/ersity ef California'
SCUDDEB. NORTH AMEEICAN CEUTHOPHILI. 17
THE NORTH AMERICAN CEUTHOPHILI^
BY SAMUEL H. SCUDDER.
Presented May 9, 1894
THE Ceuthophili are wingless Locustarians in which the tarsi are
distinctly compressed rather than depressed, with no pulvilli,* the hind
tibiae furnished on the outer margins above with spines of two distinct
grades,f the fore femora without foramina or genicular spines, the
hind femora with the angle of their insertion on the inner and not on
the outer side beneath, and the antennae strongly approximated at
base. They are all apterous.
With the exception of the genus Troglophilus Krauss, with two
species from European caverns, and the genus Talitropis Bol., with a
single species from New Zealand, placed respectively at one and the
other end of the series, they are known only from America ; and with
the further exception of Heteromallus Brunner, with two species
from Chili, they are all peculiar to the United States and Northern
Mexico. Here they include six genera and sixty-seven species, the
genus Ceuthophilus alone containing above fifty species. The larger
proportion of them, if not all (excepting Udeopsylla nigrd), frequent
dark places, such as burrows, pits, caverns, wells, hollow trees, and
especially the crevices beneath fallen logs.
They were first made known in this country by the descriptions of
Haldeman, Girard, and Harris, and before their time only a single
species from this country had been described, by Burmeister. Not a
species of the group, even the European, was known to Serville. My
first systematic paper, in 1861, was a study of " Rhaphidophora "
(Proc. Bost. Soc. Nat. Hist., VIII.) where seven of our species were
* Brunner states that Gammarotettix has a single pulvillus on the first tarsal
joint; but although the treading surface of this joint (as of the succeeding) is
broad, I can find no indication of a true pulvillus.
t This feature is obscure in Gammarotettix, where there are alternating
longer and shorter spines of such slight inequality as easily to be overlooked,
and which in the Table of Genera given below is ignored.
VOL. xxx. (N. s. xxn ) 2
18 PROCEEDINGS OF THE AMERICAN ACADEMY.
described or catalogued ; but their diversity was hardly fully recog-
nized when in the following year I published my Materials for a
Monograph of the li'orth American Orthoptera (Bost. Journ. Nat.
Hist., VII.), where eighteen species and five genera were characterized
or indicated ; since then a few more species have been described, by
Thomas, Brunner, Bruner, Packard, Walker, Blatchley, and myself,
until now the number of nominal species is twenty-eight, which must,
however, be reduced by synonymy and by reference to other genera to
In 1888, Brunner, in his Monographie der Stenopelmatiden und
Gyllacriden (Verb. Zool.-bot. Gesellsch. Wien), subjected all the spe-
cies known to him to systematic treatment ; but as the larger part of
our species and some of our genera were unknown to him, and the
number of separately described species has multiplied so greatly while
still not including, at least in Ceuthophilus, the half of our species, it
has seemed desirable to undertake a revision of the group, so far as
our native species are concerned, and in the genus Ceuthophilus to
redescribe all the older forms, as well as to publish the novelties.
Accordingly in the present paper thirty-eight additional species of
the group are characterized, together with a new genus, while I shall
further show the validity of Daihinia of Haldeman, which has been
called in question by Brunner, and shall point out first that one of the
genera thought to belong here should be separated as a member of a
distinct group. The total number of genera from North America is
therefore six, and of the species sixty-seven, while a number of other
species are known to me imperfectly by a single sex or poor examples.
Tropidischia Scudd., Bost. Journ. Nat. Hist., vii. 440-441 (1862).
in his Monograph of the Stenopelmatidae, Brunner von Wattenwyl,
from the insufficient data given in my two statements regarding the
structure of this creature, incorrectly surmised that this genus should
be placed in the Ceuthophili, and was perhaps congeneric with Hetero-
mallus, a Chilian genus. Since, however, the hind tibiae are supplied
above with spines of one grade only, it is plainly more nearly related
to the Dolichopodae, from which it may be distinguished by the simi-
larly spined margins of the under surface. It seems to form a group
apart, between the Dolichopodae and Ceuthophili, and of equivalent
value. It appears to come nearer Hadenoecus and Dolichopoda than
to any other described genera.
SCUDDER. NORTH AMERICAN CEUTHOPHILI. 19
In addition to the characters mentioned above and those given in
previous descriptions, I may add that all the legs are tetraquetrous,
with all the margins spined, the spines similar in character and uni-
formly crowded, excepting on the lower margins of the fore femora,
the inner carina of which is sparsely spiued, the outer carina unarmed;
also the lower margins of the middle femora, both carinae of which are
sparsely spined on the apical half ; and the hind femora, the four
carinae of which, even on the swollen portion, are armed excepting at
the extreme base, though both the inferior carinae are rather sparsely
spined. There are no spines on the genicular lobes of the femora,
excepting a very slight one on the posterior side of the middle femora.
There are but two pair of calcaria on the hind tibias, the upper the
longer and less than half as long as the first tarsal joint. The hind
tarsi are very strongly compressed, carinate beneath without pulvilli,
about two fifths as long as the hind tibiae, the first joint nearly as long
as the remaining joints together, the second and fourth joints of the
same length and either of them three times as long as the third.
Finally, the subgenital lamina of the male is ample, the hind margin
entire, with minute styles, consisting of a single bluntly conical joint ;
and the ovipositor is slender, gently arcuate, tapering and acuminate,
unarmed at tip.
Rhaphidophora xanthostoma Scudd.!, Proc. Bost. Soc. Nat. Hist.,
viii. 12 (1861).
Tropidischia xanthostoma Scudd.!, Bost. Journ. Nat. Hist., vii. 441
Originally described from Crescent City, Cal. (A. Agassiz). I have
since received both sexes from Mendocino, Cal., through the favor of
Mr. J. Behrens.
20 PROCEEDINGS OP THE AMERICAN ACADEMY.
TABLE OF THE GENERA OF CEUTHOPHILI.
a 1 . Last palpal joint cleft apically on the under side. Descending
lobes of the mesouotum but little longer than those of the pronotum ;
sides of fore and middle coxae externally laminate, the lamination
elevated to a denticle or compressed spine either mesially or (on middle
legs) apically, occasionally (Hadenrecus) wanting on middle legs. Fore
tibia? not sulcate above ; hind tibiae with spines of two grades on both
outer and inner margins of upper surface. Outer valves of ovipositor
unarmed above before the apex.
b 1 . Palpi long. Hind tibiae usually considerably longer than the
hind femora. Third hind tarsal joint only half or less than half as
long as the second.
c 1 . Vertex obscurely bituberculate at apex. Last palpal joint
no longer or scarcely longer than the third, and cleft beneath
only apically. Middle coxae unarmed. Middle femora with a
feeble genicular spine on posterior margin. Hind tibiae with more
than four pairs of spurs. First hind tarsal joint almost as long
as the others together. Subgenital plate of male triangular and
rather deeply and narrowly emarginate .... Hadencecus.
c 2 . Vertex not tuberculate. Last palpal joint distinctly longer
than the third, cleft beneath for almost its entire length. Middle
coxse spined mesially. Middle femora with a distinct genicular
spine on posterior margin. Hind tibiae with only four pairs of
spurs. First hind tarsal joint generally a third shorter than the
rest combined. Subgenital plate of male ample and broadly
emarginate ,.., Ceuthophilus.
Z> 2 . Palpi short. Hind tibiae shorter or at most but little longer
than the hind femora. Third hind tarsal joint hardly shorter than
the second, or (in Daihinia) wanting. (Lamination of middle coxae
produced inferiorly to the semblance of a spine.)
c 1 . Third palpal joint as long as fifth, the inferior cleft of the
latter extending over only the apical half. Middle femora un-
armed at tip or with a very feeble spine. Hind tibiae shorter or
at least no longer than the hind femora, with few spines of the
second grade on the upper surface, those of the first grade rela-
tively numerous, at least in the 9 , more or less irregularly placed
and of unequal length ; the calcana three in number on each
side, the uppermost generally a little the longest and unusually
distant from the extreme apex, so as to appear rather as an addi-
SCUDDER. NORTH AMERICAN CEUTHOPHILI. 21
tional pair of spurs. Subgenital plate of male greatly produced
and apically deeply fissured.
d 1 . Descending lobes of mesonotum slightly longer than
those of prouotum. Last tarsal joint very much shorter than
the remaining joints together, the third joint normal, though
but little shorter than the second. Subgenital plate of male
ample, rather deeply and broadly emarginate, the sides extend-
ing backwards as slender threads .... Phrixocnemis.
d 2 . Descending lobes of mesonotum no longer than those of
pronotum. Last tarsal joint about as long as the rest together ;
third tarsal joint wanting (as also on fore legs). Subgenital
plate of male immensely produced and so deeply fissured as to
form two tapering ribbons Daihinia.
c 2 . Third palpal joint shorter than the fifth, the inferior cleft of
the latter extending its whole length. Middle femora with a
genicular spine on posterior side. Hind tibia? slightly longer
than the hind femora, with numerous spines of the second grade
uniform in Ungth and pretty regularly separated; calcaria three
in number on each side, the middle one much longer than the
others. (First hind tarsal joint a third shorter than the rest
combined.) Subgenital plate of male ample, apically bitu-
a 2 . (Vertex bituberculate. Palpi short), the last joint apically with
no inferior cleft. Descending, lobes of mesonotum considerably longer
than those of pronotum ; sides of fore and middle coxae neither cari-
date nor spined. (Fore and middle femora unarmed.) Fore tibiae
sulcate above ; hind tibiae (of the same length as the hind femora)
with only one grade * of spines above on the lateral margins ; (calcaria
two in number on each side, subequal and not long. Third hind tar-
sal joint half as long as the second. Subgenital plate of male ample,
apically broadly and not deeply emarginate) ; outer valves of oviposi-
tor serrate above before the apex Gammarotettix.
See introductory remarks.
PROCEEDINGS OP THE AMERICAN ACADEMY.
ffadencecus Scudd., Bost. Journ. Nat. Hist, vii. 439-440 (1862) ;
Brunn., Monogr. Stenop., 66 (1888).
Table of the Species of ffadencecus.
Body pale testaceous. Ovipositor nearly or quite as long as the
body. Subgenital plate of $ broadly emarginate at apex.
Body dark brown. Ovipositor only half as long as the body. Sub-
genital plate of $ narrowly emarginate at apex . . puteanus.
Phalangopsis sp. Thomps., Ann. Mag. Nat. Hist., xiii. 113 (1844).
Rhapidophora cavernarum Sauss., Ann. Soc. Entom. France (4), i.
Hadencecus cavernarum Scudd.!. Proc. Bost. Soc. Nat. Hist., xii.
409 (1869); xix. 38 (1877) ; Boliv. , Ann. Soc. Ent. France (5), x.
72 (1880) ; Riley, Stand. Nat. Hist., ii. 1'84, fig. 260 (1884) ; Corast.,
Intr. Eut., 114 (1888); Blatchl., Proc. Ind. Acad. Sc., 1892, 153.
Rhapidophora subterranea Scudd.!, Proc. Bost. Soc. Nat. Hist, viii.
8 (1861); Pack., Amer. Nat, v. 745, fig. 126 (1871) ; Cope, Ibid.,
vi. 409 (1872) ; Hubb., Amer. Ent, iii. 37 (1880).
ffadencecus subterraneus Scudd.!, Bost. Journ. Nat. Hist., vii. 441
(1862) ; Walk., Catal. Derm. Salt. Brit Mus., i. 201 (1869) ; Pack.,
Guide Ins., 565 (1869) ; Glover, 111. N. A. Ent., Orth., pi. 8, fig. 6
(1872) ; Cope-Pack., Amer. Nat., xv. 882 (1881) ; Brunn., Monogr.
Stenop., 66, fig. 34 (1888); Pack., Mem. Nat. Acad. Sc., iv. 67-70,
83, 116, fig. 16, pi. 17, fig. 3 (1888) ; Id., Psyche, v. 198-199 (1889) ;
Garm., Ibid., vi. 105, fig. (1891).
Early notices of this insect by Telkampf will be found in Muller's
Arch. Anat. Phys., 1844, 318, and Wiedemann's Arch. Naturg., 1844,
384; also by Schiodte in K. Danske Vid. Selsk. Skrift. 1849, 5 ; by
Agassiz in Silliman's Amer. Journ. Sc., 1851, 127; and by Lesque-
reux in the Actes Soc. Helv. Sc. Nat, 40 Sess., 52-53 (1855).
I have studied specimens only from the Mammoth Cave, Ky. It
is also reported by Packard from many other caves in the Mammoth
Cave region, as Dixon's, White's, Salt, Ice, Diamond, Grand Avenue,
Poynters, Wetzel's, Haunted, and Emerson Spring Branch caves;
besides Mail Robbers', One Hundred Dome, Walnut Hill Spring,
SCUDDER. NORTH AMERICAN CEUTHOPHILI. 23
Short, Proctor's, Little Lithographic, and Sugar Bowl caves, and a
cave under Gardiner's Knob, all near Glasgow Junction ; also a
cave near Baker's Furnace, and John and Fred's Cave on the east
bank of Dismal Creek ; further in Carter County caves, viz. Gray
Tom's, Zwingle's, Bat, Van Meter's, Grayson Springs, and Burchell's
caves ; and finally in Nickajack Cave, Tenn. Blatchley also reports it
from Wyandotte Cave, Ind., on the authority of Cope, but it is not so
given by Cope in the references quoted ; and Walker, of course in
error, from the " west coast of America " ! I have also seen specimens
in the Museum of Comparative Zoology, Cambridge, from Turner's
Caves, Pennington Gap, Lee County, Va. (H. G. Hubbard), and
Ely Cave, Lee County, Va. (N. S. Shaler).
Hadencecus puteanus Scudd.!, Proc. Bost. Soc. Nat. Hist., xix. 37
On sides and under covering of wells in North Carolina ; also in
Ceuthophilus Scudd., Bost. Journ. Nat. Hist., vii. 433-434 (1862) ;
Brunn., Monogr. Stenop., 61 (1888).
This is one of the dominant American genera of Locustariae, con-
fined to North America and almost entirely to the United States,
embracing a large number of species in every section of the country,
of which fifty-five are here characterized. Several others are known
to me by single specimens or mutilated examples. The following
table is based principally upon the males. It has been impossible to
construct it so as to bring together the allied species, whose relation-
ship is better shown by the order in which they are described, though
even here the arrangement is far from satisfactory, nearly allied
species being sometimes separated at considerable distances in order to
bring them in closer relation with other allies. Although I have had
six hundred and fifty examples to study at this time, besides others in
alcohol, the material is still insufficient to make a satisfactory disposi-
tion of our species, and I am confident that very many more yet
remain to be discovered.
24 PROCEEDINGS OP THE AMERICAN ACADEMY.
Table of the Species of Ceuthophilus.
a 1 . Second joint of hind tarsi at least half as long again, usually twice
or more than twice as long, as the third.
A 1 . Fore femora one third or more than one third longer than the
pronotum, at least in the $ ; hind tibiae of $ almost always
straight, never greatly bowed.
c 1 . Hind tibiae of at least a tenth longer than the hind
d 1 . Ovipositor much shorter than the fore femora.
e 1 . Hind femora stout, not three times as long as broad,
at least in the $ 1. variegatus.
e 2 . Hind femora slender, four times as long as broad in
the $ .2. ensifer.
d 2 . Ovipositor much longer than the fore femora.
e 1 . Hind tibial spurs less than twice as long as the tibial
depth ; outer carina of hind femora of generally with
some spines at least half as long as the tibial spurs.
f l . Fore femora of $ three fourths as long again as
the pronotum 3. stygius.
f z . Fore femora of $ from one half to two thirds as
long again as the pronotum.
g 1 . Hind femora of $ much less than four times as
long as broad ; hind tibiae of $ very long and more
or less sinuous at base in old individuals.
A 1 . Largest spines of outer carina of hind femora
of $ simple and similar to the others.
h?. Largest spines as above greatly tumid at base.
g*. Hind femora of $ much more than four times as
long as broad ; hind tibiae of $ scarcely more than
one tenth longer than the hind femora, straight.
e 2 . Hind tibial spurs fully twice, generally much more
than twice, as long as tibial depth ; outer carina of hind
femora of $ with no spines a third as long as the tibial
f l . Armature of outer carina of hind femora of $
developed as distinct spines rather than as serrations;
ovipositor arcuate 7. secretus.
SCUDDER. NORTH AMERICAN CEDTHOPHTLI. 25
/ 2 . Armature of outer carina of hind femora of $
developed only as recumbent serrations ; ovipositor
almost or quite straight.
g 1 . Hind femora of slender, almost or quite four
times as long as broad ; hind tibiae exceptionally long,
nearly or quite one sixth longer than the femora.
A 1 . Hind femora of $ more than twice as long as
the fore femora ; ovipositor very feebly arcuate,
only two thirds as long as the hind femora.
h 2 . Hind femora of $ less than twice as long as
the fore femora ; ovipositor straight, three fourths
as long as the hind femora , . . 9. corticicola.
g 2 . Hind femora of $ less slender, being less than
three and three quarters times as long as broad ; hind
tibiae but little more than one tenth longer than the
femora 10. varicator.
c 2 . Hind tibiae of $ distinctly less than a tenth longer than the
hind femora ; ovipositor always longer than the fore femora.
d 1 . Hind tibiae of $ straight; outer carina of hind femora
of ^ never conspicuously spined.
e 1 . Hind tibial spurs nearly three times as long as the
tibial depth 11. latibuli.
e 2 . Hind tibial spurs at most less than twice as long as the
tibial depth, rarely half as long again.
f 1 . Prevailing colors blackish fuscous above, the lighter
colors being distinctly less extensive than the dark
(which is generally nearly black) and appearing as dots
or roundish spots, and sometimes also as a broad medio-
g 1 . Fore femora of $ at most scarcely more than a
third longer than the pronotum ; outer carina of hind
femora of $ serrulate, not spined.
h l . Hind femora relatively long and slender, three
and three quarters times as long as broad.
1 2. seclusus.
A 2 . Hind femora relatively stout, not over three
and a half times longer than broad.
i l . Hind tibiae but little longer than the femora,
the spurs not longer than the tibial depth, the
hind femora considerably more than twice as
long as the fore femora.
26 PROCEEDINGS OF THE AMERICAN ACADEMY.
j 1 . Hind femora less than three times as long
as broad 13. terrestris.
j 2 . Hind femora three and a half times as
long as broad 14. celatus.
*. Hind tibiae considerably longer than the
femora, the spurs half as long again as the tibial
depth, the hind femora only about .twice as long
as the fore femora 15. brevipes.
g\ Fore femora of $ nearly one half longer than
the pronotum ; outer carina of hind femora of male
spined, not serrulate 16. lapidicola.
/ 2 . The lighter colors which are more massive prevail
above, the darker appearing principally as posterior
bands to the segments and rarely darker than fusco-
castaneous, rarely with a light mediodorsal line.
g 1 . Outer cariua of hind femora of $ armed with
only a few raised points.
h 1 . Hind femora slender, nearly three and a half
times longer than broad . . . 17. arizonensis.
h 2 . Hind femora stout, about two and a half times
longer than broad 18. uniformis.
d 2 . Hind tibia? of < arcuate or sinuous; outer carina of
hind femora of always conspicuously spined.
e 1 . Hind femora very long, four times as long as broad,
the fore femora fully three fourths as long again as the
pronotum 19. lieros.
e 2 . Hind femora relatively short, not more than three and
a half times longer than broad, the fore femora consider-
ably less than half as long again as the pronotum.
f l . Inferior sulcus of hind femora of $ broadening
proximally 20. uhleri.
/ 2 . Inferior sulcus of hind femora of of uniform
width 21. blatchkyi.
6 2 . Fore femora but little if any longer than the pronotum even
in the male ; hind tibiae of male usually straight, but often bowed
e 1 . Dorsal surface of abdomen of smooth and even.
d l . Hind tibiae of arcuate or sinuate in basal half.
e 1 . Hind tibia? of considerably longer than the femora ;
hind tibial spurs usually at least half as long again as the
SCUDDER. NORTH AMERICAN CEUTHOPHILI. 27
f 1 . Hind femora of $ relatively long, three and a half
times as long as broad ; no large spines on outer carina.
f 2 . Hind femora of $ relatively stout, rarely exceeding
three, never three and a quarter, times as long as broad ;
some spines on outer carina as long as the tibial spurs.
ff 1 . Hind tibiae of $ at least a tenth longer than the
A 1 . Hind femora of $ two and a half times longer
than the fore femora; hind tibial spurs only
slightly longer than the tibial depth.
h 2 . Hind femora of $ but little more than twice
as long as the fore femora ; hind tibial spurs nearly
twice as long as the tibial depth . 45. inquinatus.
g 2 . Hind tibiae of less than one tenth longer than
the femora 22. spinosus.
e 2 . Hind tibiae of $ at most scarcely longer than the
femora; hind tibial spurs rarely longer than the tibial
f 1 . Hind femora of $ three or more than three times
as long as broad ; fore femora nearly or quite a fifth
longer than the pronotum.
g 1 . Hind tibiae of $ at most feebly sinuate at base.
g 2 . Hind tibiae of $ very strongly bowed.
f' 1 . Hind femora of $ less than three times as long as
broad ; fore femora only an eighth longer than the pro-
g 1 . Hind tibiae of $ strongly bowed, armed below
with a row of spines mounted on nodules.
g z . Hind tibiae of $ faintly sinuate at base, normally
armed beneath 51. latipes.
cP. Hind tibiae of $ straight throughout.
e 1 . Outer carina of hind femora of armed with prom-
inent conical spines, generally well separated.
f 1 . Hind tibiae of $ less than one tenth longer than
f) 1 . Dorsal surface of body almost uniformly very
dark, almost black, the lighter markings themselves
28 PROCEEDINGS OP THE AMERICAN ACADEMY.
not very light nor extensive, and therefore incon-
A 1 . Hind tibial spurs generally excessively diver-
gent, extending in nearly opposite directions on the
two sides and set at a high angle with the tibia.
h?. Hind tibial spurs rarely exceeding 120 in
divergence, and set at an angle with the tibia not
i\ Smaller species, with pallid sides, luteous
legs, and narrow dorsal stripe, the hind tibiae of
the $ a twelfth as long again as the femora.
i 2 . Larger species, with castaneous sides and legs
and broad dorsal stripe, the hind tibiae of $ not
a thirtieth longer than the femora . 26. salki.
g*. Dorsal surface of body with conspicuously con-
trasted dark and light markings, neither prevailing
over the other 47. pallidus.
y 2 . Hind tibiae of $ a tenth longer than the femora.
e 2 . Outer carina of hind femora of $ more or less deli-
cately serrate or armed with recumbent spines.
f l . Body of male very compact, short subfusiform, not
or hardly more than two and a quarter times as long as
g l . Hind femora of $ relatively stout, considerably
less than three times as long as broad, the hind tibiae
longer than the femora, and the spurs only as long as
the tibial depth 24. nigricans.
g 2 . Hind femora of $ relatively slender, three times
as long as broad, the hind tibiae shorter than the
femora and the spurs nearly half as long again as the
tibial depth .. .'.-.... 25. fusiformis.
/ 2 . Body of $ much more elongated, rarely distinctly
fusiform, over three and generally at least four times