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Psyche 5:287-294, 1888.

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PSYCHE.
THE WORK OF A DECADE UPON FOSSIL INSECTS, 1880-1889. BY SAMUEL HLJBBARD SCUDDER, CAMBRIDGE, MASS. [Annual address of the retiring president of the Cambridge Enton~ological Club, 10 Jan. 1890.1 Some years ago I published an an-
notated and tolerably complete list of
papers on fossil insects. It contained
nearly three times as many titles as
were referred to by Hagen in his entomo- logical bibliography nearly twenty years previously, but, as the multiplication
of periodical literature had brought in
a train of minor papers, largely abstracts and compilations, I remarked that the
far greater extent of my list was no
proof of an increased recent interest in this field of research, but thought it
doubtful whether in the intervening
period there had been as much activity
us when the works of Heer were open-
ing the wealth of material at hand.
So marked a change has now come
about in this respect that I venture this evening to invite your attention to a
review of the advance that has been
made during the past ten years in this
previously neglected field. In doing
this I do not by any means propose to
cite every paper that has been published, but only to call your attention to the
more important or interesting, from
whatever cause, and thus endeavor to
picture our progress as vividly as possi- ble. Indeed, the mere list of authors '
would be wearisome, for one could
make a catalogue of the writings of the
last ten years considerably longer than
the entire list given by Hagen in 1863.
To be precise, I can cite 94 authors
and about 225 papers published in
this decade, against 78 authors and about 140 papers quoted by Hagen. Or to
picture it in another way, about one
third of a complete catalogue of papers
on fossil insects would belong to the
decade just closed. Nor is the bulk of
this literature its only value ; it is quite as remarkable for its quality, for by far the most important of the discoveries
yet made in fossil insects are embodied
in the researches of the last ten years, and there is no reason to suppose that
we have reached their conclusion.
Note, first, the relatively great num-
ber of striking discoveries that have
been made within this period. The
discovery and careful study of Silurian
scorpions in several different parts of




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283 ~~Ck?.!?. January 1890.
the world,-in Swsden by Thoi-ell and
Lindstrhm, in Scotland by Peach and
Hunter, and in New York by Whitfield,
all brought out at very nearly the same
time, are unprecedented in the annals
of this division of science. These were
followed almost immediately by Brongni-
art's surprising discovery of one of the hexapods, PalaeobZatt+ in the Silur-
ian of France, still the only known true insect in this ancient deposit. Coming
down a stage later we have the i-emark-
able Devonian insect-fauna of New
Brunswick, about the nature of which
there has been so much dispute, first
announced, it is true, before our period, but only fully published with figures of the species in 188~ ; a single addition
or two has recently been made to them
by Matthew. With them must be
classed the Devonian myriopods, the
earliest known members of that group,
fully elaborated by Peach. In the car-
boniferous period we have the striking
wealth of forms from Mazon Creek and
other deposits in our country which T
have described at various times, includ- ing so extraordinary a number of blat-
tarians that I have ventured to call this period, so far as its insect-fauna is
concerned, '&the age of cockroaches."
These discoveries. largely due in this
country to the activity and zeal of Mr.
Lacoe, have been even more than pal-al-
leled by the unexampled wealth rightly
claimed for Commentry in France by
Brongniart, who as yet has published
hardly more than an outline sketch to
whet the appetite of the zealot. At
this place are found, ;is Mr. Prong-
niart informs me in a recent letter, a
considerable number of types already
signalized in America, which indeed
we had a right to anticipate by the
comparisons that had been made between
the forms already published from other
localities in the two countries, new
discoveries on one continent having
repeatedly been followed sooner or later by very similar finds on the other. The
abundance of cockroaches in both
countries is fully sustained at Commen-
try, which has yielded the vast number of nearly six hundred specimens, or many
more than are known from all other
carboniferons localities in the world
taken together. Still another striking
discovery in the carboniferous rocks is
the recent finding in Silesia of coleoptesa, the first time that these have been
signalized at this early epoch, but their description is yet to come.
These are the principal larger discov-
eries in the paleozoic series, but they
have been accompanied by the publica-
tion of many striking forms which
indicate the ancestral types of living
insects, or by the better elucidation of types already known but whose signifi-
cance had not been understood. To
specify some of these we may mention
PaZaeocanz$a and Acanthe~estes
among the myriapods, the former with
the curious and highly developed struc-
ture of the spinous hairs, the latter with its possession of segmental organs or
branchial supports as well as stigmata,
indicating a probable amphibious habit ; Anthraconiartus, Kreischeyia, and
Geralinwa, the two former examples




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January 1890.1 PSYCHE. 289
of new extinct family types of arachnids, the last the first instance of the discovery of the /edi/aZ#z' earlier than the terti- aries, and found at brief intervals on two continents ; other than this last of ~ukta's striking discoveries in the Bohemian
coal field might well be cited ; the gi- gantic ephemerid, Palififenz'a, of Bohe- mia ; Dasylefltus, an extraordinary
form of thysanura, a group not previous- ly known earlier than the tertiaries ;
CorydaZoide.~, like the preceding, one
of Brongniart's discoveries at Commen-
try, remarkable for the extensive display of branchiae on the sides of the abdomen ; Petrablattina subtilis of Kliver (Stre-
phocladus) with its strange neuration ;
M i a of England with its remarkable
coloration ; the gigantic Titam@asma,
also from Coinmentry ; the nymph of
Etoblattina Woodward has published
from England, showing the same mode
of development among the ancient as
the modern cockroaches; and, finally,
Phthanocoris, the only hemipteroid
type yet found in our own paleozoic
rocks.
All these memoranda relate to the
insects of the older formations only, but the statements regarding them in no
proper way indicate the immense strides
we have made in our knowledge of the
earlier types. The decade has been
marked not only by extensive and strik-
ing additions to known types, far more
than doubling the number that had been
previously published ; it has witnessed
also the advent of many original workers previously wholly unknown in this field, such as Beecher, Deichmuller? Karsch,
Kliver, ~usta, Matthew, Peach, Sterzel,
Thorell, and Whitfield ; but it has also seen the beginning of a new epoch in
the study of the earlier types, in that for the first time the subjects have been
treated in much more than a scattered
way, by fuller discussions of the syste- ixatic status of the insects described, by attempts to systematize our knowledge,
and by the treatment in single groups of insects from various or from all deposits, and not alone in the simple discussion
of collections from a given deposit.
Let us hope that the constantly increas- ing material and our larger knowledge
may permit in a new decade a further
correlation, by the comparative study of insects of different horizons, especially in the carboniferous age.
Previous to the last decade there
had been scarcely a single attempt at the systematic study of all the older insects, or even of any of the minor groups
found in the paleozoic rocks. Hagen,
indeed, had treated briefly of the few
termitina known over thirty years ago ;
Heer had attempted a grouping of the
cockroaches ; and Goldenberg had sum-
marized our knowledge of all by an
attempted classification ; but besides
these I do not recall n single instance
where any serious attempt had been
made to collate in a broad way our
knowledge of paleozoic insects as a
whole or in any of the parts.
Only be-
cause it has so happened that the present speaker has been perhaps the most active worker in this narrow field during the
last decade, is he obliged here to
mention mainly his own work, since




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290 PSYCHE. [January 1390.
-
it has fallen to his lot, in however im- perfect a way, to attempt a more or less monographic treatment of the extinct
type of archi$olypo&, for instance,
comprising most of the paleozoic myri-
opoda ; of the paleozoic arachnida as a
whole, in which he had been preceded
by this decade by Karsch, working on
much slenderer material and therefore
at much smaller advantage ; also on the
paleozoic cockroaches, and on the
species of Mylac* a genus of cock-
roaches known from several American
deposits ; and on the genera allied to
Dictyoneum, regarded as ancient types
of $hasnzida. Reference should here
also be made to Peach's careful work
on the carboniferous arachnida of
Scotland. In my memoir on the cock-
roaches, embracing the discussion of
fifty-eight species referred to eleven
genera, it was claimed that their differ- ences from modern types were so fun-
damental as to warrant their separation
from all subsequent and from living
cockroaches as a distinct and equivalent group, called palaeoblattariae, and
that they could be further separated into two divisions, called respectively my-
lacridae and blattinariae, of which the
former was confined to the New World.
Brauer has since questioned the value
of the palaeoblattariae as a group, and
Brongniart has recently stated that in
the enormous crowd of cockroaches
found at Corn in entry, the mylacridae
are as numerous as the hIat/inariae,
which probably means that the fauna of
Commentry is older than that of the
other carboniferous deposits of Europe
and synchronous or nearly so with most
of the cockroach-yielding deposits of
America.
Both Brongniart and myself have
also attempted new classifications of the paleozoic hexapods as a whole, which
differ considerably in character, but
which cannot yet fairly be compared ;
first because mine discusses nearly all
the known types, but includes hardly
any of those found at Commentry, then
almost wholly unknown, while Brong-
niart, writing later, confines himself
almost entirely to those of Commentry,
with only an occasional allusion to pre- viously described types ; but principally because Brongniart's work is, so far, the merest sketch with hardly any structural details, a forerunner of what he will
soon publish in extenso concerning this
wonderful fauna, while mine contains
full structural details as a basis for dis- cussion and generalization. In it I
have endeavored to point out that the
existing orders of insects were not dif- ferentiated in paleozoic times except in a feeble way, prophetic as it were of the future, so that the Palaeodictyoptera, as, after Dohrn and Goldenberg. but with
an extension of their usage, I had classed for the first time all known paleozoic
insects, could only be separated into neu- ropteroid, orthopteroid and hemipteroid
groups. These views, which I urged
also in a special paper showing the de-
velopment of the insect-type in time,
have been so strenuously opposed by
Brauer and others, that their further
discussion can hardly be profitable ex-
cept for those who have an unfortunate




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January 1Sy.3
taste for polemics, at least until the
fauna of Commentry, which will cer-
tainly double the field of observation,
gives us a fairer basis for judgment.
Meanwhile it may be said that Brongni-
art in his sketch hints by many of his
terms that he has found
the same diffi-
culties as those which faced mej and has been forced to admit a synthesis of
structure in at least some of the older
types, which indeed the very laws of
evolution would render probable.
At the beginning of this decade our
knowledge of mesozoic insects was very
limited ; it was almost entirely confined to the researches of Germar, Giebel,
Hagen and Weyenbergh on the Jura of
Eichstatt and Solenhofen ; to Heer's
account of the Liassic insects of Aargau ; and to Brodie's and Westwood's publi-
cations on the secondary insects of Eng- land. The horizon has been somewhat
extended of late years by the thorough
discussion of the Bavarian insects by
Deichmuller and by Oppenheim ; by
the careful exploitation of a new locality for Liassic insects at Dobbertin, Ger-
many, by F. E. Geinitz ; by the con-
siderable number of new generic and
specific types of cockroaches from the
secondary rocks of England described
by myself; by the repeated, though not
extensive, discoveries of Fritsch in Bo- hemia, adding interesting material for
our very meagre knowledge of creta-
ceous insects ; and by the discovery at
Fairplay, Col., of a collection of triassic cockroaches of special interest and im-
portance.
Among noteworthy contributions to
our knowledge of the insects of this
epoch may be mentioned Oppenheim's
study of the group he called rhipido-
h a w which he regarded as a distinct
order and an ancestral type of lepi-
doptera. The discussion of the structure of these insects, especially by Oppen-
heim and Deichmuller, has made clear
many points regarding the Solenhofen
insects which have always been obscure,
and brought about the agreement that
the ~~~~~~~~~hub& must be regarded as
hymenoptera and in no sense prede-
cessors of lepidoptera. Geinitz in his
study of the Liassic fauna of Dobbertin
has been able to extend considerably our knowledge of the structure of that pre-
vailing mesozoic type, Orthoph?ebia,
known entirely by its wings, and which
he regards as phryganideous. In our
own country, the triassic cockroach-
fauna of Fairplay, just referred to.
shows an interesting transition from the older to the newer forms, which goes
far to substantiate the differences I have pointed out between paleozoic and later
cockroaches ; while the study of a large number of specimens of Mormolucoides,
long but imperfectly, known from the
red sandstone of Connecticut, has en-
abled me to render it in a high degree
probable that this oldest known insect-
larva was a sialid.
In the monographic treatment of me-
sozoic insects we have only to record
the discussion of the rh/$/dorhabdi al-
ready mentioned, and a systematic re-
vision of the mesozoic cockroaches,
based on a considerable collection of
English forms new and old, lent me by




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[January 1890.
that veteran in their study the Rev. P.
B. Brodie, a work which included more
. than seventy-five species, treated after the method employed in the revision of
the paleozoic forms. The publication
of both these memoirs on the ancient
cockroaches, it may fairly be remarked,
has since brought to light many more
new forms, so that during the past de-
cade there have actually been added to
the number of pretertiary forms over a
hundred species of cockroaches, about
equally divided between paleozoic (53)
and mesozoic (57) times. A general
account of fossil cockroaches based on
these data was given in my "Cockroach
of the Past,'' in Miall and Denny's
"Structure and life history of the cock- roach" (London, I 886).
Passing now to tertiary times, we
naturally cannot expect to meet with
discoveries of equal importance and in-
terest to those which throw light upon
the origin of insect-forms. for it is a well known fact that the earliest tertiary in- sects are to all general intents and pur- poses identical with those of to-day.
They differ no doubt specifically, and
even to a considerable degree generi-
cally. Most of those so far recovered
from temperate regions indicate a then
warmer climate, but, taken as a whole,
the grand features of insect-life appear to have been essentially the same since
the beginning of tertiary times. By our
present researches upon them we no
doubt greatly widen our horizon, and
*as with modern types there always are
found problems of interest, so will there be with fossil insects, however recent.
Activity in this field can hardly be
said to be relatively so great as in the others, nor so great indeed as some
time ago when Heer and Heyden were
publishing extensively, but it neverthe- less has not been insignificant, and it is noteworthy that more special work
with groups has been undertaken ; thus
Buckton has summarized our knowledge
of the fossil aphides, Schlechtendahl
has elaborated the $sysopoda of Rott,
Gourret the arachnida of Aix, Hagen
and Kolbe the $socidae of amber, and
I the termitha of Florissant. I might
also add the butterflies of Florissant, as my paper, though not yet published,
has been months in type, and the gen-
eral results were given in a brief paper on 66Fossil butterflies" in general, in my "Butterflies of New England." Akin
to these can only be mentioned the
paper by Flach on the pleistocene cole-
optera of Hosbach, Schlechtendahl's
revision of Germar's tertiary fossils,
Williston's notice of the Florissant
Syq6hidac7 and mine of the Florissant
arachnida, my comparison of the Odo-
nata of Florissant and Green River,
the detailed study of Plunoce~ha?us
from Florissant, regarded by me as a
new and practically headless type of
I hysanura, and, finally, the discussion of the structure of this strange type and ot the supposed mite of the Rhenish brown
coal, Limnochares, both of which Bert-
kau regards as Galgulidae. To this
period also belongs my general survey of the pz\ eontology~ofFlorissant.
The additions to our knowledge of
the amber insects of Prussia during the