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PROCEEDINGS

OF THE

Washington Academy of Sciences

Vol. X

1908

WASHINGTON
January-December, 1908

I

I

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i
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OFFICERS FOR 1908

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vii

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viu

CONTENTS

PAGE

Studies in the Physiology and Morphology of some^, California
Hepaticag ; by Harry B. Humphrey ..... i

A Revision of the Tenebrionid Subfamily Coniontinae ; by Thos.
L. Casey 51

The SiDerm-Receptacle in the Crayfishes, Cambarus cubensis and
C. paradoxus; by E. A. Andrews . . . . . .167

Notices of Deceased Members . . . . . . .187

Index ........... 245

IX

ILLUSTRATIONS

PLATES

FACING PAGE

I. Ten figures showing structures of Fossotnbronia longiseta and

one of Aneura multifida major 50

II. Figures showing structure of Aneura multifida major, Lepi-
dozia attenuata, Porella bolanderi, Anthoceros pearsoni,
Fegatella conica, Fimbriaria, and Fossombronia longiseta. 50

FIGURES

1-8. Cambarus (^Procambarus) cubensis 176-177

9-12. Cambarus (^Paracatnbarus) paradoxus iSo— 183

PROCEEDINQS

OK THE

WASHINGTON ACADEMY OF SCIENCES

Vol. X, pp. 1-50. Plates I-II. January 18, 1908.

STUDIES IN THE PHYSIOLOGY AND MORPHOL-
OGY OF SOME CALIFORNIA HEPATIC^.

By Harry B. Humphrey, B.S.
Acting Instructor in Botany, Leland Stanford Junior University.

The morphology and physiology of the Hepaticse have been
treated by many authors, but their work has been confined mainly
to the group as a whole. The intimate relations of the liver-
worts to their environments, have however, received only inci-
dental treatment. Ordinarily we are inclined to associate with
the Hepaticae an environment characterized by moisture and
shade. To a certain extent we are justified in doing so, for
the majority of the known species occur in just such a habitat,
many of the larger and more striking ones, such as 3Ionoclca,
Duviortiera and some species of Aneura^ being common in the
more humid regions of the tropics. On the o|:her hand many
species are known to occur normally in parts of the world where
climatic conditions are not so evenly balanced as in the tropics ;
many thrive in extreme northern and southern regions where
they are subject to great variations in temperature, while those
growing in regions like the west coast of the United States
must adapt themselves to prolonged periods of drought alter-
nating with six or seven months of rainy weather.

It was with a view of ascertaining the nature and influence
of these various conditions common to certain California hepat-
icae that the present study was undertaken.

Proc. Wash. Acad. Sci., January, 1908. i

2 HUMPHREY

The author wishes to acknowledge his obligations to Pro-
fessor Douglas H. Campbell and Associate Professor George
J. Peirce under whose direction the work was pursued. Thanks
are also due Professor Alexander W. Evans, Dr. Marshall A.
Howe and Professor Roland Thaxter for assistance in the
determination of material and for the use of certain hepaticas
sent to the writer.

PARASITISM AND SAPROPHYTISM IN HEPATIC^.

The association of certain fungi with hepaticse was first
described in detail by Leitgeb,^ who, in his studies on PtiUdmm
ciltare, observed the infection of young sporogonia. He found
that all such sporogonia were more or less abnormal in their
mode of segmentation and inferred from this that the infected
organs were structurally effected by the action of the fungus.

Following Leitgeb, a number of writers have observed fungus
infection in other hepaticse. As early as 1879, Kny ^ discovered
sterile fungal hyphse in the rhizoids of Lunularia and Mar-
chantia. These, he states, were found to be present in rhizoids
undergoing a process of regeneration, which process may have
been stimulated by the hyphae. Cavers^ has observed that
when Marcliantia and Lunularia grow in ordinary soil free of
humus the rhizoids are penetrated by hyphae which grow up-
ward as delicate filaments showing cross-walls at rather long
intervals. These hyphaj occasionally become branched but
never, so far as he has observed, reach the tissue of the thallus.
On the other hand, he has found that when these plants grow
on humus soil the hyphee extend into the compact tissue of the
thallus, to which in fact, they are largely confined.

Golenkin '* observed the presence of endotrophic mycorrhiza in
Marchantia ^ahnata^ M. paleacea, Pf-eissia commutata^ Tar-

' Leitgeb : Untersuchungen iiber die Lebermoose, Heft 2, p- 58 ; Tafel 3,
Fig. 26.

''Kny and Bottger, 1S79: Ueber eigentliiimliche Durchwachsungen an den
Wurzelhaaren zweier Marchantiaceen. Verhandl. d. bot. Vereins d. Prov.
Brandenburg, p. 2 of Separate.

* Cavers, 1903: Saprophjtism and Mycorrhiza in IlepaticjE. The New Phy-
tologist. Vol. II, No. 2, pp. 32-33.

■•Golenkin, 1902: Die Mycorrhiza ahnlichen Bildungen der Marchanteen.
Flora, Band 90, p. 209.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 3

gionia Jiyfofhylla, Plagiochasma elongatwn and Fegatella
conica. In all these the hyphae are confined to the more com-
pact tissue of the thallus.

Jeffrey,' in his paper on the Gametophyte of Botrychium vir-
giniamini, describes an endophytic fungus associated with this
fern. His observations show that conidia develop singly within
the host cells usually at the end of a hypha. These peculiar
conidia germinate /;/ situ^ generally producing a germ-tube
which forces its way into the neighboring cells of the host.
Within certain cells of the host he observed the development of
vesicular structures along the enclosed hyphse. Similar struc-
tures have been observed by Bruchmann " in his recent studies
on Ophioglossum viilgatwn. In his Fig. 42 he has indicated
fungal structures similar to those common to the endophytic
hyphas found by the author in cells of Aneura mtiltifda major.

The same writer'^ in his work upon Botrychium lunaria
describes an endophytic fungus infecting the cells of the pro-
thallium. He states that the hyph^ usually find entrance to the
prothallial tissue through the rhizoids though in some instances
they are found penetrating the outer, cuticularized surface of
the prothallium. Once within the cells of the host, the hyphas
fill the cells with their sclerotia-like structures and vesicular
enlargements. All starch of the infected cells disappears, but
he finds oil and albumen within the hyphas which he regards as
reserve products to be utilized later by the growing embryo.
Thus he recognizes a symbiotic relationship betw^een fungus and
host, and in another part of the same paper he advances the
theory that the endophyte is a probable means in assisting the
prothallium to withstand the rigors of hot and cold weather.

'Jeffrey, 1S9S : The Gametophyte of Botrychium virginianum. University of
Toronto Studies, No. 1, p. 12.

^Bruchmann, 1904: Ueber das Prothallium und die Keimpflanze von Ophio-
glossum vidgaium L. Botanische Zeitung, Heft XII, Taf. VIII, Fig. 42 und
Fig. 42a.

* Bruchmann, 1906: Ueber das Prothallium und die Sporenpflanze von Botry-
chium lunaria Sw. Flora, Band 96, Heft i, pp. 210-21 1.

4 HUMPHREY

THE PARASITIC FUNGUS ASSOCIATED WITH FOSSOMBRONIA

LONGISETA AUST,

In the writer's recent studies on the Development of Fosso^n-
bronia longiseta ^ no reference was made to the structure of the
infecting fungus ; in fact, the material then in hand was too
scanty to make anything like a satisfactory study. Since the
publication of the above paper, however, excellent material has
been found in various stages of development, and the relation of
fungus to host has been clearly worked out.

So far as could be ascertained, the host is invaded from with-
out by hyphae; that make their entrance through the rhizoids or
directly through the cells of the host stem. The infection of
the rhizoids is very similar to that described by Cavers^ for
Lofhozia, Ce^halozia and other hepaticse. As a rule the
hyphag not only invade the rhizoids but extend throughout the
tissues of the host, regardless of the nature of the substratum,
thus differing somewhat from the nature of infection reported
by Cavers where he has observed that the degree of infec-
tion varies with the amount of humus in the soil. In Fos-
sombronia, as a rule, the hyphge infecting rhizoids alone were
confined to the inner wall of the organ, though in some instances
not a few rhizoids were found in which hyphce had penetrated
through the wall to the exterior as shown in Fig. 3. The host
thus affected was growing in soil containing little humus and
in no case were these external hyphse observed to grow longer
than those figured. The hyphae confined to the rhizoids are
very seldom branched and are rather remotely septate. In cer-
tain parts, however, usually at the end of a branch, short seg-
ments not unlike conidia develop and are abstricted from the
hypha. The subsequent behavior of these has not been
observed. It is, however, quite probable that these structures
furnish one means of transfer of infection. I did not find in
Fossoinbronia anything comparable with the conidia described
by Jeffrey^ as occurring in the endopliyte of Boi7'ychiwm vir-
gintanum.

' Humphrey, 1906 : The Development of Fossombronia longiseta Aust. Annals
of Botany, Vol. XX, No. LXXVII.
^Loc. cit.
'Loc. cit.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 5

In a few instances infected rhizoids were found to have under-
gone considerable modification of form as seen in Fig. 5. On
the other hand very similar modifications were repeatedly ob-
served in uninfected rhizoids. These were very likely due to
the stimulus of contact with compact soil. Whether the fungus
is instrumental in the development of such malformations as the
one shown in Fig. 5 cannot be determined at this writing. It
is, however, hardly probable, for the writer has observed that
when infected plants are transferred to Knop's solution the new
rhizoids though seriously infected maintain a uniform direction
of growth and are apparently unaffected by the hyph£e.

In a previous paper ^ mention was made of the fact that Fos-
sofnbronia plants in many instances developed a tuberous growth
which, on careful examination, was found to contain a more or
less complex growth of fungus hyphae. Microchemical tests
demonstrated the presence of a large quantity of starch, oil and
nitrogenous food products within the cells of this tuberous
growth and this, no doubt, in a measure accounts for the greater
development of the fungus in this region. So far as the author
has observed, none of these tuberous growths is free from infec-
tion. It is hardly probable, however, that this structure is
directly due to the activity of the fungus for it is a structure
common to a number of other hepaticie such as Gcothalliis
tuberosus Campb., Riccia canccUata Tayl., Fossombronia tuber-
ifera Goebl., and a number of others in which no fungus infec-
tion has been observed. Specimens of Geoihallus tuberosus
have been carefully examined by the writer with a view to
ascertaining the presence of infection, but in all material ex-
amined the tubers were uninfected and filled with reserve food
products as described by Campbell,- '95. No doubt the tuber
in the case of Fossombronia longiseta serves the same purpose
as in Geothallus tuberosus or in Fossombronia tubcrifcra^ i. <?.,
as a special structure to carr}'- the plant through a more or less
prolonged dry season. Howe^ has shown that in specimens of

^ Loc. cit.

"Campbell, 1896: A New California Liverwort. Botanical Gazette, Vol. 21,
No. I, p. 12.

^Howe, 1899: The Hepaticse and Anthocerotes of California. Memoirs of
the Torrey Botanical Club, Vol. VII, p. 80.

O HUMPHREY

J^ossombronia longiseta collected in southern California where
the dry season is longer, this tuberous growth is considerably
more marked than in the case of plants in the northern half of
the state. Owing to the earthquake of April i8, 1906, a break
occurred in a water-pipe not far from Stanford University. As
a consequence a considerable tract of ground was well irrigated
throughout the past, very prolonged dry summer. Upon this
thoroughly moistened soil the writer found, growing vigorousl}',
a considerable number of plants of Fossombronia longiseia^
Anthoceros -pearsoni, Targionia hypophylla, and some species
of the more common mosses. Here were growing a number
of plants accustomed to summer desiccation and it occurred to
the writer that under these conditions of increased humidity
certain structural changes might result both in the development
of the gametophyte and sporophyte.

Careful examination of a large number of these plants re-
vealed no evidence of an3'thing in the nature of a tuberous
growth in any part of the thallus though all plants examined
showed fungus infection. It would seem therefore that this
instance would lend some support to the inference that, so far
as our species of Fossombronia is concerned, these small tuber-
like structures are purely adaptive and their development
depends largely upon certain external factors. Whether growth
under similar conditions would result in the reduction or dis-
appearance of the tubers in such plants as Geothallus tubero-
sus or Anthoceros ■phymatodcs is a matter that has not 3'^et been
tested. In these forms the tuber is a well-marked and doubtless
long-established modification of the thallus and if reduced at
all would probably require a considerable period of time during
which conditions of constant moisture are allowed to act.
Peirce,' '06, has demonstrated that such a well-marked char-
acter as the dorsiventrality of certain liverwort gametophytes is
not a hereditary character as commonl}^ supposed but is pri-
marily due to the formative influence of light.

As elsewhere stated, some plants of Fossombronia longiseta
were studied in which the rhizoids were apparently free from

' Peirce, 1906 : Studies of Irritability in Plants. Annals of Botany, \o\. XX,
No. LXXX, p. 459.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 7

hyphai, though the more compact tissue of the thallus was in
many such instances thorough!}^ infected. Thus it would seem
that infection need not necessarily take place through the rhi-
zoids. Material carefully fixed in chromic acid (i per cent.)
and stained with the Fleming's triple combination seemed to
give very satisfactory results, though it was found necessary to
allow more time to the action of the stain than was customary
in staining organs or structures of the host. Sections through
the stem of infected plants almost always showed a well-de-
fined zone of cells more or less given over to hyphai. This is
not unlike the condition observed by Cavers ' ('03) in Fcgatella
and by Golenkin" ('02) in a number of forms cited above.
Golenkin states that the cells of this zone, though still retaining
their protoplasm and nuclei, are void of chlorophyll and starch.
Careful examination of infected cells of Fossoinbronia^ except
where infection was so far advanced that the cells were occu-
pied by developing sclerotia, revealed the presence of varying
amounts of chlorophyll and starch, depending upon the number
and vigor of invading hyphee.

Unlike the forms described by Golenkin, this fungus is not
necessarily confined to any given zone, for any portion of the
thallus may be invaded by hyphas extending out from the more
compact parts. Though several hundred fruiting plants have
been studied, no evidence of infection of either antheridium or
archegonium has yet been seen, and the sporophyte is appar-
ently free from the attack by fungus.

In material fixed in January, 1904, about three months after
the beginning of the rainy season, almost the entire tissue of the
stem was packed with hyphag, many branches extending through
the outer layer of cells of the host into the substratum.

Although the fungus was present in virtually the whole of
the stem-tissue, strictly speaking, it was found that at this stage
of its development the hyphas were more or less confined to the
innermost cells of the stem while those in proximity to the
growing point as well as the leaves were apparently free from
infection. The epidermal cells and those immediately beneath

* Loc. cit., p. 33.

* Loc. cit., p. 209.

8 HUMPHREY

showed evidences of comparatively few hyphee which evidently
served to connect the more functionally active interior system
with those in the rhizoids and with the substratum. Thus far
the habit of growth of this fungus agrees quite consistently with
those described by Cavers.^

At about this period in its development one may observe, scat-
tered here and there throughout the infected zone, vesicles of
considerable size and presenting a variety of form. These ap-
pear as great enlargements of the hyphae and are full of rather
coarsely granular protoplasm (Fig. 6). This protoplasm has
within it minute nuclei, at first rather few in number but as the
vesicles become older and larger increasing in number until
they quite fill the interior of the vesicle. It was further found
that in some instances at least these vesicles are connected with
the exterior by hyphge (Fig. 8) and along the course of these
hyphae may be seen certain of these nuclear bodies. Whether
or not they make their escape to the exterior as zoospores could
not be demonstrated, but their development and subsequent be-
havior is suggestive. In his recent work upon the development of
Ophioglossmn pendulum and other members of the genus,
Campbell ^ has demonstrated the presence of thin-walled vesi-
cles in the endophyte. In these the behavior of the nuclei
resembles that occurring in the vesicles of the endophytic fun-
gus found in the Fossonibronia. In Fossombronia these vesicles
completely fill the host cells in which they occur, in some cases
causing a considerable distention of the cell.

As stated above, the amount of chlorophyll or of starch and
other products of metabolism varies with the degree of infection.
Certain cells of the host seem packed with hyphae, and a micro-
chemical test fails to reveal any of the products of the normal
cell, and while a large part of the stem appears to be dying,
the leaves and the uninfected stem tissue seem to be vigorous
and unaffected by the presence of the fungus. Comparison of
infected plants with those entirely or nearly free from fungus in-
vasion seems to demonstrate that up to the vesicle stage of devel-
opment the growth of the host is not abnormally stimulated, in

^Loc. cit., p. 32-33.
' Campbell, 1907.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATICvE 9

fact an extensive examination of plants of both sorts leads one
to conclude that the presence of the fungus is of no advantage
to the host and in many instances is an evident detriment. In
March, 1906, while making some field studies upon I^ossotn-
bronia, the writer observed that certain plants appeared to be
dying. The affected plants were in some instances almost
white, apparently from loss of chlorophyll. Closer examina-
tion with a hand-lens revealed the presence of a great number
of minute black bodies along the length of the stem and in the
leaves. These were especially abundant near the base of the
leaf. On further examination with the microscope it was found
that these black bodies were sclerotia within the cells of the host
and the development of these resulted ultimately in the death of
the host. It was at first thought probable that this fungus was
distinct from the one found invading the rhizoids and stem tissue,
but a careful study seems to show that the two are identical. At
least so far as concerns the host the development of these scler-
otia marks the period of greatest activity of the fungus, and its
truly parasitic nature is manifested. Up to the appearance of
the sclerotia the life of the host is not seriously affected but as
soon as the hyphae begin to extend and to form the structures
that later develop sclerotia a very evident drain upon the vitality
of the host sets in and increases with the further development
of these structures.

It was at first thought that these might be perithecia of some
ascomycete but microtome sections demonstrated their true
nature. Wherever they occur the cells containing them become
considerably distended and completely filled. Their presence
causes no abnormal development of the cell as regards thickness
of wall or form, though the cell-contents become much modified
to the extent that all vestige of an}' starch, oil, cytoplasm, etc.,
has disappeared, even the nucleus is absorbed and the entire
cell cavity is filled by the sclerotium.

The behavior of the hypha; with reference to the nucleus and
chromatophores as described by Cavers ^ for the fungus infect-
ing Monoclca does not occur in the case of the fungus under

^Cavers, 1904: Contributions to the Biology of the Hepaticse. Part i.
Targionia, Reboulia, Preissia, Mo7toclea, p. 39.

lO HUMPHREY

consideration. In Monoclea the fungus is confined to a sharply
defined mycorrhizal zone three or four cell layers in thickness.
Many of the cells in this zone are filled with branched hyphae,
tufts of which seem to envelop the nucleus. In some cases
even the chromatophores, like the nucleus, become surrounded
by similar tufts of branching hyphai in a manner quite sug-
gesting the formation of a lichen.

In Fossomhronia the hyphse preceding the development of a
sclerotium are but sparsely ramified. The first indication of the
sclerotium is the increase in diameter of the hyphas within a cell
followed later by the profuse development of short, thick
anastomosing branches between which may at first be seen
spaces of varying size which ultimately disappear as the
sclerotium increases in extent (Fig. lo). The nucleus and
chromatophores at first visible finally disappear, doubtless being
taken up by the actively growing fungus. As yet the writer
has failed to observe how these structures are consumed.

According to Czapek ^ the tissues of Marchantia, Fegatella^
Luniilaria and other hepaticae contain an antiseptic principle
which he calls " sphagnol " because of its abundance in the
peat-mosses. He has shown that this substance exists in com-
bination with the cellulose of the cell-walls and exerts an
inhibitive influence upon the development of moulds and
bacteria. This, Cavers ^ thinks, suggests the view that in the
case of certain Fusarium-like fungi the " sphagnol " may serve
to regulate the growth of the fungus and prevent symbiosis
from passing into parasitism. In the case of Fossonib^'onia it
cannot be shown that at any time during a period of infection the
invading fungus maintains a symbiotic relation with reference
to the host, for an examination of infected cells shows that the
presence of hyphas sets up a disturbance of the cell-metabolism,
the cell becoming impaired to such an extent as ultimately to
cause its death. The degree of this impairment varies with
the activity and extent of the parasite. To be sure, many
infected plants appear to be quite as healthy and fertile as

'Czapek, 1SS9: Zur Chemie der Zell membranen bei den Laub. und Leber-
moosen. Flora, Band 86, p. 361.
^Loc. cit., p. 33.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ II

Others yet free from the fungus, but it is owing to the fact that
the degree of infection is but slight and the vigor of the host
is sufficient to throw off for a time the harmful effects of the
parasite just as the leaves of cultivated lettuce maintain apparent
full vigor during early stages of the infection by Bremta
lactiica}

Experiments designed to ascertain the effect of the fungus
upon very young plants of either Fossomb}-onia or Fimbriai'ia
have thus far failed, and it cannot be decisively stated here that
Fimbriaria is at all susceptible to infection by this fungus. The
examination of a large number of plants of all stages of develop-
ment has shown our common species, F. californica, compara-
tively free from infection b}'- any fungus. Recently, however,
the author has found a few plants of this species attacked by a
fungus which seems, in material thus far examined, to be con-
fined entirely to the cells of the first four layers on the ventral
side of the thallus. H3^ph£e in considerable numbers were ob-
served in both the smooth and tuberculate rhizoids. These
showed but slight tendency toward branching, in many instances
extending the entire length of a rhizoid without producing a
branch. The hyphas confined to the rhizoids, compared with
those within the mycorrhizal zone of the thallus, are very much
more delicate ; but upon extending into the cells of the thallus
they branch profusely and develop relatively thick and tough
walls. The fungus resembles the one described above for Fos-
sonibrotiia, in that it is filamentous and septate. Structures com-
parable to vesicles, conidia or sclerotia have not yet been
observed. The presence of the fungus seems in no wise to re-
tard the growth or affect the vigor of the thallus and no plants
have yet been found in which the hyphae have penetrated the
chlorophyll-bearing tissue.

THE PARASITIC FUNGUS ASSOCIATED WITH ANEURA
MULTIFIDA MAJOR.

While in the field collecting Aneiira the writer observed that
a considerable number of plants had taken on a rusty brown

'An illuminating discussion of symbiosis and parasitism is to be found in
Plant Physiology. Peirce. Pp. 85-92 inclusive.

12 HUMPHREY

color. Upon careful examination it was found that they were
in a dying condition owing to the presence of a fungus. A
large quantity of material consisting of plants in a normal con-
dition as well as those visibly affected by the fungus were
brought into the laboratory in September, 1906, where the rela-
tion of fungus to host could be more carefully studied. This
species of Aneura is found growing on moist surfaces of rocks
along streams, on decaying logs and moist banks of soil in
which there is considerable clay. Material collected from all
three of these sources contained a large number of infected
plants so it is hardly probable that the fungus is one confined
to plants growing on decaying wood.

Aneura multijida major ^ as well as other species of this genus,
under certain conditions produce two-celled gemmse in great
numbers. It was found that many of these after a short time
had germinated and some were infected. Figure 15 shows a
young plant resulting from the germination of a gemma, into the
older cells of which a fungus had penetrated by well-marked
haustoria. The diseased cells contained less chlorophyll and
showed unmistakable evidences of the harmful effect of the
fungus. Young plants of varying age and size were found to
be infected. It was first thought likely that infection took place
directly while the gemma was still within its mother cell, but a
very careful examination of gemmae failed to support any such
view. It seems that young plants of less than four cells are
rarely infected. The fungus develops conidia freely and it is
probable that infection is brought about by their germination.
A small number of young plants developing from germinating
spores were observed, but as yet none of these has shown the
presence of a fungus.

In the older plants, hyphse from within the cells of the thallus
grow downward extending into the rhizoids, though this is by
no means so common as in the case of Fossombronia or Fim-
brlaria. Many rhizoids whether infected or not present a
strong tendency to branch as shown in Fig. 16, a and h. This
is undoubtedly due to the influence of contact stimulus ; in fact,
rhizoids not in contact with the substratum are all quite simple
and more delicate.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 13

In his studies upon Lofhocolea hidentata. Cavers ' found that
the tissues of the gametophyte are entirely free from hyphae,
but the rhizoids which grow out in tufts from the bases of the
amphigastria penetrate the substratum of rotten wood and there
become profusely branched like the haustoria of many fungi.
This, he considers, enables the liverwort to assume a more or
less saprophytic existence. He has not however shown that the
plant actually does adapt itself to this mode of life, nor does it
appear that we can assume the branching of the rhizoids as due
to anything other than contact stimulus. Peirce and Randolph ^
have demonstrated that in the case of certain attached fresh-
water algae and in many marine forms the development of a
holdfast is directly and wholly the result of contact stimulus.
The complexity and extent of the holdfast were found to vary
with the degree of roughness of the surface of the substratum ;
young plants grown on ground glass developing much more
elaborate holdfasts than similar plants grown on smooth glass
while those grown in dust-free water developed no holdfasts at all.

In Knop's solution the writer now has plants of Fossomhronia^
Crypioviiirtuin and other liverworts growing that were removed
from a normal soil substratum some months ago. Still attached
to these plants are some of the old rhizoids which are all more
or less gnarled and modified, some exhibiting short lateral proc-
esses. Since placing the plants in Knop's solution a great
number of rhizoids have developed and in every instance they
are perfectly straight and unmodified and much more delicate
in structure than those that had grown while the plants were in
their normal habitat. We have here exactly the same behavior
exhibited by the secondary roots and root-hairs of higher plants.
If, as some botanists maintain, the rhizoids of bryophytes are
simply organs of attachment is it not probable that, as in the
case of certain algee, these branches are due to the stimulus
afforded by contact?

Aside from Aneura and Fossonibronia the writer has observed
branching rhizoids in Ccphalozia biaispidata, due probably to

' Loc. cit., p. 32.

^Peirce and Randolph, 1905: Studies of Irritability in Algae. Botanical
Gazette, 40- pp. 321-350.

14 HUMPHREY

the same stimulus, a;? they were not infected by hyphas. The
fungus associated with Aneiira differs from that attacking Fos-
sojnbroiij'a both as to development and habit of growth. In the
case of the Fossomhronia fungus we find it to be more truly
endophytic, once infection has been brought about, while the
fungus associated with Aneu7'a seems to be epiphytic in habit,
developing haustoria-like branches which in turn develop other
branches penetrating and drawing sustenance from the host
cells. The hyphas are septate and profusely branched, especially
within the tissues of the host. The hyphee within the rhizoids
are in nearly every instance unbranched and in no case were
they seen to grow through the wall of the rhizoid communicating
with the exterior as in Fosso-jubronia.

The physiological effect of the fungus upon the host is quite
as marked as any we have yet studied. The cell walls of
Aneuj'a are very much thicker than those of most of our liver-
worts, and the plants are vigorous in habit. Each cell contains
usually one large oil body and numerous chromatophores.
Fungal hyphae penetrate the cell walls without any resulting
modification and in some instances a single branch may pass
through several cells without producing haustoria or branches
of any sort. Ordinarily, however, after entrance to the host
tissues is effected, the fungus becomes quite extensive, as seen
in Fig. 13, where three affected cells are represented. In these
it will be observed that the large oil body has already disap-
peared and the number of chromatophores is somewhat below
the normal (compare Fig. 12 with Fig. 14). Ultimately the cells
become in some instances packed with hyphje and at this stage
the death of the cell rapidly ensues. Such cells when micro-
chemically tested are void of starch, no nucleus or cytoplasm
can be made out, and the chromatophores have undergone
complete disorganization ; in fact, little remains but the cell wall.
In certain plants where the fungus had reached an advanced stage
the hj^phas within certain cells had developed into knot-like
structures suggesting the beginning of sclerotia ; and in a few
instances these had, when sufficiently nourished, developed into
blackish thick-walled sclerotia completely filling and distending
the cell cavity. These, in appearance and structure, very much

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 1 5

resemble the sclerotia formed in the cells of Fosso')nhronia^
though they do not develop in such great numbers. This is
doubtless due to the fact that the comparatively early decline
and death of the host tissues impairs the vigor and health
of the parasite before these structures can develop in great
numbers.

As soon as the host cells begin to show the effects of para-
sitism there develop upon the surface of the host at indefinite
points along the surface-hyphas, perithecia-like structures (Fig.
11) the real function of which, however, has not been demon-
strated. Professor Roland Thaxter kindly attempted to iden-
tify the fungus for me but being unable to determine the true
nature of these structures was forced to wait for more advanced
stages. The writer has prepared a number of slides of micro-
tome sections which show these bodies to be hollow.

Beyond a wall no other structures have yet been observed in
connection with the interior of these somewhat globular bodies.
They seem to develop from short processes that appear here and
there along the external hyphee. They assume very early the
spherical habit and develop at various points on their surface
short spine-like processes at first pointing in different directions
but ultimately either disappearing or lying all in one plane at
the base of the structure bearing them. On certain host plants
the tissues of which have already begun to turn brown, these
perithecia-like bodies develop in such numbers as to be almost
in contact, and the complex of hypha^ on the surface and within
the host is very elaborate. Two-celled conidia were found on
the surface of the host but it could not be proved that these
spores belonged to the fungus infecting Aneura, although no
other fungus was observed in the material.

It is the intention of the writer to work out the development
and life history of this fungus and the one associated with I^os-
sonibronia more fully in a later paper. At the present moment
the main object is to discuss the relation of parasite and host.
In the case of this fungus there is less evidence of anything
comparable to a symbiotic relation than in the relation of the
Fossoinbro7iia parasite to its host. In fact, as soon as infection
is brought about, the host seems to show signs of resulting

I 6 HUMPHREY

injury. The infection spreads from plant to plant quite rapidly.
In a dinner-plate full of material, at first the majority of plants
examined were unaccompanied by fungus, but within three
weeks, on one side of the plate all the plants were turning brown ;
on examining these they were found to be infected. The fun-
gus seems usually to make its attack through the ventral side
of the thallus, though occasionally a plant would be found with
hyphas on both sides. Microtome sections failed to reveal any
vesicular structures within the host.

Anetira, being naturally less compact and vigorous than such
forms as Fegatalla or Lwiularia, furnishes a poorer field for a
parasitic fungus and seems less able to cope with the fungus.
This, however, may be partly due to greater vigor of the par-
asite. It must also be noted that infected plants behave dif-
ferently under varying conditions. Aneitra tnultifida major
as we find it here is semi-aquatic as to habitat and when grow-
ing under perfectly normal conditions may be considerably in-
fected without showing any effect other than becoming yellow-
ish green in color. If, however, these same plants are brought
into the laboratory, even though well supplied with moisture,
under a bell-jar they become brown in color finally dying,
while uninfected plants seem to thrive quite as well as when
growing out-of-doors under normal conditions. Plants grow-
ing along streams, though receiving little less than the required
amount of water, if infected, seem less able to throw off the
effects of the fungus than plants growing under perfectly
normal conditions of moisture, light, etc.

Aside from the fungi found in association with Fossombronia,
Fimhj-iaria and Aneura, the writer has had occasion to study
infected material of Anthoceros ^earsoni and Porella holanderi.
In habit of growth the fungus associated with Anthoco'os is
very much like that infecting Aneura though none of the
material examined showed any structures suggestive of fruiting
organs other than conidia which were being formed in consid-
erable numbers by abstriction along the external hyphjE. They
branch freely and are seen to anastomose not at single points
here and there but continuously for some length, both hyphae
being rather minutely septate along the length of contact. The

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATICyE 1 7

hyphae produce short lateral processes which grow in length
until they come in contact with a similar branch of a neighbor-
ing hypha effecting conjugation with it ; or in case it fails to
meet with such a branch it is seen to anastomose with a near-
by hypha and in this manner a complicated weft of hyphae may
develop over the surface of the host. The same seems to be
true with regard to the hypha; within the host cells, though to
less extent. Within certain cells (Fig. 19) dark colored scle-
rotia-like structures develop, though in the material studied
these were by no means abundant. With a limited amount of
infected material nothing definite can be said as to the relation
of the fungus to the host. All plants examined however seemed
practically unaffected by the fungus. Infected cells exhibited
the single chromatophore in every case except where sclerotia
had formed. Just what would have been the effect had the
fungus been as far advanced as in the case of Fossomhronia or
Aneura is mere conjecture. Our species of Anthoccros have,
growing within the thallus, colonies of a species of Nostoc^ but
the writer saw no evidence of fungal hyphas invading these.

Porclla holandcri is infected by a fungus of somewhat dif-
ferent habit, though its ultimate effect upon the host is not
greatly different from what has been described for Fossombronia
and Aneiira.

The principal hyphal trunks appear for the most part to be
intercellular, thus differing in this respect from the others
described above. A single hypha finding entrance to a leaf for
example, will ramify to some extent, and these various primary
branches grow along between the cells of the host, thus forming
an intercellular complex that may effect nearly every cell.
From these primary branches, haustoria find entrance to the
host cells and here they become more or less branched, with the
result that the cells lose all their contents except the nucleus ;
this seems to persist. After the disorganization and disappear-
ance of the chromatophores, the fungus structures are easily
visible and in many of these cells may be seen exceedingly
thick-walled chlamydospores, in some instances quite filling the
enclosing wall of the host cell (Fig. 17).

The fungus is not confined wholly to the interior of the tissue.

Proc. Wash. Acad. Sci., January, iqoS.

l8 HUMPHREY

In many cases a complex of hyphge over-runs the surface of a
leaf and it is not uncommon to find chlam3'dospores in connection
with these external hyphae. One-celled conidia are abstricted
in large numbers from branches of the external hyphae. Sclerotia
have not yet been found, though occasionally a cell may be seen
well filled with knots of hyphse, too loose in structure, however,
to be regarded as sclerotia. Cells in which these knots occur
are always dead and apparently empty.

The fungus behaves toward the host as a parasite. Its hyphae
produce haustoria in great numbers which in the course of time
invade all the cells of a leaf, thus impairing greatly its ability to
elaborate food materials. The younger leaves of the host are
apparently uninfected, and even though the older parts of the
thallus may be dead, growth continues at the growing point and
the plant may grow indefinitely, though very evidently diseased.

The writer has not thus far observed a fungus associated with
our species of Sfhmrocarfus or with Tai-gio7iia hypophylla,
although Golenkin ^ reports the existence of an endotrophic
mycorrhiza for the latter, and it may be that further investigation
will demonstrate the presence of a fungus in Targwnia as we
find it here. As a matter of fact it is not improbable that all
the hepaticse of this region may on further study prove to be
infected, the fungus being either a symbiont or, what is more
likely, a parasite. The writer has seen no evidence in favor of
the view that the fungus stimulates the growth or in any way
benefits the host, though the presence of hyphae may lead up to
the development of abnormal structures such as modified root-
hairs, enlarged cells, etc.

Bruchmann,^in his studies on Botrychium hinarz'a, concludes
as follows : " Der physiologische Nutzen des Zusammenlebens
von Prothallium und Pilz scheint mir auch hier nicht darin zu
bestehen, dass die w^enigen .und unscheinbaren Pilzfaden,
welche aus dem Substrat des Gamophyten in ihn emmiinden,
belanglose Humusstoffe ihm zufiihren, Sondern nur darin,
dass die von den functionsfahigen Rhizoiden der wachsenden
Schutelpartie aufgelosten und herbeigefiihrten Baustoffe mit-

^Loc. cit., p. 209.
^ Loc. cit.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 1 9

telst des Stoffwechsels des Endophyten in haltbare Reservestoffe
umgevvandelt und aufgespeichert werden, von denen namentlich
die auffallend grosse Oelmenge dem Prothallium die Fiihigkeit
verleiht, in dem sandigen Boden auch wahrend der Sommerhitze
und der Winterkalte von Austrocknung bewahrt zu bleiben."

As already stated, the author has found, in the case of all
infected forms thus far investigated, oil and of necessity
albumen, in the living cells of the gametophyte. In F'ossojii-
bronia and Anctira at a certain stage in the development of the
endophyte there is a minimum quantity of these products within
the host cells containing hyphte and a maximum amount of
these same products within the cells of the fungus. The result
is the death of the host or, at least, a very serious check to its
development, resulting in small sickly plants. In such a case
we cannot possibly regard the food products derived from the
gametophyte as reserve materials later to be given up by the
fungus as nourishment to further growth of the host.

The only forms examined not showing harmful effects from
association with a fungus v^^ere Fimbriai-ia^ Fegatella and
Anthoceros. Of the first and the last, only a very limited
amount of infected material was found, consequently little light
can be thrown upon actual relation of host to endophyte. Howe '
has reported an endophytic fungus associated with Anthoceros
olneyi Kxisi.^ the septate hyphte of which produce at the ends
of lateral branches, globular clusters within w^hich are produced
numerous dark spore-like cells bearing some resemblance to
those of the Tilletiacege. He has found the same parasite in
A. ravenellii A.\2L. (Mohr.) and the same or one very similar in
A, hallii Aus. He regards this fungus as a parasite, though no
detailed account is given relative to the character of the asso-
ciation of the two.

DEVELOPMENT OF FRUITING ORGANS OF FEGATELLA CONICA

IN CALIFORNIA.

Fegatella conica [Conocephaluvi contctmi) is described by

Howe^ as growing in moist, deeply shaded places, especially

^ Howe, 1898 : The Anthocerotaceae of North America, Bulletin of the Torrey
Botanical Club, Vol. 25, No. i.
^ Loc. cit., p. 58.

20 HUMPHREY

on Stones and rocks beside streams. It is not common in Cali-
fornia and was first collected by Professor Underwood at Fel-
ton, Santa Cruz County, in 1888. In July, 1906, it was found
in fruiting condition by the author on a trip up San Jose Cafion
in the Santa Lucia Mountains in Monterey County. So far as
is known, plants bearing receptacles had not been found in Cali-
fornia prior to the above named date, though fruiting plants
have been collected by C. V. Piper near Seattle, Wash. Con-
ditions during the rainy season there and in the parts of Cali-
fornia where Fcgatella thrives are similar. The dry season near
Puget Sound is occasionally interrupted by summer showers
which may in combination with other factors influence the
development of fruiting organs.

Fegatclla may be found in many of our near-by canons ; in
fact, it has been observed to occur along the banks of creeks
well down upon the plain of the Santa Clara valley, but almost
always it will be found in particularly well watered and densely
shaded localities, a combination of conditions very essential to
the normal development of the plant in this region of long, dry
summers.

Antheridial plants bearing receptacles have been collected
from several localities, though these plants are by no means
common. Normally, they are of such vigorous, robust habit as
to spread over a considerable area by mere vegetative growth,
and it is not uncommon to find several square feet entirely cov-
ered by Fegatella to the exclusion of everything else.

On finding female plants in fertile condition a field study was
made relative to their habitat, and conditions affecting growth
and development of fruiting organs. San Jose Canon is a deep,
narrow gorge, the floor of which is well watered by a never-
failing stream. The north wall of the canon consists very
largely of precipitous rock ledges that in many places come
abruptly to the stream's edge and are only sparsely clothed with
Artemcsia, Adenostoma and a few other plants of xerophytic
habit. The south wall rises less abruptly into hills averag-
ing seven to eight hundred feet in height. These have a
covering of rich humus and the vegetation at the base along
the creek and well up on the south slope consists of Sequoia

PHYSIOLOGY AND MORPHOLOGY OF CALH-'ORNIA HEPATIC^ 21

sempervt'rcns, Almis rhomhifolia^ Salix lasiolep/'s and other
trees which afford abundant sheUer for a variety of shade-loving
plants. Here also thrive several species of mosses and liver-
worts that in less favored localities would either die or pass into
a dormant, air-dry condition.

Fegatella occurs throughout the length of the canon on both
banks of the creek, though most abundantly on the south side
where there is a minimum of direct sunlight and evaporation.
Five trips were made during tiie past season for at least a dis-
tance of three miles toward the head of the canon and each time
a diligent search was made for fruiting plants which were found
in but few, very limited areas, where the exposure was north or
northeast. It was observed that no fruiting plants were to be
found in places void of sunlight, and the same was true when
the plants grew within six or eight inches of the water's surface
in light of the same intensity as fell upon nearby fruiting plants.
Plants bearing receptacles in greatest number were found on a
flat elevation about eighteen inches above the water in a break
between trees through which fell a flood of strong, diffused
light and at midday direct sunlight. This level bed of Fega-
tella was bathed constantly by seepage of spring-water from a
moss and fern grown cliff forming the background, upon which
were found a few fruiting plants of Fegatella and Aneu7'a -pln-
giiis. Wherever fruiting plants occurred, it was noted that the
combined conditions of light and moisture were relatively about
the same. In places where there was strong light but insuffi-
cient moisture the plants were observed to wither, and when
these conditions were reversed there was always a strikingly
vigorous vegetative development, some individual plants meas-
uring ten or more inches in length.

The lime of fertilization of our California plants is not defi-
nitely known. The antherozoids are discharged during March
and April and receptacles still containing intact antheridia have
been found in some of our canons as late as August.

In England, according to Cavers,^ the receptacles begin to
develop in early spring but do not mature until about the end of
June, while the earliest stages in the development of the sporo-

' Loc. cit., p. 98.

2 2 HUMPHREY

gonium are observed about the middle of July. Owing to the
very slow development of the sporogonium the spores are not
discharged until the following spring. Here in California our
plants behave differently. After fertilization the female recep-
tacles grow to about two to two and one half millimeters in diam-
eter. Though the plants may be amply supplied with moisture
from below throughout the summer, proper humidity and tem-
perature conditions do not prevail until the beginning of the
winter or rainy season. Throughout the summer the growth of
the receptacle is so slight as to be almost imperceptible. San
Jose Creek empties into Carmel Bay and in late summer and
autumn, the canon, being near the sea, is quite commonly filled
with heavy fogs for a part of the day, leaving all vegetation
dripping wet. While these fogs are not equivalent to the
drenching rains of winter, they do play a great role in the revi-
val of certain mosses, liverworts and lichens and doubtless stimu-
late their growth to a certain extent. At any rate, up to the
latter part of August the archegonial receptacle of FegateJla
increases but slightly in size, though the increase is sufficient to
enable one to distinguish fertilized from unfertilized receptacles.
By the first of October (before any rain had fallen) the sporo-
gonium had advanced to the spore tetrad stage and remained in
about this condition until the opening of the rainy season. Fol-
lowing this, rapid development ensues and the spores are dis-
charged in January.

This liverwort might be considered as viviparous, since the
spores germinate within the capsule. Cavers states that ger-
mination occurs within the capsule before the spores are dis-
charged, but does not say how long a time elapses between
the first evidences of germination, and the ultimate elongation
of the peduncle and dehiscence of the capsule. With our
plants germination-stages were found at least a month preced-
ing the liberation of the spores. How much earlier than this
germination actually begins cannot be said, as material for
examination was not available at that time. Germination does
not take place simultaneously in all the spores of a single cap-
sule. One may find within a capsule every stage from
the ungerminated spore to the six-cell stage, and it is to be

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 23

noted that a certain percentage of spores do not germinate at
all, but may be seen in a collapsed and shrunken condition
within the capsule.

Most of the plants examined were infected by a mycorrhizal
fungus that agrees closely in structure and habit with the one
described by Cavers. Our observations confirm this as to the
presence of starch within infected cells which is contrary to the
view of Golenkin/ who investigated a number of infected
hepaticas, including Fegatella, and states that infected cells
contain no starch or chloroplasts.

Our material was found growing on sandy soil containing
considerable humus and it was found that but few rhizoids
contained hyphae. No plants were observed in which the
ventral tissue of the midrib was infected to any extent as
shown by Cavers in plants collected near the sewer-like drains
of tanneries and similar works. All infected plants were in
vigorous, healthy, growing condition and so far as could be
ascertained, the presence of a fungus entails no such disastrous
effects as one may observe in more delicate forms such as
Fossombronia or Anctira.

ADAPTATION OF CERTAIN LIVERWORTS TO THE DRY SEASON.

It is commonly assumed that bryophytes are, in general, mois-
ture-loving plants. While this is to a great extent true it may
easily be shown that there are many important exceptions.
Among the mosses we have only to note such forms as Grim-
tnia, Iledwegia, AndrecBa and many others common to different
parts of the world growing on exposed rock surfaces, with abil-
ity to revive promptly and resume growth after the first rain.
Likewise many foliose Jungermanniaceas, common in the east-
ern part of the United States, are capable of withstanding
drought periods of considerable length, lying practically dor-
mant throufjhout the rainless interval. To what extent eastern
thallose Jungermanniacese are capable of withstanding drought
is a question, that so far as the writer is aware, has not yet been
investigated.

In this section of California, and more particularly the low-

' Loc. cit., pp. 209-220.

24

HUMPHREY

lying country about San Francisco Bay there is an interesting
hepatic flora consisting of types of most of the principal groups.
All of these forms fruit luxuriantly and are seldom found sterile
during any one growing season.

Among the more common hepaticae of this region, and espe-
cially in the vicinity of Stanford University, are certain species
of Riccia^ including R. glauca and R. t7-tchoca7'pa, both of
which grow vigorously in localities of extreme exposure ; and
especially is this true of the latter, a liverwort structurally
adapted to such a habitat. Aside from several species of Riccia
two species of Fhyibriaria [Asterella), F. califor^iica and F.
violacea are met with, the former being quite common. In cer-
tain localities, especially along the high banks of arroyos, Cryf-
tomitriimi teneriim is found to be abundant. Along with it and
in places of severer exposure, occur formations consisting almost
exclusively of the highly resistant Targionia hypophylla.

Along the banks of our coast-range streams, even where
these come down into the Santa Clara valley, occur such widel}'
distributed forms as Fegatella conica and Ma7-chantia foly-
mo7'pha. These however always grow in close proximity to
the water where throughout the year they may never be entirely
deprived of a supply of moisture. It is doubtful if these forms
would be capable of withstanding the effects of a dry season.
Three years ago the writer transferred several plants of Fega-
tella to a locality in the valley where conditions of light and
substratum were restored as nearly as could be done, at the same
time modifying and finally cutting off altogether the supply of
moisture. Bv lessening for a whole summer season the supply
of moisture it was found that growth of the thallus was greatly
checked and the resulting branches were much shorter. Plants
that were deprived of water dried up and died, all ability to
recover being lost as soon as the tissue had given up a certain
amount of the water of constitution. It cannot here be said to
what extent Marchantla polymorj^ha is capable of withstanding
drought, but since it is almost never found except where there
is a fairly constant supply of moisture it would probably endure
desiccation little better than Feg-atella conica. Other moisture-
lovinor liverworts found in moist shaded canons of the outer

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNLV HEPATIC^ 25

coast ranges are two species of Aneura {Riccardia), A. mtdtljida
major and A. pi'ngnis, the latter being far more sensitive to
change of environment than the former, but neither is capable
of withstanding much loss of water. They are both found
growing at their best when, if not actually in running water, they
are at least so situated as to receive a constant supply. The
finest, most vigorous specimens I have ever collected have been
found growing in such a position as to receive constant dripping
of fresh water from above, which so far as has been observed
does not materially influence the development of fruiting organs
as is the case when Fegalella is too abundantly supplied.

Several specimens of Riccioca^'jbiis natans found growing on
soil about a small lake in the foot-hills were brought into the
laboratory along with enough of the substratum to prevent any
shock resulting from transference. These along with some
Aneura mtiUifida major were placed in a shallow earthern
saucer and exposed to the dry air of the laboratory until in a
normal air-dry condition. In this condition they were gradually
moistened but no sign of recovery was apparent in any part of
the tissue of either plant, and after several days of normal mois-
ture supply they showed no signs of life. It is apparent from
this and several other experiments upon hygrophilous forms
that they lack the degree of adaptability to changed climatic
conditions exhibited by such forms as Riccia^ Targionia or
Porella.

Aside from several genera of Ricciaceae and Marchantiacese,
two species of Sfhcerocarfiis and one of Fossombronia, F,
longiseta, represent the thallose Jungermanniace^ for the
region about San Francisco Bay, as well as higher up into the
hills and mountains. So far as habit is concerned Sp/icero-
carpus is hardly to be compared to Fossombronia for the one is
an annual while the other remains alive throughout the entire
summer. We have here a relatively small representation of
leafy liverworts, the most common being Porella bolauderi,
and Fridlania bolauderi, while in certain isolated localities one
may find Radiila coniplanata growing on tree trunks and
Lophozia veniricosa forming reddish-brown patches on exposed
hillsides that receive abundant moisture during the rainy season.

26 HUMPHREY

The AnthocerotacejE are represented by Anthoceros fiisi-
JbrmiSy A. -pearsoni and A. ■phymatodes the first two being
very common, while the third, though not so widely distributed,
is very abundant in certain localities near Stanford University.
All three species are found growing luxuriantly in places of
severe exposure, though thriving best in slightly shaded locali-
ties. At one time it was supposed that A. fusiformis and A.
■pearsoni were annuals but they were afterwards shown by
Campbell ^ to be perennial in habit. So far as the writer has
observed Anthoceros phymatodes survives through the agency
of its conspicuous tubers. In fact, it seems that virtually all of
the thallus dies during the dry season except these structures
which are packed with food-materials and serve as very effec-
tive water-storage organs. The other two species, while surviv-
ing the dry season, resume development with but a compara-
tively small amount of the old thallus persisting, though both
species are highly mucilaginous and especially adapted to
extreme conditions as described by Campbell.^ Careful exami-
nation of a large number of recently revived plants failed to
demonstrate the presence of sex organs even in their early
stages and it is probable that these develop after the new
growth of thallus is well under wa3^ With the exception of the
two species of Sphcerocarpus it has been shown that all of our
xerophytic forms are perennial, persisting from year to year
unless artificially interfered with. Just how extended a period
of desiccation they are capable of withstanding has never, so
far as I know, been determined. In the autumn of 1903 the
writer attempted to germinate spores of Fossombronia longiseta
that had been collected in 1896. Some of the old plants along
with the spores were placed in petri dishes containing distilled
water, while others were sown upon moist earth. None of the
spores germinated nor did the plants revive. The material was
sent me from an eastern herbarium and I do not know the con-
ditions affecting the material after collection. The failure of
the plants to revive and of the spores to germinate can throw
little light upon the question.

'Campbell, 1904: Resistance of Drought bj Liverworts. Torreja, Vol. 4,
No. 6, p. 84.

* Campbell, 1895: ist ed., Mosses and Ferns, p. 117; 2d ed., 1901;, p. 123.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 27

So long as the water of constitution is not disturbed the de-
gree of desiccation seems to make no perceptible change in the
reviving powers of all our common xerophytic forms. In the
preparation of a previous paper' quantitative experiments were
made upon Fossomhronia longiseta to ascertain the actual water
content of air-dried plants and to test the vitality of plants and
spores that had been for several weeks subjected to the action
of such a powerful drying reagent as glacial phosphoric acid.
The results obtained showed that the plants of this species in
the normal air-dry condition contain an appreciative amount of
water which may be removed without the slightest injury to the
vitality of the plant, while spores subjected to the same treat-
ment germinate quite as readily as those under perfectly normal
conditions. To extend these results the same experiment was
carried out with Targionia hypophylla, Fimbriaria californica^
Riccia glauca, Porella bolanderi^ Cryptoniitrhitn tenerwn and
again with Fossomhronia longiseta. All material was collected
early in September before the first rains, was brought into the
laboratory where, with a fine stiff brush and other instruments,
it was carefully freed from any soil particles that might be
clinging to the rhizoids. Then to correct an error that might
creep in from the absorption of moisture by the plants from the
fingers or breath of the operator the plants were again subjected
to normal air-dry conditions a number of days.

Following this the five lots of each plant, were carefully
weighed on a chemical balance and then placed at once in the
receiving chamber of a desiccator over a quantity of glacial
phosphoric acid. Here the material was left from September
21, 1906, until February 9, 1907, when it was again weighed
and the respective weight losses recorded. The weights before
and after artificial drying are given below :

Fimbriaria.

Normal Air-dry
Weight.

Loss of Weight After Four
Months Desiccation.

lot I

0.05 g.

0.0025 g.

lot 2

0.05 g.

0.0007 g-

lot 3

thi

rown out as defective.

lot 4

0.072 g.

0.0091 g.

lot 5

0.105 g-

0.0047 g.

*Loc. cit.

28

Targionia.
lot I
lot 2
lot 3
lot 4
lot 5

Cryptom iiyiu m .
lot I
lot 2
lot 3

lot 4
lot 5

J?tccta glauca.
lot I
lot 2
lot 3
lot 4
lot 5

Fossombron ia.
lot I
lot 2

lot 3
lot 4
lot 5

lot I
lot 2
lot 3
lot 4
lot 5

The wide discrepancies existing between the weights of dif-
ferent lots of Fimbriaria and of Targionia are probabl}'^ due to
the fact that certain lots were largely made up of thick, bulky
plants capable of retaining more moisture than the smaller
thinner plants composing other lots. Then too, some allowance
must be made for small particles of soil clinging to the rhizoids,
though care was exercised in the removal of all foreign matter
from the plants employed in the experiment. It will be noticed
that in the case of Cryptomilriuni, Fossombronia and the others
where the plants were of more uniform size the discrepancies
are slight.

HUMPHREY

Normal Air-dry
Weight.

Lobs of Weight After Two
Months Desiccation.

0.055 g-

0.0035 g.

0.070 g.

0.0150 g.

0.052 g.

0.0066 g.

0.075 g-

0.0046 g.

0.055 g.

0.0017 g-

Normal Air-dry
Weight.

Loss of Weight After Four
Months Desiccation.

0.050 g.

0.00250 g.

0.040 g.

0.00310 g.

0.040 g.

0.00325 g.

0.060 g.

0.00230 g.

m out as defective.

Normal Air-dry
Weight.

Loss of Weight After Two
Months Desiccation.

0.030 g.

0.00170 g.

0.035 g.

0.00200 g.

0.030 g.

0.00260 g.

0.030 g.

0.00310 g.

0.025 g-

0.00330 g.

Normal Air-dry
Weight.

Loss of Weight After Four
Months Desiccation.

0.025 g-

0.0020 g.

0.020 g.

0.0020 g.

0.040 g.

0.0020 g.

0.030 g.

0.0026 g.

0,025 g.

0.0021 g.

Normal Air-dry
Weight.

Loss of Weight After Four
Months Desiccation.

0.420 g.

0.0215 g-

0.320 g.

0.0180 g.

0-350 g.

0.0150 g.

0.402 g.

0.0170 g.

0.402 g.

O.O151 g.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 29

Of the above forms Fiuibriaria^ Targiotiia, and Porella are
the most sensitively hyj]jroscopic ; at least when moistened they
are the first to become turgid and resume activity.

By dividing the total loss in weight due to artificial desicca-
tion by the air-dry weight we obtain in percentages the actual
fraction of the air-dry weiglit that is lost in drying. For exam-
ple it was found that Fimhriaria, lost 6.06 per cent, of its air-
dry weight while losses for the other five were : Targionia^
12.3 per cent.; CryptomitrmDi, 5.86 per cent.; Rtccm, 8.47
per cent. ; Fossonibronia, 8.23 percent. ; Porella, 4.57 per cent.

From these figures it will be seen that in the normal air dry
state, Targioni'a contains the greatest amount of the water of
constitution, while Fossombi'onia and Riccia contain about two
thirds as much. Poj-eUa, the only leafy liverwort used in the
experiment, had given up all but a very small percentage of its
air-dry weight followed closely by Cryptomit7'tum, one of the
Marchantiaceae.

As a rule our summers are characterized by heavy morning
fogs which drift in over the mountains from the ocean. These
hang over the valley for three or four hours after sunrise greatly
reducing the temperature and leaving behind, each morning of
their occurrence, considerable precipitation, enough sometimes
to result in dripping from eaves. The past summer (1906), how-
ever, was unusuall}' free from these fogs, even late in the season
when ordinarily they are heaviest and most frequent. Fog-
less mornings are characterized here by slight or even no pre-
cipitation of dew and are forerunners of hot, extremely dry
days. During such a season as the one just described it is
probable that our liverworts remained in an absolutely dormant
condition. It has many times been demonstrated that all our
perennial forms on being moistened revive in from half an hour
to twelve hours, resuming growth very promptly if the supply
of moisture is constant. Campbell ^ has shown in a recent paper
that this peculiar adaptation is a property not only of certain of
the hepaticse but belongs also to Gymnogramme triangularis, a
fern common to this region, which dries up in summer, remain-
ing dormant throughout the season, without the resulting death

' Loc. cit., p. 85.

30 HUMPHREY

of the leaves as occurs in the majority of ferns. Not infre-
quently one finds in the field quite fully developed prothallia of
this species early in the autumn before the winter rains have
actually set in. It is hardly possible for these to have devel-
oped from spores that had germinated at any other time than in
spring or early summer. In fact, it has been experimentally
shown by Professor Peirce ^ that the prothallia of this species
may endure prolonged desiccation without evidence of the
slightest injury, for on being moistened they very promptly re-
vive and young sporophytes develop normally from these gameto-
phytes of the preceding season. Perennial prothallia develop-
ing tubers have been reported by Goebel ^ for the allied fern
Anog}-amme cheer ofhylla^ common in southern Europe. Such
structures belong to some of our liverworts but have not yet
been shown to occur in the prothallia of any of our ferns. Aside
from the common fern Gymnogranwie, other pteridophytes native
to California, e. g., Selaginella bigloveii and S. lefidofhylla^
the latter the well known " resurrection plant," are reported by
Campbell in the paper above referred to as remaining in a dry
and dormant condition throughout the greater portion of the
year, resuming active growth during the rainy season, at first
absorbing considerable water through their leaves as do the
leaves of Gymnogratiime.

To test the vitality of those liverworts that had been naturally
and then artificially desiccated, a certain number of plants from
each lot were placed on moist earth and others in Knop's
nutrient solution. In both cases the plants revived promptly
and are now growing vigorously, producing fruiting organs in
great number. A surprisingly small portion of the thallus of
Targionta, Riccta, Fimbriaria and Porella was actually dead ;
though in Fossomhronia^ Cryflomitrium and Anihoccros fhy-
matodcs only the apical end including a small portion of the
thallus appeared to be alive. Within five hours revived plants
from each species were fixed with chromic one per cent, fixing
solution and run up through the alcohols and bergamot oil to

2 Campbell, 1904: Resistance of Drought by Liverworts. Torreya, Vol. 4,
No. 6, p. 85.

3 Goebel, 1898 : Organography of Plants. Part II, Vol. i, p. 426.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 3 1

paraffin. The same method of fixation, deh3'dration, clearing
and infiltration were employed here as detailed in a previous
paper on the development of Fossombronia longiscta, barring
slifiht modifications. It was found that the best results could
be obtained by following the method of dehydration described
by Chamberlain.' The use of diffusion shells and the " con-
stant drip" process have proved somewhat unreliable, «ot
always giving satisfactory results for such tissues as prevail
among the more delicate hepaticge. Instead of transferring
from absolute alcohol directly to a 50 per cent, solution of
bergamot oil and then to pure bergamot, a more gradual trans-
fer was employed and a similar plan was followed in running up
through the paraffins.

In the autumn of 1903 the writer revived some material con-
sisting of Fossombronia and Fiinbj'iaria and after a few hours
placed them in a fixing solution. On studying sections of each
it was observed that in the case of Fossombronia^ not only was
a considerable portion of the thallus alive but it was found that
there were nearly mature antheridia and archegonia. Sections
of Fimbriaria through the median sulcus proved the presence
of well-advanced antheridia. This suggested the possibility of
a similar early development of sex organs in other forms than
the two mentioned above. A number of slides of Riccia, Tar-
gionia, and Cryptoniitriiun as well as of Fossombronia and
Fimbriaria were prepared and in each instance, at least, anther-
idia have been found. Only in Fossombronia and Porella have
we found archegonia. In Fimbriaria and Cry^tomitriiim
these organs occur on special receptacles none of which showed
any sign of development. In Riccia and Crytomitriimi only
early stages of the antheridium were present which was rather
contrary to anticipation. A large number of plants of each
genus were sectioned and the great majority of them were
sterile. Qiiite early in the rainy season we have been in the
custom of collecting quantities of Riccia in which no difficulty
is experienced in obtaining virtually all stages in the develop-
ment of the sex organs. For this reason the writer feels that a
more exhaustive examination of this species ought to reveal

'Chamberlain, 1905 : Methods in Plant Histology, p. 22.

32 HUMPHREY

fruiting organs quite as advanced as those of 1 argioni'a or
Fimhi'iar'ia. As for Cryplomt't?'tum, it is not surprising to find
so few antheridia, for even as late as the first week of February
sections of Cryfto^niti'mm revealed the presence of mature
antheridia while in the case of Fimbriaria they may be found
almost ripe before the beginning of the wet season. Abrams '
in his studies of Cryftomitrmm found that the antheridia
matured earlier than the archegonia and informs me that he
found fairly well advanced stages of the antheridium as early
as November. In Porella not only all stages in the develop-
ment of sex organs are to be found but well-developed sporo-
phytes as well. This accounts for the very sudden appearance
of mature sporophytes shortly following a few days of wet
weather. Growing as it ordinarily does on the exposed sur-
faces of rocks and tree trunks, it is liable to severe exposure
and may be left quite dry in a few hours of wind or sunshine.
This fact may have some influence upon its reproductive habits
and, in a measure, account for the advanced sporogonia found in
revived mxaterial. At any rate Porella is scarcely to be com-
pared with such thallose forms as Riccia or Targionia. The dif-
ferences in habit and structure might well beget differences rela-
tive to the appearance and time of development of the sex organs.

In another part of this paper the writer mentions having
found Fossombronia longiseta so situated out of doors as to re-
ceive a constant supply of moisture from a leak in a water
pipe. Many of these plants were examined on November i
very near the close of the dry season and were found bearing
embryos in various stages of development and in some instances
well advanced sporogonia as shown in Fig. i. By the first of
Januar}'^ the spores and elaters had matured and some capsules
had already dehisced.

Not having observed these plants at the time of fertilization
it cannot be definitely said just when the antheridia and arche-
gonia matured. However, the approximate date may be ascer-
tained by observing the rate of development of the sporogonium
and of the ripening of the spores. In 1904 the first rain of

'Abrams, 1S99: Structure and Development of Cryptomitrium tenerum.
Botanical Gazette, Vol. 28, pp. 110-121.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 33

any consequence fell on the twenty-first of August, but liver-
worts barely revived by it were overtaken by a period of dry,
hot weather which lasted till September 23, when the rainy
season began. It might justly be assumed that the resumption
of growth, as regards Fossomhronia at least, began at this time,
More than the usual amount of rain was recorded for October
and November and growing conditions were very favorable.
Material collected in early February showed sporongia in which
the spore-mother cells w^ere preparing to pass into the tetraspore
condition, and six weeks later material was collected bearing
ripe spores and elaters. From this it will be seen that, having
a favorable rainy season, the time required to produce ripe
spores is approximately six months after the first rains of any
consequence. Judging from the advanced development in which
one finds antheridia and archegonia at the close of the summer
season, fertilization must occur within a few weeks after
resumption of growth is well established.

With regard to the plants that had grown unchecked through-
out the summer, it must be noted that temperature and humidity
conditions are widely different from those prevailing in winter.
Though the plants were well irrigated, the atmospheric condi-
tions were such as to influence to some extent the growth and
behavior of the thallus and fruiting organs. Assuming, how-
ever, the same rate of development of the sporogonia for these
plants as for those grown in winter, fertilization must have
occurred during July and later. Growing in a paraffin-coated
glass dish containing Knop's nutrient solution, the writer has
under observation several plants of Fossonibronia^ placed there
a month ago in air-dry condition as nearly free from soil parti-
cles as they could be rendered by the use of a fine, stiff brush.
Antheridia and archegonia then in the initial stages of develop-
ment are now about as far advanced as were the oldest of those
observed as soon as the plants had revived — in the course of
twelve hours. These oldest ones are now maturing. It will
be shown elsewhere, however, that Knop's solution affords a
very active stimulus to development of the thallus and fruiting
organs, probably shortening the time of their normal develop-
ment considerably.

Proc. Wash. Acad. Sci., January, 1908.

34 HUMPHREY

At any rate, it is evident that under conditions prevailing in
this part of California a second crop of antheridia appears before
the close of a single growing season, and in the case oi J^ossom-
bronia at least, when the supply of moisture is maintained, ferti-
lization and ultimate development of the sporophyte proceed
regularly, the spores ripening three or four months earlier than
those of the winter crop. It is probable that the second growth
of antheridia and archegonia begin to develop at least a month
before the close of the rainy season, and this fact is of particu-
lar importance to the plant as it insures the maturity of the
sporophytic generation well within the rainy season.

Certain of our liverworts show a tendency to develop tubers
or other structures enabling them to resist drought. For ex-
ample, in the southern part of the State, GeothalliLS tiiherosus is
a form bearing well-marked tubers, while in our own locality
Fossomhroriia develops tuber-like thickenings of the stem, and
well-defined tubers are to be found in the case of Anthoceros
■phymaiodes. All the plants employed in this study showed the
presence of scales or hairs which in some instances secrete
mucilage ; these along with the mucilage cells within the thallus
are structures undoubtedly serving as water-storage organs.

According to Howe,^ aside from Ant/iocerosphymatodes, tuber-
ous structures are not uncommon among plants of A.fearsom\
both species growing ordinarily in exposed places. In other
parts of the world, hepaticae bearing tubers have been found.
For example GoebeP describes two species of Anthoceros^ A.
argentinus and A. dichotomus and a species of Fossomhronia^
F. tuberifera, common in certain parts of Chili. He regards
the organ in the last named species a true tuber not unlike the
same structure developed by our Anthoceros phymatodes, but
considers the tubers oi A. argentinus and A. dichotomus as trans-
formed branches of the thallus, the ends of which, have become
swollen and filled with reserve food products, very similar to
what prevails in our common species A. pearsoni. Goebel'
also reports certain species of Riccia as developing tubers,

^ Loc. cit., p. 184.
^ Loc. cit., p. 293.
^ Loc. cit., p. 70.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 35

notably an Italian form in which he observed entire portions of
the thallus developed into tuber-like structures. Stephani ^ has
also described similar structures for Riccia bulbifcra. None of
our Californian Riccias has yet been described as bearing
tubers or tuber-like growths, though Riccia trichocarpa, men-
tioned elsewhere in this paper, is well adapted to withstand
severe exposure, the thallus being densely clothed with tawny
setae .3 to .65 mm. long, with those toward the apex in as many
as eight to twelve series, affording a very effective protection
against excessive drying. Though these tuber-like structures
wherever they have been examined are always found to con-
tain large quantities of reserve food, there is little doubt that
they are instrumental as water-storage organs.

STUDIES IN THE GERMINATION OF SPORES.

In a previous paper the writer has described and figured the
germination of the spores of Fossombronia loiigiseta. The
results of experiments then carried out showed the early devel-
opment of this species to agree closely with certain forms, not-
ably SphcBTOcarpiis cali/ornicns, investigated by Campbell.^

Most of the hepaticas of this region produce spores character-
ized by a heavy, more or less sculptured exosporium which
Leitgeb^ found to consist of two parts or layers of which the
inner belongs to the spore itself and is the exine, while the
perinium or outer layer is formed later and consists of parts of
the sporocyte. Leitgeb suggests the probability of the perinium
as a protection against drought but in the light of more recent
studies of certain forms producing thick-walled spores, e. g.y
Corsinia, Preissia^ Anthoceros and SfhcBvocarpus^ it may be held
that the perinium can certainly have nothimg to do with a rest-
ing period, for spores of these forms germinate readily within a
few days after being sown. As to the thick exosporium fur-
nishing a protection against excessive dryness there can be little
doubt, although such forms as Corsinia and the aquatic Ric-

' Stephani, 1S9S : Species Hepaticarum. Bull, de I'Herb. Boiss., Vol. VI, p.

333-

''■ Campbell, 1S96 : Notes on Sphserocarpus. Enthea, Vol. IV, No. 5, PI. 2.
* Leitgeb, 18S4, Ueber Bau und Entwicklung der Sporenhaute, Graz.

36 HUMPHREY

ciete produce spores of similar character. Goebel,' however,

suggests that localities favorable to Corsinia may not always be

moist, and in the case of the aquatic Riccie^e the perinium very

effectively serves as a protection against invasion of the spore by

fungal hyphas.

Although the great majority of our xerophilous liverworts

produce thick-walled spores there are such exceptions as Po7'-

ella bolanderi 2iV\di Radiila complanata whose spores are protected

by a thin, frail exosporium and are incapable of withstanding

prolonged drying. These liverworts grow on tree trunks or

exposed surfaces of rocks where they are subject to sudden

drying out and we should naturally expect their spores to be
better protected.

The character of the spore-w^all, however, might suggest the
possibility of these forms being naturally more h3'^grophilous in
character, while their adaptation to a dry habitat is merely one
phase of the extreme adaptability common to both forms.

In order to get at the relationship of the exospore to the spore
a study of the life conditions of the several species is necessary
and some light will be thrown upon the subject w^hen we ascer-
tain at what time, in nature, the spores of a large variety of
forms germinate. With a large number of forms it has been
repeatedly shown that spores sown on soil under conditions as
nearly normal as possible germinate only after a long period of
rest.

In June, 1904, the writer sowed on sterilized earth, a quantity
of spores of Fossombi'onia that had been gathered a few days
before sowing. The culture was made in an earthen saucer
and placed within another close-fitting earthen saucer, in which
was maintained a supply of water so that the dirt was kept con-
stantly moist from below. The culture was placed on a table
near a window where it could receive a nearly normal supply
of light. During the latter part of August a few spores had
germinated but not until October did they germinate in any
great number. The spore of Fossombronia, while perhaps not
as well protected as that of Finibriaria or Targionra^ nevertheless
is furnished with a resistant exosporium and is well adapted to

' Loc. cit., 107.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 37

the xerophilous conditions under which the plant grows. The
spores are positively known to maintain their vitality for at least
two years but for how much longer, cannot be said. The thin-
walled spores of such forms as Aiieura, Fcgatella^ Radula and
Porella withstand but little drying, losing all germinating power
within a few months at most, which fact would seem to be evi-
dence in favor of the statement that the thick exosporium is a
protective structure.

Spores of Anthoceros fiisij-ormis one year and eight months
old, when sown on sterilized soil germinated copiously within
two weeks. Here we have an instance of spores that are known
not to require a resting period before germinating, 3'et, unlike
Aneura, the spores of Anthoceros are known to endure at least
two years desiccation. Spores of Fimbriai'ia californica and
Targionia hypo-phylla two years of age were sown on sterilized
earth and kept well watered. These germinated within twelve
and fifty-six days respectively. This wide discrepancy is diffi-
cult to account for. The soil was kept as free from fungi as
possible : however, only a few of the Targionia spores ger-
minated and this fact may have been due to reduced vitality from
prolonged drying. Further experiments with spores of this
genus are necessary ; in fact, germination experiments conducted
out of doors are the ones of greatest value. Such a series of
culture experiments was started more than a year ago in the
Stanford Arboretum but before results of any value were ob-
tained the cultures were accidentally destroyed by a workman.
It was then too late in the season to start another series, it being
very near the close of the rainy season. A fresh collection of
spores was made from the 1906 crop and several artificial cul-
tures were started indoors, using both sterilized and non-steril-
ized earth and Knop's nutrient solution. The spores used in
the various cultures were of the followint; forms : Ricci'a o'huica^
Fimbriaria violacea, F. californica^ Targionia hypofhylla^
Fossombronia longiseta, Cryptomitrium tcneriun and Aneura
multifida major.

No spores of the first mentioned plant have yet germinated,
though sown six months ago after a resting period of five
months. Spores of both species of Fimbriaria failed to ger-

38 HUMPHREY

minate, thouirh sown at the same time as those in the case of
Riccia. A very few of the spores of either Fossombro7iia or
Cryptotmtriuni germinated. Though he did not duplicate in
every case the cultures of the writer, Professor Peirce had a
similar experience with all spores of the same (1906) crop.
This apparent loss of vitality is difficult to account for, unless
due to the unusual character of the weather during the spring
months. Ordinarily our last rains occur in April, by which time
the spores of the above-named hepaticae have either been dis-
charged or are maturing. An examination of the daily record
of precipitation from April i to the close of the season may be
instructive. Up to April i from January i the total rainfall
amounted to 10.04 inches. From April i to June 15 the record
was as follows:

Amount of Amount of

Date. Precipitation. Date. Precipitation.

April 1 0.06 May 14 0.02

" 10 o.oi " 25 0.34

" 23 o.ii " 26 0.04

" 24 0.03 " 27 0.51

" 27 0.29 " 28 O.II

" 29 trace " 28 o.ii

May 10 0.01 June 3 0.02

" II 0.02 " 5 0.73

15 0.24

" 26 trace

From the above record it may readily be seen that from
April I to April 23 the total precipitation amounted to but .07
of an inch — scarcely more than a trace. Between the twenty-
third and twenty-seventh enough rain fell to revive the already
dried mosses and liverworts to a condition of perfectl}^ normal
turgidity and they remained so at least a week after the last rain-
fall. Now during April the spores of a number of our liver-
worts are only beginning to mature, and it is barely possible
that the prolonged dry period during that month interfered with
the proper ripening of the spores. On the other hand, spores
of certain forms, for example, Fossomhronia, which had fully
matured during March but were not gathered till after the
rainy season, failed to germinate ; and this suggests the possibility
of the vitality of the spores having been impaired by successive

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 39

rains and rapid desiccation following one upon the other. What-
ever be the cause of impairment the fact remains that almost
no spores germinated even under most painstaking precautions
as to culture conditions. To get at the root of the matter, how-
ever, a series of experiments is necessary in which the student
should duplicate as nearly as possible the conditions prevailing
from April i to June 15 of 1906.

It is an unsettled question as to whether the spores of many
liverworts actually require a prolonged resting-period, and there
is still need of investigation along this line. We are familiar
with the fact that the spores of most xerophilous forms do lie
dormant several weeks before germinating but it has not been
shown in very many instances that this resting-period is inti-
mately connected with the preparation of the spore for germina-
tion. The writer has demonstrated beyond doubt that the spores
of Fossomhronia longiseta, under certain conditions, do not re-
quire a resting-period. Cultures were prepared using different
media with a view to testing, if possible, the relative influences
of the various media upon germination. One lot of spores col-
lected on January 10, 1906, were sown January 12 on pulver-
ized earth that had been thoroughly steam-sterilized and well
watered with distilled water. Care was exercised to select soil
on which Fossombronia grows vigorously out of doors ; this
was placed in a crystallization dish and covered with one half
of a petri dish to keep the culture as free from dust as possible.
A number of crystallization dishes were thoroughly cleaned
and then covered on the inside with a thin coating of parafiin
to prevent the water used from taking up any impurities that
might exist in the glass. In three of these, Knop's nutrient
solution of the following proportions was placed :

HjO 500 c.c.

Calcium nitrate 2.00 grams.

Magnesium sulphate 0.5 "

Potassium nitrate 0.5 "

Potassium phosphate 0.5 "

Other solutions of varying strength were employed as follows :
Wfi 500 c.c.

Calcium nitrate 2.00 grams.

Magnesium sulphate 0.5 "

Potassium nitrate 0.5 "

Potassium phosphate omitted.

40

HUMPHREY

Other stock solutions were made up in which the amounts of
the various sahs used in a normal solution were reduced in
quantity one fourth and one eighth, though the quantity of dis-
tilled water was kept the same, 500 c.c, in all the solutions.
It will be noticed that from one solution the acid salt, potassium
phosphate, was left out with a view to testing its value as a
stimulus to germination. Spores of the same lot were sown in
the various solutions the same day they were collected ; the cul-
ture dishes were carefully covered and placed on a north window-
sill where a uniform amount of diffused light fell upon each
culture. In another dish that had been paraffin-coated, water
that had been twice distilled was placed and on the same date
spores of the same lot were sown. These germinated in con-
siderable numbers within twelve days. Some of the water used
in this culture was taken from the supply bottle and evaporated
on a slide. On examination, a precipitate of organic particles
was found, showing that certain compounds had passed over
with the water in distilling, and it was thought probable that
these acted as a stimulus, for the spores in this culture germi-
nated before those in the culture containing none of the acid
salt. In fact these latter seemed to show no indications of any
preparation for germination, such as change in color of exospor-
ium from very dark brown to a lighter more translucent shade.
The distilled water used in the preparation of the various cul-
ture solutions was from the same stock bottle as that used in the
distilled-water culture. Why spores collected at the same time
and sown at the same time on these two media should have
germinated earlier in distilled water containing organic impuri-
ties than in a culture solution void of one of its constituent salts
is difficult to account for unless in the latter we have a physio-
logically unbalanced solution which has been shown by Loeb ^
to exercise a certain toxic effect upon certain marine and fresh-
water animals and by Osterhout^ in his experiments upon algae.
This, however, needs far more extensive investigation than has
yet been given to it.

'Loeb, 1905: PflUger's Archiv, 107: 252.

* Osteihout, 1906 : On the Importance of Physiologically Balanced Solutions
for Plants. University of California Publications. Botany, \o\. 2, No. 10, pp.
229-230.

THYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC^ 4I

The spores in the unbalanced culture solution germinated
within six da3's after those in the normal Knop's solution, which
would indicate the existence of some stimulating factor in that
solution. Another lot of distilled water was then prepared by
allowing the first fourth or fifth part to distil as waste water, thus
getting rid of the larger part of the volatile substances. Only
the middle three fifths was saved and this was redistilled in the
same manner. The best portion of this was again distilled and
only the middle portion of the product was collected ; in this way
water of a high degree of purity was obtained. This was used
in the preparation of another distilled water culture and spores
of Fossombronia sown eighteen days ago show no evidence of
germinating. Spores sown on Knop's solution, normal, germi-
nated within fourteen days producing vigorous germ tubes with
rather more than the usual amount of chlorophyll. These young
plants have grown more rapidly and vigorously than those in
distilled water that germinated approximately at the same time.
Spores sown in the normal solution germinated two days later
than those in either of the reduced solutions but in the former
there was a far greater number of young plants two days after
the first observation of germination.

Spores of Fimbriaria californica collected in March, 1905,
were sown in Knop's solution (normal) and germinated promptly
within twelve days. Young plants in this culture that are now
sixteen to twenty days old are characterized by the unusual
length of the germ-tube, in some instances measuring more than
one and a half millimeters from the basal end to the first cell
cut off at the distal end of the germ-tube. In some instances
two germ-tubes were seen to emerge from a single spore as
figured by Campbell ^ for Targionia hyfo^hylla. Spores of
the same species were sown in an earthen saucer filled more
than half-full of finely pulverized non-sterilized soil. These
germinated promptly and in great numbers. Both cultures
were so placed as to receive light from a north window : the
Knop's solution culture received somewhat the greater amount
of light being in a glass dish, coated part way up with paraffin.
None of the young plants of the soil culture produced germ-

' Loc. cit., 1st ed., p. 65 ; 2d ed., p. 66.

42 HUMPHREY

tubes of more than usual length while some showed the forma-
tion of cross-walls very early. The young plants in the nutri-
ent solution have grown rapidly, producing in some instances
as advanced a development as is seen in Fig. 21, within eigh-
teen days. A very few young plants in which there was only
a slight germ-tube development had already begun to assume
the dorsi-ventral form common to this genus. Desiccated
plants of F'imbriaria, Riccia and Fossomb^-onia when placed
in Knop's solution revived quickly producing within a few
days a greatly increased number of rhizoids and in the case of
jFossojubronm, many more than the usual number of antheridia,
closely covering the dorsal surface of the young growing shoot.
From these and other results it seems highly probable that
Knop's nutrient solution exerts a stimulative influence upon the
development of the thallus, rhizoids, and reproductive organs
as well as the germination of the spores. Further investigation,
however, with solutions of varying strength and composition is
needed in order definitely to determine the nature and chemical
composition of the salt or combination of salts that act as a
stimulus to growth.

According to Goebel,^ germination in the Marchantiese when
compared with other thallose liverworts presents certain char-
acteristic differences. He separates the product of germina-
tion of a spore of Preissia for example, into three structures,
/. (?., the germ-tube, the '* pro-embryo " and the young plant.
He finds that the "pro-embryo" is positively heliotropic and
develops at its end a flattened cell mass which he terms the
germ disk. This forms at right angles to the direction of the
light-rays and the young plant arises from one quadrant of the
germ disk as shown in his Fig. 95.

On the same page he states that a similar germination is
common in the Riccieae contrary to the description and figures
of Campbell^ who studied the ^<txvs\\\\'AX\onoi Riccia tricho car fa
and a number of other forms. He refers to Campbell's Fig. 9,
p. 38, as failing to bear out his statement that the axis of growth
in the young plant is continuous with that of the germ-tube.

' Loc cit., p. III.

^Loc. cit., isted., p. 39; 2ded., p. 38.

PHYSIOLOGY AND MORPHOLOGY OF CALH-ORMA HEPATIC^ 4

An examination of the figure referred to, however, will show
that although an angle was present in the young gametophyte
it developed in the germ-tube itself, the upper portion of which
lies in the direct axis of growth of the young plant, no such
structure as a pro-embryo being present. In his study of Fhn-
briaria and Targionia his figures show an early development
of the young plant not unlike that in the case of Riccia tricho-
carpa. The writer's investigations relative to the germination
of both Fossonihronia and Fimbriaria agree in detail with those
of Campbell in the forms studied by him (Figs. 22 and 23).

MacDougal • has shown that, given the necessary intensity of
light, its direction will vitally influence the form and habit of a
living plant as well as the disposition of various organs incident
to its development. This being true, we might, a -priori^
assume that Goebel's Preissia and Riccia spores on germinat-
ing take on the angular form shown in his Fig. 95 because of
certain external factors and, most of all, light, instead of trying
to account for it on grounds of heredity or of certain inherent
factors. In fact, Peirce,^ Rosenvinge,^ and others have experi-
mentally shown that in the case of certain algag, germinating
spores are profoundly influenced by the direction and intensity
of light. Eliminating so far as possible all other external factors
it was found that the young plants always grew towards the
light, while the rhizoid or the holdfast vvas negatively helio-
tropic. Peirce has demonstrated that in addition to this, light
exerts a directive influence upon the formation of cross-walls
in sporelings of algas, liverworts and certain ferns, these walls
always forming at right angles to the direction of the light ra3's.*

With ferns and liverworts he has further shown that when the
long axis of the germ-tube lies in the direction of the light rays,
the continued growth of the young plant will be in the same
direction. More than this, he has demonstrated that when the
germ-tube ruptures the spore wall on the side away from the

'MacDougal, 1903 : The Influence of Light and Darkness upon Growth and
Development. Memoirs of the New York Botanical Garden, Vol. II.

2Loc. cit., p. 338.

'Rosenvinge, 1S89 : Influence d'agents exterieurs sur I'organisation polaire
et dorsiventral des'plantes. Revue Generate de Botanique, i.

* Peirce, 1906 : Studies of Irritability in Plants, p. 453.

44 HUMPHREY

light there is a strong tendency to turn toward the light, and if
this is not accomplished by the germ-tube, the further develop-
ment of the young plant will at once or gradually assume the
direction of positive heliotropism.

DURATION OF VITALITY OF CERTAIN GREEN OR THIN-
WALLED SPORES.

A failure to germinate the spores of Aneura multijida major
after they had been allowed to become several weeks old,
induced the writer to prepare a series of cultures to test the life
duration of the spores of Aneura^ Radula^ Poi-ella and
FegateUa.

The first spore cultures of all these forms were started on the
same day of collection in order to determine whether the spores
were ready for germination under normal conditions. Practi-
cally all spores germinated in cultures consisting of sterilized
earth, Knop's solution and distilled water. I next allowed
spores of FegateUa to dry for three days, by placing them on a
sheet of paper on the laboratory table. These were then sown
in the various culture media. Many of the spores had already
germinated within the capsule but these along with all spores
still ungerminated failed to show the slightest sign of any
recovery after days of constant moisture and favorable light.
Later, Professor Peirce demonstrated that one hour's drying
under normal conditions was sufficient to kill all spores, and
further experiments with the spores of this species were re-
garded unnecessary. The spore wall in the case of FegateUa
is very thin, allowing rapid loss of the water of constitution,
resulting in a general contraction of spore contents and collapse
of the spore wall.

On January 12 cultures were started, sowing spores of Radula
complanata collected on the ninth of the same month. Prac-
tically all of these germinated within a few days, so another
trial was not made till the twenty-eighth of January. Approxi-
mately ten per cent, of the spores of this sowing failed to
recover, the cytoplasm and other spore contents having con-
tracted towards the center forming a brownish mass.

On February 11 a third culture was started, the spores now

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC/E 45

having dried thirty-two days. Between 20 and 25 per cent,
of these spores failed to recover. The last lot of spores were
sown on March 3 — fifty-two days after collecting. Approxi-
mately forty per cent, of these were desiccated beyond recovery.
Time does not permit at the present further testing of the spores
of this species but judging from the results of the above cultures
very few if any spores could endure longer than a few months
at the most.

Porella bolanderiv^z.?, then experimented with, cultures being
started at various intervals using the same media as in preced-
ing cultures. The results obtained were not greatly different
from those in the case of Radiila though in my last culture
which was made on March 5 not more than half the spores have
recovered and fewer still have germinated after eleven days.
The spores in this culture had been allowed to dry thirty-eight
days, or fourteen days less than was allowed spores used in the
last Radula culture. Making due allowance for the possibility
of toxic effects due to the activity of bacteria or fungi that might
have been in the various cultures it is highly improbable that
the failure of constantly increasing numbers of spores to revive
could have been entirely due to any such cause and we are jus-
tified in the conclusion that the spores of these forms and prob-
ably some others not yet investigated are incapable of enduring
more than a comparatively slight amount of desiccation.

Summary.

1. Plants of Fossomhronia longiscta, as it occurs here, are
seldom free from infection by a fungus, which, in its relation
to the host, acts as a true parasite. In early stages the fungus
is confined to the rhizoids and compact tissue of the stem, ulti-
mately extending throughout all vegetative tissues of the gameto-
phyte. Large, black sclerotia develop within the cells of the
leaves and peripheral cells of the stem, resulting in the death
of the host.

No instance of infection of sex organs or sporogonium has
been observed.

2. Fimbriaria calif ornica has been found to"be infected by
a fungus, the hyphee being confined to the outermost three or

46 HUMPHREY

four layers of ventral cells and to both sorts of rhizoids. Its
relation to the host is apparently symbiotic.

3. A fungus, evidently epiphytic in habit, was found associ-
ated with Anctira 'nmltijida major. Its relation to the host is
that of a parasite.

4. Fungi were also found associated with Anthoceros fearsoni
and Porella bolanderi. In both cases the fungus is epiphytic,
sending hausteria-like branches into the tissues of the host.

5. Fertilization in Fcgatella conica in California occurs in
early spring. At the close of the dry season the spores are in
the tetrad stage. With the commencement of the rainy season,
the growth of the female receptacle is resumed and the spores
mature early in January.

6. When hygrophilous forms such as Aneura -pingids or
Fegatella conica are transferred to a dry habitat and allowed
to become desiccated they are unable to recover.

7. All of our xerophilous forms such as Fimbriaria cali-
Jbrntca, Targionia hyfoyhylla^ several species of Riccia, Fos-

sombroma, and others resume growth promptly after months of
extreme drought. When artificially desiccated over glacial
phosphoric acid until further loss of weight is imperceptible, on
being moistened they revive quite as promptly as when nor-
mally air-dried.

8. Artificial drying does not impair germination of spores of
xerophilous forms.

9. In Porella bolanderi material examined at the close of
the dry season exhibits holdover antheridia and archegonia and
well-advanced sporogonia. Other xerophilous forms of peren-
nial habit show sex organs in various stages of development.

10. Fossombronia growing in a habitat of constant moisture
matured spores in March and December to February.

11. Mucilage secreting structures are common to many forms
and especially to our species of Anthoceros. A. fearsoni and
A. fhymaiodcs are rendered resistant to drought by the presence
of tubers. Similar structures, though less developed, are com-
mon to Fossombronia longiseta in dry regions.

12. Spores of xerophilous forms are known to retain their
vitality at least two years and probably longer. In nature they
pass through a resting period of several weeks.

PHYSIOLOGY AND MORPHOLOGY OF CALIFORNIA HEPATIC/E 47

13. Spores of Fossoinbronia longiscta sown in Knop's solution
and in impure distilled water germinated in the former within
fourteen days and in the latter within twelve days after removal
from the capsule. Spores sown on sterilized soil show no evi-
dence of germination after sixty-two days.

14. The spores of Fcgatella conica, Aneura pingtiis, Porella
bolanderi and Radnia comflanata were found incapable of
withstanding prolonged desiccation. Germination occurs nor-
mally within the capsule or very soon after dispersal.

BIBLIOGRAPHY.
Abrams.

1899 Structure and Development of Cryptomitrium teiicrian (Hook.) Aust.

Botanical Gazette, Vol. 3S.

Bruchmann.

1904 Ueber das Prothallium und die Keimpflanze von Ophtoglossumvulgattim
L. Botanische Zeitung, Heft XII, Tafel VIII.

1906 Ueber das Prothallium und Sporenpflanze von Botrychium lunaria Sw.
Flora Oder Allgemeine Botanische Zeitung. Band 96, Heft i.

CampbeU.

1896 The Development of Geothallus tuberosus Camp. Annals of Botany,

Vol. IX.
1896 Notes on Sfhcerocarpus. Erythea, Vol. IV, No. 5, PI. 2.
1895 Mosses and Ferns, ist ed. 1905, 2d ed.
1904 Resistance of Drought b}- Liverworts. Torreya, Vol. 4, No. 6.

1907 Studies on the Ophioglossaceae. Annales du Jardin Botanique, Vol.
XXI.

Cavers.

1903 On Saprophytism and Mycorrhiza in Hepaticae. The New Phytologist,
Vol. II, No. 2.

1904 On the Structure and Biology of Fegatella conica Corda. Annals of
Botany, Vol. XVIII, No. LXIX, Plate VI.

1904 Contributions to the Biology of the Hepaticse. Part i, Targionia, Re-
botilia, Preissia, Monoclea.

Chamberlain.

1905 Methods in Plant Histology.

Czapek.

1889 Zur Chemie der Zell-membranen bei den Laub und Lebermoosen.
Flora, Band 86.

Goebel.

1898 Organography of Plants, Part II, Vol. i.

Golenkin.

1902 Die Mychorrhiza ahnlichen Bildungen der Marchantiaceen, Flora, Band^
90.

48 HUMPHREY

Howe.

1898 The Anthocerotacese of North America. Bulletin of the Torrej Botan-
ical Club, Vol. 25, No. I.

1899 The Hepaticse and Anthocerotes of California. Memoirs of the Torrey
Botanical Club, Vol. VII.

Humphrey.

1906 The Development of Fossombronia lotig-tseta Aust. Annals of Botany,
Vol. XX, No. LXXVII.
Jeffrey.

1898 The Gametophjte of Botrychium virginianum. University of Toronto
Studies, No. i.
Kny and Bottger.

1879 Ueber eigenthiimliche Durchwachsungen an den Wurzelharren zweier
Marchantiaceen. Verhandl. d. bot. Vereins d. Prov. Brandenburg.
Leitgeb.

Untersuchungen iiber die Lebermoose, Heft 2, Tafel 3.
1884 Ueber Bau und Entwicklung der Sporenhaute. Graz.
Loeb.

1905 Pfliiger's Archiv, 107.
MacDougal.

1908 The Influence of Light and Darkness upon Growth and Development.
Memoirs of the Nevv^ York Botanical Garden, Vol. II.
Osterhout.

1906 On the Importance of Physiologically Balanced Solutions for Plants.
University of California Publications. Botany, Vol. 2, No. 10.

Peirce.

1906 Studies of Irritability in Plants. Annals of Botany, Vol. XX, No.
LXX.
Peirce and Randolph.

1905 Studies of Irritability in Algse. Botanical Gazette, No. 40.
Rosenvinge.

1889 Influence d'agents exterieurs sur I'organisation polaire et dorsiventrale
des plantes. Revue generale de Botanique. i.
Stephani.

1898 Species Hepaticarum. Bulletin de I'Herbier Boissier, Vol. VI.

Proc. Wash. Acad. Sci., April, 190S.

EXPLANATION OF FIGURES IN PLATES I AND II.

Fig. I. Diagram of a young Sporogonium of Fossomhronia lotigiseta collected
November i, 1906, near close of the dry season. Found in a locality of
constant moisture.

2. y^^Xwr^ ^\z.-s\\. oiFossombronia longiseta. Had grown all summer. Col-
lected January 6, drawn January 7.

3. Living rhizoid of F. lotigiseta perforated by hyphae. X560.

4. Hyphae passing from cells of thallus into rhizoid. X560.

5. Abnormal rhizoid of F. longiseta containing hyphse. X s6o.

6. Thin-walled vesicle of the fungus associated with^i^. longiseta , vf'xWnn
stem tissue of the host. X 560.

7. Hypha complex within cells of the host, F. longiseta. X 560.

8. Vesicle and hyphae apparently communicating with the exterior. X
560.

9. (a) and {b). Formation of conidia by abstriction of hypha of the fun-
gus infecting F. longiseta. X S^o.

ID. Development of sclerotia in cells of a leaf of F. longiseta. X 5^0.

11. Perithecia-like structures developing from hyphae on the surface of
Aneura multijida major.

12. Uninfected cells oi Aneura multijida major showing oil-bodies, o. b., and
chromatophores. X335'

13. Infected cells of Aneura multijida major. Note the absence of oil-bodies
and small number of chromatophores. X 335-

14. Three cells of Aneura multijida major showing knots of hyphae. Chlo-
rophyll and other cell contents wanting; cells, dead. X335-

15. Infection of a young plant of Aneura tnultijida 7najor that has developed
from a gemma. X S^o.

i6a. Branching rhizoids of Aneura multijida major. X 80.
i6b. Branching rhizoids of Lepidozia attenuata. X So.

17. Longitudinal section of young sporogonium of Porella bolanderi near
close of the dry-season. X about 50.

18. Portion of an infected leaf of Porella bolanderi showing chlamydospore
(c). X560.

19. Developing sclerotia within leaf cells of Porella. X 560.

20. Infection of tissue of Anthoceros pearsoni, showing sclerotium. X 560.

21. Portion of plant of Fegatella co?tica showing receptacle. Collected
December 28, IQ06; drawn January 2, 1907. Ad. nat.

22. Apical end of a young Fimbriaria plant. X 5^0.

23. Young plant oi Fossombronia longiseta. X 5^0.

All drawings except Fig. i, 16 and 23 were made with the aid of an Abb^
camera lucida.

(50)

Proc. Wash, Acao. Sci,, Vol, X,

Plate

HUMPHREY DEL,

rHC EcxEnr L'-'MtyC.«i

siiroN.o.c

Proc. Wash. Acad, So., Vol. X.

PUVTE II.

HUMPHREY DEL,

PROCEEDINGS

OF THE

WASHINGTON ACADEMY OF SCIENCES

Vol. X, pp. 51-166. April 25, 1908.

A REVISION OF THE TENEBRIONID SUBFAMILY

CONIONTIN^.

By Thos. L. Casey.

Such genera as Eiisattus, Coniontis^ Ccelus, Branchus and
Praocis possess a peculiar and consistent habitus distinguishing
them as a special group in a somewhat heterogeneous assem-
blage, brought together under the subfamily name Asidinas by
LeConte and Horn. The only characters considered to be of
subfamily weight by the latter authors are those relating to the
coriaceous posterior margin of the penultimate abdominal seg-
ments and the presence or absence of a trochantin, but there are
assuredly some other important characters, the utilization of
which would serve to segregate the multitude of genera forming
the first subdivision of the family into more homogeneous
groups, that, in several cases, might be regarded as subfamilies
without doing any great violence to precedent in other parts of
the Coleopterous series. I have in mind more especially the size
of the mentum and certain characters relating to the hind cox£e.
In regard to subfamily division among those Tenebrionids which
have no coriaceous hind margins of the abdominal segments,
the following scheme might be proposed : —

Mentum large, covering both the maxillae and ligula; hind coxae
variable, generally transverse, feebly oblique in some Euryme-
toponids and strongly oblique in the Zophosini ; trochantin in-
visible throughout Tent yriin^e

Mentum smaller, never concealing both the maxillae and ligula' 2

'The genera Astrotus and Ologlyftus constitute a remarkable exception to this
rule, the entire buccal opening being closely filled by the mentum ; but in all
other characters, including facies, these genera are purely Asidid, showing how dif-
ficult it is to select characters constant through large groups in the Tenebrionidse.
Proc. Wash. Acad. Sci., April, 190S. 51

52 CASEY

2 — Last abdominal segment unmodified 3

Last abdominal segment transversely excavated 4

3 — Posterior coxae oblique and long, approaching the sides of the

body; trochantin distinct Coniontin^

Posterior coxae transverse and short, more remote from the sides of the
body; trochantin inconstant, invisible to distinct Asidin^

4 — Trochantin invisible; epipleurse usually wholly undefined.

ZOPHERIN^

The Tentyriinas, as above limited, are divisible into several
groups of rather more than tribal value, among which may be
cited those genera having the hind coxae much extended trans-
versely, linear and frequently feebly oblique, the body generally
winged ; then those with the hind coxae transverse and more or
less oval, and, lastly, those with the hind coxae very oblique and
with the ante-coxal grooves still more oblique, known tribally
as the Zophosini. If the Zophosini, which are as completely
isolated in the old world fauna as the Coniontinae are in North
and South America, were placed last in the Tentyriid series, the
Coniontinag would naturally follow them, having many sugges-
tive points of resemblance. By this arrangement, which it will
be observed, greatly subordinates the importance of the trochan-
tin, many minute forms without trochantin, such as Arceoschtztis
and Stenosis would be included with the Asidinse. The Zophe-
rinae are so isolated as to assume subfamily rank, very obviously
it would seem.

The Coniontin^e as a group have the body oblong or oblong-
oval in form, compact, with the prothorax at base as wide as the
elytra, usually enveloping the humeri at the sides to a more or
less noticeable degree, the epistoma emarginate or sinuate at the
middle, the eyes at the sides of the head, transverse, rarely
divided and the mandibles short, stout and bifid at tip. The
mentum is rather small, obtrapezoidal and sinuate at tip and
there is at the apex of the post-mental surface a transverse area,
defined by a rapid divergence of the gular sutures. The ligula
is large, transverse, sinuate and densely chitinized and the palpi
of the usual Tenebrionid t3'pe, the last joint obtriangular. The
antennae differ from those of Asida in having a free, subequal
club, generally of three joints. The legs are rather stout, well
developed, the anterior tibiae more or less obtriangular, with the

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 53

outer angle frequently prolonged, the tarsi always spinulose be-
neath, the anterior coxte transversely oval, the intermediate very
moderatel}"^ separated, the prosternal process more or less ex-
tended and porrect and the mesosternum between the coxae im-
pressed and concave, these latter chara.ters, with the long and
well developed tibial spurs, elongate basal joint of the hind tarsi
and oblique hind coxaj, recalling the Zophosini very forcibly.
The fourth abdominal segment is very short when compared
with the three anterior segments. The members of the Conion-
tinge are neo-subarctic and neo-subantarctic exclusively and are
represented on the Pacific slope of North America by the
Coniontini and Coelini, on the Atlantic slope by the Branchini
and in South America by Praocts and allied genera. In the
old world, as previously intimated, they are replaced by the
Zophosini, which might be assumed to have almost if not quite
subfamily rank.

The present revision omits the Praocini, as I have no repre-
sentative of this South American tribe. The other tribes may
be outlined briefly as follows : —

Intercoxal process of the abdomen moderately broad, truncate; elytra
embracing the sides of the body beneath ; antennae well developed,
elongate; hind coxai rather more feebly oblique, body generally
ridged above and roughly sculptured Branchini

Intercoxal process narrow, more pointed or rounded and projecting
further into the metasternum 2

2 — Antennae long and slender, very well developed; basal joint of
the anterior tarsi variable, elongate or short, in the latter case
more or less eccentrically triangular and in some cases very promi-
nent at the inner apical angle, suggesting the spur-like process of
the next tribe Coniontini

Antennae very small, compact, with the joints transverse; basal joint
of the anterior tarsi greatly prolonged internally at apex, forming
a spur which sometimes extends almost to the end of the tarsus.

Coelini

The Coniontini are extremely abundant, having a widely ex-
tended distribution almost coincident with that of Eleodes^ but
the Coelini are less numerous and are restricted to the seashore
of California and Lower California, occurring also on the
neighboring islands.

The form of the prosternal process is not always of deci-

54 CASEY

sive importance in the Tenebrionidae, as for example within
the tribe Epitragini, but I have never noticed any species of the
Coniontince in which this process is not prolonged and subhori-
zontal, and but few of the Asidin£e in which it is not strongly and
rapidly deflexed to the posterior margin of the prosternum, this
being a rather consistent feature distinguishing these two sub-
families, in addition to the form and extent of the mentum and
hind coxae and structure of the antennae. All of these differential
characters are as pronounced in Branc/ms as they are in Eiisat-
tus, Coniontts and Cceltis, and it is therefore somewhat surpris-
ing that the true affinities of Branchus were not more fully
dwelt upon by LeConte and Horn, who make but little mention
of the subject. The gular characters given by LeConte (Class.,
I ed., p. 219) to distinguish the Branchini from the Coniontini
appear to be inconstant and of but little importance. As to the
Praocini of Lacordaire, that author, it is true, includes certain
genera with deflexed prosternal process, making of them a
separate section of the tribe, but it is possible that they may
properly be excluded from the subfamily Coniontinas, leaving
Praocis and its subgenera alone to be included among the
South American forms.

Tribe Branchini.

This tribe, which has little or no relationship with Asi'da, is
very limited in the number of its species and genera and is the
Atlantic coast representative of the western Coniontini and
Coelini, extending from Florida to Central America, although
one species has recently been announced, by Mr. Champion,
as inhabiting the western Mexican State of Guerrero. I have no
example of our Florida or Bahama species, but a specimen of
Branchus obscurtis Horn, from Nicaragua, very kindly given
me by Mr. Wickham, is a pure Eusattid in every way, except
in having the hind coxae more widely separated and the basal
joint of the anterior tarsi not materially larger than those that
follow it. One of its most striking points of resemblance with
the Eusatti and with Praocis is the form of the epipleurae, which
are narrow very nearly to the base, where they suddenl};- ex-
pand to the sides of the elytra, forming a prominent point

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 55

at the humeri, and the acute side margins of the elytra are
exactly as in Discodcmus and Mcgasattus ; the basal angles of
the prothorax are prolonged backward very distinctly but to a
much slighter degree than in any of the Eusatti. The scutellum
is moderately developed and triangular to obsolete, the head and
eyes nearly as in Coniontini, except that the notch at the ex-
tremity of each oblique suture is much more pronounced, the
general form recalling Asfda, and all the oral organs as well as
the antennae are purely Coniontid. The pronotum is finely
costate along the median line and reflexed at the sides, the basal
joint of the hind tarsi much elongated and the anterior tibiae
minutely dentate externally at tip.

Tribe Coniontini.
It appears to the writer eminently illogical to consider the
remarkably diversified elytral epipleurai, among the forms allied
to Eiisattus^ as having anything less than generic value. It is
unfortunate that our nomenclatorial system should be compelled
to absorb so many names, but there is nothing to prove that the
various and radical modifications are inconstant and I have seen
no intermediate forms. The epipleurse are perfectly constant
throughout the genera £usattus, with numerous species, Disco-
dcnms, with some seven apparently specific forms and elsewhere.
A very noteworthy circumstance concerning this tribe, as well
as the next, relates to the results of isolation on the islands off
the coast of California, a large proportion of the forms occur-
ring there being generically different from those of the main-
land. The genera, which form two distinct subtribal groups
clustering about Eitsatttis and Coniontis respectively, may be
defined as follows : —

Elytra widely embracing the sides of the body, the epipleuras variable ;
anterior tibias with everted external angle at apex ; basal joint
of the anterior tarsi long, slightly dilated at or toward tip ; pro-
thorax always prolonged backward at the sides and enveloping
the humeri; scutellum nearly obsolete. [Subtribe Eusatti] — 2

Elytra narrowly embracing the sides of the body, the epipleurae con-
stantly narrow and occupying the entire inflexed part; anterior
tibiae with uneverted or very feebly everted external angle at
apex, the basal joint of the anterior tarsi short, obliquely truncate
at tip ; prothorax variable at base but generally more truncate;
scutellum generally well developed. [Subtribe Coxiontes] — 8

56 CASEY

2 — Epipleurje very wide, occupying the entire inflexed sides of the

elytra ; intercoxal process of the abdomen acute ; antennae slender,
dilated moderately toward apex. Habitat insular. [Type Eu-
sattus robustus Lee] Nesostes

Epipleurae narrow, occupying much less than the entire inflexed sides
of the elytra, except sometimes at the immediate base 3

3 — Epipleura abruptly dilated at base to the lateral limits of the

elytra; intercoxal process of the abdomen acute; body oblong or
oblong-oval, moderately convex 4

Epipleurae gradually becoming wider basally, sometimes extend-
ing to the sides of the elytra at base ; sides of the elytra always
obtusely rounded in section, never acutely margined ; antennae
slender, dilated apically, the joints loose and obtriangular to sub-
cylindric 7

4 — Sides of the elytra acute and reflexed almost to the apex ; elytra

conspicuously sculptured , 5

Sides obtusely rounded in section as in Eusattus 6

5 — Antennae less slender, compressed, more glabrous and more com-
pact, gradually and more feebly enlarged distally, the joints ob-
trapezoidal, the subapical transverse, the eleventh narrower than
the tenth; tarsi long and slender; femora and propleurre gla-
brous. Habitat Sonoran of the Lower Rocky Mountains. [Type
Zophosis reticulata Say] Discodemus

Antennas slender, the joints obtriangular, gradually wider distally, the
last joint as wide as the tenth and longer ; tarsi shorter and stout ;
femora and propleurae with bristling hairs. Habitat Lower Cali-
fornia. \TyT^e Eusattus erosus Horn] Megasattus

6 — Elytra without coarse sculpture or punctuation of any kind, sub-
opaque. Habitat Lower California. [Type. Eusattus Icevisl^ec.'].

Eusattodes

7 — Intercoxal process of the abdomen acute, the coxae throughout nar-
rowly separated ; body oblong-oval to parallel, moderately convex ;
propleurag with or without bristling hairs. Habitat widely ex-
tended in the Sonoran and Pacific regions. [Type E. difficilis
Lee.]. = Conipittus Lee Eusattus

Intercoxal process obtuse, the coxze more widely separated throughout ;
body more broadly rounded, very convex, the sculpture more muri-
cate ; propleurce with more conspicuous bristling hairs. Habitat
as in Eusattus. [Type Eusattus tutiricatus Lee] ...Sphaeriontis

8 — Scutellum well developed, triangular 9

Scutellum subobsolete, very minute, triangular 12

9 — Posterior angles of the prothorax strongly posteriorly produced,
as in the preceding group of genera; palpi more elongate than
usual ; last three joints of the antenntc moderately dilated ; basal
joint of the anterior tarsi rather longer than the next two com-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 57

binetl, obliquely prominent internally at tip, two to four transverse,
rapidly diminishing in size. Habitat insular. [Type Conioutis
lata Lee. ] Coniontides

Posterior angles of the prothorax feebly produced posteriorly, the
thoracic base frequently subtruncate ; anterior tarsi as in Co7i-
ioiitidcs I o

Posterior angles not at all produced, the thoracic base transversely sub-
truncate II

10 — Eyes transversely reniform, entire, moderately coarsely faceted

as usual ; body larger in size as a rule. Habitat widely extended
from eastern Colorado to British Columbia, San Diego and the
islands off the Pacific coast. [Type C viatica Esch.].

Coniontis

Eyes completely divided; body smaller, the legs and antennjB shorter.
Habitat nearly as in Cottiontis^ excepting the immediate coast of
the Pacific. [Type Cofiiontis obcsa Lee. J Coniontellus

1 1 — Body small in size, oblong, the eyes as in Coniontis ; legs longer,

the tarsi long and filiform, the four basal joints of the anterior
subequal and slightly elongate, the basal joint of the posterior
relatively much shorter than usual, but little longer than the next
two combined ; mesosternum scarcely so much as impressed on its
anterior slope ; palpi and antennae elongate and slender; frontal
margin of the head more nearly as in Eusatttis. Habitat north-
ern Pacific coast regions. [Type C. rectus Csy.] Conisattus

12 — General habitus as in Coniontis but muricately and generally

densely punctured, the front and eyes similar ; base of the protho-
rax feebly and very gradually produced backward at the sides;
legs, antennae and palpi as usual; anterior tarsi moderate, the
basal joint as long as the next two combined, strongly and rapidly
dilateti and very obliquely, sinuously truncate at apex, the inner
angle pronounced and prominent though obtuse, the inner surface
apically impressed, joints two to four subequal, feebly transverse.
Habitat insular. [Type C. punctulata Horn] Coelotaxis

The only one of the above genera not represented before me
at present is Eusattodes, founded upon a very rare species in-
habiting the lower extremity of Lower California near Cape San
Lucas. Coelotaxis makes the closest approach to the Coelini in
sculpture, retraction of the scutellum and form of the first ante-
rior tarsal joint, but is a true Coniontid nevertheless. Coni-
pinus Lee, was founded upon Eiisattus dubius, a very small
oblong convex species, but I cannot find any structural character
to distinguish it from the other members of the true Eusatttis,
except a more oval and less pointed form of the last antennal
joint. At first it seemed possible that its smooth surface and

Proc. Wash. Acad. Sci., April, 1908.

58 CASEY

sparse, even, muricate punctures might give it validity, but it is
connected in this respect so gradually, by way oi pt'oductus and
difficilis^ with the coarsely reticulate species of the convexus
type, that this attempt also proves futile and we are compelled
to suppress the name as a virtual synonym of Eusatitis. In Dr.
Horn's later tables of £usaitus, which included all of the sub-
tribe Eusatti as defined above, he gave far too much weight to
the margining of the prosternal lobe, bringing thereby into jux-
taposition species that have no close mutual relationship what-
ever; this is especially noticeable in his latest table (Proc. Cal.
Acad., Ser. 2, IV, p. 423). Although wholly without signifi-
cance in delimiting groups, the beaded margin of the prosternal
lobe is useful in defining species if interpreted broadly, due
allowance being made for a certain amount of variability, or
more especially accidental malformation. It becomes particu-
larly valuable in the great genus Contontis, so deficient in
marked structural diversity, as will appear subsequently.

It is interesting to note, in view of the many points of resem-
blance between the Coniontinae and Zophosini, that one of the
first of our species to be described should have been assigned to
the genus Zophosis by Say, presumably upon descriptive evi-
dence only. It may also be observed in connection with state-
ments made above, that more than a fourth of the genera of
Coniontini are confined exclusively to the islands off the coast
of California.

Nesostes n. gen.

This genus is represented at present by one or two species,
among the largest of the entire subfamily and possessing many
peculiarities of structure and sculpture. The epipleuraa are
very wide and almost even in width in about basal half, be-
tween the simple inner margin of the inflexed part of the
elytra and the acute and very narrowl}' reflexed lateral mar-
gins, and the elytral sculpture is notably different from any-
thing occurring elsewhere. The type may be described as
follows : —

Body broadly oblong-oval, convex, glabrous, with a few fulvous hairs
near the external margin of the hypomera posteriorly and on the
epipleuraj, except externally, deep black, the upper surface aluta-

REVISION OF TENEHRIONID SUBFAMILY CONIONTIN^ 59

ceous ; head rather finely but strongly, moderately closely punc-
tured; antennie rather slender, moderately dilated apically, the
third joint but little longer than the fourth ; prothorax about tvvoand
a third times as wide as its median length, minutely and sparsely
punctured throughout, the punctures but little more distinct or
closer laterally, simple, tlie sides subexplanate, with the edges
heavily beaded, the bead extending entirely across the apex, but
becoming very feeble medially; scutellum obsolete; elytra but
little longer than wide, nearly three times as long as the prothorax,
the sides arcuately converging in about apical half, the apex ob-
tusely ogival ; surface nearly smooth, minutely, remotely punctate,
the punctures slightly larger but only slightly less remote though
muricate laterally and apically, the interspaces toward apex be-
coming closely, evenly granulose and polished; anterior tibiie
strongly produced and angulate externally at apex, the basal joint
of the anterior tarsi less elongate than usual, but little longer than
the next two joints combined; prosternum alutaceous, the process
thickly margined and impunctate. Length 16.0-16.5 mm. ; width
10.5-11.omm. Island of San Clemente robustus Lee.

A — Form more oblong-subparallel and narrower than in robustus,
similarly sculptured, the sides of the prothorax more rounded
and relatively more converging anteriorly and more parallel
behind the middle ; elytra more gradually and less obtusely sub-
ogival behind; antennas less slender; anterior tibiae a little
shorter, relatively somewhat broader at tip ; marginal bead of
the prothorax not subequal in thickness throughout, as in
1-obustus^ but still thicker apically than basally. Length 17.0
mm. ; width 10.5 mm. Island of San Clemente.

postremus n. subsp.

In the subspecies -postremus, which is described from a single
female specimen, the form is obviously different from that of the
more evenly subelliptic and broader robustus and there are
several other minor differences as enumerated above ; it micrht
be added, also, that the hind angles of the prothorax are more
abruptly acute and more prolonged posteriorly. The abruptly
polished area on the posterior declivity of the otherwise sub-
opaque elytra is a rather remarkable character, peculiar to this
genus.

Discodemus Lee.

The components of this genus are moderately numerous but
so closely allied among themselves as inevitably to give rise to
discussion concerning their true status. I have given them spe-
cific rank, by no means confidently, although ihey are recog-
nizably definable and do not seem to intergrade, in fact consti-

6o CASEY

tilting a t3^pical example of divergence due to local environment
in what may possibly be a comparatively recently evolutionized
type. The genus is well represented in almost every collection
and nothing further in way of description is called for at pres-
ent, except to draw attention to the form of the antennae as a
generic character hitherto overlooked. The body is smaller,
more oval and much less convex than in the preceding genus
and the basal joint of the anterior tarsi is longer, being almost as
long as the next three combined ; the outer apical angle of the
anterior tibiae is not so strongly everted but is much more
acute at tip; the scutellum is similarly obsolete. The specific
or subspecific modifications known to me may be outlined as
follows : —

Body moderately convex, stout, very obtusely rounded behind, with a
short obtuse sutural projection 2

Body narrower, more depressed, the sides more gradually, arcuately
converging behind to the subacute apex 7

2 — Upper surface of the body only partially opaque, the elevations of

the elytra rather strongly shining 3

Upper surface opaque throughout, the elevations of the elytra at most
alutaceous , 6

3 — The irregular anastomosing depressions forming broad lines be-

tween the narrower, feebly elevated and irregular subelevated
lines of the elytra, deep and better defined by the small asperulate
and minutely setigerous punctures along their anterior margins. .4
The anastomosing depressions very shallow, small in size and but little
less shining than the remainder of the surface, the entire upper
surface more shining than in any other species 5

4 — Form oblong-suboval, convex, deep black, tlie head dull and dis-

tinctly micro-reticulate, the pronotum more feebly and less dull,
the former finely but distinctly, not densely punctate, the latter
very minutely sparsely so throughout and only a little more dis-
tinctly or less sparsely laterally, the apex circularly emarginate
and evidently more than half as wide as the base, the sides broadly,
evenly rounded, the surface broadly explanate toward base, nar-
rowly so anteriorly, the marginal bead fine and almost completely
obsolete, the base broadly bisinuate and about two and a fourth
times as wide as the median length ; elytra but little longer than
wide, feebly inflated, the broadly arcuate sides becoming rapidly
converging and more arcuate in apical two-fifths, the apex very
obtusely ogival, the reticuliform erosions deep and opaque, very
confluent and irregular, the punctures on their anterior edges fine
and only moderately distinct, feebly asperate, the six discal ridges
on each elytron subcqual and rather narrow, irregular ; presternum

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 6l

finely, sparsely punctate, the process margined at the sides but
only at its base between the coxa^ ; basal joint of the hind tarsi as
long as the entire remainder. Length 12.0 mm. ; width 7.8 mm.
New Mexico. [= Zophosis reticulata Say] reticulatus Say

Form and general characters nearly similar but more broadly oval and
less oblong, the eroded depressions of the elytra large, less nu-
merous and less anastomosing, the interspaces and longitudinal
lines shining, the erosions deeper than in any other species and
more abruptly defined b}' transversely irregular lines of close-set
and strongly muricate punctures; inflexed sides of the elytra
rather coarsely but sparsely punctured, the punctures much more
distinct than in reticulatus. Length 1 2.8-13.8 mm. ; width 8.7-
9.5 mm. Texas (near El Paso) corrosus n. sp.

5 — Form broadly oblong-oval, inflated posteriorly, strongly convex,
deep black, the anterior parts alutaceous ( 9 ) or opaque (cJ*), the
head and pronotum very minutely and sparsely punctate, the
former much more finely than in reticulatus ; sides of the latter
rather broadly, subequally explanate throughout (9) or scarcely
at all explanate (cf ) ; elytra shorter than in any other species,
much shorter than wide in both sexes, the longitudinal elevations
very faint and ill-defined, stronger in the male, most of the sur-
face rather strongly shining, the muricate punctures finer, less
close-set and not forming such distinct transverse lines as in cor-
rosus; inflexed sides of the elytra only very finely, remotely and
indistinctly punctulate. Length 10.8-13.5 mm. ; width 7. 5-8. 6
mm. Arizona (Grand Canyon of the Colorado), — T. Mitchell
Prudden brevipennis n. sp.

6 — Body oblong-oval to oblong, convex, deep black, dull and opaque
in lustre; head and prothorax alutaceous, faintly and sparsely
punctulate as in brevipeiitiis^ the sides of the latter more narrowly
and more concavely explanate, the reflexed part subequal in
width throughout; elevated parts of the elytra strongly alutaceous,
the erosions rather deep, irregularly confluent, the asperulate
punctures fine, sparse and moderately distinct but not defining the
erosions, the six longitudinal lines on each elytron unusually evi-
dent, especially in the male ; elytra unusually elongate, evidently
longer than wide in both sexes, the inflexed sides sparsely and
sometimes distinctly punctate. Length 11. 8-13. 8 mm.; width
7.6-9.7 mm. Arizona (San Bernardino Ranch, Cochise Co.),
— F. H. Snow elongatulus n. sp.

Body oblong-oval, smaller than in elongatuhis and less convex, more
abbreviated, the lustre subopaque, the prothorax more trans-
verse, with the apex not more than half as wide as the base, less
concavely explanate at the sides and sometimes sensibly more
broadly toward base; elytra about as long as wide, tlie elevated
lines less distinct, the eroded foveas large, rather deep, impressed
and anastomosing, not at all defined by the small punctures,
which are sparse and not definitely muricate, usually hardly

62 CASEY

visible, the inflexed sides more shining, variably but feebly and
sparsely punctulate. Length lo. 5-1 2.8 mm.; v^'idth 7.4-9.0
mm. Arizona (more especially southern) depressulus n. sp.

Body oval, rather smaller than in depressulus and much more convex,
subopaque throughout above, the elytra with sericeous lustre ;
head rugulose anteriorly, with the margin more reflexed than
usual ; prothorax less transverse, nearly as in reticulattis but more
narrowed from base to apex, the sides evenly and concavely
explanate; elytra as long as wide, inflated, the erosions small,
feeble and isolated, more punctiform, the small punctures fine
and very feeble, scarcely muricate and clustered in small groups
of two to four at the erosions, the general surface almost smooth,
the alternate elevated lines obsolete, leaving three verv feeble
lines on each elytron, the inflexed sides rather opaque and scarcely
punctate. Length 11.6 mm.; width 8.0 mm. Arizona, —
Warren Knaus subsericeus n. sp.

7 — Form much more narrowly elongate-oval than in any of the pre-
ceding and gradually pointed behind, deep black, subopaque;
head flat throughout above, moderately sinuate at apex, finely
and sparsel)' punctate ; antenna stout and strongly compressed as
usual ; prothorax barely two and one-half times as wide as its
median length, the sides evenly converging and arcuate from base
to apex, the latter somewhat more than half as wide as the base,
minutely, sparsely punctate, slightly less sparsely and less obscurely
toward the sides, which are moderately explanate, with a feeble
thickened bead ; elytra fully three times as long as the prothorax,
evidently longer than wide, very feebly subcostulate and with
large subisolated and feebly impressed opaque areas suturally,
which completely coalesce laterally, forming a broad opaque
irregular area bestrewn with very small muricate punctures, the
side margin with a bilaterally symmetric sinus, corresponding with
an epipleural impression, slightly behind the middle, — a character
which is probably accidental in the single specimen from which
the description is drawn; — under surface and epipleunt less
opaque, minutely, sparsely punctulate. Length 1 1.7 mm. ; width
7.8 mm. Colorado (Fremont Co.), — Warren Knaus.

knausi n. sp.

The male is usually a little smaller, narrower and more par-
allel in form than the female, but in brevipennis it is very much
smaller and rather more dilated than the female. Individuals
appear to be more abundant than in any other genus of the
Eusatti and good series of several species are before me.

Megasattus n. gen.

Two species, one of which is the largest known North Amer-
ican member of the subfamily Coniontinie, are the only repre-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 63

sentatives of this genus thus far discovered. While allied more
closely to Discodemus than to any other, it differs markedly in
antennal structure, the antennae being notably slender basally
and decidedly dilated distally, in the form of the lateral margins
of the elytra, which are not evenly acute but acutely rounded
in section, sculptured and uneven and not smoothly cariniform
and in the more parallel and oblong form of the body, with the
prothorax not widest at base as in Discodemus but slightly
behind the middle. It also inhabits a very different faunal
region, which amply accounts for the present structural diver-
gencies, though apparently springing from the same archetype
at no very distant time in the geologic past, as indeed did all
these allies of Eusatttis. The species may be defined as
follows : —

Form oblong, strongly convex, deep black, glabrous, the head evidently
and the prothorax very feebly alutaceous in lustre, the elytra
polished except in the erosions, the latter being opaque ; head
rather strongly, moderately closely punctate, the anterior margin
slightly reflexed, moderately sinuate at the middle and feebly
emarginate laterally; antennae a little longer than the prothorax,
the third joint much the longest, the tenth obtrapezoidal, slightly
transverse, broadly angulate at tip, the eleventh as wide as the
tenth and longer, acutely angulate ; prothorax two and one-half
times as wide as its median length, the deep apical sinus trans-
verse medially, the angles prominent but rounded, the basal
angles rather abruptly but only moderately produced posteriorly,
acute, the lateral sinuses of the base deep, relatively small, the
median lobe very broad; sides evenly rounded, strongly, moder-
ately thickly beaded, narrowly explanate ; punctures fine and
sparse, becoming rather close, larger and muricatulate laterally;
elytra decidedly longer than wide, obtusely ogival behind, feebly
dilated, the sides slightly arcuate, each with two distinct obtuse
ridges, the intermediate ridges very feeble, the surface pitted with
numerous small deep irregular and isolated erosions, with their
anterior margins rendered very rough and jagged by the rather
strong and very asperate, minutely setigerous punctures, the
flanks just above the acute side margins more deeply and con-
tinuously eroded and opaque, with the included punctures less
asperate ; inflexed sides asperately punctate ; prosternal lobe
dilated and spatulate apically, laterally margined only along the
coxas ; anterior tibiae with the external apical projection very
acute. Length 19.0 mm.; width 11. 5 mm. Lower California.

erosus Horn

Form rounded-oval, convex, black, subopaque, the prothoi-ax finely and
sparsely punctured medially and densely though very faintly

64 CASEY

granulate laterally, the margin not thickened; elytra subcostate,
the intervals coarsely punctured and with small, smooth and
rounded tubercles; inflexed sides densely and coarsely punctured.
Length 16.5 mm. ; width 10. o mm. Lower California.

costatus Horn

Costatns is unknown to me and the description here given is
essentially a copy of the original ; it occurs in the same locality
as erostis. The single male specimen representing the latter
species in my cabinet and described above has not been com-
pared with the original type, but appears to be an authentic

example.

Eusattodes n. gen.

The chief distinctive character of this genus seems to be the
abruptly and basally dilated epipleurae, the latter at base extend-
ing laterally to the sides of the body, as in the preceding genera,
but here combined with rounded Etisatttis-X\\i^ sides of the elytra
in section, without trace of the acute margin there prevailing.
It also differs from all except Eusatttis seciitus in the completely
impunctate integuments and probably in several other charac-
ters, which must remain unknown to me for the present, in the
absence of typical examples. The t3^pe and only described
species is the following : —

Body oval, convex, black, subopaque above and beneath, less rounded
in outline than in Discodemus, with the sides of the elytra almost
straight from the base to beyond the middle and nearly parallel ;
upper surface entirel}' without sculpture; elytra slightly inflated
posteriorly, impunctate, suddenly declivous at the sides; thoracic
angles posteriorly produced, the anterior tibiiE obliquely prolonged
at the outer angle, which is rounded at the tip. Length 14.0 mm. ;
width S.o mm. Lower California (Cape San Lucas).

laevis Lee.

According to Horn the prosternum is not margined in Icevis
and the tip of the process is rounded and smooth.

Eusattus Lee.

This is a large genus, embracing considerable variety in the
form and sculpture of the body, but the epipleurce throughout
are virtually constant in extent and configuration, this being one
of the strongest of the reasons inducing me to believe that these
radical epipleural modifications must be given full generic weight

REVISION OF TENKBRIONID SUBFAMILY CONIONTINyE 65

and not lightly passed over as advocated by Horn, the latter
author even goinof so far as to advance such a trivial character
as the margination of the prosternal lobe over epipleural struc-
ture in inserting Icevis between the mutually congeneric species
■politus and dubiiis in his latest table (1. c, p. 423). The species
of Eusattiis are widely distributed from western Kansas and
Oregon to Lower California and are smaller in size than in any of
the preceding genera ; those known to me either actually or by
description may be recognized by the following characters : —

Elytra absolutely impunctate as in Eusattodes Icevis. Form oblong-
oval, nearly as in dtibuis^ black, moderately shining ; head opaque,
sparsely punctate, the clypeus nearly entire ; prothorax rather more
than twice as wide as long, widest at base, the side arcuate, the
margin not fimbriate and the hind angles acutely prolonged pos-
teriorly; surface convex, completely impunctate; elytra smooth,
impunctate, the epipleurae gradually and but little wider at base,
absolutely smooth ; presternum coarsely punctured between the
coxie, the tip of the process rounded and with a distinct marginal
bead; abdomen very sparselv and finely punctate, shining.
Length 9.0-10.0 mm. Lower California (El Taste and vSan Jose
del Cabo) secutus Horn

Elytra with sparse but evident, more or less muricate punctures 2

Elytra with larger, subconfluent to small and irregular but isolated,
punctiform erosions, without evident muricate punctures, and
having uneven, usually elevated impunctate lines; prosternal
process never margined apically 14

2 — Form oblong-elongate, the elytra smooth or nearly so, with the

punctures small or moderately small in size; prothorax in the
male very transverse, wider than the elytra 3

Form oblong or oblong-oval, much less elongate, the clypeus never
so distinctly emarginate at the middle of the rounded sides, at the
ends of the oblique sutures, as in the preceding group.... 9

3 — Body small in size, strongly convex, the elytra very smooth, the

punctures very small, sparse, evenly but irregularly distributed
throughout, generally becoming distinctly muricate only toward
the sides; tarsi slender. \_= Conipmt(sl^ec.~\ 4

Body much larger, relatively less convex, the elytra nearly smooth but
sometimes with vestigial longitudinal areas of greater elevation,
the punctures less minute and strongly muricate throughout, with
longitudinal and very feebly defined lines, corresponding with
those of greater convexity, in which they become sparse or want-
ing ; tarsi stouter 6

4 — Color black or piceous-black, the prothorax less transverse and with

less converging sides, generally but little more than twice as wide

66 CASEY

as long. Body oblong-elongate, very convex, shining to feebly
alutaceous, glabrous, the flanks sparsely puberulent ; head
strongly but not densely punctate, the apical margin only moder-
ately reflexed, sinuate at the middle, the lateral emarginations small
and feeble; prothorax with the apex nearly three-fourths as wide
as the base, the angles slightly acute and but little blunted, the
surface convex, minutely, sparsely punctate medially, much more
coarsely but rather sparsely and non-muricately laterally, the sides
narrowly explanate and finely beaded ; posterior angles small, not
greatly prolonged though acute ; scutellum wholly obsolete ; elytra
a fourth longer than wide, as wide at the middle as any part of
the prothorax ((J*) orwider(9)» the sides feebly arcuate, the apex
obtuse; surface almost vertically declivous posteriori}', with
small impressed and very sparse but distinct simple punctures, be-
coming but little larger or closer and only very feebly muricate on
the flanks; hypomera ciliate ; prosternum sparsely pubescent, the
process impunctate, pointed and unmargined at tip ; anterior
tibiae serrulate externally except apically, the external process long,
rather acute; epipleurae narrow. Length 6.3-7.9 mm.; width
3.3-4.7 mm. California (Kern Co.) nanus Csy.

Color piceous-black to pale piceo-testaceous, the prothorax much more
transverse, always much more than twice as wide as long, much
wider than any part of the elytra (^(^) or scarcely obviously so
(9), the sinuate apical margin of the head more strongly reflexed
and the lateral emarginations very distinct 5

5 — Body elongate, very convex, polished, glabrous though with a few
small hairs laterally ; head finely, sparsely punctate, densely near
the transverse suture, impunctate and polished in the apical con-
cavity behind the reflexed margin; prothorax two and one-half
times as wide as its median length, the apex two-thirds as wide as
the base, the angles and side margins throughout nearly as in
nanus, the sides much more converging from base to apex, the
punctures very minute even laterally and much less distinct;
elytra narrower and more elongate, almost a third longer than wide,
the sides less arcuate, the apex obtuse, the punctures fine but dis-
tinct, sparse, feebly asperate medially, distinctly so laterally; pro-
sternum with only a few short hairs, the process impunctate, ob-
tusely rounded or subtruncate and unmargined at tip ; anterior
tibise only feebly serrulate externally, the apical process moderate,
not so prolonged as in nanus. Length 7.9-8.5 mm. ; width
4.3-4.8 mm. Arizona (southwestern) and California (Colorado
Desert) dubius Lee.

Body nearly similar in form and size but less convex and less polished,
slightly alutaceous in lustre, rufo-piceous throughout ; head nearly
as in dubius but more angularly prominent at the sides before the
eyes, less strongly reflexed along the clypeal margiri and more
coarsely and densely punctured toward tlie transverse suture, the
punctures also extending further anteriorly toward the reflexed

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^E 67

margin ; prothorax similar but less narrowed at apex and more
parallel toward base, with the side margins more concave and
the apical sinus more transverse medially, the punctures small but
less sparse and more distinct, not differing much laterally, the
basal angles rather more prolonged backward, acute; elytra simi-
lar in outline but less convex, slightly narrower than the protho-
rax, the sides parallel and feebly arcuate, rounding at base within
the thoracic angles as usual, punctured nearly as in dubius but
somewhat less sparsely ; prosternal lobe broadly rounded at tip,
impunctate, very faintly and finely margined throughout its apical
contour; anterior tibiae crenulate externally, the external apical
projection unusually feeble, much smaller than in dubius. Length
8.0 mm. ; width 4.3 mm. California (Lancaster), — H. F, Wick-
ham oblongulus n. sp.

6 — Prosternal process not margined except along the coxal cavities... 7

Prosternal process margined at the rounded apex and also along the
cavities 8

7 — Explanate lateral margins of the pronotum deeply concave. Body

(cj*) cuneiform, moderately convex, black throughout, glabrous,
though with a few small hairs on the flanks, the hypomera roughly
punctured and ciliate, the femora f ulvo-pubescent ; upper surface
feebly shining, alutaceous ; head finely, not densely punctate, the
clypeus concave anteriorly, subangularly sinuate at the middle,
with the margin only moderately reflexed, the lateral emargina-
tions feeble ; prothorax very transverse, much narrowed anteri-
orly, the apex circularly sinuate and about three-fifths as wide as
the base, the angles less than right but blunt, the basal angles
only moderately produced and not very sharp, the sides arcuate;
surface minutely punctate, remotely so medially, more closely but
not much less finely laterally ; scutellum much larger and more
transverse than in the dubius group, though similarly confined to
the basal slope ; elytra about a third longer than wide, parallel,
with the sides feebly arcuate, not very obtuse at tip, much nar-
rower than the prothorax, the asperate punctures sparse and un-
even in distribution, closer but not much larger laterally ; surface
with some very feeble and irregular longitudinal impressed lines;
anterior tibite very finely, closely serrulate along the external edge,
the apical angle acute but only very moderately developed,
Length 10.5 mm.; width 5.8 mm. (cj*). Arizona (southwest-
ern). . , productus Lee.

Explanate lateral margins nearly flat. Body (cf ) rather more elon-
gate and less cuneiform, piceo-castaneous in color, alutaceous,
glabrous as in prodtictus and similarly punctured, except that the
punctures of the head and pronotum are even more minute and
indistinct and those of the elytra larger and stronger; head nearly
similar; prothorax similar but somewhat less transverse, scarcely
two and one-half times as wide as its median length ; elytra two-
fifths longer than wide, parallel and nearly straight at the sides.

68 CASEY

broadly ogival at tip, evidently naiTovver than the prothorax but
not so markedly as in prodiicttis, the surface with more numerous
lines devoid of punctures, the lines just visibly more convex than
the rest of the surface ; body (?) larger, notably stouter, more
parallel and more shining, the elytra fully as wide as the pro-
thorax or a little wider, with the sides more arcuate, also having
more numerous flavate hairs on the sloping sides of the pronotum ;
hypomera and anterior tibiae in both sexes nearly as in prodiic-
tus. Length ii. 4-1 1.8 mm. ; width 6.0-6.9 mm. (d^, 9). Cal-
ifornia (Colorado Desert) explanatus n. sp.

8 — Body (d^) nearly as in explanatus but rather more abbreviated

and somewhat smaller in size, feebly shining, piceous-black ; head
and prothorax nearly as in explanatus^ the antennae somewhat
shorter and thicker; elytra not more than a fourth longer than
wide, scarcely narrower than the prothorax, parallel, with the
sides feebly arcuate, the surface more even, without longitudinal
lines of greater convexity but with the punctures, which are still
stronger, closer and more coarsely muricate, especially on the
flanks, arranged in very irregular longitudinal areas of greater or
less approximation; anterior tibia) similar; basal joint of the hind
tarsi similarly not longer than the last two combined. Length
10.4 mm.; width 6.0 mm. (^J*). Arizona (southwestern).

vicinus n. sp

Body more elongate than in vicinus^ less black and more castaneo-
piceous in color, almost similarly sculptured; head very slightly
larger, similarly opaque and finely, sparsely punctate but having
the apical reflexed edge continuous and not interrupted at the
sinus ; prothorax slightly less transverse and having the basal lobe
not very broadly rounded but more narrowly rounded and promi-
nent than in any other species, otherwise as in vicinus^ except
that the apical angles are less blunt and more prominent, the sides
just behind them very faintly sinuate ; elytra decidedly more elon-
gate, almost a third longer than wide, more evidently narrower
than the prothorax than in vicinus but otherwise nearly similar,
the sides less arcuate and the punctures sparser ; prosternal process
narrower and more feebly margined at apex ; anterior tibiae less
abruptly and only moderately prominent externally at tip; intro-
mittent spicule stouter and more punctate, the coriaceous lobe be-
tween the lateral alae of the genital segment canaliculate along
the middle and not evenly convex as it is in vicinus. Length 1 1 .0
mm.; width 5.8 mm. (cJ^). Arizona (southwestern) .

lobatus n. sp.

9 — Elytral surface smooth. Body oval, very little longer than wide,

convex, black, shining; head very sparsely punctate in front, the
clypeus not incised [at the sides] ; prothorax neither explanate
nor ciliate at the sides, verv narrowly margined [in original de-
scription], without marginal line [Proc. Cal. Acad., 2, IV, p.
423], the base bisinuate, the hind angles obtuse, the surface

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^E 69

sparsely punctate at the sides, nearly smooth medially; elytra
with very line punctures, very remotely placed ; epipleurec grad-
ually wider from apex to base; prosternum coarsely punctured in
front, the process margined throughout; abdomen ver}' sparsely
and finely punctulate. Length 10. o mm. California (Sta.
Barbara) politus Horn

Elytral surface more or less uneven 10

10 — Prosternal process margined throughout 11

Prosternal process without trace of marginal bead, except basally
along the acetabula ; elytra with irregular impunctate lines as in
prodzictus 13

11 — Prothorax very broadly emarginate at apex, the sinus transversely

rectilinear medially. Body in form and size nearly as in difficilis
Lee, oval, feebly shining; head finely punctate, the clypeus
broadly emarginate, rounded at the sides, the margin narrowly
reflexed ; prothorax with the apical angles very obtuse, the basal
comparatively short and obtuse, the sides finely margined, the
surface narrowly and feebly impressed along the margin, broadly
rounded and converging before the middle, subparallel behind the
middle; disk alutaceous, sparsely and finely punctate; elytra at
the base slightly wider than the pi^othorax, without lateral margin,
feebly carinate, the surface alutaceous, with small sparse submu-
ricate granules; epipleurte gradually wider to the base, sparsely
punctate and hairy; prosternum sparsely punctate, the process
narrowly oval at tip ; abdomen sparsel}' punctate ; legs with short
sparse setae. Length n.0-11.5 mm. ; width 6.0 mm. California
(Los Angeles Co.) coquilletti Linell

Prothorax more narrowly, deeply and circularly emarginate at tip,
with the apical angles more acute and narrowly rounded, the
basal angles more posteriorly produced and acute, more evidently
embracing the humeri, the sides evenly converging from the base,
evenly and broadly arcuate; scutellum rather more evident than
usual ; elytra with more or less evident ridges which are scarcely
less punctate than other parts of the surface , 12

12 — Form broadly oblong-oval, moderately convex, deep black, sub-
glabrous above, ciliate at the sides, the femora with short and
stiffer hairs; upper surface opaque (c^) or moderately shining
( $ ) ; head rather strongly, moderately sparsely punctate, the
clypeus rather feebly reflexed, sinuate at the middle, the lateral
incisures obsolete ; prothorax scarcely two and one-half times as
wide as its median length, convex, moderately widely declivo-
explanate at the sides, with the marginal bead fine but elevated,
the punctures small but distinct, only moderately sparse, becom-
ing rather coarse but well separated and minutely setigerous later-
ally ; elytra barely longer than wide, the marginal line obtusely
protuberant at tip, equal in width to the prothorax in both sexes,
the sides feebly arcuate, the surface rugose, rather coarsely, evenly

70 CASEY

and not very closely punctate, the punctures sharply muricate and
minutely setigerous ; a few very irregular and more convex
longitudinal lines are evident; prosternum sparsely, shallowly
punctate ; anterior tibiae strongly but very obliquely prolonged
externally at tip, the external edge but feebly sinuate and finely,
inconspicuously spinulo-serrate ; epipleurse subconcave basally,
sparsely punctate and hairy throughout. Length S.o-ii.o mm. ;
width 4.8-6.6 mm. California (San Diego) difficilis Lee.

Form nearly as in difficilis but less broadly oblong-oval, opaque,
glabrous, the punctures less evidently micro-setigerous, the sides
beneath much less ciliate, the hairs of the epipleurie and femora
much shorter and less conspicuous; head nearly similar but with
the reflexed clypeal margin still more feeble and very shallow,
the lateral notches very small but more evident ; prothorax nearly
similar but with the sides rather less strongly converging from
base to apex, the punctures almost similar; elytra more evidently
elongate, as wide as the prothorax, parallel, nearly straight at the
sides, with more numerous and narrower longitudinal lines of
slight convexity, the surface less coarsely and more feebly rugose,
with much smaller, sparser and rather less strongly muricate
punctures ; prosternum densely and rather finely punctato-rugose ;
anterior tibiae nearly similar but more strongly sinuate externally
beyond the middle; epipleuras flatter, with the outer margin less
deflexed basally, rather less coarsely, very sparsely punctate ;
flanks of the elytra above the epipleurjE more prominently rounded
in profile than in other species except difficilis. Length 9.7 mm. ;
width 5.9 mm. California (Los Angeles Co.) ...agnatus n. sp.

13 — Body oblong, convex, rather narrow (c?) to very stout (?), dull
in the former, more alutaceous and with the elytra rather shining
in the latter, glabrous, ciliate at the sides, the femora with short
and sparse yellow hairs; head with moderately strong punctures,
gradually close toward the transverse suture, the apex of the
clypeus very feebly reflexed, not at all so near the median sinus,
the lateral notches obsolete; prothorax nearly as in difficilis but
less transverse, strongly narrowed with evenly arcuate sides from
base to apex, the apex evenly, deeply, circularly emarginate, with
rather pronounced and only narrowly blunt angles, the basal
angles and explanate sides nearly as in difficilis., the explanate
surface flat but diminishing in width at the basal angles, the fine
raised margin similar; surface convex, more minutely, sparsely
puncturetl, the punctures slightly larger and micro-setigerous
laterally though smaller and much sparser than in difficilis;
scutellum rather evident; elytra slightly elongate, equal in width
to the prothorax, with some feebly convex longitudinal lines, the
punctures, irregularly disposed for the most part between these
smooth lines, unusually coarse, numerous, moderately muricate
and micro-setigerous; prosternum sparsely punctate, especially
strongly so on the process, feebly and sparsely fulvo-pubescent ;

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 7 1

anterior tibia; nearly as in difficilis ; epipleuraa nearly similar but
with the flanks of the elytra above them evenly and broadly
rounded in section and not prominent. Length lo. 0-10.3 trim';
width 5.7-5.75 mm. (cf). Length 10.7 mm.; width 6.4 mm.
(?). Oregon compositus n. sp.

14 — Erosions small, more or less isolated and punctiform, unevenly
distributed, generally better defined than in the following group
but never muricate 15

Erosions of the elytra larger, generally confluent and poorly defined,
the general sculpture like that prevailing in Discodemtis iS

15 — Body oblong-oval and parallel in form, not ventricose even in
the f ema le 16

Body more rounded and ventricose in the female, the male generally
oblong-oval and more parallel 17

16 — Form stout, convex, glabrous, deep black, strongly alutaceous
or subopaque, the elytra of the female much more shining; hy-
pomera with a few short hairs posteriorly but otherwise not ciliate
at the sides nor on the femora; head finelv but distinctly punc-
tured, rather closely toward the suture, the clypeus rather broadly
sinuate, with broad but obvious lateral sinuses, the apex not
reflexed but with the surface broadly impressed behind the
rounded apical lobes ; prothorax distinctly more than twice as wide
as its median length, much narrowed and deeply, circularly emar-
ginate at apex, with the angles right and blunt, the basal angles
only moderately prolonged, acute and sharp ; sides moderately
converging and feebly arcuate to before the middle, then more
converging and arcuate; surface moderately concavo-explanate
laterally, narrowly anteriorly, more broadly basally, the bead very
feeble, the punctures very minute and inconspicuous even later-
ally ; elytra but little longer than wide, nearly straight at the sides,
the margin acute and reflexed at the humeri, thence disappearing
as usual from a dorsal viewpoint, very irregularly and shallowly
punctate, the flanks rather abruptly inflexed, the under part much
less punctate, the epipleurae nearly impunctate, j^oHshed and
glabrous ; prosternum finely, sparsely punctate ; anterior tibiae
strongly, acicularly everted at tip. Length 9.2-11.0 mm. ; width
6.0-6.9 i^'ii' New Mexico (Las Vegas) congener n. sp.

Form nearly as in congener but slightly narrower, deep black, gla-
brous, the anterior parts opaque, the elytra rather shining; head
minutely, indistinctly punctate, the clypeus very feebly sinuate
medially, the lateral notches subobsolete, the margin not at all
reflexed at any point but with the edge very slightly thickened;
prothorax shorter and more transverse but otherwise nearly similar,
the sides more strongly converging from the base and but little
more converging or arcuate anteriorly, the punctures extremely
minute and indistinct throughout, the basal angles much produced
and very acute; elytra nearly similar but more decidedly longer

72 CASEY

• than wide, tlie punctures smaller and very feeble, the inflexed
sides, prosternum, tihiie and epipleurae similar. Length 9.5 mm. ;
width 6.3 mm. Colorado acutangulus n. sp.

17 — Body oblong, small and alutaceous (d) or ventricose and with
the elytra rather shining ($ ), glabrous, deep black; head alu-
taceous, finely and rather sparsely but distinctly punctate, the
clypeus broadly sinuate medially, very obsoletely so laterally, not
reflexed at apex but broadly and very obsoletely impressed behind
the rounded lobes ; prothorax short and transverse, very strongly
narrowed from base to apex, the latter circularly sinuate, with blunt
angles as usual, the hind angles moderately produced, acute and
sharp, the sides explanate and nearly flat, rather widely behind,
very narrowly at apex, the bead fine but distinctly elevated, the
punctures sparse and very minute throughout, sometimes more
evident in slight impressions laterally; elytra scarcely longer than
wide, with rounded sides, and, at the middle, evidently wider
than the prothorax (?) or parallel and not wider (c^), the punc-
tures coarse and irregular (?) or smaller and feebler (d^), the
flanks, epipleurae and prosternum as in the two preceding species,
the anterior tibiae with a much smaller and less everted though
very acute external angle at apex. Length 8.5-10.0 mm. ; width
5.5-6.7 mm. New Mexico (Las Vegas). [Colorado, near
Long's Peak, — LeConte] convexus Lee.

Body evenly and broadly elliptical, convex, rather shining though
alutaceous, deep black, ghibrous; head nearly as in convexus
though smaller, with the apical sinus rather narrower and deeper ;
prothorax similar throughout but more evidently and rather less
sparsely punctate laterally; elytra about as long as wide, rounded
at the sides, the width at the reflexed humeri evidently greater
than that of the prothorax, the surface strongly convex and dif-
fering from any of the three preceding species in having no ele-
vated lines, the latter being traceable only by absence of punc-
tures, the latter small, feeble, rather sparse and very unevenly
distributed throughout, the flanks not very abruptly inflexed and
almost smooth ; epipleurte impunctate, more distinctly though
gradually wider from apex to base than in the three preceding
and similarly concave basally; prosternum finely, sparsely punc-
tate; anterior tibias as in convexus. Length 9.4 mm. ; width 6.0
mm. Colorado (north of Fort Collins), — Warren Knaus.

rotundus n. sp.

18 — Integuments rather shining to alutaceous in lustre 19

Integuments densely opaque throughout above 20

19 — Form elliptical, moderately elongate, convex, glabrous, deep
black, rather shining throughout; head minutely, not conspicu-
ously punctate, the clypeus broadly sinuate medially, obsoletely
at the sides, not reflexed but with the margin somewhat thickened ;
prothorax two and a fourth to iialf times as wide as its median

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 73

length, very strongly narrowed from base to the deeply and cir-
cularly sinuate apex, with the sides very evenly arcuate, the angles
rather less than right and but slightly blunt, the basal angles
much produced and acute ; surface scarcely evidently punctate
medially, minutely and sparsely so laterally, the margin flatly
explanate, rather broadly basally, very narrowly apically, the
bead fine and feebly elevated ; elytra slightly elongate, equal in
width to the prothorax, the sides nearly straight, the reflexed
humeral pai't of the epipleurse arcuately prominent ; surface with
stronger longitudinal ridges than in any other species, about six
on each elytron, the concave intervals very obscurely, coarsely and
confluently foveate and subopaque, becoming more shining and
more clearly punctate laterally, the flanks very abruptly inflexed
along an obtusely convex line, the under part sparsely punctate ;
epipleurae rapidly broader than usual from apex to base, with a
few fine punctures and very small hairs internally, the outer
edge much deflexed basally; hypomera with a few small decum-
bent hairs posteriorly. Length 9. 0-9. 8 mm. ; width 5.6-6.2 mm.
Arizona subnitens n. sp.

Form oval, subinflated posteriorly, larger and stouter, convex, deep
black, glabrous, dull and alutaceous, the elytra less strongly so;
head nearly as in subnite7is but more closely punctate; prothorax
nearly similar, except that the sides become more converging and
straighter anteriorly than near the base, the lateral explaniture
more concave, the apical angles more obtusely rounded and the
punctures near the sides much larger and closer, though feebly im-
pressed and rather indistinct ; elytra but little longer than wide
in the apparently female type, somewhat inflated, with broadly
arcuate sides, at the middle slightly wider than the prothorax,
more abruptly and veiy obtusely rounded at apex, the reflexed
humeral arcuation less prominent; surface with the longitudinal
lines less prominent and less shining, the opaque irregular fovese
of the intervals rather better defined, punctiform laterally, the
inflexed sides and epipleurae nearly similar. Length 10.6 mm.;
width 7.0 mm. Kansas turgidus n. sp.

20 — Body smaller in size, stout, oblong-oval, convex, deep black,
glabrous; head flat, minutely, not densely punctate, the clypeus
broadly sinuate medially, more feebly but evidently emarginate at
the ends of the oblique sutures, not in the least reflexed, thickened
or impressed apically, the sutures forming very fine and less
opaque unimpressed lines ; prothorax formed as in subnite?is, with
strongly converging, evenly and moderately arcuate sides through-
out, the explanate sides and basal angles similar, though with the
marginal bead very much thicker than in any other species of this
group, the apical angles less narrowly blunt, the surface so densely
opaque as to obscure all trace of punctuation ; elytra scarcely
longer than wide, as wide as the prothorax, the sides parallel and
straight, broadly rounding about the very obtuse apex in apical

Proc. Wash. Acad. Sci., April, 190S.

74 CASEY

two-fifths, the reflexed humeral part of the epipleurae narrow and
scarcely visible from above, not at all prominent; surface with
the longitudinal lines obsolete basally, becoming gradually distinct
posteriorly, opaque, the shallow opaque anastomosing foveae very
indistinct basally, becoming larger and more evidently defined
apically ; flanks rather gradually inflexed, the under side less
opaque but scarcely punctured ; epipleurae nearly as in the pre-
ceding species ; anterior tibiae only very moderately produced
externally at tip, very feebly sinuate externally ; under surface
more shining, alutaceous. Length 8.4 mm. ; width 5 S mm.
Kansas (Fort Dodge), — Warren Knaus peropacus n. sp.

Body similarly small in size but much narrower and more gradually
and acutely produced behind than in any other species, deep black,
opaque ; head flat, finely, sparsely punctulate, broadly sinuate at
apex, the sutures very fine ; prothorax nearly as in peropaais but
much less transverse and less explanate at the sides, being only
feebly declivo-explanate even posteriorly, the marginal bead much
finer and feebler; elytra fully a fifth longer than wide, at base
equal in width to the prothorax, the sides thence parallel nearly
to the middle, then gradually arcuate and converging to the
acutely rounded and prominent apex ; surface convex, with distinct
and close-set though feeble ridges which are wholly lost on the
flanks and toward apex, the intervals with very indistinct and
shallow anastomosing foveee, which are still more opaque than the
ridges; under surface rather shining though evidently alutaceous,
the prosternal process flat, evenly, circularly rounded at tip; basal
joint of the hind tarsi a little longer than in peropacus and about
as long as the entire remainder. Length 9.0 mm.; width 5.4
mm. (cf). Kansas (Logan Co.), — Warren Knaus... acutus n. sp.

There are three evident groups of species in this genus, com-
posed, first, of small forms allied to dubius, forming the Conipi-
nus group, inhabiting southern California ; second, the productus
group, including two distinct divisions represented by productus
and difficilis, depending upon bodily form and character of the
elytral sculpture, very satisfactorily connected by covipost'lus,
which has the parallel form, ver}'^ coarse elytral punctures and
rugose surface of difficilis and the longitudinal impunctate lines
and considerable sexual divergence characterizing productus^
explanatus and vicinus; this second group of the genus also
peculiar to the California fauna and extending only slightly into
Arizona ; thirdly, the last eight species of the table form a dis-
tinct group inhabiting the southern Rocky Mountain region,
extending into Mexico but never occurring in California. Se-
cuius,p>olitus and coquilletti are unknown to me, but it is inferred

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 75

that they are properly placed in the table from such characters
as have been published.^

Sphaeriontis n. gen.

In this genus the epipleurae have a form intermediate between
the unusually narrow type of Etisattus dubiiis and the broader
form, with the outer edge deflexed basally, represented by
Eiisattiis convextis and allies, being nearly similar to the devel-
opment occurring in ^rodtcctus, though never observable at the
sides of the body when viewed dorsally. The fringe of yellow
hairs at the apical and basal margins of the prothorax are much
more developed than in any other genus, and the sides of the
body are more conspicuously ciliate ; the femora, also, bristle
with longer fulvous hairs. The genus is isolated principally by
the very broadly oval form and great convexity of the body, the
more widely separated coxae, even surface of the elytra and the
more prolonged outer apical angle of the anterior tibiae. The
large and very broad scutellum sometimes makes a very shallow
sutural notch at the base of the elytra, but, as a rule, is invisible,
being covered by the base of the pronotum when the latter is
tightly closed upon the elytral base, as usual in the Eusatti.
The five known species may be defined as follows : —

Prosternal lobe not margined apically, the upper surface more or less
shining, the elytral punctures muricate and never more than micro-
setulose 2

Prosternal lobe margined continuously about its contour; sculpture
and vestiture variable.. 3

2 — Form broadly rounded, very convex, rather shining in both sexes,
deep black ; head finely rugose and opaque, with finer isolated

' Since this was written I have had an opportunity to study two of the original
cotypes of coquilletti, and have received an excellent photograph of the type from
the skillful hands of Mr. H. S. Barber, of the National Museum. It appears
that the transversality of the bottom of the apical thoracic sinus is by no means
so pronounced as might be inferred from the original description and that the
species is really closely allied to agnatus, from the same zoological region. It
differs in the more swollen reflexed humeri, causing the width of the elytra at
that point to be decidedly greater than that of the prothorax, as mentioned in the
description ; also in its slightly shorter and broader form, distinctly greater con-
vexity when viewed in profile from the rear, rather less feebly costulate and de-
cidedly more shining elytra, with the muricate punctures more distinct, and
finally in the more decidedly explanate sides of the prothorax.

76 CASEY

granules posteriorly, the clypeus rather narrowly and deeply sinu-
ate medially, rounded and with minute feeble emargination at the
sides, broadly and rather strongly reflexed except at the sinus;
prothorax very transverse, gradually though moderately narrow^ed
from base to apex, with evenly arcuate, very finely and feebly
beaded sides, the apical sinus broad, subtransverse at the bottom,
the angles moderately prominent, rounded, the basal angles pos-
teriorly produced but rather blunt; surface convex, alutaceous,
abruptly and lather widely plano-explanate at the sides, almost
equally throughout the length, the punctures fine and sparse medi-
ally, rather close, strong and setulose laterally, muricate through-
out though feebly so medially ; elytra about as long as wide,
gradually and evenly convexo-declivous posteriorly, the outline
evenly rounded from about basal third to the apex, which is very
obtusely subogival ; surface even, very convex, rather coarsely,
moderately closely and strongly muricate, each asperity with a
minute posterior puncture, micro-setigerous laterally, the asperi-
ties tending sometimes to arrange themselves in transverse irregu-
lar lines; under surface rather hairy, the epipleurs sparsely
punctate and fulvo-jDubescent, the abdomen nearly glabrous, finely,
sparsely punctulate. Length 10.5-1 2.0 mm. ; width 7-0-7-S mm.
Oi-egon muricata Lee.

Form globose, large in size as in jmiricata^ posteriorly subacute,
black, shining; head less densely rugose than in muricata^ the
prothorax deeply sinuate at each side of the base as usual, the
sides margined, the apex and base flavo-ciliate ; surface very
smooth, finely granulate at the sides; elytra with the granules
less elevated than in niuricata and transversely confluent ; epi-
pleurae very densely and finely punctulate. Length 11.2 mm.
California (Colorado Desert) dilatata Lee.

Form nearly as in 7}i7<ricata^ but very much smaller in size, similar
in color, lustre and distribution of the hairs and fringes; head
similar but less opaque and less completely rugulose, finely but
strongly punctato-granulose, densely toward the transverse suture,
sparsely so elsewhere; prothorax similar throughout, though
shorter and even more transverse, nearly three times as wide as
the median length, the minute and feeble punctures completely
wanting along the middle, finely muricate but sparse laterally,
the sides rather less broadly explanate ; elytra scarcely as long as
wide, similar in form and similarly sculptured though more ru-
gose, very convex and polished (?) or duller and more flattened
medially (cJ*), at basal third rather broader than the prothorax in
the former but not in the latter, where the basal angles of the
prothorax are more widely free ; under surface nearly similar, the
epipleurae finely, sparsely punctate, with the hairs very short and
inconspicuous. Length 8.0-8.5 mm. ; width 5.2-5.85 mm. New
Mexico acomana n. sp.

3 — Form oval, convex, black, moderately shining, slightly more

REVISION OF TENEBRIONID SUBFAMII.Y CONIONTIN^ 77

obtuse behind, the margin of the prothorax and the legs ciliate
with long yellowish hairs; head sparsely, finely punctate, the
clypeus deeply and broadly emarginate, with a moderately deep
incisure at each side, the entire front margin reflexed ; prothorax
more than twice as wide as long, much narrowed in front, the sides
arcuate, the side margin explanate, the hind angles slightly- pro-
longed but not acutely; surface smooth, with a few fine piliferous
punctures near the sides; elytra with minute, sparse, submuricate
granules, each with a short hair, the interspaces very minutely
alutaceous ; epipleurje gradually wider from apex to base, sparsely
punctate and ciliate; prosternum sparsely punctate, ciliate with
yellow hairs, the tip of the process narrowly oval ; abdomen very
sparsely, finely punctate ; legs ciliate with moderately long yel-
lowish hairs. Length 11.5 mm. Lower California (Big Canyon,
Tantilles JMts.) ciliata Horn

Form more oblong than in muricata, very much smaller in size and
more pointed behind, the sides arcuately converging from about
the middle of the elytra, similarly very convex, black ; head simi-
larly sinuate laterally, feebly emarginate and apically reflexed,
the surface subopaque, finely, sparsely and muricately punctate,
each puncture with a small decumbent yellow hair, the punctures
broadly wanting along the middle, narrowly so anteriorly ; pro-
thorax less transverse, barely two and one-half times as wide as
the median length, evenly though much less strongly narrowed
from base to apex than in any other species, the sides evenly
and feebly arcuate, finely beaded, similarly but rather less broadly
explanate, the apex more circularly emarginate, the angles simi-
larly bluntly rounded, the basal produced, slightly less than right
and not blunt; surface alutaceous, impunctate except near the
sides and narrowly along the basal margin, where the punctures
are rather sparse but distinct and muricate, each bearing a small
yellow hair; elytra as long as wide with rounded sides, before
the middle rather wider than the prothorax, convex, densely
opaque though even, not very coarsely, rather sparsely punctate,
the punctures but feebly muricate, each bearing a small and very
coarse, decumbent fulvous hair; under surface more glabrous
than usual, shining, the epipleuras finely, sparsely punctate, sub-
glabrous and polished ; tarsi very slender ; antennse only moder-
ately developed as usual in this genus. Length 7.6 mm. ; width
4.6 mm. Texas (Sta. Rosa), — H. F. Wickham...puberula Lee.

The species defined above, by compilation of the descriptions
of LeConte and Horn, under the name dilatata, appears to have
been wrongfully reduced to synonymy, the densely punctulate
epipleurse and more or less confluent granules of the elytra
amply distinguishing it from onuricata, aside from its radically
different habitat. Pubcriila is identified from description only,
but the specimen here described is probably typical ; it repre-

78 CASEY

sents a very distinct group in the genus because of the more

oblong form of the body and different outline of the prothorax,

as well as by reason of its opaque elytra, with pubiferous and

barely muricate punctures. The species described by Dr.

Horn under the name Eiisathts ciliatiis belongs without much

doubt to this genus and is defined above from published

characters.

Coniontides n. gen.

So far as known, this group of the true Coniontes is confined
to the islands off the coast of southern California ; it is allied
closely to Coniontis but has a very different facies due to the
marked prolongation backward of the thoracic angles, recalling
the Eusatti, and in every way similar to the form there prevail-
ing. There are no other very radical differences that I have
observed, excepting the more elongate maxillary palpi and the
apically more strongly narrowed and more deeply and circularly
sinuate apex of the prothorax, which, with the prolonged basal
angles, gives the few species thus far discovered a decidedly
peculiar habitus. The four species known to me may be dis-
tinguished as follows: —

Form broadly oblong-oval, moderately and evenly convex. Piceous-
black in color, rather more reddish beneath, the legs evidently
rufous, the antennae dark piceous, shining, the elytra alutaceous,
glabrous; head small, strongly though rather sparsely punctured,
prominent at the sides before the eyes; antennas long and very
slender; prothorax fully four-fifths wider than long, the sides
broadly rounded and parallel in basal third, gradually strongly
converging and straighter thence to the right and narrowly
rounded apical angles, the basal angles slightly blunt at tip; base
deeply sinuate laterally ; surface convex, evenly so to the line mar-
ginal bead, rather coarsely, deeply, moderately closely and evenly
punctured throughout from side to side; scutellum somewhat
punctate, shining; elytra a fourth longer than wide, equal in
width to the prothorax, the sides parallel and straight, arcuately
converging behind the middle to the broadly ogival apex, evenly
convex, with narrowly reflexed margins, the surface rather
strongly, moderately closely punctate discally, though less
coarsely so than the pronotum, the punctures becoming smaller
and more shallow though similarly separated laterally; under sur-
face with short, sparse and decumbent hairs, the prosternal
process broadly and obtusely rounded at tip, not margined and
rather finely though strongly, sparsely punctate. Length 11.3
mm.; width 5.9 mm. Island of San Clemente latus Lee.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 79

Form much more narrowly oval 2

2 — Body elongate-oval, shining, the elytra only faintly alutaceous,
black, the legs piceous-black, glabrous ; head only moderately
small, finely, sparsely punctate, the sides before the eyes not evi-
dently more prominent than the posterior canthus and very much
less so than in latiis ; antennie shorter and much thicker; pro-
thorax much less transverse, about two-fifths wider than long,
the sides evenly converging, evenly and feebly arcuate from base
to apex, the latter two-thirds as wide as the base, with the angles
only very narrowly rounded, the basal slightly blunt at tip; sur-
face evenly convex, finely, rather sparsely and equidistantly
punctate throughout, rather less finely toward the sides, the lat-
eral bead very fine ; scutellum broadly triangular, polished, slightly
punctulate toward tip ; elytra fully two-fifths longer than wide,
otherwise nearly as in latus^ except that the rather sparse punc-
tures are much finer and subequal to those of the prothorax medi-
ally, though becoming much finer but not at all closer laterally ;
under surface with much more minute and scarcely observable
hairs, the very obtuse prosternal process similarly unmargined.
Length 10. i mm. ; width 5.0 mm. Exact habitat unrecorded.

finitimus n. sp.

Body nearly similar but noticeably cuneiform posteriorly, glabrous,
shining throughout above, the elytra not duller, piceous-black, the
elytra, under surface and legs rufescent ; head very small, strongly,
rather closely punctate, less prominent at the sides than in latus
but more so than xnjinitimtis^ noticeably more so than the poste-
rior canthus; prothorax more transverse, rather more than one-
half wider than long, the sides subevenly and strongly converging
from base to apex and evenly arcuate, more strongly so than in
ii7iitimus^ the angles nearly similar, the apex barely three-fifths as
wide as the base ; punctures only slightly less coarse and close
than in latiis^ stronger than '\x\Jinitinius^ even in size and spacing
throughout ; elyti^ fully two-fifths longer than wide, at base equal
in width to the prothorax, the sides thence feebly converging and
just visibly arcuate to about apical two-fifths, becoming gradually
thence strongly arcuato-convergent to the more narrowly ogival
tip, the punctures very fine, rather sparse and evenly distributed
throughout, minute laterally, very much smaller than those of
the pronotum, the contrast being much greater than in either of
the preceding species ; under surface finely, sparsely punctulate and
just visibly micro-setigerous, the prosternal process less obtuse but
wholly unmargined. Length 9.9 mm.; width 4.6 mm. Island
of Sta. Cruz insularis Csy.

Body somewhat cuneiform behind the prothorax, almost as in insularis
throughout, except that the prothorax is shorter and more trans-
verse and the punctures, especially those of the elytra, much more
close-set, deep black throughout, the legs piceo-rufous distally ;
head nearly as small as in insularis^ evenly convex, similarly

8o CASEY

evenly punctate and apically sinuate but having the sides before
the eyes much more prominent and very much more so than the
posterior canthus ; prothorax two-thirds wider than long, the sides
evenly converging and broadly, evenly arcuate from base to apex,
more strongly so than in insularis^ the apex narrower, not more
than half as wide as the base, the punctures not quite so large and
a little closer, the hind angles not quite so acute ; scutellum some-
what smaller and less broadly triangular ; elytra formed almost
exactly as in insiilaris but less shining, more uneven and with
the punctures not quite so small though distinctly smaller than
those of the prothorax and much denser than in insularis, simi-
larly finer and feeble laterally; prosternal process moderate,
rounded at tip, spatulate, finely and sparsely punctate ; basal joint
of the hind tarsi longer than in insularis^ being distinctly longer
than the remaining joints combined. Length 9.5-9.8 mm. ;
width 4.3-4. 8 mm. Island of San Clemente...clementinus n. sp.

It is my impression that the type oijinitimus was taken on the
island of Sta. Rosa, though this is little better than a surmise ;
it differs notably from iiisularis in form and punctuation, /w-
sularis is not properly a variety of latus as originally described.

Coniontis Esch.

This genus, by far the most extensive of the Coniontinfe, is
essentially peculiar to the Pacific coast of temperate North
America, extending to the eastward in the more northern lati-
tudes and thence southward along the Rocky Mountains to
southern Colorado. It has not yet been reported from north-
western Mexico and I have seen not a single example from New
Mexico and but one from Arizona, probably collected in the
extreme western part ; it appears to occur also on some of the
islands off the southern California coast, although probably as
a comparatively recent migrant. The body is oblong to oblong-
oval in form, convex, stout to elongate, strongly punctured to
almost smooth and glabrous to rather conspicuously setose.
The basal margin of the prothorax is never more than feebly
bisinuate, and the basal angles are always very much less prom-
inent posteriorly than in the preceding genus or the Eusatti.
The male is generally much smaller, less dilated and occasion-
ally somewhat more posteriorly attenuated than the female, but
does not differ much otherwise.

There is probably no group of Coleoptera that to the system-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 8l

atist has been more of a "bete noire," figuratively as well as
literally, than the genus Coniontis. This has arisen in great
measure from the preposterous extent of variation assumed to
affect the species, which for this reason, have been usually con-
sidered to be virtually undefinable. I concluded therefore to
test this variability by carefully collecting series of what were
indubitably single species, in each case from one isolated en-
vironment, and studying them carefully specimen by specimen.
The result of this study shows that the various forms, whether
species or subspecies, do not go beyond the average of the Cole-
optera in point of intraspecific plasticity, but are unusually sub-
ject to modification and consequent division into closely allied
forms by local conditions, which will account for the conclusions
formerly drawn from study of miscellaneous material. The
genus, being distributed over a greatly extended and climatic-
ally varied geographic region, is naturally composed of very
numerous forms which are structurally but slightly diversified,
— the usual condition in wingless and feebly migratory genera
of Coleoptera, — and, although perhaps more conventionally
distinguishable in the majority of cases as subspecies, varieties,
aberrations, anomalies and monstrosities of the German school,

1 feel unable with the material at hand to make these assign-
ments satisfactorily and therefore announce them as species for
the present. The genus may be divided primarily into the
three following subgenera : —

Metasternum flat, margined anteriorly; elytra always much longer
than wide 3

Metasternum longitudinally convex, declivous anteriorly, not mar-
gined ; elytra as wide as long 3

2 — Basal joint of the anterior tarsi stout, shorter, obliquely obtrian-

gular ; third palpal joint distinctly though not very gi'eatly shorter
than the fourth ; mesosternum between the coxse narrow, deeply
impressed on its anterior slope; eyes very short, strongly trans-
verse and reniform ; body more or less strongly convex.

Coniontis

Basal joint of the anterior tarsi slender and more elongate, not triangular ;
third palpal joint very much shorter than the last, the latter unusu-
ally elongate ; mesosternum a little wider between the cox£e, more
feebly impressed on its anterior slope ;• eyes much less transverse,
suboval, only feebly sinuate anteriorly; body very small, subde-
pressed, elongate-oval, the legs and antennte longer than usual.

Crypticomorpha

82 CASEY

3 — Body extremely sliort and convex, small in size, the eyes, mesoster-

num and basal joint of the anterior tarsi nearly as in typical Coji-
ionii's, the legs, and especially the antennie, shorter and stouter,
the last antennal joint relatively more developed ; maxillary palpi
with the third joint relatively somewhat more abbreviated though
not so notably so as in the preceding section Brachyontis

These divisions are extremely unequal in extent, the last two
being known at present by but a single species each.

Coniontis in sp.

For convenience the various components of this division may

be assigned to eight groups, defined on broad lines of general

facies as follows : —

Punctuation of the upper surface more or less sparse, fine and incon-
spicuous 2

Punctuation conspicuous, especially that of the elytra which is gener-
ally very coarse 5

2 — Body oblong, never much more than twice as long as wide 3

Body elongate, much more than twice as long as wide 4

3 — Body large in size I {abdomiJialt's^

Body small in size II (^opaca')

4 — Body parallel inform in both sexes Ill {viatica)

Body more or less distinctly cuneiform in both sexes but particularly
in the male IV {cuneata)

5 — Body decidedly more than twice as long as wide, oblong 6

Body shorter and generally more oval, never much more than twice as
long as wide 8

6 — Form subdepressed or much less convex than usual, the punctures

bearing each a more or less elongate, subdecumbent and very evi-
dent hair V {sjibpjibescens)

Form norm ally convex 7

7 — Punctures of the elytra moderate, bearing each a stiff erect and

sometimes very conspicuous seta VI {setosa)

Punctures generally larger and more conspicuous, nude or bearing each
a small inclined and less conspicuous coarse hair, the elytral
sculpture more or less rugose VII (^eschscholtzi)

8 — Elytral punctures never conspicuously setose, usually completely
nude, very variable in size, the sculpture vai'ying from compara-
tively smooth to extremely rugose VIII (^ova/is)

These groups also correspond fairly well with geographic
distribution.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 83

Group I. — Type abdoininalis.

This group comprises the largest species of the genus ; they
are oblong-suboval in form, very convex, generally sparsely and
feebly punctured above, variably so beneath and are peculiar to
the southern coast regions of California, but few occurring at
any great distance inland. Whenever the locality of the indi-
vidual species is more accurately known it will be mentioned
after the respective descriptions, the exact locality of many
being unrecorded ; they are, as known to me at the present
time, some seventeen in number, as follows : —

Abdomen more or less closely and strongly punctured and rugulose,
at least at the middle of the basal segment; prosternum strongly
punctured, the process margined throughout; body glabrous 2

Abdomen very finely, sparsely punctured and smooth throughout,
even at the middle of the basal segment 9

2 — Elytral punctures strong and distinct though rather sparse 3

Elytral punctures equally sparse, smaller and especially shallower and
much less evident 8

3 — Elytral surface even in convexity or having each about three feebly

suberoded impressed lines 4

Elytral surface very rugose, each elytron with about six more close-set,
very irregular and subcostuliform lines of slight elevation 7

4 — Prothorax transverse, evidently more than one-half wider than

long and apparently a little wider than the elytra in both sexes,
the sides more inflated and rounded than usual though similarly
rounding and converging anteriorly. Head and pronotum alu-
taceous, the elytra and under surface shining, the head strongly
and rather closely punctured, the surface feebly impressed at the
apical lobes, the anterior canthus of the eyes obtuse and equally
prominent with the posterior ; prothorax rather finely but strongly,
sparsely punctate medially, more closely and decidedly coarsely
laterally, the margin narrowly but strongly reflexed ; elytra barely
one-half longer than wide, parallel, obtuse behind, with the
margins rather abruptly and strongly reflexed, the surface feebly
rugose, strongly so at the apex, the punctures unusually coarse
for this section but sparse; prosternum densely and the abdomen
sparsely, coarsely punctured and rugose. Length 15. 0-17.0 mm. ;
width 7.3-S.o mm. Near Monterey. [Fort Tejon, — LeConte] .

abdominalis Lee

Prothorax about one-half wider than long, not inflated and not at all
wider than the elytra ; abdomen more finely, sparsely punctate
and rugulose only medially or toward the abdominal process at
base 5

84 CASEY

t; — Sides of the prothorax almost evenly converging and evenly, very
distinctly arcuate from base to apex, the latter rather narrowly,
deeply and circularly emarginate, three-fifths as wide as the base,
with the angles unusually broadly rounded. Body smaller and less
dilated than in abdominalis^ similarly deep black, alutaceous, the
elytra less strongly so ; head similar though finely and sparsely
punctate ; prothorax very minutely, sparsely punctate, still rather
finely and sparsely but more distinctly toward the sides, the mar-
gin moderately elevated ; elytra similar though rather more pointed
behind, the surface moderately rugose, strongly so near the tip,
the sides similarly reflexed, the punctures sparse, less coarse or
deep but distinct; prosternum closely and strongly punctured,
more finely and sparsely anteriorly, the abdomen distinctly but
not very coarsely punctate and with only traces of rugosity even
mediall}' throughout the length. Length 13. 8 mm. ; width 7.0
mm. Los Angeles Co tristis n. sp.

Sides of the prothorax parallel and straight or very nearly so, converg-
ing and rounded anteriorly, the apical angles not rounded though
slightly blunt 6

6 — Form broad, nearly as in abdominalis^ deep black, rather shining,
the head and pronotum barely at all alutaceous, the former rather
strongly and closely punctured, not evidently impressed at the
apical lobes, the anterior and posterior canthi subequally promi-
nent; prothorax fully one-half wider than long, the sides parallel
and straight, becoming more abruptly converging and broadly
arcuate only well before the middle, the apex nearly three-fourths
as wide as the base, broadly and circularly emarginate, the angles
right and with thickened bead, the surface very minutely, sparsely
punctate, with a feebly defined impunctate line medially, the
punctures becoming rather coarse and closer laterally, the margin
narrowly explanate medially, with the bead strongly elevated;
elytra as in the pieceding species but with the individual apices
rounded, more narrowly reflexed at the sides than in abdomiitalis^
the punctures only moderate and sparse, each with an extremely
minute silvery seta, the surface slightly rugose, strongly so at tip;
prosternum not coarsely and decidedly sparsely punctate through-
out, the abdomen with the punctures sparse, strong only medially,
strongly rugose only near the basal process; tarsi stout. Length
16.0 mm.; width 8.1 mm. Sta. Barbara strenua n. sp.

Form distinctly less broad, the single female being evidently a little
more than twice as long as wide, deep black, rather shining, the
anterior parts but feebly alutaceous; head rather sparsely, not
very coarsely but strongly punctate, deeply impressed behind
each apical lobe and with the margins before the eyes somewhat
deplanate, as prominent as the hind canthus; prothorax barely
one-half wider than long, the sides evenly and gradually more
arcuate and converging anteriorly, becoming parallel and straight
only in about basal half, the apex narrower, scarcely more than

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 85

three-fifths as wide as the base, deeply and broadly parabolic, the
angles slightly obtuse and not thickened; surface similarly punc-
tured and with the side margins almost similarly concave and
reflexed ; elytra almost similar but with the apical angles not so
individually rounded; presternum with well separated strong
punctures and having a densely, rugosely punctured area at the
base of the process; abdomen strongly, more closely punctured
and rugose medially. Length 16.0 mm.; width 7.8 mm. Near
Monterey gravis n. sp.

7 — Form oblong, twice as long as wide, convex, shining, scarcely at

all alutaceous anteriorly, deep black, glabrous ; head rather finely
but perforately, not densely punctate, not at all impressed an-
teriorly, the anterior canthus rounded, slightly more prominent than
the posterior ; antennae slender, joints three to seven much elon-
gated, the eighth thicker but much longer than wide, the ninth a little
longer than wide ; prothorax nearly three-fifths wider than long,
the sides parallel and straight in basal half, then gradually rounded
and strongly convergent to the broadly, circularly emarginate apex,
which is fully two-thirds as wide as the base, the angles right
and bluntly I'ounded ; surface rather finely, sparsely punctured,
becoming rather coarsely, more closely and conspicuously so
laterally, the side margins concave and reflexed ; elytra barely as
wide as the prothorax, scarcely one-half longer than wide, par-
allel, obtuse at tip, the sides concave and reflexed, the surface
coarsely and strongly rugose but polished, less rugose toward
base as usual ; prosternum rather finely but strongly, not very
closely punctate, the process very wide, obtuse, transversely con-
vex, less coarsely margined at tip ; abdomen strongly but not
very coarsely, rather sparsely punctured, very obsoletel}' rugulose,
alutaceous in lustre. Length 15.0 mm. ; width 7.3 mm.

rugosa n. sp.

8 — Form oblong, convex, twice as long as wide, deep black, slightly

alutaceous throughout ; head strongly, rather closely punctured,
finely and sparsely basally, the apical sinus rather shallow, broadly
rounded, the surface obsoletely impressed behind each apical
lobe; anterior canthus narrowly rounded, as prominent as the
posterior; prothorax fully one-half wider than long, formed as in
rugosa^ the punctures much more minute, almost invisible and
sparse medially, rather sparse and not coarse though distinct
laterally, the margin much more narrowly and feebly reflexed,
the fine bead strongly elevated ; elytra rather less than one-half
longer than wide, equal in width to the prothorax, parallel, ob-
tuse at tip, the surface alutaceous, not rugose but with about six
very feeble and broadly tumescent lines on each, coming together
in some broad rugosity at apex ; the punctures shallow and very
feeble suturally, deeper, closer and more distinct laterally, the
margin deeply concave and narrowly reflexed, each puncture with
a very minute silvery seta not projecting beyond its limits; pro-
sternum rather finely, strongly and sparsely punctate ; abdomen

86 CASEY

nearly smooth, the punctures sparse and fine for this division of
the group. Length 13.8 mm. ; width 6.9 mm. Sta. Barbara.

tenebrosa n. sp.

Cf — Prosternal process margined throughout as in the preceding sec-
tion. Body broadly oblong, black, alutaceous, the elytra shining,
glabrous ; head perfectly flat throughout, not impressed anteriorly,
rather finely but strongly and unusually closely punctate, the an-
terior canthus not more prominent than the posterior; prothorax
nearly three-fifths wider than long, as wide as the elytra, the sides
parallel in about basal half, very gradually arcuate and more con-
verging anteriorly, very narrowly reflexed and with strong elevated
bead, the apex broadly, circularly emarginate, fully two-thirds as
wide as the base, the angles obtuse and rather bluntly rounded, the
surface minutely, remotely punctate, the punctures still very small
and sparse and each bearing a minute hair laterally ; scutellum not
very broadly triangular; elytra a little less than one-half longer
than wide, obtusely angulate at apex, smooth, very feebly rugose
apically, each with three extremely obsolescent longitudinal ridges,
the punctures very fine and sparse throughout, the sides very finely
reflexed; prosternum with moderate, strong and rather close-set
punctures ; femora somewhat finely but strongly punctate, each
puncture with an obvious short decumbent hair ; tarsi not very
stout. Length 13.8 mm.; width 7.0 mm pectoralis n. sp.

Prosternal process not at all margined at its apex , 10

10 — Elytra glabrous, the punctures never bearing setae which project
materially beyond their own limits 11

Elytra sparsely pubescent, the hairs stiff, decumbent and small in size
but quite obvious 15

II — Elytra parallel, with nearly straight sides, obtusely angulate at
apex 12

Elytra somewhat inflated at the middle, wider than the prothorax, with
parallel and feebly arcuate sides, the apex more prolonged and
acutely angulate than in any other of this group 14

12 — Tarsi decidedly stout, the integuments strongly shining. Head
flat, not impressed anteriorly, rather finely, moderately closely
punctate, the anterior canthus rather more prominent than the
posterior ; prothorax transverse, nearly two-thirds wider than long,
subparallel basally and gradually rounded and more narrowed
apically (c?), or broader, somewhat inflated and with more
rounded sides (?), the margin finely reflexed, the apex broadly
emarginate, with more or less blunt apical angles; surface ex-
tremely faintly alutaceous, very minutely, remotely punctate, the
punctures still very fine, nude and sparse laterally though more
distinct; scutellum moderate; elytra scarcely one-half longer
than wide, smooth and polished, each having about three fine,
very feeble and irregularly sinuous eroded lines, sending out a few
still more obsolete vermiculate branches, the punctures extremely

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 87

minute and remote throughout, the sides very finely reflexed ;
prosternum finely, sparsely punctate ; abdomen smooth and pol-
ished, with extremely minute and remote punctulation throughout.
Length 14. 6-15.0 mm. ; width 7.0-7.25 mm. Southern Cali-
fornia integer n . sp ,

Tarsi moderately slender, the upper surface distinctly alutaceous, the
elytra generally somewhat shining 13

13 — Form very stout, convex, oblong, glabrous, alutaceous through-
out above, the elytra but little less dull than the anterior parts;
head not impressed, rather finely but strongly, closely punctate,
the anterior canthus scarcely as prominent as the posterior;
prothorax fully three-fifths wider than long, the sides rounding
and converging anteriorly, becoming almost straight and parallel
in about basal half, very finely reflexed and with elevated bead,
the apex broadly, circularly emarginate, two-thirds as wide as the
base, the angles obtuse but only slightly rounded ; surface very
minutely, sparsely punctate, still finely and sparsely though more
distinctly so laterally; scutellum moderate, not very broadly tri-
angular; elytra not quite one-half longer than wide, broadly an-
gulate at apex, smooth, feebly undulato-rugoseapically, very finely
and sparsely punctate, finely and narrowly reflexed at the sides;
prosternum finely, rather sparsely punctate, the abdomen minutely,
feebly and remotely punctulate. Length 13. 0-14. 7 mm. ; width
6.0-7.0 mm. Cab. Levette levettei n. sp.

Form oblong, rather less convex, smaller in size and narrower, the
ejytra more shining, the body throughout blackish-piceous in
color; head nearly similar throughout but more narrowly and
deeply sinuate at apex ; prothorax similar and likewise very
minutely and sparsely punctate, though with the lateral punctures
not nude as they virtually are in the preceding species but bearing
each a very small hair; scutellum similar; elytra narrower, about
one-half longer than wide, the surface similarly smooth and finely,
very sparsely punctate but with the punctures less fine and more
distinct than in levettei ; prosternum strongly, somewhat closely
but not coarsely punctate, the abdomen very minutely and sparsely.
Length 12.7—13.5 mm. ; width 6.3-6.6 mm. Cab. Levette.

picescens n. sp.

Form still narrower, convex, the elytra rather shining, deep black,
glabrous ; head finely, closely and strongly punctate, not impressed
anteriorly, the anterior canthus much less prominent than usual
and not extending laterally further than the posterior, the eyes
less sunken than in the preceding ; prothorax nearly similar but
rather less transverse, one-half wider than long, the sides finely
reflexed, the punctures extremely minute, remote and barely
visible, still very fine, sparse and nude but more distinct laterally;
scutellum rather small, triangular ; elytra about one-half longer
than wide, smooth, very obsoletely rugulose posteriorly, some-
times with feeble traces of longitudinal lines of greater convexity,

88 CASEY

finely reflexed at the sides, the punctures remote, fine, though
much larger and more distinct than in levettei and slightly more
so than \\\ pices cens ; prosternum finely, rather sparsely punctate,
the abdomen minutely and obsolescentlv so. Length 12. 5-13. 5
mm. ; width 5. 8-6. 35 mm. Probably San Diego Co.

elliptica Csy.

A — Form and sculpture nearly as in the preceding but narrower,
smaller in size and rather less convex, the elytra generally with
some excessively obsolete and broadly tumescent lines, deep
black, the elytra slightly more shining than in elliptica. Length
12. 0-12. 3 mm.; width 5.4-5.7 mm. Island of Sta. Catalina,
— Warren Knaus catalinae n . subsp.

14 — Body narrower and more convex than usual, glabrous, deep

black, strongly shining throughout ; head broadly sinuate at apex,
not impressed, finely punctured posteriorly, rather coarsely,
densely and subrugosely so anteriorly, the anterior canthus broadly
rounded, much more prominent than the posterior; prothorax
nearly three-fifths wider than long, the sides straight and parallel,
broadly rounding slightly before the middle, converging and
less arcuate apically, the apex broady and circularly emarginate,
almost three-fourths as wide as the base, with the angles right
and slightly blunt; surface very minutely, sparsely punctate, but
little less minutely or remotely so laterally, the edge strongly
beaded ; scutellum but little wider than long, moderate, tri-
angular; elytra rather more than one-half longer than wide,
convex, smooth, with three very feeble eroded lines apically,
where also there are some coarse creases, very minutely, remotely
punctate, the lateral margin finely reflexed ; prosternum not closely
punctate, strongly and rather finely so anteriorly, very minuetly
on the process; abdomen smooth. Length 13.5 mm.; width
6.2 mm. Kern Co laevigata n. sp.

15 — Form oblong, more parallel, moderately convex, deep black, only

slightly alutaceous throughout above, still more shining beneath;
head rather finely but strongly, moderately closely and subevenly
punctate throughout, deeply and broadly sinuate at apex, the lobes
feebly impressed, the anterior canthus very broadly rounded,
slightly more prominent than the posterior; prothorax three-fifths
wider than long, the sides feebly converging from the base nearl}''
to the apex and slightly arcuate, then gradually more arcuate and
convergent to the apex, which is broadly sinuate and two-thirds
as wide as the base, the angles right, narrowly rounded ; surface
minutely, sparsely punctate, gradually rather strongly, deeply
and much less sparsely so and micro-pubiferous laterally ; scutellum
large and broadly triangular, obtuse at tip ; elytra as wide as the
prothorax, fully one-half longer than wide, the sides straight, with
the margin more widely reflexed basally, the apex angulate ; sur-
face rather gradually declivous behind, nearly smooth though each
with about six very obsolete lines of greater convexity, the apex

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 89

feebly undulato-rugose, the punctures fine and remote, more dis-
tinct laterally, the fulvescent hairs not at all conspicuous; pro-
sternum more strongly, rather closely punctate, the abdomen
smooth, moderately sparsely punctulate. Length 13. S mm. ; width
6.5 mm. Pomona Mts luctuosa n. sp.

Form a little shorter, broader and more convex, dull in lustre through-
out above, more strongly punctured than in any other species
of the group, deep black; head rather coarsely, deeply, not very
closely and rather unevenly punctate, not impressed, the anterior
canthus rather narrowly rounded and somewhat more prominent
than the posterior; prothorax a little wider than the base of the
elytra, transverse, fully two-thirds wider than long, the sides sub-
parallel and feebly arcuate, gradually more arcuate and converg-
ing anteriorly, the apex broadly sinuate and nearly three-fourths
as wide as the base, the angles right and blunt ; surface finely but
strongly, rather sparsely punctate, gradually coarsely, somewhat
closely and very conspicuously so laterally, each puncture with a
small hair, the margin strongly beaded, the attendant channel
deep also at apex laterally; scutellum moderate, not very trans-
verse, triangular; elytra not quite one-half longer than wide, ob-
tusely angulate at tip, the sides parallel and very feebly arcuate,
finely reflexed ; surface of each with four feebly tumescent ridges,
with intermediate ridges also, though very obsolete, the four ridges
very strong on the apical declivity, the punctures sparse, strong
and not very fine medially, becoming very coarse, closer and con-
spicuous laterally, the fulvous hairs distinct ; prosternum rather
coarsely and closely punctate, the abdomen polished, minutely,
sparsely punctulate. Length 13.0 mm.; width 6.7 mm. San
Bernardino Mts costulata n. sp.

Form stout, not more than twice as long as wide, not pubescent, con-
vex, parallel, the integuments above subopaque, the pronotum
and elytra very densely, rather coarsely and aciculately punctate,
the elytra obscurely tricostate, with the intervals obsoletely reticu-
late ; [abdomen minutely, sparsely punctulate]. Length 15.5
mm. Fort Tejon robusta Horn

Form oblong, moderately elongate, with the sides feebly arcuate, rather
strongly convex, very highly polished, smooth, the vestiture mi-
nute, fulvous in color ; head rather coarsely and densely punctured,
the sides as prominent as the posterior canthus of the eyes, the
antennae slender ; prothorax one-half wider than long, the apex
nearly two-thirds as wide as the base, the angles rather broadly
rounded; base transverse, very feebly sinuate laterally, the angles
slightly rounded and not prominent, the sides strongly arcuate
anteriorly, feebly so posteriorly ; surface very finely punctured,
the punctures becoming denser at the sides, the bead strong ; ely-
tra at least twice as long as the prothorax, the punctures fine
though larger than those of the pronotum, rather sparse, the sides

Proc. Wash. Acad. Sci., April, 190S.

90 CASEY

feebly arcuate; abdomen strongly convex, polished, feebly and
sparsely punctate. Length 12.5 mm.; width 6.0 mm. Califor-
nia (Calaveras Co., — at an elevation of 2,300 feet).

sanfordi Blaisd.

The form which I have identified as abdonimalis displays
very Httle sexual difference in the form of the prothorax, while
in integer this difference is more pronounced than in any other
known species of the genus. In the definition of robnsta, as
given above, I have made use of all the characters published by
its author, assuming as the type the specimen numbered " i "
(Rev. Ten., p. 296) and abandon the species, represented by an
assumed cotype in the LeConte cabinet, described under this
name in my previous revision (Col. Not., II, 1890, p. 375). The
species represented by this latter specimen undoubtedly has a
completely margined prosternal process and may possibly be
unconsciously included above, among the forms from abdonii-
nalis to 7'tigosa in the table ; it is probable that the process is not
apically margined in the true robnsta, although the author
records nothing concerning a character of which he subse-
quently made constant use in Eusattus. The sculpture of ro-
bnsta departs to a very remarkable degree from the usual fine
sparse punctures of this first group of the genus, but is suggested
in the coarser punctuation of costulata, which is a markedly
aberrant species. The species described by Blaisdell under the
name sanfordi (Ent. News, VI, p. 235) probably belongs in
this group, although rather discordant in habitat; it is described
above from the published characters.

Group II. — Type opaca.

The probably even more numerous species of this group,
reproduce the general form and sculpture of their more stren-
uous relatives of the abdominalis group, but are alwa3^s very
much smaller in size, including in fact some of the smaller
species of the genus. They inhabit also a distinctl}^ different
region, being virtually confined to the southern Sierras and ad-
jacent regions, not occurring, so far as known to me, very near
the coast; more definite localities when recorded, will be stated
below. Those known at present, representing probably but a
moderate proportion of all actually existing forms, may be thus
distinguished among themselves : —

REVISION OF TENEBRIONID SUBFAMILY CONIOXTIN^E 9I

Prosternal process strongly and unequivocally margined throughout its
contour 2

Prosternal process not margined apically 7

2 — Elytral punctures rather small and sparse but very distinct 3

Elytral punctures excessively minute, observable with difficulty under
a hand lens 6

3 — Elytral punctures not sensibly piliferous 4

Elytral punctures each with a very small but evident decumbent hair.. 5

4 — Body oblong, only moderately convex, deep black, rather shining

throughout ; head somewhat finely but strongly, not densely
punctate, the lobes very feebly impressed, the anterior canthus
very broadly rounded, as prominent as the posterior; prothorax
not more than one-half wider than long, the sides just visibly
converging and nearly straight to about the middle, then rapidly
more rounded, then more strongly converging and but feebly
arcuate to the apex, which is nearly three-fourths as wide as the
base, circularly sinuate, with the angles right and but slightly
blunt; surface very finely, sparsely punctate, becoming much less
finely, more closely and rather strongly so laterally, the marginal
bead very fine; elytra parallel, not quite one-half longer than
wide, as wide as the prothorax, obtusely ogival at tip, nearly
smooth, the margins finely reflexed ; prosternum rather finely but
strongly, somewhat sparsely punctate, the process broad, ogival
at tip ; abdomen and hind femora finely, rather sparsely punctate.
Length 9.6 mm. ; width 4.5 mm. Arizona (probably the extreme
western part) histrio n. sp.

Body rather stouter in outline and decidedly more convex, deep black,
duller in lustre, feebly alutaceous ; head more strongl}- and closely
punctate, not impressed at the apical lobes, the anterior canthus
more narrowly rounded, slightly less prominent than the posterior ;
antennte similarly slender; prothorax nearly similar but more
transverse, fully three-fifths wider than long, much duller, the
fine punctures stronger and less sparse medially, becoming much
larger, perforate and very close-set laterally ; scutellum rather
larger and more broadly triangular ; elytra scarcely more than
two-fifths longer than wide, as wide as the prothorax, very obtuse
at tip, the surface feebly undulato-rugose apically, elsewhere nearly
smooth but with very obsolete broadly convex lines, the punctures
rather stronger and more close-set than in histrio., very distinct,
although the surface is somewhat dull ; prosternum rather sparsely
and finely punctate, the process narrower, impressed along its
median line, the impression terminating in a deep discal fovea near
the apex — perhaps an accidental character ; abdomen and hind
femora very finely, sparsely punctate, polished. Length 9.5
mm.; width 4.65 mm. San Bernardino Co ancilla n. sp.

5 — Form stout, rather convex, oblong, deep black, only very feebly

alutaceous ; head unusually short and transverse, rather coarsely and

92 CASEY

closely punctate, much more finely so basally, unimpressed, the
anterior canthus somewhat narrowly rounded and very much less
prominent than the posterior; antennae blackish, slender ; pro-
thorax fully two-thirds wider than long, the sides broadly rounded
and converging anteriorly, becoming nearly straight and parallel
from well before the middle to the base, the apex three-fourths as
wide as the base, broadly, circularly sinuate, the angles right and
but slightly blunt ; punctures fine but strong and moderately close-
set, becoming still stronger and a little closer, perforate and con-
spicuous laterally, the marginal bead very fine ; scutellum as in
ancil/a, broadly triangular, the elytra also similar in outline, very
obtuse behind, the surface nearly smooth, slightly rugulose pos-
teriorly, the punctures sparse but subperforate and very distinct;
prosternum unusually finely, sparsely punctate, the process rather
wide, circularly rounded at tip, broadly and feebly impressed
along its median line ; abdomen very minutely, sparsely, the hind
femora rather distinctly and less sparsely, punctate. Length 9.4
mm.; width 4.4 mm. California (Lancaster), — Mr. H. F.
Wickham. A single female oblonga n. sp.

6 — Form somewhat elongate, parallel, moderately convex, rather

dull and strongly alutaceous, black with a feeble piceous tinge,
glabrous, the elytral punctures with very small inconspicuous
hairs; head rather finely but strongly, not densely punctate, un-
impressed, the anterior canthus rather narrowly rounded, but
little less prominent than the posterior; prothorax nearly as in
oblonga though a little less transverse, fully one-half wider than
long, the sides scarcely becoming parallel posteriorly but very
feebly converging from the base though less arcuate, the punctures
very fine and rather sparse medially, becoming much larger, deep
and moderately close-set laterally, the marginal bead very fine ;
elytra very nearly one-half longer than wide, more gradually and
acutely ogival behind than usual, as wide as the prothorax, virtu-
ally smooth but unusually opaque throughout, the sparse punc-
tures but slightly evident even laterally; prosternum finely, not
densely punctate, the process narrower; abdomen and hind
femora finely, more or less sparsely punctate. Length 9.7 mm. ;
width 4.25 mm. California. A single female.

obsolescens n. sp.

7 — Hind femora finely, more or less sparsely or inconspicuously

punctate 8

Hind and other femora coarsely, closely and very conspicuously
punctate 13

8 — Pronotal punctures not particularly close- set laterally, separated

by from three to five times their own diameters 9

Pronotal punctures small but deep, unusuall}' close laterally, where
they are separated by from one to three times their own diameters ;
punctures throughout with extremely short decumbent hairs. ..11

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 93

9 — Elytral punctures with very evident small decumbent hairs.

Body somewhat elongate, parallel, moderately convex, rather
dull and alutaceous in lustre, black with a pronounced piceous
tinge, the legs frequently rufescent, the antenn^E pale throughout
and slender; head finely but strongly, rather sparsely punc-
tate, not impressed, the canthi about equally prominent; pro-
thorax nearly three-fifths wider than long, the sides broadly and
moderately arcuate and converging anteriorly, becoming very
feebly arcuate and but just visibly diverging thence to the base,
the apical angles very bluntly rounded ; punctures very minute
and sparse medially, becoming much larger, deep and distinct
though still well separated laterally, the marginal bead very fine ;
scutellum only moderate in size, broadly triangular; elytra
scarcely one-half longer than wide, as wide as the prothorax,
parallel, gradually parabolic at apex, virtually smooth though
more opaque than the pronotum, the punctures fine, rather sparse,
becoming more distinct laterally; presternum rather strongly,
more closely punctate, the abdomen remotely and extremely
minutely so, shining. Length S.1-8.S mm. ; width 3-7-3-9 mm.
Southern California pallidicornis Csy .

Elytral punctures without evident decumbent hairs; size much larger.

10

10 — Form broadly oblong, rather strongly convex, deep black through-

out, shining ; head finely, sparsely punctate, the apical lobes feebly
impressed, the anterior canthus moderately narrowly rounded, fully
as prominent as the posterior; antennae slender, black, with the
tip piceous; prothorax transverse, more than three-fourths wider
than long, the sides broadly, subevenly arcuate, rather more so
and more converging at apex, becoming parallel behind the
middle, the apex circularly sinuate, two-thirds as wide as the
base, w'ith the angles right and but slightly blunt; punctures
sparse and very minute, becoming strong but still rather fine and
sparse laterally, the lateral bead very fine ; scutellum rather small,
but slightly transverse, triangular; elytra as wide as the pro-
thorax, barely two-fifths longer than wide, the sides nearly
straight, the tip gradually and rather acutely ogival ; surface
smooth, rugulose apically, only moderately sparsely punctate,
finely so suturally, rather strongly and somewhat coarsely toward
the sides, the margin very finely reflexed ; prosternum minutely,
sparsely punctate, the abdomen and hind femora still more
sparsely, minutely and feebly. Length 10. o mm.; width 4.9
mm. Locality unrecorded but probably southern California.

degener n. sp.

Form relatively less broad, the largest species of the group, convex,
black, rather dull in lustre, the elytra subopaque ; head somewhat
finely, closely punctate, minutely and less closely posteriorly,
scarcely impressed at the apical lobes, the anterior canthus very
broadly rounded, not quite as prominent as the posterior; pro-

94 CASEY

thorax barely one-half wider than long, the sides parallel and
straight, becoming broadly rounded before the middle and
strongly convergent and less arcuate apically, sinuate as usual at
apex, the angles obtuse and bluntly rounded; surface alutaceous,
extremely minutely, sparsely punctulate, the punctures becoming
more visible but still minute and unusually sparse laterally, the
side margins rather strongly channeled and more coarsely beaded
than in any other species of the group, recalling the abdominalis
group somewhat; scutellum moderate; elytra as wide as the pro-
thorax, fully one-half longer than wide, gradually rather acute at
apex, the surface with numerous feeble sinuous impressed lines
and generally a few broader, irregular and eroded longitudinal
lines, the punctures very sparse and minute, still fine and sparse
laterally; punctures of the prosternum and abdomen fine and
sparse, of the hind femora rather closer, more distinct and some-
what pubiferous. Length 11.0-11.6 mm.; width 5.0-5.1 mm.
California verna n. sp.

II — Elytra short, barely two-fifths longer than wide, somewhat in-
flated behind the middle, where they are evidently wider than the
prothorax, with the sides broadly arcuate, the apex very obtuse.
Body short, strongly convex, not very intense black, dull or sub-
opaque in lustre throughout above ; head finely but deeply, not
closely punctate, unimpressed, the canthi equally prominent;
prothorax transverse, three-fourths wider than long, the sides
broadly rounded and converging before, becoming parallel and
straight behind, the middle, broadly sinuate at apex, with the
angles right and but slightly blunt ; punctures medially very
minute and rather sparse, less minute and very close-set laterally,
the marginal bead very fine; scutellum broadly triangular; elytra
smooth, minutely, rather sparsely punctured suturally, much more
closely and strongly so laterally though still rather finely, the
margin very finel}' reflexed ; prosternum finely but strongly, rather
closely punctate, the abdomen minutely, feebly and more sparsely,
the hind femora finely and less sparsely. Length 8.4 mm. ; width
4.0 mm. California (Owen's Valley). Described from the
female opaca Horn

Elytra longer, parallel, never inflated and less obtuse at apex 12

12 — Form very broadly oblong. Body moderately convex, black,
dull and subopaque in lustre; head rather finely and closely,
strongly punctate, the apical sinvis deep and rather narrow, the
apical lobes unimpressed or very obsoletely so, the canthi sub-
equally prominent; prothorax of peculiar form, transverse, fully
two-thirds wider than long, the sides rounded and strongly con-
vergent anteriorly, becoming parallel in basal two-thirds and
rather deeply and evenly sinuate at each side at about basal
fourth, the apical sinus broadly rounded, the angles slightly ob-
tuse, bluntly rounded, the basal angles not pronounced, the lateral
sinuses of the base moderate ; surface minutely but stronglv and

REVISION OF TENEBRIONID SUBFAMILY CONIONTINyE 95

rather closely punctate medially, move strongly and very closely
laterally, the marginal bead fine; scutellum rather small; elytra
about two-fifths longer than wide, more obtusely ogival at tip than
in the two following forms, smooth or with a few very feeble and
irregular impressed lines, the punctures minute and sparse sutur-
ally, stronger, closer and very distinct laterally ; prosternum finely,
sparsely punctate, the process wide, convex, broadly arcuato-
truncate at tip; abdomen extremely minutely and obsoletely but
rather closely punctulate, the hind femora rather distinctly though
not closely punctate. Length 9.8 mm. ; width 4.7 mm. Cali-
fornia (Amedee, Lassen Co.), — H. F. Wickham. Described from
a single male specimen lassenica n, sp.

Form more narrowly oblong-elongate, rather more convex. Body
black, dull and strongly alutaceous in lustre ; head finely but
strongly, not very closely punctate, deeply and rather narrowly
sinuate at apex, the lobes flattened, the canthi subequally promi-
nent, the anterior rather narrowly rounded ; prothorax less trans-
verse, three-fifths wider than long, the sides more broadly and
evenly rounded, more gradually converging anteriorly and be-
coming parallel and straight only behind the middle, otherwise
nearly as in lassenica^ the basal sinuations very feeble and the
inner side of the hind angles scarcely oblique; elytra parallel, as
wide as the prothorax, one-half longer than wide, more gradually
and acutely ogival at apex, smooth, with some feeble and uneven
scratches, very minutely and sparsely punctate suturally, very
distinctly and much more closely near the sides, the margin very
finely reflexed ; prosternum finely but strongly, rather closely
punctured and micro-setigerous, the abdomen shining, very mi-
nutely, rather sparsely and obsoletely punctulate, the hind femora
unusually finely, obsoletely and remotely punctulate. Length 9.8
mm. ; width 4.5 mm. Nevada (Reno). A single specimen of
undetermined sex taken by the writer on the banks of the Truckee
River nevadensis n. sp.

A — Body very nearly as in 7ievadensis but with the prothorax very
slightly less transverse and with the basal sinuations stronger,
so that the hind angles project posteriorly for a very short
distance, with their inner sides oblique; sculpture similar, the
elytra slightly smoother, with the fine punctures near the sides
relatively rather less distinct ; hind femora notably more strongly
and less sparsely punctate and a little more pubescent. Length
9.5 mm.; width 4.5 mm. Nevada (Carson City), — H. F.
Wickham carsonica n. subsp.

13 — Body oblong, parallel, moderately convex, deep black, alutaceous
in lustre, the elytra more strongly ; minute setae of the punctures not
projecting beyond the limits of the latter; head somewhat coarsely
but not very closely punctate, unimpressed at the lobes, the apical
sinus rather deep and abruptly formed ; anterior canthus rounded,
slightly more prominent than the posterior ; antennae rather

g6 CASEY

slender, black, paler at tip; prothorax fully three-fourths wider
than long, wider behind the middle than at base, the sides alinos
evenly arcuate, slightly more converging anteriorly, the apex very
broadly and circularly sinuate, more than three-fourths as wide as
the base, the angles barely right and slightly blunt, the hind angles
right, not at all blunt; surface rather finely and sparsely but
strongly punctate, becoming more closely, unusually coarsely and
somewhat irregularly so laterally, the marginal bead fine; scu-
tellum rather small, not strongly transverse, triangular; elytra
nearly one-half longer than wide, as wide as the prothorax, the
sides straight, the apex gradually narrowed to the obtuse tip ; sur-
face smooth, sparsely and rather strongly punctate suturally, some-
what coarsely and less sparsely laterally; prosternum rather
strongly j^unctate, the process sparsely so, unusually declivous
apically; abdomen very minutely, feebly and sparsely punctate.
Length 9.3 mm.; width 4.25 mm. (cJ*). California (Barstow).

punctipes Csy.

In the catalogue of the Coleoptera of Baja California, p.
349, Horn cites j[)allidicornis as occurring there, without more
definite indication of locality; the identification, moreover, is
open to a good deal of doubt, as nearly all the species are very
local in habitat. Verna has some of the characters of Group
I, and is undoubtedly an intermediate form ; it has more espe-
cially the grooved and narrowly reflexed thoracic side margins
of the allies of abdoniinalis, but its much smaller size prompts
me to include it with Group II, for convenience of identifica-
tion ; dcgene7' is likewise aberrant, having some of the charac-
teristics of the ovalis group. The type of lasscnica has, besides,
the sinuation of the sides of the prothorax near the base, two
oblique impressed discal lines from lateral fourth very near the
base to lateral sixth near basal fifth ; these lines are observable
occasionally in other species of all the groups and seem to be
adventitious to some extent but they are bilaterally symmetric.

Group III. — Type viatica.
This is one of the largest groups of the genus and the most
difficult in the delimitation and definition of specific forms.
The group inhabits the coast regions from San Francisco Bay
well into Oregon, from which State I have a single species, and
but few of its components occur as far to the eastward as the
foothills of the Sierras; more definite localities will be men-
tioned when known. The very small species allied to ^//;/r//-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 97

collis^ form a tolerably distinct section by themselves and have
occasionally some unusual modifications of the marginal line of
the prothorax. The others, forming the larger part of the group,
have a superficial interresemblance which is very confusing,
necessitating prolonged and careful study before even attempt-
ing to identify them by published characters. In a number of
forms it is difficult to say whether the prosternal lobe is mar-
gined or not, there being some variability in the character, and,
when the margin becomes feeble, as in o^acicolh's and longi-
colli's, the surface must be held so as to reflect the light at grad-
ually varying angles in order to recognize the marginal con-
vexity ; in others, however, the margin takes the form of a
strong convex bead, defined by an unequivocal groove, but the
character is not altogether satisfactory.

The forms at present in my cabinet may be known as fol-
lows : —

Larger species, never under lo.o mm. in length 2

Small species, never exceeding lo.o mm. in length 15

2 — Prosternal lobe margined throughout its contour 3

Prosternal lobe margined at the sides but not at apex 12

3 — Prothorax broadly rounded at the sides, sensibly wider behind the

middle than at base. Form parallel, convex, deep black, polished
throughout, glabrous ; head rather strongly but sparsely punctured,
not impressed at the lobes, the anterior canthus slightly less
prominent than the posterior; prothorax one-half wider than long,
the sides gradually more rounded and converging apically, the
apex barely three-fifths as wide as the base, the sinus circularly
rounded but not deep, the angles obtusely rounded ; surface very
fineh^ and remotely punctate, less finely but still sparsely so later-
ally, the marginal bead rather fine, not at all incurvate at base;
scutellum large, broadly triangular; elytra parallel, as wide as the
prothorax, rather more than one-half longer than wide, very
obtusely rounded at tip, the surface nearly smooth, somewhat
finely though strongly, very remotely punctate, but little more
coarsely or less sparsely so toward the sides, the margins very
finely reflexed ; prosternum finely, sparsely punctate, the process
stronglv grooved and beaded about its entire contour; hind femora
rather finely but deeply, sparsely punctate, the abdomen minutely,
very remotely. Length 13.5 mm. ; width 5.9 mm. California.

rotundicollis n. sp.

Prothorax not conspicuously rounded at the sides, though a little
narrower at base than at a short distance before the latter. Form
narrower and slightly less convex, parallel, deep black, not so

98 CASEY

shining, feebly alutaceous throughout, glabrous; head rather
strongly, not very closely punctate, very obsoletely impressed at
the apical lobes, the anterior canthus fuller than in the preceding,
not quite as prominent as the posterior; prothorax one-half wider
than long, the sides broadly and moderately rounding and not
very converging anteriorly, subparallel thence to the base and
feebly arcuate, the apex much more broadly though moderately
sinuate in circular arc, nearly four-fifths as vv^ide as the base, the
angles right and but slightly blunt; surface finely, sparsely punc-
tate, more coarsely but still rather sparsely laterally, the marginal
bead moderate; elytra similar though less obtuse and more ogival
at tip, the surface very feebly undulato-rugulose, with vague traces
of two or three elevated lines, remotely punctate, the punctures
rather coarse for the present group and closer and still stronger
laterally; prosternum finely, rather sparsely punctate, the process
margined about its entire contour by a distinct groove and unusu-
ally fine but distinct bead ; femora and abdomen nearly as in the
preceding species. Length 13.3 mm.; width 5,4 mm. Mokel-
vunne Hill, Calaveras Co blaisdelli n. sp.

Prothorax with the sides rounded and converging anteriorly, becoming
straight or nearly so and parallel or subparallel thence to the
base 4

Prothorax with the sides converging from base to apex ; marginal bead
of the prosternal process rather fine but strong, generally entire.. 1 1

4 — Marginal groove and beading of the prosternal process strong

and distinct 5

Marginal groove and beading very feeble, sometimes observable with
difficulty S

5 — Sides of the prothorax very gradually more rounding anteriorly,

not at all subangulate 6

Sides more or less abruptly rounded or subangulate at or before the
middle 7

6 — Prothorax more transverse, about one-half wider than long. Body

elongate, moderately convex, rather shining, with short sparse
pubescence, especially distinct on the elytra ; head rather coarsely,
not densely punctate, the lobes scarcely impressed, the transverse
suture unusually broadly impressed, the anterior canthus broadly
rounded, much less prominent than the posterior ; prothorax rather
strongly narrowed anteriorly, the sides parallel and nearly straight
in basal half, the apex moderately sinuate, but little more than
three-fifths as wide as the base, the angles obtuse and blunt ; sur-
face very minutely, remotely punctate, somewhat strongly but
sparsely so laterally, the marginal bead fine; scutellum moderate;
elytra nearly as in blaisdelli but more rounded and less angulate
at apex, the surface almost smooth, much more finely and very
remotely punctate, more strongly but still finely and sparsely so
laterally, the edge very finely reflexed ; prosternum finely, sparsely
punctate, the process very minutely; abdomen and hind femora

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 99

finely and sparsely but rather strongly punctate. Length 14.0
mm.; width 5.75 mm. California innocua n. sp.

Prothorax more elongate, evidently less than one-half wider than long.
Body elongate, parallel, convex, deep black, glabrous, polished,
the pronotum sometimes feebly alutaceous ; head strongly and
rather sparsely but not very coarsely punctate, more finely and very
sparsely toward base, the lobes broadly rounded, unimpressed,
the anterior canthus narrowly rounded, slightly less prominent
than the posterior ; prothoi"ax with the sides only moderately con-
verging and broadly rounded anteriorly, the apex nearly three-
fourths as wide as the base, the circularly rounded sinus moderate
in depth, the angles rather obtuse and bluntly rounded ; surface
minutely, remotely punctate, much more strongly and less sparsely
laterally, the marginal bead moderate; scutellum triangular, not
very transverse ; elytra nearly three-fourths longer than wide,
about as wide as the prothorax or slightly less, rather abruptly
subangulate at tip, the surface nearly smooth, rather finely, re-
motelv punctate suturally, less remotely and more strongly toward
the sides, which are finely reflexed ; prosternum rather finely and
sparsely punctate, the process with a distinct and entire marginal
groove and moderately coarse bead ; hind femora and abdomen
minutely, very S])arsely punctate. Length i 2.2-15.0 mm. ; width
5.0-6.0 mm. (cJ^, 9). Benicia. Collected by the writer in con-
siderable number elongata Csy.

A — Similar to elongata throughout though a little stouter, the an-
terior canthus fuller and less angulate at the eyes but similarly
less prominent than the posterior, the elytra barely more than
one-half longer than wide and more obtuse at tip (?), and the
prosternal process broader, with relatively finer marginal bead,
its apex similarly circularly rounded. Length 14.8 mm. ;
width 6.7 mm. California. Cab. Levettc.limatula n. subsp.

7 — Form relatively rather stout, the largest species of the group, pol-
ished throughout, glabrous and deep black above, piceo-rufous
beneath; head coarsely, deeply and not densely punctate, the
punctures becoming sparse and very minute toward base, the lobes
unimpressed, the sinus broad and circularly rounded, the canthi
nearly as in elongata; prothorax shorter, fully one-half wider than
long, the sides subparallel and nearly straight from the base to
about apical third, where they are rather strongly rounded, thence
strongly converging and almost straight to the angles, which are
right and somewhat blunt; apex two-thirds as wide as the base,
with the sinus only moderately deep, the base sensibly wider than
the base of the elytra ; surface very obsoletely ridged along the
median line, very minutely, remotely punctate, more distinctly
but still sparsely laterally, the marginal bead moderate; scutellum
as in elotigata ; elytra evidently more than one-half longer than
wide, somewhat gradually acutely ogival at apex, nearly smooth,
everywhere finely and very remotely punctate ; prosternum finely,

lOO CASEY

rather sparsely punctate, the process not very broad, rounded at
tip, the marginal bead moderate; abdomen and hind femora
minutely, remotely punctate. Length 15.5 mm. ; width 6.6 mm.
(9). Mt. Diablo. A single specimen taken by the writer.

obsidiana n. sp.

Form much more slender, more convex, parallel, deep black, the tarsi
piceous, glabrous, alutaceous, the elytra polished ; head finely,
rather sparsely punctate, the sinus shallow and parabolic, the lobes
unimpressed, the anterior canthus rounded, much less prominent
than the posterior; prothorax short, a little more than one-half
wider tlian long, the sides parallel and straight to slightly before
the middle, then broadly rounded, becoming rather strongly con-
vergent and but feebly arcuate thence to the angles, which are ob-
tuse and bluntly rounded, the apex two-thirds as wide as the base,
with the circularly rounded sinus shallow; surface sparsely, ex-
tremely minutely and feebly punctate, only a little more distinctly
punctured and still rather sparsely and very finely so toward the
sides, the marginal bead fine; scutellum moderate, transversely
and obtusely triangular; elytra relatively very long, as wide as
the prothorax and almost three times as long, fully three-fourths
longer than wide, the apex rapidly narrowed and obtuse, ogival,
the surface nearly smooth, very minutely, remotely punctate
throughout; prosternum sparsely, extremely minutely punctate,
the process parallel, rounded at tip, strongly margined; hind
femora and abdomen very minutely, sparsely punctate. Length
13.0 mm. ; width 5.25 mm. California. Cab. Levette.

cylindrica n. sp.

8 — Anterior canthus less prominent than the posterior 9

Anterior and posterior canthi subequally prominent 10

9 — Body parallel, moderately convex, black, alutaceous, the elytra
less so ; punctures bearing very small, decumbent and rather dark
hairs; head coarsely, rather closely punctate, more finely toward
base, the lobes broadly vounded, thick, unimpressed, the sinus circu-
larly rounded, shallow; anterior canthus rather broadly rounded;
prothorax one-half wider than long, the sides parallel and nearly
straight, becoming gradually rounded and strongly convergent
before the middle, the apex moderately sinuate, two-tliirds as wide
as the base, the angles obtuse though only slightly blunt ; surface
sparsely, very finely punctate, gradually not very coarsely but
strongly and still rather sparsely so laterally, the lateral bead
moderately thick; scutellum moderate, sharply triangular, but
slightly transverse; elytra two-thirds longer than wide, gradually
and rather acutely ogival at tip, the sides straight and parallel in
basal two-thirds; surface nearly smooth, not very finely, very
deeply and sparsely punctate, a little less sparsely though not much
more coarsely laterally ; prosternum finely, sparsely pimctate, the
process rather wide, constricted atthecoxre, rounded at tip, where
the beaded margin becomes subobsolete ; hind femora rather finely

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN.-E lOI

but strongly, sparsely, the abdomen minutely, punctate. Length
12.6 mm.; width 5.5 mm. Lake Co genitiva Csy.

Body nearly similar but rather more convex, black, densely subopaque,
the elytra and under surface sliining ; punctures without distinct
short hairs; head rather finely but strongly, not densely punctate,
the sinus rather deep, subparabolic, the lobes not impressed, the
anterior canthus more oblique and narrowly rounded; prothorax
more transverse and much more narrowed apically, somewhat
more than one-half wider than long, the sides broadly rounded,
becoming gradually more converging apically and parallel only
near the base ; apex rather deeply sinuate, only about three-fifths
as wide as the base, the angles broadly rounded ; surface rather
sparsely, very minutely punctate, becoming rather closely and more
strongly but still finely so laterally, the marginal bead fine; scu-
tellum large, triangular, very transverse; elytra relatively very
elongate, ahiiost three times as long as the prothorax, somewhat
parabolically rounded behind, two-thirds longer than wide, the
surface nearly smooth, becoming somewhat rugulose apically,
finely, sparsely punctate, a little more strongly so laterally; under
surface nearly as in the preceding, the femora more minutely and
sparsely punctate, the prosternal lobe more narrowly and para-
bolically rounded at tip. Length 13.0 mm. ; width 5.8 mm. Not
labeled in Cab. Levette Opacicollis n. sp.

10 — Form rather stout, strongly convex, deep black, only moderately
shining, glabrous ; head rather coarsely but loosely punctate, very
minutely so at base, the sinus broad, subevenly rounded, the lobes
unimpressed, the anterior canthus broadly rounded; prothorax
unusually elongate, about two-fifths wider than long, the sides
broadly, rather feebly rounded though a little more converging
apically but barely becoming parallel even at base, the apex
evenly and moderately sinuate, almost three-fourths as wide as the
base, the angles right and but slightly rounded ; surface very mi-
nutely, sparsely punctate medially, becoming rather coarsely and
deeply though not much more closely so toward the sides, the
marginal bead rather coarse; scutellum relatively rather small,
strongly transverse ; elytra but little more than twice as long as
the prothorax, parallel and straight at the sides, gradually and
arcuately narrowed and ogival in apical two-fifths, three-fifths
longer than wide ; surface nearly smooth, with a few obsolete
eroded lines, very finely, sparsely punctured throughout ; proster-
num finely, sparsely punctate, the process gradually slightly wider
posteriorly, with almost straight sides, its apex broadly rounded,
obsoletely reflexed, broadly concave on the disk ; hind femora
rather finely, remotely punctate. Length 14.0 mm. ; width 6.25
mm. A single example, labeled "Texas "but undoubtedly in
error and probably from the coast i^egions north of San P^rancisco.

longicollis n. sp

Form more slender and very much smaller in size, convex, black,
feebly alutaceous, the elytra and under surface polished, gla-

I02 CASEY

brous ; head finely but deeply, loosely punctate, the sinus broad,
moderately deep and subparabolic, the lobes unimpressed, rather
narrowly rounded, the anterior canthus rounded ; prothorax
rather long, perceptibly less than one-half wider than long,
the apex moderately sinuate and fully two-thirds as wide as
the base, the angles right and rather broadly rounded ; sides
very moderately, subevenly arcuate throughout, slightly more
converging anteriorly but barely becoming parallel even at base ;
surface rather sparsely punctate throughout, the punctures ex-
cessively minute medially and more visible, though still very fine
laterally, the marginal bead rather fine; scutellum moderate;
elytra evidently more than twice as long as the prothorax, fully
three-fifths longer than wide, parallel and straight at the sides,
gradually and obtusely ogival behind; surface nearly smooth, with
traces of close-set longitudinal costules, very finely, sparsely punc-
tate throughout; prosternum minutely, sparsely punctate, the
process but little dilated posteriorly, with its tip rather prolonged
and narrowly rounded; abdomen and hind femora minutely, re-
motely punctate. Length lo.o mm. ; width 4.12 mm. San
Francisco. A single male taken by the writer in the suburbs.

timida n. sp.

II — Body rather small, elongate, strongly convex, deep black, gla-
brous, rather shining, the pronotum at most very feebly aluta-
ceous; head finely, but strongly, rather sparsely punctate, the
sinus rather broad and shallow, the lobes unimpressed; anterior
canthus oblique, not broadly rounded, much less prominent than
the posterior ; prothorax visibly less than one-half wider than long,
the sides somewhat strongly converging, subevenly and very feebly
arcuate from base to apex, the latter feebly and circularly sinuate,
two-thirds as wide as the base, the angles right and bluntly
rounded ; surface sparsely, extremely minutely j^unctate, still very
finely and rather sparsely, though more distinctly, toward the sides,
the lateral bead fine; scutellum moderate; elytra a little more
than twice as long as the prothorax, more than one-half longer
than wide, parallel, gradually ogival at apex; surface nearly
smooth, very finely, remotely punctate, the punctures scarcely less
remote or more distinct laterally; prosternum sparsely, very
minutely punctulate ; hind femora and abdomen remotely punctate,
the former somewhat finely but strongly, the latter extremely
minutely. Length 10. 6-1 1.2 mm.; width 4.5-4.65 mm. San
Francisco. Two male specimens taken by the writer in the
suburbs conicicollis n. sp.

12 — Anterior canthus very oblique, much less prominent than the

posterior ; sides of the prothorax more or less evidently converg-
ing from base to apex 13

Anterior canthus much fuller and broadly rounded, as prominent as
the posterior; punctuation more conspicuous than usual 14

13 — Form moderately narrow and convex, alutaceous, the elytra less

I

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ IO3

obviously so and more shining, black throughout, glabrous ; head
finely but strongly, rather sparsely punctate, the sinus broad and
shallow, the obtuse lobes unimpressed; prothorax fully one-half
wider than long, the sides broadly arcuate, strongly so and very
convergent anteriorly, becoming almost parallel basally, the sinus
shallow, two-thirds as wide as the base, the angles much rounded;
surface sparsely, extremely minutely punctate, more distinctly
though still finely and rather sparsely toward the sides, the bead
moderately fine ; scutellum moderate, broadly triangular; elytra
between two and three times as long as the prothorax, two-thirds
longer than wide, gradually ogival at apex; surface not quite
smooth, remotely and very finely punctate throughout, but little
more closely or distinctly so laterally; prosternum minutely,
sparsely punctate, the process margined almost to the tip, which is
obtusely rounded, the bead sometimes virtually entire ; hind femora
and abdomen remotely and minutely punctate. Length 10. 5-13. 7
mm. ; width 4.25-5.4 mm. San Francisco. A moderate series
taken by the writer but not received from other sources.

viatica Esch.

Form shorter, broader, more convex longitudinally, much more shin-
ing, not alutaceous anteriorly, glabrous, deep black ; head rather
sparsely, very finely punctate, the apical sinus narrower and rather
deeper, the antennae stouter ; prothorax a little more elongate,
evidently less than one-half wider than long as a rule, the sides
subsimilar though frequently subangulate at the middle, the api-
cal sinus similarly shallow and the angles well rounded; surface
polished, very minutely, sparsely punctulate, in general but little
less finely so laterally ; elytra much less elongate, more obtuse at
tip, barely one-half longer than wide, sparsely but less finely and
more strongly punctured than in viatica^ the punctures impressed ;
prosternum sparsely, very finely punctate, the process broad, ob-
tusely rounded at tip, the marginal bead rather fine, generally
extending almost to the apex but in some cases entire; hind
femora and abdomen minutely, remotely and scarcely distinctly
punctulate. Length 11. 5-12. 7 mm.; width 4.9-5.S mm. Cali-
fornia. Cab. Levette lucidula n. sp.

Form rather less slender than in viatica^ larger in size, blacK, the
under surface and legs piceous or rufescent, shining, the anterior
parts somewhat alutaceous, glabrous ; head very finely, sparsely
punctate, the apex rather broadly sinuate in circular arc, the
lobes obtusely rounded, not impressed ; prothorax barely one-
half wider than long, somewhat conical, the sides very feebly
and subevenly arcuate from base to apex, becoming parallel only
very near the base, the apical sinus shallow, three-fifths as wide
as the base, the angles obtuse and moderately rounded; surface
sparsely, extremely minutely punctate, more distinctly but still
sparsely and very finely so laterally, the marginal bead moderate ;
elytra about two and a third times as long as the prothorax,
parallel, gradually ogival behind; surface nearly smooth, finely,

104 CASEY

remotely punctate throughout, more strongly near the sides, the
punctures impressed as in the two preceding; presternum very
minutely, sparsely punctate, the process broadly rounded, margined
nearly to the apex ; hind femora finely but strongly, moderately
sparsely punctate, the abdomen minutely, very remotely so.
Length 14.2 mm. ; width 5.8 mm. Berkeley, Alameda Co.

protensa n. sp.
14 — Body rather stout for the present group, strongly convex, black
throughout, alutaceous, the elytra less so, glabrous; head rather
finely and sparsely but strongly punctate, the sinus very broad
and somewhat parabolic, the broadly obtuse lobes unimpressed;
prothorax rather large, fully one-half wider than long, the sides
parallel and nearly straight in about basal half, gradually rounded
and strongly converging thence to the angles, which are right
and slightly blunt ; apical sinus moderately deep, three-fifths as
wide as the base, the latter posteriorly obliquely sinuous laterally ;
surface sparsely, very minutely punctate, becoming more closely
and much more strongly, though not coarsely so, laterally, the
bead rather strong, the fine but strong groove flexed along the
apex to lateral fourth; scutellum broadly and obtusely angulate ;
elytra two and a third times as long as the prothorax, rather
more than one-half longer than wide, parallel, unusually obliquely
and gradually narrowed to the narrowly obtuse tip ; surface some-
what uneven and irregularly crumpled, the punctures sparse but
rather large for this group, impressed as in the preceding; pro-
sternum finely but strongly, sparsely punctate, the process obtuse,
with a strong lateral margin not extending behind the coxas ; hind
femora rather coarsely, deeply and not very sparsely punctate,
the abdomen minutely, remotely so, more strongly and closely
on the basal segment. Length 14.8 mm. ; width 6,3 mm.
California thoracica n . sp .

Body less stout, elongate, parallel, rather convex, deep black, gla-
brous, somewhat strongly shining, scarcely alutaceous anteriorly;
head rather finely but strongly, unusually densely punctate, the
sinus not very broad, rather deeply parabolic, the lobes broadly
obtuse, unimpressed ; prothorax fully one-half wider than long,
a little wider behind the middle than at base, the sides parallel
and broadly arcuate, somewhat sinuate toward base, moderately
rounding and converging before the middle, the apex with shallow
circular sinus, three-fourths as wide as the base, the angles some-
what obtuse but not broadly rounded; surface finely and rather
sparsely but distinctly punctate, not very closely though much
more strongly and relatively coarsely so laterally, the marginal
groove along the beading flexed inward at apex; scutellum not
very transverse, sharply triangular; elytra parallel, fully two-
thirds longer than wide, the sides more arcuately and less
obliquely converging at apex; surface slightly uneven, undulate
and with traces of three longitudinal ribs on each apically, the
punctures less sparse than usual and much larger thougli not

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ IO5

coarse, deep and less impressed, still coarser toward the sides;
prosternum finely but strongly, sparsely punctate, the process
broadly arcuato-truncate at tip, the beaded margin not extending
quite to the apex ; hind femora rather strongly and distinctly
punctate, the abdomen very minutely and remotely punctulate.
Length 13.3 mm.; width 5.3 mm. Oregon (Medford), — H. F.
Wickham proba n. sp.

15 — Side margin of the prothorax never more than slightly curved at

base 16

Side margin inflexed at base, extending along the basal margin 3i

16 — Anterior canthus very full and broadly rounded 17

Anterior canthus, because of the more oblique and straighter sides of
the front, coming to a bluntly rounded point adjoining the eye. .30

17 — Upper surface virtually glabrous, the hairs borne by the punctures

ver}' minute and inconspicuous i S

Upper surface evidently clothed sparsely with short and decumbent
hairs 19

iS — Form subcylindric, strongly convex, strongly shining throughout,
deep black, the legs and antennae dark rufous or piceous ; head
finely but strongly, closely punctate, the sinus broad, moderately
shallow, subparabolic, the anterior canthus broadly prominent and
but little less so than the posterior ; antennae rather stout, com-
pressed; prothorax two-fifths to three-fifths wider than long, the
anterior sinus circular, rather shallow, the sides strongly rounded
and converging before, becoming nearly straight and parallel be-
hind, the middle; surface finely but very strongly, distinctly and
closely punctate, still more strongly and closely though only a
little more coarsely so laterally, the beaded margin extremely fine,
curving at base about the hind angles only and then abruptly dis-
appearing; scutellum rather small, but little wider than long,
triangular, with the apex obtusely rounded ; elytra fully one-half
longer than wide, parallel, as wide as the prothorax, the apex
obliquely and arcuately narrowed, the extreme tip narrowly ob-
tuse ; surface very finely punctate, the punctures close-set in longi-
tudinally lineate areas separated by narrower and comparatively
impunctate lines; side margin very finely reflexed ; under surface
and femora very minutely, sparsely punctulate. Length 8.7-10.0
mm. ; width 3.5-4.3 mm. San Francisco Co., — Chas. Fuchs.
[Benicia, — Leconte] puncticoUis Lee.

Form a little shorter and less cylindrical, smaller in size, convex, deep
black, with the legs rufo-piceous, the antenuee blackish-piceous ;
surface less shining, faintly alutaceous ; head finely, densely punc-
tate, nearly as in puncticoUis but with the broadly rounded an-
terior canthus much less prominent than the posterior; prothorax
a little shorter, transverse, with the sides very feebly converging to
parallel from the base to the middle and usually nearly straight,
then more strongly convergent and moderately arcuate to the right

Proc. Wash. Acad. Sci., April, 1908.

I06 CASEY

and rounded apical angles, the sides generally with a distinct sub-
angulation at the middle; apical sinus broad and shallow, two-
thirds as wide as the base ; surface more minutely and less closely
punctate than in the preceding, the punctures however unusually
close-set, the fine margin not curved at base so completely about
the basal angles; scutellum nearly similar but less obtuse; elytra
nearly similar but a little shorter and especially more gradually
and acutely ogival posteriorly, the even finer punctures similarly
disposed, the subimpunctate lines often irregularly costuliform ;
under surface and femora minutely, rather sparsely punctate.
Length 8.2-9.2 mm.; width 3. 2-3. 85 mm. San Francisco.
Taken in the suburbs by the writer and not known from other
sources exigua n. sp .

Form still shorter, subcylindric, strongly convex, much more obtuse
before and behind, deep black, the legs rufo-piceous ; upper sur-
face alutaceous ; head finely but not very closely punctate, the
sinus shallow, the oblique side margins sinuous at the ends of the
oblique sutures — an unusual chai^acter in this genus, — the frontal
parts rufescent as in the two preceding; prothorax rather elongate,
evidently less than one-half wider than long, the sides very feebly
converging and almost straight from the base nearly to apical third,
then rounded and strongly converging but not angulate, the apex
broadly and shallowly sinuate, nearly four-fifths as wide as the base,
the angles broadly rounded ; surface finely but strongly, not very
closely and unevenly punctate, becoming a little more strongly
and closely so laterally, the marginal bead fine, arcuate at base;
scutellum rather small ; elytra short, barely one-half longer than
wide and but little more than twice as long as the prothorax, par-
allel, very obtusely ogival at tip ; surface with fine but rather
strong, moderately close-set punctures disposed as in the preced-
ing, the subimpunctate lines tumescent; under surface finely punc-
tate, the prosternum rather strongly and somewhat closely.
Length 7.9 mm. ; width 3.5 mm. California. Cab. Levette.

paupercula n. sp.

19 — Body moderately elongate, parallel, convex, deep black, the legs
in great part and the antennas apically rufescent ; upper surface
smooth and dull, strongly alutaceous ; head rather small, very
finely, not densely punctate, the sinus rather deep and narrower,
the oblique sides straight; anterior canthus about as prominent as
the posterior; prothorax short, three-fifths \vider than long, the
sides subevenly rounded, rather strongly converging anteriorly,
becoming parallel in about basal half, the apex barely more than
three-fifths as wide as the base, with moderately deep sinus and
broadly rounded angles ; base almost truncate, with obtuse angles ;
surface minutely, rather sparsely punctulate, more strongly but
still not ver}' closely so laterally, the marginal bead extremely fine,
only slightly arcuate at base; scutellum moderate, polished; ely-
tra not quite one-half longer than wide, two and one-half times as
long as the prothorax, parallel, gradually moderately acute and ogi-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^E IO7

val in posterior two-fifths ; surface very smooth, with barely a trace
of tumescent lines, the punctures minute, moderately close in
broad lines as in the preceding species; under surface very mi-
nutely, remotely punctulate and polished, the presternum more
opaque and minutely, rugosely punctate, the process rather nar-
row, margined strongly almost to the tip. Length 7.5 mm. ;
width 3.2 mm. Alameda Co microsticta n. sp.

20 — Form rather stout for the present group and moderately convex,
glabrous, shining and onlv very faintly alutaceous, deep black,
the legs rufo-piceous, the antennae nearly black ; head finely but
not very closely punctate, the sinus moderate, the anterior canthus
much less prominent than the posterior ; prothorax formed nearly
as in 7}iicrostlcta but much less transverse, barely one-half wider
than long, the surface finely but strongly, moderately sparsely
punctate, becoming still more strongly and rather closely but un-
evenly so laterally, the marginal bead rather fine, strongly curving
about the hind angles; scutellum rather small ; elytra barely one-
half longer than wide, two and a third times as long as the pro-
thorax and equally wide, the sides straight and parallel, gradually
rounding and converging behind the middle, the apex narrowly
parabolic ; surface with fine but unusually strong, moderately
close-set punctures, with the less punctate lines narrower and dis-
tinctly tumescent ; under surface minutely, sparsely punctate
throughout, the prosternal process rather wide, very faintly mar-
gined at the sides along the coxae, its apex unusually deflexed and
convex. Length 9.2 mm. ; width 3.82 mm. California.

picipes n. sp.

21 — Lateral bead of the prothorax very fine, rounding the hind angles
and extending inward along the base for only a short distance.
Body rather strongly convex, subcylindric, glabrous, deep black,
the legs dark rufous, the antennae nearly black ; surface rather
shining though somewhat alutaceous; head rather small, finely,
strongly, somewhat closely punctate anteriorly, more finely and
sparsely behind the suture, the sinus moderately broad, the anterior
canthus moderately full, less prominent than the posterior ; pro-
thorax as in picipes but rather more rapidly narrowed anteriorly,
the angles obtuse but rather less rounded; surface very minutely,
rather sparsely punctate, still very finely and rather sparsely so
laterally ; scutellum small ; elytra nearly as in picipes but nar-
rower, the surface similar but with the punctures still smaller and
feebler ; prosternal process narrower, finely though very evidently
margined along the coxae, similarly much defiexed and convex at
its apex. Length 7.7 mm. ; width 3.35 mm. San Francisco.
A single specimen taken by the writer in the suburbs.

inflexula n. sp.

Lateral bead even finer, extending along the basal margin almost to the
middle. Body rather more elongate, convex, feebly alutaceous;
coloration as in injiexula^ the upper surface bearing short but

I08 CASEY

very evident and ratlier coarse glistening hairs ; head unusually
small, finely but very strongly, rather closely punctate, the sinus
rather narrow, moderately deep, the anterior canthus full and
rounded, almost as prominent as the posterior; prothorax barely
one-half wider than long, the sides moderatel}' converging and
broadly, feebly rounded from base to apex, slightly more converg-
ing toward the apex, which is rather deeply sinuate and three-
fifths as wide as the base, with the angles right and evidently
rounded ; surface finely but strongl}', rather closely punctate,
unusually strongly and closely so toward the sides, with a feeble
impression along the sides behind the middle but not attaining
the base; scutellum small; elytra as in injiexula but still more
gradually and arcuately narrowed behind to the rather acutely
ogival apex; surface more uneven than usual, with distinct
tumescent lines, the punctures less fine or close-set than usual but
similarly disposed; under surface nearly similar. Length 8.4
mm. ; width 3.6 mm. California. Cab. Levette.

inconspicua n. sp.

In the above series the margining of the prosternal lobe be-
comes so singularly inconstant as to lose a great deal of its
supposed value elsewhere, and the divisions of the table based
upon this character are somewhat arbitrary, the prosternal char-
acters only applying to the majority of the representatives of the
several species. In clongata, for example, the greater number
of examples, all females, have an entire beaded margin, but
one of the three males in the series has the lobe completely
unmargined except toward its base. So again, in lucidula^
most of the specimens have the apical part of the lobe com-
pletely immarginate, while one, not differing otherwise to any
noticeable extent, has a strong and entire beaded margin. I
had separated a few forms upon this apparently important and
certainly most striking character, but was obliged afterward to
suppress them.' Some years ago Dr. F. E. Blaisdell gave me

'These cases are very puzzling and there are some seven or eight still unde-
scribed forms, each represented by from one to three examples, before me, in the
neighborhood of elongata, viatica and lucidula, which possess in each case
some peculiarity of habitus, sculpture or structure of the prosternal lobe. Al-
though they would seem to represent distinct species or subspecies, the material
is far too limited to enable me to form any definite opinion at present, and as
three of them are from the same locality as the types of elongata, the exact fixa-
tion of their taxonomic status is rendered thereby all the more uncertain. I
now think that the male example referred to cloiigata^ having an unmargined
prosternal lobe, mentioned above, will prove to be specifically distinct, as there
are some obvious differences in the oedeagus distinguishing it from the adopted

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ IO9

a specimen, taken by him at Mokelumne Hill, and I had been
under the impression that it represented the species which he
subsequently described as san/'ordi, but on studying the descrip-
tion of the latter it is impossible to discover a single harmoniz-
ing character to bear out the surmise, and I therefore attach to
it another name as above ; it is the only species of the present
group known to me as occurring on the eastern side of the
central valley, although the exact locality of a number of the
species described in the table is unknown. Saiifordi, by its
published dimensions, must be included in the abdoininah's
group, where, however, it is wholly isolated as before inti-
mated. The small species of the second section of the table are
distinguishable, as a group, by the peculiar segregation of the
elytral punctures into broad uneven longitudinal areas lying
between the narrower and usually somewhat elevated smooth
lines ; they might even be considered a distinct group by them-
selves.

Group IV. — Type cuneata.

In reality this group is only a continuation of the preceding,
but, as the cuneiform outline of its few known members is a
rather striking character common to both sexes, though much
more feebly developed in the female, it may therefore be con-
sidered distinct for convenience in identification. The three
known species are the following : —

Body deep black in color, more or less shining 2

Body dark piceous in color, dull and densely alutaceous, more southern
in habitat 3

2 — Form cuneate, widest anteriorly, the male shorter and more
strongly cuneiform, convex, moderately elongate, highly polished
and deep black throughout, glabrous; head somewhat coarsely,
moderately closely punctate throughout, the sinus unusually deep,
rather narrow, the anterior canthus rather full, rounded, fully as
prominent as the posterior ; prothorax one-half wider than long,
the sides evenly rounded and rather strongly convergent apically,

true male of elongata, and some of the others also represent distinct species in
all probability, but 1 will leave this discussion for a future occasion. Perhaps
after all, the margination of the prosternal lobe may not be so unstable a char-
acter in this group as I surmised on concluding the table of its species, but it
should be stated that, under the circumstances, it was deemed best to distin-
guish by name only those forms markedly isolated in general habitus and there-
fore assumably having the status of true species or subspecies.

no CASEY

becoming parallel and straighter in about basal half, the apex
moderately sinuate in circular arc, two-thirds as wide as the base,
the angles right and only narrowly rounded ; surface finely and
sparsely but distinctly punctate, decidedly coarsely, very deeply and
much more closely so toward the sides, the marginal bead rather
thick, perfectly straight to the a])ex of the basal angles ; scutellum
well developed, transversely triangular ; elytra one-half longer
than wide (?) or less (cT), the sides feebly converging from the
base and nearly straight almost to apical third, then strongly
rounded and converging to the obtusely ogival tip; surface nearly
smooth or with a suspicion of broadly tumescent lines, not coarsely
but very strongly, sparsely punctured, decidedly more coarsely
and less sparsely toward the sides; prosternum minutely, sparsely
punctate, polished, the process moderately wide, feebly dilated
and rounded apically, with a continuous beaded edge (d^, 9);
hind femora and abdomen finely, remotely punctate. Length
11.0-11.7 mm.; width 5.1S-5.3 mm. California (Humboldt
Co. , — Hoopa Valley) atronitens n. sp.

Form nearly similar, smaller in size, convex, glabrous, the legs piceous
distally ; surface scarcely at all alutaceous ; head finely, not densely
punctate, the sinus very much broader and relatively shallower than
in the preceding, the sides more oblique, the anterior canthus much
less prominent than the posterior and coming to an obtuse point
at the eye; prothorax shorter, fully three-fifths wider than long,
similar in general form, the apex moderately sinuate, fully three-
fifths as wide as the base, the angles right and only slightly blunt;
surface sparsely, excessively minutely and scarcely visibly punc-
tulate, more visibly but still sparsely and very finely toward the
sides, where the bead is finer than in atro7iiteiis though bordered
internally by a broader groove; scutellum somewhat smaller;
elytra one-half longer than wide (d^), the sides more converging
from the base to apical third than in atronitens (d^), straight,
more gradually converging and acutely ogival at apex ; surface
nearly smooth, very sparsely and finely punctate, somewhat more
strongly and less remotely toward the sides ; prosternum minutely,
rather sparsely punctate, the process a little broader, more dilated,
convex and obtusely angulate at apex, finely margined and only
along the coxse ; hind femora finely but less sparsely punctate, the
abdomen excessively minutely, remotely so. Length 10.3 mm. ;
width 4,6 mm. California (locality unrecorded) sparsa n. sp.

3 — Form broader, less convex, strongly cuneiform ; under surface,
legs and base and apex of the antennie rufescent, subglabrous, the
setre extremely minute though visible; head very finely but
strongly, evenly and rather closely punctate, the sinus moderately
broad and deep, parabolic, the oblique sides rounded evenly from
the obtusely pointed canthus at the eye, much less prominent than
the posterior, to the apical lobes, which are rather angulate ; pro-
thorax fully two-thirds wider than long, the sides evenly round-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ III

ing and converging before basal third, straight and posteriorly
converging behind basal third and continuous in direction with
the straight and converging sides of the elytra; apex moderately
sinuate, nearly two-thirds as wide as the base, the curve of the
sinus becoming anteriorly arcuate and confused with the rounded
angles laterally; base almost truncate, the sinuses subobsolete;
surface sparsely, extremely minutely punctate, more evidently but
still finely and very sparsely toward the sides, the bead fine
though attended by a distinct narrow internal gutter; scutellum
rather small ; elytra relatively small, much narrower than the
widest part of the prothorax near basal two-fifths of the latter and
barely twice as long, gradually rounding and rather acutely ogival
behind ; surface nearly smooth, very finely, sparsely punctate, but
little more closely or coarsely so laterally, where the margins are
rather less narrowly reflexed than in the two preceding ; pro-
sternum finely but strongly, not very sparsely punctate, the process
wide, strongly dilated and obtusely angulate at apex, strongly
beaded but only along the coxae ; hind femora rather strongly but
sparsely punctate, the abdomen minutely. Length 10.8 mm. ;
width 5.3 mm. California (SanDiego), — G. W. Dunn.

cuneata n. sp.

The last of the above species is one of the more distinct of the
genus and differs from the first two, which are northern forms,
in being evenly narrowed posteriorly from behind the middle of
the prothorax and not solely from the base of the elytra ; it is
represented in my cabinet by a single male specimen.

Group V. — Type siibpubescens.

This is probably not a perfectly natural group, as it comprises
a few species from nearly all the zoological regions of Cali-
fornia, differing a good deal among themselves in facies. It is,
in general, characterized by a less convex form of body than
usual in the genus, although the latter may be stout or as elon-
gate as in viatica, but the surface is generally dull and alu-
taceous and always distinctly, though more or less sparsely,
pubescent, with the hairs decumbent and not erect and bristling
as in the setosa group. In some species, as for example mon-
iana, the male is distinctly cuneiform as in the ctmeata group
preceding, but I have not noticed this character as pertaining to
both sexes as it does there. The elytral punctures are more or
less small in size and the surface never so rugose as in the
eschscholtzi group. The seven species at present known may
be differentiated very clearly as follows : —

112 CASEY

Body larger, alwaysover lo.onim. in length, the head well developed

2

Body small, always much under lo.o mm. in length, move oblong-
oval in form, the head small 7

2 — Hind femora minutely and remotely punctate 3

Hind femora coarsely, more closely and conspicuously punctured, at
least toward base 6

3 — Sides of the head before the eyes more or less broadly rounded

and as prominent as the posterior canthus or nearly so ; body
broader in form • 4

Sides more oblique, coming to a more point-like canthus, which is
much less prominent than the posterior; body narrower 5

4 — Body oblong, stout and parallel (?) or feebly cuneiform and

much narrower (c?), moderately convex, strongly alutaceous
above, black, the tarsi and antennal apex rufescent ; pubescence
sparse, long, fulvous in color; head coarsely, very deeply but not
densely punctate, the sinus moderate, rather deep, triangular;
anterior canthus a little less prominent than the posterior ; pro-
thorax transverse, two-thirds wider than long, the sides broadly
rounded and converging anteriorly, becoming parallel and straight
or feebly arcuate in about basal half ; apex moderately sinuate,
three-fourths as wide as the base, the angles slightlv obtuse but
not much rounded ; surface very finely and remotely punctate
medially, becoming notably coarsely, deeply and much more
closely so toward the sides, the marginal bead moderately strong;
scutellum well developed, only moderately transverse, evenly tri-
angular; elytra two and one-half times as long as the prothorax,
the sides parallel or feebly converging and nearly straiglit to a
little behind the middle, then unusually gradually converging,
becoming straight to the acute apex ; surface nearly smooth, with
some obsolescent irregular grooves, the punctures moderately
coarse, deep and very sparse, becoming not much larger but closer
laterally; prosternum with moderate, perforate, more close-set
punctures, the process convex, margined except at the middle of
the apex ; abdomen very finely but distinctly, sparsely punctate.
Length 10.S-13.0 mm.; width 4.9-6.2 mm. California (Lake
Tahoe). Collected only by the writer so far as known.

montana Csy.

Body oblong, still less convex, somewhat piceous-black, the tarsi and
entire antennae ferruginous; upper surface alutaceous, the hairs
rather long, more close-set and conspicuous on the elytra than in
montana; head coarsely, loosely punctate, more finely and more
densely on the base of the vertex, the parabolic sinus rather deep,
the anterior canthus very broadly rounded, fully as prominent as
the posterior; prothorax formed as in montana but with the hind
angles slightly more posteriorly prominent; surface less finely,
more closely and very distinctly punctate medially, becoming
coarsely and still more closely so laterally, the lateral bead simi-

REVISION OF TENEBRIONID SUBFAMILY CONIONTINyE II3

lar; scutellum more transversely triangular; elytra parallel and
straight at the sides in almost basal two-thirds (?), more rapidly
and arcuately narrowed to the more obtuse tip ; surface broadly
undulato-rugose, strongly so apically, the punctures rather fine
but deep, much more close-set than in iiwniana, but little closer
though sensibly coarser laterally ; prosternum rather coarsely and
closely punctate, the process rounded behind, moderately strongly
margined throughout ; abdomen not very finely, rather sparsely
but conspicuously punctate medially, more minutely and remotely
laterally. Length lo.S mm. ; width 4.75 mm. (?). California
(Marin Co.). [San Jose, — LeConte ; the original type, 9.2 x 4.3
mm., is probably a male, being smaller and more slender than my
female representative; the locality " San Diego" under the orig-
inal description is an error, a different species being involved] .

subpubescens Lee.

5 — Form elongate, very moderately convex, parallel, alutaceous, the

elytra more feebly so, deep black ; hairs rather short, very sparse,
fulvous but less conspicuous than in the two preceding ; head
rather coarsely, deeply and sparsely punctate, more closely toward
the suture, the sinus not very broad, deep and angular; prothorax
more elongate, distinctly less than one-half wider than long, the
sides parallel and nearly straight fully to apical two-fifths, gradu-
ally rounding and converging thence to the apex, which is broadly
sinuate and three-fourths as wide as the base, the angles right and
only slightly blunt ; base rather strongly bisinuate ; surface some-
what finely and sparsely but strongly punctate, becoming coarsely,
conspicuously and somewhat less sparsely so laterally, the bead
moderate, a median line subimpunctate ; scutellum moderate;
elytra not quite two and one-half times as long as the prothorax,
the sides straight and parallel, gradually arcuately converging in
apical two-fifths to the rather acutely ogival tip ; surface nearly
smooth, rugulose apically, sparsely but strongly punctate, more
coarsely so laterally and apically ; prosternum rather finely but
strongly, moderately closely punctate, the process margined
throughout; abdomen minutely, rather sparsely punctate. Length
11.25 mm. ; width 4.9 mm. California (Hermitage, Mendocino
Co.). A single specimen taken by the writer parallela Csy.

6 — Form, lustre, coloration and pubescence nearly as in fnontana but

narrower; head coarsely, deeply, somewhat closely punctate, the
sinus rather deep and angulate, not very broad, the anterior can-
thus rounded and rather full, just visibly less prominent than the
posterior ; prothorax nearly as in tnontana but not so transverse,
three-fifths wider than long, the base more broadly and strongly
bisinuate, the punctures medially sparse but not so minute, very
distinct, coarse, close and very conspicuous laterally, the bead
moderately thick; scutellum rather small, sharply triangular, not
much wider than long; elytra more than two and one-half times
as long as the prothorax in the apparently female type, the sides

114 CASEY

straight and parallel to apical two-fifths, then gradually arcuate
and converging to the rather acutely ogival tip ; surface nearly as
in tnontana^ the strong and moderately large punctures less sparse,
becoming somewhat close-set and coarse laterally ; prosternum
rather coarsely but sparsely punctate, the process dilated and
rounded behind the coxae, the marginal bead rather feeble but en-
tire; abdomen rather finely but very distinctly, sparsely punctate ;
hind femora strongly, conspicuously, evenly but not very closely
punctate throughout. Length 13.2 mm.; width 5.2 mm. Cali-
fornia. Cab. Levette canonica n. sp.

Form much broader, more oblong-oval, only moderately convex, deep
black, the tarsi and antennae picescent, the upper surface dull and
strongly alutaceous, the hairs rather long, moderately sparse, ful-
vous and distinct ; head strongly, rather sparsely, not very coarsely
punctate, the sinus narrovv^, deeply angular ; anterior canthus
full and broadly rounded, as prominent as the posterior; prothorax
as in mo7itana but more narrowed anteriorly, the apex more
deeply sinuate and only three-fifths as wide as the base, the base
rather more strongly and more broadly sinuate laterally, the punc-
tures and lateral bead nearly as in canonica; scutellum as in the
latter; elytra shorter, less than one-half longer than wide, the
sides parallel and straight to apical two-fifths, then moderately
rapidly converging and arcuate to the subparabolic apex ; surface
smooth, feebly undulato-rugose posteriorly, rather finely but dis-
tinctly, sparsely punctate, more coarsely though still not very
deeply and but little less sparsely laterally; prosternum strongly,
moderately closely punctate, the process feebly dilated and rounded
behind the coxas, convex, the beaded margin obsolescent at apex ;
hind femora strongly and rather closely punctured toward base
but sparsely and less coarsely so thence distally; abdomen tinely,
feebly and sparsely punctulate. Length 11.7 mm.; width 5.8
mm. California (South Fork of the Kaweah River).

perspicua n. sp.

7 — Body oblong-suboval, moderately convex, dull and alutaceous,
rather grayish-black, the legs, antennce, mouth-parts and clypeus
rufescent ; pubescence coarse, moderately long, fulvous and dis-
tinct though rather sparse ; head somewhat coarsely and closely
perforato-punctate, the sinus moderately narrow and parabolic,
the anterior canthus narrowly rounded, about as prominent as the
posterior ; prothorax one-half wider than long, the sides broadly
rounded, strongly converging apically, becoming nearly parallel
toward base, the apex deeply sinuate, barely three-fifths as wide
as the base, the angles right and scarcely even blunt, very dis-
tinct ; surface rather sparsely but strongly punctate, coarsely and
less sparsely so laterally, the marginal bead fine, arcuate near the
base; scutellum moderate, picescent; elytra about two and a third
times as long as the prothorax, rather obtusely ogival with strongly
arcuate sides in apical two-fifths, parallel thence to the base ; sur-
face nearly smooth, undulato-rugose apically, strongly, somewhat

REVISION OF TENEBRIOMD SUBFAMILY CONIONTIN^ II5

sparsely punctate, becoming moderately coarsely and rather less
sparsely so laterally; under surface and hind femora finely,
sparsely punctulate, the presternum more strongly, with the
process unmargined except feebly along the coxae ; tarsi unusually
slender. Length 7.0-S.o mm.; width 3.4-3.9 mm. California
(San Diego). Two specimens taken by the author and one sub-
sequently received, the latter having the scutellum much more
transversely triangular and the body a little more convex.

parviceps Csy.

Body somewhat less cylindrical, oblong-oval, distinctly more de-
pressed, nearly similar in coloration, lustre and pubescence; head
even smaller and less developed than in any other species, less
coarsely but strongly, rather closely punctate, the sinus rather
broader, the anterior canthus more obliquely pointed, as promi-
nent as the posterior ; prothorax much shorter but otherwise
nearly similar, two-thirds wider than long, the deep apical sinus
parabolic, almost similarly sculptured but not quite so coarsely,
the marginal bead fine, arcuate at base ; scutellum rather small,
similar, only feebly transverse and sharply triangular ; elytra about
two and one-half times as long as the prothorax, less than one-
half longer than wide, more gradually and acutely ogival in apical
two-fifths; surface more strongly undulato-rugulose, especially
behind, with a medial vaguely eroded line on each, the punctures
rather smaller and less sparse than in parviceps ; prosternum
rather strongly, moderately closely punctured, the process strongly
margined, obsolescently so around the broadly arcuate apex; hind
femora and abdomen finely and rather sparsely but strongly and
distinctly punctate. Length 7.8 mm. ; width 3.7 mm. Califor-
nia (Poway, San Diego Co.) filiola n. sp.

The last two species of the table form a rather isolated type,
peculiar to the Sonoran regions of California.

Group VL — Type setosa.

The following five species form a natural group leading to
eschscholtzi and allies, having the same rather stout and oblong
convex body, but with the elytral sculpture comparatively fine
and always less coarse or rugose and the more or less con-
spicuous vesliture erect and bristling on the elytra ; they have
a geographic distribution differing greatly from that of the
eschscholtzi group but coincident with that of the ovalis group
and may be thus defined : —

Prosternal process not or but feebly dilated behind the coxje 3

Prosternal process more constricted at the coxie and always more or
less strongly dilated apically 5

2 — Elytral vestiture short 3

Il6 CASEY

Elytral vestiture long, shaggy and very conspicuous 4

3 — Body rather narrowly oblong or subelongate, convex, deep black,
the antennae black, the legs piceous distally ; surface rather shin-
ing, the elytral setae small and not very conspicuous; head vv^ell
developed, rather finely and sparsely punctate, the sinus rather
deep, sharply parabolic, the lobes broady rounded, the anterior
canthus angularly rounded, as prominent as the posterior; pro-
thorax transverse, nearly three-fourths wider than long, the sides
broadly rounded, more converging anteriorly, becoming almost
parallel toward base, the apex modei'ately sinuate, fully three-
fifths as wide as the base, with the angles evidently rounded ;
surface very finely, sparsely punctate, becoming gradually rather
coarsely and closely so laterally, the bead rather fine ; scutellum
moderate ; elytra one-half longer than wide, parallel, very obtusely
and broadly, parabolically rounded behind, generally almost
smooth but sometimes rugose, with obsolete indications of longi-
tudinal tumescent lines occasionally, strongly, rather closely
punctate, more coarsely and still more closely so laterally and
also apically, where the surface is more opaque ; prosternum
finely, rugosely sculptured, the process polished and more
sparsely and strongly punctate, strongly margined at the sides
but immarginate at tip ; hind femora strongly though very
sparsely punctate, the abdomen minutely, remotely so, more
strongly toward the sides ; tarsi slender. Length 10. 0-10.3 mm.;
width 4.S-5.0 mm. Idaho (Co2ur d'Alene). One specimen is
labeled " Cal." but probably in error setosa Csy.

Body somewhat stouter and more abbreviated, deep black, moderately
shining, the bristling vestiture distinct, fulvous; head rather
coarsely and densely punctured toward the suture, coarsely
apically and sparsely and finely from before the eyes to the base,
the sinus broad, parabolic, the anterior canthus rather obtusely
pointed, much less prominent than the posterior; oblique sides
feebly emarginate at the ends of the oblique sutures ; prothorax
more transverse, nearly four-fifths wider than long, the sides sub-
evenly and moderately arcuate throughout, becoming parallel
only near the base; apex broadly sinuate, two-thirds as wide as
the base, with the angles right and but slightly blunt; surface
finely, sparsely punctate, becoming broadly rather coarsely but
not much more closely so toward the sides, the bead rather fine,
the base broadly, evenly bisinuate; scutellum rather small, trans-
versely triangular; elytra parallel, very broadly and obtusely
rounded in apical third, two-fifths longer than wide, nearly
smooth, finely, i-emotely punctate, more coarsely but shallovvly
and not much less sparsely so toward the sides, subopaque pos-
teriorly; prosternum rather strongly punctate, the process broad,
subparallel, unmargined except finely along the coxae; hind
femora and abdomen rather finely but more or less strongly,
sparsely punctate. Lengtli 10. o mm. ; width 5.3 mm. Utah
(Clear Lake), — IL F. Wickham obtusa n. sp.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN-(E II7

4 — Form still stouter, oblong-oval, convex, deep black, somewhat

alutaceous, the elytra more shining; head rather finely, very
sparsely punctate, more closely so near the transverse suture, the
sinus moderately broad, deeply subangulate, the anterior canthus
full, somewhat less prominent than the posterior; prothorax
three-fourths wider than long, the sides very feebly converging
and almost straight, or with a feeble ante-basal sinus, from the
base about to the middle, then rounding, becoming strongly con-
verging and just visibly arcuate to the apex, which is broadly
sinuate and two-thirds as wide as the base, the angles right and
rather narrowly rounded; surface very minutely, sparsely punc-
tate, becoming broadly somewhat coarsely but not very closely so
laterally; base somewhat more strongly bisinuate than in obtusa;
scutellum small, transversely triangular; elytra as in obtiisa but
barely a third longer than wide, nearly smooth, subopaque pos-
teriorly, having numerous fine and generally subtransverse, an-
astomosing creases throughout, the punctures sparse but rather
strong, becoming quite close-set and subrugulose or with trans-
versely sublineate extensions laterally ; prosternum rather finely
but strongly punctate, the process long, the post-coxal part oblong
with but feebly arcuate sides and rounded tip, broadly constricted
at the cox£e, inimargined except finely along the coxae ; hind
femora and abdomen very sparsely punctate. Length 10.8 mm. ;
width 5.75 mm. Nevada (Carson City), — H. F. VVickham.

lanuginosa n. sp.

5 — Body oblong, deep black, the tarsi piceous ; upper surface alu-

taceous, the elytra more shining though posteriorly opaque, head
rather sparsely but somewhat strongly punctured, finely toward
base, the sinus nearly as in the preceding, the anterior canthus
rather broadly rounded and almost as prominent as the posterior;
prothorax two-thirds wider than long, the sides subevenly and
moderately arcuate throughout, not quite becoming parallel even
at base, the apex rather deeply sinuate, scarcely more than three-
fifths as wide as the base, the angles right and very distinct,
scarcely at all blunt ; surface sparsely, extremely minutely punc-
tate, becoming broadly quite coarsely but scarcely less sparsely so
laterally; base rather strongly sinuate near each side ; scutellum
small; elytra not quite one-half longer than wide, parallel,
obliquely and rapidly narrowed in posterior third to the apex,
which is much more angulate than in the preceding, the pu-
bescence very conspicuous, fulvous; surface nearly smooth, the
anastomosing fine creases only distinct posteriorly, finely, remotely
punctate, more strongly and less sparsely so laterally but still
comparatively finely; prosternum rather strongly punctate, un-
margined except finely along the coxa? ; hind femora and abdomen
rather finely but unusually distinctly, though remotely, punctate;
tarsi moderately slender. Length 10. 9-1 1.5 mm.; width 5.7—
5.85 mm. Utah (Marysvale), — H. F. Wickham.

wickhami n. sp.

Il8 CASEY

Body nearly similar in form, rather strongly shining throughout, pale
piceo-rufous, the elytra, under surface, legs and antennae still
paler, ferruginous — possibly partially owing to immaturity; —
pubescence abundant and conspicuous posteriorly though only
moderate in length as in tvickhami ; head somewhat coarsely and
closely punctate thi'oughout, the sinus broadly angulate, the lobes
rounded, impressed; anterior canthus rather broadly rounded,
visibly less prominent than the posterior ; prothorax fully three-
fourths wider than long, the sides broadly rounded and strongly
converging anteriorly, becoming parallel and nearly straight in
about basal half, the apex rather shallowly sinuate, fully two-
thirds as wide as the base, the angles right and only slightly
blunt; surface very finely, rather closely punctate, becoming
coarsely and unusually densely so very broadly toward the sides,
the bead fine ; basal angles slightly prolonged, right ; scutellum
moderate; elytra one-half longer than wide, the sides gradually
converging and broadly, evenly arcuate from slightly behind the
middle to the narrowly parabolic apex, nearly smooth, finely,
rather sparsely, submuricately punctate, becoming rather coarsely,
densely, muricately so, with the punctures transversely subcon-
fluent, laterally ; prosternuni strongly, closely punctate, the process
nearly circular in form behind the very strong intercoxal constric-
tion, unmargined except moderately along the coxcE ; femora and
abdomen as in ivickhami \>wt less strongly punctured. Length
1 1.2 mm. ; width 5.4 mm. Washington State — pubifera n. sp.

Wickhami is a larger species than obtusa, with much less
transverse prothorax and angulate, and not broadly rounded,
elytral apex ; the posterior part of the elytra in -pubifera differs
again from anything else known in the group, and this is an
unusually well differentiated species, in which the pale colora-
tion is in all probability not entirely due to immaturity, a parallel
case being J'af-alloiii'ca, of the following group.

Group VII. — Type eschscholtzi.

This is one of the larger groups of the genus, and, at the same
time, one of the most restricted in habitat. It occurs exclu-
sively in the coast regions from San Francisco Bay probably
through Oregon, being geographically coincident with the
viatica group ; to the northward and eastward of this region it
is replaced b}' the sctosa and ovalis groups, and, to the south-
ward, by the ahdominalis group. The species are oblong or
oblong-suboval, usually strongly convex in form and generally
have very coarse and deep elytral punctures, separated by
convex interspaces, giving to the surface a more or less pro-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ II9

nounced rugosity. Pubescence is frequently noticeable but is
always very short, sparse and decumbent and never so con-
spicuous as in the two preceding groups. The various forms in
my collection may be known by the following characters : —

Elytra! punctures bearing short decumbent and evident, though more
or less inconspicuous hairs 2

Elytra! punctures virtually nude or l^earing ver}'^ minute setaj, usually
not projecting beyond tlieir limits 17

2 — Larger and strongly convex species, seldom less than ii.omm.

in length 3

Smaller, less convex species, though rather more convex than in sub-
pjibescois and allies, never much more than 10. o mm. in length
and generally very much less 13

3 — Species occurring in California, probably in all cases near San

Francisco Bay ; prosternal process strongly beaded throughout the
periphery in all the specimens at hand ; anterior canthus fully as
prominent as the posterior 4

Species of Oregon; prosternal process margined feebl}" at the sides
but not at apex ; anterior canthus less prominent than the pos-
terior 12

4 — Sides of the prothorax rounding and strongly converging anteri-
orly, becoming parallel and almost straight in about basal half... 5

Sides of the prothorax evenly and moderately or subevenly arcuate from
base to apex , 10

5 — Clypeal sinus only moderate in width, deep and parabolic 6

Clypeal sinus broad, very shallow, evenly rounded in circular arc — 9

6 — Prosternal process broadly and rather abruptly arcuato-truncate at

apex 7

Prosternal process narrower and evenly, semicircularly rounded at
apex S

7 — Form very broadly oblong, deep black, moderate!}' shining, the

anterior parts alutaceous ; head strongly but somewhat loosely and
irregularly punctured, the anterior canthus rounded though not
very broadly ; antennae slender, rufescent at tip as usual ; protho-
rax short, nearly four-fifths wider than long, the apex moderately
sinuate, two-thirds as wide as the base, the angles somewhat ob-
tuse and well rounded, the base transverse, not medially lobed,
becoming very feebly and posteriorly sinuato-oblique laterally;
surface finely but strongly, sparsely and somewhat unevenly punc-
tate, gradually quite coarsely and more closely so laterally, the
bead coarse and polished ; scutellum large, very transversely tri-
angular; elytra barely one-half longer than wide, parallel, the
sides very evenly rounded and converging in posterior two-fifths,
the apex obtusely ogival ; surface very coarsely, moderately closely
punctate and rugulose ; prosternum strongly but rather sparsely

I20 CASEY

punctate and with short pubescence, the process wide, subpar-
allel ; hind femora rather strongly but sparsely, the abdomen
minutely and remotely, punctate. Length 14.2 mm. ; width 6.4
mm. (9). San Francisco Co., — Chas. Fuchs expansa n. sp.

Form much narrower and relatively more convex, polished throughout
and veiy much more shining, deep black; head rather finely but
strongly, moderately closely punctate, the anterior canthus rather
narrowly rounded though not at all pointed; prothorax much less
transverse, but little more than one-half wider than long, the apex
moderately deeply sinuate, nearly two-thirds as wide as the base,
the angles obtuse but rather narrowly rounded, the base broadly
arcuate medially, just visibly and transversely sinuate laterally;
surface finely, rather distinctly but sparsely punctate, rather
strongly though not coarsely, sparsely so laterally, the bead moder-
ately thick ; elytra of peculiar form, fully one-half longer than
wide, the sides gradually evenly arcuate and converging posteriorly
to the acutely ogival apex, becoming parallel only toward base,
very coarsely but rather sparsely punctured, densely so apically,
the surface rugose; prosternum polished, very minutely, remotely
and inconspicuously punctate, the process more constricted inter-
coxally than in expansa and with the apical angles more rounded ;
hind femora and abdomen very finely, sparsely punctate. Length
12.7 mm. ; width 5.7 mm. San Francisco. A single specimen
taken by the author in the suburbs franciscana n. sp.

8 — Body very broadly oblong, rather strongly convex, deep black,
polished, the anterior parts somewhat alutaceous ; head with rather
small but strong, moderately close punctures, the anterior canthus
rather narrowly rounded ; prothorax short, fully three-fourths
wider than long, the sides more evenly rounded anteriorly than in
expansa^ the apex similar but with less thickened bead laterally,
three-fifths as wide as the base, the latter similarly truncate but
less prolonged backward laterally, being very feebly, transversely
sinuate toward the sides; surface finely, sparsely punctate, be-
coming strongly and somewhat coarsely though still sparsely so
laterally, the marginal bead rather fine; scutellum smaller, much
less transversely triangular; elytra nearly as in cxpaiisa but with
the sides becoming arcuately convergent behind at about basal
third and with the apex more acutely ogival, the punctures simi-
larly very coarse and the surface rugose ; prosternum very minutely,
sparsely and inconspicuously punctate, the process but feebly
constricted intercoxally ; hind femora and abdomen very finely,
remotely punctate. Length 12.4 mm.; width 5.8mm. Near
San Francisco, — G. W. Dunn ..truncata n. sp.

Body much narrower and more elongate, parallel, similarly convex,
deep black, strongly shining, the anterior parts slightly alutaceous ;
head similarly though much more sparsely punctate, the anterior
canthus more rounded and less oblique; antennae not very slender,
the outer joints not much enlarged; prothorax not more than one-

REVISIOIS OF TENEBRIONID SUBFAMILY CONIONTIN^ 121

half wider than long, the anterior sinus moderately shallow, nearly
three-fourths as wide as the base, the angles obtusely rounded,
the base transverse, just visibly and very broadly bisinuate; sur-
face minutely, very remotely punctate, more strongly but still com-
paratively finely and sparsely so laterally, the bead rather fine;
scutellum moderate, transversely triangular; elytra fully one-half
longer then wide, the sides parallel and straight to well behind
the middle, then gradually rounded and converging to the rather
acutely ogival apex ; surface of each with a few remote and feebly
tumescent lines, of which one margins the suture, the punctures
coarse but unusually sparse, with less convex interspaces, denser
and more rugose posteriorly ; prosternum sparsely pubescent, finely
but rather strongly, sparsely punctate, the process broader than in
trziticata, feebly constricted intercoxally ; hind femora and abdo-
men very finely, remotely punctate. Length 13.0 mm.; width
5.7 mm. (cf). Cab. Levette SUturalis n. sp.

9 — Form moderately stout, convex, alutaceous, the elytra much more

shining, deep black; head I'ather finely and sparsely but strongly
punctate, the anterior canthus obliquely and obtusely pointed ;
antennse rather slender ; prothorax three-fifths wider than long,
the apex shallowly sinuate, two-thirds as wide as the base, the
angles obtuse and rather broadly rounded, the base very broadly
and feebly bisinuate; surface rather sparsely but not very finely,
strongly punctate, becoming somewhat coarsely and less sparsely
so laterally, the bead moderately fine ; scutellum moderate, sharply
triangular, not very transverse; elytra one-half longer than wide,
the sides gradually rounding and converging from rather before
the middle posteriorly, the apex acutely ogival ; surface coarsely
but not very closely punctate, the interspaces not notably convex,
except apically, where they are narrower; prosternum very finely,
sparsely punctate, the process almost impunctate and subparallel,
broadly rounded at tip ; hind femora and abdomen minutely, re-
motely and very inconspicuously punctate. Length ii.omm.;
width 4.9 mm. Cab. Levette audax n. sp.

10 — Tarsi unusually slender. Form oblong-oval, rather shining

throughout, deep black ; head somewhat coarsely punctate, very
sparsely posteriorly, moderately closely anteriorly, the sinus not
very broad, deeply parabolic ; anterior canthus oblique and rather
pointed but blunt ; prothorax shorter than in the three following,
three-fifths wider than long, the apex only very moderately sinuate,
three-fifths as wide as the base, with the angles obtusely rounded ;
sides unusually converging from base to apex, broadly arcuate ; base
rectilinearly truncate, becoming conspicuously oblique posteriorly
at the sides ; surface very sparsely but somewhat strongly punctate,
more coarsely but still sparsely so laterally, the bead modei^ately
thick, curving strongly at base entirely about the hind angles ;
scutellum very transversely triangular, rather large ; elytra as
wide as the prothorax, one-half longer than wide, the sides
strongly rounding behind the middle, the apex obtusely ogival ;
Proc. Wash. Acad. Sci., April, 190S.

122 CASEY

surface very coarsely punctate and rugulose, the punctures well
separated except apically ; prosternum finely, sparsely punc-
tate, the process rather wide, only very feebly constricted at the
coxae, semicircularly rounded at apex ; hind femora and abdomen
very finely, remotely punctate. Length 11.3 mm.; width 5.2

mm. (cf). Precise locality not recorded convergens n. sp.

Tarsi moderately stout as usual; prothorax less transverse 11

II — Form evenly elongate-elliptical, strongly convex, deep black,
alutaceous, the elytra polished; head large, nearly half as wide as
the prothorax, strongly, rather closely punctate, the sinus only
moderately deep and wide, broadly parabolic, the lobes broadly
rounded, the anterior canthus narrowly rounded, very prominent,
somewhat more so than the posterior; prothorax unusually elon-
gate, less than one-half wider than long, the sides evenly con-
verging from base to apex, evenly and moderately arcuate, the
apex shallowly sinuate, fully two-thirds as wide as the base, the
angles right and narrowly rounded or blunt, the base broadly^
evenly, very feebly bisinuate ; surface finely and sparsely but
rather strongly punctate, somewhat coarsely and less sparsely so
laterally, the bead rather coarse, perfectly straight to the basal
angles; scutellum moderate, sharply triangular, not very trans-
verse ; elytra less than one-half longer than wide, scarcely more
than twice as long as the prothorax, the sides becoming rounded
behind the middle, the apex very obtuse from above; surface
very declivous posteriorly, very coarsely, moderately closely
punctate and rugose; prosternum very finely, sparsely punctate,
the process wide, broadly rounded at tip, feebly constricted; hind
femora finely but distinctly, remotely, the abdomen minutely,
punctate. Length 12.0 mm.; width 5.75 mm. San Francisco.
A single specimen, apparently female, from the suburbs, collected
by the writer symmetrica n. sp.

Form somewhat more oblong and scarcely so convex, deep black,
polished throughout ; head smaller, distinctly less than half as
wide as the prothorax, less coarsely, more sparsely punctate, the
sinus narrower and deeper, parabolic, the lobes less broadly
rounded, the anterior canthus similarly oblique and narrowly
rounded and very prominent though not more so than the poste-
rior; prothorax nearly as elongate, barely one-half wider than
long, nearly similar but more narrowed from base to apex, the
latter similar but only three-fifths as wide as the base, with the
angles more broadly rounded, the sides of the base more posteri-
orly oblique; surface very sparsely, rather finely but strongly
punctate, gradually rather coarsely and less sparsely so laterally
though becoming dense near the hind angles, the bead similar but
rather less coarse; scutellum nearly similar; elytra longer, one-
half longer than wide, more ogival at tip from above ; surface
less declivous posteriorly, a little less coarsely but otherwise sim-
ilarly punctate and rugose ; prosternum minutely, sparsely punc-
tate, the process semicircularly rounded at tip, moderately con-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 1 23

stricted ; hind femora rather coarsely, closely and conspicuously
punctured basally, more remotely thence to the apex, the abdomen
very minutely and remotely; hind tarsi much shorter than in
sy7n9netrica. Length 10.8 mm.; width 5.3 mm. (c?). Exact
locality unrecorded anxia n. sp.

Form moderately stout, oblong, convex, alutaceous, the elytra pol-
ished, deep black ; head not very coarsely but strongly, rather
sparsely punctate, the sinus broad, moderately deep, parabolic,
the lobes obtuse though rather narrowly rounded, the anterior
canthus nearly as in anxia but more obtusely rounded ; prothorax
shorter, a little more than one-half wider than long, the sides sub-
evenly and much more strongly arcuate, becoming parallel near
the base, strongly converging anteriorl}', the apex shallowly sin-
uate, fully three-fifths as wide as the base, the angles right and
moderately rounded, a good deal deflexed, the base broadly, feebly
bisinuate ; surface sparseh', very finely punctate, gradually strongly,
moderately sparsely to rather closely and coarsely so toward the
sides, the bead moderately thick, straight to the base; scutellum
as in the two preceding ; elytra as in anxia though still a little
more acutely ogival at apex, the punctures similarly very coarse
but w^ell separated and the surface rugose ; prosternum similar,
the process subparallel (9), semicircularly rounded at tip; hind
femora strongly and conspicuously punctured toward base, the
abdomen with small but distinct, sparse punctulation. Length
II. 5-13.0 mm.; width 5.4-6.1 mm. Exact locality unrecorded.
[Benicia, — LeConte] affinis Lee .

A — Nearly as in affinis but more narrowly elongate-suboval in
form, the punctures toward the sides of the prothorax much sparser
and less coarse, the sides of the latter less strongly and more un-
evenly arcuate, being relatively more strongly arcuate apically,
the set« of the elytral punctures altogether inconspicuous except
apically, where they are much shorter, passing but little beyond
the confines of the punctures ; vmder surface nearly similar,
except that the hind femora are minutely, sparsely and incon-
spicuously punctate throughout. Length 13.0 mm.; width
5.78 mm. (9). "Cal." patruelis n. subsp.

12 — Body oblong, convex, deep black throughout, the antennse scarcely
even picescent apically, stout ; surface feebly alutaceous anteriorly,
the elytra more shining ; head coarsely but loosely punctate, the
sinus wide, rather deep, parabolic, the anterior canthus oblique,
subacute, scarcely prominent ; prothorax one-half wider than
long, the sides rounded and convergent anteriorly, becoming
faintly sinuate and parallel in basal three-fifths, the apex shal-
lowly sinuate, two-thirds as wide as the base, the latter broadly
and rather strongly bisinuate, the basal angles unusually pro-
longed posteriorly and rather acute, the anterior obtuse though
not very broadly rounded ; surface sparsely but strongly punc-
tate, unusually coarsely, moderately closely and somewhat unevenly

124 CASEY

SO laterally, the bead moderately coarse, not at all cur\'ed at base ;
scutellum moderate, not very transverse, convexly declivous an-
teriorly ; elytra scarcely one-half longer than wide, parallel, the
sides rounding rather abruptly near apical third, thence obliquely
converging and less arcuate to the ogival apex ; surface very
coarsely, deeply but not very closely punctate, the interspaces not
notably convex, rugose apically ; prosternum coarsely, strongly,
very closely and conspicuously punctate throughout, pubescent,
the process but feebly constricted, strongly rounded at apex ; hind
femora strongly, closely punctate basally, more sparsely elsewhere,
the abdomen not very finely and unusually strongly so. Length
ii.o mm.; width 5.3 mm. (d^). Oregon (Albany), — H. F.
Wickham oregona n. sp.

13 — Base of the prothorax abruptly posteriorly oblique at the sides,

the angles obviously prominent posteriorly 14

Base of the prothorax broadly, subevenly bisinuate, not abruptly oblique
laterally, the angles barely at all prominent posteriorly ; antennae
notably slender, the joints much elongated; size very small 16

14 — Head very sparsely but unevenly punctate. Body oblong-oval,

moderately convex, subalutaceous, not very deep black, the legs
and antennce rufescent ; pubescence sparse, fulvous ; head moder-
ately large, the punctures somewhat coarse, more aggregated near
the transverse suture, very sparse elsewhere, the sinus rather
narrow, deeply parabolic, the anterior canthus broadly rounded,
about as prominent as the posterior ; antennae rather slender, the
joints moderately elongate ; prothorax one-half wider than long,
the sides rounded and converging anteriorly, becoming gradually
very feebly arcuate and almost but not quite parallel through basal
three-fifths ; apex rather shallowly sinuate, almost two-thirds as
wide as the base, with the angles right and only slightly blunt ;
surface rather finely but stronglv, remotely and unevenly punc-
tate, with a rather pronounced median impunctate line, gradually
rather strongly but still not closely punctate toward the sides, the
bead rather fine, not curved at base ; scutellum moderate, trans-
versely and sharply triangular ; elytra less than one-half longer
than wide, rounding at the sides behind the middle, the apex
moderately obtusely ogival ; surface only moderately coarsely,
rather sparsely punctate, strongly rugose posteriorly ; prosternum
finely but strongly, perforately and not very closely punctate, the
process strongly rounded at tip, not margined except feebly
toward its base ; hind femora very minutely, remotely punctate,
the abdomen somewhat more strongly though very finely.
Length 10.5 mm.; width 4.75 mm. (?)• " Cal."

extricata n. sp.

Head more closely, subevenly punctate throughout 15

15 — Form nearly as in extricata^ somewhat brownish-black, the tarsi
and anteimai rufescent; lustre rather shining, the pubescence very
obvious, fulvous ; head with the sinus moderately wide, not very

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 1 25

deep, broadly parabolic, the anterior canthus oblique, subacute,
as prominent as the posterior ; antennae slender, the joints notably
elongate ; prothorax as in extricata but shorter, fully three-fifths
wider than long, the punctures similar but much more close-set
throughout and without median impunctate line ; scutellum sim-
ilar ; elytra as in extricata but narrower and more nearly one-
half longer than wide, more gradually acute at apex, still less
coarsely and more sparsely punctate, similarly rugose posteriorly ;
prosternum finely but very strongly, moderately closely punctate
and puberulent, the process very strongly beaded along the sides
to the apex but not around the latter ; hind femora excessively
minutely and remotely punctulate throughout, the abdomen not
very finely though sparsely, strongly and conspicuously punctate.
Length 9.5 mm. ; width 4.35 mm. (9). San Francisco Co.

marginata n. sp.
Form narrower and more elongate, moderately convex, alutaceous, the
elytra polished, somewhat piceous black, the tarsi piceous and the
antennal apex rufous ; head moderate in size, strongly, rather
closely punctate, sparsely pubescent, the sinus moderately narrow
and deep, parabolic, the lobes broadly rounded, the anterior can-
thus oblique though obtusely rounded, as prominent as the pos-
terior ; prothorax three-fifths wider than long, the sides rounding
and convergent anteriorly, becoming subparallel and straight or
sometimes with a feeble sinus in more than basal half, the apex
rather shallowly sinuate, two-thirds as wide as the base, the angles
slightly obtuse but only narrowly rounded ; surface finely though
strongly, only moderately sparselv punctate, strongly but not
much more closely toward the sides, the bead not very thick
though prominent, slightly curving inward at base ; scutellum
moderate, transversely triangular ; elytra one-half longer than
wide and somewhat gradually acutely ogival behind (?), or two-
fifths longer than wide and more obtusely parabolic. at apex (cf),
rugose posteriorly, the punctures rather coarse and well separated,
notably unequal ; prosternum finely, sparsely punctate, puberu-
lent, the process rather narrow, somewhat strongly margined to,
but scarcely about, the rounded apex ($), or a little wider and
finely margined and only at its base (cJ*) ; hind femora and abdo-
men minutely, sparsely and more or less inconspicuously punc-
tate. Length 7.7-9.0 mm. ; width 3.5; -4.0 mm. San Francisco.
Taken in the suburbs rather plentifully by the writer and also
received from the County, collected by Mr.Fuchs.

nemoralis Esch.

Form rather shorter than in nemoralis^ still smaller in size, similar in
coloration and lustre ; head rather small, strongly, somewhat
closely punctate, the sinus moderately wide and deep, circularly
rounded, the anterior canthus oblique, subacute, less prominent
than the posterior ; antennae notably thick, the joints only moder-
ately elongate ; prothorax large, nearly three-fifths wider than
long, the sides broadly arcuate, more strongly so and very con-

126 CASEY

vergent anteriorly, slightly wider near the middle than at base, the
apex not more than three-fifths as wide as the base, the angles
obtuse and broadly rounded, more deflexed than usual ; surface
very finely, rather sparsely punctate, more strongly but still not
coarsely and almost equally sparsely so toward the sides, the bead
rather fine, slightly curving at base ; scutellum and elytra nearly
as in nemoralis^ the latter scarcely one-half longer than wide,
gradually arcuately narrowed behind the middle, the apex ogival,
though narrowly obtuse ; surface and punctures uneven, nearly as
in nemoralis ; under surface nearly similar. Length 7.25 mm. ;
width 3.3 mm. " Cal." parva n. sp.

16 — Body oblong, rather stout, very small in size, similar to the pre-

ceding in coloration but more shining, the anterior parts not at all
alutaceous ; head moderately large, strongly, subevenly but not
very closely punctate, the sinus nearly as in parva^ the anterior
canthus much more obtusely rounded and scarcely at all less prom-
inent than the posterior; prothorax fully three-fifths wider than
long, nearly as in nemoralis^ the apex more than two-thirds as
wide as the base ; surface finely but distinctly, sparsely punc-
tate, more strongly and less sparsely toward the sides, the bead
moderate, curving very slightly at base, the basal angles very
blunt ; scutellum rather small ; elytra short, two-fifths longer than
wide, otherwise nearly as in parva though smoother, rugulose api-
cally, the punctures not coarse but strong, well separated and
rather unequally distributed ; prosternum finely but strongly,
sparsely punctate, the process rather narrow, evenly and strongly
convex longitudinally, margined only at base ; hind femora and
abdomen very minutely, sparsely and inconspicuously punctulate.
Length 6.7 mm.; width 3.1 mm. (c?)- San Francisco Co., —
Chas. Fuchs minuta n. sp.

17 — Body deep black in color throughout, inhabiting the mainland

exclusively 18

Body piceous to pale testaceous in color; habitat insular 26

18 — Elytral punctures only moderately coarse, relatively fine for the

present group 19

Elytral punctui-es very coarse and generally deep 23

19 — Small species, never as much as lo.o mm. in length 20

Larger species never less than 10. o mm. in length 21

20 — Elytra unequally punctate, very remotely so, with the surface

nearly smooth, becoming more coarsely and closely with the sur-
face very rugose apically. Body oblong-oval, moderately convex,
deep black, the tarsi and antennal base and apex piceo-rufous, very
highly polished throughout ; head strongly but sparsely punctate,
the sinus only moderate in width, deep and parabolic, the anterior
canthus obliquely rounded, as prominent as the posterior ; antennae
moderately thick ; prothorax fully two-thirds wider than long, the
sides broadly rounded and converging before the middle, beconi-

REVISION OF TENEBRIONID SUBFAMILY* CONIONTIN^ I27

ing feebly diverging and almost straight thence to the base, the
apex shallowly sinuate, three-fifths as wide as the base, the angles
right and only slightly blunt ; surface rather finely but strongly,
remotely punctate, becoming coarsely and much less sparsely so
laterally, the bead thick, much thickened apically, curving but little
at base ; scutellum moderate, transversely triangular ; elytra not
quite one-half longer than wide, rounded at the sides and narrowed
to the obtusely ogival apex in posterior two-fifths, the punctures
deep but uneven in size ; prosternum strongly, rather closely punc-
tate, the process unmargined except feebly at base ; hind femora
minutely, sparsely, the abdomen rather strongly, somewhat less
sparsely, punctate. Length 9.2 mm. ; width 4.25 mm. " Cal."

perpolita n. sp.

Elytra almost equally and much less sparsely punctate, less smooth
though almost similarly rugose behind. Body more elongate, sim-
ilarly convex, less shining, the anterior parts sometimes faintly
alutaceous, deep black, the antennie and distal parts of the legs
picescent or rufous ; head strongly but loosely, unequally punc-
tate, the sinus moderately deep, not wide, the anterior canthus
obliquely subangulate or obtuse, fully as prominent as the poste-
rior ; antennae moderate ; prothorax one-half wider than long,
formed as in perpolita but less narrowed at the shallowly sinuate
apex, the latter two-thirds as wide as the base, the sides of which
are more abruptly oblique, with the hind angles not more pro-
duced posteriorly but more narrowly and abruptly so ; bead not so
thick and not thickened apically ; punctures much finer and a little
closer throughout in the same relative parts of the surface ; scu-
tellum and elytra nearly similar, the latter somewhat less rapidly
narrowed behind ; punctures strong, somewhat uneven, each^with
a small yellowish-silvery seta, more distinct posteriorly but not
sufficiently long to constitute pubescence ; under surface similar
throughout, except that the abdomen is much more minutely,
sparsely punctulate. Length 8.6-9.7 I'^i'^i- ; width 4.0-4.6 mm.
(d^, ?). San Francisco. Two specimens taken by the writer in
the suburbs, one example from the County, taken by Mr. Fuchs,
and several from the Levette cabinet pudica n. sp.

21 — Sides of the prothorax subangulate though rounded at the middle.
Form oblong-elongate, convex, deep black throughout, alutaceous,
the elytra rather strongly shining ; head very finely, rather sparsely,
subevenly punctulate, the sinus rather small but deep, subangular,
the lobes obtuse, the anterior canthus obtuse, very nearly as prom-
inent as the posterior ; antennae moderately thick ; prothorax almost
four-fifths wider than long, the sides sti'ongly converging though
only slightly arcuate anteriorly, parallel and straight in about basal
half, the apex moderately sinuate, even less than three-fifths as
wide as the base, the angles obtuse but narrowly rounded, the
base moderately and rather narrowly, evenly sinuate at each side ;
surface remotely, very minutely punctulate, with an impunctate
medial linear area, the punctures gradually stronger and closer

128 CASEY

but still rather small laterally, the bead fine, straight at base ; scu-
tellum large, transversely triangular ; elytra somewhat more than
one-half longer than wide, parallel, rather strongly rounded and
narrowed to the ogival apex in barely apical two-fifths, the oblique
sides becoming nearly straight, the base not quite as wide as the
thoracic base, the surface somewhat vnieven, deeply and somewhat
finely, densely rugose on the declivity, the punctures small, moder-
ately close-set, coarser posteriorly, the setae very minute and incon-
spicuous ; prosternum smooth, minutely, sparsely punctate, the
process large, much constricted intercoxally, margined throughout ;
hind femora and abdomen very minutely, sparsely punctulate.
Length 12.0 mm.; width 5.68 mm. Locality not recorded but
probably from northern California callida n.sp.

Sides of the prothorax subevenly rounded from base to apex 22

22 — Body oblong-suboval, rather strongly shining and deep black
throughout ; head finely, somewhat closely punctate, the sinus
moderately wide, rather deeply parabolic, the lobes broadly
rounded, the anterior canthus evenly rounded, fully as prominent
as the posterior ; prothorax almost four-fifths wider than long, the
sides broadly rounded and more converging apically, becoming a
little less rounded and almost parallel basally, the apex moderately
sinuate, two-thirds as wide as the base, with the bead strong lat-
erally, the angles right and moderately rounded ; base rather
strongly sinuate laterally, broadly lobed ; surface very finely,
rather sparsely punctate, subimpunctate along the middle, more
strongly but not very closely or coarsely laterally, the bead rather
fine, not arcuate at base ; scutellum small ; elytra not quite one-
half longer than wide, the sides converging and rounded behind
the middle to the acutely ogival apex ; surface with two or three
narrow faint ridges on each, rather smooth, rugose posteriorly,
somewhat finely but strongly, rather sparsely punctate, more
coarsely so toward the apex and a little closer laterally ; proster-
num strongly, somewhat closely punctate, the process minutely
and sparsely so, very feebly constricted, rounded at tip and mar-
gined throughout, the bead occasionally obsolete apically ; hind
femora and abdomen minutely, sparsely punctulate, the latter very
feebly rugulose. Length 10. 0-12.0 mm.; width 4.6-5.75 mm.
($). Siskiyou Co., — Albert Koebele shastanica n. sp.

Body larger than in shastanica^ more broadly oblong and rather less
convex, dull in lustre, the elytra rather less pronouncedly deep
black ; head rather finely, closely and subevenly punctate, the sinus
small, very deeply subangulate, the lobes broadly rounded, the
anterior canthus rounded but not very broadly, about as prominent
as the posterior; antennae rather thick ; prothorax very transverse,
slightly less than twice as wide as long, the sides rounded and
converging from base to apex, rather more so anteriorly, sub-
parallel toward base, the apex unevenly sinuate, being more trans-
verse medially, two-thirds as wide as the base, the latter broadly,

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 1 29

evenly bisinuate, with the angles rather acute and posteriorly pro-
duced, the apical angles moderately rounded, right ; surface finelv,
not very sparsely punctate, becoming strongly and unusually closely
so laterally, the bead moderately fine; scutellum rather small, not
very transversely triangular ; elytra one-half longer than wide,
much less gradually rounded and narrowed behind than in shas-
tanica, obtusely ogival at tip ; surface nearly smooth, with notice-
ably small and unusually close punctures, but strongly rugose and
more coarsely punctate on the declivity, where each puncture has
a very minute silvery seta entirely enclosed within it ; prosternum
nearly as in shastanica but with the process more elongate, more
constricted, strongly margined except at the rounded apex, which
is unmargined ; hind femora and abdomen finely, sparsely and in-
conspicuously punctate. Length 12.3 mm.; width 1^.9 mm.
Siskiyou Co. (Sisson), — H. F. Wickham conferta n. sp.

23 — Prothorax large, a little wnder than the elytra 24

Prothorax not at all wider than the elytra 25

24 — Form oblong-elongate, very convex and strongly shining through-

out, very deep black, the elytral punctures with sette so minute as
only to be observed under rather high power ; head somewhat finely
but strongly, moderately sparsely punctured, the sinus rather wide
and shallow, the anterior canthus broadly rounded, even a little
more prominent than the posterior ; prothorax nearly four-fifths
wider than long, the sides subparallel and feebly arcuate, round-
ing and strongly converging anteriorly, the apex broadly sinuate,
three-fifths as wide as the base or a little more (d^), the angles
obtuse and rounded ; base subtransverse, with the angles but feebly
and gradually produced and rounded ; surface excessively minutely
and remotely punctulate, more visibly but still very finely and
remotely so laterally, the sides narrowly explanate and with a
rather coarse bead ; scutellum rather large, transversely and
obtusely triangular ; elytra two-thirds longer than wide, straight
and parallel at the sides, very gradually narrowed and somewhat
acutely produced at apex, nearly smooth, very coarsely and sparsely
punctate, more closely so but only slightly rugose on the declivity ;
prosternal process but feebly constricted, strongly margined at the
sides but. unmargined at apex ; abdomen extremely minutely,
remotely punctulate, the hind femora with sparse but rather
strong — though small — punctures. Length 11,7-13.5 mm.;
width 5.4-6.1 mm. San Francisco. \^=:- incEqualis ^?>y^.

eschscholtzi Mann.

Form narrower, deep black, less shining, the elytral punctures pos-
teriorly each with a very small but more visible seta ; head very
finely, rather sparsely punctate, the sinus broad but rather deep,
parabolic, the lobes less obtuse, the anterior canthus oblique and
much less rounded, obtusely subangulate, as prominent as the
posterior ; prothorax less transverse, nearly three-fourths wider
than long, the sides more evenly rounded, gradually more con-

130 CASEY

verging anteriorly, becoming parallel and nearly straight toward
base, the apex shallowly sinuate, nearly two-thirds as wide as the
base, the angles rather deflexed, moderately rounded ; base very
broadly and feebly bisinuate, the hind angles right, not more than
slightly blunt ; surface very minutely, not very sparsely punctulate,
the punctures becoming decidedly strong and closer but still
noticeably fine laterally, the margins not at all reflexed and with
a moderately fine bead ; scutellum less obtuse though transversely
triangular ; elytra but little more than one-half longer than wide,
the sides broadly, subevenly rounded and converging from near
basal third to the more obtusely ogival apex, scarcely visibly nar-
rower than the prothorax ; surface coarsely, less sparsely and less
strongly punctate, but more densely and more rugose on the
declivity ; prosternum sparsely and very minutely punctulate, the
process unusually short, broadly rounded, very feebly constricted,
margined throughout ; hind femora and abdomen minutely, sparsely
and inconspicuously punctate. Length 10.7 mm. ; width 4.9
mm. (cJ*). " Cal." pagana n. sp.

25 — Body oblong-oval, strongly convex, black and moderately shin-
ing throughout, the anterior parts slightly alutaceous ; head rather
coarsely and closely punctured anteriorly, finely and sparsely
toward base, the sinus moderately wide, deep, evenly rounded, the
anterior canthus obtusely oblique, much more prominent than the
posterior ; prothorax not more than one-half wider than long, the
sides converging and progressively more rounded from base to
apex, the latter shallowly emarginate, nearly two-thirds as wide
as the base, the angles much deflexed and broadly rounded ; base
very feebly bisinuate, the angles but slightly projecting ; surface
extremely minutely, sparsely punctulate, becoming somewhat
coarsely but sparsely so toward the sides, the margin not reflexed
and not very thickly beaded ; scutellum very transversely triangu-
lar ; elytra visibly less than one-half longer than wide, slightly
wider than the prothorax, the sides parallel and nearly straight,
arcuately converging at base, rapidly rounded and converging in
apical two-fifths, the apex very broadly ogival ; surface very
coarsely but not closely punctate, rugose, densely and strongly so
posteriorly ; prosternum minutely, sparsely punctate, the process
short, impressed along the middle, rounded at tip, barely at all con-
stricted, very strongly margined throughout ; hind femora and ab-
domen minutely, remotely punctulate. Length 10.5 mm. ; width
5.18 mm. (9). California inornata n. sp.

Body oblong-oval, very stout, stronglv convex, very deep black and
strongly shining throughout, the elytra with very minute and in-
conspicuous setie posteriorly ; head finely, sparsely punctate, the
sinus moderately wide, rather deep and parabolic, the lobes very
widely rounded, the anterior canthus obliquely subacute, fully as
prominent as the posterior ; prothorax three-fifths wider than long,
the sides broadly rounded and converging anteriorly, becoming
nearly straight and almost parallel in about basal half, the apex

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I3I

shallowly sinuate, two-thirds as wide as the base, with broadly
rounded angles ; base very broadly and feebly bisinuate ; surface
very sparsely, extremely minutely punctulate, becoming more dis-
tinctly but still finely, sparsely so laterally, the bead moderate ;
scutellum transverse, sharply triangular; elytra inflated, very dis-
tinctly wider than the prothorax, the sides parallel and broadly
arcuate, strongly rounding and converging behind the middle, a
little less than one-half longer than wide, the apex ogival ; surface
almost smooth, moderately rugose posteriorly, the punctures coarse
though much less coarse and sparser than in inornata ; prosternum
very minutely, sparsely punctate, the process well developed, mod-
erately constricted, very strongly margined throughout, not im-
pressed ; hind femora finely and sparsely but rather strongly punc-
tate, the abdomen extremely minutely, remotely and almost im-
perceptibly so. Length 11.8 mm.; width 6.0 mm. ($). Cali-
foinia. Cab. Levette COngesta n. sp.

Body oblong-elongate, more parallel and less convex than in the two pre-
ceding, rather shining, the elytra with small but visible setas pos-
teriorly ; head rather coarsely, sparsely and unevenly punctate
throughout, the sinus moderately wide, rather deeply parabolic,
the lobes rounded, the anterior canthus obtusely rounded, more
prominent than the posterior; prothorax barely one-half wider
than long, the sides converging and feebly arcuate from the base,
more strongly near the apex, which is evenly and moderately sinu-
ate and fully three-fifths as wide as the base, the angles deflexed,
obtuse and moderately rounded ; base broadly, very feebly and
evenly bisinuate; surface finely and sparsely though distinctly
punctate, becoming rather coarsely and more closely so laterally,
the bead moderate ; scutellum moderate, not very transverse, ob-
tusely triangular ; elytra fully one-half longer than wide, equal in
width to the prothorax, the sides parallel, rounding and converg-
ing behind the middle to the broadly ogival apex ; surface not
rugose, moderately coarsely, very sparsely punctate, the punctures
coarser and denser, with the surface rugose, on the declivity ; pro-
sternum with strong and only moderately sparse punctures, the
process less abbreviated than in inornata, margined throughout
but less strongly, not impressed ; hind femora and abdomen finely
but rather strongly, sparsely punctate. Length 13.3 mm.; width
5.83. California. Cab. Levette agrestis n. sp.

26 — Form oblong-suboval, strongly convex, shining throughout, the
elytra with small hairs only noticeable posteriorly ; head very
coarsely, more or less densely and unevenly punctate, the sinus
moderately wide, rather shallow, the lobes not much rounded, the
anterior canthus obtusely rounded, more prominent than the pos-
terior but not more so than the unusually convex eyes ; prothorax
nearly four-fifths wider than long, the sides converging and nearly
straight about to apical third, then strongly rounded and more
converging to the apex, which is shallowly sinuate and two-thirds
as wide as the base, the latter broadly, feebly bisinuate, the an-

132 CASEY

terior angles broadly rounded ; surface sparsely but rather coarsely,
unevenly punctate medially, becoming unusually coarsely so and
less sparsely toward the sides, the bead thick; scutellum moder-
ate, transversely triangular ; elytra one-half longer than wide to
rather more, rapidly narrowed and with strongly rounding sides in
apical two-fifths, the apex broadly ogival ; surface with more or
less feeble evidence of a few longitudinal costte, rugose behind,
nearly smooth anteriorly, coarsely and more or less closely but
unevenly punctate; prosternum strongly, closely punctate, the
process rather narrow, moderately constricted, longitudinally con-
vex, rounded at tip, not margined except at the sides toward base ;
hind femora and abdomen with sparse but unusually strong, even
punctures throughout. Length 10. 2-12. 2 mm. ; width 4.75-5.9
mm. Farallon Islands farallonica Csy.

Some of the species allied to nemoralis might almost as natu-
rally fall near subptcbescens in the preceding group, but the
form is a little more convex and the sculpture stronger, giving
them on the whole more nearly the general habitus of the
present group. There are evidences of some natural geographic
subdivisions of this group, oregona 3.r\d/'araIloiiica, for example,
being widely different from each other and from the usual type
and the series of northern species from callida to conferta also
hold together very consistently, departing widely in general
sculpture from the others. It is quite obvious that the species
formerly described by me under the name incBqualis, a remark-
ably isolated form by reason of the sharply acute elytral apex
and in having the prothorax wider than the elytra, with nar-
rowly subexplanate sides, is, by reason of the virtual absence
of elytral setse and because of the very feeble punctuation of
the anterior parts, the species held by LeConte and Horn to
represent the true eschscholtzi ; affinis is therefore a valid spe-
cies, being one of a considerable number of more or less closely
allied forms as noted above.

Group VIII. — Type ovah's.

The rather numerous species falling under this head replace
the allies of eschscholtzi in the northern Pacific coast and Rocky
mountain regions, and are distributed thence southwardly to the
eastern foothills in central Colorado, and, to the westward, through
Utah to western Nevada, this extended range being nearly
coincident with that of the sctosa group. The body is always

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I33

strongly convex, comparative!}' coarsely sculptured and is more
oval than in the eschscholtzi group, but otherwise resembles the
prevailing form in the latter, except that it is more completely
glabrous, no species being known to me having more than
very minute setae virtually wholly enclosed within the punc-
tures ; they may be known by the following characters : —

Elytral punctures more or less coarse and very conspicuous, the in-
teguments deep black throughout and strongly shining; habitat
near the coast 3

Elytral punctures smaller and less conspicuous though invariably dis-
tinct, the integuments always deep black but frequently alutaceous ;
habitat further removed from the coast 6

2 — Sides of the prothorax strongly rounded and converging anteriorly,
becoming straight and parallel in rather more than basal half.
Form more oblong than usual, parallel, strongly shining ; head
strongly, rather closely punctate anteriorly, more sparsely toward
base, the sinus rather broad, moderately deep, subangular, the
lobes broadly rounded, not impressed, the anterior canthus broadly
rounded, more prominent than the posterior ; prothorax short,
fully four-fifths wider than long, the apex moderately sinuate,
barely three-fifths as wide as the base, the angles deflexed and
slightly obtuse but only narrowly rounded ; base transverse, feebly
oblique at the sides ; surface finely, rather closely and distinctly
punctate, becoming somewhat coarsely so, though not much more
closely, toward the sides, the bead rather fine ; scutellum small ;
elytra one-half longer than wide, barely as wide as the prothorax,
the sides gradually converging and broadly rounded behind from
about basal third, the apex rather acutely ogival ; surface rugose,
especially behind, coarsely and closely punctate ; prosternum
strongly and closely punctured throughout, the process well
developed, very feebly constricted, circularly rounded at tip, very
strongly margined throughout ; hind femora rather coarsely,
moderately closely and conspicuously punctate, the abdomen
strongly and unusually closely so. Length ii.o mm.; width
5.4^ mm. ( ?). Probably near Puget's Sound... breviuscula n. sp.

Sides of the prothorax strongly, subevenly rounded throughout, strongly
converging anteriorly, becoming parallel and less arcuate toward
base. Form oblong-oval, much more longitudinally convex ;
head not very coarsely but strongly punctured, densely toward
the suture, more sparselv elsewhere, the sinus broad and vmusually
shallow, circularly rounded, the lobes broadly rounded, impressed
near the margin, the anterior canthus evenly and broadly rounded,
more prominent than the posterior ; prothorax more than three-
fourths wider than long, the apex rather deeply sinuate, three-
fifths as wide as the base, the angles moderately deflexed, slightly
obtuse and well rounded ; base broadly bisinuate, the angles rather

134 CASEY

strongly produced ; surface strongly convex, sparsely but unusually
coarsely punctate, gradually closely though unequally and still
more coarsely so laterally, the bead moderately thick ; scutellum
moderate ; elytra scarcely one-half longer than wide, equal in
width to the prothorax, the sides straight and parallel, rather
rapidly converging and rounded in apical two-fifths, oblique and
straighter toward the rather acute tip ; surface very coarsely,
deeply and rather closely punctate and very strongly rugose
throughout ; prosternum rather strongly, closelv punctate, the
process finely so, distinctly constricted, rather acutely rounded at
apex and unmargined ; hind femora rather coarsely and conspicu-
ously, though somewhat sparsely, punctured, the abdomen but little
less coarsely and sparsely, strongly so. Length 9.4 mm. ; width
5.0 mm. (c?). Washington State, — exact locality unrecorded.

sculptipennis n. sp.

Sides of the prothorax converging from the base 3

3 — Prosternal process margined throughout 4

Prosternal process not margined, except feebly toward base 5

4 — Body very evenly subelliptic in form, convex, shining; head

strongly, rather loosely punctate, the sinus moderate in width,
vmusually deep, angulate, the lobes broadly rounded, more or less
strongly impressed within the margin, the anterior canthus
obliquely and somewhat narrowly rounded, a little more promi-
nent than the posterior ; prothorax fully three-fourths wider than
long, the sides almost evenlv and broadly rounded from base to
apex, the latter rather shallowly sinuate, with the bottom of the
sinus transverse, barely three-fifths as wide as the base, the angles
deflexed and somewhat obtuse but only very narrowly rounded ;
base broadly bisinuate, the angles somewhat produced and acute ;
surface rather finely and sparsely though very distinctly punctate,
gradually coarsely and rather closely so laterally, the bead fine ;
scutellum rather small, not very transverse, sharply triangular ;
elytra as wide as the prothorax, fully one-half longer than wide,
the sides parallel and just visibly arcuate, rounding and converg-
ing in apical two-fifths, the sides becoming nearly straight toward
the rather acutely angular tip ; surface rugose to nearly smooth,
moderately coarsely, rather sparsely punctate ; prosternum pol-
ished, strongly, rather closely punctate, the process distinctly con-
stricted, the marginal bead strong throughout ; hind femora with
moderate and rather strong but sparse punctures, those of the abdo-
men similar though closer. Length 11. 0-11.2 mm.: width 5.7
mm. Washington State (Leavenworth), — H. F. Wickham.

regularis n. sp.

Body less evenly elliptic, oblong-oval, sometimes feebly inflated pos-
teriorly, shining ; head rather coarsely and closely punctate, the
sinus nearly as in regularis^ the lobes more narrowly rounded,
more broadly and less linearly impressed, the anterior canthus
nearly similar but more rounded ; prothorax barely three-fourths

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I35

wider than long, the sides feebly converging and nearly straight
from the base to beyond the middle, there becoming rounded and
more converging, the apex more evenly sinuate, two-thirds as
wide as the base, the angles, base and basal angles nearly as in
regiilaris ; surface rather coarsely, not very sparsely punctate,
becoming unusually coarsely and densely so laterally, the bead
moderately coarse, arcuate about the basal angles ; scutellum more
transverse, with rather rounded sides and more obtuse apex ;
elytra fully one-half longer than wide, near the middle a little
wider than the prothorax, the sides parallel and sensibly arcuate,
rounding in posterior third, becoming straight to the acutely angu-
late apex; surface rugose, polished, very coarsely but rather
sparsely punctured ; prosternum moderately shining, strongly,
rather closely punctate, the process a little wider, constricted,
strongly rounded at tip, feebly margined and somewhat uncer-
tainly so near the apex ; abdomen polished though feebly rugu-
lose, with fine and rather sparse but very strong punctures.
Length 10.7 mm.; width 5.55 mm. Probably from the neigh-
borhood of Puget's Sound punctata n. sp.

5 — Form oblong-oval, rather elongate, strongly convex, alutaceous
anteriorly, the elytra polished ; head rather finely but strongly,
sparsely punctate, the sinus rather narrow, deep, angular, the
lobes rounded, broadly impressed, the anterior canthus nearly as
in punctata ; prothorax similar but much less coarsely punctured
and less closely so toward the sides, the bead finer and more feebly
hooked at base, not entirely surrounding the basal angles ; scutel-
lum small, similar in form ; elytra similar in form but not inflated
and with the parallel sides nearly straight, the surface similarly
rugose though with the rather sparse punctures not quite so coarse ;
prosternum subopaque, moderately punctured and with ver}' short
fulvescent seta^, the process well developed, very feebly constricted,
angularly rounded at apex, somewhat feebly margined along the
sides, the apex unmargined ; hind femora and abdomen sparsely
and finely but somewhat strongly punctured. Length i r .3 mm. ;
width 5.75 mm. Vancouver Island parilis n. sp.

Form much more abbreviated and more evenly elliptic, convex, strongly
shining throughout ; head with moderately small, perforate and
somewhat close-set punctures, the sinus a little wider and less
deep, angulate, the lobes rounded, strongly impressed, the anterior
canthus still more broadly rounded, much more prominent than
the posterior ; prothorax much shorter, nearly four-fifths wider
than long, the sides more strongly converging and nearly straight
to before the middle, then rounding, strongly converging at apex,
the latter barely three-fifths as wide as the base, more deeply
and subevenly sinuate, the angles more deflexed and much more
broadly rounded, the base and the produced angles similar ; sur-
face with larger, strong and rather sparse punctures and a broader
impunctate line, the punctures relatively less enlarged, sparser,.

136 CASEY

equally coarse and still more unevenly distributed laterally, the
bead moderately fine, almost straight, not curved at base ; scutel-
lum smaller than in any other species observed ; elytra much less
than one-half longer than wide, more oval, the sides rounding and
converging behind the middle to the more obtusely angulate apex,
the surface similarly rugose but still less coarsely, sparsely punc-
tate; prosternum opaque, finely, not conspicuously punctate, the
process abnormally short, broadly rounded, moderately con-
stricted, very sparsely punctate, unmargined except very finely
at the sides toward base ; hind femora rather closely, some-
what strongly punctured, the abdomen unusually closely and
strongly so. Length 9.8 mm; width 5.3 mm. Vancouver Island,
— H. F. Wickham vancouveri n. sp.

6 — Prosternal process margined throughout 7

Prosternal process not margined at apex but with an unusually broad,

though flat, margin at the sides 13

7 — Species of the Rocky mountain system 8

Species of the eastern foothills of the Sierras 13

8 — Anterior canthus about as prominent as the posterior .9

Anterior canthus much less prominent than the posterior 11

9 — Body oblong in form, convex, polished throughout, deep black;

head rather finely, subevenly and loosely punctate, the sinus rather
deeply parabolic, the lobes rounded, unimpressed, the anterior
canthus strongly rounded, a little more prominent than the pos-
terior ; prothorax four-fifths wider than long, the sides broadly
rounded and moderately converging anteriorly, becoming nearly
parallel toward base, the apex evenly and moderately sinuate, two-
thirds as w'ide as the base, the angles deflexed, right and but little
rounded, the base broadly, feebly bisinuate, with the angles rather
abruptly and moderately produced ; surface very finely, sparsely
punctate, becoming moderately coarsely, not very closely so later-
ally, except anteriorly where they are still rather fine and sparse,
the bead fine ; scutellum not very transverse, moderate, somewhat
obtusely angulate ; elytra one-half longer than wide, the sides
parallel and nearly straight to apical two-fifths, then strongly
rounded, becoming straight to the obtusely angulate and promi-
nent tip ; surface smooth, finely, densely rugose behind, the punc-
tures moderately fine, rather sparse ; prosternum densely punctate
and rugulose, the process polished and sparsely punctulate,
rounded at tip, strongly constricted ; hind femora and abdomen
strongly but not very coarsely punctate, the latter the more closely.
Length 9.2 mm.; wndth 4.8 mm. Utah (Nephi), — H. F.
Wickham uteana n. sp.

Body more evenly elliptic in outline 10

10 — Form rather narrowly elliptic, polished, only moderately convex,

deep black ; head very finely, sparsely punctate, the sinus only
moderately deep, broadly parabolic, the lobes broadly rounded,

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I37

scarcely impressed, the anterior canthus broadly rounded, copromi-
nent with the posterior ; prothorax two-thirds wider than long,
the sides strongly converging from base to apex, subequally and
rather strongly arcuate, more so at apex, the latter evenly sinuate
and but little more than half as wide as the base, the angles de-
flexed, right and rather distinctly rounded ; base broadly bisinuate,
the angles but little produced; surface minutely, sparsely punctu-
late, becoming strongly but not very coarsely and still sparsely
punctate laterally, the bead fine, perfectly straight and unmodified
at the basal angles ; scutellum moderate, obtuse ; elytra two-fifths
longer than wide, the sides rounding and converging posteriorly
to the regularly ogival tip, becoming parallel only toward base,
the surface smooth, rather finely rugose posteriorly, strongly and
sparsely but not very coarsely punctured, more coarsely and
densely behind ; prosternum nearly as in uteana but with the
process narrower, less constricted and more acutely rounded at
apex ; hind femora with rather small but strong and remote
punctures, the abdomen minutely, sparsely and inconspicuously
punctulate. Length 9.0 mm.; width 4.5 mm. (cf). Colorado
(Salida), — H. F. Wickham inepta n. sp.

Form more broadly elliptic and more convex, much larger, not quite
so shining, black ; head finely, sparsely punctate, the sinus rather
deep, subangular, the lobes not very broadly rounded, unimpressed,
the anterior canthus similar ; prothorax three-fourths wider than
long, the sides as in itiepta^ the apex also nearly similar but three-
fifths as wide as the base; base rather strongly bisinuate, the
angles moderately and gradually produced ; surface minutely,
sparsely punctulate, gradually more strongly but still rather finely
and only moderately closely toward the sides, the bead rather fine ;
scutellum rather small ; elytra formed nearly as in inepta but with
the sides becoming parallel in about basal half ; surface rugose
throughout to nearly smooth, always more strongly and deeply
rugose on the declivity, the punctures rather small but strong,
sparse, larger and close posteriorly; prosternum as in t?iepta,
somewhat shining, closely and somewhat asperately punctate, the
process similarly rather narrow and elongate, smooth and mod-
erately constricted but more broadly and evenly rounded at tip, the
abdomen and hind femoi'a finely, sparsely and rather inconspicu-
ously punctate. I^ength 10.2-11.5 mm.; width 5.1-5.6 mm.
Probably from northern Idaho oblita n. sp.

Form more elongate, convex, alutaceous, the elytra more shining, deep
black ; head finely, sparsely punctured, the sinus rather narrow,
moderately deep, parabolic, the lobes broadly rounded, obsoletely
impressed, the anterior canthus strongly rounded, a little more
prominent than the posterior ; prothorax three-fifths wider than
long, the sides sensibly converging and more or less nearly straight
to beyond the middle, then more strongly rounded and converging
to the apex, the latter moderately sinuate and fully two-thirds as
wide as the base, the apical and basal angles nearly as in uteana^
Proc. Wash. Acad. Sci., April, 1908.

138 CASEY

the former only narrowly rounded ; surface finely but distinctly,
rather sparsely punctate, becoming more closely and strongly so
toward the sides though still not coarsely so, the bead rather fine,
straight to the base ; elytra one-half longer than wide, with the
sides nearly straight and parallel, rounding gradually behind to the
acutely ogival and prominent tip ; surface rather finely, sparsely
punctured, nearly smooth, moderately rugose posteriorly ; pro-
sternum, legs and abdomen nearly as in oblita. Length 10.3
mm. ; width 5.2 mm. (?). Utah (southwestern), — C. J. Weidt.

arida n. sp.

Form stouter and differing greatly in being inflated posteriorly, black,
alutaceous, the elytra rather shining ; head somewhat coarsely,
loosely punctate, the sinus broad, not very deep, parabolic, the
lobes broadly rounded, unimpressed, the anterior canthus rounded,
coprominent with the posterior ; prothorax three-fourths wider
than long, in outline as in the preceding but more strongly nar-
rowed at apex, the latter rather more deeply sinuate and barely
three-fifths as wide as the base, which is more strongly bisinuate
than in arida^ with the sides more gradually and feebly oblique ;
apical angles but slightly rounded ; surface punctured nearly as in
arida but more strongly and more coarsely and much more closely
so toward the sides ; scutellum much smaller ; elytra barely a third
longer than wide, behind the middle evidently wider than the pro-
thorax, the sides broadly rounded, rapidly rounded to the very
obtusely subangulate apex; surface nearly smooth, rather finely
and closely punctate, densely rugose and more coarsely punctate
on the declivity ; prosternum differing greatl}', being somewhat
coarsely and very closely punctate even on the process, which
is shorter and broader, rounded at tip and moderately constricted ;
abdomen and hind femora rather finely but strongly, sparsely
punctate. Length i r.i mm. ; width 5.6mm. (? ). Utah (south-
western), — C. J. Weidt weidti n. sp.

II — Form oblong-oval, moderately convex, deep black, rather shining ;
head rather strongly, unevenly punctate, the sinus rather broad, not
very deep, parabolic, the lobes unusually broadly rounded, unim-
pressed, the anterior canthus somewhat oblique and narrowly
rounded; prothorax four-fifths wider than long to a little less, the
sides converging from the base and feebly arcuate, becoming
strongly rounded and converging anteriorly, the apex moderately
sinuate, three-fifths as wide as the base, which is ver}' feebly bisin-
uate, the angles but slightly produced, the anterior angles de-
flexed and moderately rounded ; surface rather strongly and unu-
sually closely punctate, becoming decidedly coarsely but not
materially more closely so laterally, the bead moderate, straight at
base ; scutellum well developed ; elytra evidently less than one-
half longer than wide, as wide as the prothorax, the sides parallel
and nearly straight, rather rapidly rounding in apical two-fifths to
the moderately obtusely ogival tip ; surface slightly uneven,
strongly and somewhat sparsely punctate, rugose posteriorly ; pro-

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I39

sternum shining medially, with strong and moderately separated
punctures, the process only moderate in width, rounded at tip,
moderately constricted ; abdomen unusually strongly and closely
punctate, the hind femora more sparsely. Length 10. 2-1 1 .0 mm. ;
width 5.2-5.6 mm. Montana (Bitter Root Mts.) ...acerba n. sp.

Form more oval and convex, polished, deep black, the punctures pos-
teriorly with very small silvery setas ; head rather finely but
strongly, not very closely punctate, the sinus moderately narrow,
deep and subangulate, the lobes much less broadly rounded, even,
the anterior canthus broadly rounded ; prothorax from less to more
than four-fifths wider than long, in outline nearly as in acerba
throughout, the base similar ; surface finely though rather strongly,
not very sparsely punctate, gradually coarsely and much more
closely so laterally, the bead moderately fine, straight to the base;
scutellum rather small, transversely and obtusely triangular ; elytra
nearly as in acerba but rather more gradually and acutely angu-
late at tip, the sides parallel and very feebly arcuate; surface
slightly uneven, rugose behind, the punctures strong though not
very coarse, closer and coarse posteriorly ; prosternum shining
though strongly and densely punctured, the process less closely
so, moderately broad, rounded behind and distinctly constricted ;
hind femora and abdomen finely and stronglv punctured, the latter
more closely. Length 9.0-10.5 mm. ; width 4.4-5.5 mm. Col-
orado (Boulder Co.), — Mrs. F. Y. Parker anita n. sp.

12 — Body stout, convex, oblong-oval, dull and alutaceous, deep black,

the elytral punctures with minute silvery setas ; head finely but
strongly, moderately closely punctate, the sinus moderately nar-
row, deep and bluntly angulate, the lobes rounded and feebly im-
pressed, the anterior canthus narrowly rounded, much more
prominent than the posterior; prothorax three-fourths v\'ider than
long, the sides rounded and converging anteriorly, less arcuate and
more or less nearly parallel toward base, the apex deeply sinuate,
two-thirds as wide as the base, the angles not much deflexed and
well rounded ; base broadly, feebly sinuate toward the sides ; sur-
face finely, rather sparsely punctate, with a more or less evident
impunctate line, becoming strongly and closely but not very
coarsely so laterally, the bead fine, straight to the base ; scutellum
small; elytra parallel, one-half longer than wide to a little less,
angulate in apical two-fifths ; surface nearly smooth, strongly but
not very coarsely, rather closely punctate, strongly rugose and
coarsely, closely punctured posteriorly ; prosternum strongly,
densely and submuricately punctate, the process nearly smooth
and very sparsely, simply punctate, broadly rounded, strongly
constricted ; hind femora and abdomen strongly and moderately,
equally sparsely punctate. Length 8.8-10.7 it^""*' i width 4.7-5.1
mm. Nevada (Reno). A moderate series taken by the writer
along the Truckee River alutacea Csy.

13 — Body narrower, oblong-oval, convex, deep black and alutaceous,

140 CASEY

the elytra shining, with very minute sctaj within the punctures ;
head finely, deeply and closely punctate, the sinus moderately
narrow, deep, angulate, the lobes not broadly rounded, unim-
pressed, the anterior canthus obliquely rounded, sensibly more
prominent than the posterior; prothorax three-fourths wider than
long, the sides strongly and almost evenly rounded, strongly con-
verging apically, becoming parallel basally, the apex rather shal-
lowly sinuate, fully three-fifths as wide as the base, the angles de-
flexed and obtuse but rather narrowly rounded, the base feebly sin-
uate laterally ; surface somewhat finely and sparsely though very
strongly punctate, becoming coarsely but loosely so laterally, the
bead rather fine, slightly curving at base ; scutellum small ; elytra
barely two-fifths longer than wide, toward base narrower than the
prothorax, the sides parallel and rounded, becoming more con-
verging but rounded posteriorly to the ogival apex ; surface slightly
uneven, rugose posteriorly, strongly, rather sparsely punctate ;
prosternum medially finely but strongly, not densely punctate,
opaque anteriorly, the process rather narrow, moderately con-
stricted, the broad flat lateral margin differing from anything else
observed in the genus ; hind femora and abdomen finely, sparsely
punctate. Length 9.35 mm. ; width 4. 75 mm. Colorado (Boulder
Co.) corvina n. sp.

Form oval, rather abbreviated, posteriorly subacute, black, shining;
prothorax shorter than wide, less finely pvmctate than in affinis^
the anterior angles strongly deflexed, the sides very finely mar-
gined, the posterior angles produced, acute ; elytra densely,
rugosely punctate. Length 10.25 mm.; width 5.5 mm. Oregon.

ovalis Lee.

The species described by LeConte under the name ovalis^ was
probably collected at some point in the Willamette Valley, and
will prove to be different from any at present represented before
me ; the original description is given above.

Subgenus Crypticomorpha nov.

Form elongate-oval, rather depressed, piceous, the under surface, legs
and antennae more or less pale and rufescent ; integuments leather
shining, clothed thinly with fine decumbent pubescence ; head
moderate, rather finely, not densely punctate, the sinus moderate
in width, broadly rounded, the lobes obtuse but not very broadly
rounded, the oblique sides arcuate, less prominent than the unusu-
ally convex and subentire eyes; antcnuie long, very slender; pro-
thorax nearly three-fifths wider than long, somewhat wider behind
the middle than at base, the sides subevenly and rather strongly
arcuate, converging anteriorly, becoming parallel basally, the apex
moderately and evenly sinuate, three-fifths as wide as the base,

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I4I

the angles right and narrowly rounded, the base broadly bisinuate,
with the angles but slightly produced ; surface minutely, sparsely
punctate, becoming but slightly more strongly or less sparsely so
laterally, the bead extremely fine ; scutellum rather small, but
slightly transverse, sharply triangular; elytra equal in width to
the prothorax, rather more than one-half longer than wide, the
sides parallel and straight, gradually strongly rounded in about
apical two-fifths to the somewhat sharply ogival tip ; surface
smooth throughout, alutaceous posteriorly, very finely, evenly and
not densely punctate ; prosternum extremely minutely punctulate,
the process unmargined, moderately constricted, long, rounded at
tip ; femora and abdomen minutely, sparsely, inconspicuously punc-
tate : tarsi long and very slender. Length 6.7 mm. ; width 2.9
mm. (cT). California, — the exact locality unrecorded, — J. J.
Rivers tenuis n. sp.

Subgenus Brachyontis nov.

Form exti-emely short and stout, very convex, feebly shining, black,
the legs rufous, the antennae piceous ; upper surface clothed rather
thinly with coai'se decumbent fulvous hairs ; head moderate, rufes-
cent anteriorly, rather finely, loosely punctate, the sinus moderate,
broadly rounded, the lobes obtuse, not very broadly rounded,' the
anterior canthvis rounded, less prominent than the posterior ; an-
tennae rather short, thick ; prothorax fully four-fifths wider than
long, the sides almost evenl}- rounded, more converging anteriorly,
becoming parallel near the base, the apex strongly sinuate, two-
thirds as wide as the base, the angles much deflexed, obtuse but
not broadly rounded, the base transverse, straight, feebly curved
posteriorly toward the sides, the hind angles rounded ; surface
finely, closely punctate, more strongly and still more closely so
toward the sides, the bead fine, curving inwardly at base ; scutellum
moderate, polished, obtuse ; elytra not longer than wnde, very
strongly declivous behind, at the middle a little wider than the pi"o-
thorax, the sides parallel and broadly arcuate, the apex very
broadly obtuse; surface smooth even posteriorly, minutely, rather
sparsely punctate ; prosternum finely, closely punctate, the process
short, moderately constricted, rounded and convex apically, not
margined except feebly at the sides basally ; abdomen almost im-
punctate. Length 6.4-6.:; mm. ; width 3.7-4.22 mm. California
(Tahichipi Pass) globulina Csy

It should be borne in mind that the groups into which the
genus Coniontis is divided above, are, in considerable meastire,
arbitrary and intended more to facilitate identification than any-
thing else ; the abdominalis, opaca and stihfubescens groups are
perhaps the least natural aggregates, while the viatica^ setosa,
cschscholtzi and ovalis groups are more in the nature of quasi-
subgeneric geographic sections of the genus, more sharply cir-
cumscribed in general habitus and are more homogeneous.

142 CASEY

Coniontellus Csy.

The small convex species constituting this genus have a
facies very similar to that of Coniontis^ but, besides the com-
pletely divided eyes, they differ in their shorter legs and smaller
antennas. They are divisible into two clearly separated groups,
the first, having stiff hispid vestiture, being confined as far as
known to the eastern slopes of the central Sierras, all that I
have seen having been taken in the valley of the Truckee
River ; the other group, smaller and glabrous or virtually so,
are very widely distributed, but, unlike Coniontis^ are entire
strangers to the fauna of the coast regions. The eight species
in my collection may be described as follows : —

Elytra bristling, more especially toward the sides and tip, with stiff
erect fulvous setae 2

Elytra glabrous, or at most with very minute, decumbent and incon-
spicuous hairs 5

2 — Integuments castaneous in color 3

Integuments black 4

3 — Body oblong, convex, rather shining, coarsely sculptured ; head

strongly but not coarsely, loosely punctate, more finely and densely
so before the suture, the sinus deep, angulate, the lobes broadly
rounded ; prothorax more than four-fifths wider than long, the sides
broadly rounded and converging before the middle, becoming
parallel or nearly so and straighter thence to the base, the apex
strongly sinuate, fully two-thirds as wide as the base, the latter
transversely truncate, gradually curved posteriorly at the sides, the
apical angles deflexed, right and narrowly rounded, the basal but
slightly produced and bhmt ; surface finely, sparsely punctured,
becoming coarsely, closely so and with the punctures elongate
laterally, the bead fine ; scutellum moderate, not much wider than
long, triangular; elytra less than one-half longer than wide, not
inflated apparently in either sex, the sides parallel and straight,
abruptly rounding in apical third to the obtuse tip ; surface almost
vertically declivous behind, slightly uneven, coarsely, very deeply,
evenly and closely punctate throughout ; prosternum and process
strongly and closely punctate, the latter margined at the sides but
not at the rounded apex ; abdomen rather finely, strongly, some-
what sparsely punctate. Length '].2-'].'] mm. ; width 3.6-3.9
mm. Nevada (near Reno), — H. F. Wickham hystrix n. sp.

Body oblong, more elongate, larger in size, shining, less strongly
sculptured ; head rather finely and closely, strongly and subevenly
punctured throughout, the sinus nearly similar, the lobes still more
broadly rounded, the sides more broadly rounded before the eyes;
prothorax twice as wide as the median length or nearly so, formed

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I43

otherwise nearly as in hystrix^ but with the coarse punctures near
the sides not elongate and with the narrow interspaces less smooth,
and also differing in having the sloping flanks somewhat sub-
explanate basally and the marginal bead coarser ; scutellum similar
though a little larger ; elytra much more elongate, fully one-half
longer than wide, nearly three times as long as the median line of
the prothorax, less declivous posteriorly, rounded in apical third,
the apex feebly produced, the sides parallel and straight (cf ) or
feebly arcuate and somewhat inflated posteriorly ( $ ) ; surface
very smooth, the punctures smaller, only moderately coarse but
perforate and much more widely separated ; prosternuin similarly
strongly, closely punctate, the process larger, more constricted,
the lateral beading much finer ; abdomen with rather small but
strong, moderately sparse punctures. Length 7.8-8.2 mm. ;
wadth 3. 8-4. 6 mm. Nevada (near Reno), — H. F. Wickham.

longipennis n. sp.

4 — Form stout, convex, deep black, somewhat alutaceous in lustre;
head punctured nearly as in hystrix^ the sinus rather small,
moderately deep, broadly angular, the lobes broadly rounded ;
sides obtusely prominent before the eyes ; prothorax four-fifths
wider than long, the sides very moderately rounded and not very
strongly converging anteriorly, becoming parallel and straighter
posteriorly, the strongly sinuate apex three-fourths as wide as the
base, the angles and base as in the preceding species ; surface not
coarsely but strongly, sparsely punctured, the punctures becoming
coarse, dense and slightly elongated laterally, the bead fine, the
hind angles more narrowly produced and more acute than in the
preceding ; scutellum very small ; elytra feebly inflated posteriorly
(c?) or strongly so (?), the sides broadly arcuate at or behind
the middle, distinctly wider than the prothorax, the apex obtusely
ogival and somewhat prolonged ; surface coarsely and deeply but
rather sparsely punctate, much more finely so on the opaque pos-
terior declivity ; prosternum coarsely, rather closely punctate, the
process longitudinally and strongly convex ; abdomen sparsely but
strongly punctate, polished. Length 6.6-7.0 mm. ; width 3.25—
3.7^ mm. Nevada (Reno). A small series taken by the writer
on the banks of the Truckee, in the suburbs of the town.

inflatus Csy.

Form still stouter, less convex, oblong, black, moderately shining, larger
in size; head finely but strongly, subevenly and closely punctate,
the sinus moderate, deep, subangular, the lobes broadly rounded ;
prothorax twice as wide as its median length, inflated, much wider
just behind the middle than at base, the sides strongly, subevenly
rovmded, more converging anteriorly, the apex strongly sinuate,
fully two-thirds as wide as the base, the angles right, rather nar-
rowly rounded, the basal rather narrowly but abruptly and moder-
ately produced and acute ; surface finely and sparsely punctate,
becoming coarsely and densely so, with the punctures just visibly
elongate, laterally, the bead rather fine ; scutellum very small, as

144 CASEY

in inJJatus^ almost equilateio-triangular ; elytra evidently less than
one-half longer than wide, not at all wider than the j^rothorax,
the sides parallel and but very feebly arcuate, the apex very obtusely
rounded, the tip angular; surface with moderate, deep and per-
forate, rather sparse punctures, which are not notably smaller on
the posterior declivity, the latter not differing in lustre, shining ;
prosternum deep black, strongly punctured, the process rather
short and somewhat strongly constricted, rounded, not margined
even at the sides ; abdomen rather finely and sparsely though
strongly punctate. Length 8.0 mm. ; width 4.2 mm. Nevada
(near Reno), — H. F. Wickham ampliatus n. sp.

5 — Elytra with very small hairs observable posteriorly. Body oblong,

stout, convex, deep black, the legs and antennae dark rufous, the
surface only feebly shining ; head finely, closely punctate, espe-
cially anteriorly, the sinus moderately deep, broadly angular, the
lobes broadly rounded ; prothorax twice as wide as its median
length, the sides evenly rounded and converging anteriorly, becom-
ing parallel in about basal half, the apex deeply and evenly sinuate,
three-fourths as wide as the base, the angles deflexed, right and
not rounded ; base transversely rectilinear, curving rather abruptly
to the rear near the sides, the angles rather prominent posteriorly
and not blunt ; surface strongly and closely punctate, becoming
more coai"sely and still more closely so laterally, the bead very fine ;
scutellum moderate, sharply equilatero-triangular ; elytra less than
a third longer than w^ide, equal in width to the prothorax, the sides
parallel and straight, rounding to the very acutely ogival tip in
apical two-fifths ; surface but slightly uneven, rather coarsely,
deeply, moderately closely and conspicuously punctate, the punc-
tures very close laterally, becoming smaller only very near the
acute tip ; prosternum strongly but not densely punctate, the pro-
cess unmargined ; abdomen not coarsely but strongly, somewhat ,
closely pvmctured. Length 6.6 mm.; width 3.1 mm. Colorado
(central). [:= Coniontis obesa Lee] obesus Lee.

Elytra completely glabrous, the setce so minute as to be invisible under
moderate power 6

6 — Form oblong, rather more elongate than in obesus., similar in

coloration and slightly more shining; head similarlv punctured
and almost equally short and transverse, the sinus smaller and still
feebler, the lobes very broadly rounding and scared}- differentiated
from the rounded sides, similarly with a very faint emargination
at the ends of the oblique sutures ; prothorax similar but less trans-
verse, not more than four-tifths wider than long, similarly punc-
tured and even more closely toward the sides ; scutellum similar
though notably smaller ; elytra similar though a little more elon-
gate, the punctures sensibly less close-set and decidedly less coarse ;
under surface nearly similar throughout. Length 5. 8-6. 75 mm. ;
width 2.8-3.2 mm. Montana (Helena) subglaber Csy.

Form similar to that of subglaber^ deep black, more or less alutaceous,

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I45

the legs and antcnnie piceo-rufous ; head nearly similar, the sinus
broader, moderately deep, angular, the lobes broadly rounded ;
prothorax almost exactly as in subglaber^ the deflexed anterior
angles blunt and by no means so sharply defined as in obesus^ the
punctures very fine and rather sparse but almost similarly strong
and close-set laterally, the hind angles a little less prolonged and
blunter ; scutellum similar, the elytra still more elongate, parallel
in both sexes, nearly one-half longer than wide, shorter in the
male, the punctures fine and sparse though distinct throughout ;
prosternum strong!}-, rather closely punctate, the process some-
times feebly margined at the sides but never at tip ; abdomen
finely, sparsely punctate, a little more coarsely but still sparsely
so toward the sides (?) or closely and more coarsely so (cJ*).
Length 6.2-7.0 mm. ; ^vidth 3. 15-3.35 mm. California (Bodie,
— elevation S475 feet) argutus n. sp_

Form nearly similar but more convex and with the elytra inflated,
probably more or less in both sexes, polished, deep black, the
legs and antennas rufo-piceous ; head nearly as in the preceding
species, strongly and closely punctured; prothorax as in argtitus^
the deflexed anterior angles right and blunt ; surface rather finely
but strongh', not very closel}- punctate, strongly and closely so
laterally; scutellum a little larger, similarly equilatero-triangular ;
elytra at the middle much wider than the prothorax, not a third
longer than wide, the sides parallel, evenly and rather strongly
arcuate, very rapidly rounding behind to the ogival but less prom-
inent tip ; surface smooth, very finely, sparsely punctured ; under
surface nearly as in the preceding species, the abdomen strongly,
moderately sparsely punctured, more coarsely so laterally. Length
6.5 mm.; width 3.45 mm. Colorado (Buena Vista, — elevation
about 8000 feet), — H. F. Wickham micans n. sp.

Some time ago I suppressed subglaber, as a synonym of obesus,
but more careful study shows that this course was premature ;
they are not by any means the same species. Mr. Warren
Knaus kindly sent me a specimen of the species named argti-
tus above, and I had also previously secured a series of four
specimens from Mr. ^Vickham, giving a sufficiently complete
succession of individuals for study ; there is very little variation,
but the male is notably less elongate than the female, some-
times having a more shining surface and stronger abdominal
punctures. It is interesting to note that the inflation of the
elytra is not necessarily a sexual character, though, when it
occurs in both sexes, it is more pronounced in the female, and
also to observe that some species in both groups of the genus,
otherwise closely allied, may have the elytra inflated or without
trace of inflation even in the female.

146 CASEY

Conisattus Csy.

While the general organization of this genus is essentially
similar to that of Coniontis, there are some characters that
recall the Eusatti, such as the reflexed margin of the head,
narrowly explanate sides of the pronotum and feebly prominent,
though obtuse, external apical angle of the anterior tibias. It
differs from the Eusalti, and is more closely affiliated with
ConiontiSy in the unproduced basal angles of the prothorax and
well developed, sharply triangular scutellum. In the simple
and filiform anterior tarsi, with the first four joints subequal
and but slightly longer than wide, shorter basal joint of the
posterior tarsi and almost wholly unimpressed mesosternum, it
is however peculiar, proving it to be an isolated type of the
subfamily ; there is but a single species as follows : —

Form oblong, convex, subalutaceous, rufo-piceous, the legs dark
rufous and the antennae pale ferruginous, glabrous, with a few^
short hairs on the hypomera and some longer bristling setae at the
sides of the epipleura; toward base ; head finely punctured, sparsely
behind, more close!}'' before, the suture, the sinus moderately wide,
rather shallow and parabolic, the margin of the oblique, rounded
and medially emarginate sides moderately reflexed ; antennae long
and very slender ; prothorax fully twice as wide as long, widest be-
hind the middle, subparallel, the sides subevenly arcuate, somewhat
more converging anteriorly, the apex broadly and strongly sinuate,
nearly three-fourths as wide as the base, the angles not deflexed
and evidently rounded ; base transverse ; surface rather finely and
sparsely but strongly punctate, becoming strongly and more
coarsely but not densely so laterally, the margins narrowly and
equally explanate throughout the length, the bead very fine, the
basal angles very feebly produced, right and not rounded ; scutel-
lum but slightly wider than long ; elytra scarcely as w'ide as the
prothorax, parallel and very feebly arcuate at the sides, arcuately
and gradually narrowed in apical two-fifths, the tip narrowly and
slightly prolonged and rather acute; surface nearlv smooth, finely,
feebly, submuricately and sparsely punctured, more closely but
not at all more coarsely toward the sides; prosterniuu moderately
finely, submuricately punctate, the process not margined ; abdo-
men rather gradually acute posteriorly, shining, finely and sparsely
punctate and sparsely puberulent ; tarsi long and very slender.
Length 6.5 mm.; width 3.25 mm. Oregon (northwestern).

rectus Csy.

A few examples were taken by Mr. Schwarz at Squally Hill,

which seems to be in the neighborhood of Astoria. The epi-

REVISION OF TENEHRIONID SUBFAMILY CONIONTIN^ I47

pleurae are more narrowed toward apex than in Coniontis and
are concave toward base ; they are very finely, feebly and
sparsely punctate.

Coelotaxis Horn.

This genus is a development from the archetypes of ConiontiSy
by reason of isolation on the solitary Island of Guadalupe,
probably during the time elapsing between some epoch in the
late Tertiary and the present time ; it is in fact simply Coniontis
with feebly ciliate sides of the body, shallow and submuricate
punctuation, subexplanate sides of the pronotum, rather more
developed and more obliquely truncate basal joint of the an-
terior tarsi, somewhat less elongate basal joint of the hind tarsi,
less impressed mesosternum and extremely minute scutellum,
the last being the most important and striking differential feature.
It is scarcely more closely allied to Coelus than is Coniontis
itself, any apparent resemblances, such as the ciliate sides of
the body, being merely superficial or directly due to some
similarity in habits. It is rather remarkable, when we con-
sider the very circumscribed habitat of the genus, that several
species, more or less closely allied it is true, should have been
gradually evolved ; these as represented in my cabinet by large
series, in all except two of the species, may be defined as
follows : —

Epistomal sinus larger, in the neighborhood of a third as wide as the
maximum width of the head; anterior angles of the prothorax
somewhat deflexed, the lateral bead more or less fine 2

Epistomal sinus very small, barely a fifth as wide as the head and very
shallow, the lobes broadly rounded ; anterior thoracic angles not
deflexed, the lateral bead unusually thick 6

2 — Elytra shining, rather sparsely and much less conspicuously punc-
tured, the pubescence shorter and barely visible. Form broadly
oblong-oval, strongly convex, dark blackish-brown, the under
surface, legs and antennte paler ; head coarsely, densely punctate,
broadly rounded, the sinus moderately shallow, subangular, the
sides broadly rounded, very much more prominent than the eyes or
the posterior canthus ; antennte long and very slender, the third joint
much longer than the fourth; prothorax slightly more than twice
as wide as its median length, the sides broadly, subevenly rounded
and converging from base to apex, a little more strongly so toward
the apex, which is three-fifths as wide as the base, deeply and
evenly sinuate, the angles but little deflexed and merely somewhat

148 CASEY

blunt ; base transversely truncate, the angles very feebly prolonged
backward and blunt ; surface coarsely, deeply and closely punc-
tate, more densely so sublaterally and again more finely and sparsely
near the sides, where it becomes rather broadly subexplanate pos-
teriorly, though narrowly so anteriorly, the bead not very thick,
strongly elevated ; elytra about a fourth longer than wide, the
sides subparallel and very feebly arcuate, rounding a little at base,
strongly rounding posteriorly to the broadly ogival tip, the surface
nearly smooth, the punctures rather small and feeble, varying but
little in size, becoming faintly muricate laterally and smaller
apically, much smaller and less deep than those of the pronotum ;
prosternum strongly, rather closely punctate, the process w^ell
developed, not margined at apex ; hind femora coarsely and con-
spicuously, though very remotely, punctate, the abdomen finely,
sparsely so medially, more coarsely and closely, muricately so
laterally. Length 8. 8-1 1.2 mm.; width 4.6-5.9 mm.

punctulata Horn

Elytra densely dull, unevenly and closely punctured, the pubescence
coarser and more conspicuous, though almost equally thinly dis-
tributed, the moderately long erect fulvous setae along the sides of
the body more developed 3

3 — Body broadly oblong-oval 4

Body narrowly oblong, with the sides more parallel 5

4 — Form very broad, moderately convex, dark castaneous, the under

surface, legs and antennas paler, dark rufo-fen^uginous ; head less
coarsely, very deeply and unequally punctate, very densely so cen-
trally, much less densely before the suture, nearly as in punctu-
lata in outline but less broadly rounded and less prominent at
the sides before the eyes ; antennae similar ; prothorax similar
throughout, except that it is a little shorter and much more than
twice as wide as long, with the apical sinus rather less deep, the
almost equally coarse and perforate punctures still denser ; ehtra
similar but with the parallel sides very slightly more arcuate, the
punctures about as large as those of the pronotum but not so
deep and intermixed with smaller punctures, very close-set and
muricate throughout ; under surface nearly similar, except that
the coarse and very conspicuous punctures of the hind femora
are generally less sparse, the basal joint of the anterior tarsi very
slightly less stout and somewhat less obliquely sinuato-truncate at
tip. Length 8.2-10.7 "i"^- 5 width 4.4-5.7 mm.

muricata Horn

Form rather less broad and more evenly elliptic, more convex, blackish-
brown in color, the legs piceous-black, the abdomen still blacker,
the sterna and antennie paler, rufo-ferruginous ; head as in 7)iuri-
cata and similarly punctate but more broadly and less obliquely
rounded at the sides, the antennie similar; prothorax less trans-
verse, barely twice as wide as its median length, the sides much
less converging from the base and more narrowly and more feebly

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I49

«

subexplanate, the bead finer, otherwise similar in outline and in
its very dense and perforate punctuation ; elytra more elongate,
more gradually and acutely ogival at tip, fully a third longer
than wide, the punctures similarly muricate and uneven in size
but not quite so dense ; prosternum less coarsely and more sparsely
punctate, the process narrower, the abdomen similar, the coarse
punctures of the hind femora remote, the basal joint of the anterior
tarsi stout and nearly as \\\ punctulata. Length i i.o mm. ; width
5.65 mm densa n. sp.

5 — Body narrow and parallel, strongly convex, blackish-brown or
paler, the under surface, legs and antennae rufous; head strongly
and densely, moderately coarsely punctui^ed, nearly as in muricata
but with the sinus broader and deeper, parabolic, the sides before
the eyes only moderately prominent, obliquely rounded, the an-
tennae nearly similar ; prothorax twice as wide as long to very
slightly more, formed as in densa and likewise very narrowly
subexplanate at the sides, coarsely, very deeply and densely punc-
tate, the punctures at the sides sparse and mingled with smaller
punctures ; elytra nearly a third longer than wide, the sides par-
allel and very feebly arcuate, not rounding at base, gradually and
arcuately narrowed posteriorly to the obtusely ogival tip, the sur-
face smooth, with the close-set, muricate and unequal punctures
of the preceding species; prosternum strongly, closely punctured,
the hind femora coarsely and sparsely, the abdomen finely but
strongly, sparsely so and with the punctures scarcely larger or
denser toward the sides. Length 7.6-9.7 mm.; width 3.75-5.0
mm angustula Csy .

6 — Form narrower, more elongate and more convex than in punctu-
lata or muricata^ rather less narrow and more elliptic than in
ang'Hstula.i brownish-black, the legs and antennse more or less
rufo-picescent, dull in lustre above; head broadly rounded at
apex, coarsely, densely punctate, especially toward the subobsolete
transverse impression, the sides more broadly rounded than in any
other species except detisa^ much more prominent than the eyes;
prothorax scarcely more than twice as wide as its median length,
the sides subevenly and distinctly converging and broadly arcuate
from base to apex, the apical angles prominent, very narrowly
blunt, the surface more coarsely and densely punctate than in any
other species, with a partial elevated impunctate line along the
middle, the sides narrowly concavo-explanate and obsoletely so at
base as usual ; elytra fully a fourth longer than wide, the sides
rounding and converging to the rather obtusely ogival tip behind
the middle, rather rough, with feeble traces of a ridge near and
oblique to the suture, not extending to base or apex, the surface
more coarsely muricate than in densa or muricata and equally
closely, with numerous small, suberoded and more opaque areas ;
under surface punctured as usual but more strongly ; pubescence
and lateral fimbriae as in muricata^ sparser and less fulvous than
in atigustula. Length ii.o mm. ; width 5.4 mm.

frontalis n. sp.

150 CASEY

The series of puncttilata and imiricata before me are com-
posed of twelve and nineteen specimens respectively, that of
angustula consisting of seven specimens, rendering the differ-
ences alluded to above patent at a glance ; the types of densa
and frontalis are still unique. In his original description of
Ccelotaxis (Trans. Am. Ent. Soc, V, 1876, p. 200), Dr. Horn
makes no mention of its most conspicuous distinguishing char-
acter, which is the minute and subobsolete scutellum, and, in
the accompanying table, interposes Cceliis between Ccelotaxis
and Eusattus. It is true that the very great obliquity of trun-
cature of the basal anterior tarsal joint, which varies but little
throughout the genus, gives rise to the appearance of a short
spur-like process homologous with the long process of Ccehis,
but this is simply a variation of the structure of this joint char-
acterizing Coniontis^ due to the more developed burrowing
habits of Ccelotaxis^ as shown by the fimbriate margins of the
body in the latter genus. At the same time, the long spur of
Cceliis is merely a still further development of this character,
surviving in more or less developed form from the common arche-
types of the subfamily Coniontinae, wholly lost in Eusattus
and allies and most developed in the burrowing types, such as
Ccelus. It simply proves that Coelus and Coniontis are descended
from the same ancestors, the remoteness of which can only be
conceived on considering the very radical antennal differences,
which must have taken untold ages to develop, and of which no
trace of any surviving intermediate form has yet come to light.

Tribe Ccelini.

In proposing a separate tribe based upon virtually a single
structural character, such as obtains in the case of Ctrlus with
one or two allied genera, much should depend upon the nature
of this differential feature, its degree of development and its
constancy or consistency. The antennae throughout the series
of genera discussed hitherto in this paper, constituting the Con-
iontini, are greatly developed, long and generally slender, while
in Ccelus and allies these organs assume a totally different
form, being very small, thicker and more compact. If there
were any intermediate living forms, and I cannot regard the

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I5I

case of certain far-distant exotics of the Praocini mentioned by
Lacordaire as being a case in point, there might be ground for
beHeving that these two groups of genera should not be tribally
separated, but there is no evidence whatever of such inter-
mediates and the proper course would therefore appear to be
the tribal separation here suggested. It is of course possible to
surmise that this very great reduction of the antennas in Coelus,
and especially Coclomorpha^ may be due simply to their bur-
rowing habits, but that this is not wholly true is proved by the
fact that in Ccelotaxis^ which, as shown by the lateral fimbriae
and by the more extended inner angles of the basal anterior
tarsal joint, has at least some burrowing propensity, there is no
tendency whatever to a shortening of the antennas, they being-
even longer and more slender than in the prevailing type of
Coniontis. At the same time, there is undoubtedly an unusu-
ally strong bond of affinity between the Coniontini and Coelini
in the modification of the basal anterior tarsal joint, proving
beyond question a common origin, but the elimination of all
intermediates in antennal structure decisively isolates the Coelini
at the present epoch of their history, and this is the only point
to consider in delimiting genera, tribes and other higher groups.
There are two genera of Coelini as follows : —

Body briefly oblong-oval, very convex, ciliate at the sides, the basal
angles of the prothorax obtuse but not much rounded ; antennae 1 1 -
jointed, the basal joint very long, arcuate, two-thirds as long as
the remainder ; submentum well developed ; eyes transversely
reniform, emarginate anteriorly ; process of the basal anterior
tarsal joint very long and stout. [Type C ciliatus Esch.].

Coelus

Body nearly similar in form but much smaller, similarly ciliate at the
sides, the basal angles of the prothorax well rounded ; antennae
lO-jointed, the basal joint much smaller, barely a third as long as
the remainder ; submentum not distinct ; eyes less transverse, not
emarginate anteriorly ; process of the anterior tarsus much smaller.
[ Type CcelomorpJia maritzjtia Csy.] Ccelomorpha

The members of these genera are very abundant individually,
and are only found on or near the ocean beaches burrowing
into loose sand.

152 CASEY

Ccelus Esch.

In this genus the body is oval to oblong-oval and always
strongly convex in form, with a dense fringe of erect fulvous
setse along the sides and with bristling hairs of the same nature
on the legs, along the anterior margin of the pronotum and on
the head behind the suture, but otherwise subglabrous. The
head is sinuate anteriorly, with a deep transverse suture delimit-
ing the epistoma, and the sides project laterally far beyond the
eyes, which are basal and more or less concealed by the pro-
thorax, the latter transverse, narrowed at the apex, which is deeply
sinuate, the base truncate and having a fine coriaceous margin
and the hind angles not at all produced posteriorly. The scu-
tellum is minute and triangular, being occasionally obsolete
and the elytra are rounded, feebly embracing the sides of the
body beneath, with the epipleurae narrow, gradually wider and
more or less concave basally. The legs are short and stout,
all the tibiae subequally dilated externally at tip, the terminal
spurs long and hollowed or flattened on their under surface, the
tarsi very slender, moderately long, with the joints swollen dis-
tally and bearing each a terminal corona of stiff fulvous setae,
the hind coxee narrowly separated as in Coniontis. The species
are rather numerous, and, so far as known, may be defined as
follows : —

Elytral punctures strongly muricate ; species inhabiting the mainland
of central and southern California exclusively. [Coelus insp.]...3

Elytral punctures simple, not muricate, the lateral fimbritB less devel-
oped ; species coniined to the islands off the coast of southern
California. [Pseudoccelus n. subgen.] 12

2 — Epistoma very deeply sinuate 3

Epistoma more broadly and less deeply, though conspicuously, sinuate,
the sinus trapezoidal in form, the bottom narrowly transverse ;
species more northern in habitat 5

Epistoma broadly and very feebly sinuate, the sinus generally subevenly
rounded 7

3 — Pronotum subevenly punctured throughout. Body broadly oblong-

oval, black to piceous in color, polished, moderately small in size;
head strongly and densely punctate, with a median impunctate area
at base as usual ; antennas rufous, having a compact fusiform club
of about three joints ; prothorax two and three-fourths times as
wide as its median length, the sides rather strongly converging
from base to apex and broadly rounded, more strongly so basally,

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I53

the apical sinus very deep, the angles not deflexed, rounded ; sur-
face coarsely, deeply and closely punctured throughout, not dif-
fering much near the sides, which are explanate, the bead fine and
feeble ; elytra but just visibly longer than wide, rounded at the
sides and obtuse behind, the immediate apex sharply ogival, the
sides becoming parallel toward base ; surface vertically declivous
and then slightly inflexed behind in profile, the punctures strong,
moderately separated, muricate, becoming granular posteriorly ;
prosternum densely punctato-rugose, the process moderately con-
stricted between the cox£e ; abdomen finely but strongly, more or
less closely punctate. Length 5.S-7.8 mm. ; width 3.7-4.9 mm.
San Diego globosus Lee .

Pronotum very unevenly punctate, with large patches devoid of punc-
tures , 4

4 — Body large in size, oblong-ovoidal, blackish-castaneous to rufo-
piceous in color, shining ; head rather coarsely, deeply and densely
punctate, with the usual basal impunctate area ; antennal club
fusoid, of about four joints ; prothorax scarcely two and one-half
times as wide as the median length, the sides much more parallel,
broadly rounded, more converging anteriorly, the apical sinus
very deep as usual, the angles more broadly rounded than in glo-
bosus; surface sparsely and rather finely but very deeply punctate,
the punctures becoming close and mingled with larger punctures
on the explanate lateral part, the bead rather coarse though only
feebly elevated and distantly punctured ; elytra but little longer
than wide, at the middle evidently wider than the prothorax, the
sides parallel and broadly arcuate, rapidly converging and rounded
behind, the surface nearly as in globosus but more rugose, the
punctui'es strongly muricate, rather sparse, less sparse and gi^anu-
lose posteriorly as usual ; pubescence long, subdecumbent and
conspicuous toward the sides posteriorly and much more conspic-
uously so than in globosus ; prosternal process only moderately
constricted between the coxae ; abdomen rather sparsely but dis-
tinctly punctate. Length 11. 0-12.0 mm.; width 6.4-7.0 mm.
Probably San Pedro, — G. W. Dunn grossus Csy.

Body similar to the preceding in general characters but uniformly
smaller and relatively narrower, polished, castaneous in color;
head notably smaller in size but otherwise similar ; prothorax a
little shorter and more narrowed at apex but otherwise similar ;
elytra as rw grossus^ except that they are less inflated and never more
than just visibly wider than the prothorax, the sculpture somewhat
less coarse and notably less close ; under surface nearly similar,
except that the prosternal process is less abbreviated and with the
punctuation less coarse, the metasternum, especially, more sparsely
and much less conspicuously so. Length 9.4-10.7 mm.; width
5.5-6.0 mm. Sta. Barbara, — G. W. Dunn solidus n. sp.

Body shorter, broader and more evenly oblong-oval in form than in
grossus and smaller in size, though unusually large for the genus,
Proc. Wash. Acad. Sci., April, 1908.

154 CASEY

deep black throughout, the antennae piceous with a testaceous
fusoid club of about three joints ; head densely but less coarsely
punctate ; prothorax nearly two and three-fourths times as wide
as long, the sides rounded and converging anteriorly, becoming
parallel, though rounded, toward base and converging toward the
very obtuse basal angles as usual, the apical angles less broadly
rounded than in ^r(95'5'?/5; surface more coarsely and much more
closely punctured, scarcely differing on the explanate lateral part,
the bead fine but well defined, not punctate ; elytra about as wide
as long, not wider than the prothorax, formed nearly as in glo-
bosus and similarly sculptured, sparsely pubescent toward the sides
posteriorly ; prosternal process strongly constricted, its neck gen-
erally but little more than half as wide as the dilated posterior
portion; abdomen polished as usual, not coarsely but strongly,
rather sparsely punctate. Length 7.6-9.6 mm. ; width 5.0-6.0
mm. Redondo Beach saginatus n. sp.

5 — Head well developed, very nearly half as wide as the prothorax.
Body oblong-oval, more elongate than usual, strongly convex,
polished, black, with the under surface piceous to piceous-brown
throughout ; head coarsely punctured behind, more finely and
sparsely before, the transverse groove, which is coarser and deeper
than in the preceding species, the epistoma more swollen trans-
versely toward the middle and more impressed or flattened at the
lobes ; antennal club fusiform as usual, of about four joints ; pro-
thorax rather less than two and one-half times as wide as long,
the sides moderately and subevenly converging from base to apex
and subevenly, moderately arcuate, the angles not more than blunt,
very much less roiuided than in the four preceding ; surface rather
finely, very sparsely punctate, the punctures unequally distributed,
having occasional vacant areas, rapidly declivous laterally and only
narrowly explanate at the sides, where there are some scattered
coarser punctures in addition, the bead fine and feeble ; elytra but
very slightly longer than wide, rounded at the sides, becoming par-
allel only basally, only just visibly wider than the prothorax, obtuse
and vertically declivous posteriorly, coarsely, sparsely punctato-
muricate, more closely and finel}^ granulate posteriorly, the fulvous
pubescence conspicuous near the apex; prosternal process strongly
constricted ; abdomen very finely and remotely punctate, the last
segment more coarsely and closely so as usual. Length 5.5-7.5
mm.; width 3.2-4.4 mm. San Francisco, — found only on the
sea-beaches ciliatus Esch.

A — Similar in general to ciliatus hwi with the body materially
larger and notably more elongate, the elytra much longer than
wide ; prosternal process even somewhat more strongly con-
stricted, the hind femora more coarsely and conspicuously punc-
tured. Length S.o mm. ; width 4.7 mm. San Francisco.

longulus n. subsp.

Head much smaller, very evidently less than half as wide as the pro-
thorax 6

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I55

6 — Form nearly as in ciliaUis but much smaller in size, blackish-cas-

taneous to much paler piceo-testaceous, polished ; head similar
though less coarsely, densely punctured behind the suture and
more finely, rather more closely before the latter ; prothorax simi-
lar in form and sculpture but somewhat more finely and densely
punctate toward the sides ; elytra similar in general form but
rather less coarsely and more closely punctato-muricate and still
more finely and closely granose posteriorly; under surface nearly
similar, the prosternal process equally strongly constricted. Length
4.S-5.8 mm.; width 3.8-3.4 mm. San Francisco, — found bur-
rowing in loose sand on Blue Mt., near the ocean beach.

debilis n. sp.

Form very much more dilated than in either of the preceding, oblong-
oval, polished, black, the under surface and legs piceo-rufous, the
antennae yellow-brown ; head small, rough and punctate behind,
and at the sides before, the suture, subimpunctate medially before
the suture, the epistoma much less tumid medially than in ciliattis ;
prothorax shorter and more transverse, the sides more strongly
converging and more strongly arcuate from base to apex, the ante-
rior angles rather more deflexed and more rounded ; surface simi-
lar and almost similarly finely and remotely, though less unequally,
punctured ; elytra not longer than wide, not at all wider than the
prothorax, the sides feebly converging and feebly arcuate from the
base, rapidly and obtusely rounded at apex, sculptured as in
ciliatus ; under surface nearly similar, except that the prosternal
process is less strongly constricted. Length 5.25 mm.; width
3.35 mm. Near San Francisco curtulus Csy.

7 — Punctures of the pronotum unequally distributed, leaving large

areas devoid of punctuation 8

Punctures of the pronotum almost evenly distributed 9

8 — Body rather broadly oblong-oval, polished, blackish, the elytra

dark red-brown, the under surface and legs rufo-piceous, the
antennae paler and more flavate as usual ; head coarsely and
densely punctate behind and with a mixture of large and small
punctures, sparse medially before the suture, the entire basal region
impunctate, the epistoma moderately convex medially, flat later-
ally ; antennae gradually clavate distally as usual ; prothorax two
and one-half times as wide as long, the sides strongly converging
from base to apex and broadly arcuate, gradually more rounded
basally, the apical angles very evidently rounded ; surface sparsely
and very strongly, but not very coarsely punctured, the punctures
closer and mingled with some much coarser punctures laterally,
the sides not very widely declivo-explanate, the bead rather strong,
especially around the apical angles ; elytra scarcely at all longer
than wide, not at all wider than the prothorax, the sides parallel
and broadly arcuate, rounding to the obtuse apex in scarcely apical
two-fifths, the immediate tip sharply ogival as usual; surface with
the usual strong, sparse and muricate punctures, becoming finer

156 CASEY

and closer granules on the alutaceous posterior declivity ; pro-
sternal process only moderately constricted between the coxae ;
abdomen finely but strongly, sparsely punctate, more coarsely,
densely so on the last segment. Length 8.4 mm.; width 5.1
mm. California (San Pedro) arenarius Csy.

Body more narrowly elongate-elliptic, strongly convex, polished, black
to rufo-piceous in color, the under surface and legs always some-
what paler ; head nearly as in arettarius^ very coarsely, densely
punctate behind the suture, more finely, clearly and densely so on
the flattened apical lobes, the median tumescent part of the epi-
stoma rugosely but less densely punctate ; prothorax less abbre-
viated, the sides less strongly converging, broadly and subevenly
arcuate from base to apex, the apical angles more distinct and less
rounded ; surface similarly very sparsely punctate, the larger lat-
eral punctures less coarse and the sides more broadly declivo-
explanate, the bead similar ; elytra not at all wider than the pro-
thorax, very slightly longer than wide, the sides converging and
rounding from before the middle, the apex obtusely and evenly
rounding from above, the surface inflexed near the apex, the
apical angle sharply ogival as usual, the sculpture somewhat less
coarse than in arenarius ; prosternal process very strongly con-
stricted between the coxte, the neck scarcely more than half as
wide as the dilated and rounded posterior part ; abdomen almost
similarly punctate, the last segment relatively less densely. Length
8.0-8.6 mm. ; width 4.7-4.8'; mm. Sta. Barbara, — G, W. Dunn.

sternalis n. sp.

9 — Body evenly elliptic in form, the prothorax not at all wider than

the elytral base 10

Body broadly oblong-suboval, the prothorax more swollen basally, and,
at a short distance before the base, distinctly wider than the base
of the elytra 11

10 — Form rather elongate, black, the antennae and tarsi paler; head

with irregular, dense, mingled coarse and finer punctures behind
the suture, the epistoma but feebly tumid medially, more or less
sparsely punctured ; prothorax two and one-half times as wide as
long, the sides strongly converging, strongly and very evenly
arcuate from base to apex, the angles only very narrowly
rounded ; surface rather finely but strongly, sparsely punctate,
more coarsely and with admixture of smaller close-set punctures
laterally, the sides rather narrowly, subequally explanate, the bead
very fine throughout ; elytra scarcely visibly longer than wide,
evenly rounded behind from above, the sides becoming parallel
only toward base, the punctures as usual, the posterior fulvous
pubescence conspicuous ; prosternal process moderately con-
stricted, the neck fully two-thirds as wide as the dilated posterior
part ; abdomen finely, sparsely punctate, more strongly and
densely on the last segment as usual. Length 6.9 mm. ; width
4.15 mm. Los Angeles Co obscurus n. sp.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN-<E I57

Form relatively a little broader, castaneous in color, the anterior parts
more blackish ; head nearly similar though notably smaller, barely
two-fifths as wide as the prothorax, the antennae only moderately
enlarged distally ; prothorax similar though a trifle more trans-
verse and similarly sparsely and strongly, though not coarsely,
punctate, more sparsely so laterally and with much less difference
in the size of the larger and smaller punctures, the sides less
abruptly, more declivously and somewhat more broadly explanate ;
elytra similar though not at all longer than wide, the punctures
throughout similarly sparse but not quite so coarse ; prosternal
process slightly more strongly constricted. Length 5.6-6.5 mm. ;
width 3.6-4.1 mm. Redondo SCOlopax n. sp.

II — Body stout, oblong, but little longer than wide, dark piceo-testa-
ceous throughout; head small, scarcely more than a third as wide
as the prothorax, densely punctate throughout, more coarsely
behind, more transversely rugose and medially swollen before, the
transverse suture ; antennae relatively more dilated apically or slender
basally than usual ; prothorax fully two and three-fourths times as
wide as its median length, inflated and with strongly rounded
sides basally, the sides very strongly converging and less arcuate
thence to the apex, the angles narrowly rounded ; surface closely,
rather coarsely punctate, the latter not differing in density but com-
posed of mingled coarse and finer punctures laterally, broadly
declivo-explanate at the sides, the bead fine but strong ; elytra
barely as long as wide, very broadly rounded behind from above,
the sides becoming subparallel and broadly arcuate in about basal
half, the sculpture more coarsely and closely muricate than in the
two preceding species, granulose and smaller posteriorly but having
the surface between the granules not at all alutaceous ; pro-
sternum coarsely and densely punctured, the process strongly con-
stricted ; abdomen finely and sparsely punctured medially. Length
4.9-7.0 mm. ; width 3.0-4.5 mm. San Diego, — G. W. Dunn.

amplicoUis n. sp.

Body similarly very stout and subquadrate, larger in size, deep black
in color, the antenuce pale, the legs piceous, similarly polished
throughout ; head larger, fully two-fifths as wide as the prothorax,
similarly densely sculptured, coarsely behind the suture, rather
more loosely so and finely rugose before the latter, the antennas
nearly similar ; prothorax a little less transverse, the sides similarly
inflated posteriorly but less converging anteriorly, the angles dis-
tinctly though not verv broadly rounded ; surface similarly coarsely
and closely, though a little more loosely, punctate, less abruptly
and rather less widely declivo-explanate laterally, the bead simi-
larly strong but not so elevated ; elytra similar, scarcely as long
as wide, the sides becoming parallel and more feebly arcuate
before the middle, the very coarse sculpture almost similar ; pro-
sternal process somewhat less strongly constricted, the neck about
two-thirds as wide as the rounded and dilated posterior portion ;

158 CASEY

abdomen rather strongly, moderately sparsely punctate, the last
segment densely. Length 7.6 mm. ; width 5.0 mm, San Diego.
Taken by the writer on the ocean beach latus Csy.

12 — Body broadly oblong-elliptical, moderately convex, piceous-black,
the surface polished ; epistoma broadly sinuate ; antennae with a
3-jointed club ; prothorax equal in width to the elytra, a little more
than twice as wide as its median length, widest immediately be-
fore the base, the sides rather feebly arcuate and strongly converg-
ent, moderately densely, evenly punctate throughout ; elvtra twice
as long as the median line of the prothorax, not longer than wide,
equally densely but more finely punctate than the pronotum, the
punctures not in the least asperate on the disk and only very feebly
so on the declivity and along the lateral margin ; process of the
first anterior tarsal joint extending under the next three ; marginal
fringe of the prothorax noticeably shorter and finer than in any of
the foregoing species. Length 7.0 mm,; width5.o mm. Island
of San Nicolas pacificus Fall

Body very convex, piceous-black, the legs and elytra brown ; epistoma
broadly sinuate; antennal club 4-jointed ; prothorax similar in out-
line to that of pacificus but shorter, the surface subopaque, densely
and coarsely punctate ; elytra shining, finely, densely punctate,
without trace of asperity ; process of the first anterior tarsal joint
extending beneath the next two ; marginal fringe longer than in
pacificus but thinner than usual. Length 6.5-7.0 mm. ; width
4.0-4.5 mm. Island of San Clemente remotus Fall

Seven of the above species are represented by large and very
complete series, showing with unmistakable clearness that they
are truly delimited from each other and in scarcely any way
opinionative ; intermediate individuals are in fact wholly want-
ing. The laws of intraspecific variation can be observed so
readily from these series, furthermore, that there can be little
or no doubt concerning the specific validity of those at present
represented by one or two specimens. Some of the species vary
prodigiously in the size of the body, to an exceptional degree
indeed, even among the Coniontinge, and there is a very remark-
able inconstancy in the development of the scutellum, which
varies from quite distinct to wholly obsolete within the limits
of probably all the species ; Lacordaire records a correspond-
ing inconstancy in one species of the Praocini (Gen. Col. V,
p. 216). In fact the scutellum is not very constant anywhere in
the Coniontinae, except in Ccelotaxis, where it is never more
than vestigial or very minute, and that in a group of genera
where, elsewhere, it has its maximum development, giving

I

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ I59

Calotaxis a notable isolation among the genera more closely
affiliated with Conioniis. It would appear almost certain that
the radically different elytral sculpture of the last two species
of the table, taken in connection with the more feebly developed
burrowing habits, as recorded by Mr. Fall (Can. Ent., 1897,
p. 242), is an ample reason for considering them at least sub-
generically removed from the normal forms.

Coelomorpha Csy.

This genus, while allied rather closely to Cceltis, differs in a
number of important structural characters, such as the lO-jointed
antennge with more abbreviated basal joint, in the less transverse
and virtually unemarginate eyes, which are similarly very
coarsely faceted, in the very feeble transverse epistomal suture,
in certain features of the prothorax, which, though possessing
the coriaceous basal margin of Ccehis, has the basal angles
rounded and the side margins without vestige of a beaded edge,
and in its more slender legs, all the tibiae, however, being simi-
larly dilated at apex. The process of the first anterior tarsal
joint is small, though slender, and very much less developed
than in Ccehis. The scutellum is very minute and similarly tri-
angular. The species, which are the smallest of the subfamily
Coniontina2, resemble Ccelus altogether in habits and facies and
may be described as follows : —

Form oblong-oval, strongly convex, rather shining, black to dark rufo-
testaceous, the sides of the body bristling with coarse and erect
but only moderately dense fusco-flavate setae ; head quite evidently
less than half as wide as the prothorax, rather closel}', finely and
very muricately punctate, not transversely impressed at the suture,
the epistoma not tumid, the emargination rather deep and abruptly
formed, circularly rounded, the surface bristling throughout with
stiff erect setse ; prothorax two and two-thirds times as wide as its
median length, the sides subevenly and moderately rounded, a little
more converging apically, the apex very deeply sinuate, with the
angles narrowly rounded ; base broadly, evenly and feebly arcuate,
the angles very broadly rounded ; surface with a few scattered
punctures, almost wholly impunctate before the middle medially,
more closely, asperately punctate, broadly declivo-explanate and
bristling with stiff erect setse laterally ; elytra just visibly longer
than wide, at the middle about a fifth wider than the prothorax,
very broadly rounded behind from above, the sides parallel and
broadly arcuate ; surface as in Ccelus but with the sculpture very

l6o CASEY

feeble, sparse and faintly muricate, more granuliform on the de-
clivity though still feeble ; epipleurae broader medially than at
base, rapidly becoming narrow posteriorly ; prosternal process
moderately constricted ; abdomen finely, sparsely punctate, the last
segment not at all more coarsely or closely so ; basal joint of the
hind tarsi but little longer than the next two combined. Length
2.8-3.4 mm. ; width i. 65-1. 85 mm. (cJ*, $). Lower California.

maritima Csy.

Form more broadly and more regularly subelliptic, less oblong, larger
in size, shining, pale flavo-testaceous in color, similar in pubes-
cence and in the bristling setas of the head and sides of the body ;
head relatively a little smaller, otherwise nearly similar ; prothorax
almost three times as wide as its median length, the lateral length
exceeding the median much more markedly than in maritima 'the
sides evenly and rather strongly arcuate, more converging from
near the base to the apex than in the preceding, the surface almost
similarly sculptured ; elytra at basal third fully a sixth wider than
the prothorax, inflated basally, more narrowed thence joosteriorly
and more narrowly rounded apically than in mai'itima^ the sides
more arcuate ; surface nearly similar though somewhat more
strongly muricate, approaching more closely the sculpture of
Coelus ; under surface and legs similar, the anterior tibiae with very
long erect setae internally, the posterior with shorter but more
numerous and very stiff erect setas externally and a few much less
conspicuous internally. Length 3.8 mm.; width 3.2 mm. (?).
Lower California pallens n . sp .

The material serving for the above descriptions was collected
by Mr. G. W. Dunn in various unrecorded parts of Lower
California.

The species above defined, as well as those of Ccelus^ Coni-
ontis and in fact quite generally throughout the subfamily, differ
from each other not so much by reason of any clearly defined
peculiarities of structure, as in a certain characteristic habitus
due to the summation of many minor features, such as general
form and outline, size, sculpture and vestiture. The study of
large series, where available, shows that this composite facies
is as persistent as elsewhere in the Coleoptera, and, in my
opinion, the various described forms are constant and true-
breeding, conforming to the usual definition of a species.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ l6l

ADDENDA.

I.

Having recently received a number of species of Praocini and
Nycteliini from Mr. Rolle, it becomes evident, even upon
cursory examination, that the subfamily Coniontinag should in-
clude both of these tribes and that the definition of the subfamily
previously given must be so generalized as to receive them.
The hind coxae, for example, very extended, oblique and com-
paratively slender in the North American tribes, become much
more abbreviated, subtransverse and ovoidal in the South
American groups, having there very nearly the prevailing form
of the Asidinee. The isolation of the Coniontinae as a subfamily
will therefore depend, as in so many other subdivisions of the
Tenebrionidie, upon the peculiar combination of a number of
characters, no one of which is absolutely constant but together
constituting a characteristic facies or habitus. Among the more
salient of these subfamily features may be mentioned antennal
structure, prolongation backward of the posterior thoracic
angles, porrect prosternal process, more or less pronounced
obliquity of the hind coxae, sparsely spinulose tarsi, rather
small, emarginate and obtrapezoidal mentum, generally exposed
and densely chitinized emarginate ligula, bifid mandibles,
sinuate clypeus and exposed emarginate labrum, well developed
tibial spurs and frequent basal dilation of the epipleuras. I
would propose the following arrangement of such tribes as are
represented by the material in my collection : —
Antennae long and slender, with or without a loose 3-jointed club... 3

Antennae very small, compact, with a feeble, compact and fusoid club,
the basal joint unusually developed 5

2 — Ligula wholly concealed ; gular sutures united and forming a
single furrow for a considerable distance behind the mentum ;
legs long; posterior coxae nearly as in As /da, widely separated
and subtransversely oval ; antennte peculiarly hispid, not clavate,
the last joint smaller, oval, pointed ; elytra widely embracing the
body, the epipleurae dilated basally. Southern South America.

*Nycteliini

Ligula largely exposed, densely chitinized; legs much shorter 3

I 62 CASEY

3 — Gular sutures uniting anteriorly, forming a single groove for a

short distance behind the mentum ; hind coxae feebly oblique, long
and cuneiform, distinctly separated, the abdominal process trun-
cate ; antennie with a 3-jointed club ; elytra widely embracing the
body, the epipleurae abruptly dilated at base. Tropical Atlantic
North America Branchini

Gular sutures rapidly diverging apically, delimiting a transverse piece
at the base of the mentum 4

4 — Hind coxae oval, subtransverse, distinctly separated though very

moderately, the abdominal process truncate to rounded ; elytra
rather widely embracing the sides of the body, the epipleurae
rapidly dilated at base; antennae with a 3-jointed club. Peru and
Chile *Praocini

Hind coxae slender, much longer, strongly oblique and cuneiform,
narrowly separated, the abdominal process acute or narrowly
rounded ; elytra widely to very narrowly embracing the sides of
the body, the epipleurae sometimes abruptly dilated at base ; an-
tennae with a 3-jointed club.^ Western temperate North America.

Coniontini

5 — Ligula and gular sutures nearly as in Coniontini, the hind coxae

also similar ; fourth abdominal segment relatively much less ab-
breviated, somewhat as in Praocis ; pronotum differing from any
of the preceding tribes in having a coriaceous basal margin ;
elytra narrowly embracing the sides of the body, never acutely
margined at the sides, resembling in this way only certain of the
subtribe Eusatti of the Coniontini ; legs rather short. Sea-
beaches of Southern California Coelini

The gular sutures in Coniontini are rather inconstant, some-
times being very distinct behind the mentum and sometimes
wholly obsolete ; it was this inconstancy that occupied my atten-
tion when on page 54 (ante), I stated that these gular characters
seemed to be rather equivocal in separating the Coniontini and
Branchini, but I am now convinced that they will serve this
purpose as suggested by LeConte. The tribe Nycteliini, as
outlined above, will comprise at least two genera which may be
defined as follows : —

' In the Coniontini tlie antennal club becomes a very variable feature. In the
subtribe Coniontes the antennae have an obvious, though feebly enlarged, 3-
jointed club. In the Eusatti, however, the character is very inconstant. Disco-
demus and Megasattus having no trace of a terminal club any more than in
Nyctelia ; Nesostcs and SpJiceriorith have a feeble 3-jointed club, but Eusattus is
composite, dubius having an obvious club of three joints, while the species from
froductus to the end of the genus have no tr.ice of an apical enlargement. I
think therefore that the genus Conipinus, for dubius, nanus and oblongulus may
have to be maintained.

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 163

Form oval, the hind body not inflated, the elytra deeply striate, the
sides of the prothorax fimbriate; habits fossorial. Atlantic Ar-
gentina. [Type Zophosis ttodosa Germ.] *Nyctelia

Form ovoidal, the hind body much inflated, the elytra not striate;
sides of the prothorax not fimbriate ; habits non-fossorial. South-
ern Andes. [Type JV. explanata n. sp.] *Nyctelioma

The body in Ny delta Latr., is smaller than in Nyctelioina
and there can be but little doubt that the habits are fossorial as
in Ccelus and other fimbriate species, including certain Praocids.
The t3''pe of Nyctelioma (n. gen.) may be described as follows :

Form convex, deep black, glabrous, rather strongly shining ; head with
some scattered coarse punctures laterally, the surface broadly im-
pressed along the sides ; prothorax a little more than twice as wide
as its median length, densely aureo-fimbriate at apex and base,
the sides parallel and almost straight nearly to the middle, then
broadly i^ounding, becoming strongly converging and straight
anteriorly, the apical sinus transvei'se at the bottom, the angles
porrect, acute and well defined ; base strongly, evenly bisinuate,
the surface gradually and feebly convex medially, with some
sparsely scattered punctures apically and broadly toward the sides,
the marginal bead fine, obliterated behind the middle ; scutellum
wholly obsolete ; elytra dilated, oval, at the middle nearly a third
wider than the prothorax, a little longer than wide, produced at
tip, evenly convex, gradually and rather broadly explanate at the
sides before the middle, the margin thickened and rugose ; surface
smooth, even, with some small and widely scattered granules
which become lineate in arrangement suturally and apically ;
hypomera with some minute, sparse, setigerous granules, smooth
and polished, the prosternum smooth, becoming rugose medially
and throughout the process, the latter rounded and thickly though
feebly beaded about its entire contour. Length 17.5 mm. ; width
II. 3 mm. Argentina (Mendoza) *explanata n. sp.

The above species is allied to latissima BL, but differs in
having no trace of undulation along the sides of the elytra. Of
a species called latissima Bl., Mr. Rolle sent me a considerable
number of examples which differ among themselves in very
confusing manner, showing that the delimitation of species in
this genus is fully as difficult as in any part of Coniontis.

II.

Dr. Georg Seidlitz (Ber. wiss. Leist. Ent., 1905, p. 162),
finds some fault with my recent papers because of the omission

164 CASEY

of bibliographic references, in contrariety with usual custom in
systematic works of this character. I can only state, in reply
to this, that those who are engaged in observing and recording
new facts in the vast and inexhaustible realm of nature in which
we find ourselves, have but little time for finding out what
others have done, except enough to avoid the duplication of
names or to point out an occasional error, for it seems to be far
more incumbent upon us — that is many of us, to correct the
errors of our esteemed coworkers than to allude to anything
good that they may be guilty of, leaving the recording of
what has been accomplished to those having special aptitude
and taste for such matters. The vocation of the bibliographer,
of whom the late H. Carrington Bolton was one of the highest
types, is a most important and useful one, and we all hail with
delight every new catalogue giving dates and references to
places of publication. But bibliography is becoming a specialty,
like everything else, and it is doubtful if any careful biblio-
grapher would accept the references and dates published by a
taxonomic specialist, but would go to original sources for such
data, so that work of this kind performed with considerable
loss of time by the searcher for basic facts, would have to be
reduplicated in any event.

In this connection it would evidently be better if figures could
be given of every described species, but these, in order to be
useful, must be rigorously exact and, as the original investigator
cannot as a rule spare the time and extra tax upon the eyes to
do this work, it must also be done by another very helpful
specialist, known as the biologic artist or draftsman, and it
becomes simply a question of halting all original scientific in-
quiry or publishing without illustrations, for entomological artists
are almost as rare as the proverbial denies gallinartmi, and those
that we have in this country are continuously employed in
museums at fixed salaries and consequently inaccessible to the
amateur, to whom scientific advancement has hitherto been in
large part due.

While referring to the subject of type figures, it should be
added that some Zoological Congresses have gone so far as to
state that descriptions of new species, more especially con-

I

REVISION OF TENEBRIONID SUBFAMILY CONIONTIN^ 1 65

chological, unaccompanied by an illustrative drawing, shall not
be considered valid, — a wholly impossible and illogical dictum,
as a good description in words is not increased a particle in value
by a poor figure, and the degree of excellence which the figure
must possess is not stated definitely in the edicts of the congress.
Another congressional dictum, to the effect that no description
of a new species, more especially botanical, can be considered
valid that does not involve at least a short Latin diagnosis, is
equally illogical and absurd. A description following a name
properly formed and printed in Roman characters, written wholly
in Japanese or in any other decipherable language, would be
perfectly valid. This might be very inconvenient for European
taxonomists, it is true, tending to halt scientific progress until
the descriptions could be translated, but it cannot properly be
assumed that every investigator of nature must be familiar with
some particular language as a condition for the recognition and
acceptance of his work, and, furthermore, to be of any real
value a description must extend beyond the few words of the
old-fashioned diagnosis.

III.
Remarks and Corrections Relating to the Revision

OF THE TeNTYRIIN^ IN THE PRECEDING VoLUME

OF THESE Proceedings.

Page 289 — The genus DitapJironotus was added to the table after the
latter had been drawn up and was inadvertently given an errone-
ous position. It should come immediately after Hylocrinus^
under the number "10" and referred to previously under
"7 — Surface glabrous."

Page 365 — The original type of Auchmobitis Icevis Lee, was lost at
sea, together with the original type of Ciyptadius injiatiis and
some other species, while being sent by LeConte to Lacordaire
for study.

Page 395 — The word " female " in the first line of the description of
plui7ibeus should read male. It was the impression of Le-
Conte that the two thoracic ridges of Epitragus {^Bothrotes^
pertain to the male and that the pronotum of the female was
even in convexity. The facts are the reverse of this, but I

I 66 CASEY

neglected to change the language of the original description in
that respect.

Page 463 — For " rcticidatzis^" in the 7th line, read nodulosus.
Page 499 — For " delapidated," in the 20th line, read dilapidated.
Page 522 — For " W. F. Rainbow," in the 2d line under IV, read
W. I. Rainbow.

PROCKKDINGS

OF THE

WASHINGTON ACADEMY OF SCIENCES

Vol. X, pp. 167-1S5. September 16, 1908.

THE SPERM-RECEPTACLE IN THE CRAYFISHES,
CAMBARUS CUBENSIS AND C. PARADOXUS.

By E. a. Andrews.

In some of the Crustacea all of the sperm that is to fertilize
the eggs is left by the male on the outside of the female and in
some others it is deposited in external receptacles. Both these
exceptional modes of transferring the sperm are met with in
the crayfishes and lobsters.

As far as known in all the crayfishes of the world, except
those of North America east of the Rocky Mountains, the male
deposits the sperm on the outside of the shell of the female, and
not in any receptacle. But in the above crayfishes of the
central and eastern parts of North America the sperm is de-
posited in special receptacles in the shells of the females, and
the same is true for the American lobster.

Before describing the sperm-receptacle in a Cuban and a
Mexican crayfish in which it was necessary to find it in order
to prove its general occurrence in all members of the genus
Cambarus, we will make a comparison of the receptacles in
the crayfish and lobster.

In both animals there is but one receptacle on each female
and this lies on the under side of the body, between the fourth
and the fifth pairs of legs.

The under side of the shell of the thorax of these animals

may be thought of as made of more or less fused plates, one

on each side and one on the middle of each somite. In the

somites of the first, second and third legs the imagined plates

Proc. Wash. Acad. Sci., August, 1908 167.

i68

E. A. ANDREWS.

are all fused into one mass, forming a groove, with the middle
plates as the bottom and the side plates as the elevated edges
to which the legs join. But between the fourth legs the middle
plate is recognizable as a distinct part of the sternal skeleton
and we will call it the annular plate as in the female crayfish
of the genus Ca?nbarus it is specialized as the so-called annulus
ventrali's, that contains the sperm-receptacle. In the next
somite, which bears the fifth legs, the middle plate is separate
and often spine-like while the side plates stand up on edge as
diagonal wings.

While the receptacle in Cambarus is contained within a single
middle plate, in the lobster it is formed by three plates, one
middle plate and two side plates, and thus though it has the
same position and the same physiological value it has not the
same morphological value. This will be more evident after a
description of the receptacle in the lobster, where it was first
recognized by Bumpus in 1897 (J. M., Vol. V). This organ is
made by two side wings or plates of the fourth leg-bearing somite
together with a wedge-like middle piece. These three plates
surround a middle space which is several mm. long, somewhat
less in width and more in depth. Bumpus states that it maybe
filled, in part, by a mass of sperm and largely by a wax-like
substance.

In the lobster the legs are inserted so close to the middle line
that the side plates come into contact at the middle line between
the fourth legs, but passing diagonally backwards flare apart
so that the middle or wedge-plate pushes in between them.
The receptacle is just above the wedge-plate which underlies
it and shuts it off from the exterior, as it runs forward like a
slanting shelf from its broad posterior base to its small, free
tip. The receptacular space would thus be entirely shut off
and inaccessible were it not for the fact that the two side plates
in front of the tip of the wedge may easily be forced apart by
inserting between them the hard, spatula-like tips of the male
stylets, or first abdominal appendages, which are very well
fitted for this purpose. Once inserted these organs are not
readily removed since the special ridge each bears, catches on
the face of the lateral plate. The lateral plates are normally

THE SPERM-RECEPTACLE IN CAMBARUS. 1 69

kept close together by the elasticity of the soft cushions formed
by the thin inner face of each plate. While the wedge-plate is
especially well calcified, the walls of the side plates, or wings,
remain membranous over most of their inner faces, so that much
of the lining of the receptacle, as well as the entrance to it, is
not brittle shell, but thin chitinous material.

Looked at from the inside of the body the protruding plates
of the under surface of the body are not solid thickenings of the
shell, but hollow protuberances, and from this standpoint the
receptacle may be described as an external space embraced on
the sides by two pouches of the shell, the right and the left
wing, and by a middle floor, a long, pointed pouch, the wedge.

It should be emphasized that the receptacle is merely an en-
closed region outside the body of the lobster, and that, as far as
known, it has no communication with any internal structure.

To determine the homology of this organ it is important to
know if all its parts belong to one somite or not. At first sight
the wedge-plate seems to belong to the somite bearing the fifth
pair of legs, as it sticks forward from the sternal ridge joining
these legs, while the side wings belong to the somite bearing the
fourth legs. But there are some reasons for holding that the
wedge belongs to the somite bearing the fourth legs, though
Herrick (The American Lobster, 1895), from a study of several
stages in the formation of the receptacle in young lobsters, con-
cluded that the wedge-plate belonged to the somite of the fifth
legs. However, an examination of his Fig. 5, Plate 33, sug-
gests that the cross line between the fourth and the fifth legs is
homologous with the similar line, which we have shown to be
the boundary of the annulus in young crayfishes (Biol. Bull.,
1906), and as the annulus seems to belong to the somite of the
fourth legs, it may be that Herrick was in error in assigning the
wedge to the somite of the fifth legs. Again, in the series of
middle and side plates, the middle plate stands a little posterior
to the line joining the side plates of its somite, as far as one can
judge from the adult condition, so that a recession of the middle
plate from its side plates is what is to be expected, while the
existence of a middle plate in advance of the side plates would
be exceptional. Thus there is a middle plate posterior to the

170 E. A, ANDREWS.

wings of the fifth legs, and there should be a middle plate pos-
terior to the wings of the fourth legs, and as such a plate we
may assume the wedge that projects forward from the line
joining the fifth wings.

In the male lobster the wedge of the female is represented by
a flat plate that is somewhat grooved along the middle. This
plate of the male may very readily be regarded as a middle
plate of the fourth leg region shoved back onto the somite of the
fifth legs.

Another mode of judging of the homology of the wedge and
its morphological position is a study of the internal skeleton, or
endophragmal system. In the crayfish there are transverse
ridges on the inside of the shell, between the side wings and the
middle plates, and from these arise the long internal bars, or
endosternites, that run up into the interior of the body, one on
the right and one on the left. In the male it is easily seen that
the endosternites come in between the wing plates and the
middle plate of the somite bearing the fourth legs and also be-
tween the wings and middle plate of the somite of the fifth legs.
In the female the same is true except that the wide annulus
takes the place of the narrow annular plate of the male.

The endosternites cannot be taken as exact bounds of the
somites, else the middle plate of the last thoracic somite would
be reckoned as on the abdomen, which the general conformation
renders unlikely.

In the lobster the extreme narrowness of the sterna brings the
endosternites close together at the middle line : but in the male
one can see that there is a pair of endosternites between the
middle plate and the wings in front of it on the last somite and
also between the hollowed out wings of the somite bearing the
fourth legs and a middle plate which may be regarded as the
annular plate. In the female the only difference is that the
peculiar development of the wings and the wedge raises the en-
dosternites of the boundary between these plates up onto the top
of the seminal receptacle, where they were noticed by Herrick
as parts of the endophragmal system.

It should also be noted that in the male lobster the receptacle
is represented by a triangular pit, some two mm. wide, deep

THE SPERM-RECEPTACLE IN CAMBARUS. I7I

and long, between the wings of the fourth legs and a rather flat
plate that may be regarded as belonging to that same somite.

Deciding that the wedge-plate really belongs to the somite of
the fourth legs we find no difficulty in homologizing it with the
annulus of the crayfish. We will call the wedge-plate an an-
nular plate. The only reason that this is not self-evident is that
the annular plate of the lobster is secondarily pushed far back
and fused to the sternal skeleton of the fifth leg somite. Even
in the crayfishes there is a marked recession of the annular
plate and in the lobster which has not the freedom of motion
between the fifth and fourth somites that is found in the crayfish,
the backward migration and fusion could take place to a greater
extent.

However, to establish the homology of the wedge with the
annular plate it is not necessary to show that they are on the
fourth leg somite, but only that they are on the same somite :
so that if the endosternites be taken as the exact bounds of the
somites both the annulus and the wedge would belong to the
somite of the fifth legs, but still be homologous.

While the receptacle of the lobster is a space between the
side plates and an annular plate the receptacle of crayfishes is
entirely within the annular plate itself : hence the two recepta-
cles are not homologous, though one element of each is the
same, the annular plate.

The general facts regarding the receptacle of the crayfish
are as follows : In the crayfishes of Europe, which are all of the
genus Astactis, it is said that the sperm is scattered over the pos-
terior part of the sternum of the thorax of the female, enclosed
in tubes of paste-like material, the spermatophores. Yet there
is, in both sexes, an annular plate, slightly marked off from the
great fused sternal mass that ends between the fourth legs. The
same is presumed to be true of all the crayfishes of Asia and of
North America, west of the Rocky Mountains as they are all of
the same genus, Astacus. On the other hand in the three score
and more crayfishes of the genus Cambarris, found east of these
mountains the annular plate is variously sculptured and provided
with a suture. Hagen, who discovered this annular plate
(Monograph N. A. Astacidae, 1870), supposed it had some gland-

172 E. A. ANDREWS.

ular function, but he recognized that it was a good aid in de-
scribing species of Cambarus and called it the annulus ventralis.
Since then it has been constantly made use of as a specific char-
acter. In 1895 we showed that the male of Cambarus affinis
deposits the sperm within a cavity in the annulus and subse-
quently we described the seminal receptacle in several species of
Carnbarus as a narrow pocket in the wall of the annular plate,
which arose in the young female as a shallow epidermal pit that
later deepened as a zigzag pocket.

In all crayfishes except Cambarus no receptacle is known
and no specialization of the annular plate is known outside of
the females of Catnbarus and of the American lobster. Yet
some sort of receptacle may yet be found in other crayfishes.
Thus the crayfishes of eastern Asia have the annular plate hol-
lowed out posteriorly, the male stylets armed wih complex
points and the legs provided with hooks, all of which leads one
to predict that a renewed search will discover some kind of
sperm-receptacle in these crayfishes. In that event the resem-
blances that led Faxon to call these crayfishes Ca^nbaroidcs, as
being like Cambarus^ would be strengthened in a way that
might add to the puzzling nature of the problem here presented
of close resemblances between animals in the eastern and west-
enr areas of the continents. North America and Eurasia along
with generic differences between the east and west of each con-
tinent.

Having shown that in Ca7nbarus affinis the sperm put by the
male into the annulus will remain alive all winter and that the
eggs laid in the spring will develop if this sperm is present
till then, though they did not develop w^hen the annulus was
removed, it was concluded that the sperm-pocket of the annulus
was an essential link in the chain of reproductive organs, without
which Cambari would come to an end.

To make way for a consideration of the possible mode of
origination of this peculiar organ, which seems to have no exact
homologue in other Crustacea nor in other animals, as it is an
unpaired, ventral, and not segmentally repeated organ, used
only for the storage of sperm, it seemed important to make more
sure that the annulus in all kinds of Cambari contained a sperm-
receptacle of the same nature.

THE SPERM-RECEPTACLE IN CAMBARUS. 1 73

Ortmann having shown that the genus Camharns (Proc.
Wash. Acad. Sci., 1906) could be divided into six subgenera;
the existence of the sperm-pocket in random members of all
subgenera would approximately establish its presence in all the
members of the genus. The annulus was found to contain a
sperm-pocket of the same fundamental structure (but with
specific differences) in species of the subgenera, Faxonius^
Bartoniiis and Camharns (Proc. Boston Soc. Nat. Hist.,
1906) and later the same was found true for a species of the
subgenus Camba7'ellus (Biol. Bull., 1908).

In the present paper the annulus is described in crayfish of
the two remaining subgenera, P}-ocambarus2,nd. Pa7'acambarus^
and the same sperm-pocket found in all.

Owing to the kindness of Dr. Ortmann, I have been able to
study the annulus in six specimens of Ca?nbarus (Procambariis)
cubensis Erichs., and in one of his type specimens of Cambarus
{Paracambarus) paradoxus Ortm. The former came to him
from the Rio Almendares, Calabazar, Prov. Habana, Cuba,
and the latter from the state of Puebla, Mexico, through the
Paris Museum. These specimens of Procambarus had so many
lost and regenerating limbs that this species would seem to be
an especially favorable one for the study of regeneration.

In this Cuban crayfish the annulus is known from a descrip-
tion of Faxon (Revision Astacidae, 1885) to be " composed of a
larger anterior, bilobed tubercle and a smaller posterior tubercle "
and Ortmann found that it showed an S-shaped fissure on the
posterior tubercle. This fact and the figure given by Ortmann
left little doubt that, when looked for, a sperm-pocket would be
found within this annulus. However there was doubt as to
what should be regarded as the real annulus since Faxon gave
the name to all three tubercles, while Ortmann stated that it
seemed to him only the posterior tubercle ought to be regarded
as the annulus.

The following illustrations, in the light of the above consider-
ations as to the morphology of the annular plate, will show that
although the receptacle is all within the posterior tubercle, yet
that alone is not the annulus, but all three tubercles together
form the annulus.

174

E. A. ANDREWS.

The width of the whole set of tubercles is about 2 mm. in a
female 48 mm. long, say 3 per cent, to 4 per cent, of the length,
which is the usual proportion of annulus to body length. But
the posterior tubercle, or " subannulus," as it might be called,

Fig. I. Ventral view annulus and neighboring region of female 50 mm.

long, 2flo-

Fig. 2. Posterior view
annulus and neighboring
organs of a female 35 mm.
long, right handed, 2aQ.

.^cTT^i

Fig. 3. Median lengthwise section of an-
nulus and neigliboring organs. Same speci-
men as in Fig. 2, 2.90 mm., A.

is only .75 mm. wide and .5 mm. long and thus not the right
size for an annulus.

Fig. I shows, right and left, the basal segments of the third,
fourth and fifth legs with the elliptical openings of the oviducts

THE SPERM-RECEPTACLE IN CAMBARUS.

175

on the third legs. Between the fourth legs are the flaring side
or wing plates, and between the fifth legs are two similar plates
which being set on edge show better from a posterior view,
Fig. 2. Between the fourth and the fifth legs is the mass of
three tubercles, shown enlarged in Fig. 4. Between the fifth
legs there is a small tubercle serially homologous with the above
set of three that form the annular plate.

The structures indicated in Fig. i are fringed with setae but
these are represented only in Fig. 2, which is a view from
behind. These setae add to the difficulty mentioned by Faxon
and by Ortmann, of finding the annulus.

Fig. 2 shows in the foreground the low middle plate, or
spine, of the sternum of the fifth legs and right and left of tha

Fig. 4. Ventral view of annulus of female 38 mm. long, 2.90 mm., A.

the side wings of the same somite. In the background are the
larger wings of the fourth leg somite and at the center the
tubercles. The posterior tubercle stands out free, attached
only along its contact with the paired tubercles, so that it can be
moved up and down, while the anterior tubercles are pretty
firmly soldered to the fused sternal mass anterior to them. The
sides of the anterior tubercles reach under the flaring wing
plates, as seen in Fig. 4, so that the entire set of tubercles has
the proper form and connections for an annular plate. The
peculiarity is that the posterior part of the annulus is set off as
a rounded tubercle, which is not known to be the case in any
other crayfish. As the posterior lobe is found to contain the

176

E. A. ANDREWS.

sperm-pocket it is not a new formation, fundamentally, but only
a specialization of the posterior part of an annular plate. The
two other tubercles seem to correspond to the paired elevations
met with on the anterior part of the annuli of some of the higher
Cambari.

In a median, lengthwise section. Fig. 3, the posterior lobe of
the annulus is seen to stand out from the rest of the annulus like
a fungus from a tree, forming a rounded shelf. Dorsal to this
shelf there is a large space and the black line that represents the
shell coming from the head, toward the right of the observer,
rises up over the protuberant anterior part of the annulus, between
the anterior tubercles, and sweeps around a deep cavity, to

Fig. 5. Ventral view of posterior lobe of annulus of female 45 mm. long, left

handed, 2A.

finally emerge over the summit of the small spine of the somite
of the fifth legs. This figure also shows the epidermis that
lines the shell and the vascular, sponge-work tissue that fills out
the annulus and all the adjacent region up to the nerve cord,
on which is represented the ganglion of the somite that bears
the fourth pair of legs.

The peculiarity of this annulus is that it contains the essential,
sperm-receptacle in the projecting posterior lobe.

This receptacle agrees so closely in structure with the pockets
that contain sperm in the higher species of Camhartis that
there is no doubt that it is used as a sperm-receptacle, though
no sperm was found in it in the five females examined. The
appearance of the pocket as seen in the posterior lobe of the
annulus made translucent, is represented in Fig. 5. The pocket
is an oblique slit that leads off to the observer's right as far as

THE SPERM-RECEPTACLE IN CAMBARUS.

177

the broken line, which represents its bottom. It is a very narrow
and not very deep pocket. The dotted lines indicate the walls
of the pocket and these are the continuations of the thick shell,
running in to line the pocket.

Fig. 6. Dorsal view of same specimen, 2A.

As shown in Fig. 6, this pocket comes to an end on the dor-
sal face of the posterior lobe of the annulus, after curving
around the posterior face. Coming up from the left of the ob-
server the mouth of the pocket runs a very short distance and
the part of the pocket directly at this end is very short, but

Fig. 7. Posterior face of posterior
lobe of annulus of the same specimen,

2A.

Fig. S. Section of part of
posterior lobe of annulus of
left-handed female 30 mm.
long, 2Z).

through the thickness of the annulus the other, or ventral end
of the pocket, is seen above and to the right of the figure.
This becomes easy of comprehension when the sperm-pocket is
viewed directly from the rear, as in Fig. 7. From this view the

178 E. A. ANDREWS.

posterior lobe of the annulus is seen to have a convex ventral
surface, a somewhat concave dorsal surface and rather sharp
edges, right and left. The mouth of the sperm-pocket is the
somewhat crescentric narrow cleft, represented by the black
line. The bottom of the pocket is represented by the broken
line and the walls by the dotted lines. The pocket thus passes
in obliquely to the right, and its ends where they pass over onto
the top and bottom are bent towards the middle plane.

The sperm-pocket here is more simple in its curves and more
easily seen to be a simple flat pocket than in any of the higher
Cambari yet studied.

In more highly magnified sections, as in the coronal section,
of which part is seen in Fig. 8, the sperm-pocket is seen to be
made by an invagination of the simple epidermis, but the cavity
of this pouch is largely filled in by the thick shell which the
epidermis has made and which is continued in from the thick
shell covering the rest of the annulus. The shell keeps its
usual character, having a thick, laminated inner part near the
epidermis, a much thinner outer part that is represented as clear
in the figure, and an outermost cuticle that is indicated by the
bounding line of the figure. The cavity lined by this shell is
very narrow and but slightly dilated at the bottom, yet it is
ample to contain innumerable sperms. It is probable that in
life, before the action of reagents, the two sides of the pocket
are so firmly in contact that the cuticular layer on each side may
allow no water to pass into the receptacle.

The sperm-receptacle of this crayfish is thus both very simple
and restricted to a small part of even the posterior lobe of the
annulus. In Fig. 3 the small area actually occupied by the
pocket is indicated by the little impitting in the posterior face
of the posterior lobe of the annulus, though in an exactly median
section this invagination would not show, as it is a little to one
side of the middle on the posterior edge. Fig. 7.

The great bulk of the interior of the annulus is filled with
areolated, spongy tissue represented in Figs. 3 and 8, and this
is full of blood and scattered corpuscles, as indicated in Fig. 8.
No glandular nor muscular tissues were found and no nerves,
though special search should show nerves for the few setae
found on the ventral surface of the annulus, Fig. 5.

THE SPERM-RECEPTACLE IN CAMBARUS. 1 79

One unexpected feature of the annulus of the genus Cainbarus
is the fact that it enables one to recognize a dimorphism amongst
the females of some, if not all, the species. The females are
either right handed or left handed in the sense that the sperm-
pockets are so placed and bent that some females have sperm-
pockets which are the mirror images of the pockets of the other
females of the same species. Indications are that about one
half are of one kind and one half of the other.

Though the sperm-receptacle is, in general terms, a median
structure it never lies entirely along the exact middle plane nor
is it exactly balanced, right and left, as are so many organs in
the anthropods.

In this Cuban Cambarics we find another illustration of this
dimorphism expressed in the annulus of the females. Of the
six specimens studied, four were left and two right handed.
The lengths of the right-handed ones were 35 and 38 mm, ; of
the left handed 30, 45, 45, 48. Thus the dimorphism is prob-
ably not a matter of age nor something that alternates in suc-
cessive periods of the same female as does the dimorphism,
so-called, of the males of the other Cambari. The differences
between the two forms of females are as follows : In the left-
handed females the sperm-pocket lies a little to the animal's left
of the middle of the body, Figs, i, 5, 6, 7. In the right-handed
female it lies to the right. Figs. 2, 4. Moreover in the two
cases the pockets slope in opposite directions, so that the right-
handed pocket is the mirror image of the left-handed one shown
in Fig. 7 ; that is, it looks like that one seen through the paper
from the other side. The crescent-like mouths of the receptacles
in the left-handed females open to the right and those of the
other to the left, so that they are to one another as new moon
and old moon.

The entire posterior lobe of the annulus is also right or left
handed, that is to say, in the left-handed females the most
prominent part of the posterior face of the annulus is slightly to
the right of the middle and the right side is the bigger. Figs.
i> 5> 6, 7. In the right-handed females the more protuberant
part is on the animal's left, as in Fig. 4.

Thus taking the species as a unit there exists a symmetrical

i8o

E. A. ANDREWS.

pair of sperm-pockets : each female possessing but one of a
symmetrical pair, while its mirror image is to be found upon
some other female.

The sperm-pocket of this species of crayfish, representing
the subgenus Procambat'uSy is thus fundamentally identical with
the sperm-pockets in all the other Cambari, thus far studied.

Turning now to the only specimen of the subgenus Para-
canihariis as yet examined with reference to the sperm-pocket

•x^

Fig. 9. Ventral view of annulus and neighboring organs of female

45 mm. long, 2«o.

within the annulus we find the general appearances indicated
in Fig. 9. The annulus itself is not so remarkable but the
spine between the fifth legs attains a height not elsewhere
known. The figure shows the bases of the second to fifth legs
inclusive and the grooved sternal mass that ends between the
fourth legs with flaring wing plates and a concave middle plate.
In the concavity so formed lies the annulus, which differs from
the usual form chiefly in rising up rather more in the middle

THE SPERM-RECEPTACLE IN CAMBARUS.

i8i

Fig. io. Side view of annulus
and post-annular spine, left half of
body removed, 2ag.

and in being decidedly concave on the posterior face. When
isolated it has the form indicated in Fig. ii, with the mouth of
the sperm-pocket lengthwise along its high middle part.

Posterior to this annulus the middle plate between the fifth
legs is developed as a very high spine, which is shown in its
true proportions and relations to other adjacent organs in Fig.
IO. In this side view, with one half of the body cut away, the
tall, conical, or somewhat flat-
tened spine juts forward to over-
hang the annulus, which in turn
stands up above the general level
of the mid-region of the sternum.
The spine bears at its apex several
long and sparsely plumose set£e
and a scant fringe of much shorter
setae is found along the edges of
the wing plates of the fourth and
fifth legs. In the background of the figure are the bases of the
fourth and fifth legs. The side plates are here seen in their
true elevation, while in Fig. 9 they are seen on edge.

In crayfishes the last thoracic somite is movable and the hard
middle plate or spine of this somite may be readily shoved
against the hind edge of the annulus. In Cambarus affinis
there is reason to believe that the female so shoves the middle
plate against the annulus as to liberate the sperms when they
are needed to fertilize the eggs, as these glide out of the open-
ings on the third legs and thence back over the annulus. In
Procambarus cubensis the low simple spine, Figs, i, 2, 3, may
thus act against the low posterior face of this annulus and, on
the other hand, in Paracambarus paradoxus the very tall spine
may be useful against the unusually high annulus, which is
concave on its posterior face, as if to receive the spine. While
the very tall spine may thus find its use in connection with a
very high annulus, as yet no use has actually been demon-
strated, and the great development of the spine in this Para-
cambarus may not have any value. The spine is a specializa-
tion of the middle plate of the last thoracic somite and as such
is to be regarded as homologous with the annulus, so that the

l82

E. A. ANDREWS.

unusual height of the spine may be correlated with the height of
the annulus as variations of serially homologous structures,
whether there is any use for the spine or not. Returning to the
annulus we find in it the usual sperm-receptacle, of much the
same character as in the higher species and more complex than
in Cambariis cubensis. The mouth of the pocket is a sinuous
groove between elevated lips running across the convex ventral
face and extending a little distance onto the anterior and poste-
rior faces. Beginning on the anterior face, Fig. ii, the narrow
mouth passes back from the middle line toward the left of the

Fig. II. Ventral face of annulus, 2A.

animal under the edge of a high, rounded tubercle. When the
annulus is seen from the front this tubercle is the highest point
of the annulus and from it the annulus slopes right and left
more steeply than is indicated in Fig. 11.

On the convex ventral face the mouth makes a U-shaped curve
to the animal's left and then a like one to the animal's right, to
finally pass onto the posterior face. Embraced in the second
curve is a second tubercle, less elevated than the anterior one
and on the opposite side of the mouth. On the posterior face
the mouth is seen as a short curved line coming down over the
edge about on the middle line and with quite an elevation on the
observer's right, close to the mouth.

As is faintly outlined in Fig. 11 the pocket into which this
doubly bent mouth opens lies below it and not off to one side as
much as in C. cubensis^ yet a more careful sketch, Fig. 12,
shows that the plane leading from the mouth to the bottom of
the pocket does slant somewhat and is not at right angles to the
surface of the annulus. Fig. 12 represents the sperm-pocket as

THE SPERM-RECEPTACLE IN CAMBARUS.

183

seen in a specimen made transparent ; the broken line stands for
the bottom of the pocket and the parallel rulings indicate the
cavity of the pocket, which is about the same thing as a plane
passed from the mouth to the bottom, since the width of the
pocket is so little. It will be seen that the cavity of the pocket
inclines first to the right, then to the left and again somewhat to
the right, nearest to the observer. The bottom is thus more
sinuous than the mouth ; the broken line in the figure more bent
than the continuous line that represents the mouth. Such in-
clination of the sperm-pocket is the

rule and the S-shaped curve is com- ^i::::^^^^-^ - — ■

mon in the higher species. In /' x^

another point this sperm-pocket sug-
gests the complex conditions in the
annuli of C. virilis and C. affinis,
and that is, the conformation of the
posterior part of the pocket. The
bottom of the pocket is not coexten-
sive with the mouth, so that the
mouth runs along the surface beyond
the limits of the bottom, both at the
anterior and the posterior ends. At
the posterior end the connection of
the mouth with the bottom is by an
inclined passage, sloping from the
surface forward. This passage is
slightly dilated right and left, as
indicated in Fig. 12. This little
chamber so formed is quite near the surface and farther from
the bottom of the pocket. It seems to be the same thing as the
'* recess " in which the sperm remains longer than elsewhere in
the annulus of C. affinis. But this chamber is easily understood
in this species, while in C. affinis it is obscure at first view.

The thick shell walls of the pocket are indicated in the figure
by the dotted lines ; the shell that lines the pocket was removed,
in preparation, from the epidermis and then it was seen that the
epidermis that had formed this pocket was an S-shaped, deep
groove, out of which the S-shaped shell represented by the dotted

Fig. 12. Ventral view of
sperm-pocket, as translucent
object, 4^.

184 E. A. ANDREWS.

lines, had been pulled. Thus, here, as elsewhere the shell-
pocket shown in Fig. 12 is cast in an epidermal mould of like
form, a mould that exists as an epidermal groove. The epider-
mal groove secreted so thick a shell as to fill up the groove, all
but a narrow slit, and it is this narrow cavity that doubtless is
filled with sperm by the male, though in this single specimen no
evidence of sperm was found.

A cross-section of this sperm-pocket would be essentially
like Fig. 8.

From this one female it is impossible to say whether this
species is dimorphic or not, but one may confidently predict
that an examination of many specimens would show that there
are both right- and left-handed forms. Some would have the
sperm-pocket the mirror image of that shown in Fig. 12, so that
these females would have the suture reversed and the pocket
inclining the opposite way at each turn, as compares with
Fig. 12.

We have thus demonstrated that the same sort of sperm-
pocket is found in these two crayfishes, Camharus cubensis and
Cambarus -paradoxus^ as in all other species of Cambariis yet
examined and have now found the same essential structure in
the annuli of representatives of all the six subgenera of Cayn-
bariiSy so that the sperm-pocket may be regarded as the funda-
mental feature of the annulus of all the Cambari.

SUMMARY.

The sperm-receptacles found on the ventral surface of the
lobster and of the crayfish of the genus Cainbartis are not
homologous with one another, though they have the same use
and location. In the lobster the receptacle is an external space
covered over by the annular plate of the seventh thoracic somite ;
in Catnbariis the receptacle is a narrow pocket invaginated into
that same annular plate.

It is shown that in each of the six subgenera of Cambarus
the annular plate contains the same sort of pocket and hence it
is most probable that sperm is stored up in such pockets in all
species of Cambari.

THE SPERM-RECEPTACLE IN CAMBARUS. 185

The middle plate of the eighth thoracic somite is homologous
with the annulus and is variously modified as a hard plale or
spine that may be pushed against the annulus. Probably the
female brings about the discharge of the sperm from the annulus
by use of this plate or spine, when the eggs are being laid.
Baltimore, March 21, 1908.

EXPLANATION OF FIGURES.

Drawn with camera and the Zeiss lenses indicated, reduced ^ diameters.
Figures 1-8, Cambarus {Procambarus) cubensis.
Figures 9-12, Cambarus {Paracambarus) paradoxus.

PROCEEDINGS

OF THE

WASHINGTON ACADEMY OF SCIENCES

Vol. X, pp. 187-248. December 24, 1908.

NOTICES OF DECEASED MEMBERS.

William Harris Ashmead.
1855-1908.

William Harris Ashmead, Assistant Curator Division of
Insects, United States National Museum, one of the foremost
American workers in systematic entomology, died in Washing-
ton October 17, 1908, after a lingering illness. Although his
death had been expected for some months, owing to the character
of the malady that laid him low, it was none the less a great
shock to his wide circle of friends among the scientific com-
munity of Washington and to the members of the Washington
Academy of Sciences, of which he was a charter member, and
of which he had been on several occasions Vice-President from
the Entomological Society of Washington.

Doctor Ashmead was born in Philadelphia September 19,
1855. He was the son of Captain Albert Ashmead and Eliza-
beth (Graham) Ashmead, and came of fine old colonial ancestry
on both sides. He was educated in the private and public
schools of Philadelphia, and early in life entered the publishing
house of J. B. Lippincott Company, of that city. Some years
later he went to Jacksonville, Fla., and with his brother
established a printing house for the publication of agricultural
books and other matter. He founded an agricultural weekly,
and a daily entitled The Florida Dispatch. He edited the
scientific department of the weekly, devoting himself chiefly to

Proc. Wash. Acad. Sci., December, 190S. 187

l88 WILLIAM HARRIS ASHMEAD.

the investigation of injurious insects. Through his interest in
this field, which became very great, he gradually built up a
large collection of insects ; and, experiencing great difficulty in
securing determinations, began the study of systematic ento-
mology. From the very first he was a tremendous worker and
produced results with astonishing rapidity. His contributions
to the scientific journals began in 1879, ^^^ from that date until
the time of his final collapse he produced a long series of
contributions to science, comprising more than 250 titles and
including many papers of great systematic value.

In 1887 he received an appointment as Special Field Ento-
mologist to the Division of Entomology of the United States
Department of Agriculture, for the investigation of certain
Florida problems. In 1888 he was appointed Entomologist to
the State Agricultural College and Experiment Station at Lake
City, Fla., and while holding that position, published one of
the very first bulletins produced by an entomologist of an agri-
cultural experiment station under the Hatch Act. It was en-
titled "Notes on Various Injurious Insects." In 1889 he was
made an assistant entomologist and investigator of the Division
of Entomology of the United States Department of Agriculture.
During the winter of 1889-90, on leave of absence, he went
abroad and studied for several months in Berlin. On his re-
turn he continued his work under the Division of Entomology,
and in 1895 was appointed Assistant Curator of the Division of
Insects, United States National Museum, which position he held
until a few months before his death. He donated his large
private collection to the Museum about 1898.

As a worker Doctor Ashmead was possessed of an enthusiasm
and of an industry that has rarely been equaled. For many
years he allowed himself but five hours, or a little more, sleep,
devoting the remainder of his time, with the exception of that
needed for eating, to an incessant study of the forms in which
he was for the time interested. The amount of work thus
accomplished was enormous. Either of his two main works,
namely, his Monograph of the North American Proctotrypid^e,
published as Bulletin 45 of the United States National Museum,
a work covering some 500 pages, or his Classification of the

WILLIAM HARRIS ASHMEAD. 189

Chalcid Flies or the Superfamily Chalcidoidea, published by
the Carnegie Museum, Serial No. 21, a quarto volume of 335
pages, would have been enough to have monopolized the work-
ing part of the lifetime of any one ordinary man. But aside
from these he left nearly ready for publication a great mono-
graph of the Braconid^, and he had published very many
smaller classificatory papers of high standing.

Doctor Ashmead was given the degree of Master of Science
by the Florida State Agricultural College, and in 1904 was
made a Doctor of Philosophy by the Western University of
Pennsylvania, the Monograph of the Chalcidoidea just men-
tioned having been submitted as his thesis. He was prominent
in scientific circles. He was a fellow of the American Associ-
ation for the Advancement of Science, and a corresponding
member of the American Entomological Society of Philadelphia.
He had been Vice-President of the Biological Society of Wash-
ington, President of the Cambridge Entomological Society
(1894), President of the Entomological Society of Washington
(189^-5), Vice-President of the Washington Academy of Sci-
ences (1888, 1893, 1894), honorary member of the Entomolog-
ical Society of Ontario, and Vice-President of the Association
of Economic Entomologists (1892). He married in Philadelphia
in 1878, Harriet, the daughter of Thomas O. Holmes. He
leaves a widow and one married daughter. He was a member
of the Cosmos Club, of Washington, and of its very important
committee on admissions.

When he came to Washington he was a man of large prop-
erty, which however, was greatly reduced by the disastrous
Jacksonville fire. This, however, did not appear to prey upon
his mind and he remained until the end the same cheerful, tire-
less worker in the field of pure science. Like so many inde-
fatigable workers in science, Dr. Ashmead was most helpful to
his fellow workers. His knowledge and his time were always
at the disposal of other workers, and he was noted for his help-
ful attitude towards younger investigators. He had the kindest
of hearts, and will always be remembered by those who knew
him.

L. O. Howard.

George W. Atherton.
1837-1906.

Geore W. Atherton, president of The Pennsylvania State
College, died at State College, Pa., on July 24, 1906.

He was born at Boxford, Mass., on June 20, 1837, coming
of good New England stock. Left fatherless at the age of 12,
he supported himself and aided his mother and sisters by work
in a cotton mill and later, on the farm and by teaching. He
worked his way through Philips Academy, Exeter, N. H., and
in i860 entered the sophomore class at Yale. At the outbreak
of the Civil War he responded to the call to arms, and on recom-
mendation of President Woolsey was commissioned First Lieu-
tenant in the Tenth Connecticut Volunteers. He took part in
Burnside's North Carolina expedition, where he served with
conspicuous bravery and efficiency, and was promoted to a
captaincy.

Leaving the army in 1863 on account of failing health, he
was graduated from Yale with his class, and on Christmas of
the same year was married to Frances D. W. Washburn, who,
with two sons and two daughters, survives him. For the next
four years he was a professor in the Albany Boys' Academy,
of Albany, N. Y. and for the succeeding year a professor, and
during most of the year, acting principal, of St. John's College
at Annapolis, Md.

In 1868, he was called to the newly established Illinois In-
dustrial University — since become the University of Illinois —
and took part in the organization of that institution, being closely
associated with its first president, Dr. Gregory. After a single
year's activity there, however, he accepted a very flattering
offer from Rutgers College, and for the succeeding fourteen
years filled the chair of political economy at that institution.

In 1873 he served as a member of the Board of Visitors to
the United States Naval Academy. In 1875 he was a member
of the commission to investigate the charges of corruption at

190

• GEORGE W. ATHERTON. I9I

the Red Cloud Indian Agency preferred by Prof. O. C. Marsh,
of Yale, having been added along with the Hon. Timothy
Howe, of Michigan, by personal action of President Grant, to
a commission of three, previously appointed by the Secretary
of the Interior. In 1876, much against his wish, he was made
the Republican candidate for Congress for his district. Although
the district was hopelessly Democratic, he threw himself into
the contest with characteristic energy and, while not elected,
ran considerably ahead of the Presidential ticket. In 1878 he
was appointed chairman of a commission to prepare and present
to the legislature of New Jersey a digest and revision of the
State system of taxation. While engaged in this work he
found time to study law, and was admitted to the New Jersey
bar while still carrying on his college work.

Both his studies in economics and his activit}^ in public affairs
naturally led him to interest himself in the development of in-
dustrial education, which occupied so large a share of educa-
tional thought during those years, particularly in connection
with the land-grant act of 1862. In 1873 he presented an
elaborate paper before the National Educational Association
upon the subject " The Relation of the General Government to
Education." In the course of this paper he traced in consider-
able detail the history and development of the land-grant col-
leges up to that time and emphasized those broad conceptions
of their functions in our educational system and of the impor-
tance of the type of education which they were designed to
give, to the concrete development of which he was to contribute
so largely.

In 1882 he accepted the presidency of The Pennsylvania
State College and began that work for which his previous life
had been the unconscious preparation.

This institution was one of the first fruits of that revolution in
the subject-matter and methods of education which characterized
the middle years of the nineteenth century. Its foundation was
so closely synchronous with that of the Michigan and Maryland
agricultural colleges that the degree of priority is largely a
question of definition. Chartered in 1855 and opened to stu-
dents in 1859, its first five years gave promise of a successful

192 GEORGE W. ATHERTON.

career ; but later, through official neglect and public indiffer-
ence, its affairs had reached a low ebb, and at the time when
Dr. Atherton was elected president the entire work of the college
was carried on in a single building, which also served to shelter
the families of several professors, and the total assured income
of the institution was $30,000 per year from the Morrill fund.
For the following college year, 1882-3, the number of collegiate
students dropped to 34 and the total enrollment to 87, while the
graduating class numbered 5.

The growth of the college in numbers and in material equip-
ment in the twenty-four years of Dr. Atherton's administration
was nothing short of marvelous. The enrollment of 87 in
1882-3 became 800 in 1905-6; the graduating class, on whom
he conferred their degrees .at his last official act, numbering 86.
The faculty increased in the same period from 16 to 66, besides
20 " assistants in administration." The work which in 1882
was carried on in a single building was, at the close of his ad-
ministration, distributed among fourteen, the cost of the new
buildings erected having been nearly $1,000,000, this including
the magnificent Schwab Auditorium and Carnegie Library.
The total of the State appropriations to the college during his
presidency was nearly $1,500,000, as compared with $183,000
during all its previous history.

Despite the demands of his college work, Dr. Atherton found
time and energy for other notable public activities. He was a
leading spirit in the activities which resulted in the passage in
1887 of the Hatch act providing for the establishment of agri-
cultural experiment stations, and also in 1890 of the second
Morrill act providing further endowment for the colleges of
agriculture and mechanic arts. He was largely instrumental
in organizing the Association of American Agricultural Colleges
and Experiment Stations and served two years as its first presi-
dent. In 1887 he was appointed by Governor Beaver chairman
of a commission of five, appointed under a joint resolution of
the legislature, *'to make inquiry and report to the next legis-
lature . . . respecting the subject of industrial education."
Special reference was had in the resolution to the question of
the incorporation of industrial training into the existing system

GEORGE W. ATHERTON. I93

of public education, and to the training of teachers for this pur-
pose. The report of this commission, which was largely his
work, was recognized as the most comprehensive and thorough
treatment of the subject up to that time. In 1891 he was for
the second time appointed on the Board of visitors to the United
States Naval Academy and in 1895 he was appointed a member
of the College and University Council of Pennsylvania, of which
he continued a member until his death.

Dr. Atherton was an indefatigable worker, never sparing
himself in the service of the college or of the larger public, and
he inspired his associates with like zeal. He had the courage
and poise of the born leader of men. Never dismayed or discon-
certed by opposition or attack, accepting defeat as the stepping
stone to future success, with an open mind welcoming every
suggestion from others yet with supreme confidence in his own
carefully considered conclusions, he bore his great responsibil-
ities with a quietness and simplicity which were an inspiration
to his associates. He was an optimist in the best sense, believ-
ing profoundly that right is stronger than wrong and that high
and worthy ideals must ultimately triumph, and he showed his
faith by his works.

Henry Prentiss Armsby.

1

Wilbur Olin Atwater.
1844-1907.

Prof. Wilbur Olin Atwater, whose death occurred
September 22, 1907, at his home in Middletown, Conn., after
an illness of nearly three years, was the son of a Methodist
clergyman well known in New England, and was born in Johns-
burg, N. Y., May 3, 1844.

He received his academic training at the University of Ver-
mont and at Wesleyan University, Middletown, Conn., gradu-
ating from the latter institution in 1865. During a period of
postgraduate study at Yale University, which led to the doctor's
degree in 1869, he was associated with Professors S. W. John-
son and W. H. Brewer and had his attention called to agri-
cultural chemistry and the great possibilities this subject offered
to investigators. With an insight which characterized him
throughout his career, he recognized the future possibilities of
such work and from 1869 to 187 1 he studied agricultural and
physiological chemistry in the Universities of Berlin and Leipsic
and acquainted himself with the European agricultural experi-
ment station movement. His career as a college teacher began
immediately after his return from Europe at the University of
Tennessee and at the Maine State College, and he was thus
brought into close touch with the movement for agricultural
education and research which was at this time taking shape in
the United States. He was called to Wesleyan University in
1873, where he held a professorship of chemistry for over thirty
years and at his death was head of the chemical department in
that institution.

The first agricultural experiment station in the United States
was organized in Connecticut in 1875 largely through Professor
Atwater's efforts, and he was made its first director. Interest
in the experiment station movement spread rapidly and the pas-
sage by Congress in 1887 of the Hatch act made possible the
establishment of such a station in every state and territory. In

194

WILBUR OLIN ATWATER. 1^5

Connecticut the unds were divided between two experiment
stations and Professor Atwater was made director of the Storrs
Station and retained this position for fourteen years. During
this period a relatively large amount of scientific work along
chemical lines and other lines related to agriculture was carried
on by the station. Of especial interest were Professor Atwater's
studies on the acquisition of atmospheric nitrogen by plants,
begun several years prior to the establishment of the Storrs
Station and continued as a part of the work of that station from
1888 to 1892.

On the invitation of Commissioner Colman, of the United
States Department of Agriculture, Professor Atwater consented
to become the first director of the Office of Experiment Stations,
established in the Department of Agriculture for the general
direction of the agricultural experiment station movement,
accepting this position on the condition that he be permitted to
retain the directorship of the Connecticut Storrs Station and his
college professorship at Wesleyan University. That this Bureau
has in later years followed with great success the general poli-
cies laid down by Professor Atwater is a proof of his wisdom
and foresight in establishing this work.

A long list of articles in scientific and popular journals, in
publications of the Connecticut Storrs Experiment Station and
United States Department of Agriculture, etc., shows the contri-
butions which Professor Atwater has made to the development
of agricultural chemistry and agricultural education. Espe-
cially noteworthy is the establishment of the series of farmers'
bulletins in 1889 under his advice and direction, a class of pub-
lications which has become of the greatest importance for the
dissemination of information along agricultural lines. He was
also founder of the Experiment Station Record, an abstract
journal published in the Office of Experiment Stations, of which
the first volume appeared in 1889. This journal covers the
field of agriculture and related sciences and its circulation
among students and investigators is world wide.

The influence of Professor Atwater on the development of
agricultural research in the United States has been greatly
broadened through the men who came into direct association

196 WILBUR OLIN ATWATER.

with him as students or co-workers and who have since become
directors of agricultural experiment stations, professors in agri-
cultural colleges, and influential in other capacities in the move-
ment for agricultural education and research.

Parallel with his studies in agricultural chemistry, including
animal nutrition. Professor Atwater began early in his career to
carry on investigations in physiological chemistry with special
reference to problems in human nutrition. Between 1879 and
1883 1^^ made extensive studies of the chemical composition and
nutritive value of American food fishes and invertebrates for
the United States Commissioner of Fish and Fisheries and
carried on work along similar lines for the Smithsonian Institu-
tion. Studies of the dietaries of people in Massachusetts and Can-
ada were also made for the Massachusetts Bureau of Statistics
of Labor and published in 1886. Nutrition investigations were
also carried on during this period as part of the regular work
of the Connecticut Storrs Experiment Station.

In 1893, Congress made an appropriation for investigations in
human nutrition in cooperation with the agricultural experiment
stations, assigning the supervision of this enterprise to the De-
partment of Agriculture where it was made a part of the work
of the Office of Experiment Stations. Headquarters for this
enterprise were established at Middletown, Conn., and Professor
Atwater was made its official chief. The work was broadly
planned and steadily developed until it became the most com-
prehensive investigation on this subject ever undertaken. The
cooperation of universities, colleges, and schools, experiment
stations, public institutions, and private organizations of various
kinds was secured in different parts of the country. Hundreds
of dietary studies of people of different occupations were made
and the results of similar studies throughout the world were
collated. Numerous digestion experiments with men were
carried on and special studies were made of the nutritive value
of different cereals, meats, vegetables, fruits, and nuts, the
effects of cooking and other forms of preparation on nutritive
value, and other important food problems. Special efforts were
made to improve methods and apparatus for such work.

In 1882-3, Professor Atwater devoted considerable time at

WILBUR OLIN ATWATER. I97

the Universities of Munich and Heidelberg in familiarizing him-
self with the German methods of studying nutrition problems,
and familiarity with the Pettenkofer-Voit respiration apparatus,
gained during this period, led him to undertake the construction
of a similar device which should, however, include the measure-
ment of the income and outgo of energy in addition to the in-
come and outgo of matter. The respiration calorimeter, which
was eventually developed by him and his associates is unrivaled
in its class as an instrument of precision, useful for the study of
a very large variety of problems connected with the physiology
and nutrition of man and animals. The bomb calorimeter and
the methods of its use were also materially improved under his
direction.

The studies in human nutrition already made with the At-
water-Rosa-Benedict respiration calorimeter have been very
important and promise to be still more so. Direct evidence has
been obtained that the law of the conservation of energy holds
good in the utilization of food in the human body ; important
data have also been obtained regarding the actual nutritive
value of different foods and the relations of food to muscular
energy and mental work.

As the Chief of Nutrition Investigations in the Office of Ex-
periment Stations, Professor Atwater planned and supervised
investigations which were carried on in about twenty States.
The results of this work are embodied in about one hundred
technical and popular publications issued by the United States
Department of Agriculture and the Storrs Experiment Station.

In the judgment of competent experts, the nutrition investi-
gations conducted under Professor Atwater's direction were
more thorough in their scientific methods, more extended in the
scope and amount of investigation, and more useful in the dis-
tribution and practical application of their results than any other
inquiry of the kind ever undertaken in this country or in Europe.

Professor Atwater's aid was often sought in the study of
nutrition problems. As instances may be mentioned the ex-
tended investigations of dietetic problems undertaken for the
New York State Commission in Lunacy, studies of the nutri-
tive value of alcohol carried on for the Committee of Fifty to

IpS WILBUR OLIN ATWATER.

Investigate the Liquor Problem, and the series of investigations
undertaken for the Carnegie Institution of Washington.

Professor Atvvater's career as a teacher was markedly suc-
cessful as is evidenced by the large number of his students and
associates who are carrying on work in this and other countries
of which he was in very large measure the inspiration.

As a public official working in a wide way in the organiza-
tion and management of enterprises for the general good,
Professor Atwater has earned the lasting gratitude of his
countrymen, especially by what he did in connection with the
agricultural experiment stations and the nutrition investigations.
To this work he brought a well trained mind and a true scien-
tific purpose. He had the enthusiasm and persistence neces-
sary to impress other men with the importance of his enterprises
and to carry him successfully over periods of opposition and
discouragement. He had unusual ability in the conception and
formulation of broad lines of work, and in attracting and hold-
ing men competent to give him such .aid as he needed to com-
plete these plans, put them into successful operation, and secure
substantial results.

Professor Atwater was a member of many learned societies

and received many honors in the United States and Europe.

His influence on the development of agricultural education and

research in this country and on the study of human nutrition

has been of very great and prominent value.

A. C. True.

I

Swan Moses Burnett.
1847-1906.

Swan Moses Burnett was born in New Market, Tennessee,
March 16, 1847, and died in Washington, D.C., January 18,
1906.

He graduated in medicine from Bellevue Hospital Medical
College, New York City, now the Medical Department of New
York University, in 1870, and first settled in Knoxville, Ten-
nessee, where he was engaged in practice for 5 years. In 1873
he married Miss Frances Hodgson. In 1875 he removed to
the District of Columbia, and soon attained prominence as a
specialist in ophthalmology and otology, as well as in literary
and art circles. He is the author of a Treatise on Astigmatism,
a Treatise on Refraction of the Human Eye, and over 64 dis-
tinct articles on diseases of the eye and ear, and chapters in
text-books. He was associated with Dr. John S. Billings in
the production of the National Medical Dictionary, and with
Doctors Norris and Oliver in that of the '* System of Ophthal-
mology." He also wrote a number of magazine articles and
public addresses.

In 1878 he was appointed lecturer on ophthalmology and
otology in the School of Medicine, Georgetown University,
continuing in this capacity until 1883, when he became clinical
professor, which position he filled until 1889, since which time
until his death he was professor in those branches. In 1879 he
established a post-graduate course in ophthalmology and otology,
in connection with his hospital and private practice, and ren-
dered most distinguished services as an author, teacher and
clinician.

He was president of the attending staff of the Central Dis-
pensary and Emergency Hospital, to which institution he gave
much of his time and skill. He founded and equipped in said
hospital the " Lionel laboratory " in memory of one of his sons,
" Little Lord Fauntleroy." This laboratory was the first to be

199

200

SWAN MOSES BURNETT.

established in connection with a hospital for clinical, bacteri-
ological and pathological research in the City of Washington.

He was for many years the ophthalmologist and otologist of
the Children's and Providence Hospitals, and also a member of
the consulting staff of the Episcopal Eye, Ear and Throat Hos-
pital. In 1889 he was elected president of the Medical Society
of the District of Columbia, and was a member of the Washing-
ton Academy of Sciences, Philosophical Society, Anthropolog-
ical Society, Historical Society, the American Ophthalmological
and Otological Society. In 1896 he was elected to the presi-
dency of the Cosmos Club of Washington, of which he was one
of the founders.

His degree of Doctor of Philosophy was bestowed by the
University of Georgetown in 1890. During his service extend-
ing over 25 years in the cause of higher medical education, he
was distinguished for his devotion to his calling and was unex-
celled as a teacher, scholar and gentleman. His kind, open and
earnest manner, his clear, concise and comprehensive lectures
could not fail to impress his students, while his shining example
always upheld the ethics of the profession and the dignity of the
physician and teacher.

Doctor Burnett died of chronic myocarditis, at his residence
916 Farragut Square, Washington. His second wife and his
son Vivian survive him.

Among his literary contributions and important writings are
the following : Translation of Edmond Landoldt's Manual of
examination of the eyes. A course of lectures delivered at the
Ecole Pratique, Rev. Edit., VII, 9-312 pp., i ch., i table, 8°,
Phila., 1879. A theoretical and practical treatise on astigma-
tism, VIII, 245 pp., 8°, St. Louis, 1882.

The principles of refraction in the human eye based on the
laws of conjugate foci. 67 pp., 8°, Phila., 1904.

Study of refraction from a new viewpoint. Phila., 1905.

See also Billings, J. S. The National Medical Dictionary.
Roy. 8°, Phila., 1890. — Landolt, E. The introduction of the
metrical system into ophthalmology. 8°, London, 1876. Dis-
eases of the conjunctiva and sclera. 82 pp., 2 pi., 8°, Phila.,
1898, contained in Vol. II of S3'st. Dis. Eye (Norris & Oliver).

SWAN MOSES BURNETT. 20I

Of the 64 distinct contributions to medical literature the fol-
lowingr are mentioned :

A case of diplacusis binauralis with remarks. 10 pp., 8°,
New York, 1877. Repr. from Arch. Ophth. & Otol., N. Y.,
1876.

A case of choroiditis exsudativa. 11 pp., 8°, New York.
Repr. from Arch. Ophth. & Otol., N. Y., 1877, 8°, VI.

Double optic neuritis (choked disc) and sloughing of the right
cornea accompanying a sarcomatous tumor on the right side of
the brain. 10 pp., 8°, New York. Repr. from Arch. Ophth.
& Otol., N. Y., 1877, 8°, VI.

Results of an examination of the color sense of 3,040 children
in the colored schools of the District of Columbia. 9 pp., 8°,
New York. Repr. from Arch. Ophth., N. Y., 1879, VIII.

A systematic method for the education of the color sense in
children. 4 pp., 8°, Washington, 1879. Repr. from Arch.
Ophth., N. Y., 1879.

A case of acute chemosis. 3 pp., 8°, New York, 1880.
Repr. from Arch. Ophth., N. Y., 1880, IX.

A case of primary external inflammation of the mastoid.
Repr. from Arch. Otol., N. Y., 1880, IX.

Objective aural sounds produced by voluntary contraction of
the tubal muscles. 3 pp. Repr. from Arch. Otol. N. Y.,
1879, VIII.

Color perception and color blindness. 7 pp. Repr. from
Arch. Ophth., N. Y., 1881, X.

Otomyces purpureus (Wreden) in the human ear. Repr.
from Arch. Otol., N. Y., 1881, X.

Are there separate centers for light, form and color percep-
tion? Rep. from Arch. Med., N. Y., 1884, XII.

The comparative frequency of eye diseases in the white and
colored races in the United States. Repr. Arch. Ophth. and
Otol., N. Y., XII.

A nomenclature of ophthalmology. Repr. from Am. Jour.
Ophth., St. Louis, 1884, I.

Theories of color perception. Repr. from Am. J. M. Sc,
Phila., 1884, LXXXVIII.

Clinical contributions to the study of ringscotoma. . Repr.
Tr. Am. Ophth. Soc, Boston, 1887, IV.

202 SWAN MOSES BURNETT.

An analysis of the refraction of 576 healthy human corneas
examined with the ophthalmometer of Javal and Schlotz.
Repr. from Tr. Am. Ophth. Soc, Hartford, 1888-90, V.

Reciprocal responsibilities. An address. 15 pp., 8°, Wash.,
D. C.

The physician as a man and citizen. 24 pp., 8°. Repr.
from J. A. M. Assoc, Chicago, 1891, XVI.

Contributions to clinical ophthalmology. Rep. from Arch.
Ophth., N. Y., 1892, XXI.

The general form of the human cornea and its relations to
the refraction of the eye and visual acuteness. Repr. from Tr.
Am. Ophth. Soc, 1894-6.

Some exceptional features in cataract extraction. Repr.
from Va. Med. Monthly, Richmond, 1895-6, XXII.

The racial and geographic distribution of trachoma in the
United States of America. Repr. from Am. Ophth., St. Louis,
1896.

A study of ocular coloboma. Repr. from Am. J. Ophth.,
St. Louis, 1898.

Gangrenous ulceration affecting the face including the
lids of both eyes and destroying the eyeballs, the results of
bites by a man. Repr. from J. Am. M. Assoc, Chicago, 1899.

Removal for relief of persistent headache of an ounce rifle
bullet imbedded in the bones of the right temporal fossa, where
it had lain unsuspected for 34 years. Repr. from J. Am. M.
Assoc, Cliicago, 1899.

A case of obstructed retinal circulation with a series of pictures
showing the changes in the vascular system during its rees-
tablishment of new vessels in the retina. Repr. from Ophth.
Rec, Nashville, 1899.

A series of cases of suppurative disease of the temporal bone
with comments. Repr. from Arch. OtoL, New York, 1900.

Double nasal hemianopsia following a fall on the head.
Repr. from Arch. Ophth., N. Y., 1900.

Methyl (wood) alcohol as a cause of blindness should be
placed on the list of poisons. Repr. Therap. Gaz., Detroit,
Dec, 1901.

The position of ophthalmology in the curriculum of the

SWAN MOSES BURNETT. 2O3

modern medical school, etc. Repr. from Am. J. Ophth., St.
Louis, 1901, Febr.

Helmholtz and ophthalmoscopy. Repr. from Am. J. Ophth.,
St. Louis, July, 1901.

Circumcorneal hypertrophy (vernal conjunctivitis) in the
negro. Repr. from Am. J. M. Sci., Philadelphia, 1904.

Appearance simulating optic neuritis due to unsuspected
irregular corneal astigmia. Repr. from Am. J. Ophth., St.
Louis, 1904.

An unusual form of exudate into the anterior chamber in
iridocyclitis after cataract extraction. Am. J. Ophth., St.
Louis, 1905.

George M. Kober.

Proc. Wash. Acad. Sci., December, 190S.

James Carroll
1854-1907

James Carroll, the second in command on the Yellow
Fever Commission, United States Army, was an Englishman,
born at Woolwich, on June 5, 1854. His early years, like
those of many men whose maturity is spent in a country not
their own by birth, are little known, especially as his entire lack
of egotism prevented his dwelling upon them. He was educated
at a private school, Albion House, with a view to his entering
the British Navy as an engineer student, but shortly before the
time came for his admission, he emigrated to Canada. For
some time after he reached Canada his life was that of a farmer
in the backwoods, until, in course of time became to the States,
and in January, 1874, he enlisted in the United States Army.

Upon enlisting he was ordered to the far west, and while
serving in Montana as a hospital steward he became interested
in the study of medicine. After some difficulty he obtained
permission to attend medical lectures at St. Paul, Minnesota,
and from this time on he pursued his medical education as he
could and where he could, until, on his return to the east he
finally completed it at the University of the City of New York
and the University of Maryland, receiving his degree of M.D.
from the latter institution in 1891. He became intensely inter-
ested in the new science of bacteriology, then beginning to de-
velop in this country, and availed himself of the graduate
classes just opened at Johns Hopkins Hospital to prosecute this
line of work. In 1895 he was assigned to duty in the Army
Medical Museum at Washington, where Walter Reed was Cu-
rator, and from this time forward these two men were constantly
associated as co-laborers. In 1899 Reed and Carroll were ap-
pointed by Surgeon-General Sternberg to investigate the true
nature of the Bacillus icteroides^ which Sanarelli had just de-
clared to be the specific agent of yellow fever. Their work on
this subject naturally associated their names prominently with

204

JAMES CARROLL. 205

the great yellow fever question ; and thus when, in 1900, an
Army Medical Commission was appointed to go to Cuba and
investigate the nature and transmission of this scourge of cen-
turies, Reed was made Chairman with Carroll as second in com-
mand.

Carroll arrived in Cuba on June 25, 1900, and shortly after-
ward the preliminary experiments were begun. Early in the
course of the work it became evident to the members of the
Commission that the proposed line of work could not be carried
on without experiments upon human beings and they agreed
that the initial experiment must be made upon one of them-
selves. Carroll volunteered for this service, and he always said
that the proudest circumstance of his life was that he was the
first person to succumb to mosquito inoculation. He had a
severe attack of the disease, during which his life was despaired
of, and although he recovered, it was with an organic heart
lesion which ultimately caused his death.

Carroll's services upon the Yellow Fever Commission, apart
from the inestimable act of self-sacrifice just mentioned, were
of the most essential character, and it is not at all too much to
say that without his native force and perseverance, guided by
his scientific knowledge and training, the work of the Commis-
sion could scarcely have been carried to a conclusion. Circum-
stances obliged Dr. Reed to leave Carroll in charge of the pre-
liminary experiments while he himself returned to the United
States on business connected with them, and it was entirely
through Carroll's exertions at this time that they were brought
to a satisfactory conclusion and the demonstration completed by
the time Dr. Reed was again in Cuba. Again, in February,
1901, when the fact that yellow fever is transmitted by the
Stegomyia calopus was definitely proved, and Reed went home,
Carroll remained behind for several weeks to determine one or
two additional points necessary to the perfect completion of
their experiments. His most valuable, as well as his most inde-
pendent service, of this description, however, was rendered in
the following summer, when he returned to Cuba in order to
undertake another line of experiments, intended to determine
whether the specific agent of yellow fever is contained in the

206 JAMES CARROLL.

blood. He encountered most serious difficulties before he could
achieve his end, but it was finally accomplished, and without
it the yellow fever experiments would undoubtedly have been
far less useful to mankind. The points which were established
by Carroll's individual efforts during this visit are :

1. That the specific agent of yellow fever is present in the
blood during at least the first, second, and third days of the
disease.

2. That the specific agent is destroyed, or at any rate attenu-
ated by heating up to 55° C. for ten minutes.

3. That yellow fever can be produced by the injection of a
small quantity of diluted serum taken directly from a patient
and passed through a Berkefeld filter.

4. That, as the specific agent is capable of passing through
a Berkefeld filter, it must belong to the class of organisms known
as ultra-microscopic.

Yellow fever was not the only disease which Carroll em-
ployed his knowledge of bacteriology to investigate. In 1898
he was sent to Camp Alger to study the blood of the fever
patients there and it was he who first showed that the illness
prevailing among the troops there was typhoid fever and not
malaria. On several other occasions he was employed to inves-
tigate typhoid fever.

After his return to the United States in 1901, Carroll continued
to disseminate the valuable knowledge which he had acquired
on the subject of yellow fever through the medium of the medi-
cal press. The first paper which he published independently,
on "The Treatment of Yellow Fever," is the first contribution
to the therapeutics of the disease after its mode of transmission
became known ; his last is the section on yellow fever in the
second volume of Osier's " System of Medicine."

For some years Carroll's services received no official recog-
nition, but during the last year of his life honors began to come to
him in which he took a manly and justifiable pleasure. He was
promoted from the rank of Lieutenant to that of Major, and two
universities, the University of Nebraska and the University of
Maryland, conferred upon him the honorary degree of LL.D.
In the summer of 1907 the heart lesion, which originated in his

JAMES CARROLL. 207

attack of experimental yellow fever, asserted itself and his
health began to fail. He died at his Home at Washington on
September i6, 1907.

In reviewing the facts of Carroll's life it is plain that he had
the elements of success in him from the first. He was the
typical vir tenax propositi, and this quality in his nature mani-
fested itself in a persistence which, if not a necessary element
of genius, is at least its closest ally and is often, pardonably,
mistaken for it. His personal character was one which com-
manded respect and inspired the warmest affection. He was,
as his former student, Dr. Donally, said of him, " a good man
and a square," true and just in all his dealings, faithful in all
his relations to those in authority over him, kindly and con-
siderate to those under his command. Whatever his hand
found to do he did it with his might, measuring the extent of
his exertions by his duties and responsibilities, not by the value
to himself of success achieved. He was modest almost to a
fault and one of his most striking characteristics was a singular
simplicity and trustfulness. He had an abiding sense of justice,
his trust in human nature causing him always to believe that
this sense was as strong in others as himself, while his belief in
the final ordering of all things for good was the simple confi-
dence of a child which trusts implicitly in higher powers whose
failure to do right it cannot conceive.

Howard A. Kelley.

Emil Alexander de Schweinitz.
1864-1904.

Emil Alexander de Schweinitz was born in Salem,
North Carolina, in the year 1864. He was a son of Bishop de
Schweinitz of the Moravian church, and a grandson of the Rev.
Lewis David de Schweinitz, who is well known on account of
his many additions to the knowledge of fungi and other plants
in the United States, Emil Alexander de Schweinitz received
his early education at the Nazareth Hall High School and the
Moravian College of Bethlehem, Pennsylvania, and subse-
quently entered the University of North Carolina from which
he received the degree of Doctor of Philosophy. From the
University of North Carolina he went to the University of Ber-
lin, and later entered the Universit}' of Gottingen, receiving from
the last-named institution the degree of Doctor of Philosophy.

During the time spent in Germany the greater part of his
labors was devoted to the study of chemistry and allied sub-
jects. Upon returning to the United States he was engaged to
teach chemistry in Tufts College, Massachusetts, and after a
short while was made Professor of Chemistry in the Agricul-
tural and Mechanical College of Kentucky. In 1888 he re-
ceived an appointment as an assistant in the Division of Chem-
istry of the United States Department of Agriculture. On
January i, 1890, he was transferred from the Division of Chem-
istry to the Bureau of Animal Industry in the Department of
Agriculture, and was placed in charge of the biochemical re-
searches which were begun by the last named bureau on that
date. The work along these lines increased so rapidly that a
separate Division of Biochemistry in the Bureau of Animal In-
dustry was created and Dr. de Schweinitz was placed at its
head. This position he occupied until the day of his death.

After entering the Bureau of Animal Industry his labors were
confined almost entirely to research work concerning the meta-
bolic products of disease-producing bacteria, the chemical com-

208

EMIL ALEXANDER DE SCHWEINITZ. 2O9

position of the bodies of these bacteria and the production of
immunity therefrom. Owing to the special interest of the
Bureau of Animal Industry in those diseases which attack the
domesticated animals, his attention was directed in great part
to the study of the etiology, the treatment, and the methods of
protection from such animal scourges as tuberculosis, hog
cholera, swine plague and glanders. His most important con-
tributions to science were probably those which dealt with the
production of immunity from tuberculosis. While not the first
to note a successful vaccination of laboratory animals against
this disease, he was the first to record the use of attenuated
human tubercle bacilli for the production of immunity from
tuberculosis in cattle.

In the later years of his life he had given much time and
energy to the production of a suitable vaccine for hog cholera.
At the time of his death he had almost completed several exten-
sive experiments dealing with the intertransmissibility of human
and bovine tuberculosis. These experiments which have since
been published, are generally recognized as important additions
to our knowledge of this subject.

Dr. de Schweinitz was elected to membership in the American
Public Health Association in 1896 and was also a member of
the Section of Bacteriology and Chemistry from the time of its
organization, having served on the council and various commit-
tees of the section. He was several times vice-president of in-
ternational congresses on tuberculosis and hygiene which were
held in Paris and Berlin, and his research work was well known
and highly regarded abroad as well as in the United States.

In addition to the duties Dr. de Schweinitz performed as
Chief of the Biochemic Division of the Bureau of Animal Indus-
try, he was also Dean of the Medical School and Professor of
Chemistry and Toxicology in that department of the Columbian
University. In 1895 that institution conferred upon him an
honorary degree of doctor of medicine. A complete biblio-
graphy of his writings has been prepared by Dr. Charles War-
dell Stiles and may be found in The Columbian University
Bulletin^ No. i.

His death, which was caused by uraemia, took place on

2IO

EMIL ALEXANDER DE SCHWEINITZ.

February 15, 1904, and came as the greatest shock to his co-
laborers and friends. The Bureau of Animal Industry has lost
a faithful and highly honored official and medical science has
lost an investigator who did much toward clearing up the little-
known paths of chemical pathology. We honored him for his
scholarly achievements, and mourn his loss as that of a friend
and an investigator who was cut off in the midst of his labors.
As brilliant as was his past work, the future promised even
greater achievements.

M. Dorset.

Daniel Coit Oilman.
1831-1908.

Daniel Coit Oilman was born in Norwich, Connecticut, July
6, 183 1. In 1848, at the age of seventeen, he was admitted to
Yale College and was graduated B.A. in 1852. His residence
in New Haven was in the family of his uncle. Professor James
L. Kingsley, whose varied learning, accurate scholarship and
keen perceptions were stimulating and inspiring. In college he
took a highly honorable position in scholarship, was president
of the Linngean Society, one of the editors of the Yale Literary
Magazine^ a member of Delta Kappa, of Alpha Delta Phi, and
of the Beethoven Society, the Atalanta Boat Club, of Skull and
Bones, and of Phi Beta Kappa. In the year following his
graduation he was engaged in private teaching and literary
work at New Haven, continuing at the same time his own
studies, and was entered for some months as a resident graduate
at Harvard College, where his home was with Professor Arnold
Guyot.

In December, 1853, he and his life-long friend, Andrew
Dickson White, sailed for Europe as attaches of the American
Legation at St. Petersburg, under Ex-Governor Thomas H.
Seymour, minister-plenipotentiary. Pending the arrival of
Governor Seymour, whom he preceded by a few weeks, he
traveled in England ; and when he was not yet twenty-three
years old, under the auspices of Mr. Richard Cobden and Mr.
John Bright, at a large meeting of the National Public Associa-
tion at Manchester, he delivered an address on "Common
School Education in America," which was enthusiastically re-
ceived. His connection with the legation at St. Petersburg
afforded unusual facilities for observing the work of the great
library and other institutions of learning, of technical schools,
and reformitories, particularly for children of the Imperial Court,
and of the great fortifications at Cronstadt during the French-
English-Russian war. As a correspondent of the New York

211

212 DANIEL COIT OILMAN.

Journal of Cotnmerce^ the Independent, and the Tribune, and
as an occasional contributor to other periodicals, his letters, be-
fore the days of ocean telegraphs, not only from Russia but
also from Berlin some months later, when he was a student in
the university, were interesting and instructive. During his
residence in Berlin he established lasting friendship with many
distinguished scholars, among whom were Professor Perts, the
historian and royal librarian, and, in the department of physical
and political geography in which he was specially interested,
with the eminent Karl Ritter and F. Adolph Trendelenburg.
In 1855 he was appointed commissioner from the state of Con-
necticut to the Universal Exposition at Paris, where he became
secretary of the Board of Associated Commissioners.

Returning to New Haven at the close of 1855 he was made
assistant librarian of Yale College in 1856, and becoming libra-
rian in 1858, he held that position until he resigned it in 1865.
He was appointed secretary of the State Board of Education,
was associated with the Honorable Henry Barnard in the publi-
cation of the Connecticut Common School Journal, and coope-
rating with Prof essor Arnold Guyot, prepared a series of school
geographies and maps. He was also a contributor to Apple-
ton's American Enyclopedia under the editorship of Charles A.
Dana, and with Professor William D. Whitney and others,
assisted Professor Noah Porter in the revision of Webster's Dic-
tionary.

After resigning the office of librarian in 1865 he devoted him-
self more directly to his duties as professor of physical and po-
litical geography in the Sheffield Scientific School, to which
office he had been appointed by the corporation of Yale College
in 1863. Associated with Professor George J. Brush and others,
he was efficient in extending and developing the work of the
school of which he became practically the chief executive, se-
curing for it large subscriptions for its permanent endowment,
especially in connection with the munificent gifts of Joseph E.
Sheffield, and Oliver S. Winchester and the family of Mrs.
Cornelia L. Hillhouse, for an astronomical observatory. In
1870 he was elected President of the University of California,
but declined the office, which, however, he assumed on his re-

DANIEL COIT OILMAN. 213

election in 1872. Continuing in that position for three years he
reorganized and greatly enlarged the work of the university
and was successful in establishing it on the firm foundation
where it has continued to grow and prosper.

Called to the presidency of the newly founded Johns Hopkins
University at Baltimore in 1875, before a brick or stone had
been laid, or a teacher or student enrolled, he devoted himself
heart and soul to its organization and upbuilding, and at the end
of a quarter of a century resigned the office, leaving behind him
in the University and in the Johns Hopkins Hospital of which he
was the first superintendent, and in the medical school of the
University, enduring monuments of his genius as an organizer
and administrator, of his inspiring influence with his colleagues
and students as an educator, and of his wise discrimination in
assembling a permanent staff of brilliant instructors and emi-
nent scholars and scientists of Europe and America as occasional
lecturers. From the beginning his motto was Men before
Buildings.

He was a frequent contributor to newspapers and periodicals
in regard to social science, civil service reform, charity organi-
zation, general education and scientific research. He delivered
many academic discourses some of which were collated under
the titles *' University Problems" and " Launching of a Uni-
versity."

He was the biographer of James Monroe, in the Statesmen's
Series, and of Professor James D. Dana of Yale College ; was
editor of the works of Doctor Francis Lieber and of Doctor
Joseph P. Thompson, and of a new edition of De Tocqueville's
Democracy in America. He was a contributor to Johnson's
Universal Cyclopaedia and was editor-in-chief of the New
International Encyclopaedia. He was chairman of the Commit-
tee on Awards at the Atlanta Exposition of 1895. He rendered
efficient service as a member of the Venezuelan Commission in
1896, under appointment by President Cleveland. He was
president of the American Bible Society ; president of the
American Oriental Society ; one of the commission to draft a
charter for the city of Baltimore, especially in the sections of
Education and Charities ; president of the Civil Service Reform

214

DANIEL COIT OILMAN.

Association ; president of the board of trustees of the John F.
Slater Fund ; vice-president of the Peabody Education Fund ;
an incorporator of the General Education Board, was for three
years president of the Carnegie Institution, and became later a
trustee of the Russell Sage Foundation. He received the
honorary degree of Doctor of Laws from Harvard University
and from St. John's College, Maryland, in 1876; from Colum-
bia University in 1867 ; from Yale University and from the
University of North Carolina in 1889; from Princeton in 1896;
from the University of Toronto in 1903 ; from the University of
Wisconsin in 1904 ; from William and Mary College and from
Clark University in 1905.

In his multifarious and important duties he never sought
political preferment, personal fame, or pecuniary reward, but
through a life of great activity " held his rudder true" with an
unswerving purpose to acquire and impart useful knowledge,
and by his voice and pen and personal influence to realize the
hopes of his youth in promoting and advancing sound education
in all departments from primary and technical schools to the
highest institutions of learning.

Between 1853 and 1908 he made ten voyages to Europe
extending his travels to Algiers, Egypt and Jerusalem. The
summer of 1908 was spent for the most part in southern Europe.
He returned on October 7, seemingly in improved health, and
after brief visits to his daughter and to relatives in Newport he
went to the home of his sisters in Norwich, Connecticut, where
he died suddenly on Tuesday afternoon, October 13, 1908.

He married in 1861, Mary Keycham, daughter of Tredwell
Keycham, of New York. She died in 1869, leaving two
daughters who survive their father.

In 1877 he married Elizabeth Dwight Woolsey, daughter of
John M. Woolsey of Cleveland, Ohio, and niece of President
Theodore Dwight Woolsey, of Yale University.

His domestic relations were of the happiest, and during his
long official career the liberal and gracious hospitality of his
household to all sorts and conditions of men, from youthful
students to eminent scholars of world-wide distinction, con-
tributed not a little to the promotion of the interests which were
dear to his heart. William C. Oilman.

William Rainey Harper.
1856-1906.

William Rainey Harper died in Chicago, Illinois, Janu-
ary 10, 1906. He was born in New Concord, Muskingum
County, Ohio, July 26, 1856, of Scotch Irish ancestors. He
was born with fine mental faculties and a genius for con-
structive work. At the early age of eight years he entered the
preparatory department of Muskingum College, a small de-
nominational school in New Concord, and two years later he
entered upon his collegiate course, completing it with honors
and was graduated with the degree of Bachelor of Arts at the
age of fourteen. He early developed a love for the Hebrew
language and literature and delivered his commencement ora-
tion in that language on the day of his graduation. For three
years after his graduation he pursued his studies without the aid
of a master, holding closely and intently to his work, and at
seventeen entered Yale University as a graduate student, re-
ceiving from that institution the degree of Doctor of Philosophy
at the age of nineteen. Thus early did he discipline his fine
intellect and bring his mental faculties under subjection to that
tremendous will-power that was characteristic of him through
all his brilliant career. He married the daughter of Rev.
David Paul, President of Muskingum College, and spent a year
as principal of the Masonic College at Macon, Tennessee. In
the fall of 1876 he accepted an appointment as tutor in the pre-
paratory department of Dennison University, and immediately
his constructive and organizing power manifested itself in a
plan for the development of Granville Academy. Before this
work was commenced, however, he was called to the chair of
Hebrew at the Baptist Union Theological Seminary in Chicago.
While occupying this position he perfected a system of teaching
Hebrew by correspondence, published text-books for the study
of Hebrew, established two periodicals called the Hebrew Stu-
dent and Hebraic, and started summer schools in Hebrew. He

215

2l6 WILLIAM RAINEY HARPER.

became principal of the Chautauqua College of Liberal Arts
and later of the entire Chautauqua system.

In 1886 he was appointed Professor of Semitic Languages in
the graduate faculty of Yale University and in 1889 was ap-
pointed to the Woolsey Professorship of Biblical Literature in
the same institution. September 18, 1890, he was elected
President of the new University of Chicago. His letter of
acceptance bears date February 16, 1891, and he entered upon
his duties July i of that year. Beginning with a general out-
line of what the institution should become, he obtained funds
to purchase about ten acres of ground for a site and secured one
million dollars in money and pledges. With this beginning the
Chicago University opened its doors in 1892. It is impossible
in this brief statement even to sketch the marvelous growth of
the university under his leadership. At the time of his death
the university had over sixty acres of ground in a great city,
forty buildings, fifteen millions of dollars in endowment and
property, a faculty of three hundred and fifty professors and
teachers, and over three thousand students. This result stag-
gers belief and when one approaches the subject more closely
and considers the infinite details of the work in securing the
enormous contributions, in planning the buildings, in organizing
the educational work, in securing the teaching staff, among
whom were so many distinguished educators, and in turning to
this institution such a large body of students, the mind and
energy and consummate leadership of the man appear in their
fullness and glory. It is not extravagant to say that history
does not give a precedent or parallel of such an achievement.

But this was not all. In addition to this administrative and
constructive work he was a great teacher and a productive
writer.

The work he loved most was teaching. He was a master of
his subject and he possessed the indefinable quality, common to
all great teachers, that aroused the student to the greatest pos-
sible interest in his subject and compelled the student to zealous
work. There was an influence which ran along the channel
of his words that no words can interpret, which was an inspira-
tion to everyone who listened to him. He did not seek so much

WILLIAM RAINEY HARPER. 217

to make his students believe what he believed, but he taught
them to think for themselves and reach rational conclusions.
He brought them a feast of knowledge which they were to
make their own by assimilation. He was both exacting and
sympathetic with his students but he had no place for the man
who would not make honest endeavor.

As a lecturer he won a brilliant place. Under his inspiring
and enlightening treatment the plainest themes became alive
with interest to almost any audience. As editor of the journals
which he founded there was one purpose running through all
of his work, and that was to popularize bible study and make
higher education attractive. He wrote extensively for these
journals during his life at Yale and Chicago. The list of books
which he published upon his chosen and allied subjects and the
important articles which he published in magazines are far too
numerous to enumerate here. All this was done while he was
planning for and bearing the heavy administrative duties of the
great university. To those who knew him intimately he will
always be remembered as the scholar and teacher, while to the
world at large his creative genius in the establishment of the
university will be his immortality.

The attractiveness and power of sincerity were well illustrated
in his life. Although he did not possess the arts of oratory or
brilliancy of style, thousands listened to him with intense interest
and followed his leadership with enthusiasm.

Dr. Harper was a man of great faith. He believed in things
not seen. He accepted faith as the evidence of things hoped
for, and without doubting laid his plans and worked with tre-
mendous energy to realize his hopes and justify his faith. He
did not believe that anything worth having came by chance.
He trained his mind, disciplined his will and believed in miracles
wrought by human patience and toil. Only those who knew
him intimately had any conception of his power of^ continuous
application, of his endurance, and of the great multiplicity of
labors which he performed. As a student and author he did
a full man's part and his life would have been successful if he
had done nothing more. As a teacher he bore his daily part
and, judged by years of service and the quality of his work, he

2l8

WII.LIAM RAINEY HARPER.

measured up to the full requirements of ordinary life. In the
organization, establishment and administration of the great
university he accomplished in sixteen years what might well
have been the labor of two generations of men. Not all that
he planned and did will remain, but there is so much that will
endure that there can be little room for regrets.

" By their fruits ye shall know them," is the divine test and
measuring-rod that is ultimately applied to every man. Thus
tested and measured, Dr. Harper stands pre-eminent. He died
without worldly estate, but the historian will administer upon
his possessions and find riches that neither time nor rust can
corrupt nor the vicissitudes of human events diminish or take
away. He belonged to the elite who are born into the world
to do quickly some great work. These have little time for
what the world calls pleasures. They begin with almost super-
human faith. They labor with masterful energy and take a
direct line to the cross. But in the final accounting it is found
that the world is vastly richer, that humanity has new aims
and new impulses, and God and truth are more clearly appre-
hended because these men with dynamic souls have lived for

a few years.

Charles Willis Needham.

Samuel Pierpont Langley.
1834-1906.

Samuel Pierpont Langley, the third Secretary of the
Smithsonian Institution, astronomer and physicist, was born at
Roxbury, Massachusetts, August 22, 1834, and died at Aiken,
South Carolina, February 27, 1906.

He was educated in various private schools and the Boston
Latin and High Schools, but owing to circumstances, he was
prevented from adding to this the advantage of a college
education. Nevertheless, he was a life-long student, was well
grounded in literature and the fine arts, in modern languages
and mathematics, and was altogether, aside from his scientific
eminence, a broadly cultivated man. After leaving school, he
devoted himself to architecture and engineering, and at the age
of 23 went westward and spent the next seven years in Chicago
and St. Louis, devoting his time to his profession, through
which he gained a modest competence. In 1864 he abandoned
what he had thought his life work and returned to New Eng-
land, spending some time constructing a telescope, and later in
European travel. Upon his return to Boston, the director of
the Harvard College Observatory, Professor Joseph Winlock,
invited him to become an assistant in that observatory, an offer
which he accepted, and from that time he dated his scientific
career.

In 1866, Mr. Langley became Assistant Professor of Mathe-
matics in the United States Naval Academy at Annapolis where
he reorganized the small observatory, the work of which had
been interrupted by the Civil War. The following year he
became associated with the Western University of Pennsylvania
as Professor of Astronomy and Director of the Allegheny
Observatory, where he remained for a period of twenty years.
By his inventions and his original work on the solar spectrum,
he gained that eminence which easily ranked him among the
foremost scientific men of his day. His turn for business affairs

Proc. Wash. Acad. Sci., December, 190S. 219

220

SAMUEL PIERPONT LANGLEY.

was shown when he secured a large portion of the money
required for the equipment of the observatory by *' selling time"
to the railroads, a plan which resulted in great practical comfort
to travellers by establishing a uniform time system, and finally
by the standardization of time throughout the United States.

Mr. Langley's studies were largely directed to the sun, more
especially to that part of the sun's energy known as the infra-
red, for the investigation of which he invented a delicate instru-
ment, the bolometer, now universally employed by astronomers.
He aided and conducted numerous expeditions to observe the
eclipses of the sun, the study of the corona and other phenom-
ena, and in every case he secured successful results. Having
great charm as a popular lecturer and possessing a singularly
clear and beautiful literary style, he popularized the modern in-
vestigation of the sun and the science of astrophysics in his
work "The New Astronomy," probably the most distinguished
scientific memoir, from the literary point of view, ever produced
by an American.

While still at Allegheny, Mr. Langley conducted a series of
experiments which led to his further work in Washington in
the solution of the problem of flying machines, or aerodromes,
as he called them. After numerous experiments he succeeded,
in 1896, in causing a steam driven machine, many times heavier
than the air, to make a free flight of over three-fourths of a
mile, and this was followed by another successful flight. These
performances were afterwards repeated many times with aero-
dromes propelled by both steam and gas driven engines, so that
to Mr. Langley will forever belong the credit of being the first
to successfully maintain in the air a mechanical device many
times heavier than the atmosphere, thus practically demonstrat-
ing the possibility of artificial flight. He later constructed a
large aerodrome, capable of carrying a man, actuated by a 52
horse-power gasoline engine, which unfortunately on two occa-
sions failed to be successfully launched.

After serving for a short time as Assistant Secretary of the
Smithsonian Institution, in 1887 Mr. Langley succeeded Spencer
F. Baird in his office as Secretary. During his administration
the one large addition to the fund, since the foundation of the

SAMUEL PIERPONT LANGLEY. 221

Institution, was secured, and, largely through his personal
efforts, two new branches, the Astrophysical Observatory and
the National Zoological Park, were added to it.

The tributes in recognition of his work are almost too numer-
ous to recite. He received the degree of D.C.L. from Oxford,
D.Sc. from Cambridge, and, among numerous others, the de-
gree of LL.D. from the universities of Harvard, Princeton,
Michigan, and Wisconsin. He was awarded the Henry Draper
medal by the National Academy of Sciences, the Rumford
medal by the Royal Society of London, and the Rumford
medal by the American Academy of Arts and Sciences, as
well as the Janssen medal from the Institute of France, and the
medal of the Astronomical Society of France. He was a for-
eign member of the Royal Society of London, a correspondent
of the Institute of France, a fellow of the Royal Astronomical
Society of London, member of the Royal Institution of London,
member of the Academia dei Lincei, of Rome, of the National
Academy of Sciences, and of many others.

He was also President of the American Association for the
Advancement of Science, Vice-President of the American
Philosophical Society, member of the Council of the National
Academy of Sciences, and a Trustee of the Carnegie Institu-
tion. He was a man of a singularly retiring disposition and of
a depth of affection which only those who knew him most inti-
mately ever fathomed.

Cyrus Abler.

Adolph Lindenkohl.
1833-1904.

Adolph Lindenkohl was born at Niederkaufungen, Hesse
Cassel, Germany, on March 6, 1833, and died in Washington,
D. C, June 22, 1904. He graduated from the Polytechnische
Schule, Cassel, in 1852. He came to the United States the
same year, and in 1857 was admitted to American citizenship.
He was employed in teaching for two years after coming to this
country, and on July i, 1854 was appointed to a position in the
cartographic work of the Coast Survey, where he remained until
his death, a remarkable record of fifty years of valuable service.

During the first year of this duty, James A. Whistler was a
fellow employe for about three months. It was found to be a
matter of difficulty to get the future distinguished artist to come
to the office with that regularity expected by the government,
or when there to devote himself to topographic drawing or to
etching views for the charts, as he preferred to sketch heads and
figures on the edge of the plates. Lindenkohl told of a per-
sonal effort to assist the young artist in punctuality, when he
went one morning to his room, the walls of which were found
to be covered with sketches on the plaster ; the young artist was
still in bed and so interested Lindenkohl in telling of his work
that the only result was that neither got to the office that morn-
ing. In the last month of his service Whistler was credited
with only six and one half days' work at one dollar and a half a
day and the experiment of his employment was terminated with-
out ill feeling on either side.

With others from the Coast Survey, Lindenkohl was assigned
to duty with the army during a portion of the Civil War from
1862 to 1864. He assisted in a topographic survey on the Po-
tomac River, and served as a topographer on the defenses of
Baltimore. He also assisted in the compilation of data for
various maps for the department of West Virginia.

In his regular duties in the office of the Coast Survey, Lin-

222

ADOLPH LINDENKOHL. 223

denkohl was engaged mainly in the compilation of data for
charts and the preparation of charts for publication, and he was
known particularly for the extensive store of information which
he acquired as to the material available for the charts, as well
as for the rapidity with which he worked ; for instance in put-
ting hachures or hill shading on charts he had both great speed
and unusual skill.

He gave much attention both in connection with and outside
of his official duties, to studies on subjects related to geography
and the physics of the sea, particularly deep-sea temperatures,
densities and currents. He wrote a number of articles on these
and similar subjects, which were published in the Reports of
the Coast and Geodetic Survey, Pctermann's Miiteilicngen, the
American Journal of Science, and elsewhere.

In 1884, in a paper on the " Geology of the Sea-bottom in the
Approaches to New York," he dealt largely with the submarine
channel of the Hudson River. In a paper in 1895 on the Gulf
Stream and circulation of the Gulf of Mexico he discussed the
evaporation, precipitation and influx from rivers and currents,
the variations of temperature and density in the Gulf waters,
and the relation of these phenomena to the Gulf Stream. In
1897 he published the results of an investigation of the salinity
and temperature of the North Pacific Ocean.

His inquiries along such lines are the more interesting be-
cause owing to the natural obstacles to investigations of the sea,
in the present distribution of scientific effort the oceanographic
problems are receiving less attention than their importance
warrants.

At the time of his death Mr. Lindenkohl was the senior
draftsman in the Coast and Geodetic Survey, having been re-
warded by promotion through all the grades of his branch of the
service. He was of a kindly and unassuming disposition, which
endeared him to all with whom he came in contact.

G. R. Putnam.

Henri Louis Francois Marindin.
1843-1904.

Mr. Marindin was born at Lausanne, Switzerland, July 2,
1843, and received his early education in the Swiss schools.
He came to the United States before attaining manhood, and
finished his scholastic education in the Owego Academy at
Owego, New York, 1860-1863.

He entered the Coast Survey as aid on November 26, 1863,
and was soon assigned to duty in a party engaged in the survey
of Roanoke River, North Carolina, made at the special request
of Admiral S. P. Lee, flag officer of the North Atlantic block-
ading squadron, the party being quartered on the gunboat Sey-
mour for that purpose. In 1864 he served in a topographic
party at work in the vicinity of Bermuda Hundred, Virginia,
under the orders of Major General Butler, with Brigadier
General Weitzel in immediate charge of the work, and later in
the same year, and in January, 1865, he served in a topographic
party engaged in work along the Potomac River under the
direction of Major C. S. Stewart. He thus began his service
to the nation of his adoption by aiding in the perpetuation of
the Union, and continued to serve his country faithfully until
death ended his long and honorable career.

In 1865 he was engaged in special surveys for a canal route
through Nicaragua, and in 1870 he was engaged in similar
work on the Isthmus of Darien.

His special work in the Coast Survey was in the field of
physical hydrography, and man}'^ important features in the cur-
rents and in the development of harbors and bars along the
Atlantic and Gulf coasts of the United States have been made
known to commerce as the result of his systematic and careful
investigation of the complex problems presented as the result
of the action of winds and tides on the waters of the sea. He
spent more than eighteen years in command of vessels of the
Survey while engaged in this work.

224

HENRI LOUIS FRANCOIS MARINDIN. 225

On March 24, 1897, he was appointed by President Mc-
Kinley to represent the Coast and Geodetic Survey on the Mis-
sissippi River Commission, and after that date devoted much
time and attention to the duties thus imposed upon him, but
continued his regular work on the Survey whenever it was pos-
sible to do so. His particular ability was recognized by numer-
ous special assignments to duty in connection with harbor boards
to establish harbor lines, and in the establishment of speed trial
courses for vessels of the navy, and is shown in several scientific
discussions of various physical problems relating to hydrog-
raphy which have appeared from time to time as appendices
to the annual report of the superintendent.

His death took place March 24, 1904.

Isaac Winston.

Herbert Gouverneur Ogden.
1 846-1 906.

Herbert Gouverneur Ogden was born in New York
April 4, 1846. He descended from Revolutionary stock,
Francis Lewis, a signer of the Declaration of Independence,
being one of his ancestors.

His career in the Coast Survey commenced in 1863, when the
Civil War was at its height. In common with a number of his
brother officers, he was assigned to duty with the army and then
with the navy, participating in the dangers and vicissitudes of
active warfare.

In 1865 he served as a topographer on the Nicaragua Expe-
dition.

In 1870 he was a member of the first naval exploring expe-
dition to the Isthmus of Darien.

In 1893 he had charge of a section of the exploratory surveys
for locating the international boundary between Alaska and
British Columbia.

He was appointed by the President one of the original mem-
bers of the Board on Geographic Names, and continued a mem-
ber until the time of his death.

These were special assignments. In the regular course of
his duties in the Survey, as his experience increased with length
of service, he showed his versatility by engaging in and eventu-
ally directing nearly every one of the many branches of the
work.

In 1880 he was placed in charge of the Engraving Division
of the office. This position did not, as its name would imply,
consist solely of superintending the work of expert engravers.
It involved a knowledge of the whole range of chart construc-
tion and publication from the surveying operations in the field
to the final verification of the chart from the press. It required
an encyclopaedic memory for details both of methods and locali-

226

HERBERT GOUVERNEUR OGDEN. 227

ties, since the accuracy of the finished product depended in a
large degree on his decision.

In 1898 he was appointed Inspector of Hydrography and
Topography. His new duties were rendered the more difficult
from the fact that a change was being made in the organization
of the Survey, and also on account of the rapid expansion of
the field of work, due to the development of Alaska, the acqui-
sition of Porto Rico and the Philippine Islands.

In this position it came within the scope of his duties to for-
mulate a general plan for hydrographic and topographic sur-
veys ; to make field inspections of the work ; also of the ships,
and to supervise their repairs and maintenance. Only those
engaged in the same lines of work can fully appreciate the im-
portance and value of what Mr. Ogden accomplished. Of the
many thousands who traverse our coasts in ships, there are few
indeed who are aware to what extent they are indebted to him
for the integrity and completeness of the charts on which their
safety depends.

The leading trait of Mr. Ogden's character — the one which
endeared him most to his associates, both old and young — was
his unfailing interest in their welfare. His ever ready sympathy
attracted the confidences of his juniors, and his advice and as-
sistance was ever constantly sought by them. At the same time
his loyalty to the service never allowed his kindly feelings to
override his high sense of duty. Decided in his convictions, he
was firm in refusing ill-considered or improper requests, but he
could deny in such a tactful and considerate manner as rarely
to wound the most sensitive.

Having a genial and sanguine disposition — a man of simple
habits — his friends hoped and expected that there were many
years of activity and usefulness before him.

It was ordained otherwise, and he died suddenly February
25, 1906.

D. B. Wainwright.

William Bramwell Powell.
1836-1904.

William Bramwell Powell was born at Castile, N. Y.,
on December 22, 1836. He was of English ancestry, being the
fifth child of Joseph and Mary Dean Powell who emigrated
from England to New York in 1830.

From his parents, who were persons of far more than ordinary
force of character and intelligence, he inherited many of the
qualities that distinguished him in life. Joseph Powell, his
father, had a strong will, deep earnestness, and indomitable
courage, while his mother, Mary Dean, with similar traits pos-
sessed also remarkable tact and practicality. Both were English
born, the mother well educated, and they were always leaders
in the social and educational life of every community where
they dwelt. Especially were they prominent in religious circles,
the father being a licensed exhorter in the Methodist Episcopal
Church. Both were intensely American in their love and ad-
miration of the civil institutions of the United States and both
were strenuously opposed to slavery, which was flourishing in
America when they arrived in 1830. For a time they remained
in New York City and then removed to western New York,
finally locating in the village of Castile, where, as before stated,
William Bramwell was born. Because of the slavery question
Joseph Powell left the Methodist Episcopal Church on the
organization of the Wesleyan Methodist Church and became a
regularly ordained preacher in the latter. It was in this atmo-
sphere of social, educational, political and religious fervor that
the future school superintendent grew up. When he was three
years old the family moved to Jackson, Ohio, and then, in 1846,
went on westward to South Grove, Walworth County, Wiscon-
sin, where a farm was purchased. They were in prosperous
circumstances, and the boy was active in the management of
affairs, early exhibiting his trait for doing things well.

228

WILLIAM BRAMWELL POWELL. 229

His early education was such as the country schools afforded
but his parents ever held before him the importance of achiev-
ing the highest education possible.

In 185 1 his family removed to Bonus Prairie, Boone County,
Illinois, where a larger farm had been purchased. About 1853
the Wesleyan College was established at Wheaton, Illinois, and
the family removed there in order to take advantage of the
opportunities afforded. The father became one of the trustees
and young Powell entered the preparatory classes. With inter-
vals of teaching he continued in the college till 1855, when he
entered the preparatory department of Illinois College at Jack-
sonville, Illinois. Here he continued a year, leaving to enter
Oberlin College, Ohio. In 1858 he returned to Wheaton Col-
lege and entered the Junior year. Leaving in 1859 ^^ ^^'
gage in his chosen profession of teaching, he was not gradu-
ated ; but in 1865 Lombard University, Illinois, conferred on
him the degree of A.M.

Mr. Powell's services as teacher and school superintendent
■extended over nearly half a century. He was principal of a
school in Sharon, Wisconsin, for some time between 1854 ^^^
1861 ; in 1861-2, principal of the Hennepin (Illinois) school;
1863 to 1870, Superintendent of the Peru (Illinois) schools ;
187 1-1885, Superintendent Aurora (Illinois) schools ; 1885-1900,
Superintendent of Washington Public (white) schools. In 1901
he visited the Hawaiian and Philippine Islands and Japan to
investigate the schools and text-book needs of these countries.

Mr. Powell was a member of the National Geographic
Society, Washington Academy of Sciences, National Academy
■of Political and Social Science, Anthropological Society of
Washington and the National Educational Association. He was
the author of a number of school books which have been exten-
sively used in the public schools of the country. The chief of
these are: " How to See, How to Talk and How to Write,"
published in 1880; and a ♦' History of the United States for
Beginners," published in 1900. He was also joint author of
the "Normal Series of Readers" published in 1887, and a
"Rational Grammar of the English Language," published in
1900.

230 WILLIAM BRAMWELL POWELL.

The following estimate of Mr. Powell's character and work
as a school superintendent is quoted from an article in a Chicago
journal of education {Intelligencer ^ April i, 1904.)

"During his forty years of service Mr. Powell labored un-
ceasingly and untiringly to improve the public school system.
In his chosen profession he felt the most profound devotion, the
greatest pride and pleasure. He was an advanced thinker, a
prophet, and as such was often in advance of his time in the
scope and grandeur of his ideals. His interests were centered
in the welfare of the children, believing that the proper educa-
tional training is that which best prepares for life, not merely
from the standpoint of earning a livelihood, but that education
which elevates above sordid, material views ; that which makes
the brain and heart capable of appreciating the good and beau-
tiful, susceptible to the allurements of the larger life.

" He felt it to be the child's birthright to have the best in edu-
cation that human effort can give. Influenced by these ideals,
he was a pioneer in methods of object teaching, of training by
seeing and doing. Believing that all education is based upon
experience, and that only through the child's own experience
can he be held to understand and appreciate the experiences of
others as found in books, he strove to give the child such expe-
riences as lead to the interpretation of the natural world and to
an understanding of the social whole. To provide for these ex-
periences he early introduced into his course the study of nature
and the study of institutional life. He collected libraries that
the children might broaden their text-book knowledge with a
knowledge of and a sympathy with life."

The following passage from the same article does some justice
to Mr. Powell's work in securing manual and industrial train-
ing in the public school system under his charge :

"A firm believer in the value of hand work, he was among
the first to prepare a way for and to establish manual training
and domestic science schools. Music, drawing and physical
training also became part of the regular course in his schools ;
all of this before most schools had advanced beyond the three
R's. In the Washington schools he strove as strenuously to
provide the best facilities in manual training, in cooking and in

WILLIAM BRAMWELL POWELL. 23 1

cutting and fitting as he did to provide the best of book instruc-
tion, thus giving to these new elements in education their proper
standing. To him the genius of American civilization demands
work — work of hand, heart and brain. He contended that
getting knowledge by rational methods gives to the child mental
and physical delight."

He was married in 1865 to Miss Minnie Paul, of Peru, Illi-
nois, who with two children. Miss Maud Powell, the violinist,
and Mr. William Paul Powell, of Mount Vernon, New York,
survives him. His death occurred after a short illness at Mount
Vernon, New York, on February 4, 1904.

Prepared by U. S. Bureau of Education.

Nicholas Senn.
I 844-1 908.

Nicholas Senn was born in Buchs, Canton of St. Gall,
Switzerland, October 31, 1844, ^"^ died, at his home in
Chicago, January 2, 1908.

He came to this country with his parents in 1852 and settled
at Ashford, Wis. He was graduated from the Fond du Lac
high school in 1864, after which he taught school himself and
began the study of medicine with Dr. E. Munk, of Fond du Lac.
He entered the Chicago Medical College in 1865 and graduated
in 1868. In 1869 he married Miss Aurelia S. Muehlhauser, of
La Crosse, and began the practice of medicine in Ashford. In
1874 ^^ moved to Milwaukee and became an attending physi-
cian of the Milwaukee hospital. Returning to Europe in 1877,
he studied at the University of Munich and received the medical
degree in 1878. He then resumed practice in Milwaukee until
1893, when he moved to Chicago where he remained until his
death.

During the comparatively brief period of thirty years — the
period of his active professional life, from the time of his
graduation in Munich until the date of his decease in 1908 —
few men in any profession have performed more productive and
excellent work, and certainly but few, if any, have received
more honors in recognition of their distinguished service to
humanity in the domain of medical science and practical sur-
gery, than Dr. Senn.

In every sphere of professional activity, alike in his chosen
field of operative surgery, as in the role of teacher and author ;
in his patriotic duties as a military surgeon ; and in works of
charity and benevolence, Dr. Senn " acted well his part."

His hospital work began as interne in the Cook County Hos-
pital of Chicago. Then he became a member of the staff of
the Milwaukee Hospital ; and later Surgeon-in-Chief to the St.

NICHOLAS SENN. 233

Joseph's Hospital and Presbyterian Hospital, Chicago; and
Surgeon to the Passavant and Polyclinic Hospital.

He was always deeply interested in military matters and
military surgery. He was made Surgeon-General of Wiscon-
sin in 1888 and retained the position until he left that State. In
1892 he was commissioned by Governor Altgeld Surgeon-
General of Illinois and retained this position until his death.

At the outbreak of the Spanish-American War he assumed
charge of the mobilization camp at Springfield and ably directed
the physical examination of recruits. He was commissioned
Lieutenant-Colonel and chief Surgeon, U. S. Volunteers, May
13, 1898, and was assigned to duty with the Sixth Army Corps.
He went with the expedition under command of Brigadier-
General Guy V. Henry, U. S. Volunteers, to Santiago, Cuba,
and was assigned to duty as chief surgeon of the operating staff
with troops in the field. He resigned September 6, and was
honorably discharged September 17. In general orders from
the Adjutant-General's office, dated February 13, 1900, Lieu-
tenant-Colonel Senn was commended for his surgical work
during the Cuban campaign and for making a scientific study
of typhoid fever among the troops.

His work as a teacher of medicine began in 1884 when he
was appointed Professor of the Principles and Practice of
Surgery in the College of Physicians and Surgeons, Chicago.
Four years later he became Professor of the Principles of Sur-
gery and Surgical Pathology in the Rush Medical College, and
later Professor of Military Surgery in the University of Chicago.
He was also Professor of Surgery in the Chicago Polyclinic.

His lectures — always without notes — were eloquent, dram-
atic and attractive.

His contributions to American medical literature comprised
something over three hundred titles, twelve of these being
printed volumes, varying in size, but all of them replete with
original matter. Many of them are used as text-books and
standard works of reference in most American medical schools,
and a good number have been translated into foreign languages.
Chief among them may be mentioned his text-books on : " The
Principles of Surgery," "Surgical Bacteriology," " Experi-

234 NICHOLAS SENN.

mental Surgery," " Pathology and Surgical Treatment of
Tumors," " Intestinal Surgery," "Tuberculosis of Bones and
Joints," and "Practical Surgery." He also wrote on the sur-
gery of the pancreas, stomach and gall-bladder; as well as
contributing several books of travel in which he gives his ob-
servation of diseases, physicians and hospitals in foreign
countries.

In 1897 Dr. Senn was elected President of the American
Medical Association. He was a member of the Philadelphia
College of Physicians, the American Surgical Society, the
American National Red Cross, the Norwegian Medical Society,
the Swedish Medical Society, and others.

Also an honorary member of the D. Hayes Agnew Surgical
Society, the Philadelphia Academy of Surgery, National Asso-
ciation of Railway Surgeons, Academy of Medicine of Mexico,
Glasgow Academy of Medicine, Manila Medical Society, the
Imperial-Royal Medical Society of Vienna, and the Royal
Medical Society of Budapest. He also received the Order of
Merit of the Japanese Society of the Red Cross by the sanction
of the Emperor of Japan.

While it was not the privilege of the writer to enjoy any per-
sonal intimacy with Dr. Senn, those who were more fortunate
in this respect speak of him as a truly great man ; " master of
his profession ; always ready to sacrifice his personal interest
and comfort for the service of his adopted country ; intensely
loyal in his friendships ; generous to a fault ; and too honest to
harbor suspicions." " His greatest glory was in his extraordi-
nary capacity for work, which he held as a duty, and that work
entirely for the benefit of his fellow-men."

Among the substantial evidences of his generosity may be
mentioned his endowment of two rooms in the St. Joseph's Hos-
pital, Chicago ; his donation to Rush Medical College of the
Senn Clinical Building, and his presentation to the Crerar
Library of a valuable collection of books, including the entire
library of Dubois-Raymond and that of the late Dr. William
Baum, Professor of Surgery in the University of Gottingen.

Taking him for all in all, it may truly be said : the world is

better for his having lived.

A. F. A. King.

Samuel Edwin Solly.
1845- 1906.

Dr. Solly was born in London, England, May 5, 1845,
and died at Colorado Springs, Colorado, November 19, 1906.
He received his early education in the best schools of his native
land, and, following the footsteps of his distinguished father,
he took up a medical career. Owing to a physical breakdown
in his early career, his attention was naturally directed along
climatological lines. Through his father, he was brought into
intimate association with many of the distinguished practitioners
of London. In this way, he became acquainted with the late
Sir Morell Mackenzie, from whom he acquired his insight into
laryngology and rhinology. In his thirtieth year he found it
impracticable, from a physical point of view, to longer reside in
London, so he turned his face toward the setting sun. We
next find him in Manitou, Colorado, where he resided for some
years, being the inspiration of the English colony at this place.
While at Manitou he made an investigation of the mineral
springs of that place. Shortly after the establishment of the
city of Colorado Springs, he removed to that inviting and
thriving town. His great life work was done at Colorado
Springs. He never tired of singing its praises or writing of its
glories. Several years ago he was very much interested in a
movement started by the civic body of Colorado Springs in
sending out the secretary of that organization on a tour of the
principal cities of the country to proclaim the attractions of
Colorado Springs as a sanitarium. Though much interested in
all projects of a civic character that redounded to the benefit of
his home city, he was much more interested in those projects
which had for their purpose the recognition of the fact that it
was the world's great sanitarium for those afflicted with tuber-
cular affection. The Cragmoor Sanitarium was his ruling
passion during the last few years of his life. This institution

Proc. Wash. Acad. Sci., December, 190S. 235

236 SAMUEL EDWIN SOLLY.

was modeled on a most comprehensive system, as only the
master hand of Solly could have formulated.

Dr. Solly was a man of remarkable attainments — a repre-
sentative of the highest type of the intellectual Englishman ; in
medicine, he was a scholar with an investigative turn of mind ;
an executive of remarkable ability ; a public-spirited citizen of
the highest grade ; and a writer upon climatological subjects
whose fame and name are co-extensive with the realms of
modern medicine. His personality was the most delightful that
the writer of this article has ever known. He possessed a quiet
and dignified bearing, a modest temperament, a genial dispo-
sition, kindness of thought and action, a courtesy of manner
that never was at fault, a ready wit that never lent itself to
sarcasm, a perfect host, a most welcome guest, strong intel-
lectually, with high moral ideals, and the courage of his con-
victions — such was the character of the lovable man of whom
we write and who won his way into the hearts of all who came
within his presence.

Charles W. Richardson.

Ainsworth Rand Spofford.
1825-1908.

Ainsworth Rand Spofford at the time of his death, which
occurred at Holderness, New Hampshire, August 11, 1908,
was Chief Assistant Librarian of Congress. He was born at
Gilmanton, New Hampshire, September 12, 1825, and was the
son of the Reverend L. A. and Grata (Rand) Spofford.

He was prepared for college by private tutors, but his health
failing, he went west at this time, 1844, and located in Cincin-
nati. His fondness for books soon decided his career; for he
soon entered upon the duties of a book seller and publisher and
spent his leisure moments in the study of literature and modern
languages. In 1850, he was one of the founders of the Liter-
ary Club, of Cincinnati, a vigorous and intellectual organiza-
tion. In 1852 he married Sarah P. Partridge, who died in
1892.

His coming to Washington, D. C, was in 1861, when he
was appointed by President Lincoln Assistant Librarian to
Congress. For several years previous to this, he was Assistant
Editor, 1859-1861, of the Cincinnati Daily Co7nmercial. His
great fitness for the position of librarian was soon recognized
and in 1864 he was made Librarian-in-Chief, holding that
position until 1897, when he was relieved of the executive
burden and made Chief Assistant Librarian. He saw the
Library of Congress grow from a collection of 70,000 volumes
to that of 2,500,000 volumes. He was the first to suggest the
need of a separate building for the Library and when Congress
was slow to act, he kept the matter before it not only in his
formal reports, but by a systematic disorder in the overcrowded
old quarters — the library, that filled every space with books,
boxes, maps, bundles, etc., leaving thus but little room for the
visiting Congressmen.

As a librarian he was widely known for his comprehensive
knowledge of books and their contents and was to public men

237

238 AINSWORTH RAND SPOFFORD.

and students a veritable catalogue. It was a tradition among
the guides that Mr. Spofford read every book that came to the
library and that he knew the position of every book. A ques-
tion or two addressed to him would save hours of laborious re-
search. The writer, a year ago, had occasion to look up the
subject of medicine versus superstition and going to Mr. Spof-
ford asked if he would pick out several of the best books on the
subject. Mr. Spofford, taking a pencil, wrote offhand from
memory the title, the author, in some cases the chapter, of eleven
books of reference on the desired subject. When General Lew
Wallace was writing "Ben Hur," he appealed to Mr. Spofford
to help him in proper material for the local color and literary
atmosphere. Mr. Spofford proposed and exhibited book after
book from the Congressional Library without satisfying General
Wallace, and finally told the General that he would find what he
wanted in the Harvard Library in such a room and such a shelf,
situated " sixth from the south end."

He contributed to many newspapers, magazines and encyclo-
pedias. He was editor of Catalogues of the Library of Con-
gress, and of the Annual American Almanac from 1878 to 1889 ;
edited with others. Library of Choice Literature (10 vols.). Li-
brary of Historic Characters and Famous Events (10 vols.),
Library of Wit and Humor (5 vols.). He was the author of
Practical Manual of Parliamentary Rules, 1884 ; and A Book for
all Readers, etc.

Many of his writings dealt with the early history of our coun-
try and especially with the history of the Federal city. He was
a firm believer in the " instinctive genius " of the French engin-
eer, Major L'Enfant, who planned the city of Washington, and
in endorsing a bill, February 11, 1905, presented to Congress,
speaks of him as "the undisputed author of the grand scheme
which was ultimately adopted."

He was a charter member and Vice-President of the Colum-
bia Historical Society from its organization in 1894 to the date
of his death. Among some of the papers he contributed were :

" The Methods and Aims of Historical Inquiry," May 7,
1894.

"Life and Labors of Peter Force, Mayor of Washington,''
June 8, 1897.

AINSWORTH RAND SPOFFORD. 239

" Washington City in Literature," February lo, 1902.
" The Lyric Element in American History," December 14,
1903.

"The Eloquence of Congress: Historic Notes," November

13' 1905-

" Virginia, 300 Years Ago," April 8, 1907.

Mr. Spofford was a man of kindly and charitable disposition
and of a temperament which once known, was as peaceful and
attractive as the cooling rivulet on a summer day. Absolutely
unconscious of his distinction and never courting notoriety, he
came and mingled with us all, giving help, advice and encour-
agement to those who would drink of the Pierian sprmg.

James Dudley Morgan.

R. Stansbury Sutton.
1841-1906.

Dr. Sutton was born in Indiana, Pennsylvania, in 1841 and
died suddenly at Pittsburg, Pa., April 21, 1906. He studied
medicine in Philadelphia and graduated with honor from the
University of Pennsylvania in the class of 1865. He soon
settled in Pittsburg where he practised his profession with great
success up to the day of his death. He had his wish, to die
with his harness on, and not to linger long with a distressing
illness, a burden to himself and an anxiety to his many friends.

He was happily married and was a devoted father and a most
useful neighbor and friend. Though a successful general sur-
geon he was among the first of his confreres to adopt the specialty
of gynecology and abdominal surgery. He was actually a
pioneer in his work and fitted himself more particularly for its
delicate and responsible duties by special studies and training in
our large cities and by instruction in the clinics and hospitals
abroad, to which he frequently returned for new ideas and a
study of their especial technique.

In 1905, Dr. Sutton was appointed a delegate to the Madrid
International Congress of Gynecology.

In order more successfully to conduct the special work to
which he had dedicated his talents, he established, in 1883, one
of the first private hospitals in the country, and devoted it to
gynecology and abdominal surgery. He named it the Terrace
Bank Hospital for Women, and conducted it for twenty years.
During the recent Spanish war his ardent patriotism led him to
offer his services to the government and he was appointed Chief
Surgeon to the Second Brigade, Third Division of the First
Army Corps.

Dr. Sutton was a frequent contributor to the medical journals
and societies and was the author of a book on abdominal sur-
gery, which was chiefly a history of his own special work. He
was twice elected Vice-President of the American Gynecological

240

R. STANSBURY SUTTON. 24I

Society. He served as President of the American Academy of
Medicine, and also of the Mississippi Valley Medical Associa-
tion, and the Pittsburg Obstetrical and Gynecological Society.
In 1884 he was elected Chairman of the Obstetrical and Gyne-
cological Section of the American Medical Association. He
was also a member of the British Medical and Gynecological

Associations.

Joseph Taber Johnson.

Robert Bowne Warder.
I 848- I 905.

Robert Bowne Warder died at his home in Washington,
July 23, 1905, after an illness extending over nearly a year.

Professor Warder was born in Cincinnati, O., March 28,
1848, and spent his early life in the country home at " Aston,'*
North Bend, Ohio. His character was formed under the influ-
ence of the Society of Friends and this faith remained the
dominant feature of his life. From childhood he showed the
effect of his parents' training and example, in a broad and
catholic view of the ethics of life, and in a love of truth and
scientific investigation. This devotion to truth was an especial
characteristic and governed his life and actions throughout.

He was graduated from a Friends' institution, Earlham Col-
lege, at Richmond, Ind., in 1866, and afterwards spent some
time at the Illinois State University, at Champaign, where he
was instructor in chemistry and natural philosophy. This work
of teaching seemed to show Professor Warder his natural bent,
and his energy was thenceforth devoted to studying the broad
principles underlying all natural science. He spent some years
in traveling, chiefly in the western half of the United States, in
connection with the different State geological surveys. In 1873
he went to Harvard, where he was graduated as B.S. in
chemistry in 1874.

After graduating at Harvard he spent a year traveling in
Germany, studying at Giessen under Heinrich Will, and at
Berlin under Hofmann. His attention was, however, especially
devoted to methods of teaching chemistry in the German uni-
versities, and the application of theoretical chemistry to the
practical sciences. His chief aim was to fit himself in the
broadest sense for his work of teaching. This was his main
desire throughout life, to help others, and he never faltered.
On returning to this country he was associated with Prof. F.
W. Clarke at the University of Cincinnati from 1875 to 1879 ^^

242

ROBERT BOWNE WARDER. 243.

professor of chemistry and physics. Professor Warder early
saw the close relation between these then distinct branches of
natural science, and his papers on *'The Speed of Saponifica-
tion of Ethyl Acetate " and " Evidence of Atomic Motion within
Liquid Molecules " were pioneer investigations in the field of
physical chemistry of to-day.

He was engaged in this line of research from 1879 ^^ 1883
when he accepted the chair of chemistry at Purdue University,
where he remained until 1887. This position carried with it
the duties of State Chemist, work of a commercial character
rather foreign to his natural tastes, but to which he gave the
same painstaking devotion that characterized all his work.
Even these routine analyses were made to pay tribute to physical
chemistr}', as is shown by papers on " Influence of Time in
Fertilizer Analysis," " Speed of Dissociation of Brass," etc.

In 1884 he married Gulielma M. Borland, who also belonged
to the Society of Friends, and like himself was interested in
evangelical work. Their life together was one of perfect
harmony.

It was probably about this time that Professor Warder felt
more keenly than ever the call to help others in another field than
chemistry. His philanthropic and evangelical work had always
been foremost in his mind and labors, and in 1887 he accepted
the professorship of chemistry at Howard University in Wash-
ington. Here he labored until he died, teaching chemistry and
physics, but above all setting an example and teaching the
principles of a Christian life with an unselfish devotion.

In spite of lack of facilities, his work at this period on " Dy-
namical Theory of Albumenoid Ammonia," " RecentTheories of
Geometric Isomerism," " Cross Fertilization of the Sciences,"
and "The Major Premise in Physical Chemistry" showed his
natural inclination to this phase of chemistry.

Professor Warder's later papers were chiefly devoted to ap-
plying the laws of mass action to and showing the speed of
chemical reactions for the analytical data obtained by other
investigators.

He was essentially a critic and his devotion to truth caused
him to scrutinize the investigations of others with the same

244 ROBERT BOWNE WARDER.

zealous care with which he looked for flaws in his own work.
This high standard, coupled with an unusual modesty, often
caused a hesitation which sometimes obscured his really pro-
found knowledge.

No one went to Professor Warder for aid and was turned
away empty-handed. What he had was given freely, and he
seemed to feel that no labor was too great in his fundamental
desire to help others.

S. S. VOORHEES.

INDEX.

Note. — New names in black-face type, synonyms in Italics.

abdominalis, Coniontis 83
acerba, Coniontis 139
acomana, Sphaeriontis 76
acutangulus, Eusattus 72
acutus Eusattus 74
affinis, Cambarus 172

Coniontis 123

patruelis, Coniontis 123
agnatus, Eusattus 70
agrestis, Coniontis 131
alutacea, Coniontis 139
ampliatus, Coniontellus 144
amplicoUis, Ccelus 157
ancilla, Coniontis 91
Andrews, E. A. 167
Aneura multifida major, Fungus parasite

of II
angustula, Ccelotaxis 149
anita, Coniontis 139
anxia, Coniontis 123
arenarius, Ccelus 156
argutus, Coniontellus 145
arida, Coniontis 138
Ashmead, William Harris 187
Atherton, George W. 190
atronitens, Coniontis no
Atwater, Wilbur Olin 194
Audax, Coniontis 121

blaisdelli, Coniontis 98
Brachyontis 82, 141
Branchini 53, 54, 162
brevipennis, Discodemus 61
breviuscula, Coniontis. 133
Burnett, Swan Moses 199

California Hepaticse. Physiology and

Morphology of i
Callida, Coniontis 128
Cambaroides 172
Cambarus afiBnis 172

subgenera of 173
canonica, Coniontis 114
Carroll, Dr. James 204
carsonica, Coniontis nevadensis 95
Casey, Thomas L. 51
catalinse, Coniontis 88
ciliata, Sphaeriontis 77
ciliatus, Ccelus 151, 154

longulus, Ccelus 154
clementinus, Coniontides 80
Ccelini 53, 150, 162
Ccelomorpha 151, 159

maritima 151, 160

pallens 160
Ccelotaxis 57, 147

angustula 149

densa 149

frontalis 149

muricata 148

punctulata 148
Ccelus 151, 152

amplicoUis 157

arenarius 156

ciliatus 151, 154
longulus 154

curtulus 155

debilis 155

globosus 153

grossus 153

latus 158

obscurus 156

pacificus 158

remotus 158

saginatus 154

scolopax 157

solidus 153

sternalis 156
compositus, Eusattus 71
conferta, Coniontis 129
congener, Eusattus 71
congesta, Coniontis 131
conicicollis, Coniontis 102

Coniontellus ampliatus 144

argutus 145

hystrix 142

inflatus 143

longipentis 143

micans 145

obesus 144

subglaber 144
Coniontides 57, 78

clemontinus 80

finitimus 79

insularis 79

latus 78
Coniontinae 52

A Revision of the Tenebrionid Sub-
family 51
Coniontini 53, 55, 162
Coniontes 55
Coniontis 57, 80, 81

abdominalis 83

acerba 139

affinis 123

patruelis 123

agrestis 131

alutacea 139

ancilla 91

anita 139

anxia 123

arida 138

atronitens no

audax 121

blaisdelli 98

breviuscula 133

245

246

INDEX

callida 128
canonlca 114
catalinse 88
conferta 129
congesta 131
conicicollis 102
convergens 122
corvina 140
costulata 89
cuneata iii
cylindrica 100
degener 93
elliptica 88
elongata 99

llmatula 99
eschscholtzi 129
exigua 106
expansa 120
extricata 124
farallonica 132
flliola 115
franciscana 120
genitiva loi

(Brachyontis) globulina 141
histrio 91
inconspicua 108
inepta 137
inflexula 107
gravis 85
innocua 90
inornata 130
integer 87
Isevigata 88
lanuginosa 117
lassenica 95
lata 57
levettei 87
longicollis loi

lucidula 103

luctuosa 89

marginata 125

microsticta 107

minuta 126

montana 112

nemoralis 125

nevadensis 95
carsonica 95

obesa 57

oblita 137

oblonga 92

obsidiana 100

obsolescens 92

obtusa 116

opaca 94

opacicoUis loi

oregona 124

ovalis 140

pagana 130

pallidicornis 93

parallels 113

parilis 135

parva 126

parviceps 115

paupercula 106

pectoralis 86

perpolita 127

perspicua 114

picescens 87

picipes 107

proba 105

protensa 104

pubifera 118

pudica 127

punctata 135

puncticollis 105

punctipes 96

regularis 134

robusta 89

rotundicollis 97

rugosa 85

sanfordi 90

sculptipennis 134

setosa 116

sbastanica 128

sparsa no

strenua 84

siibpubescens 113

suturalis 121

symmetrica 122

tenebrosa 86

(Crypticomorpha) tenuis 141

tboracica 104

timida 102

tristis 84

truncata 120

uteana 136

vancouveri 136

verna 94

viatica 57, 103

weidti 138

wickhami 117
Conipitius 56, 57
Conisattus 57, 146

rectus 146
convergens, Coniontis 122
convexus, Eusattus 72
coquilletti, Eusattus 69
corrosus, Discodemus 61
corvina, Coniontis 140
costatus, Megasattus 64
costulata, Coniontis 89
Crypticomorpha 81, 140
cuneata, Coniontis in
curtulus, Coelus 155
cylindrica, Coniontis 100

debilis, Coelus 155
degener, Coniontis 93
densa, Coelotaxis 149
depressulus, Discodemus 62
de Schweinitz, Emil Alexander 208
difficilis, Eusattus 56, 70
dilatata, Spbseriontis 76
Discodemus 56, 59

brevipennis 61

corrosus 61

depressulus 62

elongatulus 61

knausi 62

reticulatus 61

subsericeus 62
dubius, Eusattus 66

elliptica, Coniontis 88
elongata, Coniontis 99

limatula, Coniontis,99
elongatulus, Discodemus 61
erosus, Ejisatlus 56

erosus, Megasattus 63
eschscholtzi, Coniontis 129

i

INDEX

247

Eusatti 55
Eusattodes 56, 64

laevis 64
Eusattus 56, 64

acutangulus 72

acutus 74

agnatus 70

compositus 71

congener 71

convexus 72

coquilletti 69

difficilis 56, 70

dubius 66

erosus 56

explanatus 68

Icpvis 56

lobatus 68

viiiricalus 56

nanus 66

oblongulus 67

peropacus 74

politus 69

prodiictus 67

robiisttis 56

rotundas 72

secutus 65

subnitens 73

turgidus 73

vicinus 68
expansa, Coniontis 120
exigua, Coniontis 106
explanata, Nyctelioma 163
explanatus, Eusattus 68
extricata, Coniontis 124

farallonica, Coniontis 132

Fegatella conica, Development of Fruit-
ing Organs of 19

filiola, Coniontis 115

finitimus, Coniontides 79

Fossombronia longiseta, Fungus parasite
of 4

franciscana, Coniontis 120

frontalis, Coelotaxis 149

Fruiting organs of Fegatella conica, de-
velopment of 19

Fungus parasite of Aneura multifida
major 11

Fungus parasite of Fossombronia longi-
seta 4

genitiva, Coniontis loi

Germination of spores, Studies in 35

Gilman, Daniel Coit 211

globosus, Coelus 153

globulina, Coniontis (Brachyontis) 141

gravis, Coniontis 85

grossus, Coelus 153

Harper, William Rainey 215

Hepaticae, Parasitism and Saprophytism

in 2
Physiology and Morphology of Some

California i
Mstrio, Coniontis 91
Humphrey, Harry B, i
liystrix, Coniontellus 142

inconspicua, Coniontis 108
inepta, Coniontis 137
inflatus, Coniontellus 143

Inflexula, Coniontis 107
innocua, Coniontis 99
inornata, Coniontis 130
insularis, Coniontides 79
integer, Coniontis 87

knausi, Discodemus 62

laevigata, Coniontis 88
laevis, Eusattodes 64

Eusattus 56
Langley, Samuel Pierpont 219
lanuginosa, Coniontis 117
lassenica, Coniontis 95
lata, Coniontis 57
latus, Coelus 158

Coniontides 78
levettei, Coniontis 87
Ivindenkohl, Adolph 222
Liverworts and dry season 23
Liverwort Spores, Germination of 35
lobatus, Eusattus 68
longicollis, Coniontis loi
longipennis, Coniontellus 143
lucidula, Coniontis 103
luctuosa, Coniontis 89

Marginata, Coniontis 125
Marindin, Henri Louis Francois 224
Maritima, Coelomorpha 151, 160
Megasattus 56, 62

costatus 64

erosus 63
Micans, Coniontellus 145
microsticta, Coniontis 107
minuta, Coniontis 126
montana, Coniontis 112
muricata, Coelotaxis 148

Spbseriontis 76
niuricatus, Eiisattus 56

nanus, Eusattus 66
nemoralis, Coniontis 125
Nesostes 56, 58

robustus 59

robustus postremus 59
nevadensis, Coniontis 95

carsonica, Coniontis 95
nodosa, Zophosis 163
Nyctelia 163
Nycteliini 161
Nyctelioma 163

explanata 163

obesa, Coniontis 57
obesus, Coniontellus 144
oblita, Coniontis 137
oblonga, Coniontis 92
oblongulus Eusattus 67
obscurus, Coelus 156
obsidiana, Coniontis 100
obsolescens, Coniontis 92
obtusa, Coniontis 116
Ogden, Herbert Gouvemeur 226
opaca, Coniontis 94
opacicoUis, Coniontis loi
oregona, Coniontis 124
ovalis, Coniontis 140

pacificus, Coelus 158
pagana, Coniontis 130

248

INDEX

pallens, Coelomorpha 160

pallidicornis, Coniontis 93

parallela, Coniontis 113

parilis, Coniontis 135 '

parva, Coniontis 126

parviceps, Coniontis 115

paupercula, Coniontis 106

pectoralis, Coniontis 86

perpolita, Coniontis 127

peropacus, EJusattus 74

perspicua, Coniontis 114

picescens, Coniontis 87

picipes, Coniontis 107

Physiology and Morphology of Some

CaHfornia Hepaticae i
politus, Eusattus 69
postremus, Nesostes robustus 59
Powell, William Bramwell 228
Praocini 53, 162
Praocis 53

protoa, Coniontis 105
productus, Eusattus 67
protensa, Coniontis 104
Pseudocoelus 152
puberula, Sphasriontis 77
pubifera, Coniontis 118
pudica, Coniontis 127
punctata, Coniontis 135
puncticoUis, Coniontis 105
punctipes, Coniontis 96
punctulata, Coelotaxis 148

rectus, Conisattus 146
regularis, Coniontis 134
remotus, Coelus 158
reticulata, Zophosis 56, 61
reticulatus, Discodemus 61
robusta, Coniontis 89
robustus, Eusatttis 56

Nesostes 59
rotundicoUis, Coniontis 97
rotundas, Eusattus 72
rugosa, Coniontis 85

saginatus, Coelus 154
sanfordi, Coniontis 90
scolopax, Coelus 157
sculptipennis, Coniontis 134
secutus, Eusattus 65
Senn, Dr. Nicholas 232
setosa, Coniontis 116

shastanica, Coniontis 128
solidus, Ccelus 153

Solly, Samuel Edwin 235 }

sparsa, Coniontis no
Sperm Receptacle in the Crayfishes,
Cambarus cubensis and C. paar-
doxis 167
Sphaeriontis 56, 75

acomana 76

ciliata 77

dilatata 76

muricata 76

puberula 77
Spofford, Ainsworth Rand 237
sternalis, Coelus 156
strenua, Coniontis 84
subglaber, Coniontellus 144
subnitens, Eusattus 73
subpubescens, Coniontis 113
subsericeus, Discodemus 62
Sutton, Dr. R. Stansbury 240
suturalis, Coniontis 121
symmetrica, Coniontis 122

Tenebrionid Subfamily Coniontinse, A

Revision of 51
tenebrosa, Coniontis 86
TentyriiuK 51

tenuis, Coniontis (Crypticomorpha) 141
Thin-walled spores, Duration of vitality

in 44
thoracica, Coniontis 104
timida, Coniontis 102
tristis, Coniontis 84
truncata, Coniontis 120
turgidus, Eusattus 73

uteana, Coniontis 136

vancouveri, Coniontis 136
verna, Coniontis 94
viatica, Coniontis 57, 103
vicinus, Eusattus 68

Warder, Robert Bowne 242
weidti, Coniontis 138
■wickhami, Coniontis 117

Zopherinse 52
Zophosis nodosa 163
reticulata 56, 61

4

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