
five-sevenths as wide as base; sides beyond basal lateral expansion 
slightly expanded to middle, rounded to moderately constricted apex; 
in profile dorsal surface slightly arcuate; punctation deep, 0.04 mm. 
in diameter, interspaces about one-half as wide as punctures, aluta- 
ceous; basal fovea deep, narrow, extending one-third length of pro- 
thorax. Elytra at humeri two-fifths as wide as prothorax, 2.8 times 
as long as prothorax, length to width as 4 : 3; intervals twice as wide 
as striae, nearly flat, generally with one row of scales; striae moderately 
deep, fine. Scutellum triangular, 0.04 by 0.04 mm., not furcate. 
Front femora 3.5 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibia 1 with minute mucro, tibiae 2 and 3 
with small, subdentate mucrones. 

Types: I hereby designate as lectotype of A. oblitum Smith the 
male specimen (USNM 1261) labeled Colorado. A cotype (MCZ 379) 
in the J. L. LeConte Collection is labeled Texas. 

I hereby designate the lectotype of A. capitatum Smith as the male 
specimen (MCZ 380) in the J. L. LeConte Collection labeled Nebraska. 


SUBGENUS TRICHAPION—KISSINGER 369 


MATERIAL EXAMINED: Lectotype and 50 specimens. 

KNOWN DISTRIBUTION: 

Unitep States: Arizona: No exact locality (USNM). Colorado: No exact 
locality (USNM); Lincoln Co., Aug. 12, 1926 (UC). Tilinois: New Canton, 
Aug. 12, 1948, Sanderson and Stannard (INHS). Kansas: Douglas Co. (UK); 
Hodgeman Co., July 17-25, 1917 (UK); Onaga July 7, 1925, R. H. Beamer (UK) ; 
Sedgwick Co., 1,291 ft., 1916, R. H. Beamer (UK); Sumner Co., 1,189 ft., 1916, 
R. H. Beamer (UK); Wallace Co., Aug. 11, 1926, W. Benedict (UK); Wichita, 
Sept. 6, 1935, R. H. Beamer (UK). Nebraska: No exact locality (USNM). 
Texas: Potter Co., July 7, 1927, R. H. Beamer (UK). 

Remarks: Although A. capitatum has line priority over A. oblitum, 
Fall (1898), as the first reviser, fixed the latter as the valid name. 

Schwitzgebel and Wilbur (1942) collected this species on ironweed, 
Veronia interior Small. 


Apion nigrum Group 


The five species comprising this group are distinct in that the 
middle of the prothorax is distinctly wider than the base and the 
sides are strongly rounded at the middle; males of these species 
exhibit more or less well developed secondary sexual modifications 
of the front legs. The five species do not overlap inrange. A.nigrum 
Herbst occurs in the eastern half of the United States; A. dolosum 
Fall occurs in the mountains of New Mexico and Arizona; A. cordatum 
Smith occurs on the Pacific Coast from Washington to southern 
California; A. heterogeneum Sharp occurs in the mountains of central 
Mexico; and A. Jativentre Béguin-Billecocq occurs in Brazil and 
Argentina. 

Of the five species, A. dolosum and A. nigrum are the most alike. 
The former differs in that the sides of the prothorax are not as strongly 
rounded, the beak of both sexes is comparatively long; the front tarsi 
of the male are not strongly dilated, and the front femora of the male 
are incrassate. A. nigrum differs from the other four species by the 
dilated, densely pubescent front tarsi of the male. The front tibiae 
of the male of A. heterogeneum are dilated, curved, and have long, 
yellowish ciliae on the inner surface. The front tibiae of the male of 
A. cordatum are minutely mucronate. A. lativentre is distinct in 
that the basal lateral region of the prothorax is prominent on either 
side of the middle. 


Apion (Trichapion) cordatum Smith 


Figure 17,a—c 
Apion cordatum Smith, Trans. Amer. Ent. Soe., vol. 11, p. 54, 1884.—Fall, Trans. 
Amer. Ent. Soc., vol. 25, p. 153, pl. 4, fig. 3, 1898. 
Description: Length, 2.00 to 2.75 mm. 
Moderately robust. Black; pubescence conspicuous, white, coarse, 


370 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


sparse, denser on sides of mesothorax. Male beak four-fifths as long 
as head and prothorax, slightly expanded laterally and ventrally at 
antennal insertion, attenuate to middle, nearly cylindrical beyond 
middle, apex three-fifths as wide as base; punctation over antennal 
insertion strong, arranged in rows, apical third finely, sparsely punc- 
tate; pubescence inconspicuous. Female beak as long as head and 
prothorax combined, similar to male. Antennae inserted at distance 
from eye equal to width of frons, at basal fourth of beak; first segment 
slightly longer than next two; second segment shorter than next two 
combined; club 0.20 by 0.09 mm. Eyes moderately prominent; 
frons moderately broad, with slight median impression, two lateral 
rows of punctures. Prothorax at base one-fifth wider than long, 
middle wider than base, apex three-fourths as wide as base; sides 
diverging to widest point at middle, then rounding to constricted 
apex; in profile dorsal surface slightly convex, flattened basally and 
apically; punctation deep, 0.03 to 0.04 mm. in diameter, interspaces 
convex, alutaceous, narrower than diameter of punctures; basal fovea 
deep, linear, extending two-fifths length of prothorax. Elytra at 
humeri one-third wider than prothorax at base, three times as long 
as prothorax, length to width as 3:2; intervals twice as wide as 
striae, somewhat convex, intervals with one or two rows of fine punc- 
tures bearing fine scales; striae deep. Scutellum triangular, 0.06 by 
0.06 mm., with slight median sulcus. Front femora 3.50 to 3.75 
times as long as wide. Claws with moderate basal tooth. 

Special male characters: Three pairs of tibiae mucronate, tibia 1 
with small, triangular mucro; tibiae 2 and 3 with moderate, sub- 
dentate mucrones. 

Lectotype: I hereby designate the lectotype of this species as the 
male specimen (MCZ 378) in the J. L. LeConte Collection labeled 
California. 

MATERIAL EXAMINED: Lectotype and 60 specimens. 

KNOWN DISTRIBUTION: 

Unirep Srates: Washington: Rochester, July 22, 1931, H. T. Peters (UK); 
Seattle, May 11, 1953, on Psoralea physodes, E. I. Smith (USNM). California: 
Colfax (UC); Bartlett Spring, Lake Co. (TLCC); Murphys, Calaveras Co., 
May 15, 1936 (CAF). 

Remarks: Material in the U. S. National Museum was associated 
with Psoralea physodes; 50 beetles were found alive on 25 plants. 


Apion (Trichapion) dolosum Fall 


Figure 17,h—-k 
Apion dolosum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 148, 1898. 


Description: Length, 2.50 mm. 


SUBGENUS TRICHAPION—KISSINGER Spl 


ZA) | 


Ficure 17.—a-c, Apion cordatum Smith: a, entire dorsal view of male; 5, lateral view of 
head and prothorax of male; c, mucro of tibia 3 of male. d-g, A. nigrum Herbst: d, 
lateral view of head and prothorax of male; ¢, mucro of tibia 3 of male; f, lateral view of 
head and prothorax of female g, entire dorsal view of male. h-k, A. dolosum Fall: h, 
mucro of tibia 3 of male; i, lateral view of head and prothorax of male; 7, lateral view of 
head and prothorax of female; &, dorsal view of head of female. Line equals 0.50 mm. 


Moderately robust. Black; pubescence evident, whitish, coarse, 
sparse, denser on sides of mesothorax. Male beak slightly shorter 
than head and prothorax, slightly, evenly curved, expanded laterally 
and ventrally at antennal insertion, attenuate from there to apex; 
basal three-fourths moderately punctured, with slight median dorsal 
carina, tip shining; sparsely pubescent in basal three-fourths. Female 
beak about one-fourth longer than head and prothorax, slightly curved, 
nearly cylindrical throughout, dull throughout, with uniform, fine, 
sparse punctation. Antennae of male inserted at distance from eye 


372 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


two-thirds greater than width of frons, of female at distance from eye 
twice as great as width of frons, in both sexes at basal one-third of 
beak; first segment of male shorter than next three, of female equal 
to next three; second segment shorter than next two; club 0.21 by 
0.10mm. Eyes moderately prominent; frons moderately wide, nearly 
flat or with vague median depression, with three lateral rows of 
punctures. Prothorax at base one-fifth wider than long, middle 
slightly wider than base, apex three-fourths as wide as base; sides 
moderately expanded laterally at base, slightly diverging to widest 
point at middle, then rounded to constricted apex; in profile dorsal 
surface nearly flat; punctures moderate, 0.03 mm. in diameter, deep, 
interspaces narrower than diameter of punctures, convex, alutaceous; 
basal fovea linear, deep at middle, shallow on each end, extending two- 
fifths length of prothorax. Elytra at humeri one-third to two-fifths 
wider than prothorax at base, 2.75 times as long as prothorax, length 
to width as 14:9; intervals twice as wide as striae, flat to slightly 
convex, with an irregular row of punctures bearing fine scales; striae 
deep. Scutellum narrow, triangular, with slight median impression. 
Front femora of female four times as long as wide, of male three times 
as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with moderately 
long, subangulate mucrones; front femora noticeably stouter than 
those of female. 

MATERIAL EXAMINED: Type, female (MCZ 25089), Williams, 
Ariz., in the Fall Collection, and 25 specimens. 

KNOWN DISTRIBUTION: 

UniTep States: New Merico: Estancia, 1925, J. R. Douglas (USNM); 
Terrence, 1926, J. R. Douglas (USNM). Arizona: Chiricahua Mts., Hubbard 
and Schwarz (USNM); Graham Mts., Graham Co., 8,000 feet, Aug. 16, 1952 
(AMNH); Pinal Mts., Gila Co. (CU); Prescott, July, G. H. Nelson (DGK); 
Williams, Hubbard and Schwarz (USNM). 

Remarks: This species has been taken on locust at Prescott, 
Arizona by H. F. Wickham (USNM) and G. H. Nelson (DGK). 
One specimen in the U. S. National Museum apparently was reared 
from a maple seed. 


Apion (Trichapion) heterogeneum Sharp 


Figure 18 
Apion heterogeneum Sharp, Biologia Centrali-Americana. Insecta, Coleoptera, 
Curculionidae: Apioninae, vol. 4, pt. 3, p. 59, pl. 3, fig. 7, 1890. 

Description: Length, 1.95 to 3.00 mm. 

Robust. Black, elytra dull, prothorax shining; pubescence fine, 
white, sparse, coarser and denser on sides of prothorax, mesothorax, 
and metathorax. Male beak slightly shorter than head and _ pro- 
thorax, very stout; slightly, evenly curved; in dorsal view as wide as 


SUBGENUS TRICHAPION—KISSINGER 373 


frons to slightly distad of middle, strongly attenuate, apical third 
parallel-sided, less than two-thirds as wide as basal portion; in lateral 
view stoutly parallel in basal half, greatly attenuate, apical third 
nearly cylindrical; basal two-thirds with moderately dense, fine 
punctures bearing fine pubescence, apical third smoother, glabrous, 
more shining. Female beak one-fourth to one-third longer than head 
and prothorax, moderately, evenly curved, slender; in dorsal view 
basal fifth stout, nearly parallel-sided, equal to width of frons, abruptly 
attenuate to basal third where it is about two-thirds as wide as at base, 
apical two-thirds cylindrical; basal fifth dull, densely alutaceous, with 
fine, dense punctures, pubescence scant, with deep, short lateral 
furrow above antennal insertion; apical four-fifths polished, very 
sparsely, minutely punctured. Antenna of male inserted at distance 
from eye two-thirds greater than width of frons, slightly behind middle 
of beak, of female at distance slightly greater than width of frons, 
at basal fifth of beak; first segment longer than next two, shorter 


ay 
bs 
yt 


h 


Ficure 18.—Apion heterogeneum Sharp: a, dorsal view of head of male; d, lateral view of 
head and prothorax of female; c, lateral view of head and prothorax of male; d, front leg 
of male; ¢, entire dorsal view of female; f, dorsal view of tarsus 2 of male; g, lateral view 
of tarsus 2 of male; 4, mucro of tibia 3 of male. Line equals 0.50 mm. 


7 


374 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


than next three; second segment as long as third; club of male, 0.34 
by 0.12 mm.; of female 0.30 by 0.12 mm. Eyes moderately promi- 
nent; frons moderately wide, with moderately deep median sulcus and 
a lateral row of punctures. Prothorax at base slightly wider than 
long, at middle one-fifth wider than base, at apex two-thirds (male) 
to four-fifths (female) as wide as base; sides slightly expanded laterally 
at base, strongly roundly expanded to middle, rounded to strongly 
constricted apex; in profile dorsal surface of male slightly arcuate, 
flattened at base and apex, of female nearly flat, apical third depressed; 
punctation moderately deep, more so toward posterior margin, 0.03 
mm. in diameter, interspaces narrower than diameter of punctures, 
alutaceous, moderately convex; basal fovea deeper at middle, linear, 
about one-fourth length of prothorax. Elytra at humeri one-half 
(male) to three-fifths (female) wider than base of prothorax, 2.5 to 
2.8 times as long as prothorax, length to width as 19:14; intervals 
twice as wide as striae, moderately convex, very densely, minutely 
punctate, with two irregular rows of slightly larger punctures bearing 
fine scales; striae deep. Scutellum triangular, 0.09 by 0.06 mm. 
Front femora about four times as long as wide. Claws with acute 
basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, sub- 
angulate mucrones, mucro on tibia 2 more curved laterally than mucro 
on tibia 3; antennae elongate, robust, with long cilia; front coxae 
enlarged, about as wide as length of antennal club; front tibiae di- 
lated, curved, set with long cilia on inner surface; third segment of 
middle tarsus very broad, with diverging lobes receiving very short, 
broad fourth segment, claws of which are abruptly curved and ap- 
plied to ventral surface of segment. 

MATERIAL EXAMINED: One male compared with type by G. A. K. 
Marshall and 20 other specimens. 

KNOWN DISTRIBUTION: 

Mexico: Guerrero: Omilteme, 8,000 ft. (type, BMNH). Michoacan: El Pueb- 
lito, Aug. 9, 1954, 6,500 feet (DGK). Distrito Federal: Desiertos de los Leones, 
May 26, 1946, J. and D. Pallister (AMNH). Puebla: Near Huauchinango, June 
1954, D. G. Kissinger. 

Remarks: Specimens were taken by the author while beating in a 
pine and scrubby oak forest at about 5,000 feet elevation near Huau- 
chinango, Puebla, Mexico. 


Apion (Trichapion) lativentre Béguin-Billecocq 


Apion lativentre Béguin-Billecocq, Ann. Soc. Ent. France, vol. 78, p. 451, 1909.— 
Wagner, Mém. Soc. Ent. Belgique, vol. 19, p. 32, 1911. 


Description: Length, 2.06 to 2.62 mm.; width, 1.00 to 1.28 mm. 
Moderately robust. Black, elytra blue, antennae piceous; pubes- 


SUBGENUS TRICHAPION—KISSINGER 375 


cence white, on dorsal surface very fine, sparse, inconspicuous, under 
eyes, on sides of prothorax, mesothorax, and metathorax longer, 
coarser, and denser. Beak of male shorter than head and prothorax 
combined, one-fourth longer than prothorax, slightly curved, more 
noticeably so toward apex; basal third stout, subcylindrical, not ex- 
panded laterally at antennal insertion, apical two-fifths subcylindrical, 
slightly depressed at tip; basal half with strong punctures bearing 
scales; apical half and dorsal surface with fine punctures. Beak of 
female slightly shorter than head and prothorax combined, three- 
fifths longer than prothorax, slightly, evenly curved; nearly cylindrical 
beyond antennal insertion; glabrous and polished beyond antennal 
insertion, with sparse, minute punctures in that region, basal region 
with sparse pubescence. Antennae inserted at distance from eyes one- 
half greater than width of frons, of male slightly behind basal third, of 
female slightly behind basal fourth; first segment of male slightly 
longer than next two, of female slightly longer than next three; 
second segment equals next two; club 0.21 by 0.07 to 0.27 by 0.09 mm. 
Eyes slightly promiment; frons equals width of dorsal tip of beak, flat 
medially, sometimes with trace of median sulcus. Prothorax with 
basal lateral region slightly anterior to basal margin prominent on 
either side of middle; at base one-fourth wider than long, middle as 
wide or slightly wider than base, apex seven-tenths as wide as base; 
sides beyond basal lateral expansion diverging slightly to middle, 
rounded to constricted apex; in profile dorsal surface fiat, immediately 
at base strongly deflexed downward; punctation 0.03 mm. in diameter, 
shallow, interspaces generally as wide as punctures, strongly aluta- 
ceous; basal fovea very deep between basal prominences, extends 
about one-third length of prothorax. Elytral sides at middle above 
ninth stria with more or less distinct depression; elytra at humeri 
two-fifths wider than prothorax at base, three times as long as pro- 
thorax, length to width as 7:5 to 4:3; intervals flat, a little more 
than twice as wide as striae, in part with two rows of scales; striae 
deep, rather fine. Scutellum triangular, 0.68 by 0.06 mm., with deep 
median furrow. Front femora about 3.7 times as long as wide. 
Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, blunt 
mucrones. 

MATERIAL EXAMINED: Specimens determined by J. Balfour-Browne 
and six other specimens. 

KNOWN DISTRIBUTION: 

ARGENTINA: Province of Buenos Aires (type) ; Santa Fé, Rio San Javier, Estancia 

La Noria, Dec. 16, 1911, to Jan. 5, 1912, G. E. Bryant (BMNH). 
ParaGuay: Independencia, June 19, 1851 (DGK). 


Brazit: Rio de Janeiro. 
483661—592 9 


376 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Apion (Trichapion) nigrum Herbst 
FicureE 17,d-g 


Apion nigrum Herbst, Natursystem aller bekanntenin- und auslandischer Insecten, 
vol. 7, p. 112; pl. 103, fig. 11, 1797.—-Germar, Mag. Ent., vol. 2, p. 239, 1817.— 
Gyllenhal, in Schoenherr, Genera et species curculionidum, vol. 1, pt. 1, 
p. 254, 1833.—Smith, Trans. Amer. Ent. Soc., vol. 11, p. 64, 1884.—Fall. 
Trans. Amer. Ent. Soc., vol. 25, p. 152, 1898.—Cotton, Ohio Nurs. Orch. 
Insp. Bull, vol. 7, p. 23, 1906.—Pierce, U. S. Dep. Agr. Bur. Ent. Bull. 
100, p. 75, 1912.—Blatchley and Leng, Rhynchophora or weevils of north- 
eastern America. p. 82. 

Description: Length, 2.0 to 2.4 mm. 

Moderately robust. Black; pubescence scant, fine, white, coarser 
and more conspicuous laterally. Male beak one-fourth longer than 
prothorax, slightly curved, moderately expanded laterally at antennal 
insertion, attenuate to apical third which is nearly cylindrical; basal 
two-thirds dull, punctured, with scant pubescence, apical third 
smoother, moderately shining. Female beak about as long as head 
and prothorax, slightly curved, attenuate beyond basal third to 
middle, nearly cylindrical to apex; glabrous, dull, moderately punc- 
tured to apex. Antennae inserted at distance from eye two-fifths 
greater than width of frons at basal two-fifths of male, of female at 
distance from eye one-balf greater than width of frons slightly distad 
of basal third; first segment of male equal to next three, of female 
equal to next four; second segment shorter than next two; club 
0.16 by 0.08 to 0.18 by 0.10 mm. Eyes prominent; frons moderately 
wide, with median smooth area flanked by two lateral rows of punc- 
tures. Prothorax at base about one-fifth wider than long, middle 
distinctly wider than base, apex two-thirds to three-fourths as wide 
as base; sides moderately expanded laterally at base, roundly expanded 
to widest point at middle, rounding to constricted apex; some females 
with sides more broadly rounded and basal constriction greatly 
reduced; in profile dorsal surface slightly arcuate, depressed basally 
and apically; punctures moderately deep, 0.03 to 0.04 mm. in diameter, 
interspaces nearly flat, less than diameter of punctures, alutaceous; 
basal fovea deep, linear, extending about one-third length of prothorax. 
Elytra at humeri one-fourth to one-third (especially of female) wider 
than prothorax at base, 2.75 times as long as prothorax, length to 
width as 7:5; intervals twice as wide as striae, moderately convex, 
with one or two rows of punctures bearing fine scales; striae deep. 
Scutellum triangular, slightly longer than wide, with median furrow. 
Front femora 3.75 times as long as wide. Claws with acute basal 
tooth. 


SUBGENUS TRICHAPION—KISSINGER 377 


Special male characters: Tibiae 2 and 3 armed with long, sub- 
dentate mucrones, front tarsus dilated and densely pubescent on 
ventral surface. 

MATERIAL EXAMINED: 150 specimens. 

KNOWN DISTRIBUTION: 


Unitep Srates: Arkansas: Crawford Co., May 14, 1939, M. W. Sanderson 
(INHS); Franklin Co., May 19, 1940, L. Warren, on black locust (INHS.) 
District of Columbia: beating pine (USNM). Georgia: Rabun Co., Leng (USNM). 
Illinois: Charleston, June 7, 1941, Ross and Mohr (INHS); Coulterville, June 7, 
1940, L. M. Smith, on black locust (INHS); Toledo, July 7, 1950, Ross and 
Sanderson (INHS); Topeka, Aug. 17, 1907 (USNM). Indiana: Clay Co., 
July 8, 1935, on locust (USNM); Blatchley and Leng (1916) state species is com- 
mon throughout the State. Jowa: Story Co. Kansas. Louisiana: Tallulah, 
May 9, 1910, G. D. Smith, on Robinia pseudacacia (USNM). Maryland: Belts- 
ville, Aug. 24, 1922, I. L. Buchanan (USNM); Chespeake Beach, Wrenn and 
Barber (USNM); Great Fails, July 2, 1919, L. L. Buchanan (USNM); Hagers- 
town, July 3, 1917, W. E. Pennington (USNM); Lakeland, September 1895, 
Chittenden (USNM). Massachusetts: West Medford, July 21, 1919, R. H. Van 
Zwaluwenburg (USNM). Mississippi: Natchez, June 6, 1895, H. E. Weed 
(USNM). New Jersey: Many records. New York: Bellport, Long Island, 
June 11, 1912, A. Nicolay (USNM); Ithaca, Chittenden (USNM); Peekskill 
(USNM); Portage, May 30, 1885, E. O. VanDuzee (CIS); Staten Island (USNM); 
West Point, May 12, 1913, W. Robinson (USNM). North Carolina: Tryon, 
W. D. Pierce (USNM). Ohio: Franklin Co., May 16, 1949, E. L. Sleeper (ELS); 
Ross Co., Hubbard and Schwarz (USNM). Pennsylvania: Glenolden, June 16 
(USNM); Philadelphia, June 18, 1897 (USNM). South Carolina: Rocky 
Bottom, May 22, 1934, J. A. Berly (USNM); Charleston, Aug. 5, 1938, W. J. Reid 
(USNM). Tennessee: Clarksville, Apr. 23, 1919, E. R. Jones (USNM); Gatlin- 
burg, June 14, 1947, R. H. Wittaker (USNM). Vermont: Bergen Co. (USNM). 
Virginia: Many records. West Virginia: “Berkley,” Hubbard and Schwarz 
(USNM); Harper’s Ferry, May 19, Hubbard and Schwarz (USNM); White 
Sulphur Springs, Aug. 1, 1914, W. Robinson (USNM). Wisconsin: Waupaca, 
June 14, 1920, L. G. Gentner, feeding on locust (USNM). 


Remarks: Blatchley and Leng (1916) state that this species 
develops in the seeds of black locust, Robinia pseudacacia L., and 
that the adults riddle the leaves of the tree. Recorded also as feeding 
on the foliage of peach and wild cherry. 


Apion rostrum Group 


The six species comprising this group are distinct in that the pro- 
thorax is as wide or slightly wider at the middle than at the base, the 
sides are not strongly rounded; the beak is attenuate to near the apex; 
and typically the male has tibiae 2 and 3 mucronate, but in three 
species the front tibiae are also mucronate. The ranges of four 
species overlap. A. confertum Smith and A. furtivum Fall occur 


378 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


together in Florida. The southern range of A. rostrum Say extends 
into Florida. A. rostrum and A. coloradense Fall overlap in range 
in Kansas and probably in surrounding States. A. commodum Fall 
is a northern species occurring in Montana and Manitoba. A. 
mexicanum Wagner occurs in the mountains of central Mexico. 

The males of three species, A. rostrum, A. coloradense, and A. mexi- 
canum, have tibiae 2 and 3 mucronate. The tarsi of A. rostrum are 
stouter than those of the other species, the second tarsal segment is 
as long as wide and not longer than the lobes of the third segment, 
and the species is large, ranging from 2.5 to 3.0 mm. in length. The 
other two species are generally less than 2.5 mm. long and the second 
tarsal segment is longer than wide and longer than the lobes of the 
third segment. A. coloradense and A. mexicanum are closely allied 
species. The prothorax of the former is slightly constricted apically 
and the beak is strongly punctured in rows. The prothorax of the 
latter is distinctly constricted apically and the beak is finely punc- 
tured beyond the middle. The males of three species, A. commodum, 
A. confertum, and A. furtivum, have all three tibiae mucronate, the 
front pair generally has a minute mucro. The dorsal margin of the 
antennal scrobe of the last species is nearly horizontal and the tibial 
mucrones are rather short. The dorsal margin of the antennal scrobe 
of the other two species is oblique and somewhat angulate. The tibial 
mucrones of the male of A. commodum are denticulate, elytral interval 
2 has two or more rows of fine scales, and the beak is not strongly 
dilated at the antennal insertion and is somewhat cylindrical apically. 
The tibial mucrones of the male of A. confertum are simple, elytral in- 
terval 2 has a single row of scales, and the beak is strongly dilated at 
the antennal insertion and is strongly attenuate toward the apex. 


Apion (Trichapion) coloradense Fall 


FicurE 19,d,e 
Apion coloradense Fall, Trans. Amer. Ent. Soc., vol. 25, p. 152, 1898. 


Description: Length, 2.00 to 2.60 mm.; width, 0.94 to 1.18 mm. 

Moderately robust. Black, antennae piceous; pubescence white, 
fine, very sparse, not denser laterally. Male beak longer than head 
and prothorax combined, one-half longer than prothorax; more 
strongly deflexed at basal third; in lateral view expanded ventrally 
at antennal insertion, apical two-fifths nearly parallel; in dorsal view 
expanded laterally at antennal insertion, basal two-fifths attenuate, 
apical two-fifths nearly parallel; with strong, elongate punctures ar- 
ranged in rows, tip smooth. Female beak one-half longer than head 
and prothorax, nearly twice as long as prothorax; moderately curved, 
more strongly so toward apical region; in lateral view nearly parallel- 
sided; in dorsal view stout and parallel in basal third, attenuate to 


SUBGENUS TRICHAPION—KISSINGER 379 


Figure 19.—a-c, Apion commodum Fall: a, entire dorsal view of male; d, lateral view of 
head and prothorax of male; c, mucro of tibia 3 of male. d, e, 4. coloradense Fall: d, 
lateral view of head and prothorax of female; ¢, entire dorsal view of female. f-/, A. 
mexicanum Wagner:f , lateral view of head and prothorax of male; g, entire dorsal view 
of male; A, mucro of tibia 3 of male. 7,7, 4. rostrum Say: 1, mucro of tibia 3 of male; 7, 
lateral view of head and prothorax of male. k-m, A. confertum Smith: k, mucro of tibia 1 
of male; 7, mucro of tibia 3 of male; m, lateral view of head and prothorax of male. Line 
equals 0.50 mm. 


380 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


middle, tip slightly expanded; moderately punctured in rows, tip 
smooth and shining. Antennae of male inserted at distance from eye 
twice as great as width of frons at basal third of beak, of female at 
distance from eye one-half greater than width of frons at basal fourth 
of beak; first segment of male shorter than next two, of female equal 
to next two; second segment shorter than next two. Eyes not promi- 
nent; frons wider than dorsal tip of beak, with slight, shallow median 
sulcus and one lateral row of confluent punctures. Prothorax at base 
one-fourth wider than long, middle and base equal in width, apex four- 
fifths as wide as base; sides minutely expanded laterally at base, 
nearly parallel to middle, rounded to apex which is very slightly con- 
stricted; in profile dorsal surface slightly arcuate; punctation deep, 
0.03 to 0.04 mm. in diameter, not round, interspaces about one-half 
diameter of punctures, nearly smooth; basal fovea deep, narrow, ex- 
tending one-fourth length of prothorax. Elytra at humeri one-half 
wider than prothorax at base, slightly more than three times as long 
as prothorax, length to width as 13 : 9; intervals nearly flat, a little 
less than twice as wide as striae, with one or sometimes two irregular 
rows of punctures, nearly smooth; striae deep, coarse. Claws with 
acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with short, nearly 
simple mucrones. 

Lectotype: I hereby designate the lectotype of this species as the 
male specimen (MCZ 25082) labeled “Colorado Springs, Col., June 
15-30, 1896, 6,000-7,000 feet, H. F. Wickham.” 

MATERIAL EXAMINED: Lectotype and 7 specimens. 

KNOWN DISTRIBUTION: 

Unitep States: Colorado: Boulder, 8. A. Rohmer (MCZ); Colorado Springs, 
June 15-30, 1896, 6,000—7,000 feet, H. F. Wickham (MCZ). Kansas: (MCZ); 


Douglas Co., 900 feet, F. H. Snow (UK). TIilinoits: Valmeyer, May 24, 1950, 
M. W. Sanderson (INHS). 


Apion (Trichapion) commodum Fall 


Figure 19,a-c 
Apion commodum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 154, 1898. 


Description: Length, 3.0 to 3.3 mm. 

Moderately robust. Black; pubescence conspicuous, white, mod- 
erately coarse, sparse, slightly denser on sides of mesothorax. Male 
beak about as long as head and prothorax, moderately, evenly curved, 
expanded laterally, but not abruptly, at antennal insertion, attenuate 
to apical third; coarsely punctate. Female beak a little longer than 
head and prothorax, slightly curved, slender, nearly cylindrical 
throughout, slightly expanded laterally at antennal insertion; punc- 
tures in basal two-thirds moderately coarse, apical third finely 


SUBGENUS TRICHAPION—KISSINGER 381 


punctate, apex shining. Antennae of male inserted at distance from 
eye one-half greater than width of frons at basal two-sevenths, of 
female at distance from eye one-third greater than width of frons at 
basal one-fourth; first segment of male shorter than next two, of 
female equal to next three; second segment of male shorter than next 
two, of female longer than next two; female club 0.24 by 0.12 mm., 
male club 0.30 by 0.12 mm.; dorsal margin of antennal scrobe angulate. 
Prothorax at base slightly wider than long, middle one-sixth wider 
than base, apex two-thirds as wide as base; sides broadly rounded; 
in lateral view dorsal surface moderately strongly convex, flattened 
basally and apically; punctures moderate, 0.04 mm. in diameter, 
interspaces convex, alutaceous, one-half as great as diameter of 
punctures; basal fovea deep, linear, extending one-third length of 
prothorax. Elytra at humeri two-fifths wider than prothorax at 
base, three times as long as prothorax, length to width as 16.5 : 11; 
intervals twice as wide as striae, nearly flat to slightly convex, with 
two or three rows of fine punctures bearing fine scales; striae deep. 
Scutellum rounded, 0.06 by 0.06 mm., with slight median furrow. 
Front femora 3.0 to 3.5 times as long as wide. Claws with moderate 
basal tooth. 

Special male characters: Three pairs of tibiae mucronate, tibia 1 
with minute, simple mucro; tibiae 2 and 3 with larger, subdentate 
mucrones. 

MATERIAL EXAMINED: Type, male (USNM), Montana, and 30 
specimens. 

KNOWN DISTRIBUTION: 

UnitTep Srates: Montana: Custer Co. (USNM). 

Canapa: Manitoba: Aweme, September 20, N. Criddle, June 9 and July 10, 
R. M. White (CNC). 

Remarks: Material in the U. 8. National Museum was taken from 
an ovary of Psoralea esculenta and material in the Canadian National 
Collection is associated with the same plant. 


Apion (Trichapion) confertum Smith 


Ficurs 19,k—m 


Apion confertum Smith, Trans. Amer. Ent. Soc., vol. 11, p. 63, 1884.— Fall, 
Trans. Amer. Ent. Soc., vol. 25, p. 154, 1898.—Blatchley and Leng, Rhyncho- 
phora or weevils of northeastern America, p. 82, 1916. 

Description: Length, 2.3 to 2.5 mm. 

Moderately robust. Black; pubescence white, sparse, moderately 
fine, slightly coarser and more conspicuous ventrally. Male beak 
about as long as head and prothorax combined, two-sevenths longer 
than prothorax; slightly, evenly curved; attenuate from laterally 
expanded antennal insertion to apical fourth; basal two-thirds dull, 


382 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


alutaceous, shallowly rather coarsely punctate, apical third shining, 
sparsely punctured, sparsely pubescent behind antennal insertion. 
Female beak similar to male but slightly longer, three-sevenths longer 
than prothorax, and a little more slender. Antennae inserted at 
distance from eye one-third greater than width of frons at basal fifth 
of beak; first segment equals next two, second segment shorter than 
next two, club 0.18 by 0.08 mm. Eyes moderately prominent; 
frons moderately wide, wider than dorsal tip of beak. Prothorax 
at base slightly wider than long, middle about as wide as base, apex 
three-fourths as wide as base; sides beyond basal lateral expansion 
slightly expanded to middle, rounded to constricted apex; in profile 
dorsal surface slightly, evenly arcuate; punctation deep, 0.04 mm. in 
diameter, interspaces not as wide as punctures, alutaceous; basal 
fovea deep at middle, narrow, extending one-half length of prothorax. 
Elytra at humeri one-half wider than prothorax at base, about 2.85 
times as long as prothorax, length to width as 10.5 :7; intervals 
nearly flat, slightly more than twice as wide as striae, with one row of 
moderately fine scales; striae moderately deep, fine. Scutellum tri- 
angular, 0.08 by 0.06 mm., not furcate. Front femora about 3.3 times 
as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 1 and 2 with small, simple mucrones, 
tibia 3 with slightly larger, subangulate mucro. 

MATERIAL EXAMINED: Type, female (USNM 1247), Georgia, and 
four specimens. 

KNOWN DISTRIBUTION: 


Unitep Srates: Florida: No exact locality (USNM); Jacksonville. Georgia: 
No exact locality (USNM). 


Apion (Trichapion) furtivum Fall 
Apion furtivum Fall, Trans. Amer. Ent. Soc., vol. 25, p. 154, 1898. 


Description: Length, 2.00 mm.; width, 0.98 mm. 

Moderately robust. Black, antennae piceous; pubescence white, 
fine, very sparse, not more conspicuous laterally. Male beak shorter 
than head and prothorax combined, one-fourth longer than prothorax, 
slightly, evenly curved; in lateral view dorsal margin abruptly 
narrowed beyond antennal insertion, attenuating to apical fourth, 
thence nearly parallel-sided to tip; in dorsal view expanded laterally 
at antennal insertion, attenuate to apical third, thence nearly 
parallel-sided to tip; base moderately pubescent, minutely so beyond 
antennal insertion, with a row of punctures above antennal insertion, 
tip smooth, shining. Female beak one-fifth longer than head and 
prothorax combined, two-thirds longer than prothorax; moderately, 
strongly curved; nearly cylindrical throughout; finely, sparsely 
punctured throughout, with a slightly stronger row of punctures 


SUBGENUS TRICEIAPION-—KISSINGER 383 


above antennal insertion. Antennae inserted at distance from 
eye one-half greater than width of frons at basal third of beak of 
male, of female at distance from eye slightly greater than width 
of frons and slightly distad of basal fourth; first segment shorter 
than next three, second segment equals next two, club 0.18 by 
0.08 mm. Eyes slightly prominent; frons wider than dorsal tip of 
beak, with moderately strong, deep, median sulcus and one lateral 
row of punctures. Prothorax slightly wider at base than long, middle 
slightly wider than base, apex three-fourths as wide as base; sides, 
beyond slight basal lateral expansion, roundly expanding to middle, 
rounding to constricted apex; in profile dorsal surface slightly arcu- 
ate; punctation deep, 0.03 to 0.04 mm. in diameter, interspaces 
less than diameter of punctures, alutaceous; basal fovea deep basally, 
narrow, extending about one-half length of prothorax. Elytra at 
humeri one-third wider than prothorax at base, 2.6 times as long as 
prothorax, length to width as 10.5:7.5; intervals flat, at middle 
twice as wide as striae, with one row of shallow, rather coarse punc- 
tures bearing scales; striae deep. Claws with acute basal tooth. 

Special male characters: Three pairs of tibiae mucronate, mucro 
on tibia 1 somewhat smaller. 

Typrs: I hereby designate the lectotype of this species as the 
male specimen (MCZ 25098) in the Fall Collection labeled with a 
blue square (representing Southeastern States) and ‘2442,’ collected 
by Zimmerman. Cotypes (MCZ 381) in the J. L. LeConte Collec- 
tion have the same data. 

KNOWN DISTRIBUTION: 


Unirep Srares: Georgia: No exact locality. Arkansas: ‘‘Holes,” May 27, 
1950, R. H. Beamer (UK). 


Apion (Trichapion) mexicanum Wagner 


Figure 19,f-h 


Apion (Trichapion) mexicanum Wagner, Arch. Naturg. Berlin, vol. 78, p. 114, 
1912. 

Description: Length, 1.50 to 2.00 mm.; width, 0.87 to 0.94 mm3 

Moderately robust. Black; pubescence conspicuous, fine, white, 
sparse, denser on sides of mesothorax. Male beak as long as head 
and prothorax, two-thirds longer than prothorax, slightly curved, 
moderately swollen laterally and ventrally at antennal insertion, 
attenuate to middle, apical half nearly cylindrical; finely, sparsely 
punctured in basal two-thirds, pubescence scant, apical third smooth, 
shining. Female beak is longer than head and prothorax combined, 
four-fifths longer than prothorax, similar to beak of male, apical third 
nearly cylindrical; dull, alutaceous, minutely pubescent behind 
antennal insertion, glabrous distad of antennal insertion, tip smoother, 


384 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


more shining. Antennae inserted at distance from eye one-half 
greater than width of frons, at basal third of beak; first segment 
shorter than next two, second segment shorter than next two, club 
0.15 by 0.07 mm. Eyes slightly prominent; frons moderately wide, 
with a slight median depression and a lateral row of four moderate 
punctures. Prothorax at base one-third wider than long, middle 
slightly narrower than base, apex three-fourths as wide as base; 
sides moderately expanded laterally at base, nearly parallel to mid- 
dle, apex broadly constricted; in profile dorsal surface slightly evenly 
arcuate; punctation 0.03 mm. in diameter, deep, interspaces variable, 
generally less than diameter of punctures; basal fovea deep in middle, 
short. Elytra at humeri one-half wider than prothorax at base, 
3.3 times as long as prothorax, length to width as 5:3.3; intervals 
nearly twice as wide as striae, nearly flat, with a single row of fine 
punctures bearing fine scales; striae narrow, deep. Scutellum 
triangular, 0.06 by 0.04 mm., with slight median furrow. Front 
femora 3.5 times as long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with angulate 
mucrones. 

MATERIAL EXAMINED: One male determined by Hans Wagner and 
27 other specimens. 

KNOWN DISTRIBUTION: 

Mexico: “Mexico” (type, BMNH); Distrito Federal: Atcapotzalco; Tacuba; 
Tlalpam (USNM). 

Remarks: Material in the U. S. National Museum is recorded as 
occurring ‘fon bean.” 


Apion (Trichapion) rostrum Say 
Figure 19,i,j 


Apion rostrum Say, Journ. Acad. Nat. Sci. Philadelphia, vol. 5, p. 253, 1826.—Say, 
Descriptions of North American curculionides and an arrangement of some 
of our known species agreeably to the method of Schoenherr, p. 6—Smith, Trans. 
Amer. Ent. Soc., vol. 11, p. 63; fig. 16, 1884.—Fall, Trans. Amer. Ent. Soc., 
vol. 25, p. 151, 1898.—Pierce, U. 8. Dep. Agr. Bur. Ent. Bull. 100, p. 75, 
1912.—Blatchley and Leng, Rhynchophora or weevils of northeasterr 
America p. 81, 1916.—Bleasdell, Iowa State Coll. Journ. Sci., vol. 11, p. 110, 
1937.—Bissell, Journ. Econ. Ent., vol. 33, p. 846, 1940.—Frost, Journ. New 
York Ent. Soc., vol. 53, p. 221, 1945.—Tuttle, Ann. Ent. Soc. Amer., vol. 47, 
p. 305, 1954. 

Apion scrobicolle Gyllenhal, in Schoenherr Genera et species curculionidum, vol. 
5, p. 374, 1839—Wencker, L’Abeille, vol. 1, p. 121, 1864.—Wagner, 
Miinchener Kol. Zeitschr., vol. 3, p. 304, 1906-08; Genera insectorum, 
Coleoptera, fam. Curculionidae: subfam. Apioninae, fasc. 130, p. 84. 


Description: Length, 2.5 to 3.0 mm. 


Robust. Black; pubescence scant, fine, white, more conspicuous on 
sides of mesothorax and metathorax. Male beak as long as head and 


SUBGENUS TRICHAPION—KISSINGER 385 


prothorax combined, slightly, evenly curved; moderately dilated 
laterally over antennal insertion, attenuate to apical third, there 
nearly cylindrical, tip depressed; sculpture shallow, moderately 
punctured throughout, with a line of punctures over antennal inser- 
tion, tip more shining, pubescence scant. Female beak one-seventh 
longer than head and prothorax, more strongly deflexed distad of 
apical third; moderately dilated over antennal insertion, attenuating 
to middle, apical half nearly cylindrical, tip depressed; basal two- 
thirds with shallow, moderate punctures, with a line of punctures 
over antennal insertion, apical third smoother, alutaceous. Antennae 
inserted at distance from eye one-third greater than width of frons, of 
male at basal third, of female at basal fourth; first segment as long as 
next three; second segment slightly shorter than next two, club 0.27 
by 0.12 mm. Eyes prominent; frons moderately wide, with deep, 
median sulcus and a lateral line of coalesced punctures. Prothorax 
at base one-sixth wider than long, middle equal or shghtly wider than 
base, apex seven-tenths as wide as base; sides with slight lateral 
expansion at base, roundly diverging to middle, rounded to slightly 
constricted apex; in profile dorsal surface moderately convex; puncta- 
tion coarse, 0.04 to 0.06 mm. in diameter, deep, interspaces generally 
half as great as diameter of punctures, strongly alutaceous, convex; 
basal fovea deep, elongate, extending at least one-third length of 
prothorax. Elytra at humeri two-fifths to one-half wider than 
prothorax at base, 2.6 times as long as prothorax, length to width as 
14:11; intervals twice as wide as striae, flat, with one row of fine 
punctures bearing fine scales; striae deep. Scutellum roundly 
triangular, 0.10 by 0.10 mm., smooth. Front femora 3.75 times as 
long as wide. Claws with acute basal tooth. 

Special male characters: Tibiae 2 and 3 armed with long, blunt, 
dentellate mucrones. 

MATERIAL EXAMINED: 100 specimens and type of A. scrobicolle from 
“Anglia” in the Riksmuseum, through the kindness of Dr. René 
Malaise. 

KNOWN DISTRIBUTION: 

Unitep Srates: Connecticut: Lyme, June 18, 1918, W. 8. Fisher (USNM); 
District of Columbia: Sept. 15, 1907 (USNM). Florida: Farmingdale, Aug. 19, 
1945, C. O. Esselbaugh (INHS); Sansond, June 20, 1940, G. W. Barber (USNM); 
Shady, July 11, 1904, C. Buhl (USNM). Georgia: Clayton, Davis (USNM); 
Lexsy, Apr. 26, 1940, P. W. Fattig (USNM); Rabun Co., July, Leng (USNM); 
Valdosta, May 28, 1946, P. W. Fattig (USNM). Indiana: Pulaski, Starke, and 
Wells Counties. Jowa: Story and Taylor Counties. Kansas: Topeka, Aug. 7, 
E. A. Popenoe (USNM). Louisiana: Natchitoches, Mar. 28, 1907, Cushman 
and Pierce (USNM). Maryland: Beltsville, June 30, 1923, L. L. Buchanan 
(USNM); Contee, May 7, 1914, A. Wetmore (USNM); Oakland, July 16, Hubbard 


and Schwarz (USNM); Odenton, June 11, 1922, L. L. Buchanan (USNM); 
Takoma Park, June 27, 1952, G. H. Nelson, to light (GHN); 2 miles north of 


386 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Priest Bridge, June 8, 1928, H. S. Barber (USNM). Massachusetts: Agawan, 
May 25, 1916, E. A. Chapin (USNM); Boston, June 4 (USNM); Nantucket 
Island (USNM); Natick, Aug. 29, 1937, C. A. Frost (USNM); Waltham (USNM); 
Woods Hole, Wickham (USNM). Michigan: Detroit (USNM). New Jersey: 
Many records. New York: Grasmere, Sept. 8, 1944 (USNM); Ithaca (USNM); 
New York City and vicinity (USNM). Pennsylvania: Berks Co., Aug. 18, 1938, 
ex seed Bapiisia tinctorum, 8. C. Schell (USNM); Foxes Run near Newtown 
Square, Sept. 30, 1947, J. W. Adams (USNM); Glenolden, June 16 (USNM); 
Hanover, June 15, 1980 (USNM); Lancaster (USNM); Lehigh Gap, June 29, 
1897 (USNM); Mauch Chunk, July 5 (USNM). Rhode Island: Watch Hill, 
July 11, 1909, W. Robinson (USNM). Texas: Anahuac, June 20, 1918, H. C. 
Harrison (USNM). Vermont: Bennington Co. (USNM). Virginia: Four-Mile 
Run, June 22, 1919, L. L. Buchanan (USNM); Falls Church, July 3, 1920 (USNM); 
Fairfax Co., June 20, 1923 (USNM); Vienna, July 14, 1921 (USNM). West 
Virginia: Elkins, F. E. Brooks (USNM); White Sulphur Springs, July 3, 1912, 
W. Robinson (USNM). Wisconsin: Madison (USNM). 

Remarks: Larvae of this species develop in the seeds of the false 
indigo, Baptisia tinctoria and B. leucantha (Blatchley and Leng, 1916; 
Bissell, 1940; Frost, 1945; Tuttle, 1954). 


References 
ANONYMOUS 
1942. Fitofilo (San Jacinto, Distrito Federal, Mexico), afio 1, No. 2, p. 54. 
(Not seen.) 
1943. Fitofilo (San Jacinto, Distrito Federal, Mexico), afio 2, No. 3, p. 11. 
(Not seen.) 
BARTOSZYNSKI, ANTONI 
1935. Beitrag zur Carabiden und Curculioniden fauna von Mascovien. 
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BepeEu, Louis 
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B&auin-BiLLeEcoca, Louis 
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BissELL, THEODORE LEMUEL 
1938. The host plants and parasites of the cowpea curculio and other legume 
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1940. Curculionidae, Bruchidae, Lepidoptera, and their parasites, infesting 
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BLACKWELDER, RIcHARD ELIOT 
1944— Checklist of the coleopterous insects of Mexico, Central America, 
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185, pts. 1-6, 1492 pp. 
BLaNcuarD, Cuarues Emiie 
1851. Fauna Chilena. Insectos. Coleépteros. In Gay, Historia fisica y 
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BLATCHLEY, WILLIS STANLEY, AND Lenc, CHARLES WILLIAM 
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BLEASDELL, G. G. 
1937. The Rhynchophora of Iowa. Iowa State College Journ. Sci., vol. 11, 
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SUBGENUS TRICHAPION—KISSINGER 387 


BoHEMAN, Cari HEINRICH 
1839. In Schoenherr, Genera et species curculionidum . . ., vol. 5, pt. 1, 
456 pp. 
Bruan, A. F. 
1947. The external male genitalia of some Rhynchophora. Great Basin 
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CHITTENDEN, FraNK HURLBURT 
1908. An injurious North American species of Apion with notes on related 
forms. U.S. Dep. Agr., Bur. Ent. Bull., No. 64, pt. 4, pp. 29-32. 
Cotton, RicHarD THOMAS 
1906. The insects affecting black locust and hardy catalpa. Ohio Dep. 
Agr. Div. Nurs. Orch. Insp. Bull. No. 7, 55 pp. (Not seen.) 
DESBROCHERS DES Loass, JULES 
1895-1896. Révision des curculionides appartenant a la tribu des apionides 
d’Europe et des pays voisin en Afrique et en Asie, 3rd Partie. 
Le Frelon, vol. 5, pp. 1-12, 69-96, 105-164. [Paper is paged con- 
tinuing from part 2 of Desbrochers’ work and according to the 
volume, so that p. 244 of Desbrochers’ work is p. 90 of the volume.] 
DonistHoRPE, Horace St. Joun K. 
1938. <A preliminary list of the Coleoptera of Windsor Forest. Ent. Monthly 
Mag., vol. 74, No. 891, pp. 166-177. 
Estapos Unipos MEXICANOs, SECRETARIA DE AGRICULTURA Y FoMENTO 
1930. El picudo del frijo. Principales plagas y enfermedades de los cultivos 
de la Reptiblica Mexicana, pp. 84-85. (Not seen.) 
Fatit, Henry CLINTON 
1898. Revision of the species of Apion of America north of Mexico. Trans. 
Amer. Ent. Soc., vol. 25, pp. 105-184, pls. 2-5. 
1918. New North American species of Apion. Journ. New York Ent. Soc., 
vol. 26, pp. 218-223. 
Faust, JOHANNES 
1887. Curculioniden aus dem Amur-Gebiet. Deutsche Ent. Zeitschr., vol. 
31, pp. 161-189. 
1889. Notizen tiber Russel Kafer. Stettiner Ent. Zeitung, vol. 50, pp. 
227-234. 
1893. Reise von E. Simon in Venezuela. Stettiner Ent. Zeitung, vol. 54, 
pp. 3138-368. 
Frost, Stuart Warp 
1945. Insects feeding or breeding on indigo, Baptisia. Journ. New York 
Ent. Soc., vol. 53, pp. 219-225. 
GEMMINGER, Max 
1871. Berichtigungen und Zusitze zum Catalogus Coleopterorum synonym- 
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GERMAR, ERNST FRIEDRICH 
1815-1817. W. Kirby’s Monographie der Gattung Apion Herbst aus den 
Englischen iibersetzt, mit Bemerkungen, und eingeschalteten Be- 
schreibungen neuer Arten. Mag. Ent., Halle, vol. 2, pp. 114-265, 
pls. 2-4. 
GERSTAECKER, Cart Epuarp ADOLPH 
1854. Beschreibung neuer Arten der Gattung Apion Herbst. Stettiner Ent. 
Zeitung, vol. 15, pp. 234-261, 265-280. 
GotpMANn, Epwarp ALPHONSO 
1951. Biological investigations in Mexico. Smithsonian Misc. Coll., vol. 
115, 476 pp., 71 pls. 


388 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


GYLLENHAL, LEONARD 
1813. Insecta Suecia. ClassisI. Coleoptera sine Eleuterata. Vol. 1, pt. 3, 


730 pp. 
1833. In Schoenherr, Genera et species curculionidum. Vol. 1, pt. 1, 381 
pp. 
1839. In Schoenherr, Genera et species curculionidum ... Vol. 5, 
pt.1, 456 pp. 
Heresst, JOHANN FReIpRicH WILHELM 
1797. Natursystem ... Vol. 7, 346 pp., illus. 


Hustacue, A. 
1931. Famille txxrx Curculionidae, Tribu: Apioninae. Tableaux ana- 
lytiques des Coléoptéres de la Faune Franco-Rhenane. 286 pp., 
77 fig. 
Kirsy, WILLIAM 
1811. Descriptions of seven new species of Apion. Trans. Linn. Soe. 
London, vol. 10, pp. 347-357. 
Kirscu, THropor Franz WILHELM 
1874. Beitrige zur Kenntnis der Peruanischen Kaferfauna auf Dr. Abend- 
roth’s Sammlungen basirt. Berliner Ent. Zeitschr., vol. 18, pp. 
385-432. (Drittes Stiick.) 
KissIncer, Davip GEORGE 
1957a. Taxonomic notes on North American Apion. Proc. Ent. Soe. Wash- 
ington, vol. 59, p. 40. 
1957b. Studies on North American Apion with descriptions of two new species 
(Coleoptera : Curculionidae). Coleopt. Bull., vol. 11, pp. 17-24. 
KUSCHEL, GUILLERMO 
1950. Nuevas sinonimias, revalidaciones, y combinaciones. Agr. Tecn. San- 
tiago de Chile, vol. 10, pp. 10-21. 
Lene, CHARLES WILLIAM 
1920. Catalogue of the Coleoptera of America, north of Mexico. 470 pp. 
MaRrcovirtcn, 8. 
1916. Insects attacking weeds in Minnesota. Minnesota Ent. Rep. No. 
16, pp. 185-152. 
McKELvEY, JOHN Jay, JR. 
1947. Apion pod weevil; a pest of beans in Mexico. Journ. Econ. Ent., 
vol. 40, pp. 135-152. 
McKELvEY, JOHN JAY, JR., BT AL. 
1951. Biologia y control de los picudos del género Apion que atacan al 
frijol en México. Folleto Técnico, Secr. Agr. Gan. México, vol. 
8, pp. 8-42. (Not seen.) 
Prerce, WititiaAM Dwicut 
1908. Studies of parasites of the cotton boll weevil. U.S. Dep. Agr., Bur. 
Ent. Bull., No. 73, 65 pp., 6 figs., 3 pls. 
1912. The insect enemies of the cotton boll weevil. U.S. Dep. Agr., Bur. 
Ent. Bull., No. 100, 99 pp., 26 figs. 
Puinipp!1, RupoLF AMANDUs, AND PHILIPPI, FRIEDRICH HEINRICH E. 
1864. Beschriebung einiger neuen Chilenischen Kafer. Stettiner Ent. 
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Say, THomas 
1825- Descriptions of new species of coleopterous insects inhabiting the 
1826. United States. Journ. Acad. Nat. Sci. Philadelphia, vol. 5, pt. 1 
(1825), pp. 160-202; vol. 5, pt. 2 (1826), pp. 237-284, 293-304. 


SUBGENUS TRICHAPION—KISSINGER 389 


1831. Descriptions of North American curculionides and an arrangement 
of some of our known species agreeably to the method of Schoenherr. 
30 pp. 
ScHILSKY, JULIUS 
1902. Jn Kiister and Kraatz, Die Kafer Europa’s, Heft 39, 100 pp. 
SCHWITZGEBEL, R. B., anp W1LBuUR, DonaLp ALDEN 
1942. Coleoptera associated with ironweed, Veronia interior Small, in 
Kansas. Journ. Kansas Ent. Soc., vol. 15, pp. 37-44, pl. 1. 
Smarp, Davip 
1890— Jn Godman and Salvin, Biologia Centrali-Americana, Insecta, Cole- 
1891. optera, Curculionidae: Apioninae, vol. 4, pt. 3, pp. 48-86, pls. 2, 3. 
Smitn, Artur C. 
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Bull., vol. 5, pp. 61-62, 1 fig. 
1951b. A colonial cocoon in the genus Apion. Coleopt. Bull., vol. 5, pp. 
74-75, figs. 1, 2. 
Smitu, JOHN BERNHARDT 
1884. Synopsis of the Apioninae of the United States. Trans. Amer. Ent. 
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TuttLe, Donatp B. 
1954. Notes on the bioncmics of six species of Apion. Ann. Ent. Soc. 
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Voss, EpUARD 
1955. Coleoptera Curculionidae Apioninae. (130. Beitrag zur Kenntnis 
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mologique du Ruanda-Urundi (Mission P. Basilewsky. 1953), 
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WaaGner, Hans 
1908. Beitrige zur Kenntnis der Gatt. Apion Hbst. IV. Miinchener Kol. 
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1912. Beitrag zur Kenntnis der Apion-Fauna Central- und Sud-Amerikas. 
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A, Heft 2, pp. 99-136. 
WAHLGREN, E. 
1951. Cecidiologiska antekningar VI. Ent. Tidskr., vol. 72, pp. 84-156. 
WatsH, Bensamin Dann 
1867. On the insects inhabiting galls of willow. Proc. Ent. Soc. Phila- 
delphia, vel. 6, 1866, p. 269. 
WENCKER, JOSEPH ANTOINE 
1863. Description de plusieurs espéces nouvelles exotiques de genre Apion 
Herbst. Ann. Soc. Ent. France, ser. 4, vol. 3, pp. 177-182. 
1864. Apionides, tribu des Curculionides. L’Abeille. vol. 1, pp. 109-270. 
WickHamM, Henry FREDERICH 
1922. Weevils of the genus Apion injurious to beans in Mexico. Proc. 
Ent. Soc. Washington, vol. 24, pp. 118-122. 
Wo.tcort, A. B., an» Montcomery, B. E. 
1933. Ecological study of the coleopterous fauna of a tamarack swamp. 
Amer. Midl. Nat., vol. 4, pp. 113-169. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U.S. NATIONAL MUSEUM 


Vol. 110 Washington : 1959 No. 3419 


A DESCRIPTION AND CLASSIFICATION OF THE FINAL 
INSTAR LARVAE OF THE ICHNEUMONIDAE (INSECTA, 
HYMENOPTERA) 


By J. R. T. SHort! 


Introduction 


In this paper are given a description and classification of the final 
instar larvae of the Ichneumonidae based on a study of species repre- 
senting 151 genera. Keys are given to the subfamilies, tribes, and 
genera. Characters of the subfamilies and tribes are described verbally, 
but the description of each genus usually takes the form of a diagram. 
Any special features are noted in the text, and when a species differs 
from that figured for a genus the difference is noted. Diagrams were 
selected as the most concise and accurate method for recording 
larval characters. 


MatTERIAL AND ACKNOWLEDGMENTS 


In work of this nature great care must be taken to insure that only 
reliably identified material is used. Most of the material on which 
this study is based comes from the U.S. National Museum, Wash- 
ington, D.C. I wish to thank the Museum for permission to borrow 
material and I am most grateful to Miss Luella M. Walkley for her 
help and advice concerning this work. I am also grateful to Mr. G. 


1 Natural History Department, Marischal College, University of Aberdeen, Aberdeen, Scotland. 


391 


392 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Stuart Walley of the Canadian Department of Agriculture, Ottawa, 
for supplying material and to Mr. J. F. Perkins of the British Museum 
(Natural History) for permission to study material and for his advice 
on the classification of the Ichneumonidae. I also wish to thank 
Prof. G. C. Varley of the Hope Department of Entomology, Oxford, 
for his many helpful suggestions and for lending me a number of slides 
from his collection; Mr. E. R. Skinner of the Imperial Forestry Insti- 
tute, Oxford, who supplied me with larvae of Rhyssa; and Dr. G. Salt, 
F.R.S., of the Zoological Department, Cambridge University, who 
supplied me with larvae of Collyria and Idechthis. 

Since living larvae often give valuable information, I have collected 
as widely as possible for this study. Most of the slide preparations 
have been deposited at the U.S. National Museum. Material borrowed 
from the Canadian Department of Agriculture and the British Museum 
has been returned. Most of the material sent by Professor Varley 
has been returned, but material of Orthopelma and Hemiteles that he 
did not wish returned has been deposited at the U.S. National Mu- 
seum. Material of Collyria and Idechthis sent by Dr. Salt and material 
of Rhyssa sent by Mr. Skinner have also been deposited at the U.S. 
National Museum. Material returned to the Canadian Department 
of Agriculture (CDA), to the British Museum (BM), and to Professor 
Varley (GCV) is indicated by initials. Material not so indicated has 
been deposited in the U.S. National Museum. 

Detailed records (by whom determined, locality, etc.) have not 
been recorded for any specimens since this information can be ob- 
tained from the data labels of the collections of bred material (now 
returned to the U.S. National Museum and the Canadian Depart- 
ment of Agriculture) from which larval skins were obtained. The 
procedure adopted in using this bred material was as follows: The 
collection of preparations of larval skins was numbered and fully 
labeled; when a larval skin was extracted from the cocoon of a bred 
specimen a label was added to the pin of the specimen reading “Larval 
skin removed; slide No. X; J.R.T.S.; 195X.’’ Thus, it is possible to 
refer from the preparation of a larval skin to the adult and its de- 
termination. 

Meruops 


Much of the material used was available as cast skins in the cocoons 
of bred ichneumonids. The contents of the cocoon, the final instar 
larval skin and the meconium, were removed by making a longitudinal 
cut in the cocoon and then boiled in a 5 percent solution of caustic 
potash. Sometimes larval skins of instars previous to the final were 
found in the cocoons. After boiling, the cleaned and softened larval 


skins were washed in distilled water, stretched, dehydrated, and 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 393 


mounted in Canada balsam. Balsam was used to ensure a satis- 
factory permanent mount, but for more rapid work larvae may be 
transferred from distilled water to Faure’s fluid. Where larval mate- 
rial was available in the host pupa, as in the Ichneumoninae and 
Pimplini, the entire pupa was softened by boiling in caustic potash 
and the pupa was cut open when soft to extract the ichneumonid 
larval skin. This method ensured that the brittle pupa was not 
not damaged. Fresh larvae were opened by a midventral cut and 
then boiled in a 5 percent solution of caustic potash. They were 
then washed, dehydrated, and mounted. 


NOMENCLATURE 


The taxonomy of the Ichneumonidae is in a confused state, and the 
nomenclature of European and American authors differs to some 
extent. Since much of the material used in this study is North 
American in origin, the nomenclature of Walkley (1958) is used. The 
names of larvae described in the literature have been altered, when 
necessary, to conform with this nomenclature. When genera were 
not listed by Miss Walkley, her advice was sought on nomenclature, 
or, if the genera were listed in Kloet and Hincks (1945), the nomen- 
clature of these authors was followed. 


Description and Classification 


Since the relationships indicated by the larval characters support 
a classification similar to that given by Townes and Townes (1951) 
in the U.S. Department of Agriculture’s synoptic catalog and by 
Walkley (1958) in the supplement to that catalog, the classification 
of these: authors has been followed. Although a somewhat different 
classification could have been constructed on the basis of larval 
structure alone, every effort was made to avoid this because informa- 
tion on both larvae and adults is essential for an understanding of 
the taxonomy of the group. A classification based solely on the 
characters of the larvae would be as unsatisfactory as a classification 
based solely on the characters of the adults. 

It was necessary to depart from the classification in the above- 
mentioned synoptic catalog in one case: The Anomalinae are here 
considered as a separate subfamily rather than as a tribe of the 
Ophioninae. Such differences are to be expected since the inde- 
pendence of larval and adult forms makes a separate evolution possible 
(Wigglesworth, 1954). Selective factors acting on the legless larvae 
are different from those acting on the mobile adults. 

Studies on larval systematics are of interest from two standpoints; 
they enable workers to identify larvae and they help in understanding 
the classification of the adults (see van Emden, 1957). It is hoped 


394 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


that the keys to subfamilies and tribes will require no great modifi- 
cation as a result of future work, which is expected to take the form 
of detailed studies within tribes and genera. As a result of such work 
the keys to genera given here will be greatly modified, for the genera 
studied were usually represented by but two or three species. This 
number of species might be satisfactory if one could assume that the 
larval characters of all the species of a genus would indicate a close 
relationship. This is very far from being the case. In some genera 
this may be due to species differing more as larvae than as adults. In 
others, as in the genus Scambus, it appears to be due to the confused 
state of the taxonomy of the adults. However, it is hoped that the 
present keys to genera will serve as a basis for future studies. 

It is generally believed that, since adult insects show a greater 
degree of differentiation than larvae, their characters are likely to 
remain more important for species identification (van Emden, 1955). 
No attempt has been made in the present study to give a key to 
species. A vast amount of material would be required for such 
work. But itis significant that species of the braconid genus Apanteles 
can be arranged into groups corresponding with the groups into 
which the adults are divided (Short, 1953). 


EXTERNAL STRUCTURE OF FINAL INSTAR LARVA 


The final instar ichneumonid larva consists of a hemispherical 
head, three thoracic segments, and ten abdominal segments. The 
cranium is lightly sclerotized, with the mandibles and certain bands 
associated with the mouthparts more heavily sclerotized. The body 
wall of the thorax and abdomen is very lightly sclerotized or un- 
sclerotized, with the exception of the spiracles and setae. The spi- 
racles are situated in most species on the prothoracic and first eight 
abdominal segments. A line of setae is present in most species encir- 
cling each thoracic and abdominaJ segment. Small projections are 
present on the skin and these may be lightly sclerotized. 

The structures used in¥this classification are the form of the 
mandibles and head sclerites, and of the antennae, spiracles, and 
skin. For the basis of the terminology of the head parts, which is 
outlined below, reference should be made to Short (1952). The 
head of ichneumonids (fig. 1) is lightly sclerotized except for certain 
more heavily sclerotized bands strengthening the cranium in the 
region of the mandibles, maxillae, and labium. The sclerotic arch, 
which is sometimes present dorsal to the mandibles (md), is called 
the epistoma (epst). The anterior tentorial pits (at) are situated in 
the epistoma. Lateral to each mandible is a pleurostoma (plst). 
The mandible articulates with two processes of the pleurostoma (fig. 
1c). The anterior pleurostomal process (app) fits into a cavity of 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 395 


the mandible and the posterior pleurostomal process (ppp) ends in a 
socket which receives the mandibular condyle. The hypostoma (fig. 
1B, hst) is a sclerotic band running posteriorly along the subgenal 
margin of the cranium. Each maxilla (mx) consists of a lightly 
sclerotized cardo (cd) and a membranous stipes (st) supported 
ventrally by a sclerotic band called the stipital sclerite (ss(q)). The 
medial lobe of the maxilla represents the lacinia (lc). The galea is 
absent. The maxillary palp (mplp) is reduced to a flattened disc. 
A sclerotic band called the hypostomal spur (hsp) projects ventrally 
from the hypostoma across the stipes. In many species it fits into 
a depression in the stipital scierite. Surrounding the posterior 
margin of the prelabium (plb) is a labial sclerite (Ibs). The labial 
palps (Iplp) are reduced to flattened discs. The silk press (sp) at 
the orifice (slo) of the silk glands is usually well sclerotized. Some 
ichneumonid larvae, such as the Campoplegini, have a Y-shaped 
sclerite, called the prelabial sclerite, on the prelabium. A sclerotic 
band, the labral sclerite (Ims), is often present marking the dorsal 
limits of the labrum (lm). 

The mandible (fig. 1c) consists fundamentally of a broad base (ba) 
articulating with the pleurostomal processes, and a more slender 
blade (bl). The base and blade of the mandible of Xorides are not 
differentiated, but usually, as for example in the Ephialtini, the 
blade is slender. The blade of the mandible may bear teeth (t) on 
its dorsal and ventral surfaces. 

The antennae (ant) may be papilliform or disc-shaped. The 
ocular lines (ol), which mark the place of the developing imaginal 
compound eyes, are often sclerotized. 

Each spiracle (fig. 1p) consists of a rounded atrium (atr) connected 
with the closing apparatus (ca) by a length of trachea which is often 
characteristic of the group to which the larva belongs. Finer details 
of spiracular structure have not been used in this classification. The 
prothoracic spiracle is usually figured but, where the preparation did 
not show the structure of this clearly, another spiracle was selected 
for illustration. The proportions of the structure of the spiracles are 
relatively constant. 

The length of the setae of the skin and the presence or absence of 
spines have been found useful taxonomic features. Spines serving as 
holdfast organs are present on the skin of the Polysphinctini and some 
Ephialtini. 

The drawings of heads which illustrate this paper are morphologi- 
cally inaccurate since they show it as a flattened object, whereas it is, 
in reality, a hemisphere. This point is important since Beirne (1941, 
pp. 149, 167), for example, has attempted to use the position and 
degree of curving of the hypostomal sclerite as a taxonomic character. 


396 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


This is unwise since the position and degree of curving of the head 
sclerites depends to some extent on the position assumed during 
mounting. However, it has been found a satisfactory method for 
taxonomic purposes to record the heads from preparations of cast 
skins and fresh larvae flattened in a standard way. By this method 
all the head parts can be shown in the one diagram. 

The size and appearance of the cocoons have not been used in this 
classification since precise descriptions, either verbally or by drawings, 
were found very difficult. The color patterns of cocoons should be 
useful in future taxonomic work within some genera. 


poc pos 


cS léec] 


Figure 1.—Xorides sp.: a, anterior view of head; B, lateral view of head; c, lateral view of 
mandible; p, spiracle. Explanation of symbols on facing page. Detailed explanation 


on pages 394, 395. 


397 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 


Larval Key to Subfamilies of Ichneumonidae (except Orthocentrinae) 


i 


Mandibles unsclerotized and, of the head sclerites, only the pleurostoma and 
each lateral part of the epistoma are sclerotized; accessory longitudinal 
tracheal commissure not present fee Se Coliyriinae 

Mandibles well sclerotized, or, if not, then labial ecleane present; accessory 
longitudinal tracheal commissure present in thorax ...... . 2 


isualral sclerite presentrc(s to3; was y ative tastings) & Ragrotioden an sice 3 


Labral sclerite absent?. . . . Agha 5 


. Spur of hypostoma meeting stipital eclenes on or near fy 2g eit end or, 


if not, then with each lateral part of labral sclerite broadened into roughly 
quadrate area, or with mandible roughly triangular in shape with no clear 
differentiation between base and blade and with two dorsal rows of large 
teeth Pimplinae other than Pimplini and Acaenitini 
Spur of hypostoma not meeting stipital sclerite close to its median end; each 
lateral part of labral sclerite not broadened into quadrate area; mandible 
with blade clearly differentiated from base ........... + 
Epistomal arch lightly sclerotized * Tryphoninae 
Epistomal arch not sclerotized‘. . Cryptinae 


2 Except Opheltes, Protarchus (Mesoleiinae, Mesoleiini). 
3 Except Idiogramma (Tryphoninae, Idiogrammatini). 


4 Except Gambrus (Cryptinae, Cryptini). 


ExpLANATION oF SYMBOLS ON Ficure 1 


ama: Anterior mandibular articulation 

ant: Antenna 

ant sk: Antennal socket 

app: Anterior pleurostomal process 

at: Anterior tentorial pit 

atr: Atrium of spiracle 

ba: Base of mandible 

bl: Blade of mandible 

ca: Closing apparatus of spiracle 

cd: Cardo 

clp: Clypeus 

cs(decl): Coronal stem of dorsal 
ecdysial cleavage line 

dfm: Cranial depression associated with 
origin of frontal muscles 

epst: Epistoma 

fm: Food meatus 

fr: Frons 

ge: Gena 

hyph: Hypopharynx 

hsp: Sclerotic spur of hypostoma 

hst: Hypostoma 

Ib: Labium 

lbs: Labial sclerite 


Ic: Lacinia 

Im: Labrum 

Ims: Labral sclerite 

Iplp: Labial palp 

md: Mandible 

mplp: Maxillary palp 

mx: Maxilla 

oc: Occiput 

ol: Ocular line 

pa: Parietal region of cranium 
plb: Postlabium 

plst: Pleurostoma 

pma: Posterior mandibular articulation 
poc: Postocciupt 

pos: Postoccipital sulcus 

ppp: Posterior pleurostomal process 
prib: Prelabium 

pt: Posterior tentorial pit 

slo: Salivary orifice 

sp: Silk press 

ss(q): Stipital sclerite 

st: Stipes 

t: Teeth of mandible 


vx: Vertex 


398 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


10. 


Las 


12. 


138. 


14. 


15. 


16. 


ke 


18. 


9s 


Epistomal arch well sclerotized® .... isi At Bae Ae 6 
Epistomal arch very lightly sclerotized or Abeent CPt Sed Lee ae ial 
. Labial sclerite at least moderately sclerotized. . ......... a 
Labial sclerite never more than very faintly sclerotized. ...... 10 
‘ibabialsclerite complete ventrally = 4 92% 2's eo ene 8 
Labial sclerite incomplete ventrally’. . . . 208 9 


. Stipital sclerite and hypostomal spur ell sulévotiedl’ Pirnplinae, Pimplini 


Stipital sclerite and hypostomal spur very lightly sclerotized. 
Pimplinze, Acaenitini 


Setae present on prelabium ............... .. Metopiinae 
Setae absent from prelabium?7 . . f ... . . Anomalinae 
Mandible with grooved blade bosreainine prominent tooth: hypostoma verv 

lightly sclerotized; labial palps not disc-shaped . . . . Orthopelmatinaze 
Mandible without grooved blade containing prominent tooth; hypostoma 

well sclerotized; labial palps dise-shaped . . . . . . . Ichneumoninae 
Labial sclerite sharply pcinted ventrally ..... . . . .. Diplazoninae 
Labial sclerite not sharply pointed ventrally ...... , Ee 12 
Mandible with base sclerotized only at periphery .... . Mesuélicbinine 
Mandible with base evenly sclerotized .. . . . . Slee 13 
Stipital sclerite not extending laterally beyond noi of ene with spur of 

hypostoma. . . . . . . . Plectiscinae 
Stipital sclerite ex ienhine iebeeale Heed ae of meeting with spur of 

hypostoma... . gt 14 


Mandible with numerous fine ee on odor side i Binder : ‘Adelognassinae 
Mandible not toothed or with short blunt teeth on dorsal and ventral sides 
of blades 405 fa: oe 15 
Hypostoma not Seats hevand lateral end ot slipitall Heese, or with 
sclerotic band connecting posterior pleurostomal processes across dorsal 
surface of food meatus, or with twelve or more setae situated largely within 
the membrane of the prelabium. .. . . . . .  Lissonotinae 
Hypostoma extending beyond lateral end of epiptieal selerite; sclerotic band 
not present connecting posterior pleurostomal processes; if numerous 
setae present on prelabium, they are scattered over a very broad labial 


sclerite as well as over the prelabial membrane . . . 3 lee 16 
Many (more than four) setae scattered over broad jabiall aclefite and on 
membrane of prelabium .. . . . . . . Ophioninae, Ophionini 
Not more than four setae on Siento ee . wake A 
Labial sclerite as wide or wider than long (in RAterOnOcrenion directa) and 
hypostoma! spur longer than blade of mandible. . . . . Ave 18 


Labial sclerite longer than wide or, if not, then length of ispostcnnel spur 

is equal to or less than length of blade of mandible. 
Ophicninze, Campoplegini 

Mandible conical in shape with small tooth at apex. 

QOphicninae, Tersilochini 
Mandible not of thisshape. ... . in nee 19 
Closing apparatus of spiracle not well Beierotived ana broader than spiracle. 
Ophioninze, Cremastini 
Closing apparatus of spiracle moderately sclerotized and as wide as or more 
narrow than spiracle. .............. . .. Mesoleiinae 


5 Except Listrodromus (Ichneumoninae, Listrodromini), Triclistus (Metopiinae), Xanthopimpla (Pim- 
plinae, Pimplini). 
6 Except Evxetastes (Lissonotinae). 7 Except Anomalon (Anomalinae. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 399 


In the following descriptions of larval structure in the various 
subfamilies, a list of species studied is given. A list is also given of 
those descriptions in the literature which were thought sufficiently 
accurate and detailed to help workers in identifying material. The 
standard of published descriptions of ichneumonid larvae is still poor 
despite Beirne’s (1941) paper, and many references to descriptions 
have been omitted since the figures given are too small and vague. 

The biological notes given for the various subfamilies and tribes 
are based on those of ‘Townes and Townes (1951) and Walkley (1958). 
The subfamilies, tribes and genera are considered in the order given 
by Townes and Townes (1951), or Walkley (1958), except in the case 
of the Mesochorinae, which are moved from a position following the 
Anomalinae and placed near the Lissonotinae for reasons stated in 
the section dealing with the Mesochorinae. The species are considered 
in alphabetical order. When genera are added to the list of Walkley 
they are placed near what were thought, on larval characters, to be 
closely related genera. 


Subfamily Pimplinae 
Figures 2-10 


On larval characters the Pimplinae fall into two groups; in one are 
the Pimplini and in the other are the Ephialtini, Polysphinctini, 
Neoxoridini, Rhyssini, Theroniini, Labenini, and Xoridini. The 
Acaenitini have a special position. The Pimplini usually have the 
epistoma well sclerotized, the hypostoma short, and the mandible 
large and without teeth. The labial sclerite is longer than wide and 
its ventral side is broad. The labral sclerite is absent. The setae on 
the skin are small. In the second group the epistoma is, at most, 
lightly sclerotized, and the hypostoma is long. The mandibles are 
relatively smaller than in the Pimplini and they usually bear teeth. 
The form of the mandible is similar in the Ephialtini, Neoxoridini, 
Rhyssini, and Labenini in having a slender blade and a dorsal and 
ventral row of small teeth. In the Theroniini and in Delomerista 
(Ephialtini) the blade of the mandible is bifurcated, one part having 
dorsal and ventral teeth and the other being toothless. The labial 
sclerite is basically triangular in shape, although in the Rhyssini and 
Neoxoridini the labial sclerite is pointed ventrally, and in the Theroni- 
ini the ventral part of the labial sclerite is broad so that the sclerite 
resembles that of the Pimplini and Acaenitini. The labral sclerite is 
present. The setae on the skin are relatively longer than in the first 
group. The Acaenitini show characters of both groups. The man- 
dible is similar to that of the Pimplini, and the absence of the labral 
sclerite and the presence of a sclerotized epistoma also suggest this 


400 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


tribe. But the hypostoma is long and the labial sclerite is roughly 
triangular in shape, although the ventral part of the sclerite is broad 
as in the Pimplini. In all tribes of the Pimplinae, with the exception 
of the Labenini and Xoridini, the hypostomal spur meets the stipital 
sclerite on or near the point of meeting of this and the labial sclerite. 
In the Pimplini this results in the sclerites around the mandibles 
having a ringlike appearance. Apart from this one character the 
groups in the Pimplinae are distinct. 

Since the Pimplini (with the possible exception of Jtoplectis) are 
endoparasites which emerge from the pupa of the Lepidopterous host, 
and the remaining tribes of the Pimplinae (with the possible exception 
of some Acaenitini) are ectoparasites, differences in larval structure 
might be expected. The Pimplini, with their large, toothless man- 
dibles and well-developed epistomal and pleurostomal sclerites resemble 
to some degree other ichneumonid larvae with similar habits (see 
pp. 502, 505, 506). 

Beirne (1941) in his classification of larval Ichneumonidae has 
placed the Polysphinctini, Rhyssini, Ephialtini, and Xoridini with the 
Cryptinae. This has not been followed in the present study for, 
although the Pimplini differ in many aspects from the Ephialtini and 
similar tribes, the position of the hypostomal spur gives a common 
character for the subfamily. Where this character is not shown 
clearly, as in the Labenini and Xoridini, the remaining characters 
indicate the relationships of the tribe. In the Labenini the form of 
the mandible closely resembles that of the Ephialtini. In the Xoridini 
the general form of the head sclerites, particularly the labial sclerite, 
resembles that of most Pimplinae, rather than any other Ichneumonid 
group. The form of the mandible of the Xoridini differs from that of 
all other ichneumonids. 

It is significant that the Acaenitini and Theroniini show characters 
resembling those of both the Pimplini and the Ephialtini and similar 
tribes. 


LarvaL Kry 


1. Setae on body longer than blade of mandible . . . . . . . Polysphinctini 
Setae on body not longer than blade of mandible. ........... 2 
2,. Labial sclerite sharply pointed; ventrally = . <:.. . +.-..17.. 4 fase) eee 
Labial sclerite not sharply pointed ventrally. .......2..2....4 
3. Ventral point of labial sclerite continuous with sclerite. . . . . . Rhyssini 
Ventral point of labial sclerite appearing as distinct plate at end of sclerite. 
Neoxoridini 
4, Each lateral part of labral sclerite expanded into quadrate-shaped area. 
Labenini 


Each lateral part of labral sclerite not expanded into quadrate-shaped area . 5 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT AQ] 


5. Ventral part of labial sclerite broad and without lobes on ventral surface. 
Theroniini 
Ventral part of labial sclerite, if broad, has lobes on ventral surface . .. 6 
6. Mandible triangular with no sharp differentiation between base and blade. 
XMoridini 
Mandible with sharp differentiation between base and blade . . Ephialtini 


Tribe Ephialtini 
Figures 2-4 


The epistoma is absent except in Tromatobia where it is present as 
a thin, lightly sclerotized arch; the pleurostoma is lightly sclerotized; 
the hypostoma is well sclerotized and may be enlarged posterior to 
the hypostomal spur; the spur of the hypostoma meets the stipital 
sclerite at its point of meeting the labial sclerite; the stipital sclerite 
is always present but it may be short; the cardo is visible in some 
genera as a lightly sclerotized plate; the labial sclerite is roughly 
triangular in shape; the labral sclerite is present and its median part 
is lightly sclerotized and often bilobed; the mandible in most genera 
has a slender blade with a row of fine teeth on the dorsal and on the 
ventral surfaces (but Tromatobia appears to have only a dorsal row 
of teeth, and in Delomerista diprionis Cushman the blade is bifurcated) ; 
the antenna is papilliform; the closing apparatus of the spiracle is 
narrower than the atrium and is usually separated from the atrium 
by a length of trachea equal in length to that of the closing appa- 
ratus, although in Alophosternum the closing apparatus is long and 
slender; the skin has well developed setae which are usually equal 
in length to the blade of the mandible; in Tromatobia the setae are 
very small and groups of spines are present; spines are absent from 
other genera of the tribe. 

LaRvAL Kry 


ho 


1. Small lobes present on ventral edge of labial sclerite® . 
Small lobes not present on ventral edge of labial sclerite : 
2. Hypostomal spur as broad as or broader than posterior part of upestania 
Calliephialtes, Ephialtes 


co 


Hypostomal spur more narrow than posterior part of hypostoma . . Seambus 
3. Mandible with blade bifureated®. .......... . .. . Delomerista 
Mandible with blade not bifureated . ... . LN xipit AMS ata atatet Ze 4 
4. Enlargement present on posterior part of hy eae renin .... . . Iseropus 
Enlargement not present on posterior part of hypostoma ....... 5 
5. Slender epistomal arch present. . ............. . Trematobia 
Epistomal arch absent ............. ~~... Alophosternum 


8 Except Scambus pomorum (Ratzeburg) (Imms, 1918; Speyer, 1926) and Calliephialtes dimorphus Cush- 
man (Sauer, 1939). 
* Except Delomerista sp. (Morris, Cameron and Jepson, 1937). 


402 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


a) 
| WR 3 ate 
i | Ss.) \ 


Figure 2.—Pimplinae:Ephialtini, head sclerites: a, Scambus hispae (Harris); 8, Alophos- 
ternum folticola Cushman; c, Calliephialtes variatipes (Provancher). (1, antenna; 2, 
spiracle} 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 403 


; uv 
Ss 
O-imm Na 


2 3 


Ficure 3.—Pimplinae:Ephialtini, head sclerites: a, Ephialtes irritator (Fabricius); 8, 
Iseropus stercorator brunneifrons (Viereck); c, Tromatobia rufopectus (Cresson). (1, 
antenna; 2, spiracle; 3, skin.) 


A404 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


(pon 


= le 
O:i mm oon Oa Sa yo 
2 3 


Figure 4.—Pimplinae:Ephialtini, head sclerites: Delomerista diprionis Cushman. (1, 
antenna; 2, spiracle; 3, skin.) 


The above key must be regarded as provisional, for, on the present 
material, it is very difficult to decide on any one characteristic of a 
genus. This may be due to the fact that Scambus, to take one exam- 
ple, is a composite genus. Dr. G.S. Walley (in litt.) has reached such 
a conclusion on Scambus from study of the adults. 

Genus Seambus.—S. hispae (Harris) has been examined (fig. 2). 

S. brevicornis (Gravenhorst) is figured by Thorpe (1930), S. detritus 
‘ (Holmgren) by Salt (1931), S. foliae (Cushman) by Dowden (1941), 
and S. pomorum (Ratzeburg) by Imms (1918) and by Speyer (1926). 
S. pomorum has neither lobes on the ventral surface of the labial 
sclerite nor an enlargement on the hypostoma as has S. hispae. S. 
brevicornis, S. foliae and S. detritus have lobes on the ventral surface 
of the labial sclerite, but an enlargement is not present on the hypo- 
stoma. The ventral surface of the labial sclerite of S. detritus is more 
pointed than in other species. These points are of interest since 
Tromera pomorum (Ratzeburg) and Epiurus brevicornis (Gravenhorst) 
are type species of genera synonymized under Scambus. 

Genus Alophosternum.—aA. foliicola Cushman has been examined 
(fig. 28). The closing apparatus of the spiracle of this genus differs 
from others of the tribe in its long slender shape (fig. 282). 

Genus Calliephialtes.—C. nubilipennis (Viereck) and C. variatipes 
(Provancher) (fig. 2c) have been examined. Both species possess 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 405 


lobes on the ventral surface of the labial sclerite, which is 
slightly pointed. The more posterior part of the hypostoma is 
slightly enlarged in C. variatipes but not in C. nubilipennis. The 
body setae are conspicuously large in C. nubilipennis. C. dimorphus 
Cushman is figured by Sauer (1939) and lobes do not appear to be 
present on the ventral surface of the labium, although the figure is 
too small to show the head structures clearly. 

Genus E’phialtes.—E. irritator (Fabricius) has been examined (fig. 
34). E. elegans (Woldstedt) is figured by Beirne (1942a), H. mani- 
festor (Linnaeus) by Baumann (1933), “. punctulatus Ratzeburg by 
Beirne (1941) and Rosenberg (1934), and #. ruficollis (Gravenhorst) 
by Thorpe (1930). All these species have numerous lobes on the 
ventral surface of the labial sclerite. 

Genus Jseropus.—The species of this genus are gregarious. J. 
californicus Cushman and I. stercorator brunneifrons (Viereck) (fig. 
3B) have been examined. The enlargement of the hypostoma is more 
marked in I. californicus than in I. stercorator brunneifrons. I. 
stercorator stercorator (Fabricius) is figured by Beirne (1941). 

Genus TJromatobia.—Species of this genus parasitize the egg 
cocoons of spiders. TJ. rufopectus (Cresson) (fig. 8c) and 7. zonata 
(Davis) have been examined. The head sclerites are slender with 
no lobes on the ventral surface of the labium and no enlargement of 
the hypostoma. A slight epistoma is visible and teeth appear to be 
present only on the dorsal surface of the mandible. Maneval (1936) 
notes the presence of groups of large hooked spines on the dorsal 
surface of the first seven abdominal segments of Zaglyptus varipes 
varipes (Gravenhorst). Similar spines are present on the abdomen 
of Tromatobia (fig. 3c3). These spines, which appear to be holdfast 
organs, closely resemble the body spines of the Polysphinctini. Like 
Trombatobia and Zaglyptus, the Polysphinctini are parasites of spiders. 

Genus Delomerista.—The species of this genus are parasites of the 
cocoons of sawflies. D. diprionis Cushman has been examined 
(fig. 44). The bifurcate mandibles may be distinctive of this genus. 
Morris, Cameron, and Jepson (1937) figure Delomerista sp., where the 
mandible appears to have the same shape as that of other Ephialtini 
and a more or less complete epistoma is present. It is unfortunate 
that the identification was not taken further. | 


Tribe Polysphinctini 
Ficure 5 


The members of this tribe are parasites of spiders. 

The setae on the head and body are conspicuously long, the setae 
on the body being at least as long as the blade of the mandible; the 
dorsal part of the epistoma is not sclerotized and the pleurostoma is 


406 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


O1mm 


Figure 5.—Pimplinae:Polysphinctini, head sclerites: a, Hymenoepimects sp.; B, Oxyrrhexis 
carbonator texana (Cresson); c, Zatypota sp., antenna not visible on preparation. (Il, 
antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 407 


lightly sclerotized ; the hypostoma is well sclerotized and in Ozyrrhezis 
and Acrodactyla (Beirne, 1941) is expanded at the posterior end; 
the hypostomal spur is well sclerotized, as is the stipital sclerite which 
is expanded at its posterior end in Hymenoepimecis and Oxyrrhexis; 
the labial sclerite is roughly square in shape in Zatypota but roughly 
triangular in other genera, although the ventral part of the sclerite 
is pointed in Zatypota and Acrodactyla; the maxillary and labial 
palps each have one sensillum in Hymenoepimecis, Oxyrrhexis and 
Acrodactyla, but there. are two sensilla to each palp in Zatypota; 
the labral sclerite is lightly sclerotized in its lateral parts but is 
unsclerotized dorsally; the mandible is slender and small teeth are 
present on the dorsal surface of the blade; the antenna is papilliform; 
the closing apparatus of the spiracle is close to the atrium; long setae 
are present on the skin; spines, which appear to serve as holdfast 
organs, are present on the dorsal surface of some body segments. It 
is not possible to distinguish the precise distribution of these spines 
from cast skins. Nielsen (1923, 1935, 1936) figures and describes 
the distribution of these spines in the tribe but gives no adequate 
figure of the head structure of the larvae. 


LARVAL Kry 


1. No enlargement present on posterior part of hypostoma . . Hymencepimecis 
Enlargement present on posterior part of hypostoma........ 2 

2. Lateral end of stipital scleriteexpanded. ... ......... Oxyrrhexis 
Lateral end of stipital sclerite not expanded. . .. ........ 3 

3. Ventral part of labial sclerite with small dorsal projection into prelabial 
ATCA mere cae Seng he ain eae epi, cuktecee Mee EMR, eb when tN odie Zatypota 
Ventral part of labial sclerite without dorsal projection. . . . Acrodactyla 


The following have been examined: Hymenoepimecis sp. (fig. 5A), 
Oxyrrhexis carbonator terana (Cresson) (fig. 5B), and Zatypota sp. 
(fig. 5c). Acrodactyla degener (Haliday) is figured by Beirne (1941). 


Tribe Pimplini 
Figures 6, 7 


The members of this tribe are endoparasites of a variety of lepi- 
dopterous pupae. 

The head sclerites are distinctive in that they appear to form a 
ring around the mouth. The epistoma is well sclerotized, except in 
Xanthopimpla; the pleurostoma is well sclerotized and the hypostoma 
is short in most species; the hypostomal spur is well sclerotized and 
the stipital sclerite is short in most species; the labial sclerite is longer 
than wide with thin lateral parts and a broadened ventral part; 
the mandible is relatively large and has no teeth on the blade; the 
labral sclerite is absent and in some genera the sensilla of the labrum 

504675592 


408 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ps O-1.mm 3 


Figure 6.—Pimplinae:Pimplini, head sclerites: a, Pimpla aequalis Provancher, antenna 
not visible on preparation; B, Apechthis compunctor (Linnaeus). (1, antenna; 2, spiracle; 


3, skin.) 


SHORT | 409 


ICHNEUMONID FINAL INSTAR LARVAE 


Figure 7.—Pimplinae:Pimplini, head sclerites: a, Xanthopimpla stemmator (Thunberg); 
B, Itoplectis behrensii (Cresson); c, Echthromorpha notulatoria (Fabricius). (1, antenna; 


2, spiracle; 3, skin.) 


410 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


are grouped on certain raised areas; the hypopharynx has numerous 
small spine-like projections; the antenna is disc-shaped with the 
circumference lightly sclerotized in some genera, and several sensilla 
may be visible in the central part of the antenna; the atrium of the 
spiracle is broad and shallow and the closing apparatus is close to the 
atrium; the skin is relatively smooth and has small setae. 


LaRvaL Kry 


1. Hypostomal spur with lightly sclerotized lateral expansion . . Xanthopimpla 
Hypostomal spur without lightly sclerotized lateral expansion. . .. . 2 

2. Hypostoma approximately equal in length to blade of mandible . Apechthis 
Hypostoma much shorter than blade of mandible. . ........ 3 

3. Stipital sclerite much shorter than blade of mandible. ..... Itoplectis 
Stipital selerite approximately equal in length to blade of mandible. . . 4 

. Broadened (ventral) part of labial sclerite about as broad as long and not 
pointed? ventrally 9 3 2%. yuh P ean 2 ee ee ee eset ees oe Pimpla 
Broadened (ventral) part of labial sclerite longer than wide and pointed 
VETIballiy ys 14s, cetaeeeerast wos Se fe? GL mee mee ce et eee eta ereiele Eehthromorpha 


= 


Genus Pimpla—P. aequalis Provancher (fig. 6A) and P. inflatus 
Townes have been examined. P. instigator (Fabricius) is figured by 
Beirne (1941) and by Meyer (1922), P. luctunosa Smith by Itawa 
(1950), P. turionellae (Linnaeus) by Thorpe (1930), Silvestri (1941), 
and Beirne (1943), and Pimpla sp. by Beirne (1942). 

Genus Apechthis—A. compunctor (Linnaeus) (fig. 6B) has been 
examined. ‘This species is figured by Beirne (1941). 


Genus Itoplectis—The species of this genus may be primary or 
secondary parasites. Professor Varley (in litt.) informs me that 
I. maculator (Fabricius) appears to be more often a primary parasite 
on Tortrix and other species, although sometimes a secondary para- 
site. J. behrensii (Cresson) (fig. 7B) and J. conquisitor (Say) have 
been examined. J. alternans (Gravenhorst) is figured by Morris, 
Cameron, and Jepson (1937), J. olivalis Thomson by Beirne (1941), 
and Jtoplectis sp. by Snodgrass (1935). Morris, Cameron, and Jepson 
(1937) state that the inner aspects of the teeth of J. alternans are 
armed with bristles. Teeth have not been recorded on the blade of 
the mandible in any other species of this tribe. These authors studied 
this species as a primary parasite of Diprion. It oviposited on pre- 
pupae and pupae within the host cocoon and fed externally. All 
other genera of the Pimplini appear to be endoparasites. 

Genus Echthromorpha.—E. insidiator (Smith) and EF. notulatoria 
(Fabricius) (fig. 7c) have been examined. £. formosa (Smith) is 
figured by Beirne (1941). 

Genus Xanthopimpla.—The broad and lightly sclerotized hyposto- 
mal spur and the lightly sclerotized epistoma are characteristic of this 
genus. The stipital sclerite is short and fused with the hypostomal 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 411 


spur. X. cera (Cameron), X. enderleini Krueger, and X. stemmator 
(Thunberg) (fig. 74) have been examined. X. pedator (Thomson), 
figured by Beirne (1941), differs from other species of the genus in 
that the lateral parts of the labial sclerite are very broad; however, 
the hypostomal spur and the stipital sclerite appear to be similar to 
those of other species. X. citrina (Holmgren) is figured by Moutia 
and Courtois (1952). 
Tribe Neoxoridini 


FIGuRE 8a 


The only species of this tribe examined was Neozorides borealis 
(Cresson) (CDA) (fig. 84). Members of this genus are parasites of 
wood-boring Coleoptera. 

The dorsal part of the epistoma is unsclerotized but the pleurostoma 
and hypostoma are well sclerotized; the hypostomal spur has a broad 
dorsal part but narrows ventrally where it fuses with the narrow 
stipital sclerite; the lateral end of the stipital sclerite is Y-shaped; the 
ventral part of the labial sclerite is broad and pointed, although the 
point appears as a separate cap on the remainder of the labial sclerite; 
the maxillary and labial palps each have two sensilla, one round and 
one crescentic in shape; only the lateral parts of the labral sclerite are 
well sclerotized; the mandible has a slender blade and a dorsal and 
ventral row of small teeth, thus resembling the mandibles of the 
Ephialtini, Rhyssini and Labenini; the antenna is papilliform; the 
atrium of the spiracle is funnel-shaped and the closing apparatus is 
close to the atrium; the skin has setae of moderate size but no spines. 


Tribe Rhyssini 
Figure 8B 


Species of this tribe parasitize wood-boring Coleoptera and sawflies. 
The only specimen examined was Rhyssa sp. (fig. 8B). 

The dorsal part of the epistoma is not sclerotized and the pleuro- 
stoma is lightly sclerotized; the hypostoma and hypostomal spur are 
well sclerotized and the hypostomal spur is broad at its dorsal end; 
the stipital sclerite is well sclerotized and is broad at its lateral end; 
the labial sclerite is sharply pointed ventrally; the maxillary and 
labial palps each have two sensilla, one round and one crescentic; the 
dorsal part of the labral sclerite is not sclerotized; the mandible has a 
slender blade with a dorsal and ventral row of small teeth; there is a 
small, lightly sclerotized plate on the clypeus; the antenna is papil- 
liform; the atrium of the spiracle is funnel-shaped and the closing 
apparatus, which has relatively thick walls, adjoins the atrium; the 
skin has setae of moderate size but no spines. 

FRhyssa sp. is figured by Beirne (1941). 


A12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 8.—Pimplinae, head sclerites: a, Neoxorides borealis (Cresson) (Neoxoridini); 8, 
Rhyssa sp. (Rhyssini). (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 413 


Tribe Theroniini 
FIGURE 9A 


The only species examined was Theronia atalantae (Poda) (fig. 94). 
The species of Theronia are usually primary and secondary parasites 
of the pupae of Lepidoptera. 

The dorsal part of the epistoma is unsclerotized and the pleurostoma 
is lightly sclerotized; the hypostoma and hypostomal spur are slender 
and well sclerotized; the stipital sclerite is lightly sclerotized and a 
very lightly sclerotized cardo is present; the lateral parts of the labial 
sclerite are slender and well sclerotized and the ventral part is broad 
and lightly sclerotized; each maxillary and labial palp has two sensilla, 
one round and one crescentic in shape; the labral sclerite is present; 
the mandible has a broad base and the blade is bifurcated with one 
part bearing dorsal and ventral teeth and the other, which projects 
posteriorly, being toothless; the antenna bears a small papillus; the 
closing apparatus of the spiracle has slender walls and adjoins the 
atrium; the skin bears setae approximately equal in length to that of 
the toothed part of the blade of the mandible and has no spines. 

The mandible and the labial sclerite of the final instar larva of T. 
atalantae are figured by Meyer (1922). 


Tribe Labenini 


FiGurE 9B 


The only species examined was Labena grallator (Say) (fig. 98). 
Species of the genus Labena parasitize wood-boring Coleoptera. 

The dorsal part of the epistoma is very faintly sclerotized; the 
pleurostoma is well sclerotized and has a lightly sclerotized lateral 
expansion; the hypostoma is well sclerotized and has a lightly 
sclerotized posterior expansion; the hypostomal spur is well sclerotized ; 
the stipital sclerite is V-shaped with its lateral part expanded and 
curved dorsally; the labial sclerite is similar in shape to that of many 
Ephialtini, but there are no lobes on the ventral surface; on each 
maxillary and labial palp there are two sensilla, both round and one 
smaller than the other; the labral sclerite is distinctive with each 
lateral part bearing a quadrate-shaped expansion and the median 
(dorsal) part being unsclerotized except for a small, lightly sclerotized 
plate; the antenna is papilliform; the atrium of the spiracle is oval in 
shape and the closing apparatus is thick-walled and situated close to 
the atrium; the skin has setae of moderate size but no spines. 


414 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 9.—Pimplinae, head sclerites: a, Theronia atalantae (Poda) (Theroniini); B, Labena 
grallator (Say) (Labenini). (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT A15 
Tribe Xoridini 
Figure 10a 


Species of this tribe parasitize wood-boring Coleoptera. 

Xorides insularis (Cresson) and X. rileyi (Ashmead) (fig. 104) have 
been examined. 

In the genus Xorides the dorsal part of the epistoma is not scle- 
rotized, but the pleurostoma, hypostoma, hypostomal spur, and 
stipital sclerite are well sclerotized; the labial sclerite is roughly 
triangular in shape; the maxillary and labial palps each bear two 
sensilla, one round and one crescentic in shape; the labral sclerite 
in both species examined does not have the dorsal part sclerotized 
and each lateral part consists of two sclerotic bands which join ven- 
trally; the mandible is distinctive with a broad blade bearing two rows 
of large teeth on the dorsal surface; the antenna is papilliform; the 
spiracular atrium is oval in shape and the closing apparatus is close 
to the atrium; small setae but no spines are present on the skin. 

Ischnoceros rusticus (Geoffroy) is figured by Berine (1941), Xori- 
descopus sp. by Ayyar (1943), X. brachylabris (Kriechbaumer) by 
Chrystal and Skinner (1931), Xorides praecatorius (Fabricius) by 
Beirne (1941), and Xorides sp. by Short (1952). All specimens de- 
scribed in the literature appear to have the labral sclerite consisting 
only of a single sclerotic band and not double as in the species figured 
in the present paper. Chrystal and Skinner show small lobes on the 
ventral surface of the labial sclerite of X. brachylabris. The labium 
thus resembles those of some Ephialtini. These authors also record 
that there is no striking difference between the larva of this species 
and that of X. irrigator (Fabricius) which they also examined. These 
lobes have not been recorded on the labium of any other species of 
this tribe, although Ayyar figures what appears to be a sclerotic plate 
beneath the ventral surface of the labial sclerite in Xoridescopus. It 
is unfortunate that this figure is poor and on bad paper. 


Tribe Acaenitini 
Figure 108 


Members of this tribe are parasites of wood-boring Coleoptera. 
The only species of this tribe examined was Arotes formosus Cresson 
(fig. 108). Cushman and Rohwer (1921, p. 392) state that the group 
is ectoparasitic. The complete epistoma, toothless mandible and 
small skin setae of Arotes suggest that it is an endoparasite, although 
the larval characters of Coleocentrus, as figured by Baumann (1933), 
suggest that this genus is an ectoparasite. 


416 PROCEEDINGS OF THE NATIONAL. MUSEUM VoL. 110 


roo. o> 
AY oan oo PN 
Oo co 


he Erol oa \y: . 
i Se eo tay, gal 
/ 
( , 


O-imm 3 


Ficure 10.—Pimplinae, head sclerites: a, Xorides rileyi (Ashmead) (Xoridini); B, Arotes 
formosus Cresson (Acaenitini), antenna not visible on preparation. (1, antenna; 2, 
spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 417 


It has been already noted that the Acaenitini have characters 
similar to those of the two groups of the Pimplinae, the Ephialtini 
and similar tribes, and the Pimplini. 

In the genus Arotes the epistoma and pleurostoma are moderately 
sclerotized and broad; the hypostoma is moderately sclerotized but 
very slender; the hypostomal spur is broad and very lightly sclerotized ; 
the stipital sclerite is short, broad and very lightly sclerotized; the 
labial sclerite is roughly triangular in shape, but the ventral part is 
much broader than the lateral parts as in the Pimplini; the maxillary 
palps each bear one large, round sensillum and a group of smaller 
sensilla; each labial palp bears one large round sensillum and one 
small sensillum; the labral sclerite is absent and there are two prom- 
inent groups of sensilla on the labrum; the mandibles resemble those 
of the Pimplini in having a large blade and no teeth; the antennae 
were not visible on the specimen; the spiracular atrium is oval, and 
the closing apparatus, which is relatively long and with thin walls, 
is situated adjacent to the atrium; the skin has very small setae but 
no spines. 

Coleocentrus excitator (Poda) is figured by Baumann (1933). The 
antenna of this species appears to be disc-shaped. The epistoma 
appears to be incomplete, but the figure given is too small to make 
this point clear. 


Subfamily Adelognathinae 
Figure 11a 


Members of this subfamily are gregarious external parasites on 
sawfly larvae. Adelognathus and related genera have been placed 
as a tribe of Tryphoninae by some authors. In the present study it 
was found necessary to consider this group as a subfamily since the 
form of the mandible, the absence of an epistoma and a labral sclerite, 
the presence of a prelabial sclerite and of a clearly papilliform an- 
tenna '’ and the absence of spines on the skin all separate Adelognathus 
from other genera of the Tryphoninae. I have been unable to dis- 
cover whether Adelognathus possesses the stalked egg that is charac- 
teristic of the Tryphoninae. The ovaries of an adult were examined 
and the eggs present did not appear to be stalked. 

Adelognathus britannicus Perkins (GCV), A. pallipes (Gravenhorst) 
(GCV) and Adelognathus sp. (@CV) were examined. 

The dorsal part of the cranium is sufficiently sclerotized to appear 
dark; the epistoma is unsclerotized except for small projections above 

10 A papilliform antenna is present in Hybophanes scabriculus (Gravenhorst) and Phytodietus pulcherrimus 
(Cresson) (Simmonds, 1947) of the Tryphoninae. The antenna was not visible in the specimen of P. 
pulcherrimus examined in the present study. In the remainder of the Tryphoninae the antenna is not 


papilliform, although a relatively large, rounded sensillum may be present in the middle of the disc-shaped 
antenna as in Jdiogramma. 


418 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


pam icilamnly all Moab 
ii Wea ea IL 


{ @ @ 
B e 


wp =O) 


ie atine: be a 
: A 


aus © Nites - 


ps 
O-imm 
1.—Head sclerites: Adelognathus pallipes (Gravenhorst) (Adelognathinae); B, 
(Tryphoninae:Phytodietini), antenna not visible on 
(Say) (Tryphoninae:Phytodietini). (1, antenna; 2, 


Ficure 1 
Phytodietus pulcherrimus (Cresson) 
preparation; c, Netelia geminatus 
spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 419 


the anterior pleurostomal processes; the pleurostoma is lightly scler- 
otized; the hypostoma, hypostomal spur and stipital sclerite are well 
sclerotized; the labial sclerite is well sclerotized and the lateral parts 
meet the ventral almost at right angles; a prelabial sclerite is present; 
each maxillary and labial palp bears two sensilla, one round and 
one crescentic in shape; the mandibles are distinctive, being large 
with a long broad blade having a row of fine teeth on the dorsal edge; 
the labral sclerite is absent; the antenna is papilliform; the closing 
apparatus of the spiracle is long and slender and is separated from the 
atrium by a length of trachea equal in length to the atrium; the skin 
bears small setae but no spines; small bubble-like projections are 
present on the skin, but these are not pointed and sclerotized as in 
the Tryphoninae and cannot be called spines. 
Beirne (1941) figures Adelognathus sp. indescr. 


Subfamily Tryphoninae 


Figures 11B—-13p 


Members of this subfamily are ectoparasites and the tribes are 
closely related. The eggs are attached to the skin of the host by a 
stalk (by other structures in some Cteniscini) and the parasite larva 
completes its development in the host cocoon. 

The head sclerites are all well developed, although the epistoma is 
only slightly sclerotized in the Phytodietini and the prelabial sclerite 
is absent; the mandibles are of moderate size and may bear teeth on 
the dorsal edge of the blade or teeth may be absent; the antenna is 
disc-shaped with one to three sensilla in the center; the closing ap- 
paratus of the spiracle is slender and is situated some distance from 
the atrium, except in Netelia where the closing apparatus is as broad 
as the atrium and adjoins the atrium; large setae and small spines 
are present on the skin, except in Hzenterus where the spines are 
conspicuously large and are about half as long as the setae. 


LarvaL Kry 


1. A lightly sclerotized plate is present on the labium ventral to the labial sclerite. 


Phytodietini 

A lightly sclerotized plate is not present on the labium ventral to the labial 
sclerite’= 242 SY SUE ESTEE CL AL Ded TEES, TEED 

2. Dorsal part of epistomia najecleratized” - 2... . . . . . Idiogrammatini 
Dorsal part of epistoma sclerotized. . . . . Sochy sthieop eee 

3. Labial sclerite slender with width much less alan ae of sania sclerite. 
Boéthini 


Labial sclerite relatively broad with width approximately equal to that of 
StiPlcalesCleritet permet cet es Beet n wey veo sema Coane! ss bcc tan neem eee 


420 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


4. Mandible without teeth; setae on body with length equal to one-third to one- 


halfthat of mandibles. 4.0 ses 2 6) se eee et eee eC Lemiseran 
Mandible with teeth," or, if without teeth, then setae on body large with length 
equal to approximately three-quarters that of mandible. . ...... 5 

5. Mandible with teeth; maxillary and labial palps each with three sensilla, or, 
if not, then with two sensilla, one round and one crescentic . . Thymaridini 
Mandible without teeth, or, if with teeth, then maxillary and labial palps each 
with two sensilla both of which areround . ..... .. . . Tryphonini 


Tribe Phytodietini 


Figure 118,c 


The members of this tribe are parasites of caterpillars. The egg is 
attached to the host by a stalk and most of the larval development 
occurs after the host spins its cocoon or makes a pupal cell in the 
eround. 

The lightly sclerotized plate situated on the labium ventral to the 
labial sclerite is characteristic of this group. In contrast to the other 
tribes of the subfamily the dorsal part of the epistoma is very lightly 
sclerotized. 

LARVAL KeEry 


1. Closing apparatus of spiracle narrow and separated from atrium by a length 
of trachea equal to length of closing apparatus. . . . . . . Phytodietus 
Closing apparatus of spiracle broad and situated adjacent to atrium . Netelia 


Phytodietus pulcherrimus (Cresson) (fig. 118), Netelia exserta (Cush- 
man), and N. geminatus (Say) (fig. 11c) have been examined. 

The labral sclerite of N. exserta differs from that of N. geminatus 
in that the dorsal part consists only of a single sclerotic band and the 
lateral parts do not end in forked enlargements. This is of interest 
since Townes and Townes (1951) place exserta in the subgenus Pros- 
thodocis, whereas geminatus is placed in the subgenus Netelza. 

There is a relatively large, round sensillum in the center of the 
antenna in Netelia. The sensillum is, however, much smaller than 
the papillus of a papilliform antenna, and the antenna of Nef¢elia is 
best described as disc-shaped. 

Phytodietus pulcherramus (Cresson) is figured by Simmonds (1947), 
and P. gelitorius (Thunberg) and Netelia vinulae (Scopoli) are figured 
by Beirne (1941). Simmonds does not figure the sclerotized plate on 
the labium of P. pulcherrimus. ‘This is presumably an oversight. 
Beirne describes a lower row of small teeth near the tip of the blade 
of the mandible in Netelia. This row of teeth was not seen in the 
species of Netelia examined. 

11 Teeth are present on the mandible of Erromenus dolichops Townes and Townes (‘Tryphonini) but they 


are very small. This species (fig. 134) may be distinguished from the Cteniscini by the shape of the 
mandible. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 4?1 


Tribe Thymaridini (= Eclytini) 
Figure 12,8 


Members of this tribe are parasitic mostly on small Lepidoptera. 
Eclytus ornatus Holmgren is a parasite of Tenthredinidae. 

The species examined were /. ornatus Holmgren (CDA) (fig. 124), 
E. pleuralis (Provancher) (CDA), Hybophanes scabriculus (Grav- 
enhorst) (GCV) (fig. 128), and Neliopisthus elegans (Ruthe) (GCV). 

All the head sclerites are present with the exception of the prelabial 
and the dorsal part of the labral sclerite is absent in Hybophanes and 
Neliopisthus; the epistoma is slender in Helytus but relatively broad, 
although lightly sclerotized, in Hybophanes and Neliopisthus; the lat- 
eral parts of the labial sclerite meet the ventral almost at right angles 
in Kelytus but the ventral part of this sclerite is rounded in Hybophanes 
and Neliopisthus; in Hybophanes and Eclytus ornatus each maxillary 
and labial palp has three sensilla, one large, one small and one minute, 
but in EF. pleuralis each palp has two sensilla, one round and one cres- 
centic; the mandible has a prominent row of teeth on the dorsal sur- 
face of the blade; the closing apparatus of the spiracle is small and 
the length of trachea which separates it from the atrium is twice as 
long as the closing apparatus in Eelytus and three times as long in 
Hybophanes and Neliopisthus; the setae on the skin are prominent, 
especially in Hybophanes and Neliopisthus, and there are small spines. 

Eclytus fontinalis Holmgren is figured by Beirne (1941). 


LARVAL KEY 


1. Setae on skin of thorax equal in length to blade of mandible . . . . Eelytus 
Setae on skin of thorax as long as or longer than hypostoma. ....... 2 
2 Epistoma sharply arched. .f so 205 eee yk ye os ew eh Hybophanes 
Epistoma not sharply. arched... syne f-fhs 2-5 swe ste SF Neliopisthus 


Tribe Idiogrammatini 


Figure 12c 


Idiogramma sp. was examined. The species of this genus are 
parasitic on the larvae of NXyela feeding in the staminate cones of 
Pinus. Idiogramma is an isolated genus showing no close resem- 
blance to other Tryphoninae. 

The dorsal part of the epistoma is unsclerotized; the pleurostoma 
is very lightly sclerotized; the hypostoma, hypostomal spur, stipital 
sclerite and labial sclerite are moderately sclerotized; each maxulary 
and labial palp has two sensilla; there is no prelabial sclerite; the silk 
press is very lightly sclerotized; a labral sclerite is present; the man- 
dibles are distinctive in shape, each having a somewhat square base 
and a curved toothless blade; the antenna has a large central sensillum 


t 


22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


sacl siete Sc Daae 
KU AAMT aan A AS A 


+——_—____+ 
aK O:imm 3 


Ficure 12.—Tryphoninae, head sclerites: a, Eclytus ornatus Holmgren (Thymaridini), 
antenna not visible on preparation; B, Hybophanes scabriculus (Gravenhorst) (Thymari- 
dini), with skin showing prothoracic setae and spines; c, [diogramma sp. (Idiogrammatini). 
(1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 423 


but is not clearly papilliform; the closing apparatus of the spiracle 
is separated from the atrium by a length of trachea approximately 
equal to the depth of the atrium; the skin bears small setae and spines. 

It is possible that Jdiogramma should be placed in a subfamily 
of its own since the shape of the mandible and the incomplete epistoma 
separate this genus from other Tryphoninae. However, since the 
epistoma is partly sclerotized dorsal to the anterior tentorial pits, a 
labral sclerite is present, the antenna is not papilliform, and the 
eggs are stalked, the genus has been placed with the Tryphoninae. 
The structure of the spiracles and the presence of distinct spines on the 
skin are characteristics similar to those of many Tryphoninae. The 
characters of Jdiogramma do not resemble those of any other sub- 
family of the Ichneumonidae. 


Tribe Boéthini 
FIGuRE 134 


Boéthes sp. was examined. Members of this genus are parasitic on 
the larvae of argid sawflies. 

All the head sclerites are present with the exception of the prelabial; 
the labial sclerite is slender and its lateral parts meet the ventral 
almost at right angles; each maxillary and labial palp bears two 
sensilla, both round and one large and one small; two groups each of 
four sensilla are present on the labrum; the mandibles are without 
teeth; the antenna is disc-shaped with a prominent sensillum in the 
center; the length of trachea separating the closing apparatus of the 
spiracle from the atrium is almost equal to that of the closing appara- 
tus; the skin bears large setae and small spines. 


Tribe Tryphonini 


FIGurRE 138,c 


Members of this tribe are parasites of the larvae of Tenthredinidae. 
Polyblastus pedalis (Cresson) (CDA) (fig. 133), Hrromenus bedardi 
Provancher (CDA), and EF. dolichops Townes and Townes (CDA) 
(fig. 13c) have been examined. 

All the head sclerites are present with the exception of the pre- 
labial; the ventral part of the labial sclerite is pointed or curved so as 
to give the sclerite an oval to round appearance; each maxillary and 
labial palp bears three sensilla in Pulyblastus and two in Erromenus; 
the mandible is without teeth in Polyblastus but has small teeth in 
Erromenus: the disc-shaped antenna has two sensilla; the distance of 
the closing apparatus of the spiracle from the atrium is about equal 
in length to the closing apparatus in Polyblastus, but the distance is 
about twice the length of the closing apparatus in Hrromenus: small 
spines are present on the skin of both Polyblastus and Hrromenus, but 

504675—59 —3 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


424 


O:! mm 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 495 


2 


Ficure 13.—Head sclerites: a, Boéthes sp. (Tryphoninae:Boéthini); 8, Polyblastus pedalis 
(Cresson) (Tryphoninae:Tryphonini); c, Erromenus dolichops Townes and ‘Townes 
(Tryphoninae:Tryphonini); p, Exenterus canadensis Provancher (Tryphoninae:Cteniscini) ; 
E, Brachycyrtus pretiosus Cushman (Cryptinae:Brachycyrtini), antenna not visible on 
preparation. (1, antenna; 2, spiracle; 3, skin.) 


426 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


the length of the skin setae is equal to three-quarters of the length 
of the mandible in Polyblastus, whereas in Erromenus the setae are 
small and equal to one-fifth of the length of the mandible. 


LARVAL Kry 


1. Mandible without teeth; three sensilla on each maxillary and labial palp. 
Polyblastus 
Mandible with teeth; two sensilla on each maxillary and labial palp. 
Erromenus 


Tribe Cteniscini 
Figure 13p 


Members of this tribe parasitize the larvae of Argidae, Tenth- 
redinidae and Diprionidae. LEzenterus abruptorius (Thunberg), £. 
adspersus Hartig, £. canadensis Provancher (fig. 13D), H. claripennis 
Thomson, and £. tricolor Roman have been examined. 

All the head sclerites except the prelabial are present; the lateral 
parts of the labial sclerite meet the ventral almost at right angles; 
each maxillary and labial palp has two sensilla, both round, one large 
and one small; the mandible is without teeth; the antenna is disc-like 
and has three prominent sensilla; the closing apparatus of the spiracle 
is long and slender and its distance from the atrium is roughly equal 
to the length of the closing apparatus; spines are conspicuous on the 
skin and their length is about half that of the setae. 

The following are figured in the literature: Hxenterus abruptorius 
(Thunberg) (Morris, 1937), E. adspersus Hartig (Beirne, 1941), £. 
tricolor Roman (Morris, Cameron, and Jepson, 1937). 


Subfamily Cryptinae (= Gelinae) 
Figures 135-27, 64 


Most members of this subfamily are ectoparasites but some are, or 
can be, endoparasites. Mr. J. F. Perkins (in litt.) has informed me 
that in Polytribaz (Aptesini) the parasites emerge from the pupa of 
the host. Rosenberg (1934) states that he reared Cryptus sexannulatus 
Gravenhorst and Agrothereutes batavus Vollenhoven from cocoons 
within the host pupae, but that when he reared the larvae of these 
species they fed externally on host larvae. 

In this very uniform group, the head sclerites are, in general, well 
developed, although the prelabial sclerite is absent and the epistoma 
is very lightly sclerotized and is slender and more or less incomplete 
dorsally, except in Gambrus where the dorsal part is broad and com- 
plete; the pleurostoma is usually very lightly sclerotized; the labial 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 427 


sclerite is roughly triangular in shape in most genera, but, as stated 
below, this sclerite may be circular or otherwise modified; the silk 
press is well sclerotized and conspicuous and two small sensilla are 
often visible just beneath the external opening of the press; each 
maxillary and labial palp usually bears two sensilla, one round and 
one crescentic in shape, although three or more sensilla are present 
in some genera; the labral sclerite is usually lightly sclerotized in its 
dorsal part and this part may be enlarged; the mandibles are rela- 
tively small in most genera, except in the Aptesini where they are 
of moderate size; teeth may be present on the dorsal and ventral 
surfaces of the blade of the mandible, on the dorsal surface only, or 
teeth may be absent; the antenna is usually papilliform, although 
it is disc-shaped in some Hemitelini; the closing apparatus of the 
spiracle may be some distance from the atrium or it may be close to 
the atrium; the skin bears setae and small projections which may be 
sufficiently sclerotized to appear as spines. 


LARVAL Kry 


1. Hypostoma posterior to hypostomal spur shorter than hypostomal spur. 
Brachycyrtini 
Hypostoma posterior to hypostomal spur at least as long as hypostomal spur. .2 
2. Mandible with a long slender blade bearing a row of fine teeth along entire 
lengthrofidorsal surfacentit “tse 6s ss ee oye eee oa 
Mandible not of thisform .. . Seen re 
3. Teeth on dorsal surface of blade ‘or mendinle eee ihan fectte on ventral 
surface!2 which are very small and few in number, except in Ihembobius 
(fig. 214) where the blade of the mandible is short, broad and well sclerotized 
with length equal to one third of that of base of mandible and width of 
lateral end equal to one third of width of base ... . . . . Aptesini 
If teeth present on dorsal and ventral surfaces of blade of eons and teeth 
on dorsal surface markedly larger than teeth of ventral surface, then ventral 
teeth numerous and blade of mandible longer and more slender than in 
Rhembobius, with length greater than one third of that of base of mandible 
and with width of lateral end less than one third of thatofbase ..... 4 
4. Closing apparatus of spiracle separated from atrium by length of trachea at 
least equal to length of closing apparatus, or, if not, then epistoma not 


sclerotized dorsal to anterior tentorial pits . ...... . . Hemitelini 
Closing apparatus of spiracle adjoining atrium, or if not, then at least part 
of epistoma dorsal to anterior tentorial pits lightly sclerotized 14. . Cryptini 


12 Except Cubocephalus where the form and position of the closing apparatus of the spiracle differentiates 
this genus from Cryptini with a similar type of mandible and either the form of the labial sclerite or the 
form and position of the closing apparatus of the spiracle differentiates this genus from Hemitelini with 
a similar type of mandible. 

18 Except Trychosis, Mallochia and Lymeon (Cryptini). The following characters will distinguish these 
genera from the Aptesini: the form of the labial sclerite in Trychosis, the sharp dorsal bend of the lateral end 
of the stipital sclerite in Mallochia, and the heart-shaped dorsal sensillum on each labial palp in Lymeon. 

14 Except Pseudischnus. 


428 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Tribe Brachycyrtini 
FIGURE 13,5 


Brachycyrtus pretiosus Cushman (fig. 13,5) was examined. 

Members of the genus Brachycyrtus are parasitic in the cocoons of 
Chrysopidae. The Brachycyrtini have been placed by some authors 
in a tribe of the Pimplinae. From a study of the adults, Walkley 
(1956) reached the conclusion that the group fits best in the Cryptinae, 
but that additional material and knowledge might prove that it 
belongs in a subfamily of its own. The larval characters support this 
conclusion. The form of the mandibles and head sclerites resemble 
those of the Cryptinae rather than any other group. The position 
of the hypostomal spur differs from that of the Pimplinae other than 
the Labenini and Xoridini. The form of the labral sclerite differs 
from that of the Labenini. The form of the mandible differs from 
that of the Xoridini in that the blade is relatively longer. The 
group is an isolated one on larval characters, and the short hypos- 
toma is distinctive. The form of the hypostoma resembles that of 
the Sphecophagini to some extent, although the hypostoma of the 
Brachycyrtini is considerably shorter than that of the Sphecophagini. 
The head of Brachycyrtus is small and delicate and the antenna was 
not clearly visible on the preparations made. It appears to be 
dise-shaped. 


Tribe Hemitelini (= Gelini) 
Fiaures 14-19, 64 


Members of this tribe attack a great variety of hosts. Many are 
occasionally or habitually secondary parasites. The most conspicuous 
characters of the tribe are the poorly developed epistoma and the 
distance of the closing apparatus of the spiracle from the atrium. 


LARVAL Kry 


1. Mandible without teeth . 
Mandible with teeth . ait : 
2. Each ventrolateral part of labial aclnite eeeaieds 
Labial sclerite not of this shape . 
3. Ventral end of hypostomal spur broader than orsale 
Isdromas peruviana (Viereck) 


Own bd 


Ventral end of hypostomal spur narrower than dorsal. . . ‘ 4 

4. Mandibles with relatively broad blades, the tips of which da non sce in 
the median line. . . . . . . . . Isdromas lycaenze (Howard) 
Mandibles with relatively narrow ipiedes the tips of which meet in the median 
line! asexcivaets So a ale See. Joos: RE AOe, diek copie ee ar 
5. Dorsal part of abel nalerite moneda bo aed dé Ac Aten ere Srilpaus 


Dorsal part of labral sclerite not pointed. ....... . . Atractodes 


10. 


Wale 


12. 


13. 


14. 


15. 


16. 


1 


18. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 429 


. Labial sclerite round in shape and with lateral parts not meeting ventral 


approximately atright angles... . ... . . Thaumatotypus 
Labial sclerite not round in shape and aii lateral parts meeting ventral 
approximately atrightangles. .. . ... . . Phygadeuvon 

. Mandible with teeth only on dorsal eeniace of pie THe Coe at, rae! TS 
Mandible with teeth on dorsal and ventral surfaces of blade ™. .... I1 

. Teeth few in number and relatively large. . . .... =... +. Bathythrix 
Teeth smallandfine. .. . Aapaney  Mtaaee et Ge ae, 

. Two sensilla present on each masilary acl fatal palp ... . . Otacustes 
Sensilla not as above. .. . Rois ha, tee teeny MUD 
Three sensilla, two large amd! one ervey meet on each maxillary and 
aisle pallor eg ses el osc ... . . Mastrus molestae (Uchida) 


One large sensillum civuated dort and two or three small sensilla situated 
ventrally in the form of a crescent on each maxillary and labial palp. 


Thysiotorus 
Labial sclerite approximately circularin shape. ..... . . Ethelurgus 
Labial sclerite not approximately circular in shape; lateral parts meet ventral 
approximately at right angles... . Ao ds ocean ble 
Papillus of antenna reduced to disc feanne three Pane sada a Say foe eel selaix 
Antenna not of thisshape. .. . UL eee areas 
Two sensilla present on each mamiltars and labial Saree SR SOM ee (5) 
Sensilla not as above. . . . HR er 
Two sensilla of equal size Poeeee eile one or tive sivatlcn seviciiln present on 
each maxillary and labial palp. ... . Marae oe es: 
One large ventral sensillum and a dorsal group Ge prince sensilla present on 
each maxillary and labial palp. . . . . . . Mastrus smithii (Packard) 
Dorsal sensillum on each maxillary and labial palp larger than ventral. 
Endasys 
Dorsal sensillum on each maxillary and labial palp smaller than ventral or 
equal in size to ventral. .... ray oop doe ees Gee LO 


Pleurostoma more lightly sclerotized than enoctonay 
Gelis tantillus (Cresson) 


Pleurostoma as well sclerotized as hypostoma............ 17 
Labral sclerite with single median expansion. . . ... . . Hemiteles 
Labral sclerite not expanded in its median part and w an two small dorsolateral 

expansions. . . . . . . Mastrus argeae (Viereck) 


Mandible with raanded oe And rebar short blade. 
Gelis bruesii (Strickland) 
Mandible with a conical base and relatively long slender blade . . Phobetes 


Of the characters used in the above key, the form of the labral 


sclerite and its degree of sclerotization is not a good one. In some cases 
the character is clear, but in others it may be obscured when the entire 
length of the sclerite has not been forced into the one plane during 
mounting. However, since the tribe is extremely uniform and the 
labral sclerite does differ in shape and degree of sclerotization in dif- 
ferent genera, it was found necessary to use this character. 


15 Teeth on the ventral surface of the blade of the mandible are usually very small and must be looked for 
with great care. 


430 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


) fronmli 


J i 
Be Ge > ye 


2 O:1mm 3 


Ficure 14.—Cryptinae:Hemitelini, head sclerites: a, Bathythrix sp., antenna not visible on 
preparation; B, Mastrus molestae (Uchida); c, Otacustes sp. (1, antenna, 2, spiracle; 
3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 431 


BA enon Od 
Ba SRR RASS 


O-1mm 


Ficure 15.—Cryptinae:Hemitelini, head sclerites: a, Ethelurgus syrphicola (Ashmead); B, 
Isdromas peruviana (Viereck); c, Isdromas lycaenae (Howard). (1, antenna; 2, spiracle; 


3, skin.) 


432 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


2 O:-1 mm 3 


Ficure 16.—Cryptinae:Hemitelini, head sclerites: a, Thaumatotypus paradoxus (Zetter- 
stedt); B, Phobetes thyridopteryx (Riley). (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 433 


wan bn nb —- 
Raieyai~ Tn laliwn 
a Nox 


Ficure 17.—Cryptinae:Hemitelini, head sclerites: a, Gelis tantillus (Cresson); 8, Hemiteles 
areator (Panzer); c, Endasys subclavatus (Say). (1, antenna; 2, spiracle; 3, skin.) 


434 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


3 

2 O-tmm 

Figure 18.—Cryptinae:Hemitelini, head sclerites: a, Thysiotorus sp.; B, Phygadeuon 
subfuscus Cresson; c, Iselix sp. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 435 


a 


Ficure 19.—Cryptinae:Hemitelini, head sclerites: Stilpnus anthomyidiperda Viereck. 
(1, antenna; 2, spiracle; 3, skin.) 


+} 
O:-{mm 3 


The following have been examined: Bathythriz sp. (fig. 14a), 
Mastrus argeae (Viereck), M. molestae (Uchida) (fig. 148), M. smithii 
(Packard), Otacustes sp. (fig. 14c), Ethelurgus syrphicola (Ashmead) 
(fig. 154), [sdromas peruviana (Viereck) (fig. 158), I. lycaenae (How- 
ard) (fig. 15c), Thawmatotypus paradoxus (Zetterstedt) (fig. 16a), 
Phobetes thyridopteryx (Riley) (fig. 168), Gelis bruesit (Strickland), 
G. tantillus (Cresson) (fig. 174), Hemiteles areator (Panzer) (fig. 178), 
Endasys subclavatus (Say) (fig. 17c), Thysiotorus sp. (fig. 184), Phy- 
gadeuon subfuscus Cresson (fig. 188), Jseliz sp. (fig. 18c), Stilpnus 
anthomyidiperda Viereck (fig. 194), Atractodes sp. (GCV) (fig. 64a). 

Of these specimens, Mastrus molestae (Uchida) differs from the other 
species of this genus examined in that the mandibular teeth are small 
and are present only on the dorsal surface of the blade. M. smithii 
(Packard) has one large sensillum and a group of smaller sensilla on each 
maxillary and labial palp and M. argeae (Viereck) two sensilla. In 
the two species of /sdromas examined the mandible is basically similar 
and the ventrolateral part of each labial sclerite is expanded. There 
are, however, many differences between these species. The larval 
characters of Gelis, Hemiteles, and Endasys indicate a close relation- 
ship. Members of the genus Gelis attack various small cocoons, 


436 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


including those of Ichneumonidae and Braconidae, and many para- 
sitize the egg cocoons of spiders. 

Phygadeuon, Iselix, Stilpnus, and Atractodes resemble each other in 
larval characters and all are parasites of muscoid Diptera. Dis- 
tinctive features of Atractodes are that the head and the base of the 
mandibles are lightly sclerotized and the well-sclerotized blade of the 
mandible is toothless, the several very small ridges present on the 
dorsal surface of the blade not being distinct enough to be called teeth. 
The antenna, like that of Stilpnus, is disc-shaped, but Stdpnus differs 
from Atractodes not only in the key characters but also in having the 
blade of the mandible more slender, the ventral part of the labial 
sclerite more sharply narrowed in the median line, and the posterior 
end of the hypostoma straight instead of slightly bent towards the dor- 
sal surface (compare figs. 19 and 64). 

The following are figured in the literature: Otacustes crassus 
patruelis (Cushman) (Clancy, 1946), Gelis melanocephala (Schrank) 
(Beirne, 1941), G. tenellus (Say) (Clancy, 1946), Hemiteles areator 
(Panzer) (Morris, Cameron, and Jepson, 1937), H. hemipterus (Fab- 
ricius) (Salt, 1931), H. simillimus (Taschenberg) (Blunk, 1952). 


Tribe Aptesini 


Figures 20-214 


Most of the members of this tribe are parasites of sawflies. Char- 
acteristics of the larvae are the development of the lateral parts of the 
epistoma dorsal to the anterior tentorial pits and the relatively large 
mandibles. The teeth on the dorsal surface of the blade of the mandi- 
ble are larger than the teeth on the ventral surface in Aptesis and 
Rhembobius but not in Cubocephalus. 


LARVAL Kry 


1. Teeth on the dorsal surface of the blade of the mandible not markedly larger 


than: teeth: on.the) ventral'surface 4.1 ..4t) dimen}. 2195: Cubocephalus 
Teeth on dorsal surface. of blade of mandible larger than teeth on ventral 
STITEACC ccs sisiate Mae yea Ive oe yon hata artes nas ha dads ec bed sos sonics Elec aeis AoE 2 

2. Blade of mandible short, broad, and well-sclerotized, with length approximately 
equal to one-third of that of base of mandible. . ... . Rhembobius 
Blade of mandible relatively slender with length approximately equal to one- 
halfi of thatoftbase of mandible 4 Ou.u WiUAsTe GEIR: 8G 28 e Aptesis 


The following have been examined: Cubocephalus sp. (fig. 20a), 
Aptesis indistincta (Provancher) (fig. 20B), A. pteronorum (Graven- 
horst), and Rhembobius abdominalis (Provancher) (fig. 214). 

The teeth on the dorsal surface of the blade of the mandible of A. 
pteronorum are smaller than in A. indistincta. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 437 


teh A (SAUL A 
on 
O:imm 3 


Ficure 20.—Cryptinae:Aptesini, head sclerites: a, Cubocephalus sp.; B, Aptesis indistincta 
(Provancher). (1, antenna; 2, spiracle; 3, skin.) 


438 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


t—————II 
O:1mm 3 


Ficure 21.—Cryptinae, head sclerites: a, Rhembobius abdominalis (Provancher) (Aptesini); 
B, Sphecophaga thuringiaca Schmiedeknecht (Sphecophagini). (1, antenna; 2, spiracle; 
3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 439 


The following are figured in the literature: Cubocephalus abdominator 
(Gravenhorst) (Rosenberg, 1934), C. brumatae (Silvestri) (Silvestri, 
1941), Aptesis basizonia (Gravenhorst) (Morris, Cameron, and Jepson, 
1937). 


Tribe Sphecophagini 
Figure 2138 


Members of this tribe are parasitic in the nests of Vespula and 
Polistes. 

The mandible is characteristic with its long slender blade bearing a 
row of fine teeth along the entire length of the dorsal surface. 

Sphecophaga thuringiaca Schmiedeknecht (BM) (fig. 2138) and 
Sphecophaga sp. (CDA) were examined. 

Sphecophaga vesparum (Curtis) is figured by Beirne (1941) and 
Sphecophaga sp. by Snodgrass (1935). 


Tribe Cryptini (= Mesostenini) 
FIGURES 22-27 


Members of this tribe are parasitic mostly on pupae and prepupae, 
or on larvae concealed in tunnels or leaf rolls. Hosts include Lepi- 
doptera, Symphyta, Aculeata, and spider eggs. 

The larvae have the lateral parts of the epistoma well developed 
and, in many genera, the closing apparatus of the spiracle adjoins the 


atrium. 
LarvaL Kry 


1. Epistoma lightly sclerotized, broad and complete. . ... . . Gambrus 
Epistoma lightly sclerotized, narrow and more or less incomplete dorsally . 2 
2. Labial sclerite with each ventrolateral part indented... . . Trychosis 
Labial sclerite not of thisform .. . Me AAPG Zar Weteo uclg har eee ete 
3. Closing apparatus of spiracle adjoining Meriuea: Sis et ERM alter e ese: 
Closing appratus of spiracle not adjoining atrium. . . Selena LO 
4. Dorsal part of labral sclerite expanded into three Henly pclenotized lobes. 
Cryptus 
Dorsal part of labial scleritenot-of thisformys.. . <<... . 205... 2°98 
Deapixisetac) On prelabium™“2-prnes sas noma es. Bech a fis Sl, odes aes) 6 
Four setae on prelabium. .. . s Aa A ot ands 
6. Median-dorsal part of labral sclerite sah Honl, pelerotiaed enlancement! 
Goryphus 
Median-dorsal part of labral sclerite without enlargement .. . . Mesostenus 
7. Dorsal part of labral sclerite divided into two lightly sclerotized bars. 
Polycyrtus 
Dorsal part of labral sclerite not divided into two bars. ....... 8 
8. Mandible with a relatively slender blade bearing a row of fine teeth on the 
dorsal surface and teeth on ventral surface few and small. . Mallechia 
Mandiblemotvotthis forms \hemae ee ff eas SR pad ee Rucae eee Ono 


504675—59—_4 


440 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


———} ; 
O:1mm 3 


2 qv 


Figure 22.—Cryptinae:Cryptini, head sclerites: a, Cryptus albitarsus albitarsus (Cresson); 
B, Mesostenus sp. nr. gracilis Cresson; c, Polycyrtus semialbus (Cresson), antenna not 
visible on preparation. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 44] 


Figure 23.—Cryptinae:Cryptini, head sclerites: a, Trychosis sp.; B, Gambrus canadensis 
(Provancher). (1, antenna; 2, spiracle; 3, skin.) 


442 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


b ° 
ee 
’ — Stes 1 


Cc 
ce 


e 


Deane 

O-{mm 2 b--nene 

Ficure 24.—Cryptinae:Cryptini, head sclerites: a, Goryphus inferus (Szepligeti); 8, Hoplo- 
cryptus incertulus Cushman, antenna not visible on preparation. (1, antenna; 2, spiracle; 


3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 443 


SS 
O-imm 
2 i) 
Ficure 25.—Cryptinae:Cryptini, head sclerites: Nippocrypius suzukii (Matsumura) 
(1, antenna; 2, spiracle; 3, skin.) 


| Bey Ae a el a el 


a a ~ =~ Ce 


9. Width of closing Se of spiracle ee equal to half width of 


SUEMUMA es ass . .. . . Agrothereutes 
Width of closing AOD atts of Asn more filan half width of atrium. 

Eripternimeorpha 

10. Ventral row of teeth on mandible poorly developed. . . . .. . Lymeon 

Ventral row of teeth on mandible well developed. . ... . ae pte meerralel| 

11. Epistoma not sclerotized dorsal to anterior tentorial pits . . . Pestiditehens 

Epistoma sclerotized dorsal to anterior tentorial pits... . eco ereap les 

12. Six large setae on labrum... . Re MEN re Bea F otc hs Hoplocryptus 

More than six large setae on jabrum ee ae eae rae 

13. Dorsal part of labral sclerite with three fences Releronzed inpes . . Echthrus 


Dorsal part of labral sclerite with one lightly sclerotized lobe . Nippoecryptus 

The following have been examined: Cryptus albitarsus albitarsus 
(Cresson) (fig. 22a), Mesostenus gracilis Cresson, Mesostenus sp. nr 
gracilis Cresson (fig. 228), Polycyrtus semialbus (Cresson) (fig. 22c), 
Trychosis sp. (fig. 234), Gambrus canadensis (Provancher) (fig. 238), @ 
stokesi Cameron, Goryphus inferus (Szepligeti) (fig. 244), Hoplocryptus 
incertulus Cushman (fig. 248), Nippocryptus suzukii (Matsumura) 
(fig. 25a), <Agrothereutes grapholithae (Uchida), <Agrothereutes sp. 
(fig. 26a), Pseudischnus oregonensis (Cushman) (fig. 268), Eripterni- 
morpha javensis Rohwer (fig. 26c), Mallochia sp. (fig. 274), Lymeon 
orbum (Say) (fig. 278), Echthrus niger Cresson (fig. 27c). 


OF THE NATIONAL MUSEUM VOL. 110 


PROCEEDINGS 


y «K——4 =A (AEA IER Lag rzh— 
2 | O:1mm 3 


? ; @ 
oot e 2 tO 

5 vw \ 

ono @ ) @ an 

& 
on ey) \ 2 v 
a A) 
rc « BAD eh Ey 
-— AVA REN, | RK 
2 O-{mm 3 


Ficure 26.—Cryptinae:Cryptini, head sclerites: a, Agrothereutes sp.; B, Pseudischnus 
oregonensis (Cushman); c, Eripternimorpha javensis Rohwer. (1, antenna; 2, spiracle; 


3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 445 


Ficure 27.—Cryptinae:Cryptini, head sclerites: a, Mallochia sp.; 8, Lymeon orbum (Say); 
c, Echthrus niger Cresson. (1, antenna; 2, spiracle; 3, skin.) 


446 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


Agrothereutes grapholithae (Uchida) differs from the species illus- 
trated (fig. 264) in that there is but one median expansion of the 
labral sclerite and there are eight large setae on the labrum instead 
of six. 

The following are figured in the literature: Cryptus inornatus Pratt 
(Simmonds, 1948), C. laborator laborator (Thunberb) (Beirne, 1941), 
CO. sexannulatus Gravenhorst (Rosenberg, 1934), Agrothereutes abbrevi- 
ator (Fabricius) (Morris, Cameron, and Jepson, 1937), A. batavus 
Vollenhoven (Rosenberg, 1934), Zschnus polytomi (Tschek) (Morris, 
Cameron, and Jepson, 1937), Echthrus reluctator (Linnaeus) (Bau- 
mann, 1933). 


Subfamily Ichneumoninae (= Joppinae, = Phaeogeninae) 
FicurEs 28-37 


This is a very distinct and uniform subfamily whose members are 
internal parasites of the Lepidoptera. They oviposit in the host larva 
or pupa but always emerge from the pupa. 

The epistoma is lightly to heavily sclerotized and the pleurostoma 
and hypostoma are always well sclerotized; sclerotized areas may be 
present lateral and dorsal to the pleurostoma and epistoma and, to a 
more limited extent, the hypostoma; sensilla are frequently present 
on the epistoma and pleurostoma; the hypostomal spur and stipital 
sclerite are absent; the labial sclerite is usually absent but may some- 
times be seen as a very lightly sclerotized strip at the edge of the pre- 
labial area; each maxillary and labial palp is in the form of a flattened 
dise bearing several sensilla, usually about five to each palp; the silk 
press is lightly sclerotized and two small sensilla are present ventral 
to the press; the prelabial sclerite is absent; groups of setae and sensilla 
are situated on the clypeolabral area, often on lightly sclerotized 
plates the shape of which is often characteristic of a genus; the mandi- 
bles are large and well sclerotized and do not have teeth on the blade; 
the antenna is disc-shaped; the spiracle has an oval atrium, and the 
large, thick-walled closing apparatus adjoins the atrium; the skin is 
smooth with small setae and no spines. 


LARVAL Kry 


1; Ely postomaisharply bent? 45 Sy Sas A yo sah cp nC Trogini 
Hypostoma may be curved but never sharply bent. .......... 2 

2. Mandible broad and length of blade is equal to or less than one-third length 
Of niandilblel © oss alk ae ere See ne Listrodromini 
Mandible less broad and length of blade is more than one-third length of 
mandibles ie cies eS ed EVR ous cee uae eae 3 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 447 


poo 
O-{mm 


Figure 28.—Ichneumoninae:Phaeogenini, head sclerites: a, Phaeogenes hariolus (Cresson); 
B, Rhexidermus huardi (Provancher). (Antenna not visible on preparation; 2, spiracle; 


3, skin.) 


448 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


3. A lightly sclerotized plate is present on the clypeus ventral to and continuous 
with the epistoma and dorsal to the anterior pleurostomal processes; in 
Phaeogenes and in Diadromus (Given, 1944) the ventrolateral edges of the 
plate are prominent as sclerotized bars .. . . . . . . Phaeogenini 

A lightly sclerotized plate of this shape is not Oreae at ontheclypeus... .4 

4. Clypeolabral region with two slender and lightly sclerotized bands curving 
dorsally and with a line of setae which follows the curve situated beneath 
eachuband, (7 eisleus . . . . . Platylabini 

Clypeolabral region Sivech Erieranzed pamide of this Shes and position 18 . 5 

5. The sclerotized area lateral to each pleurostoma is almost as well sclerotized 
as the pleurostoma so that the lateral limit of the sclerite is not clearly 
defineds+1s2 . 5s . . . . . Eurylabini 

If sclerotized area Bronce stew ee each Plemrockarn then plouensnome 
more heavily sclerotized than this area and with lateral limit clearly 
defined 7... . 2 leculke a6 

6. Setae and sensilla on elypeomsorite aon _ Suna E in ao clongateocl areas 
with a circular area containing two sensilla beneath each oval area. 

Protichneumonini 
Setae and sensilla of clypeolabral region not as above . . . . Ichneumonini 


The differences between most tribes are slight in this uniform 
subfamily. During mounting the lightly sclerotized parts of the 
cranium adjoining the epistoma are flattened into various shapes. 
The figures show this region as it appears in the preparations, but 
it is unlikely that these shapes have any taxonomic significance. 


Tribe Phaeogenini (= Alomyini) 
FIGuRE 28 


LarvaL Kry 


1. With a sclerotized bar extending across each ventrolateral edge of the lightly 
sclerotized plate on the clypeus ............ .. . Phaogenes 
Diadromus 18 

No sclerotized bar present along each ventrolateral edge of the lightly sclero- 
tized plate.on the clypeus. 7.) . .}}... ..\. ... & . “Rhexidermus 


The following were examined: Phaeogenes ater (Cresson), 
haeussleri Uchida, P. hariolus (Cresson) (fig. 284), P. hebrus (Cresson), 
P. vincibilis (Cresson), P. walshiae walshiae Ashmead, Rhexidermus 
huardi (Provancher) (fig. 288). 

The following are figured in the literature: Phaeogenes nigridens 
Wesmael (Goidanich, 1931), Diadromus collaris (Gravenhorst) (Given, 
1944), Proscus suspicax (Wesmael) (Beirne, 1941), Herpestomus 
brunnicornis (Gravenhorst) (Beirne, 1948). 

16 Except Pseudamblyteles (Ichneumonini), where there is no line of setae beneath each sclerotized band 
as in the Platylabini. 
17 Except Melanichneumon (Ichneumonini), where the numerous setae on the prelabium distinguish 


this genus from Probolus. 
18 Of Given (1944). 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 449 


Tribe Platylabini 


FIGuRE 29 


Platylabus clarus (Cresson) was examined. 

The pair of lightly sclerotized plates situated on the clypeolabra! 
region above the setae and sensilla distinguish this ichneumonine 
from all others examined except Pseudamblyteles. In Pseudamblyteles 
there is no line of setae beneath and following the curve of each 
sclerotized plate as in Platylabus. 


Tribe Listrodromini 
Ficure 30 


Members of this tribe prefer Lycaenidae as hosts. 
The broad mandibles with their very short blades are distinctive. 


LARVAL KrEry 


1. Each maxillary and labial palp with three sensilla, the middle one raised as a 


PTOTECLION ye erass temrer A Gece ends a bs wry dete +a) bop Ren eh oe eA SOMAS 
Each maxillary and labial palp with five sensilla, but none raised as a pro- 
FECEION Ty se) ak eae coset sce Ghee Bubehose Ne ns cs ef af + Luastredronms 


Figure 29.—Ichneumoninae:Platylabini, head sclerites: Platylabus clarus (Cresson) 
(Antenna not visible on preparation; 2, spiracle; 3, skin.) 


450 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


Ficure 30.—Ichneumoninae:Listrodromini, head sclerites: a, Anisobas sp. (antenna not 
visible on preparation; 2, spiracle; 3, skin); B, Listrodromus nycthemerus (Gravenhorst) 
(other structures not visible on preparation). 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 451 


The following were examined: Anisobas sp. (CDA) (fig. 30a), 
Listrodromus nycthemerus (Gravenhorst) (BM) (fig. 308). 


Tribe Eurylabini 
Figure 31 


Probolus sp. (CDA) was examined. 

The absence of a sharp line of demarcation between each pleuro- 
stoma and the sclerotized area lateral to it distinguish this ichneu- 
monine from all others examined except Melanichneumon, which is 
distinguished by the numerous setae on the prelabial region. 


K O-1mm | \ 


Ficure 31.—Ichneumoninae:Eurylabini, head sclerites: Probolus sp. (antenna not visible 
on preparation; 2, spiracle; 3, skin.) 
Tribe Ichneumonini 
Ficures 32-35 


In the following key reference is made to setae and sensilla, but 
there is no distinction between a small seta and a sensillum. 


LARVAL KEY 


1. Epistoma broad and lightly sclerotized. .......2.2.. . Carinodes 
Epistoma narrow and well sclerotized . SIN Es io PEE eee 
2. Some eighteen setae present on prelabium. . .. . . . .Melanichneumon 


No more than six setae present on prelabium........2...2.2..8 


452 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Frcure 32.—Ichneumoninae:Ichneumonini, head sclerites: a, Melanichneumon brevicinctor 
(Say); B, Cratichnewmon unifasciatorius (Say). (Antenna not visible on preparation; 
2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 453 


Figure 33.—Ichneumoninae:Ichneumonini, head sclerites: a, Aoplus velox (Cresson) 
antenna not visible on preparation; B, Chasmias scelestus (Cresson). (1, antenna; 2, 
spiracle; 3, skin.) 


454 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 34.—Ichneumoninae:Ichneumonini, head sclerites: a, Pseudamblyteles nuncius 
(Cresson); B, Amblyteles inconstans (Cresson). (Antenna not visible on preparation; 


2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 455 


ee 
| O:-i mm 


Ficure 35.—Ichneumoninae:Ichneumonini, head sclerites: a, Ichnewmon ambulatorius 
Fabricius; p, Carinodes havenensis (Cameron). (1, antenna; 2, spiracle; 3, skin.) 
504675—59 5 


456 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


3. Clypeolabral region with ten setae as in a line and two groups each of 


two sensilla beneath this line. .. . ... . . .Cratichneumon 
Setae and sensilla SES in two groups on eclypeolabrum and not in a 
tines. i 4 

4. Clypeolabral eae eoupedia as ana ae 35a). on ore lightly aclernrived mieees 
shaped as shown (fig. 354)... . . . . . .Ichneumon 
Setae and sensilla not grouped as shan (fig. 354) on n lates shaped as shown 
(fig 354) ea <a alerted 


5. Raised areas on ei neniananen containine eCneiliay narrow end airaicholl Aoplus 
Raised areas on clypeolabrum containing setae or sensilla more broad and 
not straight... . . . Sate A eo 

6. Each raised area Rontaintiie censiin on Sieiyeeniibar aenies idoese ity: 
Amblyteles 
Each raised area containing setae or sensilla on clypeolabrum not curved 
dorsally . 
7. Each raised area on Git pcamsbren with a foney. aclerorived! doreal bar and 
containing six sensilla. . ... Soke Wi or as Pseudamblyteles 
Each raised area on clypeolabrum aihout ecleronzed dorsal bar and con- 
taiming SevientSetaer, s 2 «fee ae «ia Sa te) % Gee oe OM eIaa eS 


The following have been examined: Melanichneumon brevicinctor 
(Say) (fig. 324), MZ. rubicundus (Cresson), Cratichneumon duplicatus 
(Say), C. unifasciatorius (Say) (fig. 328), Aoplus velox (Cresson) 
(fig. 334), Chasmias scelestus (Cresson) (fig. 338), Pseudamblyteles 
animosus rubellus (Cresson), P. nuncius (Cresson) (fig. 34a), 
subfuscus (Cresson), Amblyteles inconstans (Cresson) (fig. 348), A 
subrufus (Cresson), Ichneumon ambulatorius Fabricius (fig. 35a), I 
laetus Brullé, I. rufiventrus incertus Cresson, and Carinodes havanensis 
(Cameron) (fig. 358). 

The following are figured in the literature: Melanichneumon rubi- 
cundus (Cresson) (Wishart, 1949), Aoplus pictus (Gravenhorst) 
(Beirne, 1941), Amblyteles armatorius (Foerster) (Beirne, 1941), 
Ichneumon suspiciosus Wesmael (Cameron, 1950, 1951). 


Tribe Protichneumonini 
FIGURE 364 


Coelichneumon pepticus (Cresson) (fig. 364) and Protichnewmon sp. 
(GCV) were examined. 

There are no clear characters differentiating this tribe from the 
Ichneumonini. The head parts are heavily sclerotized and there is a 
large sclerotized area of the cranium lateral to each pleurostoma. In 
Coelichneumon the ocular line (shown by the antenna in fig. 36) is 
well sclerotized and conspicuous. It is also conspicuous in Protich- 
neumon. The specimen of Protichnewmon was not figured since the 
labral setae and sensilla were not clearly visible on the preparation; 


however, their arrangement appeared to be similar to those of Coelich- 
neumon. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 457 


O:imm 


Ficure 36.—Ichneumoninae, head sclerites: a, Coelichneumon pepticus (Cresson) (Protich- 
neumonini); B, Trogus pennator (Fabricius) (Trogini), antenna not visible on preparation. 
(1, antenna and ocular line; 2, spiracle; 3, skin.) 


458 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


LarvaL Kry 


1. Three prominent setae situated beneath each labial palp.. . Coelichneumon 
Six prominent setae situated beneath each labial palp.. . . Protichneumon 


Tribe Trogini 
FIGURE 36B 


Species of Trogus and Macrojoppa are parasites of Papilionidae. 
The rest of the genera attack Sphingidae. 

Trogus pennator (Fabricius) (fig. 368) was examined. ‘This species 
differs from other Ichneumonines examined in that the hypostoma is 
sharply bent. 

Subfamily Mesochorinae 


FIGURE 37 


All members of this subfamily are believed to be endoparasites and 
secondary parasties. 

Mesochorus discitergus (Say) (fig. 37) and Mesochorus sp. were 
examined. 

The larval characters of the specimens examined indicate an 
isolated group. Those head sclerites present are lightly sclerotized ; 
the epistoma is unsclerotized; the pleurostoma is present; only the 
median part of the hypostoma is sclerotized in Mesochorus, but the 
entire hypostoma is sclerotized in Astiphromma strenuum (Holmgren) 


i 


5 ee ee 3 
O:f mm 
Figure 37.—Mesochorinae, head sclerites: Mesochorus discitergus (Say). (Antenna not 
visible on preparation; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT A459 


(Beirne, 1941); the hypostomal spur is very lightly sclerotized; the 
stipital sclerite is slender and its lateral end has a small enlargement 
bearing a sensillum; the labial sclerite is slender; the silk press is not 
sclerotized; each maxillary and labial palp bears two sensilla, one large 
and one small; there is no prelabial sclerite and no labral sclerite; 
two small sensilla are present on the labrum, but labral, maxillary, 
and prelabial setae appear to be absent; the form of the mandible is 
distinctive, the base having only the circumference sclerotized and 
with the blade long, slender, and slightly curved; the width of the 
base of the mandible is approximately equal to two-thirds the length of 
the blade; thespiracle is relatively small and thelength of the thin-walled 
closing apparatus, which adjoins the atrium, is approximately equal 
to twice the depth of the atrium; the skin bears small projections but 


setae could not be seen. 
LarvaL Key 


1. Hypostoma with only median part sclerotized ........ Mesochorus 
Hypostoma with entire length sclerotized. . ....... Astiphromma !? 


The following are figured in the literature: Mesochorus fulgurans 
Curtis (Beirne, 1941), AZ. vitticollis Holmgren (Beirne, 1943). 

The Mesochorinae follow the Anomalinae in the classification by 
Walkley (1958). The larval characters of the Mesochorinae are very 
different from those of the Anomalinae and in the present study the 
Mesochorinae have been placed near the Lissonotinae. ‘The rounded 
labial sclerite of the Mesochorinae resembles that of the Lissonotinae. 
The mandible, however, is distinctive in form and differs from that 
of all other Ichneumonids. On larval characters the Mesochorinae 
are best regarded as an isolated group. 


Subfamily Lissonotinae 
Figures 38-40 


Members of this subfamily are internal parasites of caterpillars. 

The characters show relationships to those of the Ophioninae and 
Mesoleiinae. The epistoma is not sclerotized dorsally except in 
Hzetastes; the pleurostoma is lightly sclerotized to well sclerotized; 
the hypostoma is long in the Glyptini and Banchini, but in the 
Lissonotini it is short and does not extend laterally beyond the lateral 
end of the stipital sclerite; the hypostomal spur and stipital sclerite 
are well developed; the labial sclerite, which is relatively large, is 
basically round to oval and the ventral part is often lightly sclerotized 
or unsclerotized; the silk press is large; a prelabial sclerite is usually 
present and two sensilla are situated on this sclerite; each maxillary 


19 Of Beirne (1941). 


460 PROCEEDINGS OF THE NATIONAL MUSEUM vOL. 110 


and labial palp contains one large sensillum and one to three smaller 
sensilla; the labral sclerite is absent; the mandibles are relatively small 
and consist of a large base and a small slender blade; this blade may 
be toothless or may have small blunt teeth on its dorsal and ventral 
surfaces; in some genera a lightly sclerotized band is present connect- 
ing the posterior pleurostomal processes across the dorsal surface of 
the food meatus; the antenna is disc-shaped; the closing apparatus 
of the spiracle adjoins the atrium except in Glypta; the skin has small 
setae and small conical projections but no spines. 


LarvaL Key 


1. Hypostoma short and not extending laterally beyond lateral end of stipital 
sclerite! ig |. n. . . . Lissonotini 
Hypostoma long and extendas iter beyond end 3 atipital sclerite . . 2 

2. Epistoma complete, or, if not, then with more than 10 setae present on pre- 
IGNoyuboany i 4.7% .... . . Banchini 
Epistoma inom alete: ese fan 10 Seimei on prolate bo. 4 J Glyptina 


Tribe Glyptini 
FIGURE 38A 


Glypta sp. was examined. 

With its rounded labial sclerite which is incomplete ventrally and 
its lightly sclerotized prelabial sclerite, Glypta resembles the Lis- 
sonotini. The long hypostoma resembles that of the Banchini. As in 
some Banchini and some Lissonotini a sclerotized bar connecting the 
posterior pleurostomal processes is present. The mandibles are 
distinctive with their slender toothless blades and, unlike other 
Lissonotinae, the closing apparatus of the spiracle does not adjoin 
the atrium. 

Glypta fumiferanae (Viereck) is figured by Brown (1947), G. 
haesitator Gravenhorst by Cameron (1938), and @. parvicaudata 
Bridgman by Beirne (1941). 


Tribe Lissonotini 
FIGuRES 388,c, 39 


Members of this tribe are parasites of caterpillars in burrows and 
leaf rolls. 
LarvaL Kuy 


1. Labial sclerite with ventral part unsclerotized or not as well sclerotized as 


lateral parts... . 1, ME Si ee 
Labial sclerite with rontrall pare as ell Ger Guede as tera pan DR Sd 
2.’ Mandible‘ without teeth)... sO.) ci.le Gis & Ncuiue Ov. OnRbdssemeia 


Mandible: with'teeth © 5 . 3). .°.'. wos "se ae ee ee ee er 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 461 


a pan ly A Ae ha 


= A 
ms ~ A NA 


SS 3 

O-1mm 

Ficure 38.—Lissonotinae, head sclerites: a, Glypta, sp. (Glyptini); B, Lampronota sp. nr. 
sesiovora (Rohwer) (Lissonotini); c, Leptobatopsis sp. nr. lepidus (Cameron) (Lissonotini). 
(Antenna not visible on preparation; 2, spiracle; 3, skin.) 


462 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


a 8 Arn 
a a 


| Pe & 


Figure 39.—Lissonotinae:Lissonotini, head sclerites: a, Lissonota’ brunnea (Cresson); 
s, Pimplopterus rubricus (Cresson); c, Asphragis mirabilis (Cresson). (Antenna not 
visible on preparation; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 463 


3s Dorsalvends of Jabialiscleniterbroad astm csieasikes - ee Asphragis 
Dorsal ends of labial sclerite narrow .......2.+.4.-. Leptebatopsis 
4. Hypostoma not extending laterally beyond hypostomal spur . Lampronota 


Hypostoma with short lateral bar beyond hypostomal spur . Pimplopterus 


The species of Lampronota figured is peculiar in that there is a 
lightly sclerotized band running along the ventral edge of the labrum. 
This band is distinct from the band connecting the posterior pleuro- 
stomal processes across the dorsal surface of the food meatus in some 
genera. The functions of these bands are not obvious. Beirne (1941) 
does not figure a labral band in L. catenator (Panzer) although he does 
figure a band between the posterior pleurostomal processes. 

The following were examined: Lampronota sp. nr. sesiovora (Rohwer) 
(fig. 388), Leptobatopsis sp. nr. lepidus (Cameron) (fig. 38c), Lissonota 
brunnea (Cresson) (fig. 394), Pimplopterus rubricus (Cresson) (fig. 
39B), Asphragis mirabilis (Cresson) (fig. 39c). 

Lampronota catenator (Panzer) and Syzeuctus maculatorius (Fab- 
ricius) are figured by Beirne (1941). S. maculatorius is similar to 
Asphragis although teeth do not appear to be present on the mandible. 


Tribe Banchini 


Ficure 40 


LaRvAL Kry 


NemE PISGOMMA OLESEN UG, w eee ccc ae aes, hat ede Ral fia, eae bins Pee ee Exetastes 
Espistoma aAbsemb pence! a) tors i seceen nel Syed een a cola, Galler ce) os prem e-er es 2 

2. Ventral part of prelabial sclerite more lightly sclerotized than dorsal parts. 
Ceratogastra 


Ventral part of prelabial sclerite as well sclerotized as laterial parts . Banchus 


Banchus and Ceratogastra are very similar. The T-shaped prelabial 
sclerite resembles that of many Ophioninae. Lzetastes differs from 
other Lissonotinae in having a complete epistoma, but the remaining 
characters, such as the shape of the mandible, are typical of the sub- 
family. Apart from the epistoma, Hxetastes differs from the other two 
genera of the Banchini examined in that there are teeth on the man- 
dibles. However, the structure of the spiracle is basically similar in 
the three genera, the closing apparatus being as long as two or three 
times the depth of the atrium. In the Lissonotini the closing apparatus 
is approximately of the same length as the atrium. 

The following were examined: Fxetastes bifenestratus Cushman 
(fig. 40a), H. illinoiensis (Walsh), Ceratogastra ornata ornata (Say) 
(fig. 408), Banchus femoralis Thomson (fig. 40c), B. flavescens Cresson. 

The following are figured in the literature: Hzxetastes cunctrpes 
(Ratzeburg) (Beirne, 1941), Hxetastes sp. (Snodgrass, 1935), Banchus 
femoralis Thomson (Beirne, 1941). 


464 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ma A brn A Le 


A TR Ay Brel MA eae ee 
~ x 


Ficure 40.—Lissonotinae:Banchini, head sclerites: a, Exetastes bifenestratus Cushman; 
B, Ceratogastra ornata ornata (Say); c, Banchus femoralis Thomson. (1, antenna; 2, 
spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 465 


Subfamily Mesoleiinae 
Ficgures 41-44 


Members of this subfamily are internal parasites of Tenthredi- 
noidea.”” 

The dorsal part of the epistoma is unsclerotized or lightly sclero- 
tized; the pleurostoma is usually lightly sclerotized: the hypostoma 
and hypostomal spur are well sclerotized, the hypostomal spur being 
relatively longer than in most Ophioninae; the stipital sclerite is well 
developed and a lightly sclerotized cardo is sometimes present; the 
labial sclerite is present and may be incomplete ventrally; in some 
genera the dorsal ends of the labial sclerite bear small bubble-like 
projections; a prelabial sclerite is present and two sensilla are usually 
present on the dorsal part of this sclerite; the maxillary and labial 
palps are relatively large and each bears two to three sensilla; the silk 
press is conspicuous; the labral sclerite is absent, except in Opheltes 
and Protarchus; there are two main types of mandible, one having a 
rounded and lightly sclerotized base with a relatively straight blade 
which may be short, the other having a rather square base and a 
curved blade; the antenna is disc-shaped; with the exception of Huceros 
and Opheltes where the closing apparatus of the spiracle is small, the 
spiracle has a thin-walled closing apparatus with a length which is 
two to three times the depth of the atrium; the skin bears very small 
setae and small projections which in Lophyroplectus are sufficiently 
sclerotized to be described as spines. 

The Mesoleiinae have many features resembling the Ophioninae. 
In the key to the subfamilies it was found necessary to separate the 
Ophioninae into tribes in order to key out the Mesoleiinae from this 
subfamily. The larval characters of the Mesoleiinae do, however, 
indicate a fairly distinct group. The type of mandible of Rhorus, 
with its rounded base and very small blade, is distinctive. The 
mandible of Hypsantyr is basically similar although the blade is 
relatively larger. The mandible of Hypsantyz, in turn, resembles that 
of other Mesoleiini and that of the Euryproctini. The mandible of 
Scolobates and Euceros is similar to that of most of the Ophioninae 
in that there is a rather square base and a curved blade. But the shape 
of the labial sclerite of Scolobates connects this genus to the remaining 
Mesoleiinae. The labial sclerite of Huceros, with its plate-like expan- 
sion of the ventral part, resembles that of some genera in the Ophio- 
ninae, Campoplegini. In Opheltes (Mesoleiinae, Mesoleiini), however, 
there is a smaller plate-like expansion of the ventral part of the labial 
sclerite and this may connect to the condition in Huceros. The spiracle 
of Huceros, with its relatively small closing apparatus, resembles that 


20 Wuceros frigidus Cresson has been recorded as an internal’parasite of Lepidoptera. 


466 PROCEEDINGS OF THE NATIONAL MUSEUM You. 110 


of the Ophioninae, Campoplegini, and, with the exception of Opheltes, 
differs from that of the remaining Mesoleiinae. 

It has not been found possible to separate the Mesoleiini and Eury- 
proctini on larval characters. Lamachus and Hypsantyx with their 
incomplete labial sclerites, the dorsal ends of which have bubble-like 
projections, resemble the Euryproctini rather than Opheltes, Mesoleius 
and Lophyroplectus. 

Townes and Townes (1951) place Ophelies in a subtribe Pariiatane 
and Mesoleius and Lamachus in a subtribe Mesoleiina of the Mesoleiini. 
The larval characters do not appear to support this grouping. 


LARVAL Key 


1. Mandible with rounded and very lightly sclerotized base and very small 


sclerotized blade of length less than radiusof base ..... .. Pionini 
Mandible \net, of his; form. 04 esac! feo ype aed ~~ latteliee 2 

2. Labial sclerite with ventral part expanded into plate constituting almost half 
length of sclerite. . . . Whey ci gol eu le Weep mete) Ne aoe fo 1 ey CE EE SEED 
Labial sclerite not of this fee bao na ied oo 

3. A lightly sclerotized plate is present across Seer cand the labral setae are 
situated: on this) plate; .2 =: 26 0ers) -1hio.t -..+ . « Scolobatini 

A lightly sclerotized plate not een on labrum ... Pe eo re: 


4. Labial sclerite with ventral part well sclerotized or, if noe Gree as well 
sclerotized as lateral parts. 
Ophelies, Mesoleius, and Lophyroplectus of Mesoleiini 
Labial sclerite with ventral part unsclerotized or not as well sclerotized as 
lateral parts. 
Euryproctini, and Lamachus and Hypsantyz of Mesoleiini 


Tribe Scolobatini 


Figure 414 


Members of this tribe are parasitic on argid sawflies. 

Scolobates auriculatus (Fabricus) (CDA) was examined. The 
mandibles of this species resemble those of the Ophioninae, 
Campoplegini, but the remaining characters are similar to those of 
many Mesoleiinae. 

Distinctive structures are the lightly sclerotized bar across the 
labrum and a lightly sclerotized plate which appears to be present on 
the dorsal surface of the food meatus. This may be the suspensorium 
of the hypopharynx (see Short, 1952, figs. 6-8) but, as seen in a cast 
skin, it appears to be present on the dorsal surface of the food meatus 
and therefore distinct from the suspensorium. Living material 
could not be obtained for sections to decide this point. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 467 


TENA LA eb 
ied 


4 3 
O-lmm 
TA Aste an ER A 
geste 


-—_+| 
O-imrn 
Figure 41.—Mesoleiinae, head sclerites: a, Scolobates auriculatus (Fabricius) (Scolobatini); 


B, Euceros thoracicus Cresson (Euceratini); c, Rhorus clapini (Provancher) (Pionini); 
antenna not visible on preparation. (1, antenna; 2, spiracle; 3, skin.) 


468 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Tribe Euceratini 
Ficure3 418 


Species of Huceros have been recorded mainly from Tenthredinoidea, 
but E. fridigus Cresson has been recorded from Lepidoptera. 

Euceros thoracicus Cresson was examined. This species shows a 
strong resemblance to many Ophioninae, Campoplegini, but the 
mandibles also resemble those of the Scolobatini and the expansion 
of the ventral part of the labial sclerite is also seen, although to a 
lesser extent, in Opheltes. The small closing apparatus of the spiracle 
is also seen in Opheltes. 

Tribe Pionini 
Figure 41c 

Rhorus clapini (Provancher) was examined. 

This is a somewhat isolated form, but the labial sclerite, although 
very slender, does resemble that of most Mesoleiinae in being in- 
complete ventrally. The type of mandible is also found, in a modified 
form, in the Euryproctini and Mesoleiini. 

Rhorus mesoanthus (Gravenhorst) is figured by Beirne (1941). 

Tribe Mesoleiini 


FIGuRES 42, 43 


LARVAL KEY 


1. With labial sclerite complete ventrally... . 3: gee 
With labial sclerite very lightly sclerotized or neeelerotised ence : oa fee 

2. Labral sclerite present*! . .\.. =... ....+ 5 . . = 6 =»  Opheltes 
Protarchus 2 

Labralsclerite‘absent. <<. Nee we a ee a 
SINixtseLab On DrelabiUny, sion steko eres . . Mesoleius 
Four setae on prelabium... . A tk gee ‘Lophyeonleeues 
4. Posterior end of hypostoma with eal saan .... .. . Hypsantyx 
Posterior end of hypostoma without small expansion. . . . . . Lamachus 


The broad ventral end of the hypostomal spur figured in Lamachus 
virginianus (Rohwer) appears to be present only in this species. 
In other species of Lamachus the ventral end of the hypostomal spur 
is slender. In most species of Lamachus examined the dorsal ends of 
the labial sclerite had pronounced bubble-shaped projections similar 
to those of Hypsantyx and the Euryproctini. 

The following have been examined: Opheltes glaucopterus flavi- 
pennis (Provancher) (fig. 424), Mesoleius tarsilas (Cresson), 
tenthredinis Morley (fig. 428), Lamachus albopictus Cushman, L. 


21 Opheltes differs from Profarchus in that the ventral part of the labial sclerite is less pointed. 
22 Of Beirne (1941). 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 469 


An Anon yrs 


. MC pe ae 
O:imm i) 


Ficure 42.—Mesoleiinae:Mesoleiini, head sclerites: a, Ophelies glaucopterus flavipennis 
(Provancher); 8, Mesoleius tenthredinis Morley, antenna not visible on preparation. 
(1, antenna; 2, spiracle; 3, skin.) 


470 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 43.—Mesoleiinae:Mesoleiini, head sclerites: a, Lamachus virginianus (Rohwer), 
3, Hypsantyx lituratorius (Linnaeus); c, Lophyroplectus nipponensis Cushman. (1 
antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 471 


coniortionis Davis, L. eques (Hartig), LZ. lophyri (Ashmead), L. 
ruficoxalis (Cushman), L. tsugae Cushman, L. virginianus (Rohwer) 
(fig. 43a), Hypsantyzx lituratorius (Linnaeus) (fig. 488), Lophyroplectus 
mipponensis Cushman (fig. 43c). 

The following are figured in the literature: Mesoleius tenthredinis 
Morley (Beirne, 1941), Protarchus testatorius (Thunberg) (Beirne, 
1941), Lamachus sp. (Morris, Cameron, and Jepson, 1937), Hypsantyx 
lituratorwus (Linnaeus) (Beirne, 1941), Lophyroplectus luteator (Thun- 
berg) (Beirne, 1941). Beirne (1941, pp. 151, 153) figures lateral 
projections on the labial sclerite in the region of the stipital sclerite 
in Mesoleius and Lophyroplectus. These structures were not seen. 
It is probable that lobes on the median end of each stipital sclerite 
were mistaken for lobes on the labial sclerite. 


Tribe Euryprectini 
Figure 44 


Larva Key 


1. Ventral part of prelabial sclerite absent. . ......2.2.42.2.. Synomelix 
Ventral part of prelabial sclerite lightly sclerotized... ....... 2 
2. Dhree sensilla-on, éach labial-palp-.~.-. 0 wy. 2) en Pus ee 8 Ipoctoninus 
Mwosensilla on each labial palp ioc 242 fal fe CO. ee Polyterus 


These three genera are very similar. 
The following were examined: Synomeliz sp. (fig. 444), Polyterus 
olympiae (Ashmead) (fig. 448), [poctoninus striatus (Davis) (fig. 44c). 


Subfamily Collyriinae 
Figure 45a 


Collyria calcitrator (Gravenhorst) was examined. This is a very 
isolated species with the larval characters showing no relationship 
to those of any other ichneumonid. It is endoparasitic in the sawfly 
Cephus. 

The lateral parts of the epistoma are very lightly sclerotized and 
broad, forming plates on the lateral parts of the clypeus; each pleuro- 
stoma is lightly sclerotized and bears about five sensilla; there are no 
other head sclerites: the maxillary and labial palps are represented by 
small sensilla; there is no sclerotized silk press; about 12 sensilla are 
present on the labrum; the mandibles are unsclerotized and are 
without teeth; they can be detected in specimens cleared in Faure’s 
fluid, but must be stained in order to be seen clearly; the antenna is 
disc-shaped; the spiracle has a long, thin-walled closing apparatus 
with a length which is five to six times the depth of the atrium; the 
skin is very smooth with small projections and small setae. 


504675—-59 ——6 


AG PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


O-imm 


Ficure 44.—Mesoleiinae:Euryproctini, head sclerites: a, Synomelix sp.; B, Polyterus 
olympiae (Ashmead); c, Ipoctoninus striatus (Davis). (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—-SHORT 473 


— fA 4 OAS Sane 
ot OR ~ & 


g| 


Figure 45.—Head sclerites: a, Collyria calcitrator (Gravenhorst) (Collyriinae) ;B, Ortho pelma 
mediator (Thunberg) (Orthopelmatinae). (1, antenna; 2, spiracle; 3, skin.) 


A74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


C. calcitrator is figured by Salt (1931) and a large sclerotized plate 
is shown underlying the labial area. This plate is the suspensorium 
of the hypopharynx (see Short, 1952), and is figured here, although 
in a different position (fig. 454). The suspensorium is an internal 
skeletal structure forming part of the ventral wall of the cibarial part 
of the stomodaeum, and the position in which it is seen depends on 
the position into which it is forced during mounting. 


Subfamily Orthopelmatinae 
Figure 458 


Orthopelma mediator (Thunberg) was examined. The members of 
this genus are endoparasitic in the cynipoids Diastrophus on Rubus 
and Diplolepis on Rosa. 

The larval characters resemble those of the Ichneumoninae, although 
the form of the mandible is different, the hypostoma is more lightly 
sclerotized, and dise-shaped labial palps are absent. 

The epistoma and pleurostoma are moderately sclerotized and very 
lightly sclerotized areas are present dorsal to the epistoma and dorso- 
lateral to the pleurostoma; the hypostoma is very lightly sclerotized; 
the lateral parts of the labial sclerite are very faintly sclerotized and 
folds along the median part of the ventral edge of each maxilla may 
appear as faintly sclerotized bands; the remaining head sclerites are 
absent; the maxillary palps are represented by flat discs but the 
labial palps are represented by setae; a lightly sclerotized silk press is 
visible; the mandible is relatively large and well sclerotized and has 
a curved blade containing a deep groove on the inner wall of which is 
situated a prominent tooth; the anterior tentorial pits are prominent; 
the antenna is disc-shaped; the spiracle is relatively very small with 
an oval atrium and the closing apparatus, which adjoins the atrium, 
is equal in length to twice the depth of the atrium; the skin is smooth 
with small setae and no projections. 

Orthopelma mediator as figured by Beirne (1941) shows a toothless 
mandible, disc-shaped labial palps, and a well-sclerotized hypostoma. 
These structures have not been seen in any specimen of O. mediator 
examined in the present study. 


Subfamily Plectiscinae 
Fiaure 46 
This is a very isolated subfamily and little is known about its 
biology. The usual hosts are believed to be Fungivoridae. The 
larval characters suggest that the group is endoparasitic. 


Megastylus sp. (fig. 46a) and Plectiscus varius (Haliday) (GCV) 
(fig. 468) were examined. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 475 


O-imm 


Ficure 46.—Plectiscinae, head sclerites: a, Megastylus sp., antenna not visible on 
preparation; B, Plectiscus varius (Haliday). (1, antenna; 2, spiracle; 3, skin.) 


476 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


The epistoma is unsclerotized; the pleurostoma is lightly sclerotized ; 
the hypostoma is short and lightly or very lightly sclerotized; the 
hypostomal spur and stipital sclerite are fused, the stipital sclerite 
not extending laterally beyond its point of meeting with the hypo- 
stomal spur; the labial sclerite is somewhat square in shape and the 
ventral part is relatively broad; each maxillary and labial palp is 
large and each bears two sensilla, one of which is relatively large; 
the silk press is lightly sclerotized; the labral sclerite is absent; the 
mandibles are relatively small and are triangular in shape; they are 
very lightly sclerotized and are without teeth; the antenna is disc- 
shaped; in Megastylus the closing apparatus of the spiracle is slightly 
smaller than the atrium and adjoins the atrium; in Plectiscus the 
closing apparatus of the spiracle is longer than the depth of the 
atrium and is separated from the atrium by a length of trachea equal 
to the length of the closing apparatus; the skin bears small spines and 
small projections. 

Larval Kry 
1. Hypostomal spur and stipital sclerite as well sclerotized as labial sclerite. 
Megastylus 


Hypostomal spur and stipital sclerite not as well sclerotized as labial sclerite. 
Plectiscus 


Subfamily Diplazoninae 
Ficure 47 


Members of this subfamily are endoparasitic in Syrphidae. They 
oviposit into the egg or young larva of the host and emerge from the 
puparium. The subfamily is an isolated group. 

The following were examined: Diplazon laetatorius (Fabricius) 
(fig. 474), D. orbitalis (Cresson), D. seutellaris (Cresson), Zootrephus 
rufiwentrus (Gravenhorst) (fig. 47B) (the head parts other than the 
labium were damaged in this specimen), Syrphoctonus agilis (Cresson), 
and S. maculifrons (Cresson) (fig. 47c¢). 

The epistoma is unsclerotized; the pleurostoma is lightly sclerotized; 
a small part of the hypostoma is very lightly sclerotized as is a small 
part of the hypostomal spur; the stipital sclerite is absent; the labial 
sclerite is well sclerotized and of characteristic shape with a ventral 
spine-like projection; the maxillary and labial palps are present, but 
setae appear to be absent from the maxillae and labium; there is 
a faintly sclerotized area at the dorsal end of the labial sclerite which 
may represent the silk press; the labral sclerite is absent; groups of 
sensilla are present on raised areas of the labrum; the mandibles are 
usually very lightly sclerotized although the blade is lightly sclerotized 
in some species; the antenna was not seen in any cast skin and is 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 477 


O-{mm 


++ 
O-{mm 3 


Ficure 47.—Diplazoninae, head sclerites: a, Diplazon laetatorius (Fabricius); B, Zootrephus 
rufiventrus (Gravenhorst); c, Syrphoctonus maculifrons (Cresson). (Antenna not visible 
on preparation; 2, spiracle; 3, skin.) 


478 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


presumably disc-shaped with an unsclerotized circumference; the 
spiracle has an oval atrium and the closing apparatus, which has thin 
walls, adjoins the atrium in some genera but not in others; the closing 
apparatus has a length equal to five to seven times the depth of 
the atrium; the skin is very smooth and appears to lack setae and 
projections. 


LarvauL Key 


1. Lateral parts of labial sclerite produced into relatively sharp angles and not 


POUT CCAS ceteris hc oy ee ee Or nae eS re eae amar Syrphoctonus 
Lateral parts of labial sclerite less sharp and morerounded....... 2 

2 sLabial palpsilobed's (ti) iis Skee ee Be ee Promethus 2 
Labialipalps:oval "30s: ashe ge eP SE |e. AL el ence VR oe) 3 

8. Length of labial spine approximately equal to depth of main ventral part of 
labial:sclenite ries tees: Meech 1p enn ete ce ees, Bee a Diplazon 
Length of labial spine shorter than depth of main ventral part of labial sclerite. 
Zootrephus 


The following are figured in the literature: Diplazon fissorius 
(Gravenhorst) (Schneider, 1950), D. Jlaetatorius (Fabricius) (Scott, 
1939), D. tetragonus (Thunberg) (Beirne, 1941), Promethus monticola 
(Vollenhoven) (Scott, 1939), Syrphoctonus signatus (Gravenhorst) 
(Scott, 1939), S. tarsatorius (Panzer) (Scott, 1939; Beirne, 1941). 

The figures of Scott are particularly good. Schneider figures 
a labral sclerite in Diplazon fissorius (Gravenhorst). This has not 
been seen in any specimen of Diplazon examined in the present study 
and neither Scott nor Beirne figure this structure. 


Subfamily Metopiinae 
Fiaures 48, 49 


Members of this subfamily are endoparasitic in Lepidoptera. The 
parasites emerge from the pupae of the hosts. 

The epistoma is well sclerotized except in Triclistus; the pleurostoma. . 
and hypostoma are well sclerotized; the hypostomal spur is absent 
and the stipital sclerite extends dorsally to meet the hypostoma; the 
labial sclerite is incomplete ventrally; on each maxillary and labial 
palp there is one large sensillum and one or several smaller sensilla; 
the silk press is well sclerotized; there is a lightly sclerotized plate 
ventral to the press containing two sensilla; this plate may represent 
the dorsal part of the prelabial sclerite but is more probably the 
ventral part of the silk press; the labral sclerite is absent; the mandibles 
are relatively large and do not have teeth on the blade; the antenna is 
disc-shaped; the atrium of the spiracle is shallow; the closing apparatus 
of the spiracle is thin-walled and with a length approximately equal 


2 Of Scott (1939). 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 479 


on 
O-f rar 


Ficure 48.—Metopiinae, head sclerites: a, Metopius micratorius (Fabricius); B, Triclistus 
:< curvator (Fabricius); c, Exochus scitulus Provancher, antenna not visible on preparation. 


a 


2 (1, antenna; 2, spiracle; 3, skin.) 


480 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


o 0 


— Ae OO yrs — 
LOD AR OSS 


O:-imm 


Ficurs 49.—Metopiinae, head sclerites: Chorinaeus sp. (antenna not visible on preparation; 
2, spiracle; 3, skin.) 


to the depth of the atrium, except in Metopius where the closing ap- 
paratus is thick-walled and with a length three to four times the 
depth of the atrium; the skin is smooth with small spines and small 
projections. 

The head sclerites of the subfamily show a strong resemblance to 
those of the Anomalinae. They also resemble the Pimplini and the 
Ichneumoninae in having large toothless mandibles and a well- 
sclerotized epistoma and pleurostoma. These resemblances are pos- 
sibly associated with similar habits since all these groups emerge from 
the pupae of Lepidoptera. 


LarvaL Kry 


1; Epistoma incomplete dorsally =... . 2 ..'..5 0 7 a+. 2. .  Drichstus 
Epistoma complete dorsally. ... . SOS RERENG 
2. A lightly sclerotized plate is present tateral iy eae plearomenga and hypos- 
COMMS waiter Were rae, tS: 
A lightly seleronized mint is Het mreeeat Tateral ay ancl pleuroetornn and 
hypostoma ... .... . Chorinaeus 

3. A lightly sclerotized bd is present conticatite the posteroe pleurostomal pro- 
cesses across the dorsal surface of the food meatus. . . . . . . Exochus 

A lightly sclerotized bar connecting the posterior pleurostomal processes not 
PICSENG sys ie uae es) cet we ek be as els Ee Ce Le LODIEES 


The following were examined: Metopius micratorius (Fabricius) 
(fig. 484), Triclistus curvator (Fabricius) (fig. 488), Ezochus albifrons 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT AS] 


Cresson, EL. scitulus Provancher (fig. 48c), Chorinaeus ‘sp. (GCV) 
(fig. 49). 

Beirne (1941) figures Metopius anxzius Wesmael and Triclistus poda- 
gricus (Gravenhorst). 


Subfamily Ophioninae 


This subfamily has many characters resembling those of the Lissono- 
tinae and Mesoleiinae. As stated in the introductory keys to sub- 
families, it was found necessary to break up the Ophioninae into 
tribes in order to differentiate these from the two related subfamilies. 


Tribe Campoplegini 
Fieures 50-56 


This tribe includes many important endoparasites of economic 
pests. Most of the species attack lepidopterous larvae. Exceptions 
include those of the genus Olesicampe parasitic on Tenthredinidae and 
Diprionidae, of Biolysia and Bathyplectes parasitic on Curculionidae, 
and of Pyracmon parasitic on wood-boring Coleoptera. 

The dorsal part of the epistoma is unsclerotized and the lateral parts 
are lightly sclerotized; the pleurostoma is lightly sclerotized; the 
hypostoma is long and well sclerotized; the hypostomal spur is short; 
the stipital sclerite is present and its lateral end has a plate-like expan- 
sion in many genera; the labial sclerite is longer than wide in most 
genera and the ventral part is lightly sclerotized in many genera; the 
prelabial sclerite is present and is Y-shaped in most genera; the labial 
sclerite and the prelabial sclerite differ in shape in different genera; the 
maxillary and labial palps each bear two sensilla, one large and one 
small, in most genera; the silk press is well sclerotized; the labral 
sclerite is absent; the mandible is of relatively moderate size and in 
most genera the base is rather square and the blade slender; in Pyrac- 
mon and Prochas the blade is broad; the antenna is disc-shaped; the 
closing apparatus of the spiracle is very lightly sclerotized and adjoins 
the atrium; the skin has small projections and small setae, except in 
Prochas, where the setae are equal in length to the blade of the man- 
dible. 

The following key to genera is not particularly satisfactory. Some 
genera have distinctive characters, but within certain genera, the 
larval characters of the species examined differ widely. This might 
indicate that the classification of the adults is greatly in need of re- 
vision. One solution for the present study would have been to group 
the species examined on larval characters only, ignoring the relation- 
ships indicated by the U.S. Department of Agriculture’s synoptic 
catalog. This has been done to a very limited extent. For example 


482 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


rer ah ei 
may arpa oN eS 


3 


2 O-fmm 3 
Ficure 50.—Ophioninae:Campoplegini, head sclerites: a, Pyracmon xanthognatha (Rohwer); 


B, Campoplex validus (Cresson); c, Xanthocampoplex nigromaculatus (Cameron). (1, 
antenna; 2, spiracle; 3, skin.) 


as PA 
AUT a at Sa 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 483 


1 PA i is se - \, 
Pte ttal 


MATA Art oh ane 
Nc oper aes 


3 


HH 
O-fram 3 


Ficure 51.—Ophioninae:Campoplegini, head sclerites: A, 


B, Casinaria eupitheciae Viereck; c, Charops papilionis Ash 
3, skin.) 


Idechthts nigriscapus Viereck; 
mead. (1, antenna; 2, spiracle; 


484 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


2 Pe in ra are ae 
ee 3 
O:-imm 


Ficure 52.—Ophioninae:Campoplegini, head sclerites: a, Charops narangae (Cushman); 
s, Bathyplectes curculionis (Thomson); c, Biolysia tristis (Gravenhorst), antenna not 
visible on preparation. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 485 


Q : <a 8 neh eer areal ee 
4 Na} ~ Pe Rares 
-——_+ 
Cs O-1mm = 


O-f{mm 3 


Ficure 53.—Ophioninae:Campoplegini; head sclerites: A, Campoletis oxylus (Cresson); 


B, Dusona vitticollis vitticollis (Norton); c, Neptera benevola Gahan. (1, antenna; 2 
spiracle; 3, skin.) 


486 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


ra 
2 rr ee 
a 


a— 
a) Ae -& 


— 


O:imm 3 


Ficure 54.—Ophioninae:Campoplegini. a, Phobocampe disparis (Viereck), head sclerites; 
B, Horogenes comptoniellae (Viereck), head sclerites; c, Hyposoter pilosulus (Provancher), 
mandible. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 487 


oo 


2 femal 3 
O:-tmm 


Ficure 55.—Ophioninae:Campoplegini. a, Olesicampe pikonemae Walley, head sclerites 
(1, antenna; 2, spiracle; 3, skin); B, Olesicampe lophyri (Riley) (1, mandible; 2, labium); 
c, Holocremnus clandestinus (Holmgren), head sclerites (1, antenna; 2, spiracle; 3, skin.) 


504675—59——_7 


488 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficure 56.—Ophioninae:Campoplegini, head sclerites: a, Benjaminia fuscipennis (Pro- 
vancher); B, Charopsimorpha uwnicincta (Ashmead); c, Prochas theclae Walkley, new 
genus, new species. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 489 


some species of Hyposoter are listed in a different part of the key from 
others. But the key is largely a compromise in that an attempt has 
been made to fit the larval characters into a scheme of classification 
based on adult characters. A revision of the tribe in the light of both 
larval and adult characters would be the only satisfactory solution. 


LARvAL Kry 


1. Setae on body approximately equal in length to blade of mandible. 
Prochas Walkley, new genus *4 


Setae on body much shorter than blade of mandible . . . . Shae ure ate 
2. Mandible as in ae 50a with no marked differentiation ereen base and 
bladelcn.m-sare = S Se) ae oe ees  eyracmon 
Mandible with blade more alender tien Paces Rana) 5 3 


3. A lightly sclerotized plate is present lateral to each picuroston and fone 
stoma and extending dorsally to meet ventral half of antenna . Spudastica® 
A lightly sclerotized plate, if present, not of thisshape .. . Bee wee: 
4. A lightly sclerotized and lobed plate as in figure 544 present iterall to each 
pleurostoma and hypostoma and not oars dorsally to meet ventral 
halfofantenna .. . . . . .Phobocampe 
A lightly sclerotized BIAte of this Elias mer recent Seta tsp eteteyate a stmtan <cimesee SO 
5. A lightly sclerotized plate extends dorsally from pleurostoma across to meet 
ventrolateral edge of antenna... . .. . . . . Benjaminia 
A lightly sclerotized plate of this shape not avectait sere hans (eet inns, or eee 
6. Prelabial sclerite roughly triangular in shape with ventral part poorly devel- 
ODEON cats Diese care, eekee hehe : RE er ry eta / 
Prelabial sclerite Y-shaped with SS part as Saul devetoned: as dorsal parts 
9 
7. Ventral part of prelabial sclerite extending to point just ventrad of labial 
palps .........4.+.+. +... . Charops narangae (Cushman) 
Charops ganges Cushman 
Ventral part of prelabial sclerite not extending to point just ventrad of labial 
DADS ss pitt} 
8. A lightly sclerotized ea) Sent beneath ventral part: of labial raclenite, 

Bathyplectes 

A lightly sclerotized plate not present beneath ventral part of labial sclerite. 
Biolysia 
24 In order that Dr. Short may refer to this genus by a valid name a brief description is given here. R.A. 


Cushman had set aside in the U.S. National Museum collection two black campoplegine specimens that 
he had labelled as Prochas theclae, a manuscript name, 


Prochas theclae, new genus, new species 


This new genus is closely related to Charopsimorpha Viereck, 1912 but may be easily distinguished from it 
by the broader postpetiole, different abdominal sculpture, fully areolated propodeum, and by unusual head 
characters for this group of the Campoplegini. The type species, Prochas theclae, has the occipital carina 
meeting the hypostomal carina at the base of the mandible; has non-emarginate eyes (the inner margin at 
most only slightly sinuate); has the clypeus (pale in the male) protruding just before the apex and with the 
apex slightly rounded, and mandibles with flange on only the basal half of lower margin. 

Holotype female (USNM 64687), Trinidad, B.W.I., Aug. 6, 1948, Diego Martin. Allotypemale (USNM), 
Trinidad, B.W.I., Nov. 14, 1941, Brasso. Both specimens were reared from Thecla empusa Hew. on cacao 
by E. McC. Callan. 

Luella M, Walkley 
Entomology Research Division 
U.S. Department of Agriculture 
25 Of Beirne (1941). 


490 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


9. Prelabial sclerite with ventral part broader than lateral parts of labial sclerite. 
Charops papilionis Ashmead 
Prelabial sclerite with ventral part not as broad as lateral parts of labial 


sclerite . .. yin. ge isk nek LO 

10. Lateral parts of epntont prone san dently pcleratizeds . . . Helocremnus 
Lateral parts of epistoma not of thisform ......)...:8 2.4, 0 JEL 

11; Labial sclerite longer than wide2® 5: 5 . =...) 2) «5 <2) sce eet 
Labial sclerite as wide or wider than long .. . < 2 ee 

12. Ventral part of labial sclerite broad and well ealerone _Xanthocampontes 
Ventral part of labial sclerite not as above .. . is & eS 


13. Ventral part of labial sclerite forming broad, fiently Bclenieed plate and not 
distinct from any lightly sclerotized membrane in ventral part of prelabial 
area... 3: a gee 

Ventral part of Bait ecleniees nat paar end fehtly pceroreed plate and 
distinct from any sclerotized membrane in ventral part of prelabial area 27 


16 

14. Prelabial sclerite not meeting broad ventral part of labial sclerite . . Idecthis 
Campoplex 

Prelabial sclerite meeting broad ventral part of labial sclerite . ... . 15 


15. Ventral part of labial sclerite lightly sclerotized and pointed . . . Dusona 
Hyposoter parogyiae (Viereck) 

Ventral part of labial sclerite as well sclerotized as lateral parts and more 
TOUNG eC eancue cmt: . . . . Olesicampe lophyri (Riley) 

16. Ventral part of Tapia eclerite: very tightly sclerotized; lateral part of stipital 
sclerite about 114 times as broad as median part ... . . .Campoletis 
Ventral part of labial sclerite moderately sclerotized; lateral part of stipital 
sclerite about twice as broad as median part .... . . . .Hyposoter 
Horogenes 

Olesicampe pikonemae Walley, 

Olesicampe euurae (Ashmead) 


17. Labial sclerite complete ventrally. . . .... . . . . Charopsimorpha 
Labial sclerite incomplete ventrally ... . pce aie epepemilies 
18. Each lateral part of prelabial sclerite approximately eanalli in tTonewh to ventral 
jORNHB op goo. < . . Casinaria 

Each lateral nae of anleyacl Pent one- enale no onertnied eran of ventral 

PAD Go. os. dar Sees fy S450 Aub) 8a need wale lee. Soe ae ge led aes pig eee 


The following have been examined: Pyracmon xanthognatha (Roh- 
wer) (fig. 50a), Campoplex crassifemur (Thomson), C. depressus 
Viereck, C. frustranae Cushman, C. hyalinus (Provancher), C. phthori- 
maeae (Cushman), ©. polychrosidis Viereck, C. pyraustae Smith, C. 
rufifemur (Thomson), C. tosensis (Uchida), C. tricoloripes (Schmiede- 
knecht), C. validus (Cresson) (fig. 508), Campoplex sp. nr. inaequalipes 
Cresson, Xanthocampoplex nigromaculatus (Cameron) (fig. 50c), 
Xanthocampoplex sp., Idechthis nigriscapus Viereck (fig. 514), 

26 In using this part of the key care should be taken to ensure that all of the prelabial region is fully visible 
in a preparation. In some species which have the ventral part of the labial sclerite lightly sclerotized this 
part may become folded under the rest of the prelabium during mounting, with the result that the labial 
sclerite appears to be much shorter than it really is. 

27 Except Hyposoter parorgyiae (Viereck), where the prelabial sclerite meets the broad ventral part of the 
labial sclerite. 


2% Except Campoplerz tosensis (Uchida), which can be recognized by the bubble-like appearance of the 
prelabial and labial sclerites. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 49] 


canescens (Gravenhorst), Casinaria eupitheciae Viereck (fig. 518), 
C. ndubia (Morley), C. infesta (Cresson), C. tenuiventris (Graven- 
horst), Casinaria sp., Charops ganges Cushman, C. narangae (Cush- 
man) (fig. 52a), C. papilionis Ashmead (fig. 51c), Bathyplectes corvina 
(Thomson), B. curculionis (Thomson) (fig. 528), B. exiquus (Graven- 
horst), Brolysia tristis (Gravenhorst) (fig. 52c), Campoletis ozxylus 
(Cresson) (fig. 534), Dusona glauca glauca (Norton), D. quebecensis 
(Walley), D. terebrator (Foerster), D. variabilis (Franklin), D. vitti- 
collis vitticollis (Norton) (fig. 538), Nepiera benevola Gahan (fig. 53c), 
Phobocampe disparis (Viereck) (fig. 544), P. flavipes (Provancher), 
Horogenes acutus (Viereck), H. aestivalis (Viereck), H. compressus 
(Cresson), H. comptoniellae (Viereck) (fig. 548), H. eureka (Ashmead), 
H. obliteratus (Cresson), H. pterophorae (Ashmead), H. puncetorius 
(Roman), Horogenes sp., Hyposoter fugitivus fugitivus Say, H. fugitivus 
pacificus Cushman, H. nigrolineatus (Viereck), H. parorgyiae (Viereck), 
H. pilosulus (Provancher) (fig. 54c), H. rubiginosus Cushman, Olesi- 
campe euurae (Ashmead), O. pikonemae Walley (fig. 55a), O. lophyri 
(Riley) (fig. 558), Holocremnus clandestinus (Holmgren) (fig. 55c), 
H. ratzeburgi (Tschek), Benjaminia fuscipennis (Provancher) (fig. 56), 
B. euphydryadis (Viereck), Charopsimorpha tibialis (Cresson), C. 
unicincta (Ashmead) (fig. 568), Prochas theclae Walkley, new genus, 
new species (fig. 56c) : 

Pyracmon has a distinctive form of mandible with a relatively broad 
blade. Within the genus Campoplez, C. pyraustae Smith has a labial 
sclerite which is more rounded than in the other species examined, 
and in Campoplex sp. nr. inaequalipes Cresson the mandible has a 
curved but broad blade with a width equal to that of half the depth 
of the base. C. tosensis (Uchida) differs from the species figured in 
that the prelabial sclerite meets the broad ventral part of the labial 
sclerite. Idechthis canescens (Gravenhorst) is basically similar to the 
species figured, but the enlargement on the lateral end of the stipital 
sclerite is relatively smaller. The species of the genus Charops fall 
into two groups: C. ganges Cushman being very like C. narangae 
(Cushman) (fig. 52a), and C. obtusus Morley as figured by Beirne 
(1941) being similar to C. narangae. Bathyplectes and Biolysia are 
distinctive in the shape of the labial sclerite and in the triangular and 
lightly sclerotized prelabial sclerite. Horogenes, Hyposoter, and 
Olesicampe are basically similar. The labial sclerite is more rounded 
than in figure 548 in Hyposoter fugitivus fugitivus Say and H. fugitivus 
pacificus Cushman. Also, in these two subspecies the lightly sclero- 
tized area between the base of the prelabial sclerite and the ventral 
part of the labial sclerite is not present. The mandible of Hyposoter 
has a small lobe dorsal to the blade. This is seen in a slight form in 
other Campoplegini, but is more pronounced in Hyposoter, especially 


492 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


in H. pilosulus (Provancher) (fig. 54c). In H. parorgyiae (Viereck) 
the region between the prelabial sclerite and the ventral part of the 
labial sclerite has the form of a lightly sclerotized plate. In both 
Horogenes and Hyposoter the labial sclerite has a small infolding at 
each venterolateral edge. Olesicampe euurae (Ashmead) is very 
similar to O. pikonemae Walley in structure. In O. lophyri (Riley) the 
labial sclerite differs in shape from that of these two species in that the 
ventral part is broad; also the prelabial sclerite is short and the mandi- 
ble is distinctive in form, being relatively small with a rounded base 
and a small, straight blade. Prochas theclae Walkley differs from other 
Campoplegini in the length of the setae of the skin and the shape of 
the mandible; however, the head sclerites in general resemble those 
of the Campoplegini. 

The following are figured in the literature: Nemeritis bicingulata 
Gravenhorst (Beirne, 1941), Campoplez alkae (Ellinger and Sachtleben) 
(Baker, Bradley, and Clark, 1949; Goidanich, 1931), C. borealis 
(Zetterstedt) (Thorpe, 1930), C. crassifemur (Thomson) (Thompson 
and Parker, 1930), C. cultrator Gravenhorst (Beirne, 1941), C. ensator 
(Gravenhorst) (Thorpe, 1930), C. mutabilis (Holmgren) (Thorpe, 
1930), C. rufifemur (Thomson) (Thorpe, 1930), C. validus (Cresson) 
(Timberlake, 1912), IJdechthis canescens (Gravenhorst) (Daviault, 
1930), Charops obtusus Morley (Beirne, 1941), Bathyplectes exiquus 
(Gravenhorst) (Beirne, 1941), Spudastica kriechbaumeri (Bridgman) 
(Beirne, 1941), Phobocampe unicincta (Gravenhorst) (Beirne, 1941), 
Horogenes armillata (Gravenhorst) (Beirne, 1943), H. exareolatus 
(Ratzeburg) (Beirne, 1941), H. macrostoma (Thomson) (Roberti, 
1947), H. nana (Gravenhorst) (Thorpe, 1933), H. punctorius (Roman) 
(Baker, Bradley, and Clark, 1949), Horogenes sp. (Cameron, 1938), 
Horogenes sp. (Beirne, 1942), Hyposoter pilosulus (Provancher) 
(Tothill, 1922), H. tricolor (Ratzeburg) (Beirne, 1941), Holocremnus 
ratzeburgi (Tschek) (Morris, Cameron, and Jepson, 1937), Balcarcia 
bergt Brethes (Griot and Icart, 1948). 


Tribe Cremastini 


FicurEs 57, 58 


Members of this tribe are endoparasitic, mostly in microlepidop- 
terous larvae in leaf rolls and tunnels. Many of them frequent drier 
habitats than is usual for the family. 

The tribe closely resembles the Campoplegini, although the head 
sclerites are, in general, more lightly sclerotized. Also, the hypostomal 
spur is longer, the lateral part of the stipital sclerite is not expanded, 
except in Hiphosoma where the expansion is small, and the labial 
sclerite is broader than in most Campoplegini. In all species examined 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 493 


aA~y-~n — v0 ~— 
O-fmm Bes ae eas 


Figure 57.—Ophioninae:Cremastini, head sclerites: a, Pristomerus vulnerator (Panzer); 
B, Atphosomella stenomae Cushman; c, Zaleptopygus flavo-orbitalis (Cameron). (1, 
antenna; 2, spiracle; 3, skin.) 


494 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


A> as bana At 
a 
a in Raa 


_———— 
O:{ mm 


Ficure 58.—Ophioninae:Cremastini, head sclerites: a, Cremastus ferrugineus Davis; 
B, Eiphosoma batatae Cushman. (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 495 


the labial sclerite is incomplete ventrally. The skin is lightly sclero- 
tized with small projections and small setae. 


LARVAL Key 


1. Each lateral part of the labial sclerite with a small projection towards the 


MECIANGMMC. Usp rend slowest TRE pee ke hae ari ae ica Eiphosoma 
Each lateral part of labial sclerite without small projection towards median 
VaTIG irs: 5 Se rectpeeir eh ogee hen then ad ue is be aera es 2 

2. A very lightly sclerotized epistoma present. . . Zaleptopygus, Cremastus 
pistomarnot sclerotized, 4s <2 2 Bee us oC A a 3 

3. Hypostoma bent in the middle of itslength. ........ Pristomerus 


Hypostoma bent at point one third of length from median end . Xiphosomella 


Pristomerus euryptychiae Ashmead differs from the drawing in figure 
57A in that there are two groups each of two sensilla on the labrum. 
Cremastus minor Cushman differs from the drawing in figure 58a in 
that the ventral part of the labial sclerite is lightly sclerotized. 

It was not found possible to differentiate Zaleptopygus and Cremas- 
tus on larval characters. 

The following have been examined: Pristomerus austrinus (Townes 
and Townes), P. euryptychiae Ashmead, P. pacificus appalachianus 
Viereck, P. vulnerator (Panzer) (fig. 574), Xtphosomella stenomae 
Cushman (fig. 578), Zaleptopygus flavo-orbitalis (Cameron) (fig. 57¢), 
Z. gallaecola (Cushman), Z. rosae (Cushman), Zaleptopygus sp., 
Cremastus carpocapsae Cushman, C. epagoges Cushman, C. facilis 
(Cresson), C. ferrugineus Davis (fig. 584), C. forbesi Weed, C. minor 
Cushman, Hiphosoma annulatum Cresson, E. batatae Cushman (fig. 
588), H. insularis (Viereck), E. texcanum Cresson. 

The following are figured in the literature: Pristomerus vulnerator 
(Panzer) (Goidanich, 1931; Rosenberg, 1934), Zaleptopygus flavo- 
orbitalis (Cameron) (Bradley and Burgess, 1934), Cremastus interruptor 
Gravenhorst (Thorpe, 1930; Beirne, 1941). 


Tribe Tersilochini 
FIGuRE 59 


This tribe, the larvae of which are endoparasites, resembles the 
Cremastini and many Campoplegini in essential structure, but the 
mandible is distinctive with its conical shape and small tooth at the 
apex. The head sclerites are slender and lightly sclerotized. As in 
most Cremastini, the stipital sclerite has no lateral expansion. The 
labial sclerite is broader than long and is incomplete ventrally. The 
ventral part of the prelabial sclerite is absent and several large sensilla 
are present on the ventral edge of this sclerite. The maxillary and 
labial palps are relatively large. The spiracle is of the usual shape in 
the Ophioninae with a lightly sclerotized closing apparatus adjoining 


496 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


D 


Ry cee 


— 


y ¢ YEA Ase 
Da Soa: = 


2 ———| 3 


Figure 59.—Ophioninae:Tersilochini, head sclerites: Tersilochus argentinensts (Blanchard) 
(antenna not visible on preparation; 2, spiracle; 3, skin.) 


the atrium. The skin has relatively long setae, approximately equal 
in length to one-third the length of the mandible, but the projections 
on the skin are small. 

Tersilochus argentinensis (Blanchard) (fig. 59) and J. conotracheli 
(Riley) were examined. IT. conotracheli is figured by Cushman (1916) 
and Tersilochus sp. by Beirne (1941). 


Tribe Ophionini 
Fiaure 60 a, B 


Members of this tribe are endoparasitic in larger caterpillars, except 
that Ophion bifoveolatus Brullé is known to parasitize larvae of Phyl- 
lophaga (Scarabeidae). Species of Ophion and Enicospilus are noc- 
turnal or crepuscular and are frequently attracted to lights. 

This is a distinctive tribe although it resembles other tribes of the 
Ophioninae in many basic features. The head sclerites are broad 
and moderately sclerotized. The stipital sclerite has a small lateral 
expansion and a lightly sclerotized cardo is present. The ventral 
part of the labial sclerite forms a broad plate. The prelabial sclerite 
is triangular in shape with the ventral part reduced. Numerous 
setae are present on the labrum, on the maxilla lateral to the hypos- 
tomal spur and on the prelabial membrane and the labial sclerite. 
The mandibles are distinctive, being relatively small and with a 
rounded base and a slightly or strongly curved blade. The antenna 
is disc-shaped. The closing apparatus of the spiracle is lightly 
sclerotized and adjoins the atrium. The skin has very small setae 
and small projections. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 497 


2 


Ficure 60.—a, x, Ophioninae:Ophionini, head sclerites: A, Ophion idoneum Viereck; B, Eni- 
cospilus arcuatus (Felt). c, Anomalinae:Anomalini, head sclerites, Anomalon sp., antenna 
not visible on preparation. (1, antenna; 2, spiracle; 3, skin.) 


498 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


= 


ee a a 
pit CSE ts 


3 


O-1mm 


Ficure 61.—Anomalinae:Gravenhorstiini, head sclerites: a, Aphanistes hyalinus (Norton), 
antenna not visible on preparation; 8, Barylypa insidiator (Foerster), antenna not visible 
on preparation; c, Labrorychus analis (Say). (1, antenna; 2, spiracle; 3, skin.) 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 499 


See ee 
Arn er AA SARE EAT 


A AAR ARRS 


NO. 


Ficure 62.—Anomalinae: Gravenhorstiini, head sclerites: a, Atrometus clavipes (Davis); 
B, Agrypon sp.; c, Blaptocampus nigricornis (Wesmael). (Antenna not visible on prepara- 
tion; 2, spiracle; 3, skin.) 


500 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


O:imm 


Ficure 63.—Anomalinae:Therionini, head sclerites: a, Therion circumflexum (Linnaeus); 
B, Heteropelma fulvicorne Townes, antenna not visible on preparation. (1, antenna; 
2, spiracle; 3, skin.) . 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 501 


LARVAL KEY 


ie Mandible) witht bladetshehbthyycurved( $6 2.)05 if)2 fy exude) Se cya. ne Ophion 
Mandible with blade strongly curved so that the tip points dorsally. 
Enicespilus 

The following were examined: Ophion ancyloneura Cameron, 
O. bilineatus Say, O. idoneus Viereck (fig. 60a), O. tetyri Packard, 
Enicospilus americanus (Christ), /. arcuatus (Felt) (fig. 608), E. 
glabratus (Say), EH. horsfeldi Cameron, E. purgatus (Say), 7. texranus 
(Ashmead), and Enicospilus spp. 

In Enicospilus americanus the blade of the mandible, although of 
the Enicospilus form, is not so strongly curved as in figure 608. A 
lightly sclerotized plate is present on the ventral edge of the labrum 
of some species of Hnicospilus. This should not be confused with 
the suspensorium of the hypopharynx which is also visible in several 
species. 

The following are figured in the literature: Thyreodon atricolor 
(Olivier) (Cushman, 1947), Enicospilus macrurus (Drury) (Beirne, 
1941), FE. purgatus (Say) (Cushman, 1947), Enicospilus sp. (Moutia 
and Courtois, 1952), and Ophion idoneus Viereck (Cushman, 1947). 


Subfamily Anomalinae 


Fiaures 60c-63 


This endoparasitic group usually has been placed as a tribe of the 
Ophioninae. The larval characters of those species examined differ 
markedly from the characters of the Ophioninae and the group has 
therefore been treated as a separate subfamily. The form of the 
labial sclerite is generally similar in the Anomalinae and Ophioninae 
and, although this might indicate some sort of general relationship 
between the two groups, the form of the remaining sclerites and the 
mandibles differs sharply. 

The epistoma, pleurostoma, and hypostoma are broad and well 
sclerotized; the hypostomal spur is absent; the stipital sclerite curves 
dorsally from the labial sclerite to meet the hypostoma; the labial 
sclerite is relatively slender and is incomplete ventrally except in 
Anomalon where the ventral part is moderately sclerotized, and 
in Heteropelma where the ventral part is very lightly sclerotized; the 
maxillary and labial palps each have two sensilla, one large and one 
small, except in Anomalon where the two sensilla are approximately 
equal in size, the dorsal being flat and the ventral peglike; the silk 
press is sclerotized; a lightly sclerotized prelabial sclerite which lacks 
a ventral projection is present in some genera; the labral sclerite is 
absent, but large sensilla which are grouped in various patterns are 
present on the labral area; setae are reduced and labral and prelabial 
setae appear to be absent except in Anomalon; the mandibles are 
large with relatively straight blades which lack teeth except in 


502 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 110 


Anomalon, and the blade of each mandible is well sclerotized; the 
antenna is disc-shaped; the closing apparatus of the spiracle is lightly 
sclerotized and adjoins the atrium; the skin has small projections 
and small setae. 

The larval characters of the Anomalinae are strikingly like those 
of the Metopiinae. The larvae of the two subfamilies may be distin- 
guished by the presence of setae on the prelabium of the Metopiinae 
and the absence of these setae in the Anomalinae except Anomalon, 
a genus having so many special features as to be readily recognizable. 

The reduction of the labial sclerite and hypostomal spur in the 
Anomalinae, Metopiinae, and Ichneumoninae might be correlated 
with the relatively slight cocoon spun by these endoparasites. In 
the head of the final instar larva of Xorides muscles are present which 
insert on the labial sclerite (Short, 1952, p. 41, fig. 8a). The con- 
traction of these muscles tilts forward the dorsal end of the pre- 
labium with the opening of the silk glands. In this action the labial 
sclerite rotates on the extremities of the stipital sclerites. The 
labial sclerite is thus held so that movement can take place in only 
one plane. The spurs of the hypostomae, which articulate with the 
stipital sclerites, hold the maxillae rigid. The spinning of the cocoon 
has been observed in Xorides and the region of the prelabium contain- 
ing the opening of the silk glands was tilted forward during this 
process. It is therefore possible that the reduction of a cocoon is 
accompanied by a reduction of the muscles which evert the dorsal 
end of the prelabium and a reduction of the labial sclerite and hypos- 
tomal spur. One objection to this suggestion is that the Pimplini, 
which emerge from the pupa of a lepidopterous host and have only 
a slight cocoon, have a well developed hypostomal spur and labial 
sclerite. However the stipital sclerite is reduced and this might 
be associated with the reduction of the cocoon since, when being 
everted, the labial sclerite rotates on the stipital sclerite. 


LARVAL Kry 


1. Blade of mandible with teeth; setae on prelabium....... Anomalini 
Blade of mandible without teeth; setae absent from prelabium... . 2 

2. Width of epistoma along entire length and width of pleurostoma along entire 
length equal to at least half length of mandible. ...... Therionini 
Width of epistoma or part of width of pleurostoma less than length of man- 
ible Zaee Oe Vo NSE A aay adore ome ete Gravenhorstiini 
Ophionellini ” 


Tribe Anomalini 
Figure 60c 
The general form of the head sclerites, spiracles, and skin places 
Anomalon with the genera of the tribes Gravenhorstiini and Therionini; 


29 Of Cushman (1947). 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 503 


but it does not appear to be a close relationship, a conclusion which 
Walkley (in litt.) has also reached from study of the adults. Anomalon 
differs from the other Anomalinae examined in that the blade of the 
mandible bears distinct teeth; there are setae on the prelabium; and 
the ventral part of the labial sclerite, although not as well sclerotized 
as the lateral parts, is nevertheless distinctly sclerotized. It is also 
possible that the hypostomal spur might be represented in Anomalon 
by a small projection from the anteroventral part of the hypostoma. 
Anomalon therefore can be considered to have an isolated position 
within the Anomalinae. The larval habits also distinguish this genus 
from other Anomalinae in that larvae of Elateridae appear to be the 
usual hosts, whereas the hosts of the other Anomalinae examined were 
lepidopterous. 

Anomalon sp. (60c) was examined. A. eyuncidum Say is figured by 
Cushman (1947). 


Tribe Ophionellini 


Ophionellus foutst (Cushman) is figured by Cushman (1947). This 
figure is not sufficiently detailed to allow the Ophionellini to be dis- 
tinguished from the Gravenhorstiini. 


Tribe Gravenhorstiini 


Fiaures 61, 62 
LARVAL Kry 


1. Labral sensilla arranged in two separate groups. ... . . . . Atrometus 
Labral sensilla arranged in single line... . a ane” 

2. When head sclerites mounted flat on slide then siete ends a hypostomae 
extend to point approximately level with ventral end of labial sclerite . . 3 
When head sclerites mounted flat on slide then posterior ends of hypostomae 
do not extend to point level with ventral end of labial sclerite. . ... 4 

3. Width of pleurostoma at anterior pleurostomal process approximately equal 
to two-thirds length of mandible. . .. . ... . . Aphanistes 
Width of pleurostoma at anterior plearostoreal neoges much less than two- 
thirds length of mandible. . .. . . . . . . Labrorychus 

4. Hypostoma with sclerotized expansion Alone encire dorsal edge . . Barylypa 
Hypostoma without sclerotized expansion along entire dorsal edge. . . . 5 

5. Length of blade of mandible less than half width of base and arising from 
ventral surface of base. . . . . . . Blaptocampus 
Length of blade of mandible arorna tele equal Ko half width of base and 
arising from* middie of bases. 2th). 2. ee. Oe 2 SY Agrypon 


The following have been examined: Aphanistes hyalinus (Norton) 
(fig. 614), Barylypa insidiator (Foerster) (fig. 618), Labrorychus analis 
(Say) (fig. 61c), ZL. prismaticus (Norton), Labrorychus sp., Atrometus 
clavipes (Davis) (fig. 624), Agrypon sp. (fig. 628), Blaptocampus 
nigricornis (Wesmael) (GCV) (fig. 62c). 

504675—59 8 


504 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


The following are figured in the literature: Trichomma enecator 
(Rossi) (Rosenberg, 1934), Aphanistes armatus (Wesmael) (Beirne, 
1941), Blaptocampus nigricornis (Wesmael) (Beirne, 1941). 


Tribe Therionini 
Figure 63 
LarvAL Kry 
1. Ventral part of labial sclerite lightly sclerotized. ...... Heteropelma 
Ventral part of labial sclerite not sclerotized. .......... Therion 
The following have been examined: Therion morio (Fabricius), 
T. circumflecum (Linnaeus) (fig. 63a), Heteropelma fulvicorne Townes 
(fig. 638). Therion morio (Fabricius) is figured by Cushman (1947). 


Interrelationships between the Subfamilies 


Although in many cases the larval characters suggest interrelation- 
ships which are very similar to those which are believed to be shown 
by the characters of the adults, there are certain discrepancies. 

Some similar adults, such as the Pimplini and Ephialtini, have very 
different larvae. These differences may be correlated with the fact 
that the larvae of the Pimplini are endoparasitic while those of the 
Ephialtini are ectoparasitic. The larval characters of the Ephialtini 
and similar tribes of the Pimplinae resemble those of other ectopara- 
sites such as the Tryphoninae and Cryptinae rather than the Pimplini. 

There are certain structural features which appear to be connected 
with whether the larva is an ectoparasite or an endoparasite. The 
antenna in most ectoparasites is papilliform. In most endoparasites 
it is reduced and disc-like. ‘The mandible is toothed in most ecto- 
parasites where teeth are necessary for holding on to the host and 
for piercing the host skin. In most endoparasites the mandible is 
without teeth. The labral sclerite is present in most ectoparasites 
and absent in most endoparasites. The epistoma and the labral 
sclerite are rarely present together, except in the Tryphoninae, where 
the epistoma is slender. It is difficult to suggest a reason for this. 
The anterior retractor muscles of the labrum insert on the median 
part of the labral sclerite (Short, 1952, p. 39, fig. 7B). It is not 
likely that the labral sclerite could brace the cranium in any way as 
a substitute for the epistoma. The epistoma is well developed in 
larvae with powerful mandibles. 

The closing apparatus of the spiracle adjoins the atrium in most 
endoparasites, whereas in ectoparasites it may adjoin the atrium or be 
situated some distance from the atrium. There is no obvious explana- 
tion for this. It is generally assumed that the spiracles are open in 
endoparasites in the final instar larva, which is carnivorous, feeding on 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 505 


/——____ 
2 O-imm 
Figure 64.—Cryptinae:Hemitelini, head sclerites: 4tractodes sp. (1, antenna; 2, spiracle; 
3, skin.) 


the tissues of the host and liberating air from the tracheae. If the 
spiracle is open there is possibly some biological advantage in having 
the closing apparatus situated close to the atrium. If one function of 
the closing apparatus is to prevent the entry of body fluids of the host 
into the tracheal system, then it is best situated to perform this func- 
tion when it adjoins the atrium. If, however, such a precaution were 
necessary, one would more readily expect to find hydrofuge structures 
at the opening of the tracheal system, as in many aquatic insects. 

The setae of the skin of endoparasites are reduced. 

It should be noted that the Cryptinae, which contain both ecto- 
parasites and endoparasites of prepupae and pupae, show larval 
characters which are relatively uniform within the group. 

Just as some similar adults have different larvae, so do some different 
adults, such as the Metopiinae and Anomalinae, have similar larvae. 
These similarities may be correlated with larval habits. The Meto- 
piinae, Anomalinae,®” Ichneumoninae, and Pimplini all have large, 


30 Except Anomalon. 


506 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


toothless mandibles with a corresponding development of the epistoma 
and pleurostoma. In the Metopiinae, Anomalinae, and Ichneumoninae 
the hypostoma is also well developed. In the Pimplini the hypostoma 
is reduced, but the hypostomal spur is well developed. The hypostomal 
spur is absent in the Metopiinae, Anomalinae, and Ichneumoninae. 
The labial and stipital sclerites are, except in the Pimplini, relatively 
poorly developed. The reduction of these sclerites might be corre- 
lated with the relatively slight cocoons spun by these endoparasites 
(see section on Anomalinae). 

In the species examined of the Metopiinae, Anomalinae,”! Ichneu- 
moninae, and Pimplini, development takes place in, or is completed in, 
the pupa of a lepidopterous host. The Campoplegini, which are endo- 
parasites of larvae, have a different type of head structure with the 
epistoma absent and the labial and stipital sclerite relatively well 
developed. With regard to the degree of development of these sclerites 
it is significant that many Campoplegini emerge from the host larva to 
spin a well developed cocoon, although some species of Hyposoter and 
Benjaminia pupate within the host larva. 

It cannot be claimed that all Ichneumonidae which emerge from the 
pupa of the host, or even from the pupa of a lepidopterous host, will 
have a larval head of similar form. The Cryptinae have a head struc- 
ture which is very different from that of the Metopiinae, Anomalinae, 
Ichneumoninae, and Pimplini. Also, larval habits are not constant. 
Mr. J. F. Perkins (in litt.) has directed my attention to some reared 
specimens of 7richomma occisor Habermehl with a quite well developed 
cocoon and to an Ophion in which the cocoon is spun within the host 
pupa, which remains adhering to the outside of the cocoon. Thorpe 
(1930) has observed that the ectoparasite E’phialtes ruficollis (Graven- 
horst) spins a light and irregular cocoon, yet when reared in a gelatin 
capsule usually does not spin a cocoon. But it does appear that in most 
Metopiinae, Anomalinae, Ichneumoninae, and Pimplini there is some 
connection between the form of the larval head and the habits of the 
larva. It is thus possible that convergence in larval characters asso- 
ciated with a similarity in larval habits might explain the similarity of 
the Metopiinae and Anomalinae. 

From the standpoint of larval characters alone, the following inter- 
relationships are suggested among the subfamilies of the Ichneu- 
monidae. The Metopiinae and Anomalinae are related. The Ophi- 
oninae, Mesoleiinae, and Lissonotinae are also related subfamilies. 
Similarly, but to a lesser extent, there appears to be some degree of 
relationship between the Tryphoninae, Cryptinae, and the tribes of 
the Pimplinae other than the Pimplini and Acaenitini. The Ichneu- 
moninae show some likeness to the Orthopelmatinae. The Adelogna- 
thinae, Collyriinae, Plectiscinae, Diplazoninae, and Mesochorinae are 


31 Except Anomalon, which pupates within the larvae of Elateridae. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 507 


all isolated subfamilies. The Pimplini and Acaenitini are isolated 
tribes showing some degree of relationship to each other and to the 
remaining tribes of the Pimplinae. 

There was thought to be no point in attempting to construct a family 
tree. 


Relationship between the [Ichneumonidae and Braconidae 


The tribe Xoridini of the Pimplinae has a type of mandible which 
resembles that of the subfamily Braconinae of the Braconidae. The 
Braconinae possess the most generalized larval structure of the 
Braconidae. Since the larval structure of the Xoridini is also gener- 
alized, it is possible that this tribe connects the Ichneumonidae to the 
Braconidae. There is a similarity in larval habit as well as in larval 
structure since many primitive ichneumonids and braconids are 
parasites of woodboring insects (see Brues, 1921). 


References 


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Buunk, H. 
1952. Zur Kenntnis der Hyperparasiten von Pieris brassicae L. Pt. 5: 
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Brapiey, W. G., ano Burasss, E. D. 
1934. The biology of Cremastus flavo-orbitalis (Cameron), an ichneumonid 
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441, 16 pp. 


008 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Brown, N. R. 

1947. Studies on the parasites of the spruce budworm, Archips fumiferana 
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Brugs, C. T. 

1921. Correlation of taxonomic affinities with food habits in Hymenoptera, 
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CAMERON, E. 

1938. Astudy of the natural control of the pea moth, Cydia nigricana Steph. 
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1950. The biology and economie importance of Alomya debellator (F.), a 
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1951. On the identity of an ichneumonid parasite of Hepialis lupulinus (L.). 
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CurystaL, R. N., AND SKINNER, E. R. 

1931. Studies on the biology of Xylonomus brachylabris Kr. and X. irrigator 
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Cuancy, D. W. 

1946. The insect parasites of the Chrysopidae (Neuroptera). Univ. Cali- 

fornia Publ. Ent., vol. 7, pp. 403-496. 
CusHMAN, R. A. 

1916. Thersilochus conotracheli, a parasite of the plum curculio. Journ. 
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1947. A generic revision of the ichneumon-flies of the tribe Ophionini. 
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Cusuman, R. A., AND Rouwer, §. A. 

1921. Holarctic tribes of the subfamily Ichneumoninae (Pimplinae). Proc. 

U.S. Nat. Mus., vol. 59, pp. 379-396. 
Daviautt, L. 

1930. Notes biologiques sur Nemeritis canescens Gray. et sur la morphologie 

de ses divers stades. Rev. Path. Veg., vol. 17, pp. 82-93. 
Downpen, P. B. 

1941. Parasites of the birch leaf-mining sawfly (Phyllotoma nemorata). 

U.S. Dep. Agr., Techn. Bull. 757, 56 pp. 
Given, B. B. 

1944. The anatomy of the final larval instar of Diadromus (Thyraella) 
collaris Grav. (Ichneumonidae), with notes on structural changes 
during the prepupal and pupal stages. Trans. Roy. Soc. New 
Zealand, vol. 74, pp. 297-301. 

GorpaNicu, A. 

1931. Gli insetti predatore e parassiti della Pyrausta nubilal’s Hubn. Boll. 

Lab. Ent. Bologna, vol. 4, pp. 77-215. 
Griot, M., anp Icart, A. 

1948. Observaciones sobre Balcarcia bergi Brethes parasito del “bicho de 

cesto.” Rev. Inv. Agr. Buenos Aires, vol. 2, pp. 197-204. 
Imms, A. D. : 

1918. Observations on Pimpla pomorum Ratz., a parasite of the apple 

blossom weevil. Ann. Appl. Biol., vol. 4, pp. 211-217. 


ICHNEUMONID FINAL INSTAR LARVAE—SHORT 509 


Irawa, K. 
1950. Biology of ichneumon parasites of bag-worms in Japan, I. Trans. 
Kansai Ent. Soc., vol. 15, pp. 35-47. 
Kost, G. 8., anp Hincxs, W. D. 
1945. A check list of British insects. 483 pp. 
MAnNeEVAL, H. 
1936. Nouvelles notes sur divers hyménoptéres et leurs larves. Rev. 
Frangaise Ent., vol. 3, pp. 18-32. 
Meyer, H. 
1922. On the morphology of the larvae of some parasites of the family 
Ichneumonidae. Rep. Bur. Appl. Ent. Leningrad, vol. 2, pp. 25-39. 
Morris, K. R. S. 
1937. The prepupal stage in Ichneumonidae, illustrated by the life-history 
of Exenterus abruptorius, Thb. Bull. Ent. Res., vol. 28, pp. 525-534. 
Morris, K. R. 8.; Cameron, E.; anp Jepson, W. F. 
1937. The insect parasites of the spruce sawfly (Diprion polytomum Htg.) 
in Europe. Bull. Ent. Res., vol. 28, pp. 341-393. 
Movutta, L. A., anp Courrots, C. M. 
1952. Parasites of the moth-borers of sugar-cane in Mauritius. Bull. Ent. 
Res., vol. 43, pp. 325-359. 
NIELSEN, E. 
1923. Contributions to the life-history of the pimpline parasites (Poly- 
sphincta, Zaglyptus, Tromatobia). Ent. Medd., vol. 14, pp. 137- 
205. 
1935. <A third supplementary note upon the life histories of the Poly- 
sphinetas. Ent. Medd., vol. 19, pp. 191-215. 
1936. The biology of Homonotus sanguinolenius Fabr. Ent. Medd., vol. 
19, pp. 385-404. 
Rosert!, D. 
1947. Le oplocampe del Susino. I, Hoplocampe flava (L.). Boll. Lab. 
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RosENBERG, H. T. 
1934. The biology and distribution in France of the larval parasites of 
Cydia pomonella L. Bull. Ent. Res., vol. 25, pp. 201-256. 
Sat, G. 
1931. Parasites of the wheat-stem sawfly, Cephus pygmaeus Linnaeus, in 
England. Bull. Ent. Res., vol. 22, pp. 479-545. 
Saugr, H. F. G. 
1939. Biologia de Calliephialtes dimorphus Cushm. (Hym.: Ichn.) um 
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ScHNEIDER, F. 
1950. Die Entwicklung des Syrphidenparasiten Diplazon fissorius Grav. 
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Scort, E. I. 
1939. An account of the developmental stages of some aphidophagous 
Syrphidae (Dipt.) and their parasites (Hymenopt.). Ann. Appl. 
Biol., vol. 26, pp. 509-532. 


510 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


SHort, J. R. T. 

1952. The morphology of the head of larval Hymenoptera with special 
reference to the head of Ichneumonoidea, including a classifica- 
tion of the final instar larvae of the Braconidae. Trans. Roy. Ent. 
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1953. <A grouping by larval characters of some species of the genus Apanteles 
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SILVEsTRI, F. 

1941. Contribuzioni alla conoscenza degli insetti dannosi e dei loro sim- 
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Srmmonps, F. J. 

1947. The biology of Phytodietus pulcherrimus (Cress.) (Ichneumonidae, 
Tryphoninae) parasitic on Lozostege sticticalis L. in North America. 
Parasitology, vol. 38, pp. 150-156. 

1948. The biology of the parasites of Lozostege sticticalis L. in North 
America. IV. Cryptus inornatus Pratt (Ichneumonidae, Cryp- 
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Snoperass, R. E. 
1935. Principles of insect morphology. ix-+667 pp. 
Spryrer, W. 

1926. Pimpla pomorum Ratz. (Ichneumon.), der Parasit des Apfelbliiten- 
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Berlin, vol. 14, pp. 231-257. 

THompson, W. R., AnD Parker, H. L. 

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TuHorrs, W. H. 

1930. Observations on the parasites of the pine-shoot moth, Rhyacionia 
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Towness, Henry, AND Townss, MArJorRI£ 

1951. Family Ichneumonidae. Jn Muesebeck, et al., Hymenoptera of 
America north of Mexico—Synoptic catalog. U.S. Dep. Agr. 
Monogr. No. 2, pp. 184-409. 

Van Empen, F. I. 

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ICHNEUMONID FINAL INSTAR LARVAE—SHORT 511 


WaLkLeEy, Lugetia M. 
1956. A tribal revision of the brachycyrtine wasps of the world (Cryptinae- 
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1958. Family Ichneumonidae, in Krombein, ed., Hymenoptera of America 
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No. 2, suppl. 1, pp. 36-62. 
WiaGLeswortn, V. B. 
1954. The physiology of insect metamorphosis. viii-+-152 pp. 
WisnHart, G. 
1949. The biology of Melanichneumon rubicundus (Cress.) (Hymenoptera, 
Ichneumonidae). Canadian Ent., vol. 80, pp. 118-137. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


issued 4 Weer 


SMITHSONIAN INSTITUTION 
U.S. NATIONAL MUSEUM 


Vol. 110 Washington : 1960 No. 3420 


MAMMALS OF NORTHERN COLOMBIA, PRELIMINARY 
REPORT NO. 8: ARBOREAL RICE RATS, A SYSTEMATIC 
REVISION OF THE SUBGENUS OECOMYS, GENUS 
ORYZOMYS ! 


By Puitie HERsHKOVITZ? 


Arboreal rice rats are small to medium-sized cricetines of the 
genus Oryzomys (family Muridae). They are found only in tropical 
and subtropical zone forests of Central and South America. Of the 
two recognized species, the larger, Oryzomys (Oecomys) concolor, occurs 
in northern Colombia. 

The author collected 27 specimens from six localities during his 
1941-43 tenure of the Walter Rathbone Bacon Traveling Scholarship 
and 38 specimens, including six of the smaller species, Oryzomys 
(Oecomys) bicolor, in other parts of Colombia while conducting the 
Chicago Natural History Museum-Colombian Zoological Expedi- 
tion (1949-52). This material and pertinent field observations are 
the basis of the present report. 


; j Previous reports in this series have been published in the Proceedings of the U.8. National Museum as 
‘ollows: 
1. Squirrels, vol. 97, August 25, 1947. 
Spiny rats, vol. 97, January 6, 1948. 
. Water rats, vol. 98, June 30, 1948. 
. Monkeys, vol. 98, May 10, 1949. 
. Bats, vol. 99, May 1949. 
: Rabbits, vol. 100, Suiy 26, 1950. 
. Tapirs, vol. 103, May 18, 1954. 


Garator of Mammals, Chicago Natural History Museum. 


Nop et 


513 
504676—59—1 


514 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


MatTERIAL 


A total of 390 specimens was studied. This number includes vir- 
tually all arboreal rice rats preserved in American museums, and the 
types only in the British Museum (Natural History). 

The following abbreviations for museums are used in the lists of 
specimens examined. 

AMNH= American Museum of Natural History 

BM = British Museum (Natural History) 

CM = Carnegie Museum 

CNHM=Chicago Natural History Museum 

MACN = Museo Argentino de Ciencias Naturales 
MCZ =Museum of Comparative Zoology 

UMMZ= University of Michigan Museum of Zoology 
USNM =United States National Museum 

The writer expresses thanks to the authorities of the above insti- 
tutions for permitting him to study the material in their charge. 


THE ORYZOMYINE RODENTS 


The general characters of subgenus Oecomys are common to Oryzomys 
and a number of other genera which together compose the oryzomyine 
group of the subfamily Cricetinae (family Muridae). In addition 
to the aforementioned, the group includes the currently recognized 
genera or subgenera Melanomys Thomas, Oligoryzomys Bangs, 
Microryzomys Thomas, Nesoryzomys Heller, Neacomys Thomas, 
Scolomys Anthony, Nectomys Peters, Sigmodontomys J. A. Allen, 
and Megalomys Trouessart. The group has been defined elsewhere 
(Hershkovitz, 1944, Univ. Michigan Mus. Zool. Misc. Publ. No. 
58, pp. 12-17). Its more salient characters may be summarized 
as follows. 

Size and form Mus-like to Rattus-like; eyes and ears normal; hind 
feet of the generalized scansorial-terrestrial type, or moderately 
specialized for running, swimming or climbing; three middle digits 
of hind foot partly, but not always conspicuously, webbed; claws 
short, more or less recurved, never specialized for digging; tail always 
more than one-half as long as head and body combined, generally 
thinly haired, the scales always visible; mammae eight (two pairs 
pectoral, two pairs inguinal). Distance between first molars always 
more than length of M’; median longitudinal palatal ridge not present; 
bony palate produced posteriorly beyond plane of last molars; postero- 
lateral palatal fossa present and marked by a distinct pit or reticula- 
tion of pits; parapterygoid fossa shallow, its anterior corner never 
undercut. Incisors ungrooved, generally opisthodont, never proodont; 
molars generally brachyodont, never distinctly hypsodont; crowns 
crested or terraced, never plane (cf., Hershkovitz, 1955, Fieldiana, 
Zool., vol. 37, p. 649); mesoloph (id) present in all molars and fused 


ARBOREAL RICE RATS—HERSHKOVITZ 515 


with mesostyle (id) as a functional unit; procingulum of upper molars 
and postcingulum of lower molars present and usually well developed. 

Remarks: With few exceptions, all real species of oryzomyine 
rodents are well defined. Unfortunately, the multiplication of names 
has made the identification of any one of them an extremely arduous, 
sometimes nearly impossible, task. A total of 54 names, mostly of 
specific grade, have been given to representatives of Oecomys alone. 
These are shown to be applicable to only two species with a com- 
bined total of nine subspecies. 


Subgenus Oryzomys (Oecomys) Thomas 


Gcomys, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, p. 444, 1906 (subgenus of 
Oryzomys Baird); ser. 8, vol. 3, pp. 378-379, 1909 (genus). 

Oecomys, Gyldenstolpe, Kung]. Svenska Vet.-Akad. Handl., ser. 3, vol. 11, p. 38, 
1932 (characters; list of species)—Tate, Amer. Mus. Nov., No. 581, 
p. 10, 1932 (taxonomic history).—Ellerman, The families and genera of living 
rodents, vol. 2, pp. 340, 342, 357, 408, 1941 (subgenus of Oryzomys). 

Oryzomys, Goldman, North American Fauna, No. 43, pp. 15, 82, 1918 (part, 
Oryzomys tectus group). 

Typr species: Rhipidomys benevolens Thomas [=Oryzomys 
(Oecomys) bicolor phaeotis Thomas], by original designation. 

INCLUDED SPECIES: Oryzomys concolor Wagner, O. bicolor Tomes. 

Disrrinution (figs. 1, 2): Forested parts of the tropical and 
subtropical zones of Central and South America, from Costa Rica 
through Panama into Colombia, Ecuador, Peru, Bolivia, Venezuela, 
Trinidad, the Guianas and Brazil; altitudinal range from near sea 
level to approximately 2,000 meters above. 

The ranges of the two species are coextensive in Panama, the Rio 
Orinoco basin of Colombia and Venezuela, and the Amazonian region. 
So far as known, only the larger species, Oryzomys concolor, occurs in 
Costa Rica, northern Colombia, northwestern Venezuela, and Trini- 
dad. The smaller OQ. bicolor occurs alone in western Ecuador. 


Taxonomic History 


The first name proposed for an arboreal rice rat is Mus cinnamo- 
meus Pictet and Pictet, 1844. It is based on an eastern Brazilian 
representative of the larger species of subgenus Oecomys. Unfortu- 
nately, it is invalid because the name Mus cinnamomeus had already 
been used by Lichtenstein in 1830 for a species of spiny rats of the 
modern genus Proechimys. The next name for an Oecomys is Hesper- 
omys concolor Wagner, 1845. It was given to the large form discovered 
by Johann Natterer in the forests of the upper Rio Negro, northwestern 
Brazil. While the name concolor was virtually ignored by recent 
authors, the animal it represents has since been redescribed under 
27 different names. 


VOL. 110 


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PROCEEDINGS 


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518 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


The first binomial for the smaller species of subgenus Oecomys is 
Hesperomys bicolor 'Tomes, given in 1860 to a specimen from eastern 
Ecuador. As in the case of O. concolor, the true nature of bicolor 
was not immediately appreciated by subsequent workers. The 
next member of the species, collected in western Ecuador, was de- 
scribed by Thomas in 1900 as Rhipidomys dryas. This was followed 
by Rhipidomys pheotis, R. benevolens, R. rosilla and R. paricola, all 
described by Thomas on the basis of comparisons with his R. dryas. 

In 1906, Thomas discovered the essential oryzomyine characters of 
the small arboreal rice rats assigned by him to Rhipidomys. The 
close relationship between them and Hesperomys bicolor Tomes was 
also noted. Accordingly, Thomas combined them in Oecomys, a new 
subgenus of Oryzomys. Thomas also included in subgenus Oecomys 
his previously described marmosurus, mamorae, and roberti, each 
based on a local form of the large species of arboreal rice rats. All 
other named forms of arboreal rice rats, including Hesperomys concolor 
Wagner, were ignored or tacitly regarded as species of true Oryzomys. 

The name Oecomys was elevated to full generic rank by Thomas in 
1909. This classification was followed by authors until Ellerman 
(The families and genera of rodents, vol. 2, p. 342, 1941) questioned 
the validity of Oecomys even as asubgenus. It appears that Ellerman 
was misled by the current uncritical listing of some named forms of 
Oryzomys concolor as species of typical Oryzomys, and other named 
forms of O. concolor as species of Cecomys. Thus, his specimens of 
“Oryzomys”’ tectus and ‘‘Oecomys”’ osgoodi, exhibited as proof of the 
absence of important differences between the two “genera” concerned, 
are actually conspecific and identical with Oryzomys concolor. The 
issue is further confused by Ellerman’s observation that ‘“Oecomys”’ 
catherinae Thomas shares critical cranial characters with Oryzomys 
subflavus Wagner. The type of Oryzomys catherinae, however, is not 
an Oecomys (see p. 543) but merely another specimen of Oryzomys 
subflavus Wagner. The proper basis for evaluating the taxonomic 
status of Oecomys is a comparison of its type species, O. bicolor 
Tomes, with O. palustris Harlan, the type species of Oryzomys. 
Neither of these species was taken into account by Ellerman. 


ReEecoGNIZED ForMS OF THE SUBGENUS 


Oryzomys bicolor trabeatus G. M. Allen and Barbour (p. 533). 

Oryzomys bicolor occidentalis, new name for Rhipidomys dryas Thomas 
(=Oryzomys dryas Thomas), preoccupied (p. 533). 

Oryzomys bicolor bicolor Tomes (p. 534). 

Oryzomys bicolor phaeotis Thomas (p. 540). 

Oryzomys concolor concolor Wagner (p. 545). 

Oryzomys concolor speciosus J. A. Allen and Chapman (p. 553). 


ARBOREAL RICE RATS—HERSHKOVITZ 519 


Oryzomys concolor superans Thomas (p. 556). 

Oryzomys concolor roberti Thomas (p. 559). 

Oryzomys concolor bahiensis, new subspecies (replaces Mus cinnamomeus 
Pictet and Pictet, preoccupied) (p. 561). 


Status or Certain Species INcorREcTLY ASSIGNED TO Oecomys 


Ocecomys catherinae Thomas, 1909=Oryzomys subflavus Wagner, 1842. 
(See infra p. 543.) 

Oecomys rez Thomas, 1910=Oryzomys subflavus Wagner, 1842. (See 
infra p. 543.) 

Ocecomys emiliae J. A. Allen, 1916=Rhipidomys emiliae J. A. Allen. 

Ocecomys phelpsi Tate, 1939=Akodon aerosus Thomas, 1913. The 
name phelpsi is based on a skin of what may be either an Oecomys 
or a Lhipidomys (cf. Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, 
p- 194, 1939) and a mismatched skull of Akodon (Chalcomys) 
aerosus. ‘The name phelpsi was restricted to the skull by Tate 
(Journ. Mammal., vol. 26, p. 316, 1945). Oecomys phelpsi Tate, 
therefore, is a synonym of Akodon aerosus. Goodwin (Bull. 
Amer. Mus. Nat. Hist., vol. 102, p. 300, 1953) lists the skin as 
type. Tate’s decision, however, has priority and is irrevocable. 

Mus pyrrhorhinos Wied Neuwied, 1821=Wiedomys pyrrhorhinos 
Wied Neuwied. This small arboreal mouse from eastern Brazil 
was provisionally referred to Oecomys by Osgood (Journ. Mam- 
mal., vol. 14, p. 370, 1933) and is so classified by Goodwin (Bull. 
Amer. Mus. Nat. Hist., vol. 102, p. 300, 1953). Ellerman (Fami- 
lies and genera of living rodents, vol. 2, p. 349, 1941) lists it as 
the sole member of a species group of Oryzomys. It has since 
been shown (Hershkovitz, Proc. Biol. Soc. Washington, vol. 72, 
p. 5, 1959) that the mouse is type of a distinct genus most nearly 
related to phyllotine rodents. 

Mus maculipes Pictet and Pictet, 1844=Rhipidomys maculipes Pictet 
and Pictet. Incorrectly listed as an Oecomys by Tate, Amer. 
Mus. Nov., No. 581, pp. 10, 21 [footnote 1], 1932. 

Hesperomys simpler Winge, 1888=? Winge (EK Museo Lundii, vol. 1, 
No. 3, p. 11, pl. 2, fig. 1, 1888) based the name on the anterior 
portion of a skull found in owl pellets in Recent-Pleistocene de- 
posits in the caves of Lagéa Santa, Minas Gerais, Brazil. Moojen 
(Os roedores do Brasil, p. 55, 1952) treats this species as an 
Oecomys. However, it is obvious from the original description 
and figured skull, and from Winge’s own definition of Hesperomys, 
that simpler is neither an Oecomys nor an oryzomyine. The 
broad forward-projecting zygomatic plate of simplex, its narrow, 
unridged, and rather concave-sided interorbital region, long 
palate, and simplified molars, i.e. without mesoloph, point to 


520 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


phyllotine or sigmodont affinities. A more precise determination 
of the systematic position of Hesperomys simplex cannot be made 
on the basis of available data. 


CHARACTERS 


EXTERNAL CHARACTERS: Body heavy, limbs short and adapted for 
arboreal life; size from approximately that of a house mouse (Mus 
musculus) to that of a medium-sized house rat (Rattus rattus); pelage 
long and thick; dorsal surface from buffy to tawny or rufous, with a 
fine to coarse mixture of dark brown; a poorly defined dark middorsal 
band sometimes present; underparts from sharply defined white to 
dark gray, buff, or ochraceous little or not at all defined from 
sides; an ochraceous lateral line present or absent; ears comparatively 
small, length (from notch) approximately one-half to four-fifths length 
of hind foot (dry, with claw); tail never sharply bicolor, sparsely 
haired except at thickly furred base, tip with or without a conspicuous 
tuft; length of tail from 40 to 60 percent of total length; hind foot 
comparatively short and broad, sole at heel bare but half hidden by 
overlapping hairs of side of tarsus; six plantar tubercles present and 
well developed; first hind toe with claw extending to base of middle 
phalanx of second toe, fifth hind toe well developed for grasping, its 
claw nearly or quite on a level with base of terminal phalanx of fourth 
toe; claws short and recurved, the fifth exposed, the others partially 
concealed by the digital tufts. 

CRANIAL CHARACTERS (pls. 1-3): Skull moderately heavy; sides of 
supraorbital region ridged or beaded and markedly divergent pos- 
teriorward; midtransverse width of paired frontals more than greatest 
width of rostrum; nasals comparatively short; upper anterior corner 
of vertical zygomatic plate rounded and slightly or not at all visible 
when skull is viewed from above; zygomatic plate comparatively 
slender, its anteroposterior width at midpoint not more than alveolar 
length of M’!~; interparietal well developed, its median length vary- 
ing from approximately three-tenths to slightly more than one-half 
its transverse width; sphenopalatine vacuities small, slit-like, or ob- 
solete; incisive foramina moderately long and well opened, length 
more than one-half but less than nine-tenths that of diastema, the 
posterior borders terminating from slightly before to slightly behind 
anterior plane of first molars; auditory bullae small, their antero- 
posterior length, exclusive of tube, less than that of molar row. 

DENTAL CHARACTERS (fig. 3; pl. 4a,b): Upper incisors recurved; 
upper molar rows parallel-sided; inner and outer halves of unworn to 
moderately worn upper molar crowns distinctly crested, of lower 
molars bilevel, i.e., with outer cusps lower than inner cusps; major 
fold well open, a low enterostyle present at its base in M!*, and fre- 


ARBOREAL RICE RATS—HERSHKOVITZ 521 


quently in M®; first minor fold reduced or obsolete in unworn M?, 
obsolete or absent in M?’; anterior border of procingulum of unworn 
first upper molar sometimes notched; mesostyle always fused with 
mesoloph and connected with either paracone or metacone, or both; 
a well-developed paralophule usually present and often extending 
across first secondary fold to connect with mesoloph; primary folds 
often irregular in outline, the second primary usually isolated from 
the margin; first and second internal folds of M'!* discrete or confluent 
with corresponding primary folds, of M® discrete or confluent with 
primary or secondary folds; first secondary fold present in all upper 
molars, usually as an enamel island or as two, sometimes three, is- 
lands separated by the paralophule; second secondary fold present in 
M!~ as an enamel island, in M* as a discrete island, or may be coalesced 
with either second primary or second internal fold, or absent. Pro- 
cingulum of M, subovate in outline, never distinctly bilobate, the 
internal, labial and lingual folds isolated from the margin, discrete 
or coalesced with each other; apex of major fold of M,+ hardly or not 
at all extending beyond midline of tooth; first minor fold well de- 
veloped in all lower molars; ectostylid well developed in all molars; 
ectolophid always present, frequently well developed and fused with 
ectostylid; a well-defined fold between hypoconid and ectolophid 
often present and isolated from the margin; first and second internal 
folds coalesced with corresponding primary folds; first secondary fold 
of M..; obsolete or absent; second secondary fold of M,., well developed, 
usually isolated but clearly defining mesolophid, of M3; obsolete or 
absent; a small entolophulid sometimes present in one or more molars. 


CoMPARISONS 


Subgenus Oecomys is the oryzomyine most specialized for arboreal 
life. Its adaptive characters contrast sharply with those of true 
Oryzomys as represented by its typical species, the palustrine, riparian 
O. palustris Harlan. In palustris, pelage of upperparts is dark and 
glossy, middle digits of hind feet (fig. 4f) long, the outer ones short; 
interdigital webbing nearly as well developed as in the aquatic 
oryzomyine Nectomys; fifth postdigital plantar tubercle reduced or 
lost as in many aquatic Muridae; tail coarsely scutulated and often 
provided with keel hairs. The skull of O. palustris (pls. 1-4) differs 
from subgenus Qecomys chiefly by the prominence of the forward- 
projecting zygomatic plate when viewed from above and by the ex- 
tremely large sphenopalatine vacuities. The molars of palustris and 
subgenus Oceomys (pl. 4) are similar but the lower internal folds are 
usually discrete in those of palustris. Except for the presence of 
intermediate forms, typical Oryzomys and subgenus Oceomys might 
well be treated as generically. distinct. 


022 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 

Some species of the arboreal genus Rhipidomys resemble subgenus 
Oecomys in superficial characters. Both groups have evolved as 
arboreal forms along parallel lines but Rhipidomys is more special- 
ized. Its tail (fig. 56) is more hairy and, in certain species, considerably 
longer; the fifth hind toe is longer and a brown metatarsal patch is 
generally present. The very closely related Nyctomys (fig. 5a) is 
another arboreal cricetine that resembles subgenus Oecomys in external 
characters. Rhipidomys and Nyctomys are members of the pero- 
myscine group of cricetines which differs from oryzomyines chiefly by 
its short palate (pl. 2d) without pitted posterolateral palatal depres- 
sions, some minor dental characters (pl. 4), and by the mammary 
formula, 1—2=6. 

Additional comparisons of subgenus Oecomys are given under the 
individual species headings. 


Some ARBOREAL ADAPTATIONS 


Hinp Foot (fig. 4): The pes of subgenus Oecomys is actually and 
relatively shorter and broader than that of closely related terrestrial 
species of comparable body size. Measurements of the length of the 
hind foot of a series of the small species Oryzomys (Oecomys) bicolor 
are shown in table 1. They may be compared with measurements in 


EXPLANATION OF SYMBOLS IN FicurRE 3 


Names of Cusps 
Upper Molars 
1, Protocone 
2, Hypocone 
3, Paracone 
4, Metacone 
5, Mesoloph 
6, Mesosty] 
7-12, Procingulum 
7, Anteroloph 
8, Anterolabial style 
9, Anteroconule 
10, Anterolingua] style 
11, Anterolabial 
conule 
12, Anterolingual 
conule 
13, Posteroloph (post- 
cingulum) 
14, Enterostyle 
15, Paralophule 
Lower Molars 
1, Protoconid 
2, Hypoconid 


3, Metaconid 
4, Entoconid 
5, Mesolophid 
6, Mesostylid 
7-12, Procingulum 
7, Anterolophid 
8, Anterolingual stylid 
(not shown) 
9, Anteroconulid 
10, Anterolabial stylid 
11, Anterolingual 
conulid 
12, Anterolabial 
conulid 
13, Posterolophid (post- 
cingulum) 
14, Ectostylid 
15, Ectolophid 
16, Entolophulid 


Names of Folds 


mf, major fold 

nf 1, first minor fold 

pf 1, first primary fold 
pf 2, second primary fold 


sf 1, first secondary fold 

sf 2, second secondary 
fold 

if 1, first internal fold 
(shown coalesced 
with pf 1) 

if 2, second internal fold 
(shown coalesced 
with pf 2) 

aif, anterior internal fold 

asf, anterior secondary 
fold (upper first 
molar only) 

alf, anterior lingual fold 
(upper first molar 
only) 

abf, anterior labial fold 
(lower first molar 
only) 

apf, anterior primary 
fold (lower first 
molar only) 


ARBOREAL RICE RATS—HERSHKOVITZ 523 


abf 


FIRST UPPER MOLAR 
BUCCAL VIEW 
Ficure 3.—Molars of Oecomys: a diagrammatic composite of the enamel pattern. The 


basic elements and some of the variations present in the upper and lower molars of Ory- 
zomys (Oecomys) bicolor and O. (O.) concolor are shown. Explanation of symbols on 


opposite page. 


524 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


JN 
iN 


O bicolor’ ©O concolor O. laticeps 


O. meridensis O. palustris 


Ficure 4.—Right hind foot of: a, arboreal Oryzomys bicolor; b, arboreal O. concolor; c, 
semi-arboreal O. subflaous; d, terrestrial O. laticeps; ¢, terrestrial O. meridensis; f, palus- 
trine O. palustris. g, Plantar surface of murid hind foot showing position of pads (1-5= 
postdigital plantar pads; mtp=metatarsal pad). 


ARBOREAL RICE RATS—HERSHKOVITZ 525 


table 2 of a series of the equally small but terrestrial species Oryzomys 
alfaroit. Measurements in tables 3 and 4 are, respectively, of a series 
of the large arboreal Oryzomys (Oecomys) concolor from eastern 
Colombia, and those of a series of the comparably sized terrestrial 
Oryzomys (Oryzomys) laticeps from the same region. Because of 
distortions of the hind{foot in dry specimens, no reliable measurement 
can be given for the width of the metatarsus relative to foot length. 

The hind foot of Gecomys appears short in comparison with terres- 
trial types. Nevertheless, the proportions between metatarsalia and 
digits have probably diverged little from those of the scansorial type 
of primitive cricetines. On the other hand, the greater breadth of 
the metatarsus of subgenus Oecomys (fig. 4a,b) does appear to be 
a specialization for arboreal life. The broader foot provides a more 
powerful and expansive grasp for climbing, perching, and hanging. 
It also permits an effective degree of opposability between the first 
and fifth metatarsals and a considerable amount of adduction and 
abduction in the corresponding toes. 

PLANTAR TUBERCLES: Six plantar tubercles, the primitive number, 
are present and well developed in subgenus Oecomys (fig. 4a,b). The 
first four postdigital pads are large and with little space between them. 
The smaller fifth postdigital pad is also well developed, and is sepa- 
rated from the fourth by a space less than its smallest diameter. The 
metatarsal pad is long and narrow, as usual in cricetines. The function 
of the postdigital plantar tubercles has not been observed in subgenus 
Oecomys. Homer (Contr. Lab. Vert. Gen. Univ. Michigan, No. 61, p. 
13, 1954) states that in scansorial forms of Peromyscus the pads are 
used for grasping. ‘The method was not described but undoubtedly 
slender branches and twigs may be clutched between apposing 
tubercles. 

Criaws: The claws are thick, comparatively short, recurved, their 
tips sharp and raised well above the ends of the toes. These claws 
cannot serve for digging and their position does not interfere with the 
action of the toes as grasping organs. In contrast, claws of closely 
related terrestrial cricetines such as Oryzomys laticeps are slender, 
little curved, and shield the fleshy tips of the toes when used in 
scratching or digging. 

Tait: Modification of this organ as a specialized tool for arboreal 
life is not pronounced in subgenus Qecomys. It is not remarkably 
long and in many individuals is actually shorter than head and body 
combined. The average length, however, is greater than that of head 
and body combined. The tail of subgenus Oecomys appears to be 
slightly thicker than those of comparable sylvan cricetines of pre- 
dominately terrestrial habits. This character indicates greater 
power for balancing, prehension and support in climbing. The 


526 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


C d 


Ficure 5.—Comparative length of hair of terminal portion of tail of: a, arboreal peromy- 
scine, Nyctomys; b, arboreal peromyscine, Rhipidomys; c, arboreal oryzomyine, Oryzomys 
bicolor; d, terrestrial oryzomyine, Oryzomys laticeps. 


pilosity of the tail (figs. 5, 6) varies considerably but is never as great 
as in such related arboreal cricetines as Rhipidomys and Nyctomys. 
The thin pencil and elongated hairs of the terminal one-fourth or 
one-third of the tail are probably sensory and used by the mouse for 
detecting movements in the rear. 

Discussion: Hind foot length of arboreal mice of the subgenus 
Oecomys averages less than 21.5 percent of combined head and body 
length. The hind foot of predominantly terrestrial oryzomyines aver- 
ages more than 21 percent. The arboreal or terrestrial pes could, there- 
fore, have evolved from an ancestral type with average length from 21 
to 21.5 percent of combined head and body length. Specialization of 
the hind foot of the evolving arboreal cricetine is expressed first by a 
lateral broadening of the metatarsus. As the animal becomes increas- 
ingly dependent on its arboreal habitat, the metatarsus becomes 
more flexible, the first and fifth digits more powerful and opposable, 
and the plantar tubercles modified for grasping. In contrast, the hind 
foot of the highly specialized cursorial or saltatorial terrestrial crice- 
tine becomes narrower and longer relative to head and body length, 
the metatarsus rigid, and the first and fifth hind digits reduced. 

A tail length approximately equal to head and body combined 
appears to be the generalized condition in cricetines. In the evolu- 
tion of ambulatory, terrestrial species, notably volelike and fossorial 
forms, the tail becomes shorter. In arboreal, saltatorial and some 


ARBOREAL RICE RATS—HERSHKOVITZ 527 


Ficure 6.—Hair-scale relationships on basal portion of tail in Oryzomys. Hairs of one 
scale only are shown; the same number and disposition of hairs occurs on every scale. 


(X 40.) 


aquatic cricetines, the tail has become longer, at least on an average. 

The degree of specialization of hind foot and tail of cricetines may 
be measured by the amount each organ has diverged from the critical 
proportions given above. Thus, in oryzomyines the longer the hind 
foot than 21.5 percent of combined head and body length, or shorter 
than 21 percent, the more specialized it is. In other cricetines, the 
critical proportions may be a little more or less. Likewise, the longer 
or shorter a tail than combined head and body length, the greater 
the specialization. This, in effect, implies that a specialized short- 
footed species could not have evolved from a specialized long-footed 
form, or vice versa; and that a specialized long-tailed species cannot 
claim a specialized short-tailed species as its ancestor, or vice versa. 

On the basis of the above criteria, Oryzomys (Oecomys) concolor 
is more highly specialized for arboreal life than O. (Oecomys) bicolor. 
Its hind foot is comparatively shorter, its tail proportionately longer. 
The more developed temporal ridges also indicate greater special- 
ization of the masseter muscle. 


HapBits AND Hapsrirats 


The two known species of subgenus Oecomys nest in tree hollows, 
tangled masses of epiphytes or vines, palm fronds, abandoned bird 
nests and in thatched roofs of houses. They normally feed above 
ground, but famine or drought may drive them to the ground in search 


528 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


of food and water. At least the larger species, Oryzomys concolor, 
never lives far from water. I have often shined individuals at night 
running along the maze of tree trunks, vines, logs and even fences 
that connect their arboreal nests with the water’s edge. The smaller 
species is the one most commonly found in houses where arboreal 
habitats are simulated by the palm-thatched roofs and natural timber 
frameworks. A lone mouse or one or two families of either species 
may be permanent residents in the thatched roof or attic of a house. 
On the other hand, many individuals merely visit buildings for feeding 
on insects or man’s food stores. Marauding mice are obliged to cross 
the cleared ground between the forest-edge and the building. 

Oryzomys concolor frequently nests near the ground. It is commonly 
taken in traps placed on logs, along fences and on the banks of streams 
sheltered by shrubs and overhanging vines. OQ. concolor has managed 
to survive in a number of localities (e.g. Villanueva, Colombia) 
where the primitive forest cover has been reduced to small isolated 
stands, scattered trees, shrubs and thinly wooded stream banks. 
It also thrives in plantations of coffee, bananas and other fruits. 
Where living conditions are optimum, Oryzomys concolor tends to form 
colonies and can become excessively numerous. The collector Sala- 
mon Briceiio notes (MS.) that the species does considerable damage 
in coffee plantations in the district of Mérida, Venezuela. 

The smaller species, Oryzomys bicolor, is strictly arboreal and prefers 
to nest high above ground. Although one of the most common native 
house mice in tropical South America, it is not found in areas where 
forests have lost their dominance. It lives in pairs or in single family 
groups and is nowhere known to be abundant. 

Males of Oryzomys concolor exude a musky odor. The females of 
either species of Oecomys produce two to four young in a litter. The 
condition of the mammae in dry skins suggests that young are born 
the year round. 

The name Oecomys (oikos=house-+-mus) was given in allusion to 
the mice’s predilection for houses. In eastern Ecuador where ar- 
boreal rice rats are common, the local Quechua name for QO. concelor 
is “polanda ucucha” (banana rat) and for OQ. bicolor, ‘jahua uchucha.” 
The meaning of ‘“‘jahua”’ is obscure. 


Oryzomys bicolor Tomes 


Disrrisution (fig. 1): Tropical zones of Panama, Venezuela (ex- 
clusive of the Maracaibo basin), Ecuador, the Guianas, the Amazonian 
and upper Rio Paraguay basins of Brazil, and the Amazonian regions 
of Bolivia, Peru, and Colombia; altitudinal range, from near sea level 
to about 2,000 meters above. The apparent disbributional hiatus in 


ARBOREAL RICE RATS—HERSHKOVITZ 529 


northwestern Venezuela and in northern and western Colombia may 
not be real. 

EXTERNAL CHARACTERS: Dorsal surface ochraceous to tawny finely 
mixed with dark brown; underparts sharply defined white or buffy to 
ochraceous, the basal portions of the individual hairs white through- 
out, or pale gray on belly and lateral borders; tail from 40-60 percent 
of total length. 


TaBLE 1.—Oryzomys (Oecomys) bicolor bicolor Tomes: External measurements 
(in millimeters) of a series of adults from the Rio Caquetd region, eastern Colombia. 
Specimens were collected and measured in the flesh by the author. Note proportional 
length of hind foot and compare with table 2. 


Hind foot Hind foot/ Weight 
CNHM No. Tail (dry, c. u.) ore (in grams) 


Percent 
20.9 
21.0 
20.9 
21.1 
22.8 
18.9 


CRANIAL CHARACTERS (pls. 1-11): Sides of supraorbital region 
finely beaded, sometimes narrowly ledgelike; temporal ridges never 
strongly developed anteriorly, weakly defined or obsolete posteriorly; 
anterior zygomatic plate comparatively narrow, its anteroposterior 
width at midpoint always less than alveolar length of M!-’, and fre- 
quently less than length of M?~*; upper anterior corner of zygomatic 
plate not produced forward and barely or not at all visible when skull 
is viewed from above; braincase well inflated, usually convex antero- 
posteriorly at frontoparietal suture, the interparietal steeply arched; 
position of posterior borders of incisive foramina from slightly in front 
of to slightly behind anterior plane of first molars; posterior halves of 
palatines without notable excrescences. 

DENTAL CHARACTERS. Those of the subgenus. 

Comparisons: Ordinarily the difference in size between Oryzomys 
bicolor and O. concolor is sufficient for distinguishing the smaller from 
the larger species. However, a few extremely large individuals of 
O. bicolor may equal or even exceed in size small adults of some popu- 
lations of O. concolor. Where the similarity extends to color of 
underparts, length of hind foot, size of molars and width of anterior 
zygomatic plate, it may be difficult to distinguish an isolated specimen 
of the first species from that of another of the second species. 

504676—59 2 


530 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


TaBLE 2.—Oryzomys (Oryzomys) alfaroi gracilis Thomas: External measurements 
(in millimeters) of a series of adults from Valdivia, Antioquia, central Colombia. 
Specimens were collected and measured in the flesh by the author. Note proportional 
length of hind foot and compare with table 1. 


Head and Hind foot Hind foot/ Weight 
CNHM No. Sex body Tail (dry, c. u.) Head and (in grams) 

body 

Percent 
70507 a 110 119 26 23.7 40 
70508 fot 101 108 25 24.7 40 
70511 fof 114 110 26 22.8 _ 
70513 ou 104 108 26 25.0 30 
70515 a 107 104 25 23.3 40 
70516 fol 122 122 27 pede _ 
70517 foul 104 112 25 24.0 _ 
70518 ofl 107 127 27 25. 2 40 
70509 °) 101 101 24 23.7 — 
70510 °) 106 101 25 23.6 _ 
70512 2 106 101 25 23.5 30 
70514 io) 116 116 26 22.4 _— 
Mean 108 112 25 23.7 37 


Members ofthe Oryzomys longicaudatus group, or subgenus Olig- 
oryzomys, are the only other rice rats that resemble Oryzomys bicolor. 
They are smaller, primarily terrestrial”mice, with pelage compara- 
tively coarse, underparts sometimes pure white, as in 0. bicolor but 
more commonly with the color of the basal portions of the hairs dark 
gray and usually showing through at the surface; tail more slender, 
equal to or longer than combined head and body length, coarsely 
scutulated, nearly naked and without a definable pencil; hind foot 
narrow and comparatively longer, generally more than 23 percent of 
combined head and body length; sole at heel not hidden by over- 
lapping lateral tarsal hairs in the tropical zone forms; first hind toe 
with claw barely extending beyond base of second toe, fifth hind toe 
slender, its claw extending to base of middle phalanx of fourth toe, 
sometimes slightly distad; skull (pl. 7b) more elongate, braincase 
narrower, bullae more inflated; anterior zygomatic plate forward- 
projecting so that one-half its width is visible from above; supraorbital 
region narrow, the sides concave, parallel or slightly divergent pos- 
teriorward and never ridged or beaded; median longitudinal sulcus of 
nasals continuing as a well marked depression over anteromedian 
portion of frontals. 

CoLoR AND PELAGE: Geographic variation in color in Oryzomys 
bicolor is insignificant from a taxonomic point of view. This is 
remarkable in view of the extent and ecological diversity of the area 
occupied by the animal. The usual correlation between coat color 
and color of soil or ground cover apparent in most terrestrial cricetines 


ARBOREAL RICE RATS—HERSHKOVITZ 531 


TaBLeE 3.—Oryzomys (Oecomys) concolor superans Thomas: External measure- 
ments (in millimeters) of a series of adults from the Rio Caquetd region, eastern 
Colombia. Specimens were collected and measured in the flesh by the author. 
Note tail length and proportional length of hind foot and compare with table 4. 


Hind foot Hind foot/ Weight 
CNHM No. bod (dry, c. u.) Head and (in grams) 


Percent 
anit 
18.2 


ou 
ro 
rol 
rol 
oe 
fo 
a 
rot 
fof 
2 
2 
9 
9 
2 
2 
g 
g 
2 
9 


is not evident in arboreal Oryzomys bicolor. The direct relationship 
between amount of rainfall and intensity of coat color, clearly ex- 
pressed in many species of terrestrial and arboreal mammals, is but 
slightly indicated on the underparts of some individuals of OQ. bicolor. 
Temperature throughout the area occupied by O. bicolor is fairly 
uniform and has little or no effect on the length, thickness, texture and, 
indirectly, the color of the pelage. 

The underparts vary from pure white to uniformly ochraceous. 
White underparts may be marked by buff or ochraceous spots, patches 
and streaks. In underparts that appear wholly white at the surface, 
the basal portions of the hairs are generally white except, sometimes, 
along the sides where they may be gray. Where the underparts are 
tinged, the basal portions of the hairs are, as a rule, dark gray. 

In any one population, each adult may be classified as either bright 
or dark according to the color of its upperparts and sides. In the 
bright color group, the terminal portions of the cover hairs are ochra- 
ceous orange with or without a fine dark brown tipping. The 
terminal portions of the guard hairs are dark brown. The colors in 
the dark group are similar but the cover hairs are more uniformly 
tipped with dark brown and there is a greater concentration of 
guard hairs. 


532 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


The difference between the color groups are slight and in any 
large series completely intergrading. Extreme examples of the bright 
color group are nearly erythristic. On the other hand, no specimen 
of the dark group even remotely suggests melanism. 

There are some examples in present material of adults molting 
from one color group to another. 

The color of nestling young is like that of adults. As the juvenal 
coat of cover hairs on upperparts and sides becomes worn, the dark 
eray basal portions of the hairs show through. Adult pelage begins 
to replace juvenal pelage on the lower parts of the sides of the body 
and on the shoulders. Adult pelage appears next on head, nape, and 
shoulders, respectively. Evidently, the short, wholly slate colored 
pelage characteristic of most newborn terrestrial species of Oryzomys 
is either absent or ephemeral in O. bicolor. 


THE SUBSPECIES 


Four subspecies of Oryzomys bicolor are recognized, two of them 
provisionally. The first, bicolor Tomes, occupies most of the range 
of the species east of the Andes. The second, phaeotis Thomas of 
southeastern Peru and northern Bolivia, averages larger in size than 
the others. OQ. 6. trabeatus G. M. Allen and Barbour is the Central 
American representative. It appears to be completely isolated 


TasLeE 4.—Oryzomys (Oryzomys) laticeps Lund: External measurements (in 
millimeters) of a series of adults from the Rio Caquetd region, eastern Colombia. 
Specimens were collected and measured in the flesh by the author. Note tail length 
and proportional length of hind feet and compare with table 3. 


Head and Hind foot Hind foot/ Weight 
bod (dry, c.u) many (in grams) 


rou 
rot 
a 
fol 
fol 
ou 
rou 
rot 
rou 
rol 
ro 
rou 
9 
2 
2 
© 
ie) 
2 
] 


ARBOREAL RICE RATS—HERSHKOVITZ 533 


geographically from its relatives but is not known enough to be 
characterized. 0. 6. occidentalis (new name) of western Ecuador is 
distinguished by a few trivial cranial characters. 


Oryzomys (Oecomys) bicolor trabeatus G. M. Allen and Barbour 


Oecomys trabeatus G. M. Allen and Barbour, Bull. Mus. Comp. Zool., vol. 65, 
p. 262, 1923.—G. M. Allen, Bull. Mus. Comp. Zool., vol. 71, p. 266, 1931 
(type history). 

Oecomys endersi Goldman, Journ. Washington Acad. Sci., vol. 23, p. 525, 1933 
(type locality, Barro Colorado Island, Canal Zone, Panama).—Enders, 
Bull. Mus. Comp. Zool., vol. 78, p. 454, 1935 (Barro Colorado, Panama; 
habits). 

Typr: Adult male, skin and skull (MCZ 19837); collected Apr. 10> 
1922, by Thomas Barbour and W. S. Brooks. 

TYPE Loca.ity: Rio Jesusito, Darién region, eastern Panama. 

DistrisuTion: Known only from the Canal Zone to the eastern 
boundary of Panama. The range may extend into Costa Rica on the 
west and into the Colombian Chocé on the south; altitudinal range 
from sea level to about 500 meters above. 

Cuaracters: Those of the species. See ‘‘Remarks,”’ below. 

MeEasurEMENTs: Those of the type of trabeatus (from the original 
description) are followed by those of the type of endersi: head and 
body, 110, 114; tail, 120, 124; hind foot, 22 (in flesh), 24 (dry); ear, 
15, 15; greatest length of skull, 26.0, 28.1; zygomatic breadth, 15, 
15.1; rostrum, —, 5.4; incisive foramina, —, 4.8; diastema, 7.0, 
7.4; molar row, 4.0, 4.3 (alveolar); width of zygomatic plate, —, 2.5. 

Remarks: Judged by the original description, the type of trabeatus 
is a young individual of the bright color group. It was compared 
with three darkly colored individuals of Oryzomys bicolor from eastern 
Ecuador and said to differ by its ‘somewhat larger’’ size, longer tail 
and longer, thicker pelage. None of these distinctions is valid. 
Retention here of trabeatus for Central American members of the 
common species is based solely on the availability of the name and 
the absence of any records of the occurrence of the species in the 
deciduous forests of northern Colombia. It is also possible that 
additional material from Panama may reveal subspecific differences 
that cannot be accurately evaluated from one specimen. 

The type of enderst Goldman, also from Panama, is an old individual 
of the dark color group. Its size and color are within the range of 
variation of the typical form of the species. 

SPECIMENS EXAMINED: One (UMMZ, type of endersi, from 
Panama). 

Oryzomys (Oecomys) bicolor occidentalis, new name 

Rhipidomys dryas Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 5, p. 271, 1900 
(preoccupied by Oryzomys dryas Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 
2, p. 267, 1898, a member of the subgenus Microryzomys) 


534 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


[Oryzomys (Gicomys)] dryas, Thomas, Ann. Mag. Nat. Hist., ser., 7, vol. 18, 
p. 455, 1906 (“probably = bicolor’’). 

Type: Adult female, skin and skull (BM 99.12.5.4); collected May 
11, 1899, by R. Miketta. 

Type Locauity: Paramba, Rfo Mira, Imbabura, northwestern 
Ecuador; altitude, 1,100 meters above sea level. 

Distrisution: Known from northwestern Ecuador only. No 
doubt the range extends northward into western Colombia. Its 
southward extension may be limited to the Rio Guayas drainage 
system in Manabi, Bolivar, and Guayas Provinces, Ecuador. Its 
altitudinal range is from near sea level to about 1,500 meters above on 
the western slope of the Cordillera Occidental. 

CHARACTERS: Size and external characters as in ¢trabeatus and 
typical bicolor: zygomata less expanded; midtransverse width of 
paired frontals hardly more than greatest width of rostrum and com- 
paratively less than in other races. 

MeEAsuREMENTs: Those of the type of dryas Thomas (from the 
original description) are followed by those of two adults from Pambilar, 
Esmeraldas, Ecuador: head and body, 100, 104, 106; tail, 120, 115 
(dry), 130 (dry); hind foot, 22 (s.w.), 22 (dry, c.w.), 23 (dry, c.u.); 
ear, 14, 12 [?], 15; greatest length of skull, 27.0. 26.4, 27.4; zygomatic 
breadth, 15.0, 13.5, 13.4; rostrum, —, 5.1, 5.3; incisive foramina, 
4.6, 4.6, 4.5; diastema, 7.0, 6.5, 6.4; molar row, 3.8, 4.0 (alveolar), 
4.1 (alveolar) ; width of zygomatic plate, —, 2.2, 2.5. 

Remarks: The three near topotypes at hand are brightly colored. 
One is a juvenal in old pelage but with an irregular patch of bright 
new adult pelage on nape, back, and left side. 

SPECIMENS EXAMINED: Four, from the following localities in 
Ecuador: Paramba, Imbabura, 1 (BM, type of dryas); Pambilar, 
Esmeraldas, 3 (USNM). 


Oryzomys bicolor bicolor Tomes 


Hesperomys longicaudatus, Tomes (not Bennett), Proc. Zool. Soc. London (1858), 
p. 548, 1859 (Gualaquiza, Ecuador). 

Hesperomys bicolor Tomes, Proc. Zool. Soc. London (1860), p. 217, 1860. 

[Hesperomys] M[yoxomys] bicolor, Tomes, Proc. Zool. Soc. London (1861), p. 
284, 1862. 

H{esperomys (Rhipidomys)] bicolor, Thomas, Proc. Zool. Soc. London (1884), 
p. 448, 1884 (doubtfully referred to Rhipidomys). 

Oryzomys (CEcomys) bicolor, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, p. 
445, 1906 (‘‘type. .. described from a discoloured specimen with a broken 
tail’). 

Oryzomys (Oecomys) bicolor, Sanborn, Journ. Mammal., vol. 30, p. 285, 1949 
(Yarinacocha, Loreto, Peru). 

comys bicolor, J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 210, 1916 
(Colombia: Andalucfa, Huila; Florencia, Caquet4).—Thomas, Ann. Mag. 


ARBOREAL RICE RATS—HERSHKOVITZ 535 


Nat. Hist., ser. 9, vol. 19, p. 370, 1927 (Yurac Yacu, San Martfn, Peru, 
“mostly caught in house’’); ser. 10, vol. 2, p. 262, 1928 (San Jerénimo, Loreto, 
Peru). 

Oecomys bicolor, Osgood, Field Mus. Nat. Hist., zool. ser., vol. 10, p. 161, 1914 
(Tambo Yacu, near Rioja, San Martin, Peru).—Gyldenstolpe, Kung]. 
Svenska Vet.-Akad. Handl., vol. 11, p. 39, 1932 (characters; ‘“‘type locality: 
South-western Ecuado:, Guayaquil.’”—Tate, Zoologica, vol. 32, p. 66, 1947 
(Rancho Grande, Aragua, Venezuela, 1,100 meters). 

Rhtpidomys rosilla Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 14, p. 35, 1904 
(type locality, La Unién, lower Rio Caura, Rfo Orinoco, Bolfvar, Venezuela). 

[Oryzomys (Ccomys)] rosilla, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, p. 
445, 1906 (classification). 

Rhipidomys paricola Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 14, p. 194, 1904 
(type locality, Igarapé Agu, on the railway line between Belém de Pard and 
Braganca, Parad, Brazil; altitude, 50 meters). 

[Oryzomys (GEcomys)] paricola, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, 
p. 445, 1906 (classification). 

Gcomys nitedulus Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 505, 1910 
(type locality, lower Essequibo River, 13 miles above mouth, Demerara, 
British Guiana).—Anthony, Zoologica, vol. 3, No. 13, p. 275, 1921 (Kartabo, 
British Guiana). 

CGcomys millert J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 523, 1916 
(type locality, Barfo de Malgacgo, Rio Conguiaru, upper Gy-Paran4, Rio 
Madeira, Mato Grosso, Brazil). 

Gcomys florenciae J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 524, 1916 
(type locality, Florencia, Rfo Orteguaza, upper Rio Caquetd, Caquetd, 
Colombia). 

[?]@comys rutilus Anthony, Amer. Mus. Nov., No. 19, p. 4, 1921 (type locality, 
Kartabo, at junction of Cuyuni and Mazaruni Rivers, lower Essequibo, 
Demerara, British Guiana) Anthony, Zoologica, vol. 3, No. 138, p. 275, 
1921. 

Rhypidomys [sic] benevolens, Ihering (not Thomas), Rev. Mus. Paulista, Sao 
Paulo, vol. 6, p. 420, 1904 (Rio Jurud, Amazonas, Brazil; skull, 28 mm., 
molar row, 4.3). 


Type: Young adult female, skin and skull (BM 7.1.1.96) ; collected 
by Mr. Fraser. 

TYPE LocaLity: Gualaquiza, Rio Gualaquiza, Santiago-Zamora 
Province, southeastern Ecuador; altitude, 885 meters above sea level. 

Distrisution: From the Amazonian region of Brazil north into 
the Guianas, west into the foothills of the Cordillera Oriental in 
Venezuela, Colombia, Ecuador, and the Rios Ucayali and Huallaga 
basins of Peru, south into the upper Rio Paraguay drainage basin in 
Mato Grosso, Brazil; altitudinal range from sea level to possibly 
1,000 meters above. 

CuaractEers: Those of the species. 

MEASUREMENTs: See table 5. 

VARIATION: Six specimens at hand from Zamora are near topotypes. 
Three are juvenals in old pelage with the new adult pelage appearing 
along the sides of the body. The three adults are uniformly colored 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


TaBLe 5.—Oryzomys bicolor bicolor Tomes: Measurements (in millimeters) 


Locality 


COLOMBIA 
Guaicaramo 
Mambita 
Florencia 1! 
Florencia 
Mecaya 
Tres Troncos 
Ino Gaje 


EcuADOR 
Rfo Suno 
Llunehi 
Canelos 
Copataza 
Zamora 
Gualaquiza ? 


PERU 
Curaray 
Rio Mazaén 
Rio Santiago 
Rio Ucayali 3 
Satipo 


VENEZUELA 
Rancho Grande 
La Union 4 


BRITISH GUIANA 
Kartabo 
Kartabo 6 
Essequibo?7 
Holmia 


BRAZIL 
Jauaraté 
Igarap6 Bravo 
Limontuba 
Reereio 
Igarapé-acu 8 
Baréo Melgaco ® 
Bardo Melgaco 


100 

98, 100, 100 
111, 105, 115 
109, 110 


96(85-113)8 

110, 108, 107 
103, 108, 103 
108, 112 


108 
pa 
105, 105 


100 

100, 100, 100 
95, 120, — 
119, 121 


107(98-114)7 
105, 117, 113 
100, 121, 100 
111, 111 


110 
108, =, — 
120, 115 


Hind foot 


22(21.0- 22.5)5 
22 

22 

22, 22, 22 


22, 22, 23 


24 


14, 16, 13 
14, 14 


15(14-16)8 


’ , 


153515 


by italic 


Skull, greatest 
length 


28.2(25.7-29.1)5 


26.5, 27.4, 28.8 
26.5, 26.8, 28.1 
26.9, 27.6, 27.9 
26.6, 27.0, 27.5 
27.6, 28.5 

27.5 


26.5(26.1-27.2)8 
27.3 

27.6 

26.6, 27.5, 27.6 
28.2, 28.8 


26.7 


27.4(26.7-28.1)6 
24.2 
29.0 
25.2 


27.1 
25.5 
24.9 
27.2 
25.0 
24.0 
23.7 


1 Type of florenciae J. A. Allen, from original description. 
skins without skulls. 


3 Type of bicolor Tomes, from original description. 
3 Specimens from Pucallpa, Sarayacu, and Lagarto Alto, respectively. 
4 Type of rosilla Thomas, from original] description, external measurements from the dry 


skin. 


The type and paratypes are 


of adults. 
figure.) 


Zygomatic 
breadth 


14,9(14.5-15.1)4 
14.9 


14.4, 15.4, 15.3 
14,1, 14.9 
13.8 


1440 15.9 
14.1(13.1-14.6)6 
—, 14.8, 15.0 
13.3, 15.5, 14.2 
14.3, 14.9 

14.8 


13.9(13.2-14.5)7 


’ ’ 


14.7. 14.9 


14,2 
15.5 


14.7(13.6-15.5)5 
13.5 


13.0 


5 Without claw. 


ARBOREAL RICE RATS—-HERSHKOVITZ 


Rostrum 
width 


5.1(4.7-5.4)6 
5,3 


4.9, 5.2, 5.1 
4.7, 5.5 
4.8 


4.8, 5.0, 4.7 
5.0(4.4-5.4)8 
4.9, 5.1, 4.9 
4.7, 5.5, 5.0 
5.2, 4.8 


5.0(4.8-5.4)7 
5.0 

b.2 

4.8, 5.1, 5.3 
5.2, 5.7 


5.0(4.6-5.3)6 


4.3 


Incisive 
foramina 


4.8(4.5-5.0)6 
4.9 


5.0, 4.5, 5.1 
4.7, 5.3 
4.8 


4.7, 4.8, 4.7 
4.8(4.3-5.2)8 
eS aigi tok 
4.3, 4.8, 4.9 
5.0, 4.9 


4,5(4.2-4.9)8 
4.6 

4.7 

4.8, 5.1, 4.9 
4.8, 4.7 


4.9(4.6-5.4)6 
3.7 
5.2 
4.6 


4.9 
4.3 
4.1 
4.5 
4.2 
5.0 
4.6 


Diastema 


7.1(6.2-7.6)6 
det 


6.9, 7.4, 7.4 
6.8, 7.4 
7.0 


6.5, 7.0, 7.2 
6.8(5.5-7.5)8 
One 
6.5, 6.6, 6.9 
7.5, 7.4 


6.5(5.8-7.0)8 
6.8 

6.9 

6.9, 7.3, 7.4 
7.2, 7.3 


7.0(6.6-7.5)6 


6.5 


6 Type of rutilus Anthony, from original description. 
7 Type of nitedulus Thomas, from original description. 
8 Type of paricola Thomas, from original description. 
§ Type of milleri J. A. Allen, from original description 


Alveolar 
length of 
molar row 


4.2(4.0-4.4)6 
4.3 


3.8, 4.6, 4.2 
4.3, 4.2 


3.9, 4.3, 4.2 
4.1(4.0-4.3)8 
4.0, 4.0, 4.0 
4.4, 4.1, 3.8 
3.9, 4.1 


4.1(4.0-4.3)8 
4.1 

3.8 

3.9, 4.1, 3.9 
4.1, 4.1 


3.9(3.8-4.2)6 
3.4 
3.7 
3.9 


(Number of specimens, when more ihan three, indicated 


Zygomatic 
plate width 


2.5(2.3-2.7)6 
2.4 


2.5, 2.6, 2.3 
2.3, 2.4 

2.4 

2.1, 2.2, 2.5 
2.3(1.8-2.5)8 
2.2, 2.5, 2.4 
IODINE 
2.3, 2.4 
2.2(1.9-2.4)9 
2.5 

2.6 

2.8, 2.2, 2.2 
2.5, 2.4 

2.4 


537 


538 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


except for an ochraceous streak on each side of the white underparts 
. In one. 

In five individuals from Canelos taken March 8-12, 1924, the 
underparts are faintly tinged with buff; in three from the same locality, 
taken in April 1910, and in two from the Rio Copataza and another 
from Montalvo, the underparts are pure white. The three localities 
mentioned are within a few miles of each other in the upper Rfo 
Pastaza region. 

Specimens from Jollin, Jatun Yacu, Llunchi, Avila, Concepcién, 
San José and Rio Suno, all in the upper Rio Napo region, show the 
same range of variation in color as the preceding. In one specimen 
from Llunchi, however, the belly is uniformly ochraceous and hardly 
defined from the sides. 

The series of 11 specimens from the Rio Curaray, Loreto, Peru, 
includes individuals in dark and bright pelage. One is nearly 
uniformly ochraceous orange on upperparts and sides. Underparts 
are white. One specimen has a pair of small ochraceous gular patches. 

One of four specimens from Lagarto Alto, Rio Ucayali, Peru, in old 
pelage, is nearly uniformly ochraceous tawny on head and back. This 
color is defined in the form of a molt line from the darker outer sides 
of shoulders and thighs. The other three specimens of the series are 
dark in unworn pelage. Underparts, upper lips and lower half of 
cheeks in one of two mice from Satipo, Junin, are yellowish. Under- 
parts of the other specimen are sharply defined white, the lips and 
cheeks ochraceous. 

The remaining Peruvian material consisting of one or two individuals 
from scattered localities reveal no peculiarities. 

The type series of Oecomys florenciae J. A. Allen, from eastern 
Colombia, is practically indistinguishable from typical bicolor as 
represented by any series from eastern Ecuador. Other Colombian 
specimens at hand exhibit no individual or local variables of note. 

Judged by the original description, the only distinguishing character 
of the Orinocoan “Rhipidomys”’ rosilla Thomas is the “buffy ochra- 
ceous”’ belly of the type as compared with the pure white belly of the 
type of dryas (=occidentalis) of western Ecuador. A colored belly, 
however, is a common enough character in bicolor of eastern Ecuador. 
In the absence of other distinctions, rosilla must be regarded as 
identical with typical bicolor. 

Ocecomys nitedulus Thomas of British Guiana is said to be “quite like 
the Orinoco Qe. rosilla above, but the undersurface is without the 
marked ochraceous wash characteristic of that animal.’ This 
characterization implies that nitedulus is like typical bicolor. Indeed, 
our material from British Guiana is altogether indistinguishable from 
eastern Ecuadorian bicolor. I have not examined the type of Oecomys 


ARBOREAL RICE RATS—HERSHKOVITZ 539 


rutilus Anthony. Judged by the original description, it is a subadult 
of the same Kartabo series identified by Anthony as nitedulus. 

The description of paricola Thomas, from the state of Pard, Brazil, 
like that of rosilla, was based on a comparison with western rather 
than with eastern Ecuadorian representatives of bicolor. It is said 
to be ‘“‘duller” in color and smaller. Of two specimens at hand from 
the Rio Tapaj6z, Pard, one, an adult from Limontuba, is as small 
cranially as the type; the other, from Igarapé Bravo, is a subadult 
with a much larger skull. The color of either, whether dull or bright, 
is like that of bicolor from eastern Ecuador or anywhere else. 

Oecomys milleri J. A. Allen, from Baréo de Malgacgo, Mato Grosso, 
Brazil, was aptly diagnosed in the original description as “similar in 
general coloration and size to @. bicolor (Tomes).”’ A careful exami- 
nation of the type, two paratopotypes, a specimen from Urupé and 
another from Urucum de Corumbé confirms this characterization. 
The Corumba specimen is bright ochraceous on upperparts, pure 
white beneath. The topotypes are darker, with underparts yellowish 
in one, pure white in the other. The Urupd individual is darkest, 
tawny above, more buffy beneath. 

SPECIMENS EXAMINED: One hundred and seventeen, from the 
following localities: 


Cotomsia: Guaicaramo, Meta, 6 (AMNH); Mambita, eastern Cundinamarca, 
1 (AMNH); Andalucia, Huila, 1 (AMNH); Florencia, Rfo Orteguaza, Caquetd4, 
5 (including type of florenciae, AMNH); Tres Troncos, Rio Caquetd4, Caquetd, 
3 (CNHM); Rio Mecaya, mouth of Rio Caqueté, Putumayo, 3 (CNHM); Ino 
Gaje, Rio Apoporis, Vaupés, 4 (CNHM). 

Kcuapor: Below San José, Rio Payamino, Napo-Pastaza, 2 (AMNH); Rio 
Suno, Rio Napo, Napo-Pastaza, 3 (AMNH); Avila, Rio Suno, Napo-Pastaza, 
1 (USNM); Rio Jollfin, Napo-Pastaza, 2 (MCZ); Rio Jatun Yacu, Napo-Pastaza, 
1 (MCZ); Llunchi, Rio Napo, Napo-Pastaza, 8 (UMMZ); Canelos, Rio Bobo- 
naza, Napo-Pastaza, 8 (AMNH, 5; MCZ, 3); Montalvo, Rfo Bobonaza, Napo- 
Pastaza, 1 (CNHM); Rfo Copataza, Napo-Pastaza, 3 (CNHM, 2; AMNH, 1); 
Zamora, Santiago-Zamora, 6 (AMNH). 

Peru: Rio Curaray, mouth at Rfo Napo, Loreto, 11 (AMNH); Monte Alegre, 
Loreto, 1 (AMNH); Sarayacu, Rfo Ucayali, Loreto, 2 (AMNH); Yarinacocha, 
Rfo Ucayali, Loreto, 2 (CNHM); Pucallpa, Rio Ucayali, Loreto, 1 (CNHM); 
Lagarto Alto, Rio Ucayali, Loreto, 4 (AMNH); Rfo Santiago, mouth of Rfo 
Marafion, 1 (AMNH); Puerto Indiana, Rio Marafion, Loreto, 1 (AMNH); Rio 
Mazén, Loreto, 1 (AMNH); Tambo Yacu, Rioja, San Martin, 1 (CNHM); 
Chanchamayo, Junin, 1 (CNHM); Satipo, Junin, 3 (MCZ); Pozuzo, Huanuco, 
1 (CNHM). 

British Gurana: Kartabo, Demerara, 12 (AMNH); Holmia, Potaro High- 
lands, 1 (CNHM); Barakara, Canje River, 1 (AMNH); Essequibo River, 1 (BM, 
type of nitedulus). 

VENEZUELA: La Unién, Bolivar, 1 (BM, type of rosilla); Rancho Grande, Ara- 
gua, 1 (AMNH); Caifio Seco, Cerro Duida, Amazonas, 1 (AMNH). 

Braziu: Jauaraté, Rio Uapés, Amazonas, 2 (AMNH); Igarapé Bravo, Rio 
Tapajéz, Pardé, 1 (AMNH); Limontuba, Rio Tapajéz, Pard, 1 (AMNH); between 


540 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Belem and Braganza, Pard, 1 (BM, type of paricola); Recreio, Rio Majary, 
Amazonas, 1 (AMNH); Urucum de Corumb4, Mato Grosso, 1 (CNHM); Baraéo 
Malgacgo, Mato Grosso, 3 (including type of milleri, AMNH); Urupd, Mato 
Grosso, 1 (AMNH). 


Oryzomys bicolor phaeotis Thomas 


Rhipidomys phexotis Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 7, p. 181, 1901. 

[Oryzomys (Gicomys)] phxotis, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 7, 
p. 445, 1906 (classification). 

Oryzomys (Oecomys) phaeotis, Sanborn, Publ. Mus. Hist. Nat. “Javier Prado,’ 
ser. A, zool., No. 6, p. 21, 1951 (Peru: Marcapata, Cuzco; Camante, Cuzco). 

Rhipidomys benevolens Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 7, p. 369, 1901 
(type locality, Chimate, upper Rfo Beni, La Paz, Bolivia; altitude, 700 
meters above sea, level). 

[Oryzomys (Gicomys)] benevolens, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, 
p. 445, 1906 (classification). 

Oecomys benevolens, Gyldenstolpe, Kungl. Svenska Vet.-Akad. Handl., ser. 3 
vol. 11, p. 40, pl. 4, fig. 1 (skull), pl. 17, fig. 11 (molars), 1982. 


Types: Adult male, skin and skull (BM 1.1.1.23); collected July 23, 
1900, by Perry O. Simons. 

ison Locality: ‘Segrario”’ pacer upper Rio Inambari, an 
affluent of the Rio Madre de Dios, Puno, southeastern Peru. 

DistriputTion: Departments of Cuzco, Puno, and Madre de Dios 
in southeastern Peru, and Departments of La Paz, Cochabamba, and 
Beni in northwestern Bolivia; altitudinal range from approximately 
200 to 2,000 meters above sea level. 

Cuaracters: Largest of the subspecies; molars proportionately 
larger. 

MEASUREMENTs: See table 6. 

Remarks: The available Peruvian specimens of phaeotis, including 
the type and a paratype, are from four localities within a compara- 
tively small area in the drainage basin of the upper Rio Inambari, an 
affluent of the Madre de Dios. Two from Camante, Marcapata, 
taken August 1950, are as large as the type of phaeotis collected July 
1900. An individual from Quincemil, collected August 1953, and 
another from Villa Carmen, Cosfiipata, dated September 1954, agree 
with typical bicolor in size. No measurements of the paratype were 
given but Thomas declared it to be ‘about the same general size” as 
the type. The large Camante specimens, judged by cranial charac- 
ters, are not older than the others and the molars are slightly less worn. 

Rhipidomys benevolens Thomas, from the upper Rio Beni, Bolivia, 
was characterized as larger than dryas (occidentalis) and smaller 
than phaeotis. Representative specimens from the upper Rio Mamoré 
include one subadult and three juvenals. They agree with phaeotis 
in their over-all larger size and proportionately larger molars as com- 
pared with typical bicolor of northeastern Peru and eastern Ecuador. 


541 


ARBOREAL RICE RATS—HERSHKOVITZ 


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042 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


A skin without skull from Ticunhuaya, Bolivia, not far from the 
type locality of benevolens, appears to belong here. 

The largest specimens of O. bicolor phaeotis are practically in- 
distinguishable from small adults of O. concolor from other parts 
of tropical America. On the other hand, the size difference between 
O. bicolor phaeotis and O. concolor of the same general region leaves 
no doubt regarding the separate identity of the two species. 

SPECIMENS EXAMINED: Eleven, from the following localities: 

Peru: Sagrario, Rio Inambari, Puno, 1 (BM, type of phaeotis); Hacienda Villa 
Carmen, Cosfiipata, Cuzco, 1 (CNHM); Camante, Marcapata, Cuzco, 2(CNHM); 
Quincemil, Marcapata, Cuzco, 1 (CNHM). 

Bouivia: Chimate, La Paz, 1 (BM, type of benevolens); El Palmal, Chaparé, 
Cochabamba, 1 (MACN); Marbdn, Rio Mamoré, Beni, 3 (MACN); Ticunhuaya, 
La Paz, 1 (AMNB#). 

Oryzomys concolor Wagner 

Distrinpution (fig. 2): Tropical and subtropical forested zones of 
Costa Rica, Panama, Colombia, Venezuela, Trinidad, the Guianas, 
Brazil, and the Amazonian regions of Bolivia, Peru, and Ecuador; 
altitudinal range from near sea level to approximately 2,000 meters 
above. 

EXTERNAL CHARACTERS: Dorsal surface buffy to tawny or rufous, 
with a fine to coarse mixture of dark brown; a poorly defined dark 
middorsal band sometimes present; underparts sharply or hardly 
defined from sides, basal portions of individual hairs white or gray; 
tail from 49 to 60 percent of total length. 

CRANIAL CHARACTERS (pls. 1-6, 8-11): Sides of supraorbital region 
prominently ridged, often ledgelike; temporal ridges well developed 
in adults for entire length of parietals; anterior zygomatic plate 
moderately broad, its anteroposterior width at midpoint. usually less, 
rarely equal to or slightly more, than alveolar length of M'?; upper 
anterior corner of zygomatic plate slightly projecting; braincase 
moderately inflated, flat or slightly convex anterodorsally at fronto- 
parietal suture; posterior borders of incisive foramina usually termi- 
nating anteriad to, rarely slightly behind, anterior plane of first molars; 
posterior half of palatines often marked by a pair of small transverse 
cornuate processes, one on each bone (one or both processes are 
frequently lost in the cleaning operation but the median burr or 
rugosity remains). 

DENTAL CHARACTERS (pls. 46, 126): Essentially as in O. bicolor but 
with first and second internal folds of M'!~? more frequently confluent 
with corresponding primary folds. In extremely worn teeth, however, 
these folds may be secondarily discrete. 

Comparisons: Oryzomys concolor is essentially an outsize O. bicolor. 
In most localities where the two species are sympatric the difference 
in size is obvious. On the other hand, many small northern represen- 


ARBOREAL RICE RATS—HERSHKOVITZ 543 


tatives of concolor are hardly larger than some large southern individ- 
uals of O. bicolor (pl. 5). In the Guianas, size alone is not a decisive 
character. Generally, the heavier more “adult” skull of concolor 
with its better developed supraorbital and temporal ridges, less inflated 
braincase, and more gray or buff underparts are characters that help 
separate the larger from the smaller species where the two occur 
together. 

Representatives of Oryzomys subflavus Wagner® of Brazil and the 
Guianas have sometimes been confused with QO. concolor. The two 
species are similar in color, size, and taillength. In subflavus, however, 
the pelage is comparatively lax and harsh; hind foot (fig. 4c) long, 
stout, with outer digits shorter, first hind toe with claw barely ex- 
tending beyond base of second toe, fifth hind toe with claw extending 
only to base of middle phalanx of fourth toe; sole of heel exposed; 
excision of dorsal border of antorbital foramen deep, one-half or more 
of width of anterior zygomatic plate exposed to view when skull is 
viewed from above; supraorbital region of skull (c of pls. 8-11) broad, 
its sides ridged or beaded and divergent as in concolor; zygomata less 
expanded anteriorly; incisive foramina generally narrower but with 
their posterior borders not extending behind anterior plane of first 
molars; sphenopalatine vacuities more open but less so than in QO. 
zanthaeolus (compared below); bullae more inflated; enamel pattern 
of molars (pl. 12¢) essentially as in Oecomys. 

Oryzomys xanthaeolus Thomas is a semiarboreal species with a 
superficial resemblance to O. concolor. It occupies the dry coastal 
area of western Peru and southwestern Ecuador where Oecomys is not 
known to occur. Its pelage is comparatively lax, upperparts and 
sides of body buffy or olivaceous, lateral line absent, underparts 
not pure white; tail long as in concolor but untufted and bicolor for 
most or all its length; hind foot broader than in strictly terrestrial 
species but narrower than in concolor, the outer toes shorter and less 
robust, interdigital webbing more developed; general shape of skull 
(d of pls. 8-11) and sides of supraorbital region as in concolor but 
anterior zygomatic plate projected forward so that at least one-half 
its width is visible from above; incisive foramina longer, their length 
at least three-fourths that of diastema, and terminating posteriorly 
well behind anterior plane of first molars; sphenopalatine vacuities 
large, as in O. palustris (pl. 2c), posterior border of palate without 
excrescences; bullae well inflated ; enterostyles and ectostylids (pl. 12d) 
rudimentary or absent. 


3 “Oecomys’’ catherinae Thomas is a synonym of Oryzomys subflavus. ‘‘Oecomys’’ rer Thomas, 1910, of 
British Guiana, is also a member of the same species but probably subspecifically distinct. The name, 
however, clashes with ‘‘Calomys” [= Oryzomys] rer Winge, 1888. Because of the homonymy, the name of 
the form described by Thomas may be changed to Oryzomys subflavus regalis (new name). 


544 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Oryzomys laticeps * is a terrestrial species ranging coextensively 
with Oecomys. Its resemblance to Oryzomys concolor is superficial 
and restricted to body size and, in some cases, to color and texture of 
pelage. In laticeps, however, the tail averages shorter, the hind foot 
(fig. 4d) longer, the ear larger. The outer toes of the hind foot of 
O. laticeps are shorter than in all others with which comparisons have 
been made. A dark juvenal color-pelage phase and a distinct sub- 
adult color-pelage phase are well marked and more consistently 
present in laticeps than in other oryzomyines. The skull (e of pls. 
8-11) of O. laticeps is comparatively longer than in concolor, the 
supraorbital region narrower, the sides ridged or beaded but not as 
widely divergent; rostrum with a more decided downward flexion; 
anterior zygomatic plate projecting well forward; anteroposterior 
width of zygomatic plate often more than length of M!~?; incisive 
foramina extremely short, generally less than 60 percent of length of 
diastema, their posterior borders separated from anterior plane of 
first molars by a distance approximately equal to length of first molar; 
sphenopalatine vacuities absent or small as in concolor; bones of 
posteropalatal region more cancellate with the excrescences or cornu- 
ate processes noted in concolor generally present and, usually, more 
developed; internal folds of molars (pl. 12¢) usually absent; second 
secondary fold absent in M’; second secondary folds of lower molars 
greatly reduced, sometimes coalesced with major fold of opposite side. 

CoLor AND PELAGE: The range of variation in color and character 
of pelage in Oryzomys concolor is wider than in O. bicolor. Populations 
of dry localities are distinctly paler than those of humid areas. The 
pelage of high altitude forms is invariably longer and thicker than that 
of their relatives of lower levels. As in O. bicolor, there are bright and 
dark color phases. The tone of the bright phase in a darkly colored 
population generally equals that of the dark phase of a pale colored 
population. In concolor, individual molt from one color phase to the 
other is obvious in a large proportion of the specimens. 

Pelage of nestlings is short, fine, and colored like that of adults. 
This is superseded by a longer dark gray or dark brown coat distinctly 
darker than that of adults. 


THE SUBSPECIES 


Five geographic races are recognized. The nominate subspecies, 
Oryzomys concolor concolor Wagner, occupies the greatest part of the 


4 Specific synonyms of Oryzomys laticeps include bolivaris J. A. Allen, boliviae Thomas, caracolus Thomas, 
casteneus Thomas, goeldi Thomas, legatus Thomas, macconnelli Thomas, magdalenae J. A. Allen, medius 
Robinson and Lyon, modestus J. A. Allen, mollipilosus J. A. Allen, oniscus Thomas, perenensis J. A. Allen, 
rivularis J. A. Allen, saltator Winge, sylraticus Thomas, talamancae J. A. Allen, velutinus J. A. Allen, 
and a few others. Afus capito Olfers (1818), based on the “rat & grosse tate” of Azara, is undoubtedly the 
same species and antedates Mus laticeps Lund (1841). Afus cephalotes Desmarest (1819) is another name 
for Azara’s “‘rat & grosse téte.” 


ARBOREAL RICE RATS—HERSHKOVITZ 545 


range of the species. It is a medium-sized, brightly colored form 
with an irregular pattern of geographic variation that reflects seasonal 
and local climatic differences. Nevertheless, individual variation in 
any one locality may be of the same magnitude as variation within the 
subspecies as a whole. 

The remaining four subspecies occupy peripheral and ecologically 
distinct parts of the range of the species. Oryzomys concolor superans 
Thomas, of the Amazonian rain forests along the base of the Andes, is 
the darkest and largest race. O.c. speciosus J. A. Allen and Chapman 
of the northeastern bulge of South America resembles superans in 
color, especially of the underparts, but agrees with typical concolor 
in size. The palest subspecies, 0. concolor robertt Thomas, lives in the 
subhumid southern border of the range in Bolivia and south-central 
Brazil. O. concolor bahiensis (new subspecies) is a saturate form of the 
rain forests of eastern Brazil. 

The pattern of local and geographic variation in Oryzomys concolor 
contrasts sharply with the comparative stability of O. bicolor. The 
difference between the two species in their response to color of soil 
and cover, humidity and temperature may be attributed to slight but 
significant differences in their habitat preferences. Oryzomys concolor 
generally nests lower and descends to the ground with greater fre- 
quency. It is, accordingly, less independent of factors controlling 
living conditions at ground level. 


Oryzomys (Oecomys) concolor concolor Wagner 


Hesperomys concolor Wagner, Arch. Naturg., vol. 11, pt. 1, p. 147, 1845; Abh. 
Akad. Wiss., Munich, vol. 5, p. 311, 1848 (characters) —Pelzeln, K. K. Zool.- 
Bot. Gesellsch., Wien, Beiheft 33, p. 70, 1883 (characters; habits).—Gylden- 
stolpe, Kungl. Svenska Vet-Akad. Handl., ser. 3, vol. 11, p. 151, 1932 (in- 
certae sedis).—J. A. Allen and Chapman, Bull. Amer. Mus. Nat. Hist., 
vol. 5, p. 213, 1893 (comparison with Oryzomys speciosus ‘‘with which it may 
prove to be identical’’). 

[Oryzomys] concolor, Tate, Amer. Mus. Nov., No. 579, p. 3, 1932 (taxonomic 
history). 

Oryzomys flavicans Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 14, p. 351, 1894 
(type locality, Mérida, Sierra de Mérida, Venezuela). 

O[ryzomys] flavicens [sic], Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 102, p. 310, 
1953 (lapsus for flavicans Thomas, in discussion of O. palmarius J. A. Allen). 

Oryzomys flavicans illectus Bangs, Proc. Biol. Soc. Washington, vol. 12, p. 164, 1898 
(type locality, Pueblo Viejo, Sierra Nevada de Santa Marta, Magdalena, 
Colombia, altitude, 853 meters.—Bangs, Proc. New England Zool. Club, 
vol. 1, p. 94, 1900 (Colombia: La Concepcién; Pueblo Viejo; Palomino; 
San Antonio; Oryzomys trichurus J. A. Allen, part, a synonym). 

O[ryzomys] flavicans illectus, J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 12, 
p. 206, 1899 (Minca, Santa Marta Region, Magdalena, Colombia). 

Oryzomys illectus, J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 20, pp. 436, 439, 
1904 (Don Amo, Magdalena, Colombia). 

504676—59 —_3 


546 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Oryzomys flavicans subluteus Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 2, p. 268, 
1898 (type locality, ‘‘W. Cundinamarca,” i.e., western slope of Cordillera 
Oriental, Cundinamarca, Colombia). 

O[ryzomys] subulatus [sic], Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 102, p. 302, 
1953 (lapsus for subluteuws Thomas, in discussion of Oryzomys helvolus). 

Oryzomys trichurus J. A. Allen, Bull..Amer. Mus. Nat. Hist., vol. 12, p. 206, 1899 
(type locality, El Libano, near Bonda, Santa Marta, Magdalena, Colombia, 
altitude 500 feet); vol. 20, p. 487, 1904 (Bonda, Colombia).—Tate, Bull. 
Amer. Mus. Nat. Hist., vol. 76, p. 190, 1939 (‘“‘juv. and synonym of illectus’’).— 
Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 102, p. 301, 1953 (type history). 

Rhipidomys marmosurus Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 4, p. 378, 1899 
(type locality, Maipures, middle Rfo Orinoco, Vichada, eastern Colombia). 

[Oryzomys (Cicomys)] marmosurus, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, 
p. 445, 1906. 

Oecomys marmosurus, Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 194, 1939 
(listed). 

G@[comys] marmosurus, Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 187, 1910 
(comparisons). 

Oryzomys tectus Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 8, p. 251, 1901 (type 
locality, Bugaba (misspelled ‘‘Bogava”’), Chiriqui, Panama, 250 meters alti- 
tude). 

Oryzomys tectus tectus, Goldman, North American Fauna, No. 43, p. 84, 1918 
(revision; Boruca, Costa Rica)—Goodwin, Bull. Amer. Mus. Nat. Hist., 
vol. 87, p. 394, 1946 (Costa Rica: Boruca, Puntarenas; San Gerénimo de 
Pirris, San José). 

Oryzomys klagest J. A. Allen, Bull. Amer..)Mus. Nat. Hist., vol. 20, p. 827, 1904 
(type locality, El Llagual [Yagual], near Maripa, lower Rfo Caura, Bolivar, 
Venezuela).—Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, pp. 190, 191,1939 
(listed as member of “trinitatis group”? of Oryzomys (p. 190)).—Goodwin, 
Bull. Amer. Mus. Nat. Hist., vol. 102, p. 301, 1953 (type history; ‘‘probably a 
subspecies of O. flavicens [sic] Thomas’’). 

Rhipidomys (?) klagest, Gyldenstolpe, Kungl. Svenska Vet.-Akad. Handl., ser. 3, 
vol. 11, p. 50, 1932 (type a Rhipidomys [!] according to Tate, ex litt). 

Ccomys tapajinus Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 3, p. 378, 1909 
(type locality, Santa Rosa, Rio Jamanchin, Rio Tapajéz, Pard, Brazil) ; 
ser. 8, vol. 9, p. 87, 1912 (Marajé6, Pard, Brazil); ser. 9, vol. 6, p. 277, 1920 
(Brazil: Villa Braga, Rio Tapajéz, Par4; Monte Alegre, Rio Solimées, Para; 
Manacapurt, Rio Solimées, Amazonas). 

Oryzomys frontalis Goldman, Smithsonian Mise. Coll., vol. 56, p. 6, 1912 (type 
locality, Corozal, Canal Zone, Panama). 

Oryzomys tectus frontalis, Goldman, North American Fauna, No. 43, p. 85, pl. 4, 
figs. 4, 4a (skull), pl. 5, fig. 10 (mandible), pl. 6, figs. 7, 7a (molars), 1918, 
(Panama: Cana; Corozal, Canal Zone). Smithsonian Mise. Coll., vol. 69, 
p. 101, 1920 (Panama: Cana; Corozal; Tacarcuna). 

Oryzomys helvolus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 597, 1913 
(type locality, Villavicencio, upper Rfo Meta, Meta, Colombia); vol. 35, p. 
212, 916 (Colombia: Villavicencio; Buenavista—Goodwin, Bull. Amer. 
Mus. Nat. Hist., vol. 102, p. 302, 1953 (‘probably a subspecies of O. flavicens 
[sic]; may equal O. subulatus [sic] Thomas’’). 

Oryzomys vicencianus J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 598, 
1913 (type locality, Villavicencio, Meta, Colombia); J. A. Allen, vol. 35, p. 
212, 1916 (Villavicencio, Colombia) —Goodwin, Bull. Amer. Mus. Nat. 
Hist., vol. 102, p. 302, 1953 (type history and measurements). 


ARBOREAL RICE RATS—HERSHKOVITZ 547 


[Oryzomys] vicencianus, Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 190, 1939 
(‘juv. and synonym of helvolus’’). 

Ccomys mincae J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 603, 1913 
(type locality, Minca, Santa Marta region, Magdalena, Coiombia).—Good- 
win, Bull. Amer. Mus. Nat. Hist., vol. 102, p. 301, 1953 (type history). 

Cicomys caicarae J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 32, p. 603, 1913 
(type locality, Caicara, Rio Orinoco, Bolivar, Venezuela). 

Oecomys caicarae, Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 102, p. 300, 1953 
(type history and measurements). 

Oecomys guianae caicarae, Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 193, 1939 
(Venezuela: Mt. Duida; Casiquiare). 

Oryzomys trinitatis, Tate (not Allen and Chapman), Bull. Amer. Mus. Nat. Hist., 
vol. 76, p. 190, 1939 (part; Venezuela: Agitita; Mt. Duida). 


Type: Presumably in the Vienna Natural History Museum; col- 
lected Aug. 4, 1831, by Johann Natterer; original number, 174. 

Type Locauiry: Rio Curicuriari, an affluent of the upper Rio 
Negro, entering from the right below Sao Gabriel, Amazonas, Brazil. 

DistrisuTion: Tropical and subtropical forested zones of Costa 
Rica, Panama, northwestern Colombia, and eastern Colombia in 
the Rio Orinoco drainage system, western and southern Venezuela in 
the Lake Maracaibo and the middle and upper Rio Orinoco basins, 
the Amazonian rain forest of Brazil and their extensions into extreme 
northern Bolivia, eastern Peru, and eastern Ecuador; altitudinal 
range, sea level to approximately 2,000 meters above. 

CHARACTERS: Generally brightly colored, upperparts ochraceous 
buff to tawny finely ticked with dark brown, chest and belly white to 
buff, throat and chin white. 

MEASUREMENTS: See table 7. 

VARIATION: Two specimens from the type region are at hand. 
One, from Jauaraté, Rio Uaupés, above the type locality, is larger 
than the type, its belly with a light buffy wash. The other, from 
Yauanari, Rio Negro, below the type locality, is smaller, with belly, 
chest, and sides ochraceous buff. Underparts of the type are sharply 
defined white. The same range of variation in color of underparts 
obtains in most populations of the species. 

The type of Oecomys tapajinus Thomas and two other specimens 
from Tauary, in the type region on the Rio Tapajéz, Brazil, do not 
differ from concolor of the Rio Negro-Rio Orinoco basin. The under- 
parts of one of the two Tauary specimens is creamy, most notably on 
chest, throat, and chin; in the other, the same parts are sharply defined 
white. A specimen from Lago Cuiteud, on the opposite side of the 
SolimGes, is practically identical. A young individual from the Boca 
do Igarapé Piaba, also on the north bank of the Solimées, shows the 
old pelage being replaced by a more saturate new pelage. 

The type of marmosurus Thomas, from the middle Rio Orinoco, 
Colombia, is not significantly different from any other individual of 


548 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


TaBLE 7.—Oryzomys concolor concolor Wagner: Measurements (in millimeters) 


Locality Head and body Tail Hind foot Ear mere 
Costa RICA 
San Gerénimo 125 149 28 — 34.5 
E1 General 146 142 27 — 32.6 
PANAMA 
Boruca 118, 125 140, 123 27, 28 _ 31.0, 31.1 
Bugaba ! 140, 122 142, 147 27, 29 18, — —, 33.2 
Cana § 148 161 30 — 35.5 
Cana 126 (114-137)6 148(127-162)6 28 (26-29)6 | — 31.4 (28.8-34.0)7 
COLOMBIA 
Socorré 129, 130, 143 154, 140, 154 28, 28, 29 18, 17, 18 33.7, —, 33.9 
Catival 132 146 28 17 31.4 
Muzo 128 140 29 16 31.8 
Pueblo Viejo ¢ 132 160 25 5 17 _- 
Pueblo Viejo 136(122-145)6 145(140-155)5 26(25-27)5 | 17(15-18)5 30.4(29.8-31.1) 4 
Palomino 140 145 26 16 32.3 
Don Diego 127(111-137)8 149 (136-165)8 26(25-28)8 | — 31.6(30.5-32.3)5 
Bonda 120 155 _ _ 28.8 
E] Lfbano 6 121 160 25 —_ 30.0 
Minca? 126 160 25 _— — 
Pueblo Bello 122, 114 136, 147 24, 26 17, 20 30.5, 31.4 
E! Orinoco 129, 120 145, 135 24, 25 18, 17 29.2, 30.2 
Villanueva 120(116-127) 4 151 (133-165) 4 24(23-25)4 | 16.5(15-17)4 | 29.7, 30.2 
Sierra Negra 125 (116-136) 4 141 (133-153) 4 25(24-27)4 | 18(18-19)4 30.1(29.4-31.2)8 
Marimondas 128, 140 151, 150 25, 26 17, 19 30.4, 32.8 
Buena Vista 125(115-131)5 135(123-167)5 27 (25-28)5 | — 31.8(31.2-32.6) 4 
Guaicaramo _ = 28 _ 35.2 
Villavicencio § 134 143 27 _ 34 
Villavicencio 1° 120 130 27 — 29 
VENEZUELA 
Cogollo 130 154 26 _— 31.8 
Azulita 136, 146 148, 169 28, 29 — 32.5, 34.4 
Mérida 1! 116 129 245 — _ 
Mérida _ — — — 30.7 (28.3-31.6) 14 
Caicara 13 122 152 25 16 30.2 
Caicara 119 144 26 _ 30.7 
Yagual 4 127 121 25 _ 32.0 
Casiquiare 107 136 26 = 31.3 
E] Merey 129 156 26 —_ 30.8 
Esmeralda 122, 121 140, 145 28, 26 _ 30.6, 30.9 
Ihuapo 120 132 26 _ 31.2 
Ocama 125 155 2 _ 33.0 
BRAZIL 
Curicuriari 126 124 _— _— = 
Curicuriari 1° 124 137 — _ — 
Jauaraté 137 148 28 _ _- 
Yavanari 115 125 27 _ 31.8 
Cuiteué 140 150 27 _ 33.1 
Santa Rosa 17 126 158 25 § 17 = 
Tauary 142 158 27 _ 33.8 
Manapiri 114 153 27 16 = 


SEEPS EY SRS RAST LUE SN Le eh ee SU ee ee! 
1 Type of tectus Thomas, from original description, followed by a paratype (USN M). 
2 Possibly crown length. 
3 Type of frontalis Goldman, from original description. 
4 Type of illectus Bangs, from original description. 
6 Without claw. 
6 Type of trichurus J. A. Allen, from original description. 
1 Type of mincae J. A. Allen, from original description. 
8 Type of helvolue J. A. Allen, from original description. 
§ Probably=5.2 mm. 


of adults. 


Zygomatic 
breadth 


17.0 
17.0 


15.8 
17.0, 16.5 
18 


16.5(15.4-17.4)7 


16.6, 16.2, 17.6 
15.4 

16.8 

16.3, 17.1 

17.3 

15.3 

15.0 

15.7, 16.8 

16.4, 16.0 
15(14.8-15.3)3 
16.1(15.2-17.1)4 
15.9, 16.4 
16.6(16.1-17.2)5 


16.3 

16.3, 17.4 
16.5(15.6-16.9) 4 
213 

16.9 

17.0 

16.4 

16.3 

16.6, 17.1 


ARBOREAL RICE RATS—HERSHKOVITZ 


Rostrum width 


5.9, 5.6, 6.2 
5.6 

5.5 
5.5(5.3-5.7) 4 
5.6 
5.7(5.4-6.3)5 
5.5 


5.6, 5.6 
5.5, 5.6 
5.4(5.1-5.6)5 
5.5(5.2-5.8) 4 
5.2, 5.5 
5.9(5.6-6.3)5 
6.5 


5.9 
5.9, 6.5 


5.7(5.3-6.2)16 


5.8 


Incisive 
foramina 


6.3, 5.7, 5.8 
5.5 

5.4 
5.4(4.7-5.8) 4 
5.6 
5.6(5.2-5.9)6 
4.8 

5.0 

5.1, 6.0 

4.9, 5.5 
5.1(4.8-5.3)5 
5.4(5.2-5.6) 4 
5.3, 5.7 
5.8(5.6-6.0)5 
5.8 


Diastema 


7.7(7.2-7.9) 4 
8.2 
7.9(7.6-8.2)6 
7.1 


7.7, 8.0 

ss atlads 
7.3(7.0-7.5)5 
7.4(7.0-7.7)4 
7.5, 8.0 
8.2(7.9-8.8)5 
9.1 


7.5(7.0-7.9)19 


7.6 


10 Type of vicencianus J. A. Allen, from original description. 
il Type of flavicans Thomas, from original description. 

13 Type of caicarae J. A. Allen, from original description. 

18 In the original description, ‘13.6 mm.” is obviously wrong. 
14 Type of klagesi J. A. Allen, from original description. 

18 Type of concolor Wagner, from original description. 


16 Type, collector’s measurements according to Pelzeln, 1833 (K. K. Zool.-Bot. Gesellsch., Wien., Beih., 


vol. 33, p. 70). 


17 Type of tapajinus Thomas, from original description. 


Alveolar length 
of molar row 


5.0 


5.1, 5.0, 5.2 
5.1 

4.9 
5.0(4.7-5.2) 4 
5.5 
4.9(4.6-5.2)8 


4.4, 4.6 
4.7(4.6-4.9)5 
4.9(4.7-5.0) 4 
4.7, 4.9 
5.1(4.9-5.5)6 
5.5 


4.7 

5.3, 4.9 

4.83 
5.0(4.7-5.4) 21 
5.0 

4.5 


549 


(Number of specimens, when more than three, tndicated by italic figure.) 


Zygomatic 
plate width 


3.9 
3.9 


3.1, 3.2 
—, 3.8 


3.3(2.9-3.9))7 


3.8, 2.7, 3.5 
3.3 

3.3 
3.2(3.1-3.4) 4 
3.7 
3.4(3.1-3.6)8 
3.0 


3.1, 3.4 

2.8, 2.8 
2.8(2.6-2.9)6 
3.3(3.0-3.6) 4 
3.5, 3.2 
3.7(3.5-3.9)5 
4.1 


3.3 

3.3, 3.4 

3.1 
3.1(2.8-3.5) 21 


3.1 
3.3 
2.9 
3.38, 3.0 
3.3 
3.3 


550 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


the subspecies concolor. Its belly is tinged with buffy, the hairs of the 
throat, chest, and groins white. Oryzomys klagesi J. A. Allen, from 
El Llagual (or Yagual), Rio Caura, farther down the Rio Orinoco, is 
nearer true concolor than speciosus, with which it was originally com- 
pared. O¢ecomys caicarae J. A. Allen is the third name given to the 
middle Rio Orinoco population of concolor. The duskier coloration 
of the upperparts in the type series indicates a cline leading to the 
darker speciosus in northeastern Venezuela and Trinidad. 

Twelve specimens from the upper Rio Meta region, Colombia 
(Villavicencio, Buena Vista, Mambita, Guaicaramo, Medina), repre- 
senting Oryzomys helvolus J. A. Allen and vicencianus J. A. Allen agree 
with concolor of the upper Rio Negro-Rio Orinoco region in size, but 
are more saturate on underparts and as dark on upperparts and sides 
as O. concolor superans. An extremely large female from Guaicaramo, 
like the type of frontalis, is equal in size to average adult superans. 
Morphologically and geographically, however, the mice of the Colom- 
bian base of the Andes in Orinoco drainage are nearest true concolor. 

Thirty-six topotypes of flavicans Thomas from Mérida, Venezuela, 
vary remarkably little inter se. Their upperparts are uniformly pale, 
their bellies whitish with a light to moderate, never a heavy, buffy 
wash. 

Three mice from La Azulita in the Maracaibo basin at the foot of the 
Sierra de Mérida are indistinguishable from the Mérida series. A 
specimen from Rio Cogollo on the Sierra de Perijé side of Lake 
Maracaibo is brighter above, more buffy beneath, and quite like 
individuals from the Colombian side of the same mountain range. 

The relationship of the intensity of body color to rainfall is shown 
by specimens from diverse localities in the Sierra Nevada de Santa 
Marta. The darkest specimens are from the humid Don Diego region 
at the base of the mountains. Topotypes of zlectus Bangs from the 
equally humid but cooler northern slope of the Sierra average paler. 
A series from the humid locality of Pueblo Bello on the southern side 
of the range is colored like topotypes of illectus. In the warmer, less 
humid El Salado, farther down the eastern slope, the mice average 
paler. Thirteen specimens from the comparatively dry Rio Cesar 
valley (Villanueva; El Orinoco) between the Sierra Nevada and 
Sierra de Perij4 are smaller and extremely pale, their underparts more 
nearly pure white. They were taken from the early part of the dry 
season in January to the beginning of the rainy season in April. The 
January specimens are darkest, their pelage in good condition. In 
the March specimens, the pelage is faded and molting. April speci- 
mens show the darker new pelage on the middorsal region. The 
contrast is quite marked between the pale, comparatively short-haired 
valley mice of the dry season and the saturate prime pelage of speci- 


ARBOREAL RICE RATS—HERSHKOVITZ 551 


mens taken during the rainy season on the comparatively cool slopes 
of the Sierra Nevada and the Sierra de Perija. 

Oryzomys trichurus J. A. Allen is founded on a juvenal and Oecomys 
mincae J. A. Allen on an adult, both from localities near Bonda in 
the semiarid northwestern corner of the Sierra Nevada de Santa 
Marta. They are like the small pale mice of the ecologically similar 
valley at the western base of the Sierra. One specimen at hand from 
Bonda is virtually a dwarf. 

The range of variation in color, size, and degree of development of 
the supraorbital ridges shown by the mice of the Sierra Nevada de 
Santa Marta region is greater than in those of any other single region. 
The tendency for each stream valley branch in each altitudinal zone 
to produce a distinctive population is one of the most striking bio- 
logical phenomena of the Sierra. 

Oryzomys flavicans subluteus Thomas from ‘‘W. Cundinamarca,” 
Colombia, was described as similar to Venezuelan flavicans “but dis- 
tinguished by its darker belly.” Three specimens at hand from 
Paime, Rio Minero, western Cundinamarca, with underparts lightly 
washed buffy are absolutely indistinguishable from topotypes of 
flavicans. ‘Two specimens from Muzo, western Boyacdé, and but a 
few miles downstream (north) from Paime, with underparts entirely 
buff except for the white throat and chin are exactly lke the type of 
subluteus. 

Oryzomys tectus Thomas was described as more brightly colored 
than flavicans and with “remarkably expanded supraorbital ridges.” 
The difference in color between western Panamanian tecius and Vene- 
zuelan flavicans lies well within the limits of variations common to 
any large population from either region. Supraorbital ridges are 
present in all members of the species and are more prominent in older 
than in younger individuals. It is true, nevertheless, that the ridges 
are more uniformly developed and ledgelike in most Central Ameri- 
can and western Colombian mice than in many other series of the 
subspecies. 

Western Colombian (Socorré, Puri, Catival) mice are indistinguish- 
able from their Panamanian relatives. In one specimen from Puri, 
Antioquia, the upperparts and sides are grizzled with an interspersion 
of wholly white cover hairs. A female from Socorré, upper Rio Sint, 
Cérdoba, shows the molt from dark to bright pelage. 

The name frontalis Goldman is based on eastern Panamanian mice 
that are less tawny above and less buffy beneath than the average of 
western Panamanian tectus. The difference, although locally con- 
stant, is slight. 

Remarks: The original description of Hesperomys concolor includes 
the following Latin diagnosis: “H[esperomys] fulvus, subtus abrupte 


552 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


albus, pedibus fuscentibus; cauda nuda longitudine corporis; pilis 
gastraci unicoloribus.”’ 

Measurements, presumably taken from the dry skin are: ‘‘Kérper 
[head and body], 4’’ 10’ [=126 mm.] and Schwanz [tail] 4’’ 9’’” [=124 
mm.].”’ These measurements do not tally with those taken in the 
flesh by the collector (see below). Either set of measurements, how- 
ever, lies within the range of variation for the species. Wagner 
compared the type of concolor with the larger Hesperomys anguya 
(= Oryzomys buccinatus Olfers) and noted that in the former the 
color of the upperparts are much brighter, more reddish, the hair of 
the underparts pure white, the cheeks without the gray as in anguya, 
the feet covered with brownish hair, and the tail dark and with short 


hairs. 
Field notes by the collector, Johann Natterer, published by Pelzeln 
(1883) are translated from the German as follows: 


The tarsus [of the type of concolor] is much broader than that of No. 173 [Hespe- 
romys russatus Wagner=Holochilus brasiliensis Desmarest]. My dogs found it 
in the opening of a hollow tree and I got it out with the aid of a mattock. Its 
color is exactly like that of a reddish, white-bellied mouse from Ypanema [? 
species]. Incisors ochraceous; ears round, gray brown, the base grayish buff, 
toes and upperside of tarsus apparently naked and covered with small brown 
scales on a buffy base where, on closer inspection, extremely fine short hairs 
become visible. The hairs are somewhat longer at the roots of the claws but 
do not extend as far as the tips of the claws. Only 6/’’’ [13 mm.] of the base of 
the tail is covered with fur. Total length, 10’ [=261 mm.], tail 5%’’ [=137 
mm.]. 


The size, color, broad hind foot, and relatively long and thinly 
haired tail of concolor are diagnostic of Oecomys. There is no other 
cricetine in the upper Rio Negro region with which it can be confused. 

SPECIMENS EXAMINED: One hundred and sixty-one, from the follow- 
ing localities: 

Costa Rica: San Gerénimo, San José, 1 (CNHM); El General, Puntarenas, 
1 (UMMZ); Boruca, 2 (CM). 

Panama: Bugaba, Chiriqui, 2 (BM, type of tectus; USNM, 1); Barro Colorado, 
Canal Zone, 1 (UMMZ); Cana, Darién, 11 (USNM, 9 including type of frontalis; 
CNHM, 2). 

Cotomstia: Catival, Cérdoba, 1 (CNHM); Socorré, Rfo Sint, Cérdoba, 3 
(CNHM); Purif, Rio Cauca, Antioquia, 1 (CNHM); Muzo, Rfo Minero, Boyacé, 
2 (CNHM); “W. Cundinamarca,” 1 (BM, type of subluteus); Paime, Rio Minero, 
Cundinamarca, 3 (AMNH); Pueblo Viejo, Magdalena, 5 (AMNH, 3; MCZ, 2); 
Concepcion, Magdalena, 1 (MCZ); Palomino, Magdalena, 1 (MCZ); Don Diego, 
Magdalena, 8 (CM); Bonda, Magdalena, 3 (AMNH); Mamatoca, Magdalena, 1 
(AMNH); El Lfbano, Magdalena, 1 (AMNH, type of trichurus); Minea, Mag- 
dalena, 1 (AMNH, type of mincae; Pueblo Bello, Magdalena, 3 (USNM); El 
Salado, Magdalena, 5 (USNM); El Orinoco, Rfo Cesar, Magdalena, 7 (USNM); 
Villanueva, Magdalena, 6 (USNM); Sierra Negra, Sierra de Periji, 4 (USNM); 
Las Marimondas, 2 (USNM); Mambita, Cundinamarca, 2 (AMNH); Medina, 
Cundinamarca, 1 (AMNH); Buenavista, Meta 5 (AMNH); Guaicaramo, Meta, 2 


ARBOREAL RICE RATS—HERSHKOVITZ 553 


(AMNH); Villavicencio, Meta, 2 (AMNH, types of helvolus and vicencianus) ; 
Maipures, Rfo Orinoco, Vichada, 1 (BM, type of marmosurus). 

VENEZUELA: Rfo Cogollo, Mérida, 1 (CNHM); La Azulita, Mérida, 4 (CNHM); 
Mérida, Mérida (includes Cafetos de Mérida, Milla, Lourdes, Chama, Montes de la 
otra banda) 37 (BM, type of flavicans; AMNH, 23; CNHM, 3; USNM, 10); 
Caicara, Rio Orinoco, Bolivar, 4 (AMNH, includes type of catcarae); El Yagual, 1 
(AMNH, type of klagesi); Rio Casiquiare, Amazonia, 1 (AMNH); Rfo Casi- 
quiare, near Raudal Tamasd, Amazonia, 1 (AMNH); El Merey, Rio Casiquiare, 
Amazonia, 2 (AMNH); Esmeralda, Rio Orinoco, Amazonia, 3 (AMNH); Ihuapo, 
Rio Orinoco, Amazonia, 1 (AMNH); Rfo Ocama, mouth, at Rio Orinoco, Ama- 
zonia, 3 (AMNH). 

Braziu: Jauaraté, Rio Uaupés, Amazonas, 1 (AMNH); Yavanari, Rio Negro, 
Amazonas, 1 (AMNH); Igarapé Piaba, mouth, Rio Amazonas, Pard, 1 (MCZ); 
Lago Cuiteud, Rio Amazonas, Pard, 1 (MCZ); Rosarhinho, Lago Miguel, Rio 
Madeira, Amazonas, 1 (AMNH); Tauary, Rio Tapajéz, Pard, 2 (MCZ); Ilha 
Manapiri, Rio Tocantins, Pard, 1 (AMNH); Bardo de Melgago, Mato Grosso, 
3 (AMNB#). 


Oryzomys concolor speciosus J. A. Allen and Chapman 


Oryzomys speciosus J. A. Allen and Chapman, Bull. Amer. Mus. Nat. Hist., 
vol. 5, p. 212, 1893; vol. 9, p. 18, 1897 (Caparo, Trinidad).—Goodwin, Bull. 
Amer. Mus. Nat. Hist., vol. 102, p. 300, 1953 (type history and measurements). 

Oryzomys trinitatis J. A. Allen and Chapman, Bull. Amer. Mus. Nat. Hist., vol. 5, 
p. 213, 1893 (type locality, Prince’s Town, Trinidad); vol. 9, p. 18, 1897 (Trini- 
dad: Caparo; Caura).—Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 190, 
1939 (Bartica, British Guiana; Arabupu, Mt. Roraima, and Sucre, Venezuela; 
and Trinidad. Synonyms: palmarius, fulviventer)—Goodwin, Bull. Amer. 
Mus. Nat. Hist., vol. 102, p. 301, 1953 (type history and measurements). 

Oryzomys palmarius J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 12, p. 211, 1899 
(type locality, Quebrada Seca, Rfo Manzanares, Cumanacoa, Sucre, Venezuela; 
another specimen from Los Palmales, Sucre).—Goodwin, Bull. Amer. Mus. 
Nat. Hist., vol. 102, p. 301, 1953 (type history; ‘‘may be a subspecies of O. 
ffavicens [sic =flavicans] Thomas’’). 

Oryzomys fulviventer J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 12, p. 212, 1899 
(type locality, Quebrada Seca, Cumanacoa, Sucre, Venezuela).—Goodwin, 
Bull. Amer. Mus. Nat. Hist., vol. 102, p. 301, 1953 (type history and measure- 
ments). 

Ccomys guianae Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 187, 1910 (type 
locality, Supinaam River, Demerara, British Guiana).—Anthony, Zoologica, 
vol. 3, No. 18, p. 274, 1921 (Kartabo, British Guiana). 

Oecomys guianae guianae, Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 198, 1939 
(listed). 

CGcomys splendens Hayman, Ann. Mag. Nat. Hist., ser. 11, vol. 1, p. 381, 1938 
(type locality, Mayaro, southeastern Trinidad). 

Oecomys auyantepui Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, p. 193, 1939 
(type locality, southern slopes of Mt. Auyan-tepuf, Rio Caroni, Bolivar, 
Venezuela, altitude, 3,500 feet)—Goodwin, Bull. Amer. Mus. Nat. Hist., 
vol. 102, p. 302, 1953 (type history and measurements). 


Type: Adult female, skin and skull (AMNH 5942/4672) collected 


Apr. 26, 1893, by Frank M. Chapman. 
TYPE LocauLity: Prince’s Town, Trinidad. 


554. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Disrrisution: Trinidad, the Guianas, and eastern Venezuela, 
including the coast and delta and basin of the lower Rfo Orinoco; 
altitudinal range from sea level to about 1,500 meters above. 

Cuaracters: Like concolor but darker throughout; upperparts 
tawny finely ticked with dark brown; underparts gray, sometimes 
whitish or buffy with the plumbeous basal portions of the hairs 
showing through. 

MEASUREMENTS: See table 8. 

Variation: Underparts vary from whitish to dark gray with or 
without a light buffy wash. In all specimens, however, the basal 
portions of the hairs of belly and chest are plumbeous and show 
through the surface. A pectoral streak is present in the two speci- 
mens from Caura, Trinidad, and in the specimens from Quebrada 
Seca and Neveri, Venezuela. 

Remarks: Of a series of four specimens collected in Prince’s Town, 
Trinidad, Allen and Chapman described one as Oryzomys speciosus 
and the others as O. trinitatis. The basal portions of the belly hairs 
in the first are more nearly white than in the others. The authors 
noted that “in size, proportions and coloration [speciosus] strongly 
suggests Hesperomys concolor Wagner.” Paradoxically, they failed 
to recognize the same relationship to the specimen they named 
trinitatis. 

Oryzomys palmarius J. A. Allen and Oryzomys fulviventer J. A. Allen 
are likewise based on differently colored individuals of a small series 
taken from Noy. 23 through Dec. 10, 1898, in Quebrada Seca, Sucre, 
Venezuela. In the first, the underparts are typically grayish white 
with the dark basal color of the hairs showing through. In the second, 
the belly is buffy, the throat whitish. 

When Thomas erected the subgenus Oecomys in 1906, he failed to in- 
clude in it the named forms speciosus, trinitatis, palmarius and fulviven- 
ter. A consequence of the omission was a dual system of classification 
and the multiplication of names for the eastern Venezuelan and Guianan 
race of Oryzomys concolor. Thus, Oecomys guianae Thomas, described 
in 1910, was compared only with marmosurus of Venezuela and tapa- 
jinus of Brazil, while Oecomys splendens Hayman was described in 1938 
as the first Trinidad record of arboreal rice rats. 

The two specimens from Auyan-tepui, on which the description 
of Oecomys auyantepur Tate is based, are colored like other mainland 
representatives of speciosus. ‘Their shorter hind feet and narrower 
zygomatic plates, however, are more nearly as in O. bicolor. A skull 
only from Kartabo, British Guiana, also has the narrow zygomatic 
plate. Its small molars also indicate affinity with O. bicolor. Osten- 
sibly, the large and small species of arboreal rice rats have not diverged 
as widely in the Guianan region as they have elsewhere. 


505 


ARBOREAL RICE RATS—HERSHKOVITZ 


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556 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


SPECIMENS EXAMINED: Twenty-eight, from the following localities: 


TRINIDAD: Prince’s Town, 4 (AMNH, includes types of speciosus and trinitatzs) ; 
Caura, 2 (AMNH); Mayaro, 1 (BM, type of splendens); Cumaca, 1 (AMNH); 
St. Pats, 1 (AMNH); San Rafael, 1 (CNHM). 

VENEZUELA: Quebrada Seca, Sucre, 5 (AMNH, includes types of palmarius 
and fulviventer); Neveri, Sucre, 3 (AMNH); San Rafaél, Sucre, 1 (CM); Rfo 
Yurudn, Bolivar, 1 (AMNH); Auyan-tepui, Bolivar, 2 (AMNH, includes type 
of auyanteput); Arabupu, Mt. Roraima, Bolivar, 2 (AMNH). 

British Guiana: Kartabo, 4 (BM, type of guianae; AMNH, 3). 


Oryzomys concolor superans Thomas 


(Ecomys superans Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 8, p. 250, 1911; ser. 
10, vol. 2, p. 261, 1928 (San Jerénimo, Peru, 1,000 feet altitude; palmeri Thomas, 
a synonym). 

Oryzomys (Oecomys) superans, Sanborn, Publ. Mus. Hist. Nat. ‘Javier Prado,” 
ser. A, Zool., No. 6, p. 21, 1951 (Quincemil, Cuzco, Peru). 

comys palmeri Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 8, p. 251, 1911 (type 
locality, Canelos, Rfo Bobonaza, upper Rio Pastaza, Napo-Pastaza, eastern 
Ecuador; altitude 2,100 feet). 

Ccomys melleus Anthony, Amer. Mus. Nov., No. 139, p. 4, 1924 (type locality, 
Zamora, Santiago-Zamora, eastern Ecuador, altitude 3,250 feet.—Goodwin, 
Bull. Amer. Mus. Nat. Hist., vol. 102, p. 302, 1953 (type history and measure- 
ments). 

(comys osgoodi Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 14, p. 287, 1924 (type 
locality, Moyobamba, Amazonas, Peru; altitude about 820 meters above sea 
level); ser. 9, vol. 19, p. 369, 1927 (Yurac Yacu, San Martin, Peru); ser. 9, vol. 
20, p. 601, 1927 (Tingo Maria, Peru, 2,000 feet altitude; measurements). 


Oecomys osgoodi, Laurie, Ann. Mag. Nat. Hist., ser. 12, vol. 8, p. 276, 1955 (Rio 
Tigre, Ecuador, 2,000 feet altitude). 

Oryzomys auriventer, J. A. Allen (not Thomas), Bull. Amer. Mus. Nat. Hist., vol. 
35, p. 118, 1916 (Zamora, Ecuador) [= type series of O. melleus Anthony]. 
Typre: Adult female, skin and skull (BM 11.7.19.12) collected 

Apr. 28, 1910, by M. G. Palmer. 

Type LocaLity: Canelos, Rio Bobonaza, Napo-Pastaza Province, 
eastern Ecuador; altitude, 640 meters above sea level. 

DistriBpuTION: Base, lower slopes, valleys, and foothills of the 
Andes drained by the Amazon in Colombia, Ecuador, and Peru, from 
approximately 200 to 1,200 meters above sea level. 

CuHaractsers: Largest and darkest of the subspecies; upperparts 
and sides tawny, the back evenly mixed or heavily lined with dark 
brown; underparts gray to buff or ochraceous orange, the dark 
plumbeous basal portions of the hairs showing through; throat and 
chest not markedly different from belly. 

MEASUREMENTS: See table 9. 

VARIATION: Twenty-five specimens of superans taken during the 
dry season (January-March) along the upper and middle Rio Caqueta 
(Florencia; Mecaya; Tres Troncos) vary considerably inter se. A few 
in prime pelage show either the dark color phase or the bright color 


ARBOREAL RICE RATS—HERSHKOVITZ 557 


phase on upperparts and sides. Most of the other specimens are in 
old worn pelage of one color phase with patches of new pelage of the 
other phase. Underparts range from ‘‘dirty gray,” sharply defined 
from sides, to gray washed with buff on chest and belly and hardly or 
not at all defined from sides. 

The arboreal rice rats of the rain forests of eastern Ecuador vary 
like the preceding but average darker, especially on the underparts. 
Twenty-two adults and subadults at hand collected in February, 
March, June, August, October, and November reveal no seasonal 
molt pattern. One specimen collected in each of the months of 
February, August and October is in old pelage. Others taken at the 
same times and during other months of the year are in good to prime 
pelage. 

Specimens from higher altitudes in the valleys and slopes of the 
Andes (Zamora and Chonta Urcu in Ecuador; Moyobamba, Amazonas, 
and Hacienda Exito, Huanuco, in Peru) are smaller, have longer 
pelage and are more richly colored, especially on underparts, than 
their relatives from the base of the Cordillera. 

REMARKS: Oryzomys concolor superans is the most distinctive race. 
The transition in eastern Colombia from concolor of Orinoco drainage 
to the larger darker superans of Amazonian drainage seems to be 
abrupt. Likewise, the contrast between smaller white-bellied roberts 
of northwestern Bolivia and swperans of southeastern Peru is sharp. 

Oecomys palmeri Thomas is merely a small topotype of superans 
collected the same day by the same collector. Four specimens from 
Zamora, eastern Ecuador, first recorded by J. A. Allen as Oryzomys 
aurwenter, were described as Oecomys melleus by Anthony. They 
average slightly smaller with underparts more generally ochraceous 
orange. Individually, however, they are indistinguishable from other 
eastern Ecuadorian representatives of superans. The type of Oecomys 
osgoodi Thomas is another small individual of superans. Its author 
compared it with O. palmeri only. In 1927, Thomas (Ann. Mag. 
Nat. Hist., ser. 10, vol. 2, p. 261) observed that palmeri was ‘not 
really distinct from @. superans’’ and that the Peruvian ‘‘bush rat” 
was also the same. He concluded that ‘‘throughout the tropical re- 
gions of S. America almost every district has two members of this 
genus—a large one, the size of a rat [=concolor], and a much smaller 
one, that of a large mouse [=bicolor].”’ 

SPECIMENS EXAMINED: Sixty-three, from the following localities: 

Cotomsia: Florencia, Caquetdé, 1 (CNHM); Tres Troncos, Rio Caquetd, Putu- 
mayo, 7 (CNHM); Rio Mecaya, Rio Caquetdé, Putumayo, 17 (CNHM). 

Ecuapor: Rio Napo, Napo-Pastaza, 1 (UMMZ); Llunchi, Rio Napo, Napo- 


Pastaza, 1 (UMMZ); Chonta Urcu, Napo-Pastaza, 6 (MCZ, 4; CNHM, 2); 
Canelos, Rio Bobonaza, Napo-Pastaza, 3 (BM, types of swperans and palmeri; 


VOL. 110 


PROCEEDINGS OF THE NATIONAL MUSEUM 


558 


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ARBOREAL RICE RATS—HERSHKOVITZ 559 


AMNH, 1); Montalvo, Rfo Bobonaza, Napo-Pastaza, 3 (CNHM); Rio Pindo 
Yacu, Napo-Pastaza, 5 (CNHM); Yana Rumi, Napo-Pastaza, 3 (CNH M); Capa- 
huari, Napo-Pastaza, 2 (CNHM); Zamora, Santiago-Zamora, 4 (AMNH, in- 
cludes type of melleus). 

Peru: Moyobamba, Amazonas, 1 (BM, type of osgoodt); Hacienda Exito, Rfo 
Cayumba, Huanuco, 2 (CNHM); Satipo, Junin, 1 (MCZ); La Pampa, Puno, 
1 (MCZ); Quineemil, Cuzco, 3 (CNHM); Itahuania, Madre de Dios, 1 (CNHM);: 
Boca Colorado, Madre de Dios, 1 (CNHM). 


Oryzomys concolor roberti Thomas 


Rhipidomys roberti Thomas, Proc. Zool. Soc. London (1903), No. 2, p. 237, 1908. 

[Oryzomys (icomys)] Roberti, Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, p. 
445, 1906. 

(E[comys] robertt, Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 3, p. 378, 1909 
(comparison). 

Oecomys robertt, Vieira, 1945, Arg. Zool. Sio Paulo, vol. 4, p. 428, 1945 (Fazenda 
Aric’, Rio Aric&, Mato Grosso, Brazil). 

Oryzomys (Gicomys) mamorae Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 18, p. 
445, 1906 (type locality, ‘‘Mosetenes,” upper Rio Mamoré, Yungas, Cocha- 
bamba, Bolivia). 

(Ecomys mamorae, Osgood, Field Mus. Nat. Hist., zool. ser., vol. 10, p. 206, 1916 
(Todos Santos, Rio Chaparé, Cochabamba, Bolivia). 

Typ: Adult male, skin and skull (BM 3.7.7.67) collected July 6, 
1902, by Alphonse Robert. 

TypE Locauity: Santa Ana de Chapada, Mato Grosso, Brazil; alti- 
tude, 800 meters above sea level. 

Distrisution: Northern Bolivia in the departments of Beni, 
Cochabamba, Santa Cruz, and Chuquisaca west into Mato Grosso, 
Brazil, in the Rio Paraguay basin; altitudinal range approximately 
200 to 1,500 meters above sea level. 

CHARACTERS: Smaller and paler than superans; upperparts buffy 
to tawny mixed with dark brown, underparts entirely white and 
sharply defined from sides, or, sometimes, with a buffy or ochraceous 
wash on chest and belly; skull with supraorbital ridges less developed 
and divergent posteriorly than in other races. 

M#ASUREMENTsS: See table 10. 

VaRIATION: The type and two specimens at hand from Mato Grosso, 
Brazil, are the palest representatives of the species east of the Andes. 
Specimens from the Bolivian departments of Chuquisaca, Beni, and 
Santa Cruz show a heavier mixture of dark brown on upperparts and 
sides. Six specimens from Todos Santos, Rio Chaparé, collected by 
R. H. Becker, average nearly as dark on upperparts as swperans. 
The underparts, however, are pure white in two, more or less buffy 
on belly and midline of chest in the others. 

Remarks: The clear white or buffy underparts distinguish every 
member of robertz from superans. On the other hand, subspecies 
roberti is hardly separable from Amazonian concolor by its paler color, 


VOL. 110 


PROCEEDINGS OF THE NATIONAL MUSEUM 


560 


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ARBOREAL RICE RATS—HERSHKOVITZ 561 


slightly larger size, and, more significantly, by its narrower supra- 
orbital region and weaker temporal ridges. 
SPECIMENS EXAMINED: Twenty-three, from the following localities: 


Braziu: Santa Ana de Chapada, Mato Grosso, 1 (BM, the type of roberti); 
Descalvados, Mato Grosso, 1 (CNHM); Urucum de Corumbdé, Mato Grosso, 1 
(CNHM). 

Bouivia: ‘‘Mosetenes,’’ Cochabamba, 1 (BM, type of mamorae); Todos Santos, 
Cochabamba, 10 (AMNH, 5; CNHM, 5); Buenavista, Santa Cruz, 6 (CNHM, 5; 
MACN, 1); Rio Yapacani, Santa Cruz, 1 (MACN); Marbdn, Rio Mamoré, 
Santa Cruz, 1 (MACN); Ticucha, Rio Capirenda, Chuquisaca, 1 (CNHM). 


Oryzomys concolor bahiensis, new subspecies 


Mus cinnamomeus Pictet and Pictet, Notices sur les animaux nouveaux ou peu 
connus du musée de Genéve, p. 64, pl. 19 (animal), pl. 238, fig. 5 (molars), 
1844 (type locality, Bahia, Brazil; name preoccupied by Mus cinnamomeus 
Lichtenstein, 1830, Darstellung neuer oder weniger bekannter Saugethiere, 
Berlin, pl. 36, fig. 2 [==Proechimys myosuros Lichtenstein)). 

Oecomys cinnamomeus, Moojen, Os roedores do Brasil, p. 53, 1952 (Ilhéus, Bahia, 
Brazil). 

Type: Adult male, skin and skull (CNHM 63780) collected 
Sept. 19, 1944, by the Servico de Estudos e Pesquisas Sobre a Febre 
Amarela; original number M 19330. 

Typr LocaLity: Fazenda Almada, Ilhéus, Bahia, Brazil. 

DistrisuTion: Known only from eastern Bahia, Brazil, but 
possibly ranges south into Minas Gerais and north and west into 
the Rio Sao Francisco basin. 

Cuaracters: Darker throughout than concolor; upperparts tawny, 
underparts gray with a buffy wash, the slaty basal portions of the 
hairs of belly, chest and limbs showing through; supraorbital region 
broad, the ridges well developed and ledge-like. 

Measurements: Of the type and paratype, respectively, as follows: 
Head and body, 140, 130; tail, 145, 150; hind foot, dry with claw, 
27, 28; weight, —, 57 grams; greatest length of skull, 33.0, 36.6; 
zygomatic breadth, 18, 16.7; width of rostrum, 5.9, 6.0; nasals, 11.7, 
11.5; incisive foramina, 6.0, 5.9; diastema, 8.0, 7.9; alveolar length of 
molar row, 5.3, 5.3; zygomatic plate, 3.8, 3.4. 

Measurements of the type of cinnamomeus Pictet and Pictet, given 
in the original description, are as follows: head and body, 148; tail, 
121; hind foot, 27. 

Remarks: In the type, the upperparts are ochraceous tawny 
mixed with the dark brown of the guard hairs; a faint ochraceous buff 
lateral line is present; belly and underside of limbs gray, washed with 
buff, the slaty basal portion of the hairs showing through; posterior 
half of chest like belly, anterior portion and throat nearly pure white; 
tail brown, upperside of metacarpus and metatarsus brownish, digits 
silvery gray. A paratopotype is hardly distinguishable from the type. 

504676—59—_~—-4 


062 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


The present specimens agree closely with the original description 
of Mus cinnamomeus. The colored plate of cinnamomeus, however, 
shows. the wrist and adjoining part of the arms wholly white. This 
condition is not only anomalous for Oecomys but is not in accord with 
the text where it is stated that the anterior and outer sides of the fore- 
limbs are colored like the dorsal surface of the head and body. The 
name Mus cinnamomeus Pictet, 1844, is preoccupied by Mus cinna- 
momeus Lichtenstein, 1830, a Proechimys. 

SPECIMENS EXAMINED: Two, from Brazil: Almada, Ilhéus, Bahfa, 
1, the type (CNHM); Urucutuca, Ilhéus, Bahia, 1 (CNHM). 


GAZETTEER 


(With Name of Collector and Name of Species) 


Bouivia 


Buenavista (400-500 m.), 17°27’ S., 
63°21’ W. 
Rio Yapacani, Ichilo, Santa Cruz. 
F. and J. Steinbach; Oryzomys 
concolor. 
Chimate (700 m.), 15° S., 68° W. 
Upper Rio Beni, La Paz. 
P.O. Simons; O. bicolor. 
El Palmal (Palmal), 17°05’ S., 
65°23’ W. 
Chaparé, Cochabamba. 
F. Steinbach; O. bicolor. 
Marban (240 m.) 
Rio Mamoré, Beni. 
F. Steinbach; O. bicolor, O. con- 


color. 

‘‘Mosetenes’”’ (probably a mis- 
spelling) 

Upper Rio Mamoré, Yungas, 
Cochabamba. 

L. Balzan; O. concolor. 

Ticuche, (1,500. m.) 20°25! #85, 
63°58! W. 


Rio Capirenda, Chuquisaca. 
J. Arduz Tardio; O. concolor. 
Ticunhuaya (1,500 m.), 15°35’ S., 
68°10’ W. 
iti La:Paz: 
G. H. H. Tate; O. bicolor. 
Todos Santos (250 m.), 16°40’ §., 
6b° 15" W. 
On Rio Chaparé, Cochabamba. 
R. H. Becker; O. concolor. 


Bouivia—Continued 
Yapacani, Rio 
Same general collecting locality as 
Buenavista, Santa Cruz (q.v.). 
F. Steinbach; O. concolor. 


BrRaAziIuL 


Almada (Fazenda) (near sea level), 
14°50’ S., 39° W. 
Tihéus, Bahia. 
Rockefeller Institution; O. con- 
color. 
Arica (Fazenda) (203 m.), 15°45’ S., 
55°46’ W. 
Rio Aric’, Mato Grosso. 
O. concolor. 
Baraio de Melgaco (850 m.), 11°52’ 
S., 60°48’ W. 
Rio Commemoracao, upper Rio 
Gy-Parand, Mato Grosso. 
L.. E. Miller; O. bicolor, O. :con- 
© 2 color. 
Descalvados (142 m.), 17°15’ S., 
57°48’ W. 
Mato Grosso. 
C. C. Sanborn; O. concolor. 


Igarapé-agu (50 m.), 1°00’ S., 
47°35’ W. 
Between Belém and Braganza, 


Para. 
A. Robert; O. bicolor. 
Igarapé Bravo 
Rio Tapajéz, Para. 
A. M. Olalla; O. bicolor. 


ARBOREAL RICE RATS—HERSHKOVITZ 563 


Brazit—Continued 
Ilha de Manapiri 
Rio Tocantins, Pard. 
M. Goeldi; O. concolor. 
Jauaraté [Yauareté] (82 m.), 1°24’ 
N.; 69°12%W.. 
Rio Uaupés (left bank), Amazonas. 
Olalla Bros.; O. concolor, O. bi- 
color. 
Jurué, Rio (150.m.), 6°45’ S., 
79° W. 
Amazonas. 
Garbe; O. bicolor. 
Limontuba 
Rio Tapajéz, Para. 
A. M. Olalla; O. bicolor. 
Manacapurt, 60°40’ W. 
Rio Solimées, Amazonas. 
O. concolor. 
Marajé (Island) 
At mouth of Rio Amazonas below 
the Equator in Pard. 
O. concolor. 
Monte Alegre (10 m.), 2°00’ S., 
54°04’ W. 
Rio Solimdes, east of Santarém, 
Pard. 
O. concolor. 
Recreio, 3°30’ N., 61° W. 
Rio Majary, Amazonas. 
A. M. Olalla; O. bicolor. 


Rosarinho (about 25 m.), 3°41’ S., 
59°07’ W. 
Lago Miguel, Rio Madeira. 
Olalla Bros., O. concolor. 
Santa Ana de Chapada (855 m.), 
12°25) Ss 50°25' W. 
Mato Grosso. 
A. Robert; O. concolor. 


Santa Rosa, 5°20’ S., 55°30’ W. 
Rio Jamanchim, Rio Tapajéz, 
Pard. 
E. Snethlage; O. concolor. 
Tauary, 3°50’ S., 55°00’ W. 
Rio Tapajéz, Pard. 
A. M. Olalla; O. concolor. 
Urucum de Corumb4, 19°00’ §&., 
57°40’ W. 
Corumba, Mato Grosso. 
C. C. Sanborn; O. concolor; O. 
bicolor. 


Brazit—Continued 
Urucutuba (near sea level), 14°50’ 
S., 39°00’ W. 
Iihéus, Bahfa. 
Rockefeller Institution; O. con- 
color. 
Urup4, 10°50’ S., 61°35’ W. 
Rio Urup4, a tributary of the 
Parand, Mato Grosso. 
L. E. Miller; O. bicolor. 
Villa Braga, 4°20’ 8., 56°00’ W. 
Rio Tapajéz, Pard. 
O. concolor. 
Yauanari, 0°30’ S., 64°50’ W. 
Rio Negro, Amazonas. 
Olalla Bros.; O. concolor. 


BRITISH GUIANA 


Barakara, 5°52’ S£ 57228" Wi 
Canje River, 50 km. southeast 
of New Amsterdam. 
W. Beebe; O. bicolor. 
Bartica (near sea level), 6°24’ N., 
58°36’ W. 
At junction of Essequibo and 
Mazaruni Rivers, Essequibo. 
O. concolor. 
Essequibo River (lower), 6°-7° N., 
58°35’ W. 
S. B. Warren; O. bicolor. 
Holmia, 4°55’ N., 55°35’ W. 
Potaro Highlands. 
M. A. de Freitas; O. bicolor. 
Kartabo, 6°23’ N., 58°42’ W. 
At junction of Mazaruni and 
Cuyuni Rivers, Essequibo. 
W. Beebe; O. bicolor, O. concolor. 
Supinaam (River) 
Demerara; enters mouth of Es- 
sequibo (q.v.) from west. 
O. concolor. 


COLOMBIA 


Andalucia (630 im.), 
75°30’ W. 

Huila. 

L. E. Miller; O. bicolor. 

Bonda (50-80 m.), 11°17’ N., 
74°02’ W. 

Northwest slope of the Sierra 
Nevada de Santa Marta, 
Magdalena. 

Mrs. H. H. Smith; O. concolor. 


2°151S N., 


564 


CoLtomB1a—Continued 


Buena Vista (1,500 m.), 4°20’ N., 
73°40’ W. 

Above Villavicencio, Meta. 

Chapman, Cherrie, et al.; O. 
concolor. 

Catival (120 m.), 8°17’ N., 75°41’ W. 

Upper Rio San Jorge drainage, 
Cérdova. 

P. Hershkovitz; O. concolor. 

Conecepcién (La) (about 900 m.), 
1TOL NS 23 265 W. 

Rio San Antonio, northern slope 
of Sierra Nevada de Santa 
Marta, Magdalena. 

W. W. Brown; O. concolor. 

Donama. (600 mz), 11°13" N.; 
74°00’ W. 

Northwestern slope of Sierra 
Nevada de Santa Marta, Mag- 
dalena. 

M. A. Carriker, Jr.; O. concolor. 

Don Amo [= Donama (q.v.)] 
Don Diego (sea level), 11°16’ N., 
73°37’ W. 

Northern slope of the Sierra 
Nevada de Santa Marta, Mag- 
dalena. 

M. A. Carriker, Jr., O. concolor. 

El ‘Libano’ (1,250).m.), ‘11910 N.., 
74°00’ W. 

Northwestern slope of Sierra 
Nevada de Santa Marta, Mag- 
dalena. 

H. H. Smith; O. concolor. 

El Orinoco, 10°13’ N., 73°23’ W. 

Rio Cesar, Magdalena. 

P. Hershkovitz; O. concolor. 

El Salado (480 m.), 10°22’ N., 
73°29’ W. 

Southeastern slope of Sierra Ne- 
vada de Santa Marta, Mag- 
dalena. 

P. Hershkovitz; O. concolor. 

Florencia (450 m.), 1°45’ N., 75°40’ 
W. 

Rio Orteguaza, Caqueta. 

L. E. Miller; O. bicolor. P. Hersh- 
kovitz; O. concolor. 

Guaicaramo (about 250 m.), 4°35’ 
N., 73°00’ W. 

Northeast of Villavicencio, Rfo 

Upia, Meta. 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


CotomBi1a—Continued 


Hermano Nicéforo Marfa; O. con- 
color. Gonzdlez; O. bicolor. 
Jino Goje, 0°15’ S., 70°28’ W. 
Rfo Apoporis, Amazonas. 
I. Cabrera; O. bicolor. 
Mamatoca (2 m.), 11°15’ N., 74°09’ 
W. 
Three miles east of Santa Marta, 
Magdalena. 
H. H. Smith; O. concolor. 
Mambita (700 m.), 4°46’ N., 73°20’ 
W. 
Rio Guavio, eastern Cundina- 
marca. 
Gonzalez; O. concolor, O. bicolor. 
Maipures (115 m.), 5°18’ N., 67°50’ 
W. 
Rio Orinoco, Vichada. 
G. K. and 8S. M. Cherrie; O. con- 
color. 


Las Marimondas (1,000 m.), 10°52’ 
N., 72°43" W. 
Sierra de Perij4, Magdalena. 
P. Hershkovitz; O. concolor. 
Mecaya, Rio (185 m.), 0°29’ N., 
75°10! W. 

Rio Caqueté, Putumayo. 

P. Hershkovitz; O. bicolor, O. 
concolor. 

Medina, 4°30’ N., 72°45’ W. 

East slope of Eastern Andes, 
Cundinamarca. 

Hermano Nicéforo Maria; O. con- 
color. 

Minca (660 m.), 11°12’ N., 74°04’ 
W. 

Northwestern slope of Sierra Ne- 
vada de Santa Marta, Magda- 
lena. 

H. H. Smith; O. concolor. 

Muzo (1,000-1,300 m.), 5°30’ N., 
74°10’ W. 

Western slope of the Cordillera 
Oriental, Boyaca. 

P. Hershkovitz; O. concolor. 

Paime (1,038 m.), 5°25’ N., 74°10’ 
W. 

Cundinamarea. 

Hermano Nicéforo Maria; O. con- 
color. 


ARBOREAL RICE RATS—HERSHKOVITZ 565 


CoLomB1a—Continued 
Palomino (about 1,200 m.), 11°17’ 
N., 73°34’ W. 

Northern slope of the Sierra Ne- 
vada de Santa Marta, Mag- 
dalena. 

W. W. Brown; O. concolor. 

Pueblo Bello (1,067 m.), 10°24’ N., 
73°43’ W. 

Southern slope of the Sierra Ne- 
vada de Santa Marta, Mag- 
dalena. 

P. Hershkovitz; O. concolor. 

Pueblo Viejo (853 m.), 10°58’ N., 
73°26’ W. 

Northern slope of the Sierra Ne- 
vada de Santa Marta, Mag- 
dalena. 

W. W. Brown; O. concolor. 

Purif (40 m.), 7°25" N.;°75°20' W. 

Rio Cauca valley above Caceres, 
Antioquia. 

P. Hershkovitz; O. concolor. 

San Antonio, 11°00’ N., 73°26’ W. 

Rio San Antonio, northern slope 
of the Sierra Nevada de Santa 
Marta, Magdalena. 

W. W. Brown; O. concolor. 

Sierra Negra (1,265 m.), 10°36’ N., 
72°55’ W. 

Western slope of the Sierra de 
Perij4, Magdalena. 

P. Hershkovitz; O. concolor. 

Socorré (100-150 m.), 7°51’ N. 
NGoOlLi W. 

Rio Sind, Cérdova. 

P. Hershkovitz; O. concolor. 

Tres Troncos (150 m.), 0°05’ N., 
CELT W: 

Right bank, Rfo Caquetd, La 
Tagua, Putumayo. 

P. Hershkovitz; O. concolor, O. 
bicolor. 


Villanueva (274 m.), 10°37’ N., 
72°58’ W. 
Rio Cesar Valley, Magdalena. 
P. Hershkovitz; O. concolor. 
Villavicencio (498 m.), 4°10’ N., 
73°40’ W. 
Meta. 
G. M. O’Connell; O. concolor. 


a 


Costa Rica 


Boruca (500 m.), 9°04’ N., 83°27’ W. 
Puntarenas. 
M. A. Carriker, Jr.; O. concolor. 
El General (625 m.), 9°21’ N., 
83°40’ W. 
Puntarenas. 
A. Smith; O. concolor. 
San Gerénimo (about 150 m.), 
9°39’ N., 84°18’ W. 
Rfo Pirris, San José. 
C. F. Underwood; O. concolor. 


EcuapDoR 


Avila (600 m.), 0°38’ S., 77°25’ W. 
Rio Suno, Napo-Pastaza. 

E. J. Brundage; O. bicolor. 
Canelos (530 m.), 1°35’ 8., 77°45’ W. 
Rio Bobonaza, Napo-Pastaza. 

G. H. H. Tate; O. bicolor. M.G. 
Palmer; O. concolor. 
Capahuari, Rfo (300 m.), 1°55’ S., 
77°20’ W. 
A tributary of the Rio Pastaza, 
Napo-Pastaza. 
R. Olalla; O. concolor. 
Chonta Ureu (700 m.) 
Upper Rio Napo, Napo-Pastaza. 
W. Clark-MacIntyre; O. concolor. 
Chonta Urcu, east (500 m.) 
Napo-Pastaza. 
C. Olalla; O. concolor. 
Copataza, Rfo (650 m.), 1°45’ S., 


77°50’ W. 
Enters Rio Pastaza, Napo-Pa- 
staza. 


M. Olalla; O. bicolor. 
Gualaquiza (885 m.), 4°22’ N., 
78°32! W. 
Rfo Gualaquiza, Santiago-Za- 
mora. 
O. bicolor. 
Jatun Yacu, Rfo (700 m.), 1°04’ S., 
00°47" We 
Rio Napo, mouth at Puerto Napo. 
W. Clark-MaclIntyre; O. bicolor. 
Jollin, Rio (700 m.), 0°45’ S., 77°40’ 
W. 
Enters the Misagualli east of 
Tena. 
W. Clark-MacIntyre; O. bicolor. 


566 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ecuapor—Continued 
Llunchi (250 m.), 0°37’ S., 76°46’ W. 
Rio Napo, Napo-Pastaza. 
P. Hershkovitz; O. concolor, O. 
bicolor. 
Montalvo, 1°40’ S., 77°20’ W. 
Rio Bobonaza, Napo-Pastaza. 
R. Olalla; O. bicolor, O. concolor. 
Napo, Rio 
Napo-Pastaza 
L. Séderstrém; O. concolor. 
Pambilar (20 m.), 1° N., 79° W. 
Esmeraldas, 
G. Fleming; O. bicolor. 
Paramba (1,100 m.), 0°49’ N., 
78°22’ W. 
Rio Mira, Imbabura. 
R. Miketta; O. bicolor. 
Pindo Yacu [Pinto Yacu] (250 m.), 
2°33) Si 6AS8 OW: 
Rio Bobonaza near junction with 
Rio Pastaza. 
R. Olalla; O. concolor. 
San José (below) (500 m.), 0°32’ S., 
77°26’ W. 
Rio Payamino, Napo-Pastaza. 
Olalla Bros.; O. bicolor. 
Suno, Rfo (below), 0°40’ S., 77°08’ 
W. 
Enters Rio Napo, Napo-Pastaza. 
Olalla Bros.; O. bicolor. 
Yana Rumi 
Napo-Pastaza. 
R. Olalla; O. concolor. 
Zamora (1,000 m.), 4°10’ S., 78°43’ 
W. 
Rio Zamora, Santiago-Zamora. 
H. E. Anthony; O. bicolor. W. B. 
Richardson; O. concolor. 


PANAMA 


Barro Colorado, Canal Zone. 
R. K. Enders; O. concolor, O. 
bicolor. 
Bogava [Bugaba (q.v.)] 
Bugaba (250 m.), 8°28’ N., 83°38’ 
W. 
Pacific coast, Chiriqui. 
H. J. Watson; O. concolor. 
Cana [Santa Cruz de Cana] (600 m.), 
7°43’ N., 77°40’ W. 
Darién. 
E. A. Goldman; O. concolor. 


PansMa—Continued 


Corozal, Canal Zone. 
O. concolor. 
Jesusito, Rio (near sea _ level), 
8°03’ N., 78°14’ W. 
A tributary of the Sambu, 
Darién. 
T. Barbour and W. 8. Brooks; 
O. bicolor. 
Tacareuna (560-600 m.), 8°05’ N., 
77°20’ W. 
Darién. 
H. E. Anthony; O. concolor. 


PERU 


Boca Colorado (279 m.), 12°30’ S., 
70°25’ W. 
Madre de Dios. 
C. Kalinowski; O. concolor. 
Camante (2,000 m.), 13°30’ S., 
70°50’ W. 
Near Cadena, Marcapata, Cuzco. 
C. Kalinowski; O. bicolor. 
Chanchamayo (1,000 m.), 11°10’ S., 
75°20’ .W. 
Junin. 
J. M. Schunke; O. bicolor. 
Curaray, Rfo (mouth) (140 m.), 
2°26’ S., 74°04’ W. 
Loreto. 
Olalla Bros.; O. bicolor. 
Exito (Hacienda) (940 m.), 9°25’ 8. 
Rio Cayumba, Huanuco; the 
Cayumba enters the Huallaga. 
E. Heller; O. concolor. 
Itahuania, 12°40’ S., 71°10’ W. 
Madre de Dios. 
C. Kalinowski; O. concolor. 
Lagarto Alto, 5°55’ S., 74°35’ W. 
Rio Ucayali, Loreto; the coordi- 
nates are those of ‘‘Lagarto,’’ 
which may not be the actual 
collecting locality. 
Olalla Bros.; O. bicolor. 
La Pampa (573 m.), 13°38’ S., 
69°36’ W. 
On road between Limbani and 
Astilleros, Puno. 
O. Pearson; O. concolor. 
Mazdn, Rio (100 m.), 3°20’ &., 
73°05’ W. 
Tributary of Rio Napo, Loreto. 
Mendo B.; O. bicolor. 


ARBOREAL RICE RATS—HERSHKOVITZ 567 


PEeru—Continued 
Monte Alegre 
Presumably in Loreto Depart- 
ment. 
G. Tessman; O. bicolor. 
Moyobamba (820 m.), 6°03’ S., 
76°58’ W. 
At head of Rfo Moyobamba, 
Amazonas. 
L. Rutter; O. concolor. 
Pozuzo, 9°55''S.; 75°35! W. 
Huanuco. 
E. Heller; O. bicolor. 
Pucallpa (180 m.), 8°25’ S., 74°35’ 
W. 
Loreto. 
J. M. Schunke; O. bicolor. 
Puerto Indiana (100 m.), 3°20’ &S., 
72°40’ W. 
North bank of Rio Marajion, 
above mouth of Rio Napo, 


Loreto. 
Olalla Bros.; O. bicolor. 
Quincemil (680 m.), 13°17’ S., 
70°35’ W. 


Rio Marcapata, Cuzco. 
C. Kalinowski; O. concolor, O. 
bicolor. 
Sagrario (1,020 m.), 
69°41’ W. 
Upper Rio Inambari, Puno. 
P. O. Simons; O. bicolor. 
San Jerénimo (300 m.), 7°45’ S., 
74°50’ W. 
Rio Ucayali, Loreto. 
R. W. Hendee; O. concolor, O. 
bicolor. 
Santiago, Rio 
J. M. Schunke; O. bicolor. 
Sarayacu, 6°45’ S., 75°15’ W. 

Rio Ucayali, Loreto. 
Olalla Bros.; O. bicolor. 
Satipo, 11°12’ S., 74°32’ W. 

Junin. 
O. Pearson; O. concolor. 
and Gardner; O. bicolor. 
Tambo Yacu (1,000 m.), 6°00’ S., 
1°10) We 
Rioja, San Martfn. 
Osgood and Anderson; O. bicolor. 
Tingo Marfa (700 m.), 9°10’ &., 
75°45’ W. 
Rio Huallaga, Huanuco. 
R. W. Hendee; O. concolor. 


13°55! S., 


Pearson 


PreRu—Continued 
Villa Carmen (Hacienda) (600 m.), 
12°52’ S., 71°15’ W. 
Cosiiipata, Cusco. 
C. Kalinowski; O. bicolor. 
Yarinacocha (160 m.), 10°15’ S., 
74°45’ W. 
Rio Ucayali, Loreto. 
J. M. Schunke; O. bicolor. 
Yurac Yacu (800 m.), 6° S., 77° W. 
San Martin. 
R. W. Hendee; O. concolor, O 
bicolor. 


TRINIDAD 


Caparo 
Caroni, Port of Spain. 
F. M. Chapman; O. concolor. 
Caura 
Caroni. 
F. M. Chapman; O. concolor. 
Cumaca 
W. G. Downs; O. concolor. 
Mayaro, 10°20’ N., 61°00’ W. 
Southeastern Trinidad. 
O. concolor. 
Prince’s Town, 10°16’ N., 61°22’ W. 
Victoria. 
F. M. Chapman; O. concolor. 
San Rafael, 10°30’ N., 61°15’ W. 
F. Wonder; O. concolor. 
St. Pats 
W. G. Downs; O. concolor. 
VENEZUELA 
Agiita (1,000 m.), 3°22’ N., 65° 
35’ W. 
Southeast slope of Mt. Duida, 


Amazonas. 
G. H. H. Tate; O. concolor. 


Arabupu:’ (280) +m-)}'' 5°03" N.., 
60°39’ W. 

South side of Mt. Roraima, 
Bolivar. 


G. H. H. Tate and T. D. Carter; 
O. concolor. 
Auyantepuf (1,100 m.), 5°45’ N., 
62°30’ W. 
East of Rio Caroni, Bolfvar. 

G. H. H. Tate; O. concolor. 
Caicara’ (146° m.), 7°38’ N.; 66° 
10’ W. 
Right bank 
Bolivar. 
G. K. Cherrie; O.’concolor. 


of Rio Orinoco, 


568 


VENEZUELA—Continued 


Cafio. Seco\,.(700', m:),., 3221’ “N., 
65°35’ W. 
Southeast slope of Mt. Duida, 
Amazonas. 
Olalla Bros.; O. bicolor. 
Caroni, Rfo (1,100 m.), 6°00’ N.., 
62°45’ W. 
Cerro Auyantepuf, Bolfvar. 
G. H. H. Tate; O. concolor. 
Casiquiare, Rfo (80 m.), 2°48’ N., 
65°55’ W. 
Two miles west of Raudal Yamadd, 
Amazonas. 
G. H. H. Tate, Olalla Bros.; 
O. concolor. 
Chama (Cafetos de) (1,600 m.), 
8°33! N., 71°13" W.. 
Rio Chama, Mérida. 
S. Bricefio; O. concolor. 
Cogollo, Rio (100 m.), 9°55’ N., 
72°12’ W. 
Zulia, Perija. 
W. H. Osgood and H. B. Conover; 
O. concolor. 
El Llagual [Yagual], 7°12’ N., 
64°58’ W. 
Lower Rio Caura, Bolivar. 
M. A. Carriker, Jr.; O. concolor. 
El Merey, 3°06’ N., 65°53’ W. 
Rio Casiquiare, Amazonas. 
Olalla Bros.; O. concolor. 
Esmeralda (143 m.), 3°11’ N., 
65°33’ W. 
Mt. Duida, north bank of upper 
Orinoco. 
G. H. H.. Tate, Olalla Bros.; 
O. concolor. 
Thuapo, 7°00’ N., 65°27’ W. 
Rio Orinoco, Amazonas. 
Olalla Bros.; O. concolor. 
La. Azulita (1,135 m.), 8°43’ N., 
71°26" W. 
Mérida. 
W. H. Osgood and H. B. Con- 
over; O. concolor. 
La Uni6én (150 m:);),7°05’ N., 
65°52’ W. 
Rio Caura, Bolfvar. 
S. M. Klages; O. bicolor. 
Los Palmales, 10°30’ N., 64°10’ W. 
East of Cuman4, Sucre. 
F. M. Urich; O. concolor. 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 11¢ 


VENEZUELA—Continued 


Lourdes (Cafetos de) (1,800 m.), 
SoSSUAING, (Ml 20M, awe 
Mérida. 
S. Bricefio; O. concolor. 
Mérida (1,641 m.), 8°36’ N., 71°09’ 
W. 
Mérida. 
O. concolor. 
Mérida (Cafetos de) (1,630 m.), 
8$°36/Ni 71°09" We 
Mérida. 
S. Bricefio; O. concolor. 
Milla (Cafetos de) (1,630 m.), 
8°43" N;, 2°00" Wi: 
Mérida. 
S. Bricefio; O. concolor. 
Milla (Montes de) (1,630-1,670 m.), 
SP43"" Nua wl, We 
Mérida. 
S. Bricefio; O. concolor. 
Montes de la Otra Banda (1,600- 
1,630 m.), 8°37’ N., 71°10’ W. 
Mérida, Mérida. 
S. Briceiio; O. concolor. 
Neveri (750 m.), 10°06’ N., 64°37’ 
W. 
Sucre. 
G. H. H. Tate and H. J. Clement; 
O. concolor. 
Ocama, Rfo (150 m.), 2°47’ N., 
65°15’ W. 
At mouth of Rfo Orinoco, Ama- 
zonas. 
Olalla Bros.; O. concolor. 
Quebrada Seca, 10°15’ N., 63°55’ 
W. 
Rio Manzanares, Cumanacoa, 
Sucre. 
F. W. Urich; O. concolor. 
Rancho Grande (1,100 m.), 10°21’ 
N., 67°41’ W. 


Aragua. 
G. H. H. Tate and W. Beebe; 
O. bicolor. 


San Rafael (900 m.), 10°05’ N., 
64° W. 
Cumanacoa, Sucre. 
H. J. Clement; O. concolor. 
Yurudn, Rfo, 6°42’ N., 61°37’ W. 
Bolivar. 
M. A. Carriker, Jr.; O. concolor 
O. bicolor. 


PROC U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 1 


palusiris 


) 


(Oryzon ys 


OP 


—~ & 
& 28 
EOC 
Symes 
a Z, 
S$ 3 
~ 

mS 
~~ c=) 


14); b, 


84 
R} ip 1d0 mys 


=, 
3 


a, 


) } 7 olor (CNHM 


(CNHM 48455); 


omys (Oecomy. 


Ory 


a, 


| yy rsal 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 2 


a, Oryzomys bicolor; b, O. concolor; c, O. palustris; d, Rhipidomys venustus. 


<2) of same skulls shown in plate I: 


Ventral aspect ( 


PROG. US. NAT. MUS: VOL. 110 HERSHKOVITZ, PLATE 3 


Lateral aspect (2) of same skulls (with mandibles) shown in plates 1, 2: a, Orvzomys 


bicolor; b, O. concolor; c, O. palustris: d, Rhipidomys venustus. 
t B 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 4 


Upper and lower molars (10) of same specimens (except as noted) shown in plate 3: a, 
Oryzomys bicolor (CNHM 18541); 6, O. conc olor; c, O. palustris; d, Rhipidomys venustus. 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLAMTE 5 


Dorsal aspect (1-++) of skull in certain subspecies of Oryzomys concolor (a-e) and O. bicolor 


(f-7), showing variation in size with broad overlapping in the opposite extremes of each 
species. d-c, Oryzomys concolor superans: a, Male (CNHM 41459), Montalvo, Rio 
Bobonaza, Ecuador; b, female (CNHM 43257), Rio Pindo Yacu, Ecuador; c, female 
(CNHM 43252), Yana Rumi, Ecuador. d, e, Oryzomys concolor concolor: d, male (USNM 
280563), Pueblo Bello, Colombia; ¢, female (USNM 280583), El Orinoco, Rio Cesar, 
Colombia. f, g, 1, Oryzomys bicolor bicolor, males: f (CNHM 18541), Holmia, British 
Guiana; g (UMMZ 80162), 7 (UMMZ 80147), both from Llunchi, Rio Napo, Ecuador. 
h, J, Oryzomys bicolor phaeotis: h, Female (CNHM 84314), Villa Carmen, Cosnipata, Peru; 
j, male (CNHM 68638), Camante, Marcapata, Peru. 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 6 


Ventral aspect of (1-++) same skulls in same order as shown in plate 5: Oryzomys concolor 


(a-e) and O. bicolor (7 Hh 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 7 


Dorsal and ventral aspect of skull (2) of: a, Oryzomys bicolor (CNHM 84314); 5, O. lon 
icaudatus (CNHM 24593), 


HERSHKOVITZ, PLATE 8 


110 


U.S. NAT. MUS. VOLE 


PROC 


INHNO) 


5 NJOIYIUDX ‘O ye!) 


aK 


IPTST INHNO) Snevyqns ‘O° 


‘ 


=( 


‘(90648 INHNO) £42240] ‘O ‘2 *(9SF61 


ZS7EF INHINO) 40702"02 0 “4 *(HIEF8 INHNO) 4079209 Shmozhsg ‘D :(Z 


) [[nys Fo qoodse 


[estoqd 


PROG. USS NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 9 


laticep 


O 


] 


YANTNEOLUS; @, 


(OR 


as 


subflavus; 


(Oy 


h 


Oryzom AVA) bicolor: 


( 


) of same skulls shown in plate 8: 


4 


{ 


aspect 


Ventral 


PROC. U.S. NAT. MUS. VOL. 110 HERSHKOVITZ, PLATE 10 


Lateral aspect (2) of same skulls shown in plates 8, 9: a, Oryzomys bicolor; b, O. concolor; 
I i : 3 
c, O. subflavus; d, O. xantheolus; e, O. laticeps. 


PROG. WS. .NAdTIMUS. VOL. 110 HERSHKOVITZ, PLATE 11 


Lateral aspect (2) of left mandibles of same skulls shown in plates 8-10: a, Oryzom 


,O. subflavus; d, O. xantheolus; 


bicolor; b, O. concolor; ¢ 


PROC. US. NAT. MUS. VOL 110 HERSHKOVITZ, REAGE 12 


Upper and lower molars (>< 10) of same specimens shown in plates 8-11: a, Oryzomys bi- 


olor; b, O. concolor; c, O. subflavus; d, O. xantheolus; e, O. laticeps. 


PROCEEDINGS OF .THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U.S. NATIONAL MUSEUM 


Vol. 110 Washington: 1959 No. 3421 


THE RODENT-INFESTING ANOPLURA (SUCKING LICE) OF 
THAILAND, WITH REMARKS ON SOME RELATED SPECIES 


By Puy.us T. Jounson ' 


During 1952-55, Robert E. Elbel, Malaria Control Adviser in the 
North Eastern Provinces, conducted wild-mammal- and_ bird-ecto- 
parasite surveys for the U.S. Operations Missions to Thailand, 
International Cooperation Administration. This first such compre- 
hensive survey in Thailand has proved to be a major contribution to 
mammalogy, ornithology, and entomology. Mr. Elbel’s collections 
are the result of one of the first wholehearted attempts to sample 
populations of a large and representative percentage of the rodent and 
rodent-ectoparasite species native to Southern Asia, and have therefore 
provided medical entomology with valuable and unique information 
on this area of the world. 

The present paper is an account of the Anoplura collected by Mr. 
Elbel and his associates, and of a collection made by D. C. and 
EK. B. Thurman in northeastern Thailand. Of the 18 species repre- 
sented, 8 are new.?. To date only 4 species of ‘Anoplura have been 
recorded from Thailand. Of these 4, only 1 was not included in 
Mr. Elbel’s collections (Hoplopleura pectinata Cummings from Fattus 
surifer, Trong, Lower Thailand). 


1 Entomology Research Division, Agricultural Research Service, U.S. Department of Agriculture. 
2 One of the new species collected during the survey will be described in another paper, and is not men- 
tioned here. 


569 


570 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


I am indebted to Mr. Elbel for submitting the Thai Anoplura 
collection for study and deposit in the collections of the U.S. National 
Museum. Ernestine B. Thurman and the late D. C. Thurman, Jr., 
kindly donated their collections of Thai Anoplura, which have served 
as a valuable supplement to Mr. Elbel’s collections. 

As dictated by the distribution of its hosts, the Thai anopluran 
fauna is mainly attributable to the Oriental Region, but contains 
elements from the Palaearctic and Australian Regions as well. One 
of the species, Polyplax spinulosa (Burmeister), has a cosmopolitan 
distribution on Rattus species. 

Probably at least half the species of rodent-infesting Anoplura in 
Thailand have now been collected. ‘The Anoplura of the primates 
and ungulates in Thailand remain completely unknown, and those 
from all the surrounding areas are very poorly known. 

The description of a new species of Neohaematopinus taken from a 
flying squirrel from Borneo is included in this paper because this 
species is very closely allied to one taken from Thai flying squirrels. 
I am indebted to Lt. Col. Robert Traub, M.S.C., U.S. Army, for 
donating the specimens from Borneo. 

David H. Johnson, curator of mammals, U.S. National Museum, 
determined the hosts reported in this paper. Skins and skulls of 
these mammals are in the collections of the Division of Mammals. 
I wish to thank Dr. Johnson for his very necessary and helpful coop- 
eration. 

Unless otherwise indicated, all the specimens reported are from 
Thailand. If there are only two elements in a locality name, they 
represent the province and the village (Ban) or mountain. If three 
or more elements are included, they represent the province, the district 
(Muang), and the village or subvillage or mountain. Provincial 
names are given in small capital letters. Transliteration of the names 
follows the U.S. Army Gazetteer for Thailand (1944) with emenda- 
tions by H. G. Deignan. 

Collector’s names (Elbel and his associates) have been omitted 
except in data pertaining to the holotype of each of the new species. 
Individuals and organizations who cooperated with Mr. Elbel in 
obtaining the material were: Boonsong Lekagul and H. G. Deignan; 
and the Banpong Plague Laboratory, the Khorat Plague Laboratory, 
and the Nakhon Sawan Plague Laboratory. 


Enderleinellus corrugatus, new species 
Figures 1-5 


Typr pata: Holotype male from Tamiops macclellandi, cHAIYA- 
PHUM: Phukhieo, Ban Kaeng, Ban Lat, January 16, 1952, RE-355, 


ANOPLURA OF THAILAND—JOHNSON 571 


= lala 
An t \ > 
Toa. 


——— 

UD .. 
pe —t 
= x : 


SS 


eae 
f 
Ey 


= 


Figures 1—5.—Enderleinellus corrugatus, new species: 1, Paratype 2; 2, Holotype &; 3, 
Genitalia, holotype o&; 4, Paratergal plates II-VI, paratype 2 ; 5, Thoracic sternal plate, 
holotype oc’. 


R. E. Elbel, collector. One paratype female from Tamiops mac- 
clellandi, cHIENGMAI: Chiengmai, Ban Den, February 6, 1952, 
D. C. and E. B. Thurman collectors. 

Holotype deposited in the collections of the U.S. National Museum, 
type-catalog No. 64381. 

Diaenosis: FE. corrugatus is close to FE. malaysianus Ferris and 
E.. dremomydis Ferris. The thoracic sternal plate and male genitalia 
are much like those of malaysianus. It is easily distinguished from 
malaysianus in that paratergal plates II-V are heavily ridged and 
the spiracles are not particularly enlarged. FE. corrugatus has the 
head almost as broad as long but not noticeably longer, and has 
spiracles on abdominal segments 3-8, not 3-5 as does malaysianus. 
In corrugatus, large distinct paratergal plates are present on segments 
2-5, a very small plate is present on segment 6, and areas of sclerotiza- 
tion occur around the spiracles on segments 7 and 8. FE. malaysianus 
completely lacks plates on segments 6-8. LF. corrugatus is distinct 
from EF. dremomydis, which has paratergal plates on segments 2-6, 
in that dremomydis lacks rugosities on these plates, has the plates 
differently shaped, and has different setation. 


ie PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Description: Male: Head almost as broad as long; with marked 
postantennal angles; lateral margins slightly convergent posteriorly; 
broadly rounded anteriorly; venter very scaly; rostrum opening 
ventrally; antennae set well back from anterior head margin; seg- 
ments of antennae distinct and as broad or broader than long. Thorax 
about as long as head; sternal plate oval, heavily sclerotized laterally. 
Legs with first and second pairs with well-developed tarsal thumb 
opposing claw; coxae of first and third pairs noticeably rugose. 
Abdomen with medium-sized spiracles on segments 3-8; large par- 
atergal plates on segments 2-5; a very small plate on segment 6; 
small sclerotizations around spiracles of segments 7 and 8. Par- 
atergal plate II with one apical seta; plate III with two setae about 
half length of plate; plate IV with one seta somewhat shorter than 
plate and one minute seta; plate V with one short and one minute 
seta; plate VI with two minute setae. Dorsally with one row of 
setae per segment, a narrow plate associated with each row; setae 
on segments 4-7 large and inflated, these rows continuous across 
dorsum and containing 10, 11, 12, and 8 setae, respectively. Ventrally 
with plates associated with rows of setae on segments 2 and 3 only; 
setae of segments 3-6 inflated. Genitalia having basal plate with 
posterior arms about as long as undivided anterior part; endomeral 
piece broadly U-shaped, with median fracture; pseudopenis elongate- 
triangular, posterior part lacking rugosities. 

Female: Head, thorax, and legs asin male. Abdomen with para- 
tergal plates as in male. Veuntrally, plates associated with rows of 
setae on segments 2 and 3 only; segments 3-7 with setae inflated; 
6 setae in rows on segments 3 and 4; 9-11 setae in rows on segments 
5-6; segment 7 lacking setae medially, with 2 setae laterally on 
each side. Dorsally, plates associated with anterior 4 rows of setae; 
setae of segments 4-8 inflated ; rows on segments 5-7 containing 11-14 
setae, extending completely across dorsum. Genitalia as in figure 1, 

Lengths: Male 0.58 mm.; female 0.71 mm. 


Enderleinellus dremomydis Ferris 
Enderleinellus dremomydis Ferris, Contributions toward a monograph of the 
sucking lice, pt. 1 (1919), p. 29, fig. 16, 1920; The sucking lice, p. 108, 
1951. 
This species was described from two females found on Dremomys 
pernyi, West Szechuan, China. 
New reEcorpD: One female from Dremomys rufigenis, Lornt: Dan 
Sai, Kok Sathon, Phak Khi Nak Mountain, March 29, 1955, RE-5189. 
This Thai record is the first since the original description. The 
specimen agrees well with the original description and the female 
paratype in the U.S. National Museum collections. The male of 
dremomydis remains unknown. 


ANOPLURA OF THAILAND—JOHNSON 573 


Enderleinellus kumadai Kaneko 
FicuREs 6-9 


Enderleinellus kumadai Kaneko, Bull. Tokyo Medical and Dental Univ., No. 1, 
p. 49, figs. 1, 1-8, and 2, 1-2, 1954. 

Kaneko described this species from the holotype male, allotype 
female, and four male and seven female paratypes, collected from 
Callosciurus erythraeus thaiwanensis, O-shima, Tokyo, Japan. ‘The 
host squirrels are native to Formosa and had escaped from a zoo 
during World War II to become established on O-shima. 

New recorps: Three males, one female from Callosciurus sp., 
NAKHON SAWAN: Bon Phot Phisai, Tha Ngiu, March 30, 1953, RE- 
1740. Two females, same data, but RE-1741. One male, one 
female, same data, but RE-1743. One female from Callosciurus 
finlaysoni, CHATYAPHUM: Phukeio, Ban Kaeng, Ban Non Taolek, 
January 16, 1952, RE-361. 

The specimens reported above agree quite well with Kaneko’s 
original description and figures. HE. kumadai is related to EL. malay- 
sianus Ferris, LE. menetensis Ferris, and: /. larisci Ferris—all from 
Asian squirrels; it may be separated from malaysianus in having 
paratergal plates on segments 2-4 (2-5 in malaysianus) ; and it differs 


Ficures 6-9.—Enderleinellus kumadai: 6, Genitalia, @'; 7, 9 ; 8, Thoracic sternal plate, o; 
9, Paratergal plates, o’. 


574 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


from menetenis, which also has three paratergal plates, in that spiracles 
are located on segments 3-5, not 3-6 (or 3-8), and the thoracic 
sternal plate has the sclerotized portion dumbbell-shaped in kumadai 
rather than entire, and in that the apical setae of paratergal plate 
ILI are longer than the plate in kumadai. FE. kumadai is closest to 
LE. larisci but is easily separated as follows: LE. larisci lacks definite 
postantennal angles on the head and has no apical setae on para- 
tergal plates IT and IV. E. kumadai has one apical seta on plate 
II and two on plate IV. In male larisci the endomeres are broadly 
separated and the pseudopenis very small, whereas kumadai has 
the endomeres joined medially (although a fracture line is visible 
in some specimens), and the pseudopenis is large and as long as the 
parameres. 

The Thai specimens of kumadai were compared with two male and 
two female paratypes in the collection of Captain John E. Scanlon. 


Enderieinellus malaysianus Ferris 


Enderleinellus malaysianus Ferris, Contributions toward a monograph of the 
sucking lice, pt. 1 (1919), p. 12, figs. 3-4, 1920; The sucking lice, p. 110, 
1951.—Werneck, Mem. Inst. Oswaldo Cruz, vol. 45, pp. 281, 302, 1947. 

Ferris described malaysianus from material collected off Calloscwurus 
caniceps and Callosciurus prevosti from Malaya, Lower Siam, and 
Borneo. The holotype is from C. caniceps, St. Lukes Island, Mergui 
Archipelago, Malaysia. 

New reEcorD: One female from Callosciurus caniceps, NAKHON 
SsAWAN: Pak Nom Pho, Khao Khat, May 4, 1953, RE-2503. 

This female has spiracles considerably smaller than the holotype 
and paratypes in the U.S. National Museum, but agrees well in other 
respects. 

Enderleineflus menetensis Ferris 


Enderleinellus menetensis Ferris, Contributions toward a monograph of the 
sucking lice, pt. 1 (1919), p. 14, figs. 5-6, 1920; The sucking lice, p. 110, 
1951.—Werneck, Mem. Inst. Oswaldo Cruz, vol. 45, p. 281, 1947. 

Ferris described FE. menetensis from a series of males and females 
taken from Menetes berdmorei rufescens, Koh Kut Island, Southeast 
‘Siam. The holotype is a male. 

New record: One male, one female from Menetes berdmorei, 
KANCHANABURI: Latya, May 17, 1952, F—752. 

These specimens agree well with the original description and figures 
except that both sexes have more numerous setae on the abdomen 
dorsaily and ventrally, chiefly by reason of having three to five setae 
in each lateral group rather than one or two. Spiracles are present 
on abdominal segments 3-8 in the present specimens. The spiracles 
are very small and hard to see unless the specimen is in a favorable 


ANOPLURA OF THAILAND—JOHNSON 575 


position. The paratype female in the U.S. National Museum col- 
lections has apparent spiracles only on segments 3-6. It is possible 
that this species has a variable number of spiracles. 


Hoplopleura malabarica Werneck 


Fiacures 12-14, 16 


Hoplopleura malabarica Werneck, Rev. Brasileira Biol., vol. 14, No. 1, p. 113, 
figs. 8-12, 1954. 

Werneck’s type series consisted of the male holotype, female 
allotype, and one male and four female paratypes from Bandicota 
malabarica Shaw, Colombo, Ceylon, July 28, 1944, collected by C. D. 
Radford. 

New recorps: Many specimens in 27 collections from Bandicota 
sp., Bandicota bengalensis ssp., and Bandicota indica ssp. from the 
following localities: cHatyApHUM: Phukhieo, Ban Kaeng, Ban Lat. 
KANCHANABURI: Tamaka, Luk Kae; Tamaka, Tharua; Latya, Ban 
Nua; Bo Phloi; and Kanchanaburi District. Loppurt: Khao Oerawan 
Mt. NAKHON SAWAN: Pak Nam Pho, Khao Khat Mt. rar Burt: 
Banpong, Pakraet; Banpong, Boekphrai; and Paktho, Phuchik. 

H. malabarica is a parasite of Bandicota species, and although 
Rattus species may be found in similar ecological situations together 
with Bandicota, in only one instance did Elbel collect malabarica 
from Rattus. 


Ficures 10-16.—Hoplopleura malabarica and H. hirsuta: 10, Paratergal plates, H. hirsuta, 
2 ; 11, Same, o; 12, Same, H. malabarica, 2 ; 13, Same, &; 14, Thoracic sternal plate, H. 
malabarica, &; 15, Genitalia, H. hirsuta, &; 16, Same, H. malabarica, &. 


576 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Werneck compared malabarica to H. hirsuta Ferris (from Sigmodon 
in the New World). The long series of individuals of malabarica 
collected by Elbel makes possible an expanded diagnosis and com- 
parison of this species with hirsuta. 

In most respects the two species are similar. The thoracic sternal 
plate is identical and the shape of the paratergal plates very similar. 
The lateral setae of the abdomen (those occurring off the tergal and 
sternal plates) are noticeably heavier in hirsuta, although a comparison 
of actual specimens is necessary to satisfactorily show this difference. 
The form of the paratergal plates and their setation is highly variable 
in both species. Female malabarica always have one long apical 
seta on plates IV-VI (on VI this seta may be very long); the second 
apical seta on the above plates is minute or absent. One abnormal 
female examined had two long setae on paratergal plate IV. In hirsuta 
females plates IV-VI always have two long setae (fig. 10). Para- 
tergal plate VI of hirsuta usually has two apical angles, although in 
occasional specimens there are no free angles. Malabarica females 
have a dorsal angle on plate VI, but usually the ventral angle is 
rounded and not free. Generally, hirsuta females have more setae 
in the abdominal rows. Werneck draws and describes the female 
of malabarica as having neither tergal nor sternal plates on the typical 
abdominal segments. This condition was not true of the Thai 
specimens, the plates being invariably present although often ex- 
tremely narrow and incomplete. In hirsuta females there are always 
three or more setae laterally off each abdominal plate both dorsally 
and ventrally, while malabarica females have but one or two setae 
in these positions. 

In males of the two species the setation of the paratergal plates 
is extremely variable. In both hirsuta and malabarica the male 
usually has one long seta on plates [V—VI with the second seta minute 
or absent (compare figs. 11 and 13). However, hirsuta males may 
have two long setae on one or all of plates [V—VI, and both types may 
be found in a single collection. None of the malabarica males exam- 
ined had more than one long apical seta on plates [V-VI. Malabarica 
males generally have fewer setae in the abdominal rows than hirsuta 
males have. Werneck’s drawing of male malabarica shows no tergal 
abdominal plates. All the Thai males have a normal number of nar- 
row tergal plates. The male genitalia of malabarica and hirsuta differ 
in small but constant fashion. In hzrsuta the parameres (fig. 15) are 
strongly curved apically so that were the tips joined by an imaginary 
line the parameres would form an almost perfect semicircle apically. 
In malabarica the parameres are not so strongly curved inward. 
The pseudopenis of hirsuta is usually not serrate, and the angle at 
its broadest point is usually about halfway between the base of the 


ANOPLURA OF THAILAND—JOHNSON BT 


pseudopenis and the inner aspect of the tip. In malabarica the pseudo- 
penis is serrate, and the angle at its broadest point is always much 
nearer the base of the pseudopenis than it is to the joining of the 
inner aspect of the tip. 

Specimens of malabarica illustrated were from Bandicota benga- 
lensis, RAT BURI, Banpong, Boekphrai. Drawings of male and female 
paratergal plates of hirsuta were taken from specimens from Florida, 
and the male genitalia from a specimen from the Panama Canal 
Zone—all from Sigmodon. 


Hoplopleura pacifica Ewing 


Hoplopleura pacifica Ewing, Bernice P. Bishop Museum Bull., No. 14, p. 9, figs. 
1b-c, 1924.—Hopkins, Proc. Zool. Soc. London, vol. 119, p. 481, 1949. 
Hoplopleura oenomydis, Ferris, Bernice P. Bishop Museum Bull., No. 98, p. 121 
(in part), 1932; The sucking lice, p. 139 (in part), 1951.—Pritchard, Journ. 

Parasit., vol. 35, p. 374, 1947. 

Hoplopleura pacijica was described from five males and seven 
females collected off Rattus hawaiiensis [Rattus exulans hawatiensis], 
Hawaiian Islands. 

New recorps: A long series of specimens from Rattus rattus ssp. 
from the following provinces: Chiengmai, Rat Buri, Chaiyaphum, 
Nakhon Ratchasima (—Khorat), and Chiang Rai. 

H. pacifica is the characteristic louse parasite of Thai F. rattus. 
Hopkins (1949) records all Hoplopleura specimens from Rattus 
species as pacifica, not H. oenomydis Ferris, and notes that H. oenomy- 
dis is restricted, so far as is known, to the African murine genus 
Oenomys. 

Diagnoses and descriptions of oenomydis and pacifica, together with 
host records, are being prepared for publication by Captain John E. 
Scanlon; therefore, no diagnoses are included in this paper. 


Hoplopleura thurmanae, new species 
Fiaures 17-19, 22-23, 25 


Tyre pata: Holotype female from Tamiops macelellandi, wort: 
Dan Sai, Na Phung, Ban Khok, Namlang Mountain, May 29, 1955, 
RE-5416, R. E. Elbel collector. Allotype male, same data, but 
June 3, 1955, RE-5512. Paratype male, same data but May 23, 
1955, RE-5290. Paratype female, same data, but May 29, 1955, 
RE-5415. Paratype female, same data, but June 6, 1955, RE-5555. 
Paratype female, same data, but June 1, 1955, RE-5460. Four 
male, one female paratypes from Tamiops macclellandi ssp., CHAIYA- 
PHUM: Phukhieo, Ban Kaeng, Ban Lat, January 16, 1952, RE-355. 
Four male, four female paratypes, same data, but January 18, 1955, 


505126—59——_2 


578 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficures 17-18.—Hoplopleura thurmanae, new species: 17, Holotype 9; 18, Allotype o. 


RE-369-371. One male, one female paratypes from Tamiops 
rodolphei, locality as above, December 19, 1952, RE-931. One 
female paratype from Tamiops rodolphei, Phukheio, Ban Kaeng, 
Ban Non Taolek, December 14, 1952, RE-900. One male paratype 
from Tamiops sp., KANCHANABURI: Bo Phloi, March 26, 1952, 
RE-563. One male, five female paratypes from Tamiops macclel- 
landi, cu1ENGMAI: Chiengmai, Ban Den, February 6, 1952, D. C. 
and E. B. Thurman collectors, TT—23—24. 

Holotype deposited in the U.S. National Museum, type-catalog 
No. 64382. 

Also examined: One male, three females from Menetes berdmorei 
ssp., KANCHANABURI: Bo Phloi, July 16, 1952, Y—308. 

This species is named for my friend and colleague, one of the 
collectors, Dr. Ernestine B. Thurman, 

Dracnosts: This species is very closely related to Hoplopleura 
erismata Ferris from Callosciurus species. The female of H. thurmanae 


ANOPLURA OF THAILAND—JOHNSON 579 


has the posterior angles of paratergal plates IV-VI produced into 
acute points that are almost as long as the apical setae. In erismata 
the apical angles of paratergal plates ['V—V are very short, obviously 
much shorter than the apical setae, and paratergal plate VI has 
scarcely any angles. In the male of thurmanae, the apical angles of 
paratergal plates 1V—VI are not so long as in the female, but definitely 
longer than in erismata. H. thurmanae also differs from ertsmata in 
the shape of the thoracic sternal plate (compare figs. 25 and 26). 
In thurmanae the plate has definite lateral and anterior angles; in 
erismata the lateral angles are smoothed out so that the lateral and 
anterior margins form a continuous curve. Male genitalia of the 
two species are quite different (compare figs. 23 and 24). 

Description: Female: Head rounded anteriorly, postantennal 
angles definite but rounded; postantennal head margins slightly 
convergent posteriorly. Thorax with thoracic sternal plate diamond- 
shaped, the posterior angle more produced than the anterior angle. 
Abdomen with outermost of the two modified setae on the first 
sternal plate of segment 3 usually definitely bent. Typical abdominal 
segment with three narrow sternal and tergal plates; both dorsally 
and ventrally with one seta lateraily on each side, off the plates. 
Paratergal plates as in figure 22; plates IV-—V produced inward 
anteriorly toward the midline both dorsally and ventrally. Plates 
IV-VI with apical angles produced mto long acute points which 
are almost as long as the apical setae; plate VII lacking posterior 
angles but not much reduced. 

Male: Head as in female except third antennal segment with a 
short stout dorsal seta. Thorax as in female. Abdomen with outer- 
most of the two modified setae on the first sternal plate of segment 3 
usually definitely bent. Sternal and tergal plates as in genus, nar- 
rowed but not strongly reduced; one or two setae laterally off tergal 
plates of segments 4—6, and one seta laterally on sternites 4-6, midway 
between the two sternal plates of each of these segments. Paratergal 
plates IV-VI and sometimes VII anteriorly produced inward toward 
midline both dorsally and ventraily. Plate VII not noticeabiy re- 
duced; plates IV—VI with posterior angles produced into points 
that are not so long as in female, being no more than half the length 
of the apical setae. Genitalia with parameres poorly sclerotized, 
particularly basally; pseudopenis with triangular flanges on each 
side at about middle, its apex triangular, slightly serrate. 

Lengths: Female 1.15-1.2 mm.; male 0.85 mm. 

One collection of H. thurmanae was taken from Menetes, but that 
this is a normal occurrence needs to be confirmed. 


580 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Hoplopleura erismata Ferris 


Figures 20-21, 24, 26 


Hoplopleura erismata Ferris, Contributions toward a monograph of the sucking 
lice, pt. 2, p. 113, figs. 72B, 72E, 72F, 1921; The sucking lice, p. 136, 1951. 
Ferris described erismata from Thailand from Callosciurus 
ferrugineus, C. caniceps, and Tamiops sp. Holotype female from 
C. ferrugineus, Southeast Siam. It is very likely that the specimens 
taken from Jamiops were actually H. thurmanae. 
New rREcoRDS: Two males, two females from Callosciurus finlaysoni 
tacherdi, NAKHON RATCHASIMA (=KHORAT): Sikhui, Pakchong, June 


Se a 
—— 
—_ 


/ 

fe 
Bw) 4 "\ 
4, (y ) 


Ficures 19-26.—Hoplopleura thurmanae, new species, and H. erismata: 19, Head, H, 
thurmanae, holotype 2 ; 20 Same, H. erismata, ? ; 21, Paratergal plates II-VI, H. erismata. 
2; 22, Same, H. thurmanae, holotype 9 ; 23, Genitalia, H. thurmanae, allotype &; 24, 
Same, H. erismata, oc’; 25, Thoracic sternal plate, H. thurmanae, holotype 9 ; 26, Same, 
H. erismata, 2. 


ANOPLURA OF THAILAND—JOHNSON 581 


6, 1952, Y-216-221. Five females from Callosciurus caniceps, CHIANG 
RAI: Chiang Saen Khao, Ban Sop Luak, March 2, 1953, RE-2340. 
One male from C. caniceps, KAMPHAENG PHET: Khlong Khlung, 
Ban Hua Thanon, April 4, 1953, RE-2365. Two males, five females 
from C. caniceps, NAHKON SAWAN: Pak Nam Pho, Khao Khat Mt., 
May 13, 1953, RE-2539. One female from C. caniceps, torr: Dan 
Sai, Kok Sathon, Phu Lom Lo Mt., February 28, 1955, RE-4824. 
A series of males and females from Callosciurus finlaysoni, Dan Sai, 
Kok Sathon, Phu Lom Lo Mt. and Phak Khi Nak Mt., February 
and March 1955, RE-4800, 5018, 5058, 5063, and 5105. A series of 
males and females from C. finlaysoni, Dan Sai, Na Phung, Ban 
Khok, Namlang Mt., May and June 1955, RE-5399, 5486, 5546, 
and 5585. 

H. erismata is easily determined by reference to the original descrip- 
tion and drawings by Ferris. It should be added that in the male, 
paratergal plates IV-VI (not IV and V as in the female) have the 
anterior angles strongly produced toward the midline of the body. 
The paired modified setae on the first sternal plate of abdominal 
segment 3 are often not so strongly developed as in most species of 
Hoplopleura, although this is not always the case, and sometimes the 
outer setae may be not only heavy but noticeably bent. 

Lengths: Male 0.95-1.1 mm.; female 1.3-1.5 mm. 


Neohaematopinus callosciuri, new species 


FIGuRES 27-28, 34b, 35c, 42-43 


Neohaematopinus sciurinus, Ferris, Contributions toward a monograph of the 
sucking lice, pt. 4, pp. 244, 245, 1923 (in part, records from Callosciturus).— 
Hopkins, Proc. Zool. Soc. London, vol. 119, pp. 459, 460, 1949 (records 
from Callosciurus). 

Type pata: Holotype male, allotype female, and two male and four 
female paratypes from Callosciurus jfinlaysoni, tort: Dan Sai, Kok 
Sathon, Phu Lom Lo Mt., February 26, 1955, RE-4800, R. E. Elbel 
collector. A series of 179 male and female paratypes from Callosciurus 
Jfinlaysoni, C. caniceps and C. erythracus from the following provinces: 
Loei, Chaiyaphum, Rat Buri, Kanchanaburi, Nakhon Sawan, Nakhon 
Ratchasima, Prachuap Khiri Khan, Chiang Rai, Kamphaeng Phet, 
and Lop Buri, collected 1952-55. Two male and one female para- 
types from C. erythraeus, Formosa: Ma-an Lio, April 12, 1952, D. H. 
Johnson collector, DJ—7274. One male and two female paratypes, 
same data, but April 19, 1952, H. J. Stanton collector, JHS-6. One 
male paratype, same data, but April 16, 1952, JHS-3. 

Holotype deposited in the collection of the U.S. National Museum. 
type-catalog No. 64383. 


582 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 110 


js NCS ) 

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>: Ne! nx A 

A, See 28 
et 
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iret So. Kx , SS zi hey 


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Ficures 27-33.—Neohaematopinus callosciurt, new species, N. scturinus, and N. semi- 
fasciatus: 27, N. callosciuri, allotype 9 ; 28, N. callosciuri, holotype &*; 29, Abnormal 
antenna, N. sciurinus, 2 ; Florida from S. niger; 30, Antenna, NV. sciurinus, 9 , Texas from 
S. niger; 31, Same, o'; 32, Antenna, N. semifasciatus, o', Alaska from T. hudsonicus; 33, 
Same, ?. 


I have also seen material of this species from Callosciurus notatus, 
North Borneo: Mount Kinabalu, Tenompak, June 18, 1951, collected 
by Traub. 

Dracnosis: N. callosciuri is very similar to N. sciwrt Jancke and 
N. sciurinus (Mjéberg), differing only in the shape of the basal 
antennal segment and size of its posteroapical enlarged seta. It is 
separable from sciurinus in that the enlarged seta on the first antennal 
segment is not as large in callosciuri (compare figs. 41 and 42). WN. 
callosciurt is distinct from sciuri in that the apical enlarged seta is 
borne on a slight prolongation of the basal antennal segment, whereas 
in sciuri the basal segment is not modified and the enlarged seta is 
set back from the margin (compare figs. 40 and 42). N. callosciurt 
differs from both sciuri and sciurinus in having a flattened, rather 
indistinct (but always present) tubercle on the venter of the basal 
antennal segment. 

Description: Male: Head broadly rounded anteriorly; postanten- 
nal angles marked; posterior head margins only slightly convergent 
posteriorly. Basal antennal segment produced posteroapically and 
bearing a stout seta; ventrally basal segment with flattened tubercle; 
third segment modified, bearing two short, stout setae apically. 
Thorax broader than long, about as long as head; sternal plate with 


ANOPLURA OF THAILAND—JOHNSON 583 


posterior angles produced into points, plate about as broad as long, 
its shape varying considerably. Legs typical of genus. Abdomen; 
ventrally, segment 2 with two plates; segment 3 with well-developed 
plate associated with anterior row of setae; vestige of a plate asso- 
ciated with second row of setae; segments 4-6 with one plate, asso- 
ciated with first of two rows of setae; lateral seta near paratergal 
plates on each side of segments 3-6. Dorsally, posterior plate of 
second segment emarginate medially and bearing several stout setae 
laterally on each side; remaining segments with one plate associated 
with single row of setae; segments 4-7 with one seta laterally off 
plate on each side. Paratergal plates with both posterior angles 
toothed; apical setae varying in length. Genitalia similar to other 
related species. 

Female: Head as in male except third antennal segment unmodified. 
Thorax and legs as in male. Abdomen having two rows of setae on 
segments 2-6 both dorsally and ventrally; plate associated with 
anterior row of each segment; posterior row with small areas of 
sclerotization around each setal base. Paratergal plates as in male. 
Genitalia not diagnostic. 

Lengths: Male 1.5-1.7 mm.; female 2.1-2.2 mm. 


Neohaematopinus cognatus, new species 


Fiaures 34a, 35d, 39, 44 


Neohaematopinus sciurinus, Ferris, Contributions toward a monograph of the 
sucking lice, pt. 4, p. 245 (in part, records from Menetes berdmoret), 1923.— 
Hopkins, Proc. Zool. Soc. London, vol. 119, p. 460 (records from M., 
berdmoret), 1949. 

Typn pata: Holotype male, allotype female from Menetes berd- 
more, LOEI: Dan Sai, Kok Sathon, Phu Lom Lo Mt., 8,000 ft., 
February 22, 1955, RE-4765, R. E. Elbel, collector. Three female 
paratypes with same data as holotype. A long series of paratypes, 
all from Menetes berdmoret, as follows: cHatyApHuM: Phukheio, Ban 
Kaeng, Ban Non Taolek, two females, January 16, 1952, RE-357. 
Phukheio, Ban Song, January 15, 1953, one female, RE-993. 
KANCHANABURI: Ban Nua, May 7, 1952, one male, three females, 
B-721, F-722, F-730. Tamuang City, May 12, 1952, one male, 
F-741. Kanchanaburi City, June 11, 1952, two females, F-878, 
F-881. Bo Phloi, March 13, 1952, three males, five females, RE-516. 
July 16, 1952, one female, Y-308. Purr BuRI: Khao Yoi, Nong 
Chum Phon, July 14, 1952, four females, F-981. NAKHON SAWAN: 
Phayuhakhiri, Ban Phayuhadaen, March 6, 1953, five males, five 
females, RE-1645. Wat Sai (forest), March 29, 1953, two males, 
one female, RE-1699. Pak Nam Pho, Khao Khat, April 10, 1953, 
one male, one female, RE-1738. May 7, 1953, five males, five 


584 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


o 


<i et 


resem 


Ficures 34-44.—Neohaematopinus callosciuri, new species, and related forms: 34, Para- 
tergal plates, 9 : a, N. cognatus; b, N. callosciuri; c, N. sciurinus; d, N. sciuri, England 
from S. vulgaris; 35, Genitalia, @: a, N. sciuri, England from S. vulgaris; b, N. sciurinus; 
c, N. callosciuri; d, N. cognatus, holotype; 36, Thoracic sternal plate, N. sciuri, 2 , from 
S. carolinensis: a, N. Carolina; b, Georgia; c, Maryland; 37, Same, N. sciurinus, ?, 
from S. niger: a, Maryland; b, Same; c, Ohio; d, Georgia; e, Texas; 38, Same, N. semi- 
fasciatus, 2 , irom Tamiasciurus: a, Alaska; b, Same; c, Montana; 39, Head, N. cognatus, 
allotype 9 ; 40, Same, N. sciuri, 9 ; 41, Same, N. sciurinus, 9 ; 42, Same N. callosciurt, 
allotype 9 ; 43, Thoracic sternal plate, N. callosciuri, allotype’ 9 ; 44, Same, NV. cognatus, 
allotype 9. 


ANOPLURA OF THAILAND—JOHNSON 585 


females, RE-2517. May 8, 1953, one female, RE-2520. May 8, 
1953, three males, two females, RE-2521. May 20, 1953, one male, 
one female, RE-2574. May 21, 1953, one male, RE-2577. May 21, 
1953, one male, two females, RE-2580. LopspBpurt: Khao Oerawan 
Mt., July 27, 1953, two males, one female, RE-2843. 

Holotype deposited in the collections of the U.S. National Museum, 
type-catalog No. 64384. 

DIAGNOSIS AND DESCRIPTION: In all ways N. cognatus is similar to 
N. callosciuri except that the basal segment of the antenna of both the 
male and female of cognatus lacks an enlarged posterior apical seta, but 
has a slightly enlarged seta set in from the apex, and that the basal 
segment is not prolonged posteroapically. 

Lengths: Male 1.4 mm.; female 1.8 mm. 


Neohaematopinus sciurinus and related forms 


Fieurses 31, 33 


For a number of years there has been confusion as to what name or 
names should apply to a group of Neohaematopinus species found on 
members of Sciurus and related genera of squirrels. Ferris (1951) 
reviewed the situation and reasoned that more than onespecies might be 
present but that final decisions would have to await the collection of 
further material. Names proposed in this group include the following: 

1. Haematopinus sciurinus Mjéberg, 1891, type host Sciurus niger, 
the North American fox squirrel. - (Acanthopinus antennatus Osborn, 
1910, from the same host, is a junior synonym of sciurinus Mjéberg.) 


2. Neohaematopinus antennatus semifasciatus Ferris, 1916, type host 
Scvurus douglasi (now known as Tamiasciurus douglasi), the pine or 
red squirrel. Ferris (1923) wrongly synonymized semifasciatus under 
sciurinus Mjoéberg. 

3. Neohaematopinus sciuri Jancke, 1931, type host Sciurus vulgaris, 
the European squirrel. 


4. Neohaematopinus callosciuri, type host Callosciuris finlaysoni 
from Thailand. 


5. Neohaematopinus cognatus, type host Menetes berdmorei from 
Thailand. 


At least a partial explanation of the confusing species tangle has 
been found through a study of the specimens described in this paper as 
N. callosciuri from Callosciurus spp., and N. cognatus from Menetes 
spp., plus numerous specimens of Neohaematopinus in the U.S. Na- 
tional Museum from North American Sciurus carolinensis, S. niger, 
and Tamiasciurus spp., and specimens from British Sciurus vulgaris 
(lent by T. Clay of the British Museum). Since very limited material 


586 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


has been collected from North American Sciurus species other than 
from S. carolinensis and S. niger, it is necessary to exclude such 
specimens from the following discussion. 

Slight but constant differences in the antennae, coupled with con- 
stant host associations, afford a reliable method for distinguishing 
species of the complex. A search for genitalic characters, size differ- 
ences, and other morphologic differences among the various species of 
the sciurinus complex has been fruitless. The thoracic sternal plate 
varies greatly in shape and size (figs. 36-38, 43, 44); the male genitalia 
are similar (fig. 35, a-d); and patterns of chaetotaxy, although some- 
what variable, are similar for all the species. 

There are two groups, based on the antennal characters: the scvuri- 
nus-like species have the basal antennal segment prolonged postero- 
apically, with an enlarged seta on the prolongation (figs. 41, 42); and 
the sciuri-like species lack the prolongation and may lack the enlarged 
apical seta (figs. 39, 40). 


Neohaematopinus sciurinus and related species 


1. N. scturinus Mjéberg is found on Sciurus niger in North America. 
It is characterized by having the basal antennal segment prolonged 
posteroapically, with this prolongation bearing a spinelike apical seta, 
by lacking a short spinelike seta on the posterior margin of the second 
antennal segment, and by having the fourth segment longer than the 
fifth (fig. 30). Specimens have been examined as follows: From 
Sciurus niger: Texas, Kansas, Florida, Georgia. South Carolina, 
Maryland, and Ohio. 

2. N. semifasciatus Ferris is found on Tamiasciurus douglasi and 
T. hudsonicus in North America. This species has the basal antennal 
segment with the posteroapical prolongation and seta as in sciurinus, 
but the second antennal segment has a short spinelike seta on the 
posterior margin, and the fourth segment is equal in length to the 
fifth (fig. 82). Specimens have been examined as follows: From 
Tamiasciurus douglasi: California and Oregon. From Tamiasciurus 
haudsonicus: Tennessee, Montana, and Alaska. 

3. N. callosciuri is found on Callosciurus species in Southeast Asia 
and Malaysia. It is characterized by having a tubercle on the venter 
of the basal antennal segment, by the posteroapical prolongation and 
its seta being smaller than in sciurinus, and by lacking a stout spine- 
like seta on the second antennal segment (fig.42). The Asian specimens 
from Callosciurus and probably those from Lariscus, reported by 
Ferris (1923) as sciurinus, are referable to N. calloscvuri. 

3 One female sciwrinus examined has a. abnormal seta on the basal antennal segment on one side (fig. 29), 
and several sciurinus specimens in three collections are recorded as from “‘red squirrel.”’ All the ‘‘red squir- 


rels’’ may have been misdetermined, and in one case the specimens were collected in Kansas where Tamias- 
ciurus (red squirrel) does not occur. 


ANOPLURA OF THAILAND—JOHNSON 587 


Neohaematopinus sciurt and related species 

1. N. sciuri Jancke apparently occurs on both European Seiurus 
vulgaris and North American Sezurus carolinensis. At least, the 
North American specimens are morphologically indistinguishable 
from the European ones. ‘This species is characterized by not having 
the posteroapical prolongation on the basal antennal segment, and 
by having a stout spinelike seta at the posteroapical angle (fig. 40). 
Specimens have been examined as follows: From Seiurus vulgaris: 
England. From Sciurus carolinensis: New York, Rhode Island, 
Maryland, District of Columbia, Virginia, North Carolina, Georgia, 
Florida, Mississippi, and Ohio. Also examined was one collection 
supposedly from “southern fox squirrel,” South Carolina. 

2. N. cognatus is found on Menetes berdmorei, Thailand. This 
species is like secur? in not having the basal antennal segment pro- 
longed posteroapically. It is further characterized by having the 
stout seta of the basal segment set in from the apex and by having a 
tubercle on the venter of the basal antennal segment (fig. 39). Thai 
specimens from MM. berdmorei reported by Ferris (1923) as sezurinus 
are referable to N. cognatus. 


Neohaematopinus capitaneus, new specries 
Ficures 45-47, 50, 56-57, 60 


Type pata: Holotype male, allotype female from Hylopetes 
phayret, CHAIyYAPHUM: Phukhieo, Ban Khon San, December 24, 1952, 
RE-953 and 959, R. E. Elbel collector. Two male, two female 
paratypes with same data as holotype. One male, six female para- 
types from Hylopetes phayrei; Phukhieo, Ban Kaeng, Ban Lat, 
January 17, 1952, RE-362. 

Holotype deposited in the collections of the U.S. National Museum, 
type-catalog No. 64385. 

Diagnosis: This species is near sciuropteri (Osborn) from North 
American Glaucomys. N. capitaneus is larger than sciuropteri (lengths: 
scvuroptert co 1.25-1.6 mm., 9 1.6-1.9 mm.; capitaneus & 1.8-2.2 
mm., 2 2.5-2.8 mm.). The thoracic sternal plate has the posterior 
angles not produced into such long points in capitaneus and is wider 
in relation to its length (compare figs. 59 and 60). The male genitalia 
differ considerably in the 2 species. N. capitaneus has the pseudo- 
penis rugose and the penis extending only about one-fourth the 
length of the parameres (compare figs. 54 and 56); and the terminal 
lobes of the abdomen are not so well developed as in seiuropteri. 
Particularly in the female of capitaneus, typical paratergal plates 
have small acute projections on either side of the apical setal bases. 
The female genitalia have the lateral setigerous lobes of the eighth 


588 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficures 45-46.—Neohaematopinus capitaneus, new species: 45, Allotype ? ; 46, Holotype c. 


segment (‘“gonopods’’) elongate in capitaneus (fig. 47), whereas these 
lobes are smaller and more nearly vertical in sciuropteri (fig. 49). 
Description: Male: Head broader than long, anterior margin 
very broadly rounded so that antennae are set near apex of head. 
Postantennal angles very prominent, postantennal margins parallel; 
occipital region strongly constricted; gular area strongly raised; basal 
antennal segment longer than broad, with stout seta dorsally, set 
well in from margin; third antennal segment modified, bearmg 2 
short stout apical setae. Thorax longer than head and somewhat 
broader. Thoracic sternal plate large, subrectangular, posterior 
corners produced into short points. Legs as in other members of the 
genus. Abdomen with 1 tergal and 2 sternal plates on each segment, 
typical sternal plate with row of 7 to 8 setae; typical tergal plate 
with row of 16-20 setae; 1 to 3 setae laterally off each plate dorsally 
and 1 seta laterally off second plate of each segment ventrally. Third 
tergal plate with posterior margin emarginate medially and with groups 
of stout lateral setae on each side. Paratergal plates large, with 2 
apical setae; paratergal plate VII extended somewhat dorsally so that 


ANOPLURA OF THAILAND—JOHNSON 589 


one of the lateral abdominal setae arises from it; plate VII dorsally 
continuous with tergal plate of segment 8. Genitalia with 
narrow basal plate; parameres of approximately same width through- 
out most of their lengths, acute apically, subapically with lateral 
triangulate lobe; penis opening above proximal fourth of parameres; 
pseudopenis with distal half rugose. 

Female: Head as in male except basal antennal segment as broad 
as long and third antennal segment not modified. Thorax and legs 
as in male. Abdomen with tergal and sternal plates well developed, 
two plates per segment dorsally and ventrally, typical ones each 
bearing a row of 6 to 10 setae; second sternal plate of segment 7 with 
3 or 4 short medial setae flanked by 2 long setae; 1 or 2 lateral setae 
off each plate. Paratergal plates much as in male, but plate VUT 
not joined to tergal plate of that segment; plates I1{-VI with pos- 
terior corners produced into short acute lobes, smaller acute lobes on 
each side of setal bases. Genitalia as in figure 47; genital plate (of 
segment 8) heavily spiculated; lateral setigerous lobes elongate. 

Lengths: Male 1.8-2.2 mm.; female 2.5-2.8 mm. 


Nechaematopinus kinabalensis, new species 
Fiaures 48, 51, 55, 61-63 


Tyre pata: Holotype male, allotype female from Hylopetes sagitta 
harrisoni, British North Borneo: Mount Kinabalu, Tenompak, 5,000 
ft., June 31, 1952, R. Traub collector, RT—B—9089. 

Holotype deposited in the collections of the U.S. National Museum, 
type-catalog No. 64386. 

Diacnosis: N. kinabalensis is closely allied to sciuropteri (Osborn) 
and capitaneus. It is somewhat larger than sciuropteri, and smaller 
than capitaneus. N. kinabalensis differs from capitaneus in that the 
basal antennal segment is as broad as long in the male, whereas in 
capitaneus this segment is longer than broad (compare figs. 50 and 51). 
The male genitalia are quite different in the two species. In kina- 
balensis the parameres are short, not enclosing the pseudopenis for 
most of its length, and their outer margins are convex (compare 
figs. 55 and 56). Females of capitaneus and kinabalensis are quite 
similar, differing mainly in the genital area. The lateral setigerous 
lobes of the eighth segment (‘“gonopods”) are smaller and more 
widely separated in kinabalensis, the lip of the vulva between these 
lobes is only slightly convex, and the genital plate is not so heavily 
spiculated (compare figs. 47 and 48). WN. kinabalensis differs from 
sciuropteri in having the thoracic sternal plate differently shaped, with 
the posterior points not so long (compare figs. 59 and 61). N. kina- 
balensis males have the basal antennal segment as broad as long, 
not longer than broad as in sciuropteri (figs. 52, 53). The male genitalia 


596 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ala 


Ww 
a, VW MWe 
Way) 


— 


59 
6l 


Ficures 47-61.—Neohaematopinus capitaneus, new species, N. kinabalensis, new species» 
and N. sciuropteri: 47, Genital plate, N. capitaneus, allotype 9 ; 48, Same, N. kinabalensis, 
allotype ? ; 49, Same, WN. sciuropter1, 9 , Florida from Glaucomys; 50, Head, N. capitaneus, 
holotype o@; 51, Same, NM. kinabalensis, holotype co’; 52, Same, N. sciuroptert, &; 53, 
Antenna, WN. sciuropteri, 2 ; 54, Genitalia, N. sciuropterit, &; 55, Same N. kinabalensis, 
holotype co; 56, Same, N. capitaneus, holotype o&; 57, Paratergal plate, N. capitaneus, 
paratype 9; 58, Same, N. scturopteri, 9 ; 59, Thoracic sternal plate, N. sciuropteri, 2 ; 
60, Same, N. capitaneus, allotype 9 ; 61, Same, N. kinabalensis, allotype 9. 


ANOPLURA OF THAILAND—JOHNSON 591 


of the two species differ in that in sciuropteri, the penis opens much 
further distad, the parameres have straight lateral margins, and the 
apical lobes of the abdomen are much better developed than in 
kinabalensis (compare figs. 54 and 55). The female genitalia differ in 
that in sciuropteri the lateral setigerous lobes of the eighth segment 
(“gonopods’’) are almost vertical rather than slanted and end anterior 
to the lip of the vulva, which is straight and not heavily spiculated 
(compare figs. 48 and 49). 

Description: Male: Head almost as broad as long, broadly 
rounded anteriorly, antennae set near apex; marked postantennal and 
occipital angles; postantennal margins straight and parallel. Basal 
antennal segment as broad as long, with short stout seta on dorsal 
surface; segment 3 modified and bearing two short stout setae. 
Thorax longer and broader than head; sternal plate roughly rectangu- 
lar and with posterior angles drawn into short acute points. Legs as 
in other members of the genus. Abdomen with two sternal plates 


Ficures 62-63.—Neohaematopinus kinabalensis, new species: 62, Allotype ¢ ; 63, Holotype 
od". 


592 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


and one tergal plate per segment, these plates well developed. Typical 
sternal plate with 7 to 9 setae along posterior margin; tergal plates 
with 15 such setae; ventrally 1 lateral seta off the plate on each 
segment; dorsally with 2 or 3 lateral setae. Paratergal plates with 
apical setae shorter than the plates; posterior margin with postero- 
lateral angles acutely pointed and with each setal base usually flanked 
by shorter points. Tergal plate of segment 8 joined with paratergal 
plates VIII. Genitalia with penis opening at about proximal third 
of parameres; parameres broadly convex laterally, their apices folded 
over pseudopenis dorsally; pseudopenis serrate distally, extending 
well beyond apices of parameres. Terminal lobes of abdomen not 
enlarged or elongate. 

Female: Head as in male except third antennal segment normal. 
Thorax and legs as in male. Abdomen with two well-developed 
sternal and tergal plates on each typical segment; plates with row 
of 7 to 10 setae. Ventrally each segment with one lateral seta off 
plates on each side; dorsally with one to two such setae off each plate 
on each side. Paratergal plates as in male except plate VIII not 
joined to tergal plate of segment 8. Genitalia: Lateral setigerous 
lobes of segment 8 (‘“‘gonopods’’) lying almost horizontally, widely 
separated, margin of vulva between them only somewhat convex; 
genital plate not heavily spiculated. 

Lengths: Male 1.67 mm.; female 2.25 mm. 


Neohaematopinus elbeli, new species 
Figures 64-68 


Type pata: Holotype male, allotype female from Dremomys 
rufigenis, LOEI: Dan Sai, Kok Sathan, Phak Khi Nak Mt., March 
19, 1955, RE-4977, R. E. Elbel collector. One male, one female 
paratypes with same data as holotype. One male paratype, Dan 
Sai, Na Haeo, Ban Bo, May 15, 1955, RE-5236. Three male para- 
types, Ban Bo, May 19, 1955, RE-5279. All specimens were taken 
from Dremomys rufigenis. 

Holotype deposited in the collections of the U.S. National Museum, 
type catalog No. 64387. 

This species is named for the collector, Robert E. Elbel, whose 
magnificent collections of Thai ectoparasites have made known a 
fauna completely neglected in the past. 

Diacnosts: N. elbeli is a member of the group of Neohaematopinus 
species that have the posterior angles of the thoracic sternal plate 
produced into points, and paratergal plates [V-VI with no more than 
two apical setae. It is closest to N. inornatus (Kellogg and Ferris), 
which also has the abdominal plates much reduced, but is distinct in 
that the head is almost as broad as long in elbeli; the thoracic sternal 


ANOPLURA OF THAILAND—JOHNSON 593 


Figures 64-68.—Neohaematopinus elbeli, new species: 64, Allotype 9; 65, Holotype @; 
66 Thoracic sternal plate, allotype ¢ ; 67, Genitalia, paratype co’; 68, Paratergal plates 
II-VI], allotype 9. 


plate is much broader than in inornatus, and the paratergal plates are 
more strongly sclerotized. 

Description: Male: Head broadly rounded anteriorly, almost as 
broad behind antennae as entire head is long; postantennal angles 
rounded, postantennal margins convex. First antennal segment not 
modified although there is one large seta dorsally, set in from the 
margin; third antennal segment modified and with two short, stout 
apical setae. Thorax noticeably broader than long. Sternal plate 
broader anteriorly than posteriorly, slightly broader than long, with 
its posterior angles produced into points. Legs as in other members of 
the genus. Abdomen entirely lacking sternal plates except for those 
associated with the genital segments. Tergal plates present on seg- 
ments 2—7, plates reduced except for the second plate, which is emar- 
ginate medially and bears a lateral group of setae on each side. Para- 
tergal plates large and well sclerotized, each with two apical setae; 
plate VIII joined dorsally to tergal plate of segment 8; apical angles 
of plates II-VI produced into short points. Genitalia as in figure 67; 
basal plate narrow, expanded posteriorly, twice as long as the para- 
meres; parameres with outer margins broadly convex (extended in 
holotype); pseudopenis narrow. 


594. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Female: Head as in male except third antennal segment not modi- 
fied. Thorax and legs asin male. Abdomen lacking plates except for 
one tergal plate on segments 2, 8, and 9, and the usual sternal plates 
on the genital segments. Paratergal plates as in male, except plate 
VIII not joined to tergal plate of segment 8. Typical abdominal 
segments with two rows of setae. Genitalia not diagnostic. 

Lengths: Male 1.2 mm.; female 1.5 mm. 


Nechaematopinus petauristae Ferris 


Neohaematopinus petauristae Ferris, Contributions toward a monograph of the 
sucking lice, pt. 4, p. 258, figs 166, 167A, 167C—E, 1923. 

Petauristophthirus petauristae, Eichler, Boll. Soc. Ent. Italiana, vol. 79, p. 12, 1949. 

Neohaematopinus petauristae, Ferris, The sucking lice, p. 195, 1951. 

Ferris described petauristae from male and female specimens taken 
from Petaurisia inornata, Kashmir. The holotype is a female. The 
record below is the first since the original! description. 

New rReEcorD: Two males, two females from Petaurista taylort, 
PRACHUAP KHIRI KHAN: Ban Khlua Klang, December 14, 1952, 
RE-2032. 


Polyplax asiatica Ferris 


Polyplax asiatica Ferris, Contributions toward a monograph of the sucking lice, 
pt. 4, p. 233, fig. 152D, 1923; The sucking lice, p. 206, 1951—Hopkins, 
Proce. Zool. Soe. London, vol. 119, p. 483, 1949—Johnson, Bull. Brooklyn 
Ent. Soc., vol. 53, p. 77, figs. 1-5, 7, 1958. 

The holotype female and allotype male of Polyplar asiatica were 
collected from Crocidura caerula, Rangoon, Burma. Ferris also 
reported several females from Nesokia indica (as N. hardwicki) from 
Baluchistan. 

New recorp: One female from Bandicota bengalensis, KAN- 
CHANABURt: Kanchanaburi City, June 10, 1952, F—-875. 

P. asiatica is probably very rare on Bandicota bengalensis, the 
normal} louse parasite of this rat being Hoplopleura malabarica Werneck. 
Ferris and Hopkins have reported asiatica from Nesokia indica, 
Rattus concolor, and Bandicoia malabarica. Nesokia, which is closely 
related to Bandicota, is probably the primary host, although aszatica 
appears to be well adapted to life on other related genera of rodents, 

ts occurrence on Crocidura, a shrew, was almost certainly fortuitous, 


Polyplax cannomydis, new species 
Ficures 69-72, 74 


Typ pata: Holotype male, allotype female from Cannomys badius 
minor (bamboo rat), RAT BURI: Banpong, Boekphrai, June 18, 1955, 
Y-236, R. E. Elbel and Banpong Plague Lab. personnel, collectors. 
Two male, four female paratypes with same data as holotype. One 
female paratype from C. badius minor, KANCHANABURI: Tamaka, 


ANOPLURA OF THAILAND—JOHNSON 595 


Tharua, February 1, 1952, RE-485. Three male, seven female para- 
types from C. badius minor, KANCHANABURI: Tha Khanun, Hin 
Laem, October 27 and 29, 1952, RE-1308 and RE-1329. 

Holotype deposited in the collections of the U.S. National Museum, 
type-catalog No. 64388. 

Diacnosts: P. cannomydis is immediately separable from all other 
described members of the genus Polyplax by the lack of any vestige of 
paratergal plates on the eighth abdominai segment, by the absence of 
plates on the abdomen, except on the genital segments and by the 
very large and flattened tarsal claw on the third leg. 

Description: Male: Head elongate, preantennal region strongly 
sclerotized and produced so that antennae are inserted well posteriad 
to the head apex; gular region apparently not raised, although a 
lateral longitudinal fold may represent the limits of a greatly broad- 
ened gular area; postantennal head margins slightly convex; occipital 
region constricted; antennae normal, basal segment not enlarged and 
third segment unmodified. Thorax broader than head and not as 
long; sternal plate obsolescent, represented by a small area of deeper 
pigmentation. First pair of legs small, with slender claw; second pair 
similar and only somewhat larger; third pair much larger than others, 
with large, flattened claw. Abdomen entirely devoid of plates except 
the ventral genital plate of segments 7 and 8. Ventrally with one 
row of 2 setae on segment 2; one irregular row of 9 to 10 setae on 
segment 3 (these setae may represent coalescence of two original 
rows); segments 4-6 with one row of 5 to 6 setae plus 1 lateral seta 
on each side near paratergal plates. Dorsally with 1 row of setae 
per segment; segments 3-5 with one median row of 10 to 12 setae, 
plus 1 or 2 lateral setae on each side; segments 6 and 7 with row of 
11 to 14 setae; segment 8 with row of 4 setae. Paratergal plates 
present on segments 2-7; plate II poorly sclerotized, with no apparent 
division into two parts; plates TII-VIE sclerotized only apically, with 
posterior angles rounded; each bearing two very long apical setae. 
Segment 8 lacking paratergal plates but with spiracles and 
usual two long lateral setae on each side below spiracle. Genitalia 
with parameres enclosing proximal half of stout wedge-shaped 
pseudopenis. 

Female: Head, thorax, and legs as in male. Abdomen lacking 
plates except the genital plate of the eighth segment. Dorsally, 
segment 2 with one row of setae; segment 3 with two rows; segment 4 
with one row, plus 2 lateral setae on each side; segments 5-7 with 
two rows, anterior row of 6 to 9 short setae, posterior row of 10 to 14 
much longer setae, and with 1 or 2 setae laterally on each side. Ven- 
trally segments 2-4 with one row of setae; 5-7 with two rows, anterior 
row of 2 tc 5 short setae, posterior rows of 4 to 8 longer setae; 


596 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Ficures 69-70.—Polyplax cannomydis, new species: 69, Allotype 9; 70, Holotype o. 


segments 3-7 with 1 or 2 lateral setae near paratergal plate on each 
side. Paratergal plates asin male. Genitalia (fig. 72) not typical of 
Polyplaz species. Genital plate (of segment 8) rectangular, lip of 
vulva smooth, not beset with fimbriate processes; lateral setigerous 
lobes of segment 8 (‘gonopods”) merely more heavily pigmented 
areas occurring laterally on genital plate, not protruding as lobes, 
with 2 or 3 setae on the pigmented area. Laterally, below vulva, 
lobe on each side bearing many setae; below these, the usual lateral 
setigerous lobes of segment 9 (‘‘gonopods’”’). Spermatheca heavily 
sclerotized, appearing as a ring-shaped sclerite. 

Lengths: Male 1.5 mm.; female 2.0 mm. 

The affinities of cannomydis, new species, are difficult to outline. 
There is a very real question as to whether it belongs in Polyplaz or 
whether it should be segregated as the type of a new genus. The 
female external genitalia of cannomydis are not of the usual Polyplaz 
type. In typical Polyplax the genital plate has a triangular median 
sclerotization flanked by definite lateral setigerous lobes (‘““gonopods”’), 
and the tip of the vulva is always fimbriate to varying degree (fig. 
73). P. cannomydis has the setigerous lobes incorporated into the 
genital plate and the lip of the vulva smooth. The well-marked 
setigerous lobes occurring laterally between the vulva and the seti- 
gerous lobes of the ninth segment are unusually well developed in 
cannomydis, although typical Polyplax species do have groups of 
setae in these positions. Not only are the female genitalia different 


ANOPLURA OF THAILAND—JOHNSON © 597 


from other Polyplaz, but cannomydis lacks paratergal plates on the 
terminal abdominal segments. 

Two genera, both closely related to Polyplax, share certain charac- 
teristics with cannomydis: 

Proenderleinellus Ewing (only included species: calvus (Waterston)) 
shows relationships to cannomydis in the form of the male genitalia, in 
lacking abdominal plates, and in having the third pair of legs greatly 
enlarged, although in calvus the claw is not flattened. The female 
genitalia are somewhat similar in the two species, but calvus has the 
lip of the vulva strongly fimbriate (fig. 75). Further, males and 
females of calvus have two rows of setae on each abdominal segment 
ventrally and dorsally while cannomydis follows the Polyplaxz pattern 
by having only one row ventrally on segments 4-7 in the male. 

Like species of Hulinognathus Cummings, P. cannomydis lacks the 
terminal paratergal plates and the female genitalia resemble £. 
denticulatus Cummings. However, species of Hulinognathus lack the 
pronotum (which, if present, is always reduced to a very narrow 
median longitudinal strip in the Anoplura); the pronotum is present 
in species of Polyplax, and the legs of cannomydis also are more nearly 
like Polyplaxz than any species of Hulinognathus. 

Although the generic concept of Polyplax must be broadened in 
order to include cannomydis, I feel that such broadening is the proper 
course at this time. Ferris (Contributions toward a Monograph of 
the Sucking Lice, pt. 4, 1923) grouped the various species of Polyplaz, 
although without discussing the groups, in what appears to be the 
most logical pattern. It would be impossible to justify the establish- 
ment of a new genus to receive cannomydis without naming four or 
five other new genera to contain the various groups of Polyplaz. 
Polyplax is certainly a complex assemblage of species but there is 
serious doubt as to whether any purposes would be served by splitting 
it into several genera. 


Polyplax spinulosa (Burmeister) 


Figure 73 
Pediculus spinulosus Burmeister, Genera quaedam insectorum . . . , Rhynchota, 
vol. 1, No. 8, 1839. 
Polyplax spinulosa, Ferris, Contributions toward a monograph of the sucking 
lice, pt. 4, p. 187, figs. 119, 120A, D, F, H, 1923. 

P. spinulosa is a parasite of Rattus species throughout the world, 
particularly PR. rattus and R. norvegicus. P. spinulosa proved to be 
a rare species on Thai Rattus, being taken in 12 collections from 
Rattus rattus ssp., once from Rattus norvegicus, and once from a 
“badger,” which probably acquired the louse from one of its Rattus 
victims. 


598 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


New recorps: Two females from R. rattus ssp., RAT BURL: Banpong, 
Pakraet, February 23, 1952, RE-464. One male, one female, same 
data, but July 2, 1952, F. 949. Two males, one female, same data, 
but RE-465. One female from AR. rattus ssp., RAT BURI: Banpong, 
2 km. west of city, June 25, 1952, F. 935. One male from R. nor- 
vegicus, Bangkok, June 1, 1952, Y. 206. One male from A#. rattus 
ssp., KANCHANABURI: Tha Khanun, Hin Laem, October 29, 1952, 
RE-1333. One female from “badger,’’? NAKHON RATCHASIMA (= KHO- 
RAT): Khorat (city), February 8, 1953, RE-1722. One female from 
R. rattus ssp., locality as above, February 10, 1953, RE-1723. Two 
males, 4 females, same data, but February 13, 1953, RE-1725. One 
female, same data, but January 30, 1953, RE-1781. 


Figures 71-75.—Polyplax cannomydis, new species, P. spinulosa, and Proenderleinellus 
calous: 71, Paratergal plates, P. cannomydis, allotype ? ; 72, Genitalia, P. cannomydis, 
allotype 9; 73, Same, P. spinulosa, 9, Tanganyika from R. ratius; 74, Genitalia, P. 
cannomydis, holotype &; 75, Genitalia, P. calvus, 9 , Tanganyika, from Cricetomys. 


U.S. GOVERNMENT PRINTING OFFICE: 1959 


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INDEX 


(New genera, species, etc., are printed in zalics. 


Page numbers of principal 


entries also in ztalics) 


abbreviator, Agrothereutes, 446 

abdominale, Apion, 250, 256, 332, 3383, 
334 (fig.) 

abdominalis, Rhembobius, 436, 438 (fig.) 

abdominator, Cubocephalus, 439 

abeli, Plecotus, 209 

abruptorius, Exenterus, 426 

Acaenitini, 397, 398, 399, 400, 414, 416 
(fig.), 506, 507 

acanonicum, Apion, 250, 254, 281, 282, 
283 (fig.) 

Acanthopinus antennatus, 585 

Acer, 211 

Acrididae, 69 

Acridium, 55 

spretis, 4, 55 
Acrodactyla, 407 
degener, 407 

acrophilum, Apion, 249 

Aculeata, 439 

acupunctatum, Apion, 250, 347, 348, 
349 (fig.) 

acutus, Horogenes, 491 

tes teas Apion, 250, 258, 282, 284, 287 


(fig.) 
Adelognathinae, 398, 417, 418 (fig.), 506 
Adelognathus, 417 
britannicus, 417 
pallipes, 417, 418 (fig.) 
sp., 417, 419 
adspersus, Exenterus, 426 
sane: Apion, 250, 257, 259, 264 
fig. 
aequalis, Pimpla, 408 (fig.), 410 
aerosus, Akodon, 519 
aestimabile, Apion, 250, 260 
aestivalis, Horogenes, 491 
affinis, Melanoplus, 3, 14, 15, 42, 45 
Melanoplus devastator, 42, 43 
agilis, Syrphoctonus, 476 
Agrothereutes, 443 
abbreviator, 446 
batavus, 426, 446 
grapholithae, 443, 446 
sp., 443, 444 (fig.) 
Agrypon, 503 
sp., 499 (fig.), 503 
Akodon aerosus, 519 
alaskanus, Melanoplus, 3, 45, 46, 48 
alba, Betula, 280 
aia eae ) Apion, 250, 252, 260, 264 
fig. 


546764—60——2 


albifrons, Exochus, 480 
albitarsus, Cryptus albitarsus, 440 (fig.), 
443 


albopictus, Lamachus, 468 
alfaroi, Oryzomys, 525 
alkae, Campoplex, 492 
alleganiensis, Corynorhinus, 137, 209 
Plecotus, 136, 137, 138, 140, 141, 
210, 213, 215 
Alnus, 211 
Alomyini, 448 
Alophosternum, 401, 404 
foliicola, 402 (fig.), 404 
alternans, Itoplectis, 410 
alticola, Apion, 250, 261 
Amblyteles, 456 
armatorius, 456 
inconstans, 454 (fig.), 4 
subrufus, 456 
ambulatorius, Ichneumon, 
456 
americanus, Mnicospilus, 501 
Amorpha canescens, 344 
amplexicaulis, Wyethia, 52 
analis, Labrorychus, 498 (fig.), 603 
aneyloneura, Ophion, 501 
angelicus, Melanoplus, 3, 14 
anguya, Hesperomys, 552 
Anisobas, 449 
sp., 450 (fig.), 451 
annulatum, Eiphosoma, 495 
Anomalinae, 398, 399, 459, 497 (fig.), 
498 (fig.), 499 (fig.), 500 (fig.), 
501, 502 (key), 505, 506 
Anomalini, 497 (fig.), 502 
Anomalon, 398, 501, 502, 503, 505, 506 
ejuncidum, 503 
sp., 497 (fig.), 502 
Anoplura (sucking lice) of Thailand, 
with remarks on related species, 
569 
antennatus, Acanthopinus, 585 
anthomyidiperda, Stilpnus, 435 (fig.) 
antiquus, Myotis, 206 
antrozoi, Basilia, 174 
Antrozoinae, 104 
Antrozous, 104, 105, 106 
pallidus, 169, 170, 172 
anxius, Metopius, 481 
Aoplus, 456 
pictus, 456 
velox, 456 


56 


455 (fig.), 


599 


600 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 110 


Apanteles, 394 
Apechthis, 410 

compunctor, 408 (fig.), 410 
Aphanistes, 503 

armatus, 504 

hyalinus, 498 (fig.), 503 
Apion subgenus Trichapion Wagner in 

the New World, A revision of, 247 

Apion, 247, 281 

abdominale, 250, 256, 332, 333, 334 


(fig.) 
acanonicum, 250, 254, 281, 282, 283 


(fig.) 

acrophilum, 249 

acupunctatum, 250, 347, 348, 349 
(fig.) 

adaetum, 250, 258, 282, 284, 287 
(fig.) 

aequabile, 250, 257, 259, 264 (fig.) 

aestimabile, 250, 260 

albidulum, 250, 252, 260, 264 (fig.) 

alticola, 250, 261 

asellum, 250, 262 

aurichaleeum, 248, 249, 250, 254, 
272, 273, 295, 296, 299 (fig.) 

auriferum, 250, 257, 326, 327 (fig.) 

auronitidum, 250, 263 

bettyae, 250, 253, 263, 264 (fig.) 

bicolor, 250, 257, 347, 349 (fig.), 
350 

brachycephalum, 250, 2538, 309, 
310, 311 (fig.) 

brunnicornis, 250, 253, 322, 323 


(fig.) 

caenum, 250, 253, 311 (fig.), 312 

calearatipes, 250, 254, 309, 311 
(fig.), 312 

capitatum, 368, 369 

centrale, 250, 256, 332, 834 (fig.), 
835 

chaiybaeum, 250, 265 

chuparosae, 250, 258, 291, 293 (fig.) 

coloradense, 251, 258, 378, 379 (fig.) 

commodum, 251, 252, 378, 379 
(fig.), 380 

confertum, 251, 252, 268, 377, 378, 
379 (fig.), 381 

cordatum, 250, 253, 369, 371 (fig.) 

coryt, 250, 252, 281, 282, 283 (fig.), 
285 

coxale, 249 

crassinasum, 289, 290 

davidis, 250, 347, 349 (fig.), 351 

dolosum, 250, 258, 369, 370, 371 


(fig.) 

eccentricum, 250, 323, 324 

enoplus, 250, 254, 309, 311 (fig.), 
314 

eppelsheimi, 279 

evustum, 249, 250, 255, 331, 332, 334 
(fig.), 337, 347 

fraternum, 306, 307 

furtivum, 251, 253, 268, 377, 378, 
382 


Apion, fusconitidum, 250, 332, 334 (fig.), 
33 


8 
elyphicum, 250, 258, 291, 292, 293 


(fig.) 

godmani, 250, 254, 262, 295, 296, 
297, 299 (fig.) 

gracilirostre, 250, 257, 326, 347, 349, 
(fig.), 352 

griseum, 250, 255, 295, 296, 299 
(fig.), 300, 306, 307, 308 

grossulum, 250, 255, 269 

guatemalense, 250, 270 

guatemalenum, 350, 351 

ae 250,257; 832; SSSi3389 

g. 

hadromerum, 250, 258, 326, 327 
(fig.), 328 

heterogeneum, 250, 255, 369, 372, 
373 (fig.) 

hystriculum, 330 

imitator, 250, 257, 261, 262, 281, 
282, 286, 287 (fig.) 

impexum, 249 

importunum, 250, 257, 348, 349 
fig.), 343 

innocens, 250, 253, 304, 309, 311 

_ (ig.), 314 

ee 250, 254, 309, 315; 316 

g. 

laminatum, 247, 250, 264 (fig.), 271 

lanuginosum, 279, 280 

latitator, 250, 253, 309, 316 (fig.), 
$17 

lativentre, 250, 258, 369, 374 

managuense, 250, 256, 348, 349 
(fig.), 354 

mandoni, 350 

mediocre, 250, 253, 261, 366, 367 
(fig.) 

meorrhynechum, 250, 255, 274, 275 

metallicum, 249 

mexicanum, 251, 258, 260, 378, 
379 (fig.), 383 

rie 250, 258, 347, 348, 355, 356 
fig.) 

mirandum, 250, 253, 309, 316 (fig.), 
318 

mirificum, 250, 253, 316 (fig.), 319 

modestum, 250, 256, 274, 276 (fig.) 

nanulum, 250, 256, 348, 356 (fig.), 
357 

nebraskense, 250, 254, 295, 301, 
302 (fig.) 

nigrum, 250, 255, 346, 369, 371 
(fig.), 376 

nitidirostre, 250, 347, 356 (fig.), 3858 

nitidum, 250, 256, 339 (fig.), 344 

novellum, 250, 254, 295, 303 

nunenmacheri, 289, 291 

oblitum, 250, 253, 366, 367 (fig.) 
368 

obseurum, 275 

opacicolie, 249 

oriotes, 250, 255, 296, 299 (fig.), 304 

oscillator, 250, 256, 265, 269, 348, 
356 (fig.), 359 


INDEX 601 


Apion, parcum, 250, 255, 295, 296, 302] Aptesis, 436 


(fig.), 305 


patruele, 249, 250, 


251, 256, 331, 


332, 333, 339 (fig.), 341 
peninsulare, 247, 250, 271 


perforicolle, 249, 250, 257, 331, 332, 


339 (fig.), 343 


perpilosum, 250, 257, 326, 327 (fig.), 
329 


persulcatum, 250, 272 
pervicax, 250, 257, 348, 360 


plectrocolum, 250, 


(fig.), 288 


porcatum, 250, 256, 331, 332, 


(fig.), 345 


proclive, 250, 258, 274, 281, 282, 287 


(fig.), 289 
propinquicorne, 
(fie) 277) 


punctulirostre, 250, 254, 322, 


(fig.), 325 
reclusum, 249 


reconditum, 249, 250, 


(fig.), 361 


rostrum, 249, 251, 257, 307, 308, 


250, 255, 274, 


257, 282, 287 
339 


378, 379 (fig.), 384 


rufinulum, 346 


rufipenne, 249, 250, 255, 331, 332, 


339 (fig.), 346 


sagax, 350 
sancti-felicis, 
(fig.), 320 


sayi, 250, 254, 275, 295, 296, 


250, 


253, 309, 316 


(fig.), 300, 301, 306 
scrobicolle, 384, 385 
semicastaneum, 346 


simile, 249, 250, 


258, 273, 274, 275, 


276 (fig.), 278, 282 


smithi, 249 


spectator, 325, 326 


spiculiferum, 250, 272 
spinitarse, 250, 254, 309, 321 


submetallicum, 250, 258, 326, 327 


(fig.), 330 
subnudum, 272 
subrufum, 249, 


250, 256, 263, 348, 


356 (fig.), 363 


subsequens, 250, 256, 348, 356 (fig.), 


364 


subtinctum, 249, 250, 256, 348, 365 
superciliosum, 278 


tabogense, 247, 250, 273 


tenebricosum, 275 
tenuirostrum, 249 


triste, 279 
troglodytes, 249 


turbulentum, 3861, 362 


vicinum, 279 


vinosum, 249, 250, 258, 291, 294 
walshii, 279, 280 


Apios tuberosa, 343 


appalachianus, Pristomerus  pacificus, 


36 (key), 437 (fig.), 


4Yv 


495 
Aptesini, 426, 427, 4 
438 (fig.) 


276 
323 


256, 348, 356 


299 


basizonia, 439 
indistincta, 486, 437 (fig.) 
pteronorum, 436 
Arbutus (madrone), 211 
Archaeonycteridae, 205 
Archaeonycteris, 205 
arcticus, Caloptenus, 3, 14, 70 
Melanoplus, 3, 15 
arcuatus, Enicospilus, 497 (fig.) 
areator, Hemiteles, 433 (fig.), 485, 436 
argeae, Mastrus, 429, 435 
argentinensis, Tersiloxhus, 496 (fig.) 
Argidae, 426 
ariel, Plecotus auritus, 132 
armatorius, Amblyteles, 456 
armatus, Aphanistes, 504 
armillata, Horogenes, 492 
Arotes, 415, 417 
formosus, 415, 416 (fig.) 
asellum, Apion, 250, 262 
asiatica, Polyplax, 594 
Asphragis, 463 
mirabilis, 462 (fig.), 463 
Astiphromma, 459 
strenuum, 458 
atalantae, Theronia, 413, 414 (fig.) 
ater, Melanoplus, 3, 41, 43 
Phaeogenes, 448 
atlanis, Caloptenus, 3, 14 
Melanoplus, 3, 10, 11, 12, 14, 15, 
18, 21, 34, 55 
Atractodes, 428, 436 
sp., 435, 505 (fig.) 
atricolor, Thyreodon, 501 
Atrometus, 503 
clavipes, 499 (fig.), 503 
aurichaleeum, Apion, 248, 249, 250, 
254, 272, 273, 295, 296, 299 (fig.) 
auriculatus, Scolobates, 466, 467 (fig.) 
auriferum, Apion, 250, 257, 326, 327 
fi . 


g. 
auritus, Plecotus, 98, 105, 107 (fig.), 
LO 1120114 (fies) 115 (fig.), 
LG, Aly ig je 1s, L20mGigs), 
121 (fig.), 126, 130, 132, 138, 
134, 135, 139, 140, 178, 209, 
212, 213, 215, 218, 224 
Plecotus auritus, 132 
Vespertilio, 127, 132 
auriventer, Oryzomys, 556, 557 
auronitidum, Apion, 250, 263 
australis, Corynorhinus townsendii, 185 
Plecotus townsendii, 142, 143, 144 
(fig.), 147, 152, 153, 185, 187, 
188, 191, 192, 221, 228 
austrinus, Pristomerus, 495 
austroriparius, Myotis, 173 
auyantepui, Oecomys, 553, 554, 555, 556 


Badger, 597, 598 

bahiensis, Oryzomys concolor, de / 
Oryzomys concolor, 517, 519 

Balearcia bergi, 492 

balticus, Juncus, 80 


602 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


Banchini, 459, 460, 463 (key), 464 (figs.) | bilituratus, Caloptenus, 3, 14 


Banchus, 463 
femoralis, 463, 464 (fig.) 
flavescens, 463 
Bandicota, 495 
bengalensis, 575, 577, 594 
indica, 575 
malabarica, 594 
sp., 575 
Baptisia leucantha, 386 
tinctoria, 386 
Barbasteila, 107, 118, 120, 205, 207 
(fig.), 208, 211, 212, 213, 215 
barbastellus, 107 (fig.) 
barbastellus, Barbastella, 107 (fig.) 
Barylypa, 503 
insidiator, 498 (fig.), 503 
Basilia antrozoi, 174 
corynorhini, 174 
basizonia, Aptesis, 439 
batatae, Eiphosoma, 494 (fig.), 495 
batavus, Agrothereutes, 426, 446 
Bathyplectes, 481, 489, 491 
corvina, 491 
curculionis, 484 (fig.), 491 
exiguus, 491, 492 
Bathythrix, 429 
sp, 4380 (fig.), 485 
Bats, revision of American, of the genera 
Euderma and Plecotus, 95 
bechsteinii, Myotis, 206 
bedardi, Erromenus, 423 
behrensii, Itoplectis, 409 (fig.), 410 
benevola, Nepiera, 485 (fig.), 491 
benevolens, Oecomys, 540 
Oryzomys, 540, 541, 542 
Rhipodomys, 515, 518, 540, 541 
Rhypidomys, 535 
bengalensis, Bandicota, 575, 577, 594 
Benjaminia, 489, 506 
euphydryadis, 491 
fuscipennis, 488 (fig.), 491 
berdmorei, Menetes, 574, 578, 583, 585, 
587 
bergi, Balearcia, 492 
bettyae, Apion, 250, 253, 263, 264 (fig.) 
Betula alba, 280 
verrucosa, 280 
bicingulata, Nemeritis, 492 
bicolor, Apion, 250, 257, 347, 349 (fig.), 
350 


Hesperomys, 518, 534 
Myoxomys, 534 
Oecomys, 518, 534, 535, 538, 539 
Oryzomys, 5138, 515, 516, 517, 523 
(fig.), 524 (fig.), 526 (fig.), 527, 
528, 529, 530, 531, 532, 533, 534, 
536-537, 542, 543, 544, 554, 555 
Oryzomys bicolor, 516, 518, 529, 
532, 534, 536-537, 540 
bifenestratus, Exetastes, 463, 464 
bifoveolatus, Ophion, 496 
bilineatus, Ophion, 501 


Melanoplus, 2, 3, 10, 12, 13 (fig.), 
15, 17 (fig.), 18, 19 (fig.), 24, 38, 
45, 46, 47, 50, 52, 55, 65, 66, 72 
Melanoplus bilituratus, 3, 4, 7, 13 
(fig.), 14, 17 (fig.), 18, 19 (fig.), 
22, 25, 27 (fig.), 28; 29, 31 (fig.), 
32, 33 (fig.), 34, 37 (fig.), 38, 40, 
43, 44, 45, 52, 64, 70 
Melanoplus mexicanus, 18 
Biolysia, 481, 489, 491 
tristis, 484 (fig.), 491 
Blaptocampus, 503 
nigricornis, 499 (fig.), 503, 504 
Boéthes sp., 425 (fig.) 
Boéthini, 419, 425 (fig.) 
borealis, Caloptenus, 3, 67, 68 
Campoplex, 492 
Lasiurus, 118, 119 
Melanoplus, 2, 3, 67, 68, 69, 74 
Melanoplus borealis, 2, 3, 4, 8, 19 
(fig.), 27 (fig.), 31 (fig.), 33 (fig.), 
37 (fig.), 49 (fig.), 59 (fig.), 67, 
18; 4, (8, 79, 82 
Neoxorides, 411, 412 (fig.) 
Bothryopteron, subg., 248 
brachycephalum, Apion, 250, 253, 309, 
310, 311 (fig.) 
Brachycertus, 428 
Brachycyrtini, 425 (fig.), 427, 428 
Brachyeyrtus pretiosus, 425 (fig.), 428 
brachylabris, Xoridescopus, 415 
Braconinae, 507 
Bradynotes chilcotinae, 48 
brasiliensis, Holochilus, 552 
brasiliensis, Tadarida, 172 
brevicinetor, Melanichneumon, 456 
brevicornis, Epiurus, 404 
Secambus, 404 
britannicus, Adelognathus, 417 
Brooks, A. R., see Gurney and Brooks, 1 
bruesii, Gelis, 429, 435 
brumatae, Cubocephalus, 439 
bruneri, Melanoplus, 3, 7, 13 (fig.), 15, 
17 (fig.), 19 (fig.), 27 (fig.), 33 
(fig.), 37 (fig.), 38, 45, 49 (fig.), 
50, 51, 59 (fig.), 69, 82, 83 
brunnea, Lissonota, 462 (fig.), 463 
brunneifrons, Iseropus stercorator, 403 
(fig.), 405 
brunnicornis, Apion, 250, 253, 322, 323 


g. 

Herpestomus, 448 
buccinatus, Oryzomys, 552 
Buckellacris chilcotinae, 48 
Bumelia sp., 366 


caenum, Apion, 250, 253, 311 (fig.), 312 
caerula, Crocidura, 594 
caeruleipes, Caloptenus, 3 
Caloptenus spretus, 52 
Melanoplus, 3, 15, 55 
Melanoplus spretus, 3 
caicarae, Oecomys, 547, 548-549, 550, 
553 


Oecomys guianae, 547 


INDEX 


603 


calearatipes, Apion, 250, 254, 309, 311 Tee re 594, 596 (fig.), 


.), 312 


calcitrator, Collyria, 471, 473 (fig.), 474] Cannomys badius minor, 594, 595 


californicus, Iseropus, 405 
Macrotus, 170 
Myotis, 169, 170, 172 
Calliephialtes, 401, 404 
dimorphus, 401, 405 
nubilipennis, 404, 405 
variatipes, 402 (fig.), 404, 405 
callosciurt, Neohaematopinus, 681, 582 
(fig.), 584 (fig.), 585, 586 
Callosciurus caniceps, 574, 580, 581 
erythraeus, 581 
erythraeus thaiwanensis, 573 
ferrugineus, 580 
finlaysoni, 573, 581, 585 
finlaysoni tacherdi, 580 
prevosti, 574 
sp., 573, 578, 585, 586 
Caloptenus arcticus, 3, 14, 70 
atlanis, 3, 14 
bilituratus, 3, 14 
borealis, 3, 67, 68 
caeruleipes, 3 
extremus, 3, 67 
mexicanus, 9 
parvus, 4, 68 
scriptus, 4, 14 
selectus, 4, 8, 9 
spretus, 4, 52 
spretus caeruleipes, 52 
calvus, Proenderleinellus, 597, 598 (fig.) 
Campoletis, 490 
oxylus, 485 (fig.), 491 
Campoplegini, 398, 466, 468, 481, 482 
(fig.), 483 (fig.), 484 (fig.), 485 
(fig.), 486 (fig.), 487 (fig.), 488 
(fig.), 489 (key), 491, 492, 495, 
506 


Campoplex, 490, 491 
alkae, 492 
borealis, 492 
crassifemur, 490, 492 
cultrator, 492 
depressus, 490 
ensator, 492 
frustranae, 490 
hyalinus, 490 
inaequalipes, 490, 491 
mutabilis, 492 
phthorimaeae, 490 
polychrosidis, 490 
pyraustae, 490, 491 
rufifemur, 490, 492 
tosensis, 490, 491 
tricoloripes, 490 
validus, 482 (fig.), 490, 492 
sp., 490, 491 
canadensis, Exenterus, 425 (fig.), 426 
Gambrus, 441 (fig.), 443 
candidissima, Cornus, 363 
canescens, Amorpha, 344 
Idechthis, 491, 492 
caniceps, Callosciurus, 574, 580, 581 


capitaneus, Neohaematopinus, 587, 588 
(fig.), 589, 590 (fig.) 
capitatum, Apion, 368, 369 
Carinodes, 451 
havanensis, 455 (fig.), 456 
carolinensis, Sciurus, 584, 585, 586, 587 
carpocapsae, Cremastus, 495 
Casinaria, 490 
eupitheciae, 483 (fig.), 491 
indubia, 491 
infesta, 491 
tenuiventris, 491 
sp., 491 
catenator, Lampronota, 463 
caterpillars, 420, 459, 460, 496 
catherinae, Oecomys, 518, 519, 543 
Oryzomys, 518 
Cecidomya sp., 280 
Cecilionycteris, 205 
centrale, Apion, 250, 256, 332, 334 (fig.), 
335 


Cephus, 471 
cera, Xanthopimpla, 411 
Ceratogastra, 463 
ornata ornata, 463, 464 (fig.) 
Chalcomys subg., 519, 543 
chalybaeum, Apion, 250, 266 
Charops, 491 
ganges, 489, 491 
narangae, 484 (fig.), 489, 491 
obtusus, 491, 492 
papilionis, 483 (fig.), 490, 491 
Charopsimorpha, 489, 490 
tibialis, 491 
unicincta, 488 (fig.), 491 
Chasmias, 456 
scelestus, 456 
chilcotinae, Bradynotes, 48 
Buckellacris, 48 
Chiroptera, 108, 205 
Chorinaeus, 480 
sp., 481 
christiei, Plecotus auritus, 132 
Chrysopidae, 428 
chuparosae, Apion, 250, 258, 291, 293 
fi 


g.) 

cinctipes, Exetastes, 463 
cinereus, Lasiurus, 118, 119 
cinnamomeus, Mus, 515, 519, 561, 562 

Oecomys, 561 
circumflexum, Therion, 500 (fig.), 504 
citrina, Xanthopimpla, 411 
clandestinus, Holocremnus, 487 (fig.), 

491 

clapini, Rhorus, 467 (fig.), 468 
claripennis, Exenterus, 426 
clarus, Platylabus, 449 (fig.) 
clavipes, Atrometus, 499 (fig.), 503 
cockerelli, Melanoplus, 3, 75 
Coelichneumon, 456, 458 

pepticus, 456, 457 (fig.) 
Coelocephalapion subg., 248 


604 


PROCEEDINGS 


coeruleipes, Melanoplus, 3, 15 
Melanoplus atlanis, 3, 14 
cognatus, Neohaematopinus, 583, 584 
(fig.), 585, 587 
Coleocentrus, 415 
excitator, 417 
Coleoptera, 247, 411, 413, 415, 481, 507 
Coleopterapion subg., 248 
collaris, Diadromus, 558 
Collyria, 392 
calcitrator, 471, 473 (fig.), 474 
Collyriinae, 397, a1; 473 (fig.), 506 
ere Apion, 251, 258, 378, 379 
(fig. 
commodum, Pi 251, 252, 378, 379 
(fig.), 3 
complex, eenaeblue mexicanus, 12 
compressus, Horogenes, 491 
comptoniellae, Horogenes, 486 (fig.), 491 
compunctor, Apechthis, 408 (fig.), 410 
concolor, Hesperomys, 515, 545, 551, 
554 
Oryzomys, 513, 515, 518, 523 (fig.), 
524 (fig.), 525, 527, 528, 529, 540, 
542, 545, 548- 549, 559, 561 
Oryzomys concolor, 517, ‘518, 544, 
546, 548-549 
Rattus, 594 
Condylura, Loy 
confertum, Apion, 251, 252, 268, 377, 
378, 379 (fiz.), 381 
conotracheli, Tersilochus, 496 
conquisitor, [toplectis, 410 
consansuineus, Melanoplus, 3, 41, 43 
conspicuus, Melanoplus, 3 
Melanoplus devastator, 
contortionis, Lamachus, 471 


42, 43, 44 


cordatum, Apion, 250, 253, 369, 371 
(fig) 
Cornus, 211 
candidissma (panicled dogwood), 
363 
corrugatus, Enderleinellus, 470, 571 
fig. 
corvina, Bathyplectes, 491 
coryt, Apion, 250, 252, 281, 282, 283 


(fig.), 285 

Corylus rostrata, 50 

corynorhini, Basilia, 174 
Trichobius, 173 

Corynorhinus, 105, 106 
alleganiensis, 137, 209 
macrotis, 161, 163, 190, 201 
macrotis intermedius, 199, 197 
macrotis pallescens, 141, 185, 190 
macrotis townsendii, 197 
megalotis megalotis, 190, 201 
megalotis mexicanus, 141, 185 
megalotis pallescens, 190 
magalotis townsendii, 197 
mexicanus, 141 
phyllotis, 130 
rafinesquei rafinesquei, 189 
rafinesquii, 161, 163, 201 
rafinesquii mexicanus, 141, 185 
rafinesquii pallescens, 190 


OF THE NATIONAL MUSEUM 


VOL. 110 


Cory norhinus, rafinesquii townsendii, 197 


tetralophodon, 140, 270 
townsendi, 190, 197 
townsendii australis, 185 
townsendii ingens, 189 
townsendii townsendii, 197 
townsendii virginianus, 201 
subg. 95, 96, 98, 108, 104, 105, 
109, T1O,. M2 11S) 14s aS: 
116, 117 (fig.), 129, 1386 (key), 
140, 143, 148; 166, 207° (is), 
209, 214 (fig. ), 215, 216 (fig.) 
coxale, Apion, 249 
crassidens, Plecotus, 208, 213, 215 
crassifemur, Campoplex, 490, 492 
crassinasum, Apion, 289, 290 
Cratichneumon, 456 
duplicatus, 456 
unifasciatorius, 456 
Cremastini, 398, 492, 
(fig.), 495 (key) 
Cremastus, 495 
carpocapsae, 495 
epagoges, 495 
facilis, 495 
ferrugineus, 484 (fig.), 495 
forbesi, 495 
interruptor, 495 
minor, 495 
Cricetinae, 514 
Crocidura, 594 
caerula, 594 
Cryptinae, 397, 400, 425 (fig.), 426, 
427 (key), 428, 4380 (fig.), 431 
(fig.), 482 (fig.), 433 (fig.), 434 
(fig.), 485 (fig.), 437 (fig.), 438 
(fig.), 440 (fig.), 441 (fig.), 442 
(fig.), 443 (fig.), 444 (fig.), 445 
(fig.), 504, 505 (fig.), 506 
Cryptini, 397, 427, 439 (key), 440 (fig.), 
441 (fig.), 442 (fig.), 443 (fig.), 
444 (fig.), 445 (fig.) 
Cryptus, 439 
albitarsus albitarsus, 440 (fig.), 443 
inornatus, 446 
laborator laborator, 446 
sexannulatus, 426, 446 
Cteniscini, 419, 420, 425 (fig.), 
Cubocephalus, 427, 436 
abdominator, 439 
brumatae, 439 
sp., 486, 437 (fig.) 
cultrator, Campoplex, 492 
Curculionidae, 481 
curculionis, Bathyplectes, 484 (fig.), 491 
curvator, Triclistus, 479 (fig.), 480 


493 (fig.), 494 


426 


davidis, Apion, 250, 347, 349 (fig.), 

dawsoni, Melanoplus, 76 

defectus, Melanoplus, 3, 38, 45 

Melanoplus bilituratus, 3, 7, 11, 12, 

13, 14, 16, 17 Gig), 19%hee2, 
27 (fig), 28, 30, 31) Gigsk ), 37 
(fig.) , 38, 45, 59 (fig.) 

degener, Acrodactyla, 407 


351 


INDEX 


Delomerista, 399, 401, 405 
diprionis, 401, 404 (fig.), 405 
sp., 401, 405 
denticulatus, Hulinognathus, 597 
depressus, Campoplex, 490 
detritus, Scambus, 404 
devastator, Melanoplus, 3, 4, 7, 9, 12, 
13) (fig.), V7 Gigs), doe Gene 2d 
(fig.), Giga; 33) Giga) 137 
(fig.), 41, 48, 54 
Melanoplus mexicanus, 12 
Diadromus, 448 
collaris, 448 
Diastrophus, 474 
Dichroplus mexicanus, 10 
difficilis, Peromyscus, 145 
dimidipennis, Melanoplus, 3, 15, 45 
diminutus, Melanoplus 3, 41, 43 
dimorphus, Calliephialtes, 401, 405 
Diplazon, 478 
fissorius, 478 
laetatcrius, 476, 477 (fig.), 478 
orbitalis, 476 
scutellaris, 476 
tetragonus, 478 
Diplazoninae, 398, 476, 477 (fig.), 478 
(key), 506 
Diplolepis, 474 
Diprion, 410 
Diprionidae, 426, 481 
diprionis, Delomerista, 401, 404 (fig.), 
405 
discitergus, Mesochorus, 458 (fig.) 
disparis, Phobocampe, 486 (fig.), 491 


dolichops, Erromenus, 420, 423, 425 
(fig.) 

dolosum, Apion, 250, 258, 369, 370, 371 
(fig.) 


douglasi, Sciurus, 585 
Tamiasciurus, 585, 586 
dremomydis, KMnderleinellus, 571, 572 
Dremomys pernyi, 572 
rufigenis, 592 
drummondi, Malvaviseus, 366 
dryas, Oryzomys, 518, 533, 534, 538, 540 
Rhipidomys, 518, 533 
duplicatus, Cratichneumon, 456 
Dusona, 490 
glauca glauca, 491 
quebecensis, 491 
terebrator, 491 
variabilis, 491 
vitticollis vitticollis, 485 (fig.), 491 


eccentricum, Apion, 250, 323, 324 
Echthromorpha, 410 
formosa, 410 
insidiator, 410 
notularia, 409 (fig.), 410 
Echthrus, 443 
niger, 443, 445 (fig.) 
reluctator, 446 
Eelytus, 421 
fontinalis, 421 
ornatus, 421, 422 (fig.) 
pleuralis, 421 


605 


ega, Lasiurus, 118, 119 
Eiphosoma, 492, 495 
annulatum, 495 
batatae, 494 (fig.), 495 
insularis, 495 
texanum, 495 
ejuncidum Anomalon, 503 
Hlateridae, 503 
elbeli, Neohaematopinus, 592, 593 (fig.) 
elegans, Ephialtes, 405 
Neliopisthus, 421 
Emballonuridae, 206 
emiliae, Oecomys, 519 
Rhipidomys, 519 
empusa, Thecla, 489 
Endasys, 429 
subclavatus, 433 (fig.), 485 
Inderleinellus corrugatus, 670, 571 (fig.) 
dremomydis, 571, 572 
kumadai, 673 (fig.) 
larisci, 573, 574 
malaysianus, 571, 573, 674 
menetensis, 573, 574 
rufigenis, 572 
enderleini, Xanthopimpla, 411 
endersi, Oecomys, 533 
enecator, Trichomma, 504 
Enicospilus, 496, 501 
americanus, 501 
arcuatus, 497 (fig.) 
macrurus, 501 
purgatus, 501 
sp., 501 
enoplus, Apion, 250, 254, 309, 311 (fig.), 
314 


ensator, Campoplex, 492 
enurae, Olesicampe, 490, 491, 492 
epagoges, Cremastus, 495 
Ephialtes, 401, 405 
elegans, 405 
irritator, 403 (fig.), 405 
manifestor, 405 
punctulatus, 405 
ruficollis, 405, 506 
Ephialtini, 395, 399, 400, 401, 402 (fig.), 
403 (fig.), 404 (fig.), 504 
Epiurus brevicornis, 404 
eppelsheimi, Apion, 279 
Eptesicus, 118, 169, 171, 204, 205, 206, 
207 (fig.) 
fuscus, 170, 172, 206 
eques, Lamachus, 471 
Eripternimorpha, 443 
javensis, 443, 444 (fig.) 
erismata, Hoplopleura, 578, 579, 480 
fi 


Erromenus, 423, 426 

bedardi, 423 

dolichops, 420, 423, 425 (fig.) 
erythraeus, Callosciurus, 581 
erythrina, Piscidea, 340 
esculenta, Psoralea, 381 
Ethelurgus, 429 

syrphicola, 431 (fig.), 435 
Euceratini, 466, 467 (fig.), 468 


606 


Euceros, 465, 468 
frigidus, 465 
thoracicus, 467 (fig.), 468 
EKuderma, 95, 96, 103, 104, 105, 106, 107 
(figs.), 108, 109, 111, 118, 119, 
120, 121 (key), 122, 207 (fig.), 
210, 212, 215 
maculata, 122 
maculatum, 98, 107 (fig.), 108, 109, 
116, 122, 123, 139, 140, 218, 224 
Eulinognathus, 597 
denticulatus, 597 
euphydryadis, Benjaminia, 491 
eupitheciae, Casinaria, 483 (fig.), 491 
eureka, Horogenes, 491 
Kurylabini, 448, 441 (fig.) 
Kuryproctini, 465, 466, 468, 471 (key), 
472 (fig.) 
euryptychiae, Pristomerus, 495 
evotis, Myotis, 170 
evustum, Apion, 249, 250, 255, 331, 332, 
334 (fig.), 337, 347 
exareolatus, Horogenes, 492 
excelsus, Melanoplus 3, 45, 46 
excerta, Netekia, 420 
excitator, Coleocentrus, 417 
Exenterus, 419 
abruptorius, 426 
adspersus, 426 
canadensis, 425 (fig.), 426 
claripennis, 426 
tricolor, 426 
Exetastes, 398, 459, 463 
bifenestratus, 463, 464 (fig.) 
cinctipes, 463 
illinoiensis, 463 
sp., 463 
exiguus, Bathyplectes, 491, 492 
Exxochus, 480 
albifrons, 480 
scitulus, 479 (fig.), 481 
extremus, Caloptenus, 3, 67 
Melanoplus, 3, 69 


facilis, Cremastus, 495 

femoralis, Banchus, 463, 464 (fig.) 

femur-rubrum, Melanoplus, 2, 5, 8, 13 
(fig.), 14, 15, 52,53 

ferrugineus, Callosciurus, 580 

Cremastus, 484 (fig.), 495 

finlaysoni, Callosciurus, 573, 581, 585 

fissorius, Diplazon, 478 

flavescens, Banchus, 463 

flavicans, Oryzomys, 545, 548-549, 550, 
551, 553 

flavicens, Oryzomys, 545 

Se Opheltes glaucopterus, 469 


g. 
flavipes, Phobocampe, 491 
flavo-orbitalis, Zaleptopygus, 495 
florenciae, Oecomys, 535, 536-537, 538 
foliae, Seambus, 404 

foliicola, Alophosternum, 402 (fig.), 404 
fontinalis, Eelytus, 421 

forbesi, Cremastus, 495 

formosa, Echthromorpha, 410 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


formosus, Arotes, 415, 416 (fig.) 
Myotis, 109 
fossilis, Miniopterus, 206 
foutsi, Ophionellus, 503 
Fragus, 211 
fraternum, Apion, 306, 307 
Fraximus, 211 
frigidus, Kuceros, 465 
frontalis, Oryzomys, 546, 548-549, 550, 
551, 552 
Oryzomys tectus, 546 
frustranae, Campoplex, 490 
fugitivus, Hyposoter fugitivus, 491 
fulgurans, Mesochorus, 459 
fulvicorne, Heteropelma, 500 (fig.), 504 
fulviventer, Oryzomys, 553, 554, 555, 
556 


fumiferanae, Glypta, 460 

Fungivoridae, 474 

furtivum, Apion, 251, 253, 268, 377, 378, 
382 


fuscipennis, Benjaminia, 488 (fig.), 491 

fusconitidum, Apion, 250, 332, 334 (fig.), 
338 

fuscus, Eptesicus, 170, 172, 206 


gallaecola, Zaleptopygus, 495 
Gambrus, 397, 426, 439 
canadensis, 441 (fig.), 443 
stokesi, 443 
ganges, Charops, 489, 491 
Gelinae, 426 
Gelini, 428 
Gelis, 435 
bruesii, 429, 435 
melanocephala, 436 
tantillus, 429, 433 (fig.), 435 
tenellus, 436 
gelitorius, Phytodietus, 420 
geminatus, Netalia, 419 (fig.), 420 
glauca, Dusona glauca, 491 
Glaucomys, 587, 590 
Glauconycteris superba, 109 
glaucopterus, Opheltes, 468 
glyphicum, Apion, 250, 258, 291, 292, 
293 (fig.) 
Glypta, 460 
fumiferanae, 460 
haesitator, 460 
parvicaudata, 460 
sp., 460, 461 (fig.) 
godmani, Apion, 250, 254, 262, 295, 296, 
297, 299 (fig) 
Glyptini, 459, 460, 461 (fig.) 
Goryphus, 439 
inferus, 442 (fig.), 443 
gracilirostre, Apion, 250, 257, 326, 347, 
349 (fig.), 362 
gracilis, Mesostenus, 440, 443 
Oryzomys alfaroi, 530 
grallator, Labena, 413, 414 (fig.) 
grandis, Plecotus, 208 
grapholithae, Agrothereutes, 443, 446 
Grasshoppers of the mexicanus group, 
genus Melanoplus (Orthoptera: 
Acrididae), 1 


INDEX 


Gravenhorstiini, 498 (fig.), 499 (fig.), 
502, 503 (key) 
griseum, Apion, 250, 255, 295, 296, 299 
(fig.), 300, 306, 307, 308 
grivensis, Rhinolophus, 208 
Vespertilio, 208 
grossulum, Apion, 250, 255, 269 
guatemalense, Apion, 250, 270 
guatemalenum, Apion, 350, 351 
guianae, Oecomys, 553, 554, 555, 556 
Oecomys guianae, 553 
gulare, eplon, 250, 257, 332, 338, 339 


(fig. 

Gurney, Ashley B., and Brooks, A. R.; 
Grasshoppers of the mexicanus 
group, genus Melanoplus (Or- 
thoptera: Acrididae), 1 


hadromerum, Apion, 250, 258, 326, 327 
(fig.), 328 

Haematopinus sciurinus, 585 

haesitator, Glypta, 460 

haeussleri, Phaeogenes, 448 

Handley, Charles O., Jr.; A revision of 
American bats of the genera 
Euderma and Plecotus, 95 

hardwicki, Nesokia, 594 

hariolus, Phaeogenes, 447 (fig.), 448 

harrisoni, Hylopetes sagitta, 589 

havanensis, Carinodes, 455 (fig.), 456 

hawaiiensis, Rattus, 577 

Rattus exulans, 577 

hebrus, Phaeogenes, 448 

Helianthus sp., 308 

helvola, Strophostyles, 308 

helvolus, Oryzomys, 546, 548-549, 550, 
553 


hemipterus, Hemiteles, 436 
Hemiteles, 392, 429 
areator, 433 (fig.), 485, 436 
hemipterus, 436 
simillimus, 436 
Hemitelini, 427, 428 (key), 430 (fig.), 
431 (fig.), 482 (fig.), 433 (fig.), 
434 (fig.), 435 (fig.), 505 (fig.) 
Herpestomus brunnicornis, 448 
Hershkovitz, Philip; Mammals _ of 
Northern Colombia, preliminary 
report No. 8: Arboreal rice rats, 
a systematic revision of the sub- 
genus Oecomys, genus Ory- 
zomys, 513 
Hesperomys, 519 
anguya, 552 
bicolor, 518, 534 
concolor, 515, 545, 551, 554 
longicaudatus, 534 
Myoxomys bicolor, 534 
russatus, 552 
simplex, 519, 520 
Heterapion, subg., 248 
heterogeneum, Apion, 250, 255, 369, 
372, 373 (fig.) 
Heteropelma, 501, 504 
fulvicorne, 500 (fig.), 504 
hirsuta, Hoplopleura, 575 (fig.), 576, 577 


546764—60——3 


607 


hispae, Scambus, 402 (fig.), 404 
Histiotus, 104, 105, 106, 122 
maculatum, 105 
maculatus, 122 
Holochilus brasiliensis, 552 
Holocremnus, 490 
clandestinus, 487 (fig.), 491 
ratzeburgi, 491, 492 
homochrous, Plecotus auritus, 132 
Hoploecryptus, 443 
incertulus, 442 (fig.), 443 
Hoplopleura, 581 
erismata, 578, 579, 580 (fig.) 
hirsuta, 575 (fig.), 576, 577 
malabarica, 575 (fig.), 594 
oenomydis, 577 
pacifica, 577 
pectinata, 569 
thurmanae, 577, 578 (fig.), 580 (fig.) 
Hordeum sp., 80 
Horogenes, 490, 491, 492 
acutus, 491 
aestivalis, 491 
armillata, 492 
compressus, 491 
comptoniellae, 486 (fig.), 491 
eureka, 491 
exareolatus, 492 
macrostoma, 492 
nana, 492 
obliteratus, 491 
pterophorae, 491 
punctorius, 491, 492 
sp. 491, 492 
huardi, Rhexidermus, 447 (fig.), 448 
hudsonicus, Tamiasciurus, 586 
Tamiops, 582 
hyalinus, Aphanistes, 493 (fig.), 503 
Campoplex, 490 
Hybophanes, 421 
scabriculus, 417, 421, 422 (fig.) 
Hylopetes phayrei, 587 
sagitta harrisoni, 589 
Hymenoepimecis sp., 406 (fig.), 407 
Hyposoter, 489, 490, 491, 492, 506 
fugitivus fugitivus, 491 
fugitivus pacificus, 491 
nigrolineatus, 491 
parogyiae, 490, 491, 492 
pilosulus, 486 (fig.), 491, 492 
rubiginosus, 491 
tricolor, 492 
Hysantyx, 465, 466, 468 
lituratorius, 470 (fig.), 471 
hystriculum, Apion, 330 


Ichneumon, 456 
ambulatorius, 455 (fig.), 456 
laetus, 456 
rufiventrus incertus, 456 
suspiciosus, 456 
Ichneumonidae, description and classi- 
fication of the final instar larvae 
of, 391 
Ichneumonidae. 397 (key) 


PROCEEDINGS 


608 


Ichneumoninae, 398, 446 (key), 


OF THE NATIONAL MUSEUM 


VOL. 110 


447 , Ischnoceros rusticus, 415 


(figs.), 449 (fig.), 450 (figs.), 451 | Ischnus polytomi, 446 
(fig.), 452 (figs.), 453 (figs.), 454] Isdromus lycaenae, 428, 431 (fig.), 435 


(figs.), 455 (figs.), 505, 506 
Ichneumonini, 448, 451 (key), 
(figs.), 453 (figs.), 454 (figs.), 
455 (figs.), 457 (figs.) 
Ichoronyssus sp., 174 
Idechthis, 392, 490 
canescens, 491, 492 
nigriscapus, 483 (fig.), 490 
Idiogramma, 397, 417, 423 
sp., 421, 422 (fig.) 
Idiogrammatini, 397, 419, 421, 422 (fig.) 
Idionycteris mexicanus, 129, 130 
phyllotis, 130 
subg., 95, 96, 104, 105, 106, 107, 
109, 110) 112) 113. 117, 128" 729, 
154, 207 (fig.), 212 
idoneus, Ophion, 497 (fig.), 501 
illectus, Oryzomys, 545, 548-549, 550 
Oryzomys flavicans, 545 
illinoiensis, Exxetastes, 463 
imitator, Apion, 250, 257, 261, 262, 281, 
282, 286, 287 (fig.) 
impexum, Apion, 249 
importunum, Apion, 250, 257, 348, 349 
(fig.), 353 
impudicus, Melanoplus, 35 
inaequalipes, Campoplex, 490, 491 
incertulus, Hoplocryptus, 442 (fig.), 443 
incertus, Ichneumon rufiventrus, 456 
inconstans, Amblyteles, 454 (fig.), 456 
indica, Bandicota, 575 
Nesokia, 584 
indistincta, Aptesis, 436, 437 (fig.) 
indubia, Casinaria, 491 
inferus, Coryphus, 442 (fig.), 448 
infesta, Casinaria, 491 
inflatus, Pimpla, 410 
ingens, Corynorhinus townsendii, 189 
Plecotus townsendii, 153, 186, 187, 
188 789, 191, °202, 221, 228 
ainnocens, Apion, 250, 253, 304, 309, 311 
(fig.), 314 
tnnocuum, Apion, 250, 254, 309, 3145, 
316 (fig.) 
inornata, Petaurista, 594 
inornatus, Cryptus, 446 
Neohaematopinus, 592, 593 
insidiator, Barylypa, 498 (fig.), 503 
Echthromorpha, 410 
insignis, Myotis, 206 
instigator, Pimpla, 410 
insularis, Eiphosoma, 495 
Xorides, 415 
interior, Veronia, 369 
intermedius, Corynorhinus 
190, 197 
Melanoplus, 3, 14, 15, 18 
Plecotus townsendii, 192, 193 
interruptor, Cremastus, 495 
Ipoctoninus, 471 
striatus, 471, 472 (fig.) 
irrigator, Xorides, 415 
irritator, Ephialtes, 403 (fig.), 405 


macrotis, 


peruviana, 428, 431 (fig.), 435 


452 | Iselix, 429, 436 


sp., 435 
Iseropus, 401, 405 
ealifornicus, 405 
stercorator brunneifrons, 403 (fig.) 
405 
stercorator stercorator, 405 
[toplectis, 400, 410 
alternans, 410 
behrensii, 409 (fig.), 410 
conquisitor, 410 
maculator, 410 
olivalis, 410 
sp., 410 


javanicus, Pipistrellus, 108 

javensis, aH inua 443, 444 
(fig. 

Johnson, Phyllis T.; The rodent-infest- 
ing Anoplura (sucking lice) of 
Thailand, with remarks on some 
related species, 569 

Joppinae, 446 

Juncus balticus, 80 

junius, Melanoplus, 3, 69, 70 

Melanoplus borealis, 69, 70 
Pezotettix, 3, 68 


Kerivoula picta, 109 
kinabalensis, _Neohaematopinus, 
590 (fig.), 591 (fig.) 
Kissinger, David G.; A revision of the 
Apion subgenus Trichapion Wag- 
ner in the New World (Coleop- 
tera: Curculionidae), 247 

klagesi, Oryzomys, 546, 548-549, 550, 
553 


589, 


Rhipidomys, 546 
kozlovi, Plecotus auritus, 32 
kriechbaumeri, Spudastica, 492 
kumadai, Enderleinellus, 573 (fig.) 


Labena, 413 

grallator, 413, 414 (fig.) 
Labenini, 399, 400, 413, 414 (fig.), 428 
laborator, Cryptus laborator, 446 
Labrorychus, 503 

analis, 498 (fig.), 503 

prismaticus, 503 

sp., 503 
Laephotis, 104, 105, 106 
laetatorius, Diplazon, 476, 477 (fig.), 

478 

laetus, Ichneumon, 456 
Lamachus, 466, 468 

albopictus, 468 

contortionis, 471 

eques, 471 

lophyri, 471 

ruficoxalis, 471 

tsugae, 471 

virginianus, 470 (fig.), 471 

sp., 471 


INDEX 


laminatum, Apion, 247, 250, 264 (fig.), 
QT 


Lampronota, 463 
catenator, 463 
sesiovora, 461, 463 
sp., 461 (fig.) 
lanuginosum, Apion, 279, 280 
larisci, Enderleinellus, 573, 574 
Lariscus, 586 
Lasiurus, 115, 118, 119, 204, 207 (fig.), 
208 


borealis, 118, 119 
cinereus, 118, 119 
ega, 118, 119 
laticeps, Oryzomys, 524 (fig.), 525, 526 
(fig.), 532, 544 
latifercula, Melanoplus, 3, 31 (fig.), 37 
(ig). 75 
latitator, Apion, 150, 253, 309, 316 (fig.), 
317 
lativentre, Apion, 250, 258, 369, 374 
lecontii, Olecotus, 161 
Synotus, 161 
leiosperma, Strophostyles, 308 
Lepidoptera, 421, 439, 446, 478 
pupae, 413, 481 
lepidus, Leptobatopsis, 461, 463 
Leptobatopsis, 463 
lepidus, 461, 463 
sp., 461 (fig.), 463 
leucantha, Baptisia, 386 
Lissonota, 460 
brunnea, 462 (fig.), 463 
Lissonotinae, 398, 399, 459, 460 (key), 
461 (fig.), 462 (fig.), 464 (fig.), 
481, 506 
Lissonotini, 460 (key), 461 (fig.), 462 
fi 


(fig.) 
Listrodromini, 398, 446, 449 (key), 450 


(fig.) 
Listrodromus, 398, 449 
nycthemerus, 450 (fig.), 451 
lituratorius, Hypsantyx, 470 (fig.), 471 
longicaudatus, Hesperomys, 534 
Oryzomys, 530 
lophyri, Lamachus, 471 
Olesicampe, 487 (fig.), 490, 491, 492 
Lophyroplectus, 465, 466, 468, 471 
luteator, 471 
nipponensis, 470 (fig.), 471 
lucifugus, Myotis, 103, 108, 170 
luctunosa, Pimpla, 410 
Lupinus sp., 290 
luteator, Lophyroplectus, 471 
lycaenae, Isdromus, 428, 431 (fig.), 435 
Lycaenidae, 449 
Lymeon, 427, 443 
orbum, 443, 445 (fig.) 
macclellandi, Tamiops, 570, 571, 577, 
57 
Macrojopps, 458 
macrostoma, Horogenes, 492 
macrotis, Corynorhinus, 161, 163, 190, 
201 


Synotus, 161, 164, 190 


609 


macrotis, Plecotus, 136, 154, 155, 156, 
161, 163, 166, 167, 197 
Plecotus rafinesquii, 152, 161, 
220, 227 
Vespertilio, 161 
Macrotus, 132 
californicus, 170 
waterhousii, 205 
macrurus, Hnicospilus, 501 
maculata, Huderma, 122 
maculator, Itoplectis, 410 
maculatorius, Syzeuctus, 463 
maculatum, Huderma, 98, 107 (fig.) 108, 
109, 116, 122, 123, 139, 140, 218, 
224 
Histiotus, 105 
maculatus, Histiotus, 122 
maculifrons, Syrphoctonus, 476, 477 
fig 


164, 


g. 
maculipes, Mus, 519 
Rhipidomys, 519 
malabarica, Bandicota, 594 
Hoplopleura, 576 (fig.), 594 
malaysianus, Enderleinellus, 571, 573, 
574 
Mallochia, 427, 439, 443 
sp., 443, 445 (fig.) 
Malvaviseus drummondi, 366 
Mammals of Northern Colombia, pre- 
liminary report No. 8; Arboreal 
rice rats, a systematic revision of 
the subgenus Oecomys, genus 
Oryzomys, 513 
mamorae, Oecomys, 559 
Oryzomys, 518, 559, 560, 561 
managuense, Apion, 250, 256, 348, 349 
(fig.), 354 
mandoni, Apion, 350 
manifestor, Ephialtes, 405 
marmosurus, Oecomys, 546, 547 
Oryzomys, 518, 546, 553, 554 
Rhipidomys, 546 
marylandica, Meibomia, 363 
Mastrus argeae, 429, 435 
molestae, 429, 430 (fig.), 435 
smithii, 429, 435 
maugei, Plecotus, 203 
Synotus, 203 
Vespertilio, 203 
mediator, Orthopelma, 473 (fig.), 474 
mediocre, Apion, 250, 253, 261, 366, 367 
fi 


g.) 
Megadermatidae, 205 
Megalomys, 514 
megalotis, Corynorhinus megalotis, 190, 
2 

Reithrodontomys, 145 

Vespertilio, 154, 155, 156, 163 
Megastylus, 476 

sp., 474, 475 (fig.) 
Meibomia marylandica, 363 
Melanichneumon, 448, 451 

brevicinctor, 456 

rubicundus, 456 
melanocephala, Gelis, 436 
Melanomys, 514 


610 


Melanoplus, 2, 4 

affinis, 3, 14, 15, 42, 43 

alaskanus, 3, 45, 46, 48 

angelicus, 3, 14 

arcticus, 3, 15 

ater, 3, 41, 43 

atlanisy3, 10) Ti 12. 14 Ab lS) 21, 
34, 55 

atlanis coeruleipes, 3, 14 

bilituratus, «<2):-3,, 10): 12; 13, (fig.), 
15, 17 (fig.), 18, 19 (fig.), 24, 38, 
45, 46, 47, 50, 52, 55, 65, 66, 72 

bilituratus bilituratus, 3, 4, 7, 13 
(fig.), 14, 17 (fig.), 18, 19 (fig.), 
ae 255 2 (ties), 628, 29, oi) (figs), 

ane (fig.), 34, oy, (fic.), 38, 40, 

re 44, 45, 52, 64, 70 

bilituratus defectus, 3, 7, 11, 12, 13, 
14, 16, 17 (ig.); 19 (fig),, 21, 27 
(fig.), 28, 30, 31 (fig.), 37 (fig.), 
38, 45, 59 (fig.) 

bilituratus vulturnus, 4, 7, 12, 138 
(fig.), 14, 17 (fig.), 19 (fig.), 26, 
27 (fig.), 29, 31 (fig.), 33 (fig.), 
37 (fig.), 56 (fig.), 65 

borealis, 3 3, 67, 68, 69, 74 

borealis borealis, 2, 3, 4, 8, 19 (fig.), 
27 (fig.), 31 (fig.), 33 (fig.), 37 
(fig.), 49 (fig.), 59 (fig.), 67, 73, 
alee 78, 79, 82 

borealis junius, 69, 70 

borealis monticola, 70, 76 

borealis palaceus, 4, 8, 31 (fig.), 33 
(fig.), 37 (fig.), 49 (fig.), 59 (fig.), 
Ws (25 71,78, Se 

borealis stupefactus, 3, 4, 8, 31 
(fig.), 33 {ig.), 37 (fg) 49 (fig.), 
59 (fig. Ve ls TOs 1S 

borealis utahensis, 4, 8, 31 (fig.), 33 
(fig.), 37 {ie 49 (fig.), 59 (fig.), 

? 

bruneri, 2 4, Por (figs) lo ypla Cues) 
19 (fig.), 27 (fig.), 33° (fig.), 37 
(fig.), 38, 45, 49 (fig.), 51, 51, 
59 (fig.), 69, 82, 83 

caeruleipes, 3, 15, 55 

cockerelli, 3, 75 

coeruleipes, 3, 15 

consanguineus, 3, 41, 43 

conspituus, 3 

dawsoni, 76 

defectus, 3, 38, 45 

devastator, 3, 4, 7, 9, 12, 13 (fig.), 
17 (fie.), 19 (fig.),; 27. fig.) 31 
(ag.), 33 (fig.), 37 (fig.), 41, 43, 


devastator affinis, 42, 43 
devastator conspicuus, 42, 43, 44 
devastator obscurus, 42, 43 
devastator typicalis, 42, 43 
dimidipennis, 3, 15, 45 
diminutus, 3, 41, 43 

excelsus, 3, 45, 46 

extremus, 3, 69 

extremus scandens, 68 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


Melanoplus, femur-rubrum, 2, 5, 8, 13 
fig.), 14, 15, 52, 53 
impudicus, 35 
intermedius, 3, 14, 15, 18 
junius, 3, 69, 70 
latifercula, 3, 31 (fig.), 37 (fig.), 75 
mexicanus, 1, 2,3, 4; 0 85, Lae 
17 (fig.), 19 (fig.), 21, 24, 27 (fig. ys 
29, 35, 37 (fig. a 44, 46, 52, 55, 
64, 65, 66 
mexicanus bilituratus, 18 
mexicanus complex, 12 
mexicanus devastator, 12 
mexicanus mexicanus, 12, 15, 18, 39 
monticola, 4, 68, 69, 70, 76 
obscurus, 4 
palaceus, 4 
parvus, 4, 69 
rugglesi, 60, 66 
sapellanus, 4, 75, 
scandens, 4, 69 
scriptus, 4, 15 
selectus, 4 
septentrionalis, 4, 37 (fig.), 69 
sierranus, 4, 15 
spretis, 4, 55 
spretus, 1, 2 bids 4 0; LON aes 
(fig.), 17 (fig.), 18, 19 (fig.), 22, 
27 (fig. 5 290037 (fig.), 52, 54, 56 
(fig.), 59 (fig.) 
spretus caeruleipes, 3 
stupefactus, 4 
typicalis, 4 
uniformis, 4, 41, 43 
utahensis, 52, 81, 83 
virgatus, 4, 42, 43 
sp., 17 (fig.) 
melleus, Oecomys, 556, 557, 558, 559 
menetensis, Enderleinellus, 573, 574 
Menetes, 579 
berdmorei, 574, 578, 583, 585, 587 
berdmorie rufescens, 574 
sp., 585 
peor: yeacra: Apion, 250, 255, 274, 
27 


76 


meridensis, Oryzomys, 524 (fig.) 
meridionalis, Plecotus auritus, 132 
mesoanthus, Rhorus, 468 
Mesochorinae, 398, 399, 458 (fig.), 459 
(key), 506 
Mesochorus, 458, 459 
discitergus, 458 (fig.) 
fulgurans, 459 
vitticollis, 459 
sp., 458 
Mesoleiina, 466 
Mesoleiinae, 397, 398, 459, 465, 467 
(fig.), 468 (key), 469 (fig.), 470 
(fig.), 472 (fig.), 481, 506 
Mesoleiini, 397, 465, 466, 468 (key), 
469 ( (fig.), 470 (fig.) 
Mesoleius, 466, 468, 471 
tarsilas, 468 
tenthredinis, 468, 469 (fig.), 471 
Mesostenini, 439 


INDEX 


Mesostenus, 439 
gracilis, 440, 443 
sp., 440 (fig.), 443 
metallicum, Apion, 249 
Metopiinae, 398, 478, 479 (fig.), 480 
(key), 505, 506 
Metopius, 480 
anxius, 481 
micratorius, 479 (fig.), 480 
mexicana, Pezotettix, 8, 9, 10 
Platyphymus, 9 
Tadarida, 170 
mexicanum, Apion, 251, 258, 260, 378, 
379 (fig.), 383 
Platyphyma, 9 
mexicanus, Caloptenus, 9 
Corynorhinus, 141 
Corynorhinus megalotis, 141, 185 
Corynorhinus rafinesquil, 141, 185 
Dichroplus, 10 
Idionyeteris, 129, 130 
Melanoplus, 1, 2, 3, 4, 7, 8, 12, 14, 
17 (fig.), 19 (fig.), 21, 24, 27 (fig.), 
29, 35, 37 (fig.), 44, 46, 52, 55, 
64, 65, 66 
Melanoplus mexicanus, 12, 15, 18, 
39 


Paradichroplus, 9 
Pezotettix, 3, 8 
Plecotus, 102, 110, 114, 137, 1388, 
139, 140, 141, 142, 143, 144 (fig.), 
145, 146, 147, 153, 166, 167, 170, 
185, 187, 192, 212, 216 (fig.),'219, 
226 
Plecotus megalotis, 143 
Plecotus rafinesquii, 141, 143, 185 
Trigonophymus, 10 
micratorius, Metopius, 479 (fig.), 480 
Microryzomys subg., 514, 533 
Microtus, 165 
milleri, Oecomys, 535, 536-537, 539 
mincae, Oecomys, 547, 548-549, 551, 552 
Miniopterus, 206, 207 (fig.) 
fossils, 206 
minor, Apion, 250, 258, 347, 348, 3564, 
356 (fig.) 
Cannomys badius, 594, 595 
Cremastus, 495 
mirabilis, Asphragis, 462 (fig.), 463 
mirandum, Apion, 250, 253, 309, 316 
(fig.), 318 
mirificum, Apion, 250, 253, 316 (fig.), 
319 
modestum, Apion, 250, 256, 274, 276 


(fig.) 
molestae, Mastrus, 429, 430 (fig.), 485 
monticola, Melanoplus, 4, 68, 69, 70, 76 
Melanoplus borealis, 70, 76 
Promethus, 478 
mordax, Plecotus auritus, 132 
morio, Therion, 504 
Muridae, 513, 514 
Mus cinnamomeus, 515, 519, 561, 562 
maculipes, 519 
musculus, 520 
pyrrhorhinus, 519 


611 


musculus, Mus, 520 
mutabilis, Campoplex, 492 
myosuros, Proechimys, 561 
Myotini, 104, 105, 106 
Myotis, 103, 105, 108, 115, 116, 117, 
118, 135, 148, 165, 171, 204, 206, 
207 (fig.), 211 

antiquus, 206 

austroriparius, 173 

bechsteinii, 206 

ealifornicus, 169, 170, 172 

evotis, 170 

formosus, 109 

insignis, 206 

lucifugus, 103, 108, 170 

mystacinus, 206 

occultus, 103 

oxygnathus, 206 

salodorensis, 206 

sancti-albani, 206, 208 

sodalis, 170 

subulatus, 126, 170 

thysanodes, 108, 169, 170 

velifer, 120 (fig.), 170, 172, 174 

volans, 169, 170 

yumanensis, 108, 169, 170 
mystacinus, Myotis, 206 
Mystipterus, 206 


nana, Horogenes, 492 
nanulum, Apion, 250, 256, 348, 356 
(fig.), 357 
narangae, Charops, 484 (fig.), 489, 491 
Neacomys, 514 
nebraskense, Apion, 250, 254, 295, 301, 
302 (fig.) 
Nectomys, 514, 521 
Neliopisthus, 421 
elegans, 421 
Nemeritis bicingulata, 492 
Neohaematopinus, 570, 585, 592 
antennatus semifasciatus, 585 
callosciuri, 681, 582 (fig.), 584 (fig.), 
585, 586 
capitaneus, 687, 588 (fig.), 589, 590 


(fig. 
cognatus, 688, 584 (fig.), 585, 587 
elbeli, 592, 593 (fig.) 
inornatus, 592, 593 
kinabalensis, 589, 590 (fig.), 591 
(fig.) 
petauristae, 594 
sciuri, 582, 584 (fig.), 585 
sciurinus, 581, 582 (fig.), 583, 584 
(fig.), 585, 586, 587 
sciuropteri, 587, 588, 589, 590 (fig.), 
591 
semifasciatus, 582 (fig.), 584 (fig.), 
586 
Neoxorides borealis, 411, 412 (fig.) 
Neoxoridini, 399, 400, 411, 412 (fig.) 
Nepiera, 490 
benevola, 485 (fig.), 491 
Nesokia, 594 
hardwicki, 594 
indica, 594 


612 


Nesoryzomys, 514 
Netelia excerta, 420 
geminatus, 418 (fig.), 420 
vinulae, 420 
subg., 419, 420 
niger, Echthrus, 443, 445 (fig.) 
Sciurus, 582, 584, 585, 586 
nigricornis, Blaptocampus, 499 (fig.), 


nigridens, Phaeogenes, 448 

nigriscapus, Idechthis, 483 (fig.), 490 

nigrolineatus, Hyposoter, 491 

nigromaculatus, Xanthocampoplex, 482 
(fig.), 490 

nigrum, Apion, 250, 255, 346, 369, 371 
(fig.), 376 

Nippocryptus, 443 

suzukii, 443 (fig.) 

nipvenousis, Lophyroplectus, 470 (fig.), 

471 


nitedulus, Oecomys, 535, 5386-537, 538, 
539 


nitidirostre, Apion, 250, 347, 356 (fig.), 
358 

nitidum, Apion, 250, 256, 339 (fig.), 341 

norvegicus, Rattus, 597, 598 

notularia, Echthromorpha, 409 (fig.), 
410 

novellum, Apion, 250, 254, 295, 303 

nubilipennis, Calliephialtes, 404, 405 

nuncias, Pseudamblyteles, 454 (fig.), 

6 


nunenmacheri, Apion, 289, 291 

Nycteribiidae, 174 

Nycterobius, 206 

nycthemerus, Listrodromus, 450 (fig.), 
451 

Nycticeini, 105, 106 

Nycticeius, 204 

Nyctomys, 522, 526 (fig.) 

Nyctophilinae, 104, 105 

Nyctophilus, 104, 105, 106 


obliteratus, Horogenes, 491 
oblitum, Apion, 250, 253, 366, 367 
(fig.), 368 
obscurum, Apion, 275 
obscurus, Melanoplus, 4 
Melanoplus devastator, 42, 43 
obtusus, Charops, 491, 492 
occidentalis, Oryzomys bicolor, 516, 518, 
533, 540 
occisor, Trichomma, 506 
occultus, Myotis, 103 
Oecomys auyantepui, 553, 554, 555, 556 
benevolens, 540 
bicolor, 518, 534, 535, 538, 539 
caicarae, 547, 548-549, 550, 553 
catherinae, 518, 519, 543 
cinnamomeus, 561 
emiliae, 519 
endersi, 533 
florenciae, 535, 536-537, 538 
guinae, 553, 554, 555, 556 
guianae caicarae, 547 
guianae guianae, 553 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 110 


Oecomys mamorae, 559 


marmosurus, 546, 547 
melleus, 556, 557, 558, 559 
milleri, 535, 586-537, 539 
mincae, 547, 548-549, 551, 552 
nitedulus, 535, 536-537, 538, 539 
osgoodi, 518, 556, 557, 558, 559 
palmeri, 556, 557, 558 
phelpsi, 519 
rex, 519, 543 
roberti, 559 
rutilus, 535, 536-537, 539 
splendens, 553, 554, 555, 556 
superans, 556 
tapejinus, 546, 547, 548-549, 554 
trabeatus, 533 
subg., 514, 615, 518, 519, 523 (fig.), 
525, 5438, 544, 552, 554 
oenomydis, Hoplopleura, 577 
Oenomys sp., 577 
Olesicampe, 481, 491 
enurae, 490, 491, 492 
lophyri, 487 (fig.), 490, 491, 492 
pikonemae, 487 (fig.), 490, 491, 492 
Oligoryzomys, 514 
subg., 530 
olivalis, Itoplectis, 410 
olympiae, Polyterus, 471, 472 (fig.) 
opacicolle, Apion, 249 
Opheltes, 397, 465, 466, 468 
glaucopterus, 468 
glaucopterus flavipennis, 469 (fig.) 
Ophion, 496, 501, 506 
ancyloneura, 501 
bifoveolatus, 496 
bilineatus, 501 
idoneus, 497 (fig.), 501 
tityri, 501 
Ophionellini, 502, 503 
Ophionellus foutsi, 503 
Ophioninae, 398, 459, 463, 465, 466, 
468, 481, 482 (fig.), 483 (fig.), 
484 (fig.), 485 (fig.), 486 (fig.), 
487 (fig.), 488 (fig.), 493 (fig.), 
495, 496 (fig.), 497 (fig.), 501, 
506 
Ophionini, 398, 496, 497 (fig.), 501 (key) 
orbitalis, Diplazon, 476 
orbum, Lymeon, 443, 445 (fig.) 
oregonensis, Pseudischnus, 443, 444 


oriotes, Apion, 250, 255, 296, 299 (fig.), 
304 
ornata, Ceratogastra ornata, 463, 464 


fig. 
ornatus, Eelytus, 421, 422 (fig.) 
Scotomanes, 109 
Orthocentrinae, 397 
Orthopelma, 392 
mediator, 473 (fig.), 474 
Orthopelmatinae 398, 473 (fig.), 474, 
06 


Oryzomys, 513, 514, 515, 518, 521, 527 
fig.) 


g. 
alfaroi, 525 
alfaroi gracilis, 520 


INDEX 613 


Oryzomys auriventer, 556, 557 

benevolens, 540, 541, 542 

bicolor, 513, 515, 516, 517, 523, 
(fig.), 524 (fig.), 526 (fig.), 527, 
528, 529, 530, 531, 532, 533, 534, 
536-537, 542, 543, 544, 554, 555 

bicolor bicolor, 516, 518, 529, 532, 
534, 536-537, 540 

bicolor occidentalis, 516, 518, 33, 
538, 540 

bicolor phaeotis, 515, 516, 518, 540, 
541 


Oryzomys vicencianus, 546, 547, 548- 
549, 550, 553 
xanthaeolus, 543 
oscillator, Apion, 250, 256, 265, 269, 348, 
356 (fig.), 359 
osgoodi, Oecomys, 518, 556, 557, 558, 
559 


Otacustes, 429 

crassus patruelis, 436 

sp., 430 (fig.), 435 
Otonycteris, 104, 105, 106 
oxygnathus, Myotis, 206 
oxylus, Campoletis, 485 (fig.), 491 
Oxyrrhexis, 407 

carbonator texana, 406 (fig.), 407 


bicolor trabeatus, 516, oS 532, 
583, 534 

buccinatus, 552 

catherinae, 518 

concolor, 513, 515, 518, 523 (fig.), 
524 (fig.), 525, 527, 528, 529, 540, 
542, 545, 548-549, 559, 561 

concolor bahiensis, 517, 519, 561 

concolor concolor, 517, 518, 544, 
545, 548-549 

concolor roberti, 517, 519, 559, 560, 
561 

concolor speciosus, 517, 518, 550, 
653 


pacifica, Hoplopleura, 577 
pacificus, Hyposoter fugitivus, 491 
palaceus, Melanoplus, 4 
Melanoplus borealis, 4, 8, 31 (fig.), 
33, (fig.), 37 (fig.), 49 (fig.), 59 
Gig) 7 i227 Ve usS 
Palaeochiropterygidae, 205 
Palaeochiropteryx, 205 
pallescens, Corynorhinus macrotis, 141, 
185, 190 
Corynorhinus megalotis, 190 
Corynorhinus rafinesquii, 190 
Plecotus macrotis, 143 
Plecotus rafinesquii, 190 
Plecotus townsendii, 142, 143, 147, 
152, 153, 154, 183 (fig.), 184 (fig.), 


186, 187, 188, 190, 198, 199, 200, 
222, 230 
pallidus, Antrozous, 169, 170, 182 
pallipes, Adelognathus, 417, 418 (fig.) 
palmarius, Oryzomys, 553, 554, 555, 556 
palmeri, Oecomys, 556, 557, 558 
palustris, Oryzomys, 518, 521, 524 (fig.), 
543 


concolor superans, 517, 519, 531, 
550, 566, 558 

dryas, 518, 533, 534, 538, 540 

flavicans, 545, 548-549, 550, 551, 
553 


flavicans illectus, 545 

flavicans subluteus, 546, 551, 552 

flavicens, 545 

frontalis, 546, 548-549, 550, 551, 
552 

fulviventer, 553, 554, 555, 556 

helvolus, 546, 548-549, 550, 553 

illectus, 545, 548-549, 550 

klagesi, 546, 548-549, 550, 553 

laticeps, 524 (fig.), 525, 526 (fig.), 
532, 544 

longicaudatus, 530 


papilionis, Charops, 483 (fig.), 490, 491 
Paradichroplus mexicanus, 9 
paradoxus, Thaumatotypus, 432 (fig.), 


mamorae, 518, 559, 560, 561 435 
marmosurus, 518, 546, 553, 554 ae ree 255, 295, 296, 302 
d ‘ \ g. , 


meridensis, 524 (fig.) 

palmarius, 553, 554, 555, 556 

palustris, 518, 521, 524 (fig.), 543 

paricola, 535, 536-537 

phaeotis, 540 

rex, 543 

roberti, 518, 559, 560 

rosilla, 535, 536-537 

speciosus, 553, 554, 555, 556 

subflavus, 518, 519, 524 (fig.), 543 

subflavus regalis, 543 

subulatus, 546 

superans, 556, 557, 558, 559 

tectus, 515, 518, 546, 547, 548-549, 
551 

tectus frontalis, 546 

tectus tectus, 546 

trichurus, 546, 548-549, 551, 552 

trinitatis, 547, 553, 554, 555, 556 


Pareptesicus, 206, 207 (fig.) 
paricola, Oryzomys, 535, 536-537 
Rhipidomys, 518, 535 
parogyiae, Hyposoter, 490, 491, 492 
parvicaudata, Glypta, 460 
parvus, Caloptenus, 4, 68 
Melanoplus, 4, 69 
patruele, Apion, 249, 250, 251, 256, 331, 
332, 333, 339 (fig.), 341 
patruelis, Otacustes crassus, 436 
pauciflora, Strophostyles, 308 
pectinata, Hoplopleura, 569 
pedalis, Polyblastus, 423, 425 (fig.) 
pedator, Xanthopimpla, 411 
Pediculus spinulosus, 597 
Pedies virescens, 9 
peninsulare, Apion, 247, 250, 271 
pennator, Trogus, 457 (fig.), 458 
pepticus, Coelichneumon, 456, 457, (fig.) 


614 


PROCEEDINGS 


OF THE 


NATIONAL MUSEUM VOL. 110 


perforicolle, Apion, 249, 250, 257, 331, | pilosulus, Hyposoter, 486 (fig.), 491, 492 


332, 339 (fig.), 343 
Perilissina, 466 
pernyi, Dremomys, 572 
Peromyscus, 165 
difficilis, 145 
perpilosum, Apion, 250, 257, 326, 327 
(fig.), 329 
persulcatum, Apion, 250, 272 
peruviana, Isdromas, 428, 431 (fig.), 435 
pee Apion, 250, 257, 348, 360 
etaurista inornata, 594 
taylori, 594 
petauristae, Neohaematopinus, 594 
Petauristophthirus, 594 
Petauristophthirus petauristae, 594 
Pezotettix junius, 3, 68 
mexicana, 8, 9, 10 
mexicanus, 3, 8 
septentrionalis, 4, 67 
stupefactus, 4, 75 
Phaeogenes, 448 
ater, 448 
haeussleri, 448 
hariolus, 447 (fig.), 448 
hebrus, 448 
nigridens, 448 
vincibilis, 448 
walshiae walshiae, 448 
Phaeogeninae, 446 
Phaeogenini, 447 (fig.), 448 (key) 
phaeotis, Oryzomys, 540 
Oryzomys bicolor, 515, 516, 518, 
540, 541 
Rhipidomys, 518, 540 
Pharotis, 104, 105, 106 
Phaseolus polystachys, 301 
retusa, 304, 305 
sp., 301 
phayrei, Hylopetes, 587 
phelpsi, Oecomys, 519 
Phobetes, 429 
thyridopteryx, 432 (fig.), 435 
Phobocampe, 489 
disparis, 486 (fig.), 491 
flavipes, 491 
unicincta, 492 
phthorimaeae, Campoplex, 490 
Phygadeuon, 429, 436 
subfuscus, 434 (fig.), 435 
phyllotis, Corynorhinus, 130 
Idionycteris, 130 
Plecotus, 98, 107 (fig.), 110, 112, 
P13, 5114 015 116, 117, Lis. eat 
(fig.),, 129; 730; 131, 139, 140, 
212, 218, 224 
Phylophaga, 496 
puyedee Psoralea, 370 
hytodietini, 418 (fig.), 419, 420 (key) 
Phytodietus, 420 
gelitorius, 420 
pulcherrimus, 417, 418 (fig.), 420 
picta, Kerivoula, 109 
pictus, Aoplus, 456 
pikonemae, Olesicampe, 487 (fig.), 490, 
491, 492 


Pimpla, 410 
aequalis, 408 (fig.), 410 
inflatus, 410 
instigator, 410 
luctunosa, 410 
turionellae, 410 
sp., 410 
Pimplinae, 397, 398, 399, 400 (key), 
402 (fig.), 403 (fig.), 404 (fig.), 
406 (fig.), 408 (fig.), 409 tae)? 
412 (fig.), 414 (fig.), 416 (fig.), 
428, 504, 505, 506, 507 
Pimplini, 397, 398, 399, 400, 407, 408 
(fig.), 408 (fig.), 409 (fig.), 410 
(key), 417, 504, 506, 507 
Pimplopterus, 463 
rubicus, 462 (fig.), 463 
Pinus, 421 
Pionini, 466, 467 (fig.), 468 
Pipistrellini, 105, 106 
pipistrellus, Pipistrellus, 135 
Pipistrellus, 108, 115, 116, 117, 118 
javanicus, 108 
pipistrellus, 135 
subflavus, 108, 170, 172 
Piscidea erythrina, 340 
Platylabini, 448, 449 (fig.) 
Platylabus clarus, 449 (fig.) 
Platyphyma mexicanum, 9 
Platyphymus mexicana, 9 
Plecautus, 132 
Plecotinae, 104 
Plecotus, 95, 96, 103, 104, 105, 106, 107 
(figs.), 108, 109, 111,412) 13; 
118, 119, 120 (fig.), 121 (key), 


127, 128 (key), 129, 173, 
176, 204, 205, 207 (fig.), 208 
abeli, 209 


alleganiensis, 136, 137, 138, 140, 
141, 210, 213, 215 

auritus, 98, 105, 107 (fig.), 110, 
112, 114, dig’), 115, (ie. te: 
117  (fiz.), 118, 120 ies) 28 
(fie.), “126, 130, 132)"°133,, 134; 
135, 139). 140° 178;>\2097" 212; 
213, 215, 218, 224 

auritus ariel, 132 

auritus auritus, 132 

auritus christiei, 132 

auritus homochrous, 132 

auritus kozlovi, 132 

auritus meridionalis, 132 

auritus mMordax, 132 

auritus puck, 132 

auritus sacrimontis, 132 

auritus teneriffae, 132 

auritus wardi, 132 

crassidens, 208, 213, 215 

grandis, 208 

lecontii, 161 

macrotis, 136, 154, 155, 156, 161, 
163, 166, 167, 197 

macrotis pallescens, 143 

maugei, 203 


INDEX 


Plecotus, megalotis mexicanus, 143 
mexicanus, 102, 110, 114, 137, 138, 
139, 140, 141, 142, 148, 144 (fig.), 
145, 146, 147, 153, 166, 167, 170, 
185, 187; 192, 212, 216 (fig.), 
219, 226 
phyllotis, 98, 107 (fig.), 110, 112, 
1135 L149 LS Ges, 121 
(fig.), 129, 130, 131, 139, 140, 212, 
218, 224 
rafinesquii, 110, 114 (fig.), 115 (fig.), 
116, 137, 188, 139, 141, 1438, 751, 
152, 163, 164, 166, 167, 168, 202, 
212, 215, 216 (fig.), 220, 227 
rafinesquii macrotis, 152, 161, 164, 
220, 227 
rafinesquii mexicanus, 141, 143, 185 
rafinesquii pallescens, 190 
rafinesquii rafinesquii, 152, 162, 
168, 220, 227 
tetralophodon, 136, 138, 139, 140, 
210, 215 
townsendii, 103, 107 (fig.), 110, 112, 
113, 114 (fig.), 115, 116, 117 (fig.), 
118, 120 (fig.), 121 (fig.), 126, 137, 
138, 
145, 
153, 


ie.) 


139, 140, 141, 143, 144 (fig.), 
146, 147, 148, 149, 151, 152, 
156, 166, 172, 185, 186, 187, 
188, 189, 190, 193, 194, 197, 198, 
210, 212, 215, 216 (fig.) 
townsendii australis, 142, 143, 144 
(fig.), 147, 152, 153, 185, 187, 188, 
191, 192, 221, 228 
townsendii ingens, 153, 186, 187, 
188, 189, 191, 202, 221, 228 
townsendii “intermedius,’’ 192, 193 
townsendi pallescens, 142, 143, 147, 
152, 153, 154, 183 (fig.), 184 (fig.), 
186, 187, 188, 190, 198, 199, 200, 
222, 230 
townsendii townsendii, 153, 186, 
191, 197, 223, 232 
townsendii virginianus, 139, 152, 
153, 186, 187, 190, 201, 223, 232 
subg., 96, 107, 109, 110, 116, 132, 
154, 207 (fig.), 213 
Plectiscinae, 398, 474, 475 (fig.), 476 
(key), 506 
Plectiscus, 476 
varius, 474, 475 (fig.) 
plectrocolum, Apion, 250, 257, 282, 287 
g.), 288 
pleuralis, Eclytus, 421 
Poa sp., 80 
podagricus, Triclistus, 481 
Podisma, 4, 75 
Polistes, 439 
Polyblastus, 423, 426 
pedalis, 423, 425 (fig.) 
polychrosidis, Campoplex, 490 
Polyeyrtus, 439 
semialbus, 440 (fig.), 443 
Polyplax, 595, 596, 597 
asiatica, 594 
cannomydis, 594, 596 (fig.), 598 (fig.) 
spinulosa, 570, 597, 598 (fig.) 


615 


Polysphinctini, 395, 399, 400, 405, 
406 (fig.), 407 (key) 
polystachys, Phaseolus, 301 
Polyterus, 471 
olympiae, 471, 472 (fig.) 
polytomi, Ischnus, 446 
Polytribax, 426 
pomorum, Scambus, 401, 404 
Tromera, 404 
porcatum, Apion, 250, 256, 331, 332, 339 
(fig.), 346 
praecatorius, Xorides, 415 
pretiosus, Brachycyrtus, 425 (fig.), 428 
prevosti, Callosciurus, 574 
prismaticus, Labrorychus, 503 
Pristomerus, 495 
austrinus, 495 
euryptychiae, 495 
pacificus appalachianus, 495 
vulnerator, 495 
Probolus sp., 451 (fig.) 
Prochas, 481, 489 
theclae, 488 (fig.), 489, 491, 492 
proclive, Apion, 250, 258, 274, 281, 282, 
287 (fig.), 289 
Proechimys, 515, 562 
myosuros, 561 
Proenderleinellus, 597 
calvus, 597, 598 (fig.) 
Promethus, 478 
monticola, 478 
propinquicorne, Apion, 250, 255, 274, 
276 (fig.), 277 
Proscus suspicax, 448 
Prosthodocis subg., 420 
Protarchus, 397, 465, 468 
testatorius, 471 
Protichneumon, 456, 458 
sp., 456 
Protichneumonini, 448, 456, 457 (fig.), 
458 (key) 
pseudacacia, Robinia, 377 
Pseudamblyteles, 448, 449, 456 
animosus rubellus, 456 
nuncias, 454 (fig.), 456 
subfuscus, 456 
Pseudischnus, 427, 439 
oregonensis, 443, 444 (fig.) 
Psoralea esculenta, 381 
physodes, 370 
pteronorum, Aptesis, 436 
pterophorae, Horogenes, 491 
Puccinella sp., 80 
puck, Plecotus auritus, 132 
pulcherrimus, Phytodietus, 
(fig.), 420 
punctorius, Horogenes, 491, 492 
punctulatus, Ephialtes, 405 
punctulirostre, Apion, 250, 254, 322, 
323 (fig.) 325 
purgatus, Enicospilus, 501 
pusillus, Sorex, 208 
Pyracmon, 481, 489, 491 
xanthognatha, 482 (fig.), 490 
pyraustae, Campoplex, 490, 491 


417, 418 


616 


pyrrhorhinos, Mus, 519 
Wiedomys, 519 


PROCEEDINGS 


quadrisetosus, Trichobius, 173 
quebecensis, Dusona, 491 
Quercus, 211 
rafinesquei, Corynorhinus rafinesquei, 
189 
rafinesquii, Corynorhinus, 161, 163, 201 
Plecotus, 110, 114 (fig.), 115 (fig.), 
116, 137, 138, 139, 141, 143, 161, 
152, 163, 164, 166, 167, 168, 202, 
212, 215, 216 (fig.), 220, 227 
Plecotis rafinesquii, 152, 162, 163, 
220, 227 
rattus, Rattus, 520, 577, 597, 598 
Rattus concolor, 594 
exulans hawaiiensis, 577 
hawaiiensis, 577 
norvegicus, 597, 598 
rattus, 520, 577, 597, 598 
surifer, 569 
sp., 570, 575, 597, 598 
ratzeburgi, Holocremnus, 491, 492 
reclusum, Apion, 249 
reconditum, Apion, 249, 250, 256, 348 
356 (fig.), 361 
regalis, Oryzomys subflavus, 543 
Reithrodontomys megalotis, 145 
reluctator, Echthrus, 446 
retusa, Phaseolus, 304, 305 
Revilliodia, 206 
rex, Oecomys, 519, 543 
Oryzomys, 543 
Rhembobius, 427, 436 
abdominalis, 436, 438 (fig.) 
Rhexidermus, 448 
huardi, 447 (fig.), 448 
Rhinolophidae, 205 
Rhinolophus grivensis, 208 
Rhipidomys, 518, 522, 526 (fig.) 
benevolens, 515, 518, 540, 541 
dryas, 518, 533 
emiliae, 519 
klagesi, 546 
maculipes, 519 
marmosurus, 546 
paricola, 518, 535 
phaeotis, 518, 540 
roberti, 559 
rosilla, 518, 535, 538, 539 
Rhorus, 465 
clapini, 467 (fig.), 468 
mesoanthus, 468 
Rhypidomys benevolens, 535 
Rhyssa, 392 
sp, 411, 412 (fig.) 
Rhyssini, 399, 400, 411, 412 (fig.) 
rileyi, Xorides, 415, 416 (fig.) 
roberti, Oecomys, 559 
Oryzomys, 518, 559, 560 
Oryzomys concolor, 517, 519, 559, 
560, 561 
Rhipidomys, 559 
Robinia pseudacacia (black locust), 377 


OF THE NATIONAL MUSEUM 


VOL. 110 


rodolphei, Tamiops, 578 

Rosa, 474 

rosae, Zaleptopygus, 495 

rosilla, Oryzomys, 535, 536-537 

Rhipidomys, 518, 535, 538, 539 

rostrata, Corylus, 50 

rostrum, Apion, 249, 251, 257, 307, 308, 
378, 379 (fig.), 384 

rubellus, Pseudamblyteles animosus, 456 

rubicundus, Melanichneumon, 456 

rubicus, Pimplopterus, 462 (fig.), 463 

rubiginosus, Hyposoter, 491 

Rubus, 474 

rufescens, Nenetes berdmorei, 574 

ruficollis, Ephialtes, 405, 506 

ruficoxalis, Lamachus, 471 

rufifemur, Campoplex, 490, 492 

rufigenis, Dremomys, 592 

Enderleinellus, 572 

rufinulum, Apion, 346 

rufipenne, Apion, 249, 250, 255, 331, 
332, 339 (fig.), 346 

rufiventrus, Zootrephus, 476, 477 (fig.) 

rufopectus, Tromatobia, 403 (fig.), 405 

rugglesi, Melanoplus, 60, 66 

russatus, Hesperomys, 552 

rusticus, Ischnoceros, 415 

rutilus, Oecomys, 535, 536-537, 539 


sacrimontis, Plecotus auritus, 132 

sagax, Apion, 350 

Salix strobiloides, 280 

salodorensis, Myotis, 206 

Samonyeteris, 206, 207 (fig.) 

sancti-albani, Myotis, 206, 208 

sancti-felicis, Apion, 250, 253, 309, 316 
(figs) 9320 

sapellanus, Melanoplus, 4, 75, 76 

sayi, Apion, 250, 254, 275, 295, 296, 299 
(fig.), 8300, 301, 306 

seabriculus, Hybophanes, 417, 421, 422 
fi 


(fig.) 
Seambus, 394, 401, 404 
brevicornis, 404 
detritus, 404 
foliae, 404 
hispae, 402 (fig.), 404 
pomorum, 401, 404 
scandens, Melanoplus, 4, 69 
Melanoplus extremus, 68 
Scarabeidae, 496 
Searcobatus sp., (greasewood), 80 
scelestus, Chasmias, 456 
scitulus, Exochus, 479 (fig.), 481 
sciuri, Neohaematopinus, 582, 584 (fig.), 
585 


sciurinus, Haematopinus, 585 
Neohaematopinus, 581, 582 (fig.), 
583, 584 (fig.), 585, 586, 587 
sciuropteri, Neohaematopinus, 587, 588, 
589, 590 (fig.), 591 
Sciurus, 585 
carolinensis, 584, 585, 586, 587 
douglasi, 585 
niger, 582, 584, 585, 586 
vulgaris, 584, 585, 587 


INDEX 


Scolobates, 465 
auriculatus, 466, 467 (fig.) 
Scolobatini, 466, 467 (fig.), 468 
Scolomys, 514 
Scotomanes ornatus, 109 
scriptus, Caloptenus, 4, 14 
Melanoplus, 4, 15 
scrobicolle, Apion, 384, 385 
scutellaris, Diplazon, 476 
selectus, Caloptenus, 4, 8, 9 
Melanoplus, 4 
semialbus, Polyeyrtus, 440 (fig.), 443 
semicastaneum, Apion, 346 
semifasciatus, Neohaematopinus, 582 
(fig.), 584 (fig.), 586 
Neohaematopinus antennatus, 585 
septentrionalis, Melanoplus, 4, 37 (fig.), 
9 


Pezotettis, 4, 67 
Sequoia, 211 
sesiovora, Lampronota, 461 
sexannulatus, Cryptus, 426, 446 
Short, J. R. T.; A description and clas- 
sification of the final instar larvae 
of the Ichneumonidae (Insecta, 
Hymenoptera), 391 
sierranus, Melanoplus, 4, 15 
Sigmodon, 577 
Sigmodontomys, 514 
signatus, Syrphoctonus, 478 
simile, Apion, 249, 250, 258, 273, 274, 
275, 276 (fig.), 278, 282 
simillimus, Hemiteles, 436 
Simonycteris, 206, 207 (fig.) 
simplex, Hesperomys, 519, 520 
smithi, Apion, 249 
smithii, Mastrus, 429, 435 
sodalis, Myotis, 170 
Sorex pusillus, 208 
speciosus, Oryzomys, 553, 554, 555, 556 
Oryzomys concolor, 517, 518, 550, 
568 
spectator, Apion, 325, 326 
Sphecophaga thuringiaca, 438 (fig.), 439 
vesparum, 439 
sp., 439 
Sphecophagini, 427, 428, 439 
spicata, Tephrosia, 344 
spiculiferum, Apion, 250, 272 
Spilogale, 109 
spinitarse, Apion, 250, 254, 309, 321 
spinulosa, Polyplax, 570, 697, 598 (fig.) 
spinulosus, Pediculus, 597 
splendens, Oecomys, 553, 554, 555, 556 
spretis, Acridium, 4, 55 
Melanoplus, 4, 55 
spretus, Caloptenus, 4, 52 
Melanoplus; 1, 2, 3, 4, 7, 10, 12, 13 
(fig.), 17 (fig.), 18, 19 (fig.), 22, 
27 (fig.), 29, 37 (fig.), 52, 54, 
56 (fig.), 59 (fig.) 
Spudastica, 489 
kriechbaumeri, 492 
Stehlinia, 206, 207 (fig.) 
stemmator, Xanthopimpla, 409 (fig.), 
411 


617 


Stenapion subg., 248 
stenomae, Xiphosomella, 495 
stercorator, Iseropus stercorator, 405 
Stilpnus, 428, 436 
anthomyidiperda, 435 (fig.) 
stokesi, Gambrus, 44 
Streblidae, 173 
strenuum, Astiphromma, 458 
striatus, Ipoctoninus, 471, 472 (fig.) 
strobiloides, Salix, 280 
Strophostyles helvola, 308 
leiosperma, 308 
pauciflora, 308 
umbellata, 308 
sp., 301 
stupefactus, Melanoplus, 4 
Melanoplus borealis, 3, 4, 8, 31 
(fig.), 33 (fig.), 37 (fig.), 49 (fig.), 
59 (fig.), 71, 75, 78 
Pezotettix, 4, 75 
subclavatus, Endasys, 433 (fig.), 485 
subflavus, Oryzomys, 518, 519, 524 
(fig.), 543 
Pipistrellus, 108, 170, 172 
subfuscus, Phygadeuon, 434 (fig.), 435 
Pseudamblyteles, 456 
Pub euEs Oryzomys flavicans, 546, 551, 
55 


submetallicum, Apion, 250, 258, 326, 
327 (fig.), 330 

subnudum, Apion, 272 

subrufum, Apion, 249, 250, 256, 263, 
348, 356 (fig.), 363 

subrufus, Amblyteles, 456 

subsequens, Apion, 250, 256, 348, 356 
(fig.), 364 

subtinctum, Apion, 249, 250, 256, 348, 
365 


subulatus, Myotis, 126, 170 
Cryzomys, 546 
Vespertilio, 198 
superans, Oecomys, 556 
Oryzomys, 556, 557, 558, 559 
Oryzomys concolor, 517, 519, 531, 
550, 666, 558 
superba, Glauconycteris, 109 
superciliosum, Apion, 278 
surifer, Rattus, 569 
suspicax, Proscus, 448 
suspiciosus, Ichneumon, 456 
suzukii, Nippocryptus, 4438 (fig.) 
Symphyta, 439 
Synomelix, 471 
sp., 471, 472 (fig.) 
Synotus, 105, 164 
lecontii, 161 
macrotis, 161, 164, 190 
maugei, 203 
townsendi, 190 
townsendii, 197 
syrphicola, Ethelurgus, 431 (fig.), 435 
Syrphidae, 476 
Syrphoctonus, 478 
agilis, 476 
maculifrons, 476, 477 (fig.) 
signatus, 478 


618 


Syrphoctonus tarsatorius, 478 
Syzeuctus aculatmorius, 463 


tabogense, Apion, 247, 250, 273 
tacherdi, Callosciurus finlaysoni, 580 
Tadarida brasiliensis, 172 
mexicana, 170 
Tamiasciurus, 165, 584, 586 
douglasi, 585, 586 
hudsonicus, 586 
sp., 585 
Tamiops hudsonicus, 582 
macclellandi, 570, 571, 577, 
rodolphei, 578 
sp., 578, 580 
tantillus, Gelis, 429, 433 (fig.), 485 
tapajinus, Oecomys, 546, 547, 548-549, 
554 


578 


tarsatorius, Syrphoctonus, 478 
tarsilas, Mesoleius, 468 
taylori, Petaurista, 594 
tectus, Oryzomys, 515, 518, 546, 547, 
548-549, 551 

Oryzomys tectus, 546 
tenebricosum, Apion, 275 
tenellus, Gelis, 486 
teneriffae, Plecotus auritus, 132 (map) 
Tenthredinidae, 421, 423, 426, 481 
ashes tea Mesoleius, 468, ‘469 (fig.), 


Teniiecinipiiees 465, 468 
tenuirostrum, Apion, 249 
tenuiventris, ‘Casinaria, 491 
Tephrosia spicata, 344 
terebrator, Dusona, 491 
Tersilochini, 398, 495, 496 (fig.) 
Tersilochus argentinensis, 496 (fig.) 
conotracheli, 496 
sp., 496 
testatorius, Protarchus, 471 
tetragonus, Diplazon, 478 
tetralophodon, Corynorhinus, 140, 2/0 
Pipeobue 136, 138, 139, 140, 210, 
15 
texana, Oxyrrhexis carbonator, 406 (fig.), 
407 
texanum, Eiphosoma, 495 
thaiwanensis, Callosclurus erythraeus, 
57 
Thaumatotypus, 429 
paradoxus, 432 (fig.), 435 
Thecla empusa, 489 
theclae, Prochas, 488 (fig.), 489, 491, 492 
Therion, 504 
circumflexum, 500 (fig.), 504 
morio, 504 
Therionini, 500 (fig.), 502, 504 (key) 
Theronia, 413 
atalantae, 413, 414 (fig.) 
Theroniini, 399, 400, 401, 413, 414 (fig.) 
thoracicus, Euceros, 467 (fig.), 468 
thuringiaca, Sphecophaga, 438 (fig.), 439 
thurmanae, Hoplopleura, 577, 578 (fig.), 
580 (fig.) 
Thymaridini, 420, 421 (key), 422 (fig.) 
Thyreodon atricolor, 501 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL, 110 


thyridopteryx, Phobetes, 432 (fig.), 435 
thysanodes, Myotis, 108, 169, 170 
Thysiotorus, 429 
sp., 434 (fig.), 435 
tibialis, Charopsimorpha, 491 
tinctoria, Baptisia, 386 
tityri, Ophion, 501 
tosensis, Campoplex, 490, 491 
townsendi, Corynorhinus, 190, 197 
Synotus, 190 | 
townsendii, Corynorhinus macrotis, 197 | 
Corynorhinus megalotis, 197 
Corynorhinus rafinesquii, 197 
Corynorhinus townsendii, 197 
Plecotus, 103, 107 (fig.), 110, 112, 
1138, i14 (fig op aa a 116, 117 
(fig.), 118, 120 (fig.), 121 (fig.), 
126, 137, 188, 139, 140, 141, 143, 
144, (fig.), 145, 146, 147, 148, 
149, 151, 152, 153, 156, 16, 172, 
185, 187, 188, 189, 190, 193, 194, 
197, 198, 210, 212, 215, 216 (fig.) 
Plecotus townsendii, 153, 186, 191, 
197, 223, 232 
Synotus, 197 | 
trabeatus, Oecomys, 533 | 
Oryzomys bicolor, 516, 518, 532, 
533, 534 
Trichapion subg., 247, 248, 249, 251, | 
252 (key), 260, 347 | 
Trichobius corynorhini, 173 
quadrisetosus, 173 
Trichomma enecator, 504 
occisor, 506 
clues Oryzomys, 546, 548-549, 551, 


55 
Triclistus, 398, 478, 480 
curvator, 479 (fig.), 480 
podagricus, 481 
tricolor, Exenterus, 426 
Hyposoter, 492 
tricoloripes, Campoplex, 490 
Trigonophymus mexicanus, 10 
ee ee 547, 553, 554, 555, 
55 


triste, Apion, 279 
tristis, Biolysia, 484 (fig.), 491 
Trogini, 446, 457 (fig.), 448 
troglodytes, Apion, 249 
Trogus, 458 
pennator, 457 (fig.), 458 
Tromatobia, 401, 405 
rufopectus, 403 (fig.), 405 
zonata, 405 
Tromera pomorum, 404 
Trychosis, 427, 439 
sp., 441 (fig.), 443 
Trypanosoma vespertilionis, 174 
Tryphoninae, 397, 417, 418 (fig.), 419 
(key), aie 422 (fig.), 423, 425 
(fig.), 504, 506 
Seles 120, 428, 425 (fig.), 426 


ey 
tsugae, Lamachus, 471 
tuberosa, Apios, 343 
turbulentum, Apion, 361, 362 


INDEX 619 


turionellae, Pimpla, 410 vulgaris, Sciurus, 584, 585, 587 
typicalis, Melanoplus, 4 vulnerator, Pristomerus, 495 

Melanoplus, devastator, 42, 43 vulturnus, Melanoplus bilituratus, 4, 7, 

12, 13 dig.) 5: 14547 Ag.) .e9 
Ulmus, 211 (fig.), 25, 27 (fig.), 29, 31 (fig.), 
umbellata, Strophostyles, 308 33 (fig.), 37 (fig.), 56 (fig.), 65 
unicincta, Charopsimorpha, 488 (fig.), 
491 walsheai, Phaeogenes walshiae, 448 
__ Phobocampe, 492 walshii, Apion, 279, 280 

unifasciatorius, Cratichneumon, 456 wardi, Plecotus auritus, 132 
uniformis, Melanoplus, 4, 41, 43 waterhousii. Macrotus, 205 
utahensis, Melanoplus, 52, 81, 83 Wiedomys pyrrhorhinus, 519 


Melanoplus borealis, 4, 8, 31 (fig.), | Wvyethi lexicaulis. 52 
jean ie 49" (ies 50 
(fig.), 78, 79, 81 xanthaeolus, Cryzomys, 543 
validus, Campoplex, 482 (fig.), 490, 492 | *@2thocampoplex, 490 


Aer nigromaculatus, 482 (fig.), 490 
variabilis, Dusona, 491 sp., 490 


aa ee Calliephialtes, 402 (fig.), 404, xanthognatha, Pyracmon, 482 (fig.), 490 


varipes, Zaglyptus varipes, 405 mone pple, 308/207 )410 


varius, Plectiscus, 474, 475 (fig.) rede ae 

velifer, Myotis, 120 (fig.), 170, 172, 174 enderleinie4ti 

velox, Aoplus, 456 pedator 411 

Veronia interior, 369 stemmator, 409 (fig.), 411 

verrucosa, Betula, 280 vs ? 
Xiphosomella, 495 

vesparum, Sphecophaga, 439 skenoniac. 495 

Vespertilio, 105, 132, 207 (fig.), 208 Z 


auritus, 127, 132 Xorides, 295, 415, 502 
grivensis, 208 Ue ae 

j irrigator 
eee praecatorius, 415 


maugei, 203 ees 
megalotis, 154, 155, 156, 168 Bar aa 


subulatus, 198 


Apes Xoridescopus, 415 
Vespertilionidae, 104, 106, 206 1s 
Vespertilioninae, 104, 106 Sr ae ates 
vespertilionis, Trypanosoma, 174 Monin 399, 400, 401 415 416 (fig.) 
Vespula, 439 428 507. : : ie 


vicencianus, Oryzomys, 546, 547, 548- 
549, 550, 553 Xyela, 421 
vicinum, Apion, 279 


vincibilis, Phaeogenes, 448 yumanensis, Myotis, 108, 169, 170 
vinosum, Apion, 249, 250, 258, 291, 294 
vinulae, Netelia, 420 Zaglyptus, 405 
virescens, Pedies, 9 varipes varipes, 405 
virgatus, Melanoplus, 4, 42, 43 Zaleptopygus, 495 
virginianus, Corynorhinus townsendii, flavo-orbitalis, 495 
201 gallaecola, 495 
Lamachus, 470 (fig.), 471 rosae, 495 
Plecotus, townsendii, 139, 152, 153, sp., 495 
186, 187, 190, 201, 223, 232 Zatypota, 407 
vitticollis, Dusona vitticollis, 485 (fig.), sp., 406 (fig.), 407 
491 zonata, Tromatobia, 405 
Mesochorus, 459 Zootrephus, 478 
volans, Myotis, 169, 170 Zootrephus rufiventrus, 476, 477 (fig.) 


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