THE CARBONIFEROUS
INSECTS.

I'.VIJT I.
PAGES 1—80; PLATES I— IV.

Ipalaeontogvapbical Society

•

A MONOGRAPH

FOSSIL INSECTS

BRITISH COAL MEASURES.

MY

HERBERT P.OLTOX, M.So., F.R.S.E.. K.d.S.

c.K I'lIK IIKIS'J'OI, M1I8EUM.

PA1IT I.

PAUKS 1— so; PLATKS 1— IV

L() X DO X :
n •

FEINTED FOE THE P A L.EONTOO E A P HIC A L SOCIETY

JULY, 10 '21.

BT AliLAKH \ N [ i M..V AX]> WEST NKWMAX. LTD., l.ciM.HN \ Ml UdKKIN'..

THE FOSSIL INSECTS OF THE BRITISH COAL

MEASURES.

INTRODUCTION.

THE first recorded Palaeozoic insect of any country appears to have been a
"British specimen, LitJioxlulis brongniarti (Mantell), which was discovered in the
Coal Measures of Coalbrookdale in the early part of last century. It was sent by
Mantell to Brongniart as a leaf impression. Brongniart in turn submitted the
fossil to Mons. Audouin, who (1S33, Andonin, 'Ann. Soc. Ent. France,' ii, Bull.,
p. 7) described it as " d'nn inserte inconnn," and allied to the Hemerobiidas, Semblis,
and especially to (%•//</<(//* and ^[«nlix. The specimen was afterwards figured and
named by Mantell (1854, ' Medals of Creation,' vol. ii, p. o7o, fig. '2).

According to Parkinson, however, Lhuyd first recognised fossil insects in the
British Coal Measures. Parkinson (' Organic Remains,' vol. iii, p. 258, 1804 — 1811)
states that Lhnyd in a postscript to a letter to Dr. Richardson wrote as follows :
"Scrips! olini suspicari me Araneorum quorundam icones, una cum lithophytis,
in schisto carbonario observasse ; hoc jam nlteriore experientia edoctus aperte
assero. Alias icones habeo, qiue ad Scarabseorum genns ([iiam proxime accedunt.
In posternm ergo non tantnm Lithophyta, sed et qusedam insecta in hoc lapide
investigare conabimur." (' Lithophylacii,' p. 113.)

["I have formerly written that 1 believed I had observed certain impressions of
spiders identical with Lithophytes in carbonaceous shales; this I now, taught by
later experience, openly assert. 1 have other impressions which approach nearest
to the family of beetles. For the future, therefore, we will endeavour to investi-
gate not only Lithophyta, but also certain insects in these shales."]

Parkinson reprints four figures given by Lhnyd in his ' Iconograph/ tab. 4.
Two of these figures show eight legs and must therefore represent the remains of
Arachnids. None of the figures show wing-structure.

o o

Interest in the occurrence of fossil insects was stimulated in 1837 by the
publication of Dean Bnckland's 'Bridgewater Treatise' on Geology, in which he
described and figured two fossils found at Coalbrookdale by Mr. Anstice (18-37,
Buckland, ' Geology and Mineralogy,' '2nd ed., vol. ii, p. 70). He determined both
specimens to be the remains of coleopterous insects — a determination since
corrected by H. Woodward (1871, ' Geol. Mag.,' vol. viii, p. 380, pi. xi), by
Scudder, and finallv by Pocock, who referred them to the Arachnida ("Terrestrial
Carboniferous Arachnida," 'Mon. Pal. Soc./ 1'Jll, pp. 39, 77).
1

FOSSIL INSISTS OF THK BRITISH COAL MKASUFJKS.

Attention was afterwards diverted from tin.- Coal Measures l>v tin1 remarkable
discoveries of insect-remains made ]>y Brodie in the 1'iirlieck and Liassic rocks,
and by similar discoveries on the continent.

.Mr. E. W. Binney (1807, ' Proc. Lit. and I'hil. Soc. Manchester,' vol. vi, p. 50)
exhibited a specimen which "bore some resemblance to the pupa state of a
coleopterous insect," and had been found in the (.'inderl'ord I>vke 1'it at Bradlev,
near Hnddersfield.

A second specimen exhibited by Binney at the same lime \vas referred to
Xylolrius sigillarice, Dawson. Binney added: " \\ e must expect great additions to
be made to the Carboniferous fauna, as doubtless the rich and luxurious
vegetation of that remote period would afford food and shelter for numerous,
insects."

Binney's notes on the Huddersfield specimens caused the Uev. 1". 1>. Itrodie to
record (1807, ' Geol. Mag1.,' vol. iv, pp. i2.So — 2Mi) that lie had in his collection ''a
wing of a gigantic Neuropterous insect in inmstone from the Derbyshire Coal
Measures."

The same year Kirkby (1807. 'Geol. -Mag.,' vol. iv, pp. 3S8— 300) reported the
finding of clearly defined insect-remains in the Durham Coal Measures. One
example consisted of " portions of the fore wing or teginina of an orthopterous
insect nearlv allied to JUiilt/i or Cockroach," and the other "of an orthopterous
insect, apparently the abortive wing of a specimen related to the Phasinid:e."
Kirkby's first specimen is the small but verv line wing described here under the
name of Phyloniylacris mantidiuides (Sternberg). The second specimen ;s not
determinable as an insect-fragment, ami may prove to be a fossil fruit, referable
to one of the higher plants of the Coal Measures.

From 1807 onwards the finding of fossil insects in the British Coal Measures
occurred at long intervals until in 1'Jd1?, the date of publication of Handlirsch's
' I'ossilen Insekten,' the following had been recorded:

PhylomylacriB mantidioides (Goldenberg).

Olilll, " ulliril til 111,1/1:1," ICil'UllV.

olim, " Blattina uiiuiftili'iitli'*."

Goldeuberg.
Lithosialis brongniarti (Maiili'll i.

olim, " GTyllaeris brongniarti," M;mli II
LitJioinantis carbonarius, Woodward.

Ai-i-li;ru/ililtl!t iiliji-ux. Si'llildiT.

lii'iiJni I'fifi'ni'i in-ill , Scudder.
.l''lii-'ijiliii>--iiiii inii/IIi'ii, Scudder.
Aphthoroblattina johnsoni ( Woodward).

''Inn, " I''/"l>/ii / 1 1 tut fohnsoniy Woodwtird.

( ISIiittuiilt'K ) ji/'urliii ( Woodward).

ciliiii. " Etoblattina peachii, ' Woodward.
Leptoblattina c.i ///,-•, Woodward.
Lithomylacris kirklit/t. Woodward.
Siiiinii/liirriit deanensis (Woodward).

iiliin. " Etoblattina <!<'/< I/I'IIKIK,' Woodward.
Pseudofouquea cambrensis (Allen).

uliiii. " Fain/ urn i-niiilii-fitxig," Allen.
Breyeria woodwardiana ( Handlirsch).

olim, ";illinilv with Lithomantis cctTbonariits"
SI,. Mis.

olim, " Stobbsia woodwaTdiana," Hundlirscli,

Tin' numerous discoveries of insect-remains in the('oal Measuresof ('
(Allier). Krance. ;iml the remarkable series made known hv Handlirsch from the

INTRODUCTION. ?>

continent generally, and from the United States, overshadowed the limited British
series, which seemed almost trivial by comparison. The present monograph shows,
however, that the fossil insect-remains of the British Coal Measures are far more
abundant than was supposed, and that they are by no means unimportant. About
seventy specimens are known, of about sixty species, and they represent five of
the great groups of fossil insects.

PalsBodictyoptera are the dominant forms, and are closely followed by Blattoids,
several of which are referable to genera occurring in the French and Belgian
coalfields. The British examples of Soomylacris are represented near Lens and
Lievin by Soomylacris lievinensis, IV., while Pliylotnylacris mantidioides has its
counterparts in Phylomylacris i/txloui, Fr., and Plnjlomylacris lujlttci, IV., from Lens.
The great Protodonata of Comment ry are represented in the Bristol coalfield by
Boltoiiif'.'x radstoekensis.

The generic identity of French and British Coal Measure insects implies that
they formed part of a general and wide-spread fauna, a view which is strengthened
by the fact that while Soomylacris <1<'<uicnsis and N. #t»i'lci occur in the Forest of
Dean coalfield, to the west, N. Imrri occurs in the Kent coalfield to the east, and
only separated by the Straits of Dover from the Coal Measures of Northern
I'Yance, in which IVuvost finds other species of the same genus.

IVuvost has also shown that in the Coal Measures of Lens and Lieviu there is
present a well-defined hori/ou of Anthmcomya phillipsii, in which that species
passes through the same developmental changes as in the Kent, coalfield.

It is extremely likely that the Kent coalfield will later yield numerous insect-
remains closely allied to those of France, and that systematic search will amplify
the list of forms already known from all the British coalfields.

The British Palaeodictyoptera, on the whole, are more varied than the French,
few forms showing the primitive condition of Stenodictija, while certain examples,
such. as Mecynoptera tubercidata, Palasomantis -macroptera, and the three genera of
Lithomantids, are highly specialised. A similar degree of specialisation is seen in
the British examples of the IVotorthoptera, while the Blattoids, by their numerous
genera and species, indicate that the group had a long history and a wide
geographical range in the British coal period.

The fossil insects already found in the British Coal Measures form probably
but a small fraction of those which remain to be discovered when attention is more
fully directed to them. The insect-fauna, however, is not usually associated with
the general fauna in the Coal Measures, but occurs in beds of lighter coloured rock
than the ordinary carbonaceous shales, and with abundant ironstone nodules, or in
the case of the Blattoids, in association with masses of drifted vegetation in thek
black shales, where the neuration of the wings so closely simulates the pinnules of
Neuropteris as to be mistaken for the latter and cast aside.

While insect-remains are usually regarded as wholly restricted to the West-

4 FOSSIL INSECTS OF THE 1MMTISH COAL .ME AST RES.

phalian and Stephanian stages of tin1 Coal .Measures, tin- fauna with which they
aiv most often associated in this country may indicate a greater age, as it, is known
to occur as low down as the Calciferous Series of Scotland. Any statement,
therefore, of the range in time of lintish Palaeozoic insects based on the present
known forms may have to be set aside by later discoveries.

The fauna with which fossil insects are usuallv associated in Great Britain is
one in which arachnids and certain of the more primitive' arthropods are the
dominant forms. Arachnids are kno\\n to occur in the ( 'alciferous Sandstone
Series of Scotland at Kedhall, near Slateford, Edinburgh, and in the Cement-stone
Group of the Lower Carboniferous at Langholme, Dumfriesshire (I'.'ll, Pocock,
' Mon. Pal. Soc.,' p. IS), and elsewhere, the genus Archasoctonus being represented by
.1. (//(///(/• and .1. tuberculatns, and the genus Ci/clophthalmns by C. euglyptus at
Redhall, Blair Point, near Dysart, and Cramon near Edinburgh.

No insect-remains are known from any of these hori/ons, but if the faunal
association seen in the Coal Measures is a trustworthy guide, they may be looked
for with some prospect of success.

The faunal association existing in the '' Soapstone Bed " of the Lower Coal
Measures at Carre Heys, Colne, Lancashire (I'.M.io, Bolton, ' Geol. Mag. ' [•>], vol. ii),
is so similar in character to the typical insect-fauna elsewhere, that it is likely
that insects lived in the Lower Coal Measure period in Lancashire.

This faunal association at Carre Heys is as follows, and may lie compared
with the faunal association in which insects have been found to occur in other
coalfields :

AETHROPODA.

Pygocephalus cooperi, Huxley.
Anihrajpaleemon ferralnf, "\Vood\v.

,. iriinjir,ii;li, Et lll'Vlil U'O

Preshvichia r<itun<l<ita, AVoodw.
Architai~bus Kiilim-uH*,

//•/, Peach.

A MPHIBIA.

In-owni, Woud\v.

PISCES.

ii, TIUIJ.

Etiiji/rinus vildi [A. S. \Vo-nl\\.).
Microsaiirian remains.

The oldest known fossil insect in the British Carboniferous appeal's to be a
fragmentary wing (Genentomum .-•/<//</<•//////»), described l>y the author from shales
at a depth of li:!7 feet below the I Vdniiiisf er (ireat Vein in the liristol Coaltield,
and therefore al a considerable depth below the Pennant Grit.

Pseiidofouquea cambrensis (A.\len} was obtained from the top of the Four-foot
Seam in the Lower Coal .Mea.-nres at the Llanbradach ( 'olliery near Cardiff ; while
the shales over the Xo. LI Khondda Seam have yielded a wing-fragmenl of
Boltoniella tenuitegminata (Bolton); and Ihe shales over the Graigola Seam have
\K'lded the wings of two Hlattoids, Hemimijlacris convexa and Or/l
lanceolata.

HABITS AND MODE OF OCCURRENCE. 5

The No. 2 Rhondda Seam and the Graigola or Six-font Vein of Swansea both
occur in the Pennant Grits, the former near the base of the series and the latter
at 200 yards below the Swansea Four-foot Seam, which forms the base of the Middle
Coal Measures in South Wales.

In Monmouthshire, insect-remains occur in shales over the Mynddislwyn
Vein, a seam at the base of the Upper Coal Measures.

The Durham and South Lancashire Coalfields have yielded insect-remains in
measures near the top of the Middle Series, while those recorded from the Derby-
shire Coalfield are on a still lower hori/on in the Middle Series. Few fossil insects
are known from the Upper Coal Measures.

HABITS AND MODE OF OCCURRENCE OF FOSSIL INSECTS.

The bodies, and more particularly the wing's, of insects, have been entombed
in various deposits under conditions difficult to determine. Whatever the condi-
tions, they must have been closely related to the habits of life. The older writers
claimed that wind-dispersal and water-carriage were the chief agents. Buckland,
for example (' Anniv. Address to the Geol. Soc.,' 1842), supposed " that multitudes
of insects have been occasionally drifted by tempests to the sea." Mantell
('Wonders of Geology,' 7th ed., ls:>7) pointed out that Westwood had drawn
attention to the fact that "the streams brought down innumerable insects at
certain periods, perhaps those of heavy rain."

Alfred Russel Wallace (' The Geographical Distribution of Animals,' 1876) and
Heilprin ('The Distribution of Animals,' 1887) alike drew attention to the Avide-
spread occurrence of living insects far out at sea, in some instances still flying
strongly. Members of the British Association on their voyage to Australia in 1914
verified these statements by the capture of locusts as their vessel proceeded down
the Red Sea and into the Indian Ocean. More than a score of locusts were
captured on the vessel by which the writer travelled, and many more must have
been driven down into the water by the fringe of a simoon into which the vessel
entered beyond Aden.

Many insects are destroyed yearly by falling into streams and rivers after the
deposition of their eggs in the water, and by becoming entangled in the
surface film.

The occurrence of whole, or almost whole, insects is more likely to furnish
surer proof of the conditions under which life was passed than is the occurrence of
wings only, because the bodies, being more compact and much heavier than the
wings, are less likely to have drifted to great distances. Sometimes, as we shall
see later when we consider special cases, such as the Coal Measures of Commentry,
France, or the remarkable faunal associations of certain of the British fossil insects,

<) FOSS1I, 1XSKCTS OF TITF BRITISH COAI- MEASURES.

valuable information is supplied liy the deposits, or by the nature of the associated
forms of life.

The great group of the Palseodictyoptera and certain of the Protorthoptera
and I'rotodonata had large wind's, and were powerful fliers. \Ve slionld therefore
expect to find their remains widely dispersed in deposits of varied nature. This
seems to lie the case. Compact heavy-bodied insects like the Hlattoids would have
a more limited range, and their bodies after death could not be carried to threat
distances. Larval forms would in most cases be included in the deposits in the
immediate neighbourhood of the area in which thev lived.

]\I. Henri Fayol, in his description of the Coal Measures of Commeiitry,
France, shows that these deposits were laid down in narrow land-locked lakes of a
trough-like form lying in depressions of older schistose and crystalline rocks.
The trampiil waters received only the finest mud in suspension, and the resultant
mudstones have yielded a large insect-fauna, in which Blattoids are most numerous.
The bodies of the insects are preserved in many cases. Certain of the insects were
strong fliers, and their occurrence with the bodies intact indicates that they, in
all probability, haunted the vicinilv of the lakes and flew over them. When
strongly-flying insects like BoUotiites radstockensis or Lithosialis bronrjniarti died
upon the land, the wings, because of their membranous and cliitinous nature,
would persist after the destruction of the softer body, and be swept off into streams
after heavy rains or Hooding of the land-surface, their great superficial area
combined with their lightness making flotation easy.

The transference of insect- wings from the land into water would be accom-
panied bv the drifting of plant-material, and the two would be buried together in
the deposit then formini;'. The wmt;' of Boltonites from Radstock was found with
plant-remains in deposits of this sort, and may be taken as a prool. supported as
it is by other examples of IVotodonate wings, that these insects lived over the
land and died upon it.

The Palseodictyoptera, with their wings capable only of an up-and-down move-
ment in one plane at right-angles to the bodv, and, when in a position ol rest,
disposed straight outwards, are not likelv to have Frequented the ground, except
in the open. These insects, like ino>( of the I'aheo/.oie forms, were all ot large si/.e,
as contrasted with living tvpes. Pruvos! assumes that the characters of the wing
unfitted these insects fora fore>t life, and that they muM have been restricted to
flight in the open neighbourhood of swamp pools. I do not wholly ai^ree \\iih
this assumption, for the branches and leaves of the Coal Measure plants do not
seem to have had so great a densit v and interlacing of Foliage as seriously to
impede the flight of powerful \\mgeil insects. There seems no reason why these
insects should not Irive lived among the brakes of 1 .epidodend roid and ('alanntean
tree-, and after death fallen or been blown into adjacent waters. The lad that
isolated wings are often found in perfed condition and without any signs of wear

CONDITIONS OF LARVAL INSECT LIFE. 7

and tear sucli as the wings of aged insects show to-day, raises tlie question whether
in some cases the wings were not shed, as in certain species of recent ants, the
shorn insect continuing its life as a ground-feeder.

The Protorfchoptera were, judging from the structure of their month-parts,
somewhat general feeders or carnivorous, and the presence of strong walking legs
suggests that thev spent much of their life on the ground, possibly along the
margins of swamps, where food would be especially abundant. They had, never-
theless, powerful wings, and some members, such as the (Edischiidse, had legs
adapted for leaping. Orthoptera, represented chiefly by Blattoid forms, were all
fitted for ilight by means of their hirge membranous hind-wings, and equally well
fitted by powerful walking legs for life on the ground. In repose tin1 hind-
wings were hidden under the modified fore-wings.

I have elsewhere given my reason for a belief that the Blattoids were not
wholly phytophagous, but in all probability carnivorous also ('Quart. .lourn. Geol.
Sec...' vol. Ixvii, p. l->5, HU1).

Blattoids may also have entered the water in search of food, for the hind
r!vin»' wiii»'S would be securely protected by the tegmina, whose broad muscular

i O «' 1 *

bases of attachment were sufficiently powerful to compress them down on the back
and prevent water entering beneath, just as in the case of the living water-beetle,
Ifi/tlrojiliiliis pieeiis. The chitinous surface of the body and of tin1 tegmina would
not hinder progress in water, for their sin-faces are no rougher than those of the
modern Ih/tt'wiix, nor would the insect on emerging bring with it so heavy a film
of water as to clog its movements. An objection may be found in the presence of
stout bristle-like hairs on the legs seen on such forms as Neeymylacris lerichei
(Bolton) (I'.'IT, 'Mem. and 1'roc. Lit. and I'hil. Soc. Manchester,' vol. Lxi, p. 15),
which might conceivably cause air-bubbles to cling in such profusion as to prevent
the insect being able to submerge. The presence of fine hairs on the swimming
legs of ]li/<1r<ii>lnlitx and .Ih/tim-ns does not hinder the immersion of these insects in
water, so that this is not a valid objection. If no hindrance to immersion was
caused by the bristle-like hairs, they may have been useful in assisting the act of
swimming.

1 think the probabilities are in favour of the Blattoids being at least semi-
uquatic as well as land insects.

CONDITIONS OF LARVAL INSECT LIFE.

The conditions under which larval life was passed are even more conjectural.
The Protodonata may be regarded as insects whose larva1 must have been aquatic,
like the aquatic larva- of the Odoiiata now living, but Tillyard (1 '.M 7, ' The Biology
of Dragonflies,' Camb. 1'niv. I'rrss, p. •'!<>(;) conjectures that since adult, Prot-
odonata are found at Commentry without the occurrence of larval forms, the latter

FOSSIL TXSKCTS OF TIIK BRITISH COAL MEASURES.

may have dwelt in (.lamp earth rather than in water, ami that ''the formation of
the larval traeheal system undoubtedly proves that this at one time was the case.
It mav well have been so in Carboniferous tunes."

Tillyard's views are well worth ((noting in full, espeeially as they support in
some measure those of t'ruvost : " \Ve mav pietnre to ourselves the giant insects
of Cominentry as inhabiting the shores of a large, shallow, nearly stagnant
lake. In the muddy oo/e around its borders grew forests of the Giant Mare's
Tail, while further buck on the sandv slopes the graceful C'ycads and other extra-
ordinary plants formed a more diversified medlev. There, amidst rotting vegetation,
these insects lived and bred. In such almost amphibious conditions it may well be
that the larva1 of Protepliemeroidea and Protodonata first began that series of
adaptive changes which finally led them to adopt a purely aquatic mode of life."

The lar\;e of Ilr/iilin and of other forms whose wings I describe under the
name of "Pteronepionites" must have lived under conditions fitted for their gradual
metamorphosis. The body was long, well segmented, and bore rudimentary
wings, which were carried well up over the thorax in an erect or semi-erect posit ion.
Though rudimentary, the wings possess features which may have determined to a
large extent the mode of life. They are attached by broad, strong bases to the
thorax, and are very muscular, as shown by the stout ridges proceeding from the
point of attachment into the wings, and the anterior margins are also thickened.
The bodies with their lateral expansions of the terga are very suggestive of those
of the Diplopoda, and like them would offer no serious obstacle to progression
through rank and rotting vegetation. That these larval insects would also
penetrate soft muds, if necessary, in search of food is possible, since the
soft-bodied caterpillars of the Hawk-moths of to-dav are able to enter hard soil
before pupal ion takes place.

The stout wing-bases and the strengthened margins of the WHILES would
prevent damage to these structures as the larva? crawled about, or sought to bury
themselves in the soil or muds. They were essentially adapted for a ground
habit. Whether they were capable of an aquatic or semi-aquat ic habit can only
be settled by a knowledge of the mode of respiration.

Lubbock, Gegenbaur and others have adduced strong reasons in favour of an
aquatic origin of the insects, ami in the Carboniferous types we should naturally
expect that t he original habits had not had time to undergo any great modification.
Larval wings ol the " Pteronepionites' Ivpe must have been living structures in
\\hich metabolism was active, and very unlike tin: dried membranous sac-like
expansions of the adult insect. The growth of the larval wind's was continued
throughout metamorphosis, ami during this period then- delicate nature, broad
expanse, and the thinness ol the inlegiiiiienl may have enabled them to assist in
the respiratory fund ion.

The researches of Comstock and Xeedham show thai larval \\ IIILI'S of recent

CONDITIONS OF LARVAL INSECT LIFE. 9

insects receive a plentiful supply of trachea! brandies at an early stage, and it is
evident that these are much in excess of any aeration the wings are likely to
require. The trachea! development seems to be a persistence from an earlier
more active condition, when the larval wing's may have played a part in assisting
the respiration. These considerations, and the presence of spiracle-like structures
in the interstitial neuration of the adult wings of many Pala?odictyoptera, lend
support to the inference that the wings functioned as organs of respiration. These
spiracle-like structures are usually oval or rounded in outline, and thickened. In
some instances they show a series of raised lines radiating from the thickened
edge into the surrounding areas, as if they had been muscular strands and capable
of expansion and contraction.

Are these structures the atrophied remains of spiracles once functional, and
fitting the larva for a more or less aquatic existence? During a recent
visit to this country Dr. Tillyard has suggested to me that they are
rudiments of sensory organs which may have been scent-glands. Scent-
glands are known to occur on the wings of many insects, as, for example, the
Green-veined White (Pieris net pa?), the Small White (Pteris rapa?) and others,
and their appearance is certainly a strong argument in favour of the view. Scent-
glands are, however, in all probability but specialised developments of previously
existing structures, and it is possible that the glandular-like organs to which I
give the name of "pseudo-spiracles," and Handlirsch the name of "pterostigmata,"
are an earlier development connected with the " tracheoles " of Comstock, or that
primary tracheolation to which Tillyard has given the name of " archyodictyon."
Tillyard does not accept the view that they had ever any connection with
respiration.

The almost total absence of structures which can be accepted as functional
gills in these fossil insect-larva? may be accounted for by the perishable nature of
such organs. Before dismissing the question of the respiratory function in its
relation to the conditions under which larval life was passed, it is desirable to draw
attention to the larval Blattoid, Lppt<>l>l<ttt!it« en7/x, YYoodw. In this insect the
abdominal segments have the dorsal hinder maryin extended into broad lamellar

o O

expansions so filmy in texture that they may have served as organs of aeration.
The lamellar expansions were longer in life than they now are, the hinder borders
showing an irregular torn edge. Their extreme thinness would permit of a ready
osmotic-like action, especially in damp vegetation, or in an aquatic or semi-aquatic
habitat. Scudder, Handlirsch, Lameere and others are all agreed in the belief
that the Blattoids frequented decaying vegetation in or near water, and under these
conditions the presence of organs of aeration similar in character to the abdominal
expansions of Lcptoblnttiita ctHix would be of the greatest value and offer no
difficulties to the habit of life. No similar structures are known in any other
larval Blattoid, so that the view cannot be pressed.
2

10 FOSSIL INSECTS OF THE BRITISH COAT, MEASURKS.

We shall not be f;ir wrong in assuming that the larva? of some of tin- Coal
Measure insects were wholly aquatic, others semi-aquatic; that the adult Blattoids
were indifferently aquatic or terrestrial ; the adults of the non-Blattoid types spent
must of their life in the vicinity of the swamp-pools in which (heir larval stages
Avere passed, and to which they might need to return to lay their eggs.

Such a view seems to accord with the known facts, and will explain the special
character of the fauna of such deposits as those of Coseley, in Staffordshire, and
the brick-clays of Sparth Bottoms, Rochdale, Lancashire. These are evidently
true lagoon or swamp-pool deposits as contrasted with the ordinarv shales and
binds of the Coal Measures.

FOOD OF COAL MEASURE INSECTS.

The nature of the food of Coal Measure insects has been much discussed, as it
is so closely associated with habits. Ilandlirsch considers that the Gymnosperms
and Pteridosperms of the Coal Measure forests were not likely to have been

frequented by insects in search of f I, as these plants do not prove attractive to

living insects. Pruvost, on the other hand (1CJ1(.>, ? l'J'20, " La Fanne Continent ale
du Terrain Honiller du Nord de la France," 'Mem. Carte Gcol. France,' pp. LMili —
207), considers that many members of the Coal Measure flora possessed in their
spores, or in the case of the higher plants, in their cones, a plentiful food supply for
insects, and he finds in the association of a l'1iijllol>l<iU« at Lens with the r»l<>nir<i
of Linopteris some support for his conclusions.

The contemporaneous rapid development of plants and insects is also quoted
by Pruvost in support of his views.

Several writers have argued that the powerful wings and consequent powers
of rapid flight of many of the insects are more in accordance with a predatorv and
carnivorous habit than witli a purely frugivorous or herbivorous one, and this
belief has led Lameere to write as follows (1(.»17, ' Hull. Soc. Zool. France,' vol. xlii,
pp. -50 — 37): "Over the lake of Commentry flew magnificent Ephemeroptera and
splendid Odonatoptera, the carnivorous larva' of which were aquatic; doubtless
the Odonatoptera, when fully grown, devoured the Kphemeroplera, of which (In-
most fully developed types, the Megasecopterid;e, which have left no descendants,
must have made great slaughter among the smaller insects.

"On the ground, in the forests, swarmed innumerable Ulattoids, which
frequented the detritus, and which had as enemies the ferocious and agile
Orthopiera, the varied counterparts of our Mantidse. These latter must have
attacked equally the large vegetarian Orthoptera, the counterparts of the Phasmag,
which probably climbed on trees, and I he bulk v Protohemiptera, which sucked (he
sap. Some of the Orthoptera jumped, anil there were some which by their
appearance recall our . \cridians, but all these beings were mute.

FOOD OF COAL MEASURE INSECTS. It

" Small amphibians, and numerous arachnidians, came to limit the swarm of
this articulate world in a country without birds or mammals."

Prnvost does not accept the view of a primitive aquatic origin of insects, but
affirms his belief in a terrestrial origin, and thinks that even if an aquatic habit
be proved for certain of the larva1 of the Coal Measure insects, the habit must
have been secondary, and derived from an earlier land ancestry (<^>. '.'//., p. 268).

Scudder has observed in the case of the Blattidse that the venation of the
tegmina very closely resembles the surface-features of the Neuropteris pinnule—
so strongly in fact as to suggest mimetism.

Pruvosfc rightly urges that a mimetism is of little value unless the mimetic

O «/ o

insect frequents the plant mimicked. At the same time, it can be urged that the
stout compact bodies of the fossil Blattoids and their powerful walking legs were
equally admirably fitted for progression among rank and decaying vegetation, and
that in these conditions the Blattoids were quite as likely to have been omnivorous,
while finding some degree of protection among the Neuropteris pinnules lying on
the ground.

The writer has previously commented on the association of the wings of Blattoids
with the leaves of <'<n-<liil<-x (101 1, ' Quart. Jonrn. Geol. Soc.,' vol. Ixvii, pp. UU—
l(i">), and has made the following comment: "While Carboniferous Blattoids
may have been wholly phytophagous, it is interesting to note that the leaves of
Corchdtcs (in the present case) are impressed with shallow pits, which shcnv faint
traces of a spiral. I have in many previous instances found that such pits owed
their origin to attached shells of /ty/ror/W.s' /I«N////(.S. Whether these leaves were
partially submerged in \vater during life is an open question ; but in all cases the
plant-tissues of the pittings are depressed, and are accurate impressions of
N///ni/V;/.s\ If the Carboniferous Blattoids were not wholly vegetable feeders, the
occurrence of Spirorbis pusillus upon the i.'m'tlnites may supply a reason for their
frequent association."

CLASSIFICATION.

The classification of fossil insects has presented great difficulties, both to the
palaeontologist, and to the systematist of living forms. Palreozoic insects show to
the systematist a series of forms not strictly referable to any modern grouping,
but presenting certain generalised details of structure which link two or more
now widely separated groups, besides other features not met with in living
forms.

The palaeontologist finds that he has not to deal with early and primitive
types, followed by a regular series showing a developmental progression, but with
an apparent sudden incursion of large series of highly modified and well-developed
insects, co-existent with others of more primitive type.

12 FOSSIL 1XSKCTS OF THE B1UTISH COAL MEASl'l.'KS.

Further discoveries will doubtless do niueli to eliminate these difficulties, but
present knowledge is such that recent entomology helps very little, and the
classification of Paheo/.oie inserts must lie largely bused on the known fossils,
realism^ that many of the intermediate forms are yet unlcnown.

The classifications of various authors vary widely, and even the broad general
facts of relationship are still uncertain. The earliest attempt at a classification
of fossil insects appears to have been that of (Soldenberg ( 1 S73- -1877, 'Fauna
Saraepontana Fossilis : Die Fossilen Thiere aus der Steinkohlenformation von
Saarbriicken.') lie recognised a new Order, Palseodictyoptera, for the inclusion
of fossil insects differing in structure and the shape of the wings from
living representatives of the Orders Xeuroptera and Orthoptera, while also
possessing characters which seem to link the two orders together. He did
not define the order, but arranged it with the other orders as follows:

Order. — PAL^EODICTTOPTEEA, Gulden! >.

Genera. — Dicti/mii'in-n, Goldrnb. ; Engsri-nii. Ildhrn; Mhin/iti, Puna ; Ili.'iiirriflin, Puna;

Hdplophlebium, Seudder.
Order. — ORTHOPTERA.

Sub-order. — Orthoptera Pseud". Neiiroptera.

Genera. — Tenm.-, Gulden!'. ; Tenititiiliiiiii, Guldenlp.
Sub-order. — Orthoptera vera.

Genus. — Blattiiiti, Gerinar.
Order. — EHYNCHOTA.

Sub-order. — Homoptera.

Genus. — Fii!/j<>riiiu, Goldeub.

Seudder (1887, article '; lusecta," ' Trait c de Palseontologie,' by Karl von
Zittel, translated by Dr. Charles Barrois, vol. ii, ' Palaeozoologie,' pp. 7H> — 833), in
his latest classification, considerably extended that of (loldenlierg, while retaining
the primitive group of the 1'aheodictyoptera. His complete arrangement is as
follows :

A.— PAL/EODICTYOPTERA,

OBTHOPTEROIUKA. Snulder.

Family. — Palseoblattariae, Si-udder.
Sul-faniily.— Mylarridic, Seudder.

Genera. — Mylacris, Scd. ; PremyJacris, Srd. ; Paramylacris, Bed. ; Lithomylacris,

Si'd. ; JKecymylacris, Seil.
Sill -iainilv. — Blatt inari.i', Srudder.

'li'iiera. — Etiililiil/iini, Sed . S/ii/,ililiilliiin, Sed. ; .•li'i-liinitj/Hi-i-is. Scd. ; Anflirtn-n-
blattina, Bed . Gerablattina, Scd. ; Hermatoblattina,Scd.;

liim. Scd.; Oi-i/r/iilil.illiiht. S -d.; ri'lrnlil.ittimi. Sed. ; rni-nl

Family. — Protophasmidae, I'lnm^.

Genera. — TitanopJiasma, Mron^. ; liil/nn-iirii. Sed.: /'/r7//n//r»iM. < ioldenli. ;
riilinjili'iiiia, Scd ; ArcJucojjtilua, SIM!.; ri-iilnjilnmiiiii, ]!nin^. ; lin'ijrriii.
dc liiurc; Meganeura, l!i'.>n^.; .l''.<l<r<iiil«iaiiut. Sed. : Goldenbergia,
S<-d. ; llii^ln[ilil<-hinni, Sed. ; 1'iiulin, Siuiili; Archegogryllus, Sed.

CLASSIFICATION. 13

NEUROPTEROIDEA, Seudder.

Family.— Palephemeridse, Sol.

Genus. — PlatepJiemera, Scd.
Family. — Homothetida?, Sod.

Genera. — Acridites, And.; E>ii\i'nns, Scd.; Gfn<nitfryx, Scd.; Genentomum, Scd.;
Diilynwjilili'ps, Scd.; Humothefug, Scd.; Mi.eoterme$, Sterzel ; Omnlia,
van Ben.
Family. — Palseopteriua, Scd.

Genera. — Minniin, Sod. ; Dieconeura, Scd. ; Strephecladus, Scd.
Family. — Xeuoneuridse, Scd.

Genus. — Xen one nra, Scd.
Family. — Hemeristina, Scd.

Genera. — Lithumantis, Woodw. ; Lifhosiulis, Scd. ; Pachytylepsis, de Borre ;

Lithentomum, Scd. ; Hemerislia, Dana.
Family. — Gerariua, Scd.

Genera. — P6lyemu8,ScH. ; Gerarus,BcA.; Adiphlebia, ScH. ; Meyattienfomnm, Scd.

HEMIPTEROIDEA, Seudder.

Geneva. — Eiujereoii, Dolirn ; Fulgorina,, Goldeiib.

COLEOPTEROIDEA, Seudder.

Palaeodictyoptera having a coleopterous aspect indicated liy Geiiiitz and Brongniarfc.

B.— HETEROMETABOLA, Packard.
ORTHOPTERA, Olivier.

Family. — Forficulariae, Latreille.
Family. — Blattariae, Latreille.

Genera. — NeortJirobhittina, Scd ; Scutinoblattina, Scd.; BlaffiiliHin, Westw. ;

Mesolilaltina, Gein.
Family. — Mantidse, Latreille.

Genus. — Mantis. Linur.
Family. — Pbasmidit', Leach.

Genera. — Agathemera, Scd. ; Pseudoperla, Pictet.
Families. — Aoridii, Latreille ; Locustidse, Latreille ; Giyllidse, Latreille.

NEUROPTERA, Linuc.

Sub-order. — Pseudoueuroptera, Erichson.
Sub-order. — Neuroptera vera.

HEMIPTERA, Linnc.

Homoptera, Latreille.
Heteroptent, Latreille.

COLEOPTERA.

C.— METABOLA, Packard.

PlPTERA.

LEPIDOPTERA.
HYMENOPTERA.

A modification of this classification was introduced by Handlirsch, in his
"Sub-phylum Insecta" in Eastman's translation of Zittcl's 'Text-book of
Palaeontology,' 1913, as follows :

14 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

Class I -PTERYGOGENEA, Brauer.

( >rdiT. — PAI,.I:<>DICTTOPTERA, Goldeuberg.

Families. — Dictyoneuridse, Megaptilidse, Hypermegethidae, Lithomantida>, Heolidse,

FouqueidfE, Spilapteridse, Lamproptilidee, Polycreagridse, Paolidu--.
Order. — MIXOTERMITOIDEA, Haudlirseh.
Order. — RECULOIDEA. Handlirsch.
Order. — PROTORTHOPTERA, Handlirsch.

Families. — Spanioderidse, Ischnoneuridse, Caloneuridee, Sthenaropodidse, (.Edisclmdse,

Geraridse, Cacurgidse.
Order. — ORTHOPTERA, Olivier.

Sub-order. — Locustoidea, Leach.
Sub-order. — Acridioidea, Haudl.
Order. — PHASMOIDEA, Leach.
Order. — DERMAPTERA, De Geer.
Order. — DIPLOGLOXSATA, De Saussure.
Order. — THYSANOPTEKA, II a lid a v.
Order. — PROTOBLATTOIDEA, Haudlirseli.

Families. — Stenoneuridee, Protophasmidse, Eoblattidse, Oryctoblattinidie, ^-Etopblebidse,

Cheliphlebidse, Eucseuidse.
On.ler. — BLATTOIDEA, Hamllirsdi.

Families. — Spiloblattinidse, Mvlacridse, Poroblattiuidse, Neorthroblattinidse, Mesobla.t-
tiniihi'. Pseudomylacridee, Dictyomylacridse, Neomylacridse, Pteridoiuv-
lacridae, Idiomvlacridte, Diechoblattinidse, Proteremidse.
Order.-— MANTOIDEA, Handlirscli.

Genus. — PaJiconwiilif. Bui ton.
Order. — SYPHAROPTEROIUEA, Handlirsch.
Order. — HAPALOPTEROIDEA. Handlirsch.
Order. — PROTOEPHEMEROIDEA, Handlirsch.
Order. — PROTODONATA, Broiitjiiiarl .
Order. — MEGASECOPTERA, Brongniart.
Order.— PROTOHEMIPTERA, Handlirsch.

The publication of Handlirsch' s great work, 'Die Fossilen Insekten/ lOOil—
T.HI8, marked an important jilia.se in the history of the stmlv uf fossil inserts.
Hamllirseli surveyed the whole field of fossil entomology, and brou«>'lit the "-real

•/ O •/ ' O O

bulk of the known forms under abruad classification. The Order Palaeodictyoptera
was much extended, defined, and made to include a lar^e series of families,
several of which, however, are clearly widely divergent in type. This
was soon recognised hy oilier workers, ;is doubtless by Handlirsch himself,
who may have considered it wiser to extend Goldenberg's order, even to
tin1 inclusion of forms not definitely related, rather than to formulate a new
classification the components of which could not be rigidly defined. Knowing
t.hal the field ol research was rapidly widening, Ilandlirsch exercised a \\IM-
iv-tramt in not adding a new classification, which could only be of a temporary
character. Subsequent events have proved the wisdom of Ins action. Since
1'JOS, (lie study of fossil insects has attracted more students, new localities and

CLASSIFICATION. 15

insect-horizons have been found, and many new types recorded. Some of these
linking forms already known, and others indicate relationships not fully under-
stood. The retention of the Order Palasodictyoptera has therefore resulted in the
formation of a somewhat heterogeneous assemblage, all members of which have
one point of agreement. They are primitive co-types, more nearly related to each
other in various ways than the}r are to recent insects, although that relationship is
not always as demonstrable as one could wish.

The most ambitions classification yet attempted is that of Prof. Lameere
(1917, ' Bull. Mus. Hist. Nat., Paris,' no. 1), who has published only a summary of
his conclusions. We are not able to determine how valid are his arguments, or
if he is justified by evidence in setting forth his new scheme. He sweeps the
Order Palreodictyoptera wholl}' away, pointing out that it consists of a hetero-
geneous assemblage, and substitutes a more detailed classification as follows:

SUBULICORNIA.

EPHEMEEOPTEEA.

Family. — Spilapteridse.

Genera. — LainprojitHia, Brong. ; E^itftJte, Handl. ; Becquerelia, Brong. ;
PaJn'iiptiliis, Brong.; Compsoneura, Brong.; Sjiiluptilus, Handl.;
Homaloneuru, Brong.; Graphiptihis, Brong.; Spila^iterii, Brong.
Family. — Megasecopterklae.

Genera. — Aspidothorax, Brong. ; Gorydaloides, Brong. ; DiapJianoptera, Brong. ;
Cycloscelis, Brong. ; Sphecoptera, Brong. ; Psilothorax, Brong. ; Mis-
clioptrra, Brong. ; Ischnoptilus, Brong.
Family. — Protephemeridae.

Genera. — Apopappus, Haudl. ; Trijilusoba, Haudl.
ODONATOPTEEA.

Family. — Fouqueidae.

Genera. — Fuuquea, Broug. ; Rhubtl<if>tilus, Brong.
Family. — Dictyoiieuridae.

Genera — Microdictya, Brong. ; StenoJivtya, Brong.
Family. — Dictyoptilidae (Protcidonata).

Genera. — Archsemegaplilus, Brong. ; Dictyoptilus, Brong. ; Poromaptera, Brong. :

Protayrion, Brong. ; Qilaonia, Broug. ; Meganeura, Brong.
EHTNCHOTA.
PEOTOHEMIPTERA.

Familv. — Homoiopteridss.

Genera. — Lycocercus, Haudl. ; Huntoioj-itera, Brong. ; Lithojptilus, Brong.
Family. — Megapt ilida?.

Genus. — A/V;/.^</ ilns, Brong.
Family. — Breyeridae.

G-ewis.—Megaptiloides, Haudl. ; Burrea, Brong.
Family. — Mecynostomidae.

Genus. — Mecynostoma, Brong.
HEMIPTERA.

Family. — Dictyocicadidse.

Genus. — Dictyocicada, Brong.

10 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

ORTHOPTERA.

Noiiioiieura (BhiHa'fnrmia, Handl.).

(a) Ulattoidea.

Families. — llyalnptilida-, Protoperlida-, Favolifllidn'. ( iryrtoMattiniilir.

(b) Mautoidea.

Families. — Stenont-uritido?, Stenoneuridae, Isi'liuoiifiirida-.

Heteront'ura. (Equivalent in part to Orfclioptera Cursoria and Orthopteroidea, Haudl.)
(</) I'hasmoidea.

Fa mil v. — Sthenaropodidse.
(li) IjixMistida1. (Equivalent in part to Orthoptera Saltatoria. )

Families. — (Ediscliiidne, CaKmeuridaj.

Lameere restricted his research to the French fossil insects, his studies being
based on the types described and figured by Brongniart (18'J^, 'Bull. Soc.
Industrie, Saint Etienne,' o ser., vii), the collections made by Fayol, and the
large series of fossil insects from the Upper Coal Measures (Stephanian) of
Comment ry, now preserved in the National Museum of Natural History, Paris.
The following remarks mav be made on his classification :

Ei'HKMKKOiTKKA. — The three families forming this division are regarded as
closely related, the genus PecguereHa of the Spilapteridse bearing certain characters
of the Megasecopteridae, while the family is also linked through the genus
Apopappus (which, is taken to supplv a natural transition between the Spilapterjdse)
to the genus Triplosoha.

ODOXATOPTliEA. — The family Fonqueidiv is held to differ from the Spilapteridse
in that transverse veins are nunierons, close together, and form a network over
the inner margin, and in the anal area a feature which brings it nearer to the
Protodonata. The family Dictyoneuridas possesses a network of veins extending
over the whole wing, as in M!ci-<i<li<:ti/<t. The remaining family, Dictyoptilidse,
contains Archsemegaptilus, which I.anicei-e considers differs only from the Dictyo-
ncnrida1 in the fusion of the median and radial veins at the base of the wing.
Protinjrlon is considei'eil nearest to the ti'iie Odonata, while Megaiteura and Gilsonia
are specialised types.

RII vxeuoTA. — The presence of a rostrum in f,iji-nr,'i-t-n^ </n/(/r///>c/v//, and the
resemblance of the head and leg of Homoioptera gigantea to the same structures in
'ini, are considered sufficient proof of the Protohemipteroid characters of
!!*, Homoioptera, and the allied genera Lithoptilus, Megaptilus, Mecynostoma,
ArcJissoptilus and Pctvciinegcqjtilns.

()i(Tiioi''i'i-:i;.\.-- l-ameere regards I landlirsch's group of I'rotorthoptera as an
assenil)lage of two I'elated but distinct types, \\hich he classifies under Xomo7ieura
and I letcmneura. The genus Stenoneurites is regarded as the connecting link
between the Abmtoidea and the ancestors of the Hlattoidea, the genus StenoneilTQ,
being also in some measure transitional between StenOHeurites and the l-chno-
aeuridse.

CLASSIFICATION. 17

Nomoneunt. — This sub-division includes the Blattseformia of Handlirsch, and
is distinguished by tlie wings having no precostal area, as contrasted with a second
division, Heteroneui'a, in which a precostal area is present, and in which the legs
are adapted for running and jumping. The Xomoneura include forms classified
bv Handlirsch under the Protorthoptera and Protoblattoidea. Lameere separates
his Nomoneura into (a) Blattoidea, and (b) Mantoidea, the former characterised by
a sub-costal which joins the outer or costal margin, a more or less lengthened
radius, and a small cubitus. The Mantoidea have the sub-costa joining the radius,
and a large preponderating cubitus.

Heteronritra. — This sub-division includes the Phasnmidea and the Locustoidea,
the former containing the faniilv (n) Sthenaropodidse, in which the legs are long
and stout, the head prognathous, the prothorax long and narrow in front and very
wide behind, and presenting two dorsal expansions.

The wing- venation of the Sthenaropodida? is such that they may have
been the ancestors of the Phasmida?. Lameere, however, does not regard the
Sthenaropodidaa as ancestors of the Phasmida?, but as arising with them from a
common ancestor.

The members of the families (Kdischiida? and Caloueuridre possess legs fitted
for jumping, but differ considerably in their wing- venation. The (Edischiidse are
possibly linked with the Locustidsv, and the Caloneuridse with the Acridida?.

Circumstantial and detailed as Prof. Lameere's classification is, the arguments
and deductions are not easily Followed, the paper being only in abstract. A study
of the fossil insect-material from Coinmentry alone is not in itself likely to yield
all the facts and premisses upon which a classification can be built applicable to the
Palaeozoic insect-fauna of all coalfields and countries. .Much more evidence is
wanted, and until the full paper is published, it is necessary to hold the classification
in suspense.

The most recent publication on the Palaeozoic insects is an extensive and
valuable memoir by Dr. P. Pruvost (HH'O, "La Faune Continental.' du Terrain
Houiller du Xoril de la France," 'Memoires pour servir a 1'Explication de la Carte
Geologique detaillee de la France,' Paris, H)20) on the fossil insects recently found
by him, and others, in the neighbourhood of Lens and Lievin in the north of
France. Dr. Pruvost adopts the classification of Handlirsch with few emendations
as follows, and by his new material he has added considerably to our knowledge of
the Protoblattoidea and Blattoidea :

Order. — PAL^ODICTYOPTERA.

Family. — Stenodictyopteridse, Brong. (Dictvoneuridse, Handlirsch ).

Family. — Spilaiiteridiv, Brong., emend. Haudlirseli.
Order.— PROTORTHOPTERA, Handlirsch.

Families. — <Edischiida\ Caloueuridse.
Order. — HAPALOPTEROIDEA. Handlirsch.

Family. — Hapalopteridae,

3

18 FOSSIL 1XSKCTS OF TIIH BRITISH COM, MEASURES.

Order. — PROTOHI.ATTOIUKA. Ha.ndlirsrli, enu-inl. Pruvost.
Sub-order.— Archiblattidse, Pruvost
Fa.milv. Cariir^klr, Hamll.
Siili (inlcr. — Archimantidse, Pnivost.

Family. — Cymenophlebidse, Pruvust.
OnU-r. -Bi.ATTouiKA, Haixllirwli.

Family. — Archimylacridse, Hiiinll.

(.Ti-uera. -Archiinylaeris, A*<-iiii>lilntt<i. Mannlilatiii, Actinoblatta, 1'lii/lnlilttttii, An-lt:fi.>-

lijilii', BarToiaiblatta, Grypoblatta, MesitoMattct.
Familv. — Mylai-riilir. Sol.

Geneva. — HemimylcLcris, Phyloniylacris, Trilophoniylacris, Soomylacria, OrthomylcLCi'is,

Slt'iiuiii i/htcrix, ami JjitJiomylacris.
Family. — Purnlilatt iniiliv, Ha mil.
s Preninoiblatta, Pniv.

1 have compared notes with M. Pruvost, and we have arrived independently
at the conclusion that for the present the classification of Handlirsch is, with few
emendations, the best to adopt, and most in keeping with the known tacts.

FAUNAL ASSOCIATION.

Various collectors in British Coalfields have discovered not only insect-remains,
but a definite faunal association, of which the significance seems to have been
overlooked, and it has therefore not received the attention it deserves.

Most of the insect-remains are found in ironstone nodules, and the beds in
which these nodules occur are usually light- coloured rocks more similar to hardened
clav than to normal shales. The nodules are in vast numbers, rangingin size from
half-an-inch to ten and twelve inches in diameter. The beds seem more com-
parable to the fireclays or seat-earths than to the ordinary fissile shales, and both
in lithological character and fossil contents stand in some measure apart from the
ordinary Coal Measure rocks. They are not restricted to one coalfield, but have a
wide distribution. \Vherea systematic search of beds of this character has been
made, the insect-remains have been found accompanied l>v a launa in which
arthropods of a more primitive type than insects are conspicuous.

The character of this fauna will be best understood by reference to the
following lists of fossils which have been recorded from certain localities:

DlKHAM CoAU'lKl .11.— " /one of .1 //l/ii'ilrini/l/il />// ill i />xii ( Will.) " ill Upper part
of the Middle Coal Measures; Claxheugh escarpment, 1 wo miles west of Sunder-
land, Durham.

.Ki'yi'Dii \. MKI.'US'I u
" Ancylus ri/ili, Kirkliv " (<•/'. "Spat'' of Belinurus ti'echmamii (\Vooil\v.).

Anthracomya phillipsii, Will.). DIPLOPODA.
Anthracoinya minium (Ludvig). .1. /.-r/'/x, Euphoberia, sp,

var. scotlcct.

FAUNA I; ASSOCIATION.

lit

OSTRACODA.

Beyricliin, McCoy.
i'rt' or Ciji>ris.

INSECTA.

Lithomylacris kirkbyi, Woodward.

Phylomylacris iiniiiliilinides (Goldenberg) .
PISCES.

Rhizodopsis sauroides (Will.).

NOTTINGHAM AMI DKUCYSHiitE CoALFi tan. — Below the Top Hard Coal, Middle Coal
Measures (1911, Moysey, ' Geol. Mag-.' [oj, vol. viii, p. oUli); Shipley Manor clay-pit,
one and a half miles north of Ilkeston, Derbyshire.

ANNELIDA.

Spiriii'liifi, sp.
PELECYPODA.

Antl/i'iicniii i/n iituilinlnrin (Sow.).
<_'<trli(in!r<il<t iii/ii!lina (Sow.).
Naiaiti/i's iiiinliiijiiria (Row.).
OSTRACODA.

Beyricliin fii-fiinta .' (Bean).
Egtheria sp.

Lenin trigonioides, Moysey.
CRUSTACEA.

Preanaspides prxcursor, Woodw.
Arthropleura nriinitn. Jordan.

,, sp. nov., Moysey.

ARACHNIDA.
Cyclii.i. sp.

jiilniKiiiil, Woodw.
Beliiinriit lii-lluhts, Ivonig.

,, kn nif/ iniiii. <, Woodw.

liiinjii-djiililuf, Woodw.
,, sp.

Preftii'ii'liin iniilinix (Prestw.).
,, birtu'etll, Woodw.

rnliimtfita (Prestw.).

sp.

ilrrliii'iiaia, Woodw.

AEACHNIDA — (continued).

En ri/ji/f /•».< ini'ii/m'i/i. Woodw.

Scorpion, post-abdominal segment, Moysey.

Geraliiiitrn britannica, Pocock.

Eiibnt/iiia Imlli, Pocook.

Anthmcosiro fritschi, Pocock.

,, woodwardi, Pocock.

,, sp.

Protolycosa, sp.
INSECTA.

Gryptovenia iimi/ni'i/i, Bolton.
Ofthoeoatu splendens, Bolton.
Pierotinlin /ilii-ufnln, Bolton.
PISCES.

ElonicJithys, sp.

EGG CAPSULES OF FISHES (?).
Fdi/nliii i-rt'itittiifii, Moysey.
., of. i/i'iilntu, Zeiller.
l'iil;i'ii,ri/rif eni'liiniiirliiit (Schimper).
., A.7/c/.-/-ii/./.'.s- (Morris).
,, prfndeli, Lesq.

Vetaca/psula johnsoni (Kidstou).

,, i-iinjieri, Mackie and Crocker.

PLANT REMAINS.

Annnhiriu ; < 'nln ii/<n'lii<//is ; Sphenophyllum
Lf}i!<li>i'liylliiin ; Cn/ttiiiiten ; ferns.

LANTASHIUK COALFIELD.— Greyish-blue shales, 13o— ISO ft-t-t above the Royley
or Arley mine, the latter at the base of the .Middle Coal Measures; Sparth Bottoms,
Rochdale, Lancashire. I am indebted lor the following list to Mr. Walter
Baldwin, F.G.S., who, with Messrs. Sutcliffe, Parker, 1'latt and others, has devoted
years to the examination of these beds.

VERMES.

SpiTorbin (Spiroglyphus) .
BRACHIOPOIJA.

Linrjiiln, sp.

PELECYPODA.

( ',irboiiii-"l<i tn-iita (Sow.).
nilinatii (Sow.).
liinjiiln (Brown).

20 FOSSIL INSECTS OF THK BRITISH COAL AlKASUJKS.

PELECYPODA — (

NaiaJ ilf* modiolaris (Sow.).

ntrinuta (Sow.)
fl'jiiijatii (Hind).
Aiil/irfii-nii/i/ii williamsoni (Brown)

EUCRUSTACEA.

ihiiliis I'uiiperi, Huxley.
,, ( .' Anthrapaltemon) parkeri

( Woodw.).

Anthrapalxmon i/iv.winVi, Sailer.
Eurypterus, sp.
Cyclus jiili nitiiiii, Woodw.

li'in'lidti/iii jiarkri'l. Woodw.
ISiliniiriif luniihix (Martin).
,, konigianus, Woodw

baJdifini, Woodw.
,, longicaiidntus, Woodw.

testudineus (Womlw.).
ARACHNIDA.

Prestwichia /'//•/»•,'///. Woodw.

rntiinclata (Prestwich).

var. major, Woodw.
in in fir, Woodw.
„ (Eupi'oops ) (In /'.•»', Meek A. Wortlieu.

n/illii-ct.c (Piv!>l wich).
iiiii* ( Mii.-.niiin ) wardingleyi, Woodw.

AKACHNIHA (continui •/ .

Anthracoscorpio buthiformis, Pocock.
., sparthensis, Porock.

<1< nil in n r<i fiitcliffri. Woodw.
Anthracomartus Ii-Huliitim. Si-udder.

sp. 1 in Plait Coll.

>p. -2 in Platt Coll.
Anthracosiro ii-multi-nnli, Pocork.
Phalaiigioiarbvs tiilim-n/lit { Woodw.).
„ (ArchitaTbiif) niti/ml

Woodw.

DlPLOPODA.

Xylobius jilnffi, Woodw.
En^hi ill friii ffrnr. Sailer.

,, (irmiijrra (Baldwin)

r.'liimtit (Baldwin).
ii'fiinl ini nl! (Baldwin).
Arcliinliif. sp.
Acantherpestes major. Meek & Worthen.

V'V" "'''"•-'. Baldwin.
INSECTA.

S/ii/iijiti'i-ii niti-lifii'i. l>olton.
Mecynoptera tnln'i-i'iiltitii, Bolion.

PISCES.

Plaiysomus tenuistriatus, Traqnair.
Itlii-iuliipfi* zauriiiJes (Will.).

INCERT.H SEDIS.

Pain "i i/rix /n, ml'-li, Lesq.

SOUTH STAFFORDSHIRE COALFIKI.D. — Hinds between the " Hi'ooch " and "Thick "
coals; Tiptun, Dudley, and C'oseley.

EUCRUSTACEA.

EupJioberia fi'rn.r. Salter.

ARACHNIDA.

.1 nthracoscorpio buthiformis, Pococls
Geralinura britannica, Pocock.

<lr:i'iililiniiii* niii/lii'iix. Porork.

I''nfli-Hi:.il itlll'irnlil, Pm-iM-k.

Arrlin'i'iiii'lii in'i'li Hi nn, 1'ucoi-k.

Curciilioides im*lii-ii, 15iu-Klund.

I'lilini-lirril illlif''/!!!, I'orork.
/Vrs//'i) i, III ilrlrlfl, INirork.

Geraphrynus angulatus, Pocock.

liimli. I'ipi-iick.
„ tuberculatus, Pocock.

i 'i'/i>t/<iiii. Pocock.

,, liilfii-lln, I'lii-iirk.

A.RACHNIDA - irnnlnmri!).

Geraphrymis inn/ //.-7»s, Pocock.
Anthracomartm hit/:!!. IV-m-k.
jirii'xli, Pocock.
Anthracosiro woodwardi, Pocock.

., irilxr/ii, I'ocock.

Trigonotctrbits johnsoui, Pocock.

|NM:I i \

I ',i lieodii't v optera.

Pritvostict specttibilis, Bolton.
Brodict priscotincta, Scuddcr.

.. f,,ri;iln (Han. II. L
"Pteronepionites" /c/i//.--, Bolton.

iinilii/liiiis. IJolton.

<ll'l'lllll'IH'll I .' I Ill'll/lt. I Illltoll.

FAUNA I, ASSOCIATION. 21

INSECTA — (continued'). INSECT A — ( cmitinuecl).
Protorthoptera. Blattoidea — (emit inn fil).

Xvroptera obtusata, Bolton. Aplithurolilnttiiia rggintoni (Bolton).

Scalsebptera recta, Bolton. Arcliimylacrig incisa (Bolton).

Coselia pahniformis, Bolton. Plti/llnblnftn transversalis (Boltoii).

Blattoidea. LARVAL BLATTOID.

Aphthoroblattina johnsoni (Woodw.). Leptoblattina efilif, Woodw.

COALBROOKDALE COALFIELD, SHROPSHIRE. — Ironstone Nodules of the Pemiystone
Series.

EUCKUSTACEA. AKACHNIDA (roil till lied) .

EupJwberia fernr, Salter. CiircitJiuii.li>* anaticii. Buckland.

Anthrapalssmon (Apim) tlnliing (Prestw.). Eophrynus prestvici (Bucldand).

ARACHNIDA. DIPLOPODA.

Prestwichia anthrax (Prestw.). Acantherpestes lirmliei, Sod.

rotitndata (Prestw.) INSECTA.
,, Iriloliitniiles, Woodw. Lithogialit bromjinarti (Mautell).

SOUTH WALKS COALFIELD. — Shales in the neighbourhood of the Mynddislwyn
Vein, base of the Upper Coal Measures.

AUACHNIDA. INSECTA.

Aphantomartne areulofuf, Pocock. Aphthoroblattina mtlcntu (Bulton).

G-rieophonus onglienf. Pucuck. Orthomylacris lanceolata (Bolton).

Muiocercus celiicus, Pocock. Archimylacris hastata (Bolton).

K ri'iscJier/a r-errucvsa, Pocock. ,; tibovata (Bolton).

The arthropod association in the lists given is significant, for no other animals
enumerated are so readily water-borne as are insects. It may be assumed that
neither the more primitive arthropods, nor the insects, have been transported to
any great distance from their former habitat. Their preservation under similar
conditions supports the belief that their habits and habitats were the same, or
closely approximated to the same conditions of entombment.

The freedom of the deposits from comminuted carbonaceous matter, such as is
usually a chief constituent of the Coal Shales, may be accounted for by the beds
having been laid do\vn in quiet lagoons or swamp lakes, into which only the finer
mud particles and floating pinnules and (//'///•/.•* of coal-plants could pass and
accumulate. Such waters were probably fresh or brackish, shallow, and limited
in area. As we have indicated elsewhere (p. 10), it has been considered that the
larva1 of many of the Coal Measure insects Avere semi- or wholly aquatic, and if
such was the case, they would be more likely to be found in the deposits accumu-
lating in quiet waters than in others exposed to movement. The presence of
Mollusca and the lower orders of Artliropoda, with such forms as Palseoxyris, and
even fish-remains, can be accounted for by the existence of occasional or
permanent passages leading to open waters, such as river channels and the
open sea.

22 FOSSIL IXSECTS OF THE BRITISH COAL MEASURES.

A succession of sucli lagunal lakes or swamp pools might be seasonal features
along a depression which, in a wet season, formed a water-course.

The paucity of Mollusca is noteworthy, only one species (" (rnio") having been
recorded by Moysey, notwithstanding his careful search of material from the
Shipley clay-pit at Ilkeston, Derbyshire; while Kirkbv, and more recently
Trechmann and Woolacott have recorded Anthracomya jilii/Hi^ii, in addition
to " Ancijlnx riitti" (now known to be the larval form of A. phillipsii), from the
insect-bearing beds at Claxheugh, near Sunderland.

The deposits at Sparth Bottoms, Rochdale, Lancashire, are remarkable in that
they have yielded three species of Oarbonicola, four species of Naiadites, and one
species of Anthracomya.

In regard to the presence of Crustacea, Thomson (1804, 'Trans. Linn. Soc.
London,' Zoology [-], vol. vi, p. o), has shown that the recent Anaspides /</*///</ <//,r
lives in freshwater pools and lakes which are wholly cut off from the sea, and
Dr. H. Woodward (I'.XiS, ' Geol. Mag.' [5] vol. v, p. 080) has described an allied
form, Preanaspides prteeiirsor, from the Coal Measures of Shipley Clay-pit.
Further research may prove that not only Preaihisjiiilrs, but such forms as
Belinurus, Prestivichia, ISwypterus and Anthrapalsemon were also fresh-water in
habit, co-existing with insect larva? in the lagoons and swamp lakes.

Observations by the writer during the visit of the British Association to
Australia in 1914 bear on this point. While collecting in the " Bush country " at
St. Margaret's Bay, Western Australia, at Warburton, S. Australia, and else-
where, examination of almost every loose sheet of bark hanging to the gum trees
revealed a colony of scorpions, millipedes, spiders and cockroaches. In the
Australian localities mentioned true "bush" conditions prevailed, and seemed
much unlike those of low-lying swamps, such as are predicated for the Coal
Measure period. Subsequent experiences along the coast of North Queensland
modified these first impressions. It was found that notwithstanding the hot
tropical day, or perhaps by reason of it, the nights brought an extremely heavy
drw, so that it was impossible to move four yards in the jungle1 before the clothing
was running with water discharged from the leaves of the jungle plants. In a
short time after sunrise the jungle was dry again, but much of the moisture must
have been caught up under projections capable of resisting the penetration of the
sun's heat. In the "bush " country, slabs of bark may give conditions which I lie
arthropod fauna, find the most tolerable during the hot season. The Australian
winter with its heavy rains would more nearly accord with swamp conditions.
Long stretches of the coastline of Queensland, north of Brisbane, the country
along the llinchinbrooke Channel, and in the neighbourhood of Townsville, are
low-lying, and covered b\ dense mangrove swamps growing out into the sea and
completely hiding the outlets of rivers.

The conditions seem similar to those of the low-level country, swamp and

FAUNAL ASSOCIATION. 23

mud flat upon which the coal forests are believed to have flourished. During the
voyage along the coast, a landing was made at Lucinda Point, and an opportunity
was afforded of entering the swamp. Part of the swamp was awash on landing,
hut as the tide receded, it was possible to reach the shore. The latter was found
to be a flat shelving beach, consisting of fine marine sand on the seaward side,
passing shorewards into a fine tenacious clayey mud in which the mangroves grew.
The sand bore numerous remains of echini, mollusca and marine ili'lris, while the
mud below high-water mark and above it was penetrated in all directions by
the roots of mangroves, by the burrows of crabs, and by a burrowing gasteropod,
Telescopium fuse urn.

Masses of leaves, branches and other vegetable material were mixed with the
mud, the latter in some places forming irregular lumps and boulder-like masses
around the vegetable material. A broad shallow depression, with a few inches of
water over a thick bed of mud littered with leaves, led into the swamp. It was
evidently the bed of a stream during the wet season, remaining as a series of
disconnected pools at other times, or drying up.

The resemblance of the physical conditions to those accepted as dominant in
the Coal Measure period was so evident that special attention was given to it with
a view to discovering discrepancies ; yet, had the mangroves been replaced by
dense groves of gum trees with their arthropod fauna, the circumstances would
have been well-nigh identical. Had the faunal association of scorpions, millipedes,
spiders and cockroaches of \Vest Australia been transported to the swamp, their
remains would have become entombed in material not unlike that at Sparth
Bottoms or Coseley.

There is ample evidence to show that the coal plants grew on ground of this
character, and we know that the remains of fossil insects are associated with the
plant-remains.

The swamp conditions of the North Queensland coast reproduce with great
exactitude the presumed Coal Measure conditions, but lack the arthropod fauna
owing to the unsuitability of the vegetation.

Scudder had some such habitat and faunal association in mind when writing-

«~

his work " Archipolypoda : a Sub-ordinal Type of Spined Myriapods from the
Carboniferous Formation" ('Mem. Boston Soc. Nat. Hist.,' vol. iii, no. 5, pp.
143' — 182). Plate X of that work is described as "an attempted restoration of a
specimen of Acantherpestes major, Scd." The specimen is represented as leaving
the water, in which the hinder part of the body is still swimming by means of its
legs, while the fore part of the body is creeping up the trunk of a Lepidodendron
(L. erst it tint). On the trunk crawls a cockroach (Etoblattina inanona, Scd.) ; and a
broken stem of Galamites fixtil, Brong., lies partly fallen in a clump of Newopteris
lii'Acliii, Lesq.

We are, therefore, not without some justification in assuming that the fauna!

24 FOSSIL TXSKCTS OF TIIK BRITISH COAL MEASURES.

association which exists at Coseley, Sparth Bottoms, and elsewhere is not d in-
to the accident of deposition or transportation, but the natural result of the
conditions under which insect life was passed.

SYSTEMATIC INSCRIPTIONS.
Order I'AF^EOlUCTYOl'TKHA (Goldenberg), Haudlirsch.

1877. Goldeulierg, Die Fossileu Thiere aus der Steinkohlenformation von Saarbriickeu. Fauna

Sarsepoutana Fossilis, pt. ii, p. 8.
1900. Haiullirsch, Die Fossilen Insekten, p. <!1.

Slender insects with moderate-sized head and biting' jaws, simple antenna?,
and two pairs of equal and similarly shaped wings. The wing-venation is not
unlike the hypothetical tracheation of the primitive nymph worked out by
Comstock and Needham ('American Naturalist,' vol. xxxii, no. o~4, p. 85, fig. 4,
1898—99).

The wings could not be folded, being outstretched laterally in the position
of rest, and only moving in a vertical plane at right-angles to the body. The
thoracic segments are three in number, with wing-like pleurites in some cases
on the first segment. The abdomen has eleven segments, the eleven! li segment
bearing cerci. Legs all similar, and fitted for running.

Goldenberg did not define the characters of the Order, but included in it a
group of Palaeozoic insects which, while somewhat related to the existing forms
of Neuroptera, are sufficiently unlike to prevent their inclusion in the latter
Order.

llamllirsch defined the Order as a primitive generalised group, probablv the
ancestors of all later insects, and wholly confined to the Palaeozoic. He considered
that the larva? were predatory and aquatic, developing their wings gradually
without resting stages, and being in other respects similar to the imago. His
diagram of the primitive Palaeodictyopteroid wing shows a slight advance upon
the primitive nymph of Comstock and Xeedham, the primary tracheation being
increased bv the development of cross-nervures, uniting to form a meshwork.

Family DKTYONKCUIIM-:, Ilandlirsch.

lyOti. Haudlirsch, Die Fossilen Insekten, p. Ii;!.

lyl'J. Hundlirsch, Revision der Paliio/oisclit-n Insekten, p. 3.

Palaeozoic insects in which the wings possess a close reticulated neuration
between the principal veins, the latter strong and parallel over the first third of
the wing. Branches of the radial sector, median and cubit us few in number, and
strongly curved back to the inner margin.

Ilandlirsch regards this family as closely related to Microdictya.

DTCTYONEURA HIGGINSII. 25

Genus DICTYONEURA, Goldenberg.
1854. Dictyoneura, G-oldenlitTi;, Palseontographica, vol. iv, p. 33.

Wing three times as long as wide. Base narrow, outer margin curving
regularly backwards beyond the middle of the wing into the wing-apex. Inner
margin rounded, and joining the apex in a blunt curve. Subeosta straight,
reaching the costal border in the outer third of the wing. Radius simple, parallel
to subcosta, and giving off a radial sector with few branches. Hinder branch
of median and first branch of cubitus uniting to form a " cubito-median Y-vein."
Anal veins with few branches. Interstitial neuration irregularly reticulate.

Dictyoneura higginsii (Handlirsch). Plate I, fig. 1.

1871. " Neuropterous Insect-Wing," Higgins, Proc. Liverpool Nat. Field Club, vol. ii, p. 18, fig. 15.

1906. (Pcil;<oiU<'t>/i>jiteron) liiijyinsii, Hundlirsch, Foss. lusekt., p. 125, pi. xiii, fig. 6.

1917. (Dii-tyoiieiirnii ) l/iijijlimli. Holt 011, Quart. Journ. G-eol. Soc., vol. Ixxii. p. 46, pi. iii, fig. 2.

1919. Sherbornicllii liii/ijinsii, Handlirsch, Revis. Palnozoischeii Insekten. p. 25.

TI//IP. — Basal portion of a left wing in counterpart; Liverpool Museum.

Horizon tnnl Locality. — Middle Coal Measures; Ravenhead railway cutting,
near St. Helens, Lancashire.

lii'xi-i'ipfioit. — The wing-fragment lies on the surface of a greyish-purple
ironstone nodule, and has a length of 32 mm., with a greatest width of 22 mm.
The total length and width cannot be exactly estimated.

Handlirsch, who named the species, did not describe it, and it is doubtful
if he ever saw it. It was first described by the late Rev. H. H. Higgins thus:
"A second and smaller specimen of insect wing was obtained by myself .and
referred to the genus Gdnjdalis. Mr. J. P. G. Smith compared it with 7'Wr/ora.
A slight sketch of it was seen by Mr. F. Smith, of the British Museum, whom
it reminded of G'ri/Uiitii//in. Mr. Benj. Cooke, of Manchester, after a careful
examination, says, ' I believe the fossil represents the basal portion, about
one-third only, of the forewing of a Chnjsopa, Goldeneye, or Lace-wing fly, or
rather Nothochri/sa, separated from Chrysopn by Mr. McLachlan on account of
the manner in which the third cubital cell is divided.' This cell is remarkably
well shown in the fossil, and though I conld only judge from memory, I believe
it is sufficient to settle its relationship."

The "cubital cell" mentioned by Mr. Benj. Cooke is an elongated area
enclosed by a union of two of the main veins in the base of the wing. The
identity of the veins enclosing the " cell " will be considered later.

The costal border is preserved for a length of 29 mm., is strongly curved
proximally and less so distally. It would seem to have been directed somewhat
4

26 FOSSIL INSECTS OF TTTR BRITISH COAL MEASURES.

backwards in its course to the.1 wing-apex. The intercostal area is verv \viili1
at the base, and diminishes in diameter towards the wing-apex, lint it is doubtful
it' it reached the latter.

The subcosta is straight, but not vrell defined. The radius is fairlv, parallel
to the sulicosta and gives off the radial sector h>\v down in the base of the win--;
the radial sector comes off from the radius at an acute angle, and keeps parallel
with it as far as preserved. Its direction is such that it must have reached the
wing-apex. The median and cubital veins merit special consideration. In the
specimen three' veins occupy the position of the normal median and cubitus.
Of these, two either arise from a common root, or so clo>e together as to lie
indistinguishable, the third having a separate origin. The innermost of the three
sends off from near its base a forwardly directed twig, which joins the second
of the two outer veins.

It is necessai'y to resolve these veins into median and cubitus. To do this
we must consider recent research on living insects. Comstock and Xeedham
concluded that the primitive median vein was a four-branched structure; more
recently Tillyard. from further studies of the wing-venation of recent nymphs,
concludes that the primitive median had an initial dichotomy, of which the outer
branch divided up into four (" M " of Tillyard, ' Proc. Linn. Soc. N.S. Wales.'
vol. xliv, I'.U'.i, p. -V>1>), and the second branch remained simple. Comstock and
Xeedham concluded that the primitive cubitus was two-branched; Tillvard considers
it (Inf. fit., ]i. ;j.ri!)) as three-branched, the outer or lirst branch having a distal
forking into two feeble twigs, and an inner feeble branch which remains simple.
The primary cubital fork is situated near the base of the wing.

Tillyard's views are not very different from those of Comstock and Xeedham,
as he admits a basal fork of the cubitus, but adds a secondary forking of the end of
the outer branch.

Tillyard has studied the relation of the two veins in the various Orders, and
finds that the " posterior arculus " of Comstock, which is supposed to be a cross-
vein from the median to the cubitus, is, as shown in the fossil Order Paramecop-
tera, not a cross-vein, but a true branch of the median. The various stages b\-
which the connection becomes established and afterwards developed into a
combined vein from the point of union, are fully stated (Inf. <•//.. p.:!.")?), and the
compound vein is named the " cubito-median Y~V(-'in." An examination of the
conditions observable in I h'rh/uiK'iirn lii</</i iitii shows that the united veins and
their single-stemmed prolongation are in position and character identical with
Tillyard's " cubito-median Y-vein." The three veins of the specimen, therefore,
are the outer median vein, the inner median uniting witli the lirst cubital vein, to
I'oi-m a " cubito-median Y-vein," and lastly, the second (inner) branch of the cubitus.

Tillyard recognises the presence ol' the " cubito-median Y-vein " in the Permian
Order Paramecoptera, but its discovery in the Coal Measure genus l>!fli/<>iteur<i

ORTHOCOSTA SPLBNDENS. 27

shows that its origin is of niucli older date. The cubito-median Y-veiu lies close
to the outer median for n short distance, and then bends obliquely inwards to the
margin of the wing. The inner cubital vein is fairly straight, and widely spaced
along almost its whole length from the cubito-median Y-vein.

Six anal veins are present, all going obliquely to the inner margin, the second
and third alone forking. The interstitial neuration. which I have been able to
study better by immersing the specimen in water, consists of numerous irregular
cross-nervnres, occasionally uniting in crossing, or forming a loose mesh work,
especial I v between the median and cubital veins.

A-ftin'ttii-x. — AVith so small a wing-fragment, and with few wings of similar
character as guides, it is not possible to form an accurate idea of the whole wing.
The radius and radial sector entered the wing-apex, and the median-cubital
elements occupied the greater part of the inner margin of the wing.

The broad intercostal space and the course of the subcosta are very similar to
those ia. Poly creagr a tlnjanx, Handlirsch, but the general trend of the main veins,
and most of all the character of the interstitial neuration, are so suggestive of
I >irh/i>iit>iir<i lilx'lhilulili'x, Goldenberg, that 1 had referred the specimen to that
genus before Handlirsch's paper was published.

Handlirsch has now (1910) referred this species to a new genus, Sherbornietta,
giving the new name " for the sake of uniformity," but without diagnosis.

Family ORTHOCOSTID.E, Bolton.
191-2. liiilUni, guiivt. Jouru. Geol. Soc,, vol. l.xviii, p. 313.

\Yings with almost straight outer margin; inner margin well rounded; costa.
subcosta and radius closely approximated ; median, cubitus and anal veins
occupying nearly the whole of the inner half of the wing.

Genus ORTHOCOSTA, Bolton.

191-J. Orfli,,<-r,*t,-<. B»lt. in. Inc. fit., p. 313.

<-}eii<>i-i<: Characters. — Radial sector diverging from radius, and with few
divisions. Median with two branches united by a commissure. Cubitus stout,
forked near base, the outer branch forking twice, and the inner once. Anal area
elliptical. Interstitial neuration forming an open polygonal meshwork.

Orthocosta splendens, liolton. Plate I, fig. 2; Text-figure 1.

1912. n>ilir><;>siri splendens, liolton. Quavt, Joiirn. (icol. Soc., vol. kviii, p. 310. pi. xxxi. faVs. 1—3.

Ti/pe. — Incomplete wing, and impression; Moysey Collection, Museum of
Practical Geology, Jermyn Street (nos. :',(», -J-J-J and ;.50,223).

28 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

Ifnri-nn mid Locality. — Middle Coal Measures (below the Top Hard Coal I ;
Shipley Manor Claypit, llkeston, Derbyshire.

Specific Characters. — Radial sector reaching the inner half of the wing-tip.
Outer branch of median four or five times divided, inner branch simpler and
forked. Cubitns dividing low down, the outer branch the stronger, and each
doubly forked. Anal veins one or two in number, alternately branched.

Ih'xi-ri/itinii. — The species is founded on a wing-fragment, the apex, a portion
of the inner margin, and the base being incomplete. The total length is 84 mm..
the width oo mm. The complete wing must have had a length of at least 10(.) mm.,
and a width of oo — 40 mm. The whole insect had in all probability a span of
wing measuring 'J-Vo cm. ( 1 0 in. or more).

The outer third of the wing is differentiated from the rest by its uniform and
gentle convexity, and by the character of the costa, snbcosta, radius, and median
veins, which pass outwards towards the wing-apex in straight lines, and show no
trace of divisions until well beyond the middle of the wing, contrasting strongly

VIG. l.~ Orthocosta S/'/OM^OIS. liolu.n ; ivsturatiuii .i| winy, showing portiuii invsi-rvril and tin-
rliara.-!iT i>f tliu intrrstitial iii-uration. natural si/,c — MHilli- C.ial Pleasures (In-low the '\'"\*
Hani Cciiil) ; Shipley Manor ( 'laypit , Ilkr.stoii. 1 Irrl.yslmv. Moysi-y t 'olhvtion, Mus. Pi-art . < ici.l.
(noa :io.L'i'L' an.l :id,L'j:i).

with the areas occupied by the marginal divisions of the median, cubital and anal
veins. The inner two-thirds of the wing is marked by deep furrows, in which lie
the marginal branches of the median vein, and the whole of the cubital and anal
veins. The areas between any two veins in this region are markedly convex.
The differences in character of the outer and inner portions of the wing are empha-
sised by a line of fracture which fairly accurately separates the two. Its occur-
rence Miggest> that it has followed a nat ural line of weakness, the more delicate
inner part of the wing breaking a\\a\ from the outer stronger portion.

The costa, subcosta, radius and median are all well-developed veins, stout in
structure and standing out in relief. The first three retain this evidence of
-treiigth over two-thirds of their length, the median vein showing signs of
attenuation beyond the proximal third.

The general structure of the wing indicates considerable powers of Might.
The outer margin, of which only a portion is preserved, appears as a stout, slight l\
elevated and well-rounded ridge.

The subcosta agrees in general character with the costal remnant, and is

PTERONIDIID/K. 2'.i

straight to the wing-tip, parallel \vitli the outer margin, and not far removed from
it. Basally it appears to be united to the radius.

The radius gives off the radial sector at about Id nun. from its base, and then
passes out to the wing-tip, at no point being more than Id mm. from the subcosta.
The radial sector diverges widely from the radius, the t\vo enclosing along, narrow
triangular area. At 50 mm. beyond its origin it divides into two equal branches,
which diverge to a distance of -\ nan. and then become parallel. The direction of
the branches of the radial sector is such that they must have ended on (he inner
half of the wing-apex, the outer branch probably forking again before the wing-
apex was reached. The median vein consists of two brandies, the common origin
of which is not shown, owing to the absence of the base of the wing. To the
middle of the wing the two branches are parallel, and both pass beyond this
point before branching. The outer branch attains a length of 66 mm. before
branching, and then gives off four branches on its outer side, the last arising close
to the margin.

The second of the four branches bifurcates. The inner branch of the median
gives off one forward branch only, which forks into a feeble twig dying out in the
polygonal mesh work, and into a stronger division which reaches the margin. The
two branches of the median are united at the base of the wing by an oblique
commissural vein which comes off at an acute angle from the outer branch, and
passes obliquely to the inner branch.

The basal portion of the cubitns has been lost, so that its branching is not
readily determinate. The main vein seems to have divided near the base into
two equal branches, which curve down to the inner wing-margin, bifurcating
twice in each case before the margin is reached. The eight marginal twigs of the
cubitus thus produced enclose the middle third of the inner margin. The anal
area is wholly destroyed on the wing, and only a trace of one vein can be distin-
guished on the counterpart. This is a narrow deeply-sunk vein which gives off
feeble off-shoots on both sides. It diminishes in strength, so that the last portion
of its course can only be determined with difficulty. The anal area is comparatively
narrow and small, and exhibits few veins.

Family PTERONIDIIIU-:, Bolton (emend. Cockerell).
1912. Bolton, Quart. Jouru. (reel. Soc., vol. Ixviii, p. 314.

Wings long and narrow. Outer margin arcuate. Median and cubitns with
divergent branches.

The attempt to classify these wings is difficult. That they possess affinities
with the genus Polycreagra is certain, but they are more simple and more
Dictyoneurid in type. As Dr. Handlirsch observes, it is not possible to refer them
to the family Polycreagridas, and they must be taken as the type of a new family.

*0 FOSSIL IXSKCTS OK TIIK BRITISH COAL MKASTRKS.

Prof. T. U. A. Cockerel] lias pointed out to me that the familv name was
originally wrongly written " Pteronidia-," and that the generic name was ante-
dated by I'li'i-nnii'li.'n, Rolnver, I'.Ml. lie thinks that the ditTerenee is sufficient to
justify retention, although it is only a single letter ('Proc. U.S. Xat. Mus.,' vol. xlix.
p. -i<><), 1915). Ilandlirsch (' Revision Palao/.oischen Insekten,' I'.'l'.l, |i. \:>) has
made the same correction, hut seems unaware that Cockerel! forestalled him.

Genus PTERONIDIA, liolton.
I'.ML', Pfi:niiiiil!n, Bnllmi. (juurt. Joiirn. Gi.'ol. Soc.. vol. Ixviii. p. :!!:;.

Generic Characters. — Wings three times as long as wide. Radial sector with
five simple, backwardlv directed twigs. Median dividing low down into two
principal branches, and ending on the inner margin in five twigs. Cubitus stout,
divided into two principal branches, the foremost with two twigs, and the hinder
ending in four. Interstitial neuration of thin oblique nervures.

Pteronidia plicatula, Bolton. Plate I, fig. :! ; Text-figure ;2.

IlU'2. Pteronidia ylicatula, Button, <L>nart. .loimi. (icol. Soc , vol. Ixviii. p. 31:!, pi. xxxii, HL;'S. I- -It.

TI//II'. — Incomplete wing ami impression; Moysey C'ollection, .Museum of
Practical Geology, .lermyn Stivet (nos. 30,-JiU and .'iO/J-J-j).

FlQ. 2.— Pteronidia plicatula, 'Bolton; ivst<.rati.ui «f l''ft winy, natural sixt-. — II i'Mh' ( 'c'.il Measures
below Top Hani Coal); Shipley Manor Claypit, Ilkeston, Derbyshire. Moysey Collection, Mvis

i'racl. (icol i in is. ;J(),L'L']. ami ;!i).^.">).

l/iiri::<in <m<l Lun/l it //. — Middle Coal Measures (below the Top Hard Coal);
Shipley Manor Claypit, Ilkeston, Derbyshire.

N//rr/'//V I 'liiintrfrrx. Wings triangular in outline, with subacute tip, strongly
plicated. Radial sector dividing into five simple twigs. Median with two prin-
cipal branches and five marginal twigs. Culiitus wide-spaced; two principal
branches, the inner doubly forked.

/ V.-.T, •i/i/iiiii. — This is a long delicate wing, of which the outer marginal portion
and the base are missing. It was probably about- 7<> — "•"> mm. long, with a basal
width of L!<» nun., reduced to S mm. near the tip.

The outer margin must have been flatly convex, while the inner margin is

BYPBRMEGETHIDJE. :li

nearlv straight. The tip of the wing is acute. The costal and subcostal veins are
not seen. The radius is represented l>y a single forked vein, not shown in the
figures. The radial sector isunbranched until well beyond the middle of the wing,
lievond which it gives off four (possibly five) backward branches, which occupy
the greater part of the tip of the wing and a portion of the distal inner margin.

The median is represented by two branches, the outer dividing into two twigs
and the inner into three.

The marginal area occupied by the median is a little longer than that of the
radial sector, and the two extend over the distal half of the inner wing'-margin.

O O

The cubitus is a relatively short and stout vein, dividing into two main
branches, the foremost having two twigs, and the hinder dividing into four by a
double bifurcation. Owing to the wide divergence of its branches the cubital
area is larger than that of the median. The anal area was large, but the whole
surface has been pitted by attempts to clear the matrix, and as a result the
presence and character of the veins cannot be distinguished with certaintv. It is
possible that certain surface-indications are evidence of two simple widely
separated veins. The wing-surface exhibits a strong plication.

Viewed obliquely, the wing shows a series of ridges formed by the vein with
V-shaped intervening sulci, which only flatten out close to the wing-margin.
Springing from the principal veins is a close series of fine cross-nervures, obliquely
disposed in the direction of the wing-margin. The portion of the wing preserved
is sufficient to indicate that as a whole it was triangular in outline.

Family HYPERMKGETHIDJE, Handlirsch.

1906. Hancllirsch, Proc. U.S. National Museum, vol. xxix, p. 672.

Costa marginal, costal area broad ; radius simple, radial sector present ; the
median probablv dividing at the base into two or more main brandies, the first
of which may become united to the radius. Cubitus forked near the base, with its
branches widely spaced. Anal veins few, and anal area not exceeding one-third
of the inner margin.

Hancllirsch established this family for the inclusion of a gigantic wing of which
only the basal half is known. The total length of the whole wing was estimated
to be 120 mm., the basal half having a length of GO mm. The discovery in the
Coal Measures of Durham of a nearly similar wing adds to our knowledge of the
group, and enables the family to be defined with more accuracy than was at first-
possible.

The general assemblage of characters found in the Hypermegethida? is, I
believe, highly suggestive of the Protodonata, but no definite conclusions can be

formulated until the whole wing is known.

:V2 FOSSIL INSECTS OF THE BTUTTSTT COAL MEASURES.
Genus HYPERMEGETHES, Handlirscli.

I!""', Hi/i^riiifiirtlii'it, llMinllirsrli, I'ror. U.S. National Must-inn, vol. \\i\. |». r.7'J.

Generic Characters. — Costal border feebly convex, snbcosta mid radius close
together along tin- greater part of tlieir length. Cubitns composed of two.
possibly more, parallel ami simply forked brandies, divided near the point of
origin. Anal veins few ; interstitial nenration of tine, irregularly anastomosing
nervures.

Hypermegeth.es northumbriae, Bolton. 1'late I. tig. I ; Text-figure :'>.

U'l". Hypermegethes n<irtliinnlri:r. Bolion. Quart. .tom-n. (leol. Soc., vol. Ixxii. p. 55, pi. iv, figs. 2

Mini :i, ;iinl woodcut in l(jxt.

TI//H'. — Portion of the basal half of a left wing, in counterpart; British
Museum (no. In. 1 s.'dl.

FIG. rt. — Hypermi gethes northumbrix, Bolton ; suggested outline of left wing, restoiv.1. shnwin

preserved, slightly less than luitui-il size — CVil Measures (sliale aliovc tin- ( 'IM\V I'.ial); Phoenix
Brickwiivks, Craw.-rook, ])urhain. Bvit. Mus. (no. In. 1S52-1'.

Horizon iniiJ L<ii'<iHh/.— (\)&] Measures (shale above the Crow Coal); Phoenix
Brickworks, Crawcrook, Din-ham.

X/jfC///c < 'luirnrh'i'x. — Costal area wide, and crossed by an irregular mesliwork
of small veins. Median united witli the radial sector basally, and giving off an
inner branch which unites with the eubitus. ('ubitus with a short stem Forking
into two equal and widely separated branches. Anal veins simple, and widely
spaced.

Description. — The fragment is little more than a third of the whole wing, and
its characters can only be determined with ditliculty. The two halves of the
nodule do not coincide, and the outline drawing of the wing is buili up Iroin
details supplied by both halves. The inner margin, anal aiea and base of the
wing have been lost, so that a little less than two-thirds of the outer portion ot
the basal half of the wing is present.

The portion of wing remaining being <i;'> mm. long, with a depth ol ol mm. at
its widest purl, the whole \\ing must have had a length of about I'-'ii mm., or

HYPERMEGETHES NORTHUMBRLE.

5 in., and the whole insect a span of wing of nearly 11 in. The wing-fragment
shows the basal portions of the costa, snbcosta, ratlins, radial sector, median, and
cubitus, and possibly a trace of an anal vein.

Much of the finer detail of the wing is not seen until the fossil is immersed in
water — a mode of treatment suggested to me by Dr. F. A. Bather, F.R.S., who had
previously photographed the wing in this manner, and brought out details of
which, by ordinary methods, I could not find a trace.

The costa is moderately convex from its base to a distance of about 30 mm.,
beyond which it becomes straight. Separated from the outer or costal border by
a wide area basally is the subcosta, an extremely feeble and hardly distinguishable
vein. It passes straight to the outer margin some distance beyond the middle of
the wing. The radius arises close to the subcosta, and is parallel with it. It gives
off two branches posteriorly, the proximal branch passing obliquely towards the
inner side of the wing-apex, while the second or distal branch arises from the
radius a little further out, and keeps parallel with it.

I had formerly considered the proximal branch of the radius to be the radial
sector, and the distal a branch of the radius ; but as the interstitial neuration now
shows the specimen to be closely related to Hypermegethes schucherti, the proximal
branching vein must be regarded as the main stem of the median which has entered
into union with the radius, and the distal branch as the radial sector.

Regarding the proximal offshoot of the radius as the median vein, it diverges
widely from the radius, giving off a forward twig parallel with the radial sector,
and then continues inwards and unites with the next vein, separating again a little
further out on the inner side. A comparison of this assumed median vein with
that of H. scltuclterti, Handl., is instructive. In the latter species the branching
of the main stem of the median arises nearer the base of the wing than does the
branching off of the radial sector. It is therefore somewhat in the position of
the starting-point of the median vein as a free structure in the present wing. The
median vein of //. xi-hnchn-ti, Handl., has, however, no union with the radius, the
main stem running out towards the wing-apex parallel with the inner branch of
the radius, giving off a backward branch of the cubitus, which passes obliquely
inwards and unites with the anterior branch. In the middle of its length it gives
origin to a twig running parallel with the main stem, and midway between
that vein and the cubitus. The course of the median vein in the specimen
here described is exactly similar to that of the branch of the median vein in
//. sclntcherti, Handl., except that the inner branch not only unites with the
cubitus, but crosses it.

The condition may be summarised by saying that, in H. schucherti, Handl., the
median vein is entirely free and gives off an inner branch dividing into two twigs,
of which the inner unites with the cubitus. In the specimen here described the
median is united with the radius for some little distance, becoming free before the

34 FOSSIL INSECTS OF THE' BRITISH COAL MEASURES.

radial sector is reached, and then forking, the inner twig uniting with and crossing
the cubitns.

The cubitns has a short, stout basal stem, forking into two equal and somewhat
widely separated branches, the outer uniting with the inner branch of the median.
The inner branch is forked just before the broken edge of the wing is reached.
Nearer the inner margin of the wing are traces of two other veins. The first has
its basal portion missing, and follows a course parallel with the inner branch of
the cubitus. It is strongly forked. The remaining vein is represented by three
detached fragments. If Handlirsch's interpretation of the wing of 11. scliucln rll
is followed, we should regard both these veins as anal. I am, however, of opinion
that, while the innermost fragmentary vein may be anal, the forked vein by its
position, its forked character and stoutness, must be regarded as a portion of the
cubitus. I am likewise of opinion that the first, and possibly the second, of the
veins marked as anal in Handlirsch's figure of II. schucherti, ought also to be
classed as cubital. Both in II. schucherti, and in this specimen, the anal area
would have an enormous development and occupy most of the wing-margin, if the
veins alluded to were wholly anal in character. I feel sure that, if the vein nearest
the cubital had been better preserved, it would be found branching off from the
cubitus.

The interstitial neuration is typically that of Hypermegethes. The intercostal
area is filled with an irregular mesh work of fine nervures, with a tendency on the
outer and inner sides to a transverse arrangement. Between the median and the
cubital veins the interstitial neuration consists of short, straight and transverse
nervures, and feeble traces of similar nervures can be seen in the median area.
The cubital area is filled with a meshed neuration, larger and more regular than
that of the intercostal area, and this seems to continue into the anal area.

Family CKYPTUVENIIILI;, Bolton.
191:2. Bolton, Quart. Journ. G-eul. Sor., vol. Ixviii, p. 31.x

short and broad. Apex rounded, costal vein marginal ; subcosta feeble,
and extending to near apex. Radius simple; radial sector and median with few
divisions; cubitus with two main branches.

Among the insect-remains discovered by the late Dr. L. Moysey at the Shipley
Manor Clavpit is a small wing, 1 (.1 mm. long, unlike anv previously known. It
is Ivpieallv I'aliBodictyopterons, and agrees remarkably well with Dr. Handlirsch's
I vpe-lignre ('Mittli. Geol. Gresell. \Vien,' vol. iii. 1'JlO, p. ;"><>5, fig. 1). ll differs from
that form in I lie greal er division of the cubitns, which ends in five- twigs instead of
t hree. Tin' greatest dept h of the wing was also, in all probability, nearer the base
than in his figure. With the <jvnus Atliymodictyn, llandlirsch (' Amer. Journ. Sri.'

CRYPTOVENIA MOYSEYT. 35

[4], vol. xxxi, 1911, p. '2\)S), the relationship is even closer, as in that genus the
costa and subcosta are close together, the radial sector arises lo\v down and is
divergent from the radius, the median is a simple vein ending in three branches,
while the cubitus is almost identical in its divisions, the difference being that the
first forking arises at a higher point, than that in Athymodicti/a [HI mi, Hand!., and
that the inner simple ramus of the outer branch comes off a little below the middle
of the wing. The anal veins number four in J. /mmt as against five in the wing
under consideration. The wings are almost equal in size. It is not possible,
however, to refer this wing to Athymodictya owing to the character of the interstitial
neuration. In Athymodictya this is reticulated, while in the British wing the
interstitial neuration is apparently made up of transverse nervures. If this point
can be clearly determined, the wing may have affinities with the Homoiopterida1
or with the Litliomantida,1, but it cannot be referred to either of these families.
Neither Dr. Handlirsch nor I could satisfactorily refer the wing to any known
family, and 1 therefore founded the family Cryptoveniidse to receive it.

Genus CRYPTOVENIA, Bolton.
1912. Cryptovenia, Bolton, Quart. Juuru. Geol. Soc., vol. Ixviii, p. 315.

< '/ntnir/i'rx. — Wings twice as long as wide. Median, cubitus and anal
veins curving sharply inwards. Interstitial neuration of straight cross-nervures.

Cryptovenia moyseyi, Bolton. Plate II, fig. 1 ; Text-figure 4.
1012. Cryptovenia moyseyi, Boltou, lor. clt., p. 315, pi. xxxii, figs. 4 — 6.

Ti/[>r. — Incomplete wing, in counterpart; Moysey Collection, Museum of
Practical Geology, Jermyn Street (nos. 30,226 and 30,227).

Horhon <unl Lnr/i/iti/. — Middle Coal Measures (below the top Hard Coal);
Shipley Manor Claypit, Ilkeston, Derbyshire.

Xy<(v//?'c Characters. — Costa feebly curved distally, subcosta reaching outer
margin near wing-apex. Radius close to and parallel with the subcosta; radial
sector arising from radius in basal half of the wing, the five branches occupying
the inner halt' of the wing-tip. Median with the first branch undivided, and the
second forking twice. Cubital vein strongly arcuate, two-branched, and ending in
five twigs. Anal veins five or more in number, and curving sharply inwards.
Wing plicated, and the interstitial neuration obscured by a mass of wrinkles.

Description. — A small delicate wing, incomplete at the base, which must have
been very narrow and with the main veins crowded together. The total length of
wing preserved is 1(> mm., and its maximum diameter, in the cublto-anal region,
is 8 mm.

36 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

The costal vein is stout and raised above the general level of the wing in the
outer third of its length, where it curves backwards into the wing-tip. The
subcostal is a, weaker vein, parallel with and very close to the costa, dying out or
joining the outer margin at the apex. The radius is a powerful undivided vein,
parallel with the subcosta and reaching the middle of the wing-apex. The radial
sector is given off about the middle of the length of the wing and forks into two
equal branches, which again fork before reaching the margin ; the outermost branch
of the second forks again and divides just before the wing-apex is reached. The

FIG. -4. — Cryptoreiiia moyseyi, Bolton ; restoration of li-ft wing reconstructed from the \\ inn'-ti'iiyinout
and counter-impression^ enlarged two-and-a-half times. — Mi«Mli' <_'oal Measure^ tin-low tin: Top
Hard Coal) ; Shipley Manor Claypit, Ilkiv-tcni. IvHiyriljiiv. .Mi>\^rv t'ollivtiuii. Jliii- 1'ract. (.leol.
(nos. 30,1'L'li and :{O,^27.)

inner half of the wing-apex is occupied by the five branches of the radial sector.
The median vein is a comparatively simple structure, forking low down below the
middle of the wing into two nearly equal branches. The outer branch remains
undivided, and gently curves to the inner margin. The inner branch divides
twice, first at a point near the middle of the wing, and again before the margin
is reached. The median vein therefore ends on the margin in four twigs, three
members of which arise from the inner of the two main branches.

The cubital vein is strongly arcuate, dividing near the base into two branches,
the outer forking twice and the inner once. The cubital vein therefore ends in five
twigs.

Family MECYNOPTERID^E, Handlirsch.
1906. Handlirsch, Die Possilen Insekten, p. Sii.

Handlirsch established this family to receive a large win»- (nun the Middle
Upper Carboniferous of Belgium, and placed it between the families llypenne-
gethidae and Lithomantidae. The characters of the familv are based mi those <>f
the type-species, Mecynoptera xfi/nnliiln, Ilandl.

Genus MECYNOPTERA, Handlirsch.

I '.ml.. Mecynoptera, ILnnllirscli, Mrin. Mus. \i«\-. Hist. Nut. Uel^., vol. iii, ]>. 7.
I '.MM;. Mecynoptera, llanillirsrli, liir Ko.xsilrn luselcten, |>. SL!

MEGYNOPTERA TUBERCULATA.

37

ii: Characters. — Wing three to four times as long as broad, veins of costal
region specially compact, and thickened basally. Costa, subcosta and radius
closely approximated and tuberculated. Radial sector and median well developed,
and occupying a considerable portion of the wing-surface. Cubit us small. Inter-
stitial neuration of transverse nervures in junction areas of principal veins, and
with a meshwork further out.

Mecynoptera tuberculata, sp. nov. Plate II, fig. 2 ; Text-figures 5 and <>.
1!M1. Stenndictija Inbatu, Baldwin (error e), Cleol. Mag. [5], vol. viii, p. 75.

Type. — Portions of two fore-wings, and the cubito-anal portion of a hind-wing
contained in a nodule of ironstone, 3 in. long, and 1^ in. wide; British Museum
(no. In. 18,576).

Horizon and Locality. — Middle Coal Measures (grey-blue shales at 135 — 180
feet above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire.

Specific Characters. — Principal veins thickened at base and finely tuberculated.
Costal margin almost straight, much thickened at base, and covered with a fine

FIG. 5. Fio. 0.

FIG. 5. — Mecijnoptci'c tiiberrirfutn. sp. nov. : diagram of remains of the two fore wings, and cubito-anal
portion of a hind-wing (the suli-costa appearing as a fine line close to the radius, and sending off
three branches to the costal margin), enlarged one-and-a-half times. — Middle Coal Measures
(above the Eoyley or Arley Mine) ; Sparth Bottoms, Rochdale, Lancashire. Brit. Mus. (no
In. 18,570).

Fin. (i. — Mecynoptera. tubei-ciihita, sp. nov. ; diagram of fragmentary impression of the two fore- wings,
enlarged one-and-a-half times. — Middle Coal Measures (above the Eoyley or Arley Mine) ; Sparth
Bottoms, Rochdale, Lancashire. Brit. Mus. (no. In. 18,570).

tuberculation. Subcosta sunken, close to costa, and extending to the wing-apex.
Radius close to costa, tuberculated, and elevated. Radial sector arising low down,
much branched. Median Avith two main branches, the first simple, the second
with three twigs. Cubital vein small.

Description. — The type-specimen comprises remnants of the two fore-wings.
60 mm. long and 18 mm. wide, and a portion of the cubital-anal area of a single
hind-wing in one ironstone nodule. A similar ironstone nodule from the same
horizon and locality shows the radius and radial sector areas of the two wings of a
second insect.

38 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

AVhen complete, the wings must have had a length of <>.j — 70 mm. They were
originally identified with those of <s'/f;/ <»//>///</ Inlmln, Brongniart, and are quoted as
such in the faunal lists of the Sparth Bottoms deposits, but with no detailed
description or figures.

\\ hen the specimens were tirst examined they presented an apparently anoma-
lous union or suppression of certain (if the principal veins. Careful development
has since shown that what was formerly considered to he the outer or costal
margin of the wings was really the thickened and tuberculated radius vein, and
that the outer margin, with the costa and subcosta, had been hidden under the
matrix. A little of the outer marginal costa and the subcosta are now uncovered,
and the wing-structure is therefore proved to be of a normal type.

The three wings in the larger, more nearly complete specimen are superposed
and fragmentary. The uppermost fore-wing is represented by two portions, one
having a little of the outer or costal margin, the subcosta, and the greater part of
the radius and radial sector. The second part consists of the median, and a
portion of the cubitus. The second fragment is displaced backwards, allowing
the radius and radial sector of the second fore-wing to be seen. The median-
cubital area of the hind-wing is very thin and closely pressed on the rest, but the
course of the veins is clearly discernible, and by their sharp inward turn indicates
that the hind-wings were much broader than the fore-wings.

Although the wings are thus broken up, displaced, and superposed, the parts
.missing in the one wing are present in the other, and it is possible to reconstruct
their general character. The outer margin is almost straight, gently rounded into
the base, and into the wing-apex. The subcosta is weak, and lies in a deep sulcus,
extending into the apex of the wing. The radius is a little elevated, thickened,
tuberculated in the basal third, and in close proximity to the subcosta] vein. The
radius is a strong vein, raised above the level of the rest of the outer margin of
the wing, so that when the latter was hidden, it nafurallv appeared to be the outer
marginal costal vein. It is much thickened in its basal third, tuberculated, and
remains parallel with the subcosta in the apex of the wing.

The wing-space taken up by these three veins is very small, and their close
approximation and the coriaceous thickening of the costa and radius serve to give a
considerable degree of strength and rigidity to the outer margin of the wing. The
radial sector passes straight outwards to the apex of the wing, giving off three inner
branches in one wing and two in the other, the first branch only forking. The first,
branch soon forks, the outer fork dividing into two twigs, and I lie inner into three
upon the distal inner half of the wing-apex. The median is a large vein, occupying
the greater part of the inner half of the wing. It divides at its point of origin into
an outer undivided branch which traverses the middle of the wing, and an inner
branch which is widely separated from its fellow, and gives off three twigs to the
wing-margin, the lii>( forking in the middle of its length. Cubital veins are rcpre-

MECYNOPTERA TUBERCTJLATA. -°.9

sented by two undivided elements which are directed obliquely to the margin.
The inner margin of the wing- is more curved than the outer, and merges into the
apex. The interstitial neuration is remarkable. BetAveen the costa, subcosta and
radius it consists of short stout nervures crossing the areas somewhat obliquely,
and in some cases arranged in V-shape. The area between the radius and radial
sector is crossed by a numerous S-shaped scries of nervures, which are joined up
into a meshwork in the outer or radial half of the area. The area itself is very
wide, and only equalled by that separating the two branches of the median.

The radial sector and the median occupy the greater part of the wing-
surface.

The remaining areas are crossed by transverse nervures near the junction of
branches with the principal veins, and further out by nervures which are joined up
by zig-zag longitudinal branches which occasionally enclose polygonal cells. The
interstitial neuration is very well developed, and must have added materially to
the strength of the wings.

Afinitiefi. — The close approximation of the costa, subcosta and radius, and the
coriaceous thickening and tuberculations of the principal veins, are characters
which may rank as of generic importance. Less distinctive, but also characteristic,
is the interstitial neural ion of the transverse nervures and mesh work, with its
minute monilation.

The characters of the genus !\lt>ci/>i«i>ter<i are in closest agreement with those
of this insect. There is the same approximation of the costa, subcosta and radius,
the radial sector is widely spaced from the latter, and the interstitial neuration
consists of transverse nervures near the junctions of the principal veins and their
branches, and of a meshwork further out. The general outline of the wing is also
the same.

M. splencliilit, Handlirsch, throws light on one point which had proved a
difficulty in the determination of the wings. It has the radial sector of large
size and much branched, the divisions occupying all the inner half of the apex of
the wing, and extending out on the inner margin into the area usually occupied by
the outer branches of the median. Prior to noting this feature in M. splendida,
1 had formed the opinion that the first division of the radial sector was the median,
which had united with the radial sector and the radius.

The broken condition of the Rochdale wings prevents absolute determination
of this point, but by analogy I conclude that the whole vein which branches off
from the radius basally is the radial sector, and the next vein the median. This
conclusion also removes the insect from the neighbourhood of jfflnigmatodes tlantdsi,
Handlirsch, a wing discovered in the Coal Measures of Mazon Creek, near Morris,
Illinois, U.S.A., which possesses the same character of interstitial neuration, but
with the divisions of the median vein stretching out to the wing-apex, and the
costa, subcosta and radius more widely separated.

40 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

Unless it can be shown by the discovery of a whole wing that, the reconstruction
now attempted is faulty, the balance of evidence is in favour of the provisional
reference of the Sparth Bottoms wing to the genus Mecynoptera.

Family INPKRT.E SKDIS.

Genus PAL^OMANTIS, Holton.
1917. Paln'oinantis, Boltori, Quart. Jouru. Geol. Sue , vol. Ixxii, p. 52.

< u'lii'i-ic Characters. — Wings short, twice as long as broad. Apex well rounded.
Radius, radial sector and median all powerful veins with few divisions. Median
with two main branches. Anal veins directed almost straight inwards. Interstitial
neuration forming an irregular meshwork.

Palaeomantis macroptera, Bolton. Plate II, tig. 3; Text-figures 7 and S.

1871. "Wing of large insect," Higgins, Fres. Add. Liverpool Naturalists' Field Club, vol. ii, p. 18.
11U7. Puln'uiiiaittlg ntucrofitei-a, Boltou, (juart. Jouru. C4eol. Sor., vol. Ixxii, p. 48. pi. iii, figs. 3 — 4,
text -figs. 2 — 3.

Ti/jie. — Remains of two wings in nodule; Liverpool Museum.

Horizon and Locality.- — Middle Coal Measures; Ravenhead railway cutting,
near St. Helens, Lancashire.

tf/iecijic Clmractfrx. — Divisions of radial sector occupying almost the whole tip
of the wing. Divisions of median and cubital veins occupying distal two-thirds of
inner margin.

Description. — The larger half of the nodule has lost a portion which contained
the tip of the right wing. The proximal third of the left wing remains, with its
dorsal surface uppermost, and its ventral surface closely applied to that of the
right wing. It is evident that the whole of the two wings was contained in the
nodule, but, when the latter was split open, a thin film of ironstone carried away
the middle and distal portions of the left wing.

One unusual feature in the position of the wings is that they lie with their
ventral surfaces apposed. To bring them into this position one must have become
bent under the body, instead of falling sideways across the thorax of the insect.
The body of the insect would thus, if the wings still remained attached, lie between
the two. No trace of the bodv can be seen, but the left wing has a deep inward
flexure, such as it would naturally acquire if the body had been carried round with
the right wing and pushed into the anal area ol the left. wing. Had the bodv been
carried round in this wav the right wing would not coincide m position with the
left, but be thrust further out, which is actually the case, the outward displacement

PAI^EOMANTIS MACROPTERA.

41

of the right wing- as compared with that of the left being at least 20 nun. The
wings are remarkably wide — a feature which we usually associate with hind-wings.

The outer third of each wing is supported by strong and fairly rigid veins,
becoming more slender as they pass towards the wing-tip. The inner margins of
the wings are more membranous, and were evidently lacking in a rigidity equal to
that of the outer margins.

A little more than the proximal third of the hi't icing is present and in good
preservation. It is 42 mm. long and about the same in width. The impression
on the other half of the nodule shows apparently all but the wing-apex. The main
stems of the costa, subcosta, radius and median are all stout, and in relief on the
wing-surface. The cubital and anal veins are but half the thickness of the former and
lie in shallow grooves. The costal vein forms the outer margin of the wing, which
is slightly convex forwards, and slopes gradually into the apex. The subcosta is

FIG. 7.

FIG. 7. — Palssomantis niai'i-niilem, Bolton ; restoration of left wing, witli restored outline, natural size.

Middle C.ial Measures; Raveuhead Railway Cutting, near St. Helens, Lancashire. Liverpool

Museum.
Km. 8. — Palieomantis macroptei'n, Bolton; restoration of right wing, natural size. — Middle Coal

Measures ; Ravenhead Railway Cutting, near St. Helens, Lancashire. Liverpool Museum.

separated from the outer margin by a costal area 4 mm. broad. It passes almost
in a straight line to the outer edge of the wing-apex, and is joined to the costa by
a series of oblique transverse branches nearly parallel with each other. The radius
arises close to the base of the subcosta, and diverges a little from it in its course
to the apex, forking once just before the broken edge of the wing is reached. The
main stem of the radial sector probably arises from the radius close to the base of
the latter, but the exact point cannot be determined, owing to the base of the wing
being broken a\vay. The radial sector forks three times, and each of the resultant
twigs again divides on the wing-apex. The median vein, which is stout basally.
divides very 1<>\\- down into two branches, the outer of which has a feeble forking
near the inner margin, while the inner branch forks three times. The cubitus is
a much weaker vein than its fellows, and divides near its point of origin into two
branches. The first branch forks once and the second twice. Seven anal veins
are present, all undivided except the third, which forks twice. The interstitial
6

42 FOSSIL INSECTS OF THE BRITISH COAT, .MEASURES.

neuration is in the form of a meshwork, except in the intercostal area, where are
the transverse eross-nervures already noted.

Much less of the rii/lil irim/ remains than of its fellow, lint sufficient is present
to show that the nenration was not quite the same. The median ends in five
divisions in place- of six. and the eubitns has seven final branches instead of six.
The greatest width of the wing is along a line drawn from the outer margin to the
middle of the culiital area on the inner margin, the width being 40 mm. The
absence of the apex in each wing is unfortunate, as il renders the outline of the
whole wing a little uncertain. The shape of the nodule indicates that very little
of the wings is missing, if thev were wholly included in the nodule, as seems
probable. The somewhat semicircular inner wing-margin and the short wide
wings indicate a. broadly rounded wing-apex.

. I /////// ii'ft.-— The interstitial neuration is much like that of. Hypermegetlte?t. The
great width of the wings is a character usually associated with the hind-wings of
members of the taimlv l.ithomantuhe, although in this case the wings lack the
distal attenuation noticed in the hind-wings of that fa mil v. With Titanodictya
i/ii'i/inJn, Scudder, there is a close relationship, both in the general character of
the main veins, tlie interstitial meshwork in all areas other than intercostal and
subcosta-radial, and in the presence of the same oblique cross-nervnres in the
intercostal area in these wings.

The true systematic position of the genus seems to lie between the Lithoman-
tid;e and the genus Titanodictya, and closest to the latter. Because of the greater
development of a meshwork neuration between the mam veins, and the limitation of
cross-nervures to the intercostal area. I regard this genus as more primitive than
any of the Lithomantidie, but closely related thereto.

Family LITUO.MAXTID.I:, Handlirsch.
I'.'IM;. Hiuullirsc'li, Prw. U.S. National Mus., vol. xxi.\, ]>. 073; ulso Die Fossilen Insekteu, [>. s-j.

This family is closely allied, by wing-structure, to the Dictyoneuridse. The
branching of the mam veins has proceeded further than in the DictyoneuridsB, and
the body, where it has been preserved, shows striking differences. The family is
represented iu the Coal Measures of Great Britain and North America and in the
Upper Carboniferous of continental Europe.

Cem is LITHOMANTIS, Woodward.

I'^fii. Lithomantis, II. Wood wai'd, Quart, .loiini. <!col. Sue., vol. \x.\ii, [>. <!0.

it/'in'i'ii- Characters.- l/ar^e insects \\ith iwo pairs of living wings; the hinder
pair double the width of the anterior pair. I'rot horax produced into a centra]

UTIIO.MAXTIS CARBONABIUS. |:i

rostral-like process, acutely pointed, and expanded into two lateral bulbous lobes.
Thorax wide. The interstitial neuration consists usually of stout transverse
nervures, occasionally forking or in a loose meshwork.

Lithomantis carbonarius, \Vood\vard. Plate II, fig. 4; Text-figure '.».

1876. Litlioniantis carboiiaring, Woodward, lac. cit., p. (id. pi. ix, tig. 1.

1893. Lithoniiiiifia I'lirli'iuiiritts. Brongniurt, Faune Eutoiu. Terr. Prim., p. 489, tin'.

190(i. Llt/itiitiiiiiHx carbonari mi. Woodward, (j-eol. Mag. [_5], vol. iii, p. '25, tig. o (no/i fig. 1).

190(3. Lithomantis c<irbon<.trin>t, Haiullirsch, Die Fossileii Iiisekten, p. 88.

— Portions of the fore- and hind-wings, with prothorax, and a left anterior
leg in nodule; British Museum (no. I. 8118).

//<//•/';.«// i/inl Lin'iilih/. — Coal Measures; Scotland.

X[>''<-'(fi<- Characters, — Hind pair of wiu^s double the width of the fore pair.
Outer wing-margin straight, costa and subco.sta close together; radius a weak
vein, giving off the radial sector far out. Median small. Cubitus a powerful
vein with numerous widely spaced branches. Anal veins six or more.

FIG. 9. — Lilhomantis carbonarius, "Woo&waxA; diagram of neuratum of tlic li't't fmv- :uid Mini-wings,
natural size. — CD;I! Mx-usures : Scotland. Brit. Mus (no I. SUM.

. — The type-specimen lies on the surface of one half of an ironstone
nodule, the counterpart being lost. It was obtained bv Mr. Edward Charlesworth
from the Coal Measures of Scotland, but the locality and hori/on are not known.

The remains consist of portions of two pairs of wing's, those of the left side
being most nearly complete. In no case is the apex of the wing or the inner
margin preserved. -Lying in front of the wings are a pair of very large convex
lobes which Dr. Woodward regards as part of the prothorax, and in front of these
is a roughly quadrangular structure prolonged forward in the middle line into a
styliform process or rostrum. Woodward describes the latter as " the small head

44 KOSSI], IXSKCTS OK TIIK UIMTIS1I COAL M KASf I! KS

with its eyes," hut 110 definite truer of the latter are observable. rriie " head " is
small, not more than 7 mm. wide, and hetweeii 5—6 mm. lony1 at the .sides ;
medially it is prolonged into the rostral process, which is ahont 8—9 mm. long
and ending in a sharp point. I am unable to distinguish any dividing line between
the "head" ami the prothorax; on the oilier hand, the margin of the latter is
continuous with and inseparable from it. The marking which "Woodward ha<
regarded as representing eyes is, I believe, the thickened margin. lam of opinion
that the whole structure is wholly prothorax, and that the head lies concealed
beneath. The main mass of the prothorax is :>() mm. wide, with a flattened margin,
best seen on the left side. Within the fiat margins rise two low dome-shaped
lobes separated by a wide hollow in front, their margins meeting in an obtuse
angle posteriorly. The median edge of each lobe dips sharply into a wide median
hollow, and from each of these edges arises a series of veins, which spread out to
the lateral margins of the lobes. The areas between the veins are occupied by a
fine mesh work of smaller veins.

A trace of the mesothorax is shown between the bases of the fore-wings as a
slight transverse liar, a small rounded tubercle lying in front and a little to the
left of the middle line.

The left fore- and hind-wings are the most nearlv perfect . t he hind-wing being
•">'• mm. long, with a greatest width of :>0 mm. The fore-wing is a little shorter
and narrower. The outer margin appears to have been straight, and the costa
and subcosta closely approximated. Traces of both veins are present. The radius
is a straight thin vein not far removed from the subcosta along its whole length,
and giving off the radial sector beyond the middle of the wing. The radial sector
comes off at an acute angle, going out to the wing-apex and keeping elo>rl\
parallel with the radius. The median is somewhat inconspicuous owing to the
great length of the main stem, and the narrow areas which bound it between the
radius and the prominent cnbitus. It gives off t\vo outer branches before reaching
the broken edge of the wing. The median vein of the hind-wing has three
outer branches, the first arising nearer the base than the point of origin of the
radial sector, whereas in the fore-wing the first branch arises distallv to the origin
ol the radial sector. The median of the hind-wing is a more important vein than
its fellow in the fore-wing and occupies a greater area owing to its greater inward
curvature. The cubitus is a powerful vein with widely spaced divisions, the tir>t
branch, both in fore- and hind-wings, coming off from the main stem on the outer
>ide, and low down near the base of the wing, and then passing in a bold convex
sweep down to the distal portion of the inner margin. On its inner side the
cubitus give> off five branches, the fourth forking in the middle of its length.
The branches arise ai irregular distances, and the main stem reaches the margin

far oiil towards the Wing-apex. The cubitus of the hind-wing gives off a large
Outer liranch which is almost equal in strength to the main stem. This arises

UTHOSTALIS. 45

even nearer the base than its counterpart in the fore-wing, but its great length
is masked bv the development of the median. Beyond the origin of the outer
branch the cnbitus gives off three inward branches, the first forking. Owing to
the strong inward curvature of the cnbitus and its branches, the succeeding anal
veins are directed almost straight inwards. The anal veins are six in number, the
first forking twice, and ending on the margin in three branches.

The remaining veins are undivided. The cubital and anal veins of the
fore-wing are more obliquely disposed than those of the hind-wing, and the
latter has an anal area much larger than that of the fore-wing.

The interstitial neuration consists of short stout cross-nervures, which
occasionally fork, and in the wider areas unite to form a meshwork.

A triangular area is marked off from the base of each wing by a deep furrow.
This Woodward correlates with a similar area in the wing of Gryllacris (Corydalis)
lirniii/itiiiiii, which Swinton (' Geol. Mag., [2], vol. i, 1874, p. :J07, pi. xiv, tig. '•>)
has described as a stridulating organ. As stated elsewhere, I am of opinion
that Swinton's conclusions in the case of (.-1. In-fnnjiiitnii were founded on a
misinterpretation of the wing, and that no stridulating organ, or similar
structure, is present. In the case of L. carbonariiis such a structure would
apparently be useless, as none of the wings could come into such close apposition
as would allow the structure to be used. I am unable to determine its purpose
or significance, unless it be a portion of the musculature attachment of the wing.

Traces of a fore-leg are present, projecting from beneath the left lobe of the
prothorax. Its structure is too indefinite for description.

The wings were probably one-fourth or one-third longer than the portion
preserved — an estimate which would make each complete wing about 70 mm. long,
with a spread of about 140 mm., or (> inches.

Genus LITHOSIALIS, Scudder.
1881. Lit/iiiKinUf. Scudder, Geol. Mag. [2], vol. viii, p. 299.

Generic Characters. — Wings three to four times as long as wide. Outer and
inner margins almost parallel; intercostal area wide basally, and diminishing to
extinction at apex of wing. Radius simple; median with two main branches,
and cubitus large; anal veins few and oblique.

The wing on which this genus is founded is interesting as being the first
discovered in British Pahvoxoic rocks, except the problematic examph s "mentioned
by Lhuyd and not otherwise known. Supposed to be a plant, the wing' i was
first sent by Mantell to Jirongniart, who in turn referred it to Audouin. The
latter recognised its insect character, and brought it before the Entomological
Society of France, the Academy of Sciences, and the Assembly of German

46 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

Naturalists at Bonn. Aiulmiiii described tin- wing- as that of an unknown
Neuropterous insect allied to Hemerobins, Xo///;//.v and especially to GoTij^aLis.
.Mantell (r Audouin) described it as closely resembling a species of living Goryilalis
of Carolina. Swinton states that Mantel! purchased the fossil at a sale ot
Parkinson's collection, although Mantel! C. Medals of ( 'real ion.' p. .*i .">-!•) says that
lie '"discovered" it in a nodule From Coalbrookdale. Possibly the nodule had
previously formed part of the Parkinson Collection. Hoth the figures liy Mantel!
and Murchison are badlv drawn, and it was not until 1 S~4 that a reliable drawing
was published by Swinton. The latter author devoted considerable attention to
a "serrated vein" at the base of the wing, which he regarded as a stridnlating
organ. lie therefore referred the wing to the Orthoptera, and to the genus
Gi'i/ilctcris. Scudder threw considerable doubt on S.win ton's conclusions, and
showed that such an organ so placed could not have been of any service. As
we shall see later, the supposed stridulatmg organ is merely a torn edge of the
base of the wing. A studv of the wing-structure convinced Scudder thai the
wiiiL!' was most clnselv related to the Lithomantis c.arljunariiis, Woodward. Being
genericallv distinct from Lithomantis, and from living types, he gave the insect
the generic name of l/itJiOsialis.

Lithosialis brongniarti (Mantell). Plate 111, tig. 1 ; Text-figure lit.

1833. Corydulix ,', Audouin, Ann. 8oc. Enl. France, vol. ii, Hull., p. 7.

]s:_!il. ( ',ii-i/ilii/in .', Audouin. J'.uckland, Brid^ewatt-r Treatise, vol. ii. p. 77.

1844. Ciiri/iliilin liriniijiiiiiiii. Mantel], Mi-da Is of ('ivatioii, ed. 1. vol. ii, p. 57S. lie,u. 1:24, tii,r. '2.

1854. "Sialid;i'." Pictet. Traitc dt- Paleontologie, ed. ~2, p. :)77. pi. \1. Ii-. 1.

lst;7. t'liri/diilia, "allied to." Murchison. Siluria, i-d. 4, ]>. :!uu, woodcut SO.

1871. Cori/clii/ix liruHijiiiur/i, Woodvvafd. Gcol. Mai;'., vol. viii, ]i. :W7 ( naiur onlvl.

1874. Oorydalis hrmiiji/iiii-ti, Swiutun, Ge(>l. Mat;'. [-2], vol. i. p. 'i-!'.', pi. xiv. fi^. :!.

lx7-">. <li-i/llin-ri.< (<'nri/t/iili*) lii-'iin/iiiiiiii, Woodward, Gi'ol. Ma^-. - , vol. ii, p. <>2i2.

1870. (.'iiri/ihilis lirmiiiiiiiirli, \Vooihvard, Quarl. Journ. Cleol. Soc.. vol. x.\\ii, p. (<-2.

Issil. Curi/dulix lirinii/iiiiiiii, Nowa.k. Jalirb. k.k. ({col. K'ciclisanst .. \Vit-n, vol. xxx, p. 7^1. pi. ii, (!•_;• I.

1881. Lifli,,*;,!/;* In-nmjnnnii. Sclld-lc]', (i,M,I. Mill,'. - • Vol. viii. p. '1W .

188o. LillinssiiiHii In-iiiK/iiiiiiii, ScMiddtT, Mem. Bost. Soe. Nat. Hist., vol. iii, p. -J-jn, pi. \\ii. ii^>. 1. •_!. S, ;i.
1885. Prutogryllacrig linnii/iiiiirli, Brougniart, lJull. Soc. Amis Sci. Nat. Iii n :! . aim. \\i. p. .'*'.'

Lilliiiiiiiiiilix liriniijiiiii rll, IJroii^iimrt. Faunc Entimi. Terr. I'rini., p. )!71. fi^s. 17, IS.

Litlittti'ilit lii-iiiii/iiini'/i. llaiidlirsch, Die Fossilen Iusrkli-u, ]>. S4, pi. x. tii,'. |:'..

'/'///"'. — A left fore-wing; British Museum (Mantell ('oil.."///// Parkinson Coll.,
no. I I/,]*.)).

ll<n-i::<»i ninl Locality. — Coal Measures; near ( 'oallirookdale. Shropshire.

Specific Characters. Radial sector \\iih not more than iliree branches; median
much branched, and ocriipvmg much of the distal inner wing-margin. Cubitus
with three divisions. Anal veins few, and very oblique.

LTTHOSFALIS BRONGNJAETI. 17

I>,-xi-i'ifiti<m. — The k't't fore-wing measures 61 mm. long and IS mm. wide
across the distal third. The apex, the inner margin and a very small portion
of the base are missing. The wing appears to have been strap-shaped, with a
blunt apex, the inner margin curving most into the apex.

The outer margin is feeble, almost straight over the greater part of its length,
and slightly sloped backwards. Basally it dips abruptly inwards to the point
of attachment. The subcosta is widely spaced from the outer or costal margin
proximallv. the two gradually approaching each other until they meet at the
wing-apex. The radius passes straight out to the apex, giving off the radial sector
before the middle of the wing is reached; the radial sector remains undivided up
to the distal third of the wing, beyond which it forks twice. The median vein
divides a little further out than the radial sector, forming two strongly divergent
branches, an outer large branch with two large forward twigs, and a small inner
branch which forks, the inner twig forking again. The median, therefore, ends
on the inner margin of the wing in six twigs.

pIG. jo. — Litlinsinlis bronii it i" rli (Mantell) ; diagram of neuration of left fore-wing, natural size.—
Coal Measures-, near ('c>!ill>r<»>kcla.]e. Shropshire . Mantell Collection, ulhii Parkinson < 'olhvtion,
Hrit. Mus. (no. ll,lil!»)

The cubitus divides into three branches, the first arising low down, and passing
in a bold sweep beyond the middle of the inner margin. The anal veins are five
in number, all but the first being undivided. The first anal forks near the base,
the outer branch forking again.

The whole wing is covered by a numerous series of strong transverse nervures,
rarely branching, and usually crossing at right-angles to the main veins. A few
are oblique or curved.

The "file" or "serrated vein," described at length by Swinton, and by him
correlated with that present in recent forms of Gryllacris, appears to be nothing
more than an irregular torn edge of the basal part of the wing, which is also lifted
up a little above the general level. The torn edge extends along the line of the
median vein for a short distance, and wholly lacks the symmetry and detail given
to it by Swiii to n.

Aljiiiifii'S. — Although closely allied to Litltointnttix rn i-li/ninr/iix, \Voodw., this
wing differs in the great Avidth basally of the intercostal area, and in the straighter
course of the cubitus and anal veins, due to the greater length of the wing. The
interstitial neuration is much the same.

48 FOSSIL IXSKCTS OF THE BRITISH COAL MKAST IIKS.

Genus PRUVOSTIA, novum.

Generic < 'linrn/'t/'rx. — Fore-wings two ;iml ;i li:ilt' times as long as wide; outer
and iniii'i1 wing-margins almost parallel; \ving-apex well rounded. Sidicosta
widelv removed from the costal margin liasally. and apparently noi connected with
thecosta or radius. Radius straight with divergent radial sector. Radial sector,
median and cubitus all distally branched Anal veins few. Interstitial neuration
of straight cross-nervure>.

Certain features of this wing- are suggestive of the Protorthoptera. These are
the wide basal, intercostal area, the basal origin of the radial sector, and the
remote branching of the radial sector, median and cubital veins.

It is, however, more nearly allied to LitJiosialis than to Metri/in minlis, Ilandh,
for example, among the Protorthopteroids. The great length before division of
the main stems of the radial sector, median and cubitus, their strong divergence,
and the many branches of the median, form an assemblage of characters not
elsewhere known, and certainly deserving of generic recognition.

It is with pleasure that I attach to this genus the name of l>r. 1'. I'ruvost, of
Lille University, in recognition of his valuable work on the fossil insects of the
north of France.

Pruvostia spectabilis, sp. uov Plate 111. tig. 2; Text-figure 11.

7'///»'. — A left fore-wing; British j\luseum (.lolmson Collection, no. I. K>,8(.>!).

Jl/n'i::"ii nnil /,<«•<///'///. — Aliddle Coal Measures (clav ironstone nodules in the
Hinds between the ''Brooch" and "'Pluck" coals); Coseley, near Dudley,
Staffordshire.

KM. 11. — Prueostiti .;/"V/,//,i/is, sp. nov. : diagram of neural inn "1 l<'t't !'«]•<• -winy, natural si/r Coal
Measures (clay ironstone nodules in tin- lands Ix-tween the " Jirom'li " and "'I'liii-K" coals);
Coseley, IKMT Dudley, Stall'm-dslim-. .l,,lni.-,i,ii ('oll.'i-ti.»n, Brit. Mus. (no. I. 15,891).

iii'r*. — Outer margin convex at base, and inclining inwards to
the apex. Subcosta straight, not reaching the apex of wing. Radius straight,
irivinu' off radial sector near base; radial sector twice branched, median divergent,
three limes branched, the first two branches forking. Cubitus dividing basally
into two, the outer forking three times, the inner branch once.

I ><:•«•,'/ i>li<:/i. — A left fore-wing -V_! mm. long and - I -111111. wide, almost covering
the surface of one half of the median plane of a reddish -brown ironstone nodule. A
little of the base of the wing is missing, while a |iarl of the wing-apex is concealed

PRUVOSTIA SPEOTABILIS. 49

liy ;i film of ironstone \vliicli cannot lie removed. Fortunately the slight film of
ironstone does not hide the outline of the wing, and it is evident that the outer and
inner wing-margins are almost parallel, while the apex is well rounded. The outer
margin merges l>y a well-rounded contour into the wing-apex. The subcosta is
well-marked, widely removed from the outer border proximnlly, and lies in a shallow
groove formed of the intercostal and radial subcostal areas. These two areas are
reduced distally to less than half their proximal diameter owing to the backward
inclination of the outer margin. The subcosta seems to die out near the radius,
about 7 mm. from the wing-apex, and gives off a numerous series of irregularly
spaced and forwardly directed nervures, which are at first straight and then curved
towards the wing-apex At their point of origin the cross-nervnres are very
distinct, but they thin, and occasionally die out, before reaching the outer margin.
The radius is a strong vein, passing perfectly straight out from the wing-base to
the outer part of the wing-apex. The radial sector arises near the base of the
radius, gradually diverging from it up to the junction of the middle and outer
thirds, where a single branch is given off, the latter taking a position parallel with
the main stem of the radial sector, and entering the middle of the wing-apex.
Kleven mm. further out and within 7 mm. of the apex a second branch is
given off, which lies evenly between the main stem and the first branch. The
median vein arises so close to the radius as to appear united with it. Like the
radial sector, it does not divide until it reaches the junction of the middle and outer
thirds of the wing. Here the first inward branch arises, and at equal distances
further out are given off two more; the first two each fork in the middle of their
length, the third remaining undivided ; the median ends therefore on the inner
wing-margin in six divisions. The cubitus consists of two, and possibly three,
main stems. Owing to the base of the wing being broken away these stems are
not seen to be in actual union. Little doubt can exist as to the union of the first
two, but if the third vein is a branch of the cubitus, it can only join the other two
by a strong outward curvature. This third vein may be the first anal, although its
manner of division is similar to that of the cubitus, in this respect agreeing with
what is seen in the first anal of Lithosialis brongniarti. The next vein is parallel
with the one succeeding, which is undoubtedly anal, while it shows an increasing-
basal divergence from the second cubital. For this reason I have regarded the
third vein as the first anal. The first branch of the cubitus diverges widely from
the median, passing obliquely to the inner margin of the wing, which is reached
just beyond the middle third. Owing to the great divergence of the first cubital
branch from the median, the area between the two, at the point where they first
branch, is very wide — almost twice the width of any other area.

The first cubital vein gives off three outward twigs. The second is parallel
to the first as far as its division into two equal twigs. The next vein is that to
which I have already alluded as a possible third branch. It is widely separated
7

50 FOSSIL INSECTS OF THE BEITISH COAL MEASURES.

from tlit1 undoubted cubitns, and l>v reason of its t\vo forward branches resembles
the cubital elements in front of it. Its wide separation from the second branch of
the cnbitns supports the view that it is the first anal vein. The undoubted anal
veins are two in number, undivided, and passing backwards very obliquely to the
wing-margin. More anal veins may have been present, but it is verv doubtful.
The wing-surface is much wrinkled across both the length and breadth, and
traversed by numerous cross-nervures, which are most evident in the median area.

Affinities. — The wing is typically Pahtodict yopteroid, and in the development
of the radius, radial sector, and cubitus, shows an affinity to Lithosialis l>r<>n//iiiitrti.
It differs markedly from that species, however, in the great length of the main
stems before divisions arise.

Dr. Tillyard, to whom I have shown my enlarged drawings, is of opinion that
Pruvostia is allied to Pseudofouqiiea rather than to the Lithomantidse.

Family KI.'KYKKIIKK, llandlirsch.
190IJ. Handlirseh, Die Fossilen Insekten. p. 95.

Wings markedly triangular, with broadened bases. Costa, subcosta and radius
brought closely together in the outer part of the wing, the median with few
divisions. Cubitus and anal veins directed almost straight inwards, at right
angles to the length of the wing.

Cenus BREYERIA, Borre.

1875. Bi'i'i/fi'in. I5orre, Arm. Soc. Eiitom. Belg., vol. xviii, p. 40.
1908. St'Mifin. Handlirsch, Die Fossilen Insekten, ]>. 1348.

Generic Characters. — Wings two and a half times as long as wide. Outer wing-
margin straight; apex curved inwards. Inner wing-margin strongly convex.
Costa marginal, subcosta parallel and joining the radius near the wing-apex.
Radius giving off the radial sector near the base of the wing, and reaching the
apex undivided. Radial sector subparallel to radius. Median boldly curved inwards
almost at a right angle. Cubitus less curved than the median. Anal veins few.
Interstitial neuration of two kinds, that of the intercostal and radial sector areas of
short straight nervures, and that of the remaining areas of irregular thin nervures
which tend to anastomose into a loose meslnvork or reticulate arrangement.

Breyeria woodwardiana ( llandlirscli ). Text-figures li!. l:>.

190:!. "Allied hi Lithomantig cartonarius," Slobl.s, (in.l. M.i^ I . \,.| \, |> 5~24.
PalaeodictyopterOll, SI)., llandlirscli. l»n- l-'ussilni InscLlen, |>. I'jli.

BREYERIA WOODWARDIANA.

51

1906. LitJiomantis carbonarius (/), Woodward, G-eol. Mag. [5], vol. iii, p. 26, fig. 1.
1908. Stobbsia woodwardianct, Handlirsoh, Die Possileu lusekteu, p. 1348, text-fly.

C. — Greater part of a left hind-win"- ; Mr. J. T. Stobbs' Collection.

Horizon <nnl Locn.Hhj. — Peacock marls overlying the Peacock Coal, and near the
top of the workable Coal Measures ; Foley, near Longton, Staffordshire.

Sfifcijii- l '/nii-iifh'1-x, — Radial sector dividing in its distal half into fine inwardly
directed twigs which end on the distal third of inner margin. Median and cubit us
with few widely spaced branches, occupying greater part of inner margin.
Anal veins about three to four in number.

Description. — The specimen has not been available For examination, and my
observations are based on the figures published by Woodward and Handlirsch. It
is probable that neither of these figures is wholly correct, Handlirsch pointing out
what appears to be an obvious error by Woodward in the character of the cubitus

FIG. 12.

FIG. 12. — Breycrut wouilirnnliiiini (HandlirsclO ; left hind-wing as restored mid figured by Dr. H.
Woodward under the name of Lithomantis carbonarius?, natural size. — Coal Measures (Peacock
marls overlying the Peacock Coal) ; Foley, near Longton, Statt'i >vd*hire. Mr. J. T. Stobbs' Collection.

Fin. 13. — Breyeria vjonilii-iirtliinin ( Handlirsch ) : same- left hind-wing as restored and figured by Dr. A.
Handlirsch under the name of Stalilixin ii'iimlti-in-tUmui. nearly natural size. A = IX, anal; Cu —
VII, cuhitus ; M = V, median : R = III, radius ; Rs = IV, radial sector ; Sc = II, subcosta : /, costa.

vein, and the position and mode of division of the radial sector being doubtful.
Notwithstanding this difficulty, I have by means of an old plaster cast been able
to satisfy myself upon the more essential details.

The fragment is 50 mm. long and '20 nun. wide, and consists of the greater
part of a left wing, of which the base and a portion of the basal third are stated
to be obscured by a pinnule of Neuropteris. The maximum width is across the
anal area, beyond which the wing rapidly narrows owing to the forward direction
of its inner margin. The costal vein is marginal and strong, passing in an almost
straight line distally until joined by the subcosta, after which it curves gradually
backwards into the wing-apex. The subcosta is parallel with the costal margin
over more than two-thirds of the wing, and then joins the costa. The base of
the radius is not shown in the figures, except a very small portion in front of the
point of origin of the radial sector, the latter arising in the basal third of the wing;
beyond the radial sector the radius passes out to the wing-apex, keeping parallel
with the costal maroin.

52 FOSSIL TXSKC'TS OF TIT K Bh'ITIRH C'OAF MKASFRKS.

The radial sector presents several difficulties. In neither of the two published
figures is this vein depicted as we might expect. In all other respects the wing
agrees remarkably closely with those of ///v//, •/•/'' and Megaptiloides, where the
radial sector sends off inwards a series of simple branches, and runs out fairly
parallel with the radius to the wing-apex. In [dace, however, of the radius
passing straight outwards, it is represented as dividing into two branches in the
distal fourth of the wing, the outer branch forking once and the inner forking
twice. The inner branch of the radial sector diverges widelv from the outer, and
its three divisions go to the inner side of the wing-apex. A small branch is shown
by Woodward as joining the radial sector to the first branch of the median, while
the same branch is shown bv Ilandbrsch as coming off from the radius immediately
in front of its division into two, and passing down towards the inner margin
between the inner branch of the radial sector and the first branch of the median.
but not uniting to the latter. The median vein arises near the radius and sweeps
out in a bold curve to the middle of the inner margin, giving off three outer
undivided branches. Dr. "Woodward, in his restoration of the base of the wing,
has inadvertently indicated the main stem of the cubitus as joining the median.
This is corrected in Handlirsch's drawing. The cubitus consists of a strongly
curved stem giving off two branches, but only the inner marginal portions of the
veins are present. The anal veins are three or four in number, and directed
backwards at right angles to the length of the wing.

o O o o

The interstitial neuration consists of feeble transverse nerviires. which either
pass irregularly across between the main veins or occasionally fork.

Affinities. — Dr. Woodward doubtfully refers the wing to Lithomantis carbo-
nari-its, Woodw. Ilandhrsch, in the earlier part of his \\ork. ' Die Fossileii
Insekten ' (p. 1 -(.'>), classed it as a " Palaeodictyopteron " only, and afterwards,
owing to its supposed relationship to ],. carbonariits, and its evident likeness to
Litlnixni/ix and ll<nlrt>ii<-/i r<i. established the genus Sluhlixiii for it, placing the specie>
in the Lithomantidse. There are, however, certain features of the wing which
militate against his view. The close apposition of the cost a, subcosta and radius
are in marked contrast to the condition in that familv. where these veins are
widely spaced, and where there is also a verv wide intercostal area. The radial
sector is also more complex. In those details in which the wing departs from the
Lithomantid;e, it approaches the characters of the genera /</v//'''''" and .l/<';/<>/<//-
loides. The resemblances to Mef/aptiloides /</•<»//>'/, IJrong.. and Breyeria borinensis,
Horre, are very close, so far as can be determined bv the distal fragment of the
lormer wing and the more than three-fourths of the latter wing. The mam
difference between (his wing and ihose of In-ri/i'i-in l/ariiifiixix and /•'. lachlani is in
the character of the siilicosta, which in the latter two species joins the radius.
Whether the publislied figures of the Staffordshire wing are correctly drawn in
tins particular \\e do not know, and as we have alreadv seen that these ligures are

SPILAITKRA. 58

wrong in other details, Ave are compelled to assign the specimen to the family
Breyeriidae, and doubtfully to the genus ///T///- rin, Avith which it, seems in agreement.

Family Snr.AiTKUiD.i-: (nro^j^m't), Handlirsch.
liioti. Hanillirscb, Die Fossilpn Inseklen, i>. 1(11.

< I

Radial sector more or less branched; median divided into two main branches,
the outer much divided ; cubitus with an outer branch sending numerous twigs to
the inner margin. Intercostal area occupied bv a, series of straight cross-nervares.

The family Spilapteridae, founded bv Hronguiart in 1885, has been re-defined
by Handlirsch and Lameere. The latter extended the group to include species
which Hrongniart had placed partly in the family Platypterida? and partly in
Protephemeriihe. All the forms thus brought together by Handlirsch agree in
the possession of a typical palaeodictyopteroid neuration and the general characters
enumerated above.

Lameere ('Hull. Mas. Hist. Nat. Paris,' 1017, no. 1), Avho rejects Handlirsch's
vieAvs and classifications (see above, p. I-")), has remodelled the family and given it
a new significance. He is of opinion that the genera Lamproptilia, Epitethe,
Becquerelia, Palseoptilus, Compsoneura, Spiloptilus, Homaloneura, Graphiptilus and
Kliiliiptri-a form a natural family, the Spilapteridae, in which a progressive
evolution in the longitudinal venation can be observed. The family is regarded
as linked to that of Megasecopteridae through the genus Becquerelia, and to the
Protephemeridae through the genus Ajinjxiji/iii*. The three families are then
grouped in his new order, Ephemeroptera. Such a classification is based on the
belief that a perfect evolutionary sequence can be made out. Unfortunately, in
presenting this classification, Lameere gives only a summary of his reasons and
evidence, and it is not possible to criticise his •argument. It is. to say the least,
very doubtful if, in the present state of knowledge, Ave can judge relationships
always correctly, Avhile the sequence of evolution is still more difficult.

Handlirsch has acknowledged that he is unable to undertake an\' division of
the family Spilapteridse, as he understands it, and is content to aAvait the discovery
of the bodies of these insects for a fuller knowledge of the family. As the more
rational vieAv, Handlirsch's definition of the family is adopted.

Genus SPILAPTERA, Brougniart.
1885. Spilaijtera, Broujjiiiart, Bull. S.>r. Amis Sci. Nat. Roneii [3], ;mu. xxi, p. 63.

< ti'iii'ftf Characters. — Insects closely resembling Pdl&uptilim ; fore-wings
narrower basally than the hind- wings ; body slender.

54 FOSSIL INSECTS OF THE HliTTISH COAL MKASI'IJKS.

Spilaptera sutcliflfei, Boltoii. 1'latc 1 1 1, tig. :\ ; Text-figure 14.

1917. Si>ilai>teru sutcliffei, Bolt oil, Quart. Jouru. (jeol. !Soc., vol. Ixxii, p. 53, pi. iv, fi<,'. 1, anJ
text-fig.

Ti/]>t'. — Basal third of a wing; Manchester Museum (no. L. Sh)~).

[fori::/m mnl Loi-nlif ij. — Middle Coal Measures (grey-blue shales 13o — 1 SO feet
above the Royley or Arley Mine) ; Sparth Hottoms. Rochdale, Lancashire.

X/>i-fijif < 'Im ructt'i-x. -Subcosta ]»arallel \vilh the costal margin; median vein
dividing near the base into two branches, of which the outer forks just beyond the
point of origin of the radial sector. Cubitus a large and much-divided vein.
Anal veins few in number.

Description. — This specimen was formerlv labelled " Stenodicti/a lobata," and is
one of three recorded by Sntcliffe, Baldwin, and others, in their papers on the
fossils found at Sparth Hottoms. The remaining two are described (p. 37) under

Km II. — ,S'jn7ri;>/,T.i s»/r/,/j'ci, Holton : restoration of left wing, natural size — MicMle (A>al Measure.-,
(shales al>ove (In- Kovley or Arley Mine) : Sparlh ]intt»ius. K'm-licbilr. Ijaiu-asliire. Manclirstci-
Museum (no. L. SUIT)!

Mecynoptera ln/><'r<-/i/<if<i, Molton. The specimen consists of the basal third of a
wing lying on the median plane of a small irregular micaceous sandy nodule. The
finer structure of the wing has not been preserved, owing, no doubt, to the coarse
grain of the matrix. The chief veins of the wing are robust structures, and these
are fortunately well marked and clear. The wing-fragment is .T> mm. in length
along the outer margin, and -7 mm. in greatest width, but as the inner margin is
broken away and lost, the total width of the wing exceeded this, and mav have
been over •'!<> mm. It belongs to a left wing, and when complete must have been
at least '.«> mm. in total length. The perfect insect must therefore have had a
spread of wing of nearly 2<l() mm., or about S inches.

The outer margin is feebly convex, the subcosta fairly parallel, sunk in a groove,
and gradually approaching the outer margin as it passes towards the wmir-apex.
The rate of approach is so gradual that the junction must have been far out near
the wing-apex. The intercostal area is crossed by a series of straight nervmv.-
which arc oblique in 1 heir course outwards.

Sl'TI.APTERA SUTCLIFFEI. -55

The radius is strong, standing up above the surface, and almost parallel with
the snbcosta. The radial sector arises from the radius at about 22 niiu. from the
base of the wing. The median vein, owing to its inward direction away from
the radial sector, is better shown. It forks very low down into two equal
branches, both of which are widely spaced. The outer branch divides almost
opposite the point of origin of the radial sector, while the inner branch forks
much nearer the base of the wing, the innermost branch curving sharply
back from its fellow, so that the area between them is wide almost from the
commencement.

The general direction of the branches of the median is such that they Avould
reach the outer half of the inner margin of the wing.

The cubitus consists of two main stems, the basal union being missing; the
outer branch passes, after twice forking, in a double curve down to the inner
margin of the wing, the inner branch being directed more directly backwards,
and forking three times in the basal third. Fragments of four anal veins are
distinguishable, the first dividing near the middle of its length into two equal
branches. The interstitial neuration seems to have consisted of comparatively few
straight and well-spaced nervures, some of which can be seen crossing the wide
area between the innermost twig of the median and its fellow. No others are
visible, except those in the intercostal area already mentioned.

Affinities. — Notwithstanding the fact that only about one-third of the wing
is known, it is yet possible to determine the generic characters with a reasonable
degree of accuracy. The mode of division of the radius and median, the manifest
importance of the latter, the wide area between the median and cubitus, and the
few widely spaced cross-nervures, are all typical Spilapteroid characters. The
wing cannot be confused with that of Stenodictya, in which the median is much less
developed, and the interstitial neuration closely reticulated. The only other genus
to which it might be referred, that of Beojuerelln, is distinguished by a union of
the anal veins, which in Spilaptera and in this specimen are distinct.

Spilaptera has hitherto comprised only three species. From S. pucl'nrJi,
Brong., the specimen differs by the subcosta being parallel with the outer margin;
by the equal separation of the subcosta, radius and median in the basal third of
the wing ; and by the division of the median into two branches close to the base of
the wing. N. lil><'llnh>i<1es, Brong., of which the distal half of the wing only is
known, has a much feebler cubitus, which divides by forking further out than in
S. sutdiffei. N. reintstn, Brong., was established on a fragment much similar to that
of N. liMlnloides, in which the subcosta is a short vein, and the radial sector arises
nearer the middle of the wing.

56 FOSSIL TXSECTS OF THE BRITISH COAL MEASURES.
Familv LAMPROPTILID/E (Brongniart), Handlirsch.

1885. Brougniart, Bull. Soe. Amis Sri. N;it. K'oiicn [3 . ami. .\\i. |>. >'<•'>.
190C. Handlirsrli. Hie Kossilcu Insi-kti-n, p. lu'.i.

Fore- and hind- wing stronglv marked. Veins of the anal and cubital groups
numerous and directed obliquely backwards. Hind-wings short and broad.

(ienus BOLTONIELLA, Handlirsch.

1885. Lamproptilia, Brongniart. Bull. Sue. Amis Sci. Nat. Rourn :! , aim. x\i. p. tl:;.
1919. Boltoniella, Handlirscli, Revision <ler Paliiozoischeii Insi-klijn, p. Jl.

Generic CJiarnrferx. — Fore-wings two and a halt' times as long as wide ; hind--
wings as long as wide. Outer margin of former curved, of hind-wings almost
straight. Apex of wings rounded. The fore-wings are almost elliptical, and the
hind-wings rectangular in shape. The costa is marginal and the costal area some-
what narrow. The subcosta joins the costa far out. Radius simple, the radial
sector arising near the base of the wing and giving off three forked branches,
which pass obliquely to the inner half of the wing-apex. Median dividing into
two main branches, the inner twice as much divided as the outer. Oubitus with
twro main branches, each breaking up into numerous twigs. Interstitial neuration
consisting of widely spaced straight cross-nervures.

Handlirsch, who has formed this genus, admits that it reminds one in many
ways of Lamproptilia, but by reason of the small si/,e and closer spaced cross-
nervures he separates it from that genus.

Boltoniella tenuitegminata (Bolton). Plate TIT, fig. 4.

1911. Lamproptllin tenuitegminata, Bolton, Quart. Journ. Geol. Soe., vol. Ixvii, p. 170. pi. x. fit;, li.

Ti/jie. — A right hind-wing, in a small fragile block of brown mudstone or shale,
crowded with plant-remains; Museum of Practical ( Jeologv, .lermvn Street, London
(no. 24509).

Hori::oit nt/il Lnrnlifi/. — Coal Measures (Xo. -2 Rhondda Seam, base of the Pennant
Scries) ; 1 :(! miles north-east of Resolven Station, Glamorganshire.

S/nrijic Characters. — Broad wings of great tenuity and blattoid-like in character.
Principal veins much divided, and fading out on the inner margin. Apex of win^
bluntly rounded. Anal veins very numerous.

Descfijilinii.. — The type-specimen consists of a right hind-wing of considerable
tenuity, the underlying plant-remains being easily traceable through the texture
of the wing. The greatest length is 'J'.t mm., and the greatest breadth 7 mm. The
neuration is so similar to thai which I have observed in the hind-wind's of blattoids

BOLTONIELLA TENUITEGMINATA. 57

from Commentry, France, that it is difficult to refrain from classing the wing as
lilattoiil. The neuration is extensively branched and the wing-boundaries are not
\vell marked, but the wing appears to have been somewhat quadrangular in outline,
\vitli a sinuous inner margin, and the base much broader than is seen in the ordinary
form of blattoid hind-wing. The outer margin seems to have been straight, a
portion of it still remaining in the middle third. The two margins merge in a
well-rounded apex. The subcostal area is narrow, strap-shaped, and probably
extended over the whole length of the outer margin. No traces of cross-nervures
can be seen on it. The radius divides near the base, giving rise to a series of
branches which curve inwards as they approach the apex of the wing. The course
of the branches is irregular, the interspaces widening and narrowing, possibly
owing to the wing having crumpled during deposition. The median divides into
two branches low down, each of which is repeatedly forked, the final divisions
becoming attenuated and nntraceable before the inner margin of the wing is
reached. The course of the cubitus is obscured by a reed-like plant, only two
basal portions being distinguishable. The anal area is filled by a broad series of
thread-like veins, which sweep obliquely inwards in a fan-shape, and occupy a large
part of the inner margin.

The inner margin of the wing to a third of its total length is quite filmy, the
veins crossing the area as faint shadowy lines. The distal two-thirds is more
strongly impressed, while in the broad base of attachment the stems of the
principal veins seem to have been more than usually robust. No trace of trans-
verse nervures or reticulation can be seen.

Affinities. — The nearest analogues to this wing are, I believe, the forms described
by Brongniart as L<intpr<>/iti/i<i <jr« n<i'<'i/r//i and L. *tirr>ij>! ('Etudes sur le Terrain
Houiller de Commentry,' vol. iii [1*93], pp. 467—70, pi. xxxv [19], figs. 7—9).
It appears to lie more closely related to L. stirrttpi than to L. r/r((ii<r<>ur//i, but is
more quadrangular, and its costal area is broader. The anal portion of the wing-
is of greater tenuity, and occupies fully half of the inner margin.

Family BRODIID^, Handlirsch.

190G. Handlirsch. Die Fossilen lusekten, p. 113.

1919. Handlirsch, Revision der Paliiozoischeu Insekten, p. 73.

Wings in which the anal area is much reduced and specialised, the radius
undivided, and the median, cubitus, and anal veins arched and directed back
towards the inner margin.

From a re-examination of the originals in the British Museum, Handlirsch has
concluded that the family does not belong to the Palasodictyoptera, but to the
Order Megasecoptera. This view seems at variance with his own definition of the
8

58 FOSSIL IXSKCTS OF THK BRITISH COAL MEASURES.

latter Order ('Die Fossilen Insekten,' p. 312), where lie states that the special
distinctive characters of the Megasecopteridse are the tendency to a reduction of the
anal area of the wing, and the partial fusion of the median and cubitns with
the base of the radius. Hr<i<li« certainly agrees with the Megasecopteridae in the
reduction of cross-nervures, but the anal area is very long, and I have not seen
a single case among the twelve or thirteen specimens in the British Museum
where there is any clear sign of fusion between the median, cvibitus, and
the base of the radius. The bases of the three veins arc- closely brought together
in the inner spatulate portion of the wing, but this seems a natural result of the
narrowing of the wing in this region.

The family Rrodiidre is more fully represented by individuals in the British
Coal Measures than any other, thirteen specimens being known of llnul/n
/irixi-oliiirtu, Scd., alone. To this family also seem to belong a series of small
larval wings. The neuration in these is immature, and the wings themselves are
far too small and weak to have supported the body in Might. The abdomen seems
long and well segmented, the segments showing evidence of well-developed
pleura, and they may also have borne tubercles. The whole integumentary
structure is thin, and if at all chitinous, only feebly so, and but faintly outlined
on the surface of the nodules. The wing-shape is much like that of Ili-nJui, and
the veins, so far as determinable, such as may reasonably be supposed to have
developed into the typical //-/Wm-type in the adult.

While o-eneral evidence and association point to these wings being those of

O 1 <— '

larval forms of Hi-o<li«, and possibly of /.'. priseotincta, Scd., it is more judicious
to retain them in a distinct group. The same difficulties occur with other larval
wings which cannot be allocated to known genera, and for which new generic
names are not advisable. I therefore class larval wings of unknown relationship
as a separate group under the name of" Pteronepionites," a name which has no
classificatory value, but merely serves to indicate their larval nature. When
evidence is forthcoming of the generic and specific relationship ol auv member
of the group, it can be removed without the necessity of reducing a generic term
which has passed into nomenclature to the rank of a synonym.

I would allot the term " Pti'r<ni>'i>i<iiiih'x " to all larval insect-wings of all
geological periods.

Genus BRODIA, Scudder.

1881. Itnnlni, Srudder, Geol. Mag. [•_>!, vol. viii, p. -J!»::.

rir Characters,- — Wings spatulate in shape, tliree I lines as long as wide.
Outer margin feebly convex. Costa and radius spinulose, radius undivided, radial
sector and median vein with lew divisions. Cubit us a single vein. Anal veins
few and widely spaced.

BRODU PRISCOTINCTA. 59

Brodia priscotincta, Sendder. Plate III, figs. 5, 6; Plate IV, figs. 1 — 3; Text-
figures 15, 16.

1881. Urtnliii jiriffotinrtn. Scudder, G-eol. Mag. [2], vol. viii, p. 293, text-tig.

1883. Brodia {>riat'i>!!>irtn, .Scudder, Mem. Boston Soc. Nat. Hist., vol. iii, p. 213, pi. xvii, figs. 3 — 7.

1885. Brodia prim-otiiii-tii, Bronguiart, Bull. Soe. Amis Sci. Nat. Eoueii [3J, aun. xxi, p. 63.

1893. Ji roil in jii-im-otini-lii, IJroiigniurt, Fauue Eutoiii. Terr. Prim., p. 528, pi. xl (24), fig. 4.

1906. Brodia priscotincta, Handlirsch, Die Fossileu Insekten, p. 113, pi. xii. tig. 13.

1917. Brodia priscotincta, Boltou, Proc. Birmingham Nat. Hist, and Phil. Soc., vol. xiv, pt. 2, p. loo,

pi. vii, figs. 3—4, text-tigs. 2 — 3.

1919. Braclifi priscotincta, Handlirsch, Revision der Paliio/oischon Insekten, p. 73, tig. 83.

1919. Broil la 8<-n<lclrri, Haudlirsch, <>/>. cit., p. 74, tig. 84, Brit. Mus., no. I. 3879.

1919. Broilitt petiohita, Handlirsch. nfi. cit., p. 74, tig. <S5. Brit. Mus., no. I. 2961.

1919. Brodia pict'qii'iiiiif, Handlirsch, op. <•//.. p. 74, fig. 86, Brit. Mus., no. I. 2961.

1919. Brntlin fum-iiitii, Handlirsch, op. cit., p. 75, fig. 87, Brit. Mus., no. I. 1557.

1919. BrofUa itebutuM, Haudlirscli, np. cit., p. 75, tig. 88, Brit. Mus., no. I. 2961.

Ti/pc. — Incomplete wing; British Museum (Brodie Collection, no. I. 3896).

llnri::nii «ml Lm-iilit//. — Middle Coal Measures (clay ironstone nodule from the
binds between the " Brooch " and " Thick " coals) ; Dudley, Staffs. (Scudder gives
the locality as Tipton).

FIG. 15. — Brodia priscotinctn , Scudder ; diagram of a complete left wing showing position of tubercles
and character of neuration, natural size. — Middle Coal Measures : Dudley, Staffordshire.

Specific Characters. — Outer margin of wing spinulose, almost straight, or feebly
convex at the most. Subcosta remote from margin at base, and extending beyond
the middle of the wing, joining neither to the costa nor to the radius. Radius a
strong vein, and spinulose. Radial sector arising in the first half of the wing, and
giving off four branches. Median with two forward branches. Cubitus a single
vein. Anal veins two. Inner margin strongly convex.

Description. — The Geological Department of the British Museum possesses no
less than eleven examples of this species in addition to the type, while two more
are in the Geological Collection of Birmingham I'niversity, one being in the Beale
Collection, and the other presented by Dr. Blake. The British Museum specimens
are registered as follows : I. 1557, I. 1567, I. 3879, I. 3896, In. 18431, all from
Coseley; I. :2961, In. 184.3:5, In. 1843 i, all from Tipton; In. 184119 from Sedgeley ;
and In. 18430, 1. 5866, I. 2902, and In. 18432. The following description and
remarks are based on a study of all these specimens.

The longest wing I have seen measures 58 mm. in length, and we are not likely

60 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

to be beyond the mark in estimating the total span of the insect as about !:><> mm..
or over 5 inches.

The outer or costal margin is straight, or feebly convex at most, over the greater
part of its length, curving distally into the wing-apex. Close to the base it swells
out into a slight hump-like elevation which is seen in all cases where the wing has
been broken off close to the body. The whole of the outer margin bears a dense series
of minute, conical, sharply pointed spinules of a black colour. These are arranged
in two rows on the proximal half, with the points of the spinules directed towards
the wing-apex. The bases of the spinules are expanded, and give a doubly corded
or monilated appearance to the wing-margin. Tillyard regards these spinules as
modifications of large hairs which lie has termed " macrotrichia," and I see no
reason to dissent from his view.

The subcosta at its origin is widely spaced from the outer margin, and passes
out beyond the middle of the wing, gradually approaching, but failing to reach it.
The subcosta is a strong, straight vein, the greater part elevated above the level
of the wing-membrane, and distally flattening into it and disappearing.

The radius is a strong vein, more convex than the outer or costal margin, and
therefore more widely separated from it at either end than in the middle. The
basal portion is parallel with the subcosta for the whole length of the latter; it
then becomes parallel with the outer margin for a short- distance, and afterwards
curves into the wing-apex. A single row of spinules can be distinguished along
the whole vein in some specimens, and gives a slight monilation to the vein-surface.
The radial sector is well marked, and comes off from the radius about the point of
origin of the first forward branch of the median. The two veins are close together
for some distance, but in the distal half of the wing they become parallel, the
interspace being equal in width to that between the radius and the outer margin.
The radial sector gives off four inward branches, which end on the inner side of
the wing-apex. The first branch arises just beyond the middle of the wing, at
an acute angle, and is separated from the second by an interval which is double
the length of that separating the second and third. The fourth branch is verv
short, and so close to the margin as to be absent in some specimens.

The median vein for the first quarter of its course lies in the middle line of the
wing, and then bends inwards in a wide curve to the distal third of the inner
margin. It gives off two outward brunches, both of which are larger and stronger
than those of the radial sector. Itotli brunches have the' same sweeping curve
possessed by the branches of the radial sector, and are parallel \\itli the latter,
while the main stem becomes almost straight.

The ciibitus is a single vein, not united at the base to the median, and passing
obliquelv to the margin. Some wings have broken oil so lur out from llie base
that the ciibifus appears lo join the wing-margin at the junction of the middle and
distal thirds. A striking feature of (he cubitus is iis isolated position upon the

BRODIA PRISCOTINCTA. <>1

margin, for while the interval between it and the stem of the median is much
wider than any other in the fore-part of the wing, the interval between the cubitus
and the next vein is nearly twice as wide. There is, in fact, a progressive
widening between the veins as they are traced from the apex to the base of the
wing, the areas between the median and its branches being wider than those
enclosed by the branches of the radial sector.

Two somewhat dissimilar anal veins are present. The first has a wide curve
to a point far out, on the margin, sometimes giving off a short branch, while the
second vein is much shorter and joins the margin at an acute angle. In some
specimens this vein is seen to give off two, or even three, short oblique branches.

The shape of the wings is very similar to that of the wings of mosquitoes
(Anopheles}, and they bear evidence of having been folded in a plicate or fan-like
fashion along their length. The first two folds are united at the wing-base, and
pass out along the radius and the first outward branch of the median. The third
fold lies along the line of the cubitus vein. The degree of plication which a wing
retained when silted up modifies considerably the apparent distance between the
several veins and their branches, and at times hides important junctions. In one
example (Brit. Mus., no. In. 18431) the wing was well flattened out before being
buried, and the origin of the veins and their true position can now easily be
determined. This specimen shows that the radial sector arises much nearer the
wing-base than the first outward branch of the median — a feature not usually
shown in the remaining specimens.

The general build of the wing is such that the bases of the costal and subcostal
veins on the outer margin, and those of the anal veins on the inner, must have
served as the main support to the distal two-thirds of the wing, the latter consisting
mainly of the radial sector and its branches, the median, and the distal half of the
cubitus. Flight must have been mainly maintained by the action of this more
distal expanded area, while the strain of movement would fall across the narrow
neck-like base of the wing, and may ultimately have led to fracture and the loss of
the wings. It is quite possible, also, that these insects were capable of finding
food among the decaying vegetation of the coal forests, and thus prolonging life
for a considerable period after the wings were lost. Such a presumptive sequence
of events would account for the total absence of any trace of the bodies or legs,
although the wings of this species are more numerous than any others in the Coal
Measures and preserved in good condition.

I'ulour Hmnlx. — Scudder mentions this species as the most striking instance
among Palaeozoic insects of the preservation of "colour bands," and as he states,
some wings show three broad irregular belts of dull umber-brown colour across
the wings. Close examination of these " colour bands " in the type-specimen, and
in other examples, leads us to doubt the correctness of his view. In all cases
where the " colour bands " do not show on the wings, the areas appear to be

62 FOSS1I, 1XSKCTS OF THE BRITISH COAL .MEASURES.

totally destitute of any traces of the wing-membrane, and the course of the vein,-
only is slio\vn across the matrix. It is much more likely that the "colour-band "
effect has been produced by conditions of preservation owing to the wing-
membrane being destroyed in those areas which do not show colour.

Aijiiiitii'x. — Although Scudder founded both genus and species, he did not
attempt anv diagnostic description of either, confining his efforts mainly to a
discussion of relationships. His figure is unusually pour, and adds nothing to the
text. Scudder's general conclusion was thai the wing was neuropteroid in
character, but "refusing Lo affiliate closely with the restricted families of the
present day." A manuscript note in Brodie's handwriting placed with the type-
specimen would seem to show that Scudder's views acquired greater definition
later. Hrodie writes under date. February, 1880: "I sent this wing to Mr.
Scudder, and he supposes it to belong to the white ants (Termitida>). or close to
the group of which (ioldenberg's Ihi-l ijnin'ii rn is the best type." The first detailed
descriptive note of the species was published iu IS'Jo by Brongniart, with an

FIG. 16. — Brodia iinscotinftn. Scudder; immature wing, twice natural size. — Middle Coal Measures
(clay ironstone nodule from binds between the " ISroncli " and "Thick" coals); I'oselev,
Staffordshire. Madeley Collection, Brit. Mus. (no. I. i'J6li). -4, anal; 0, costa ; (.'», cubitu.-;
M, median; U, radius ; Rs, radial sector ; Sc, subcosta.

excellent enlarged drawing. Brongniart very doubtfully assigned the wing to the
Protodonata, and " alongside the Campyloptera."

Immature ]riu<jx. — The collection of insect-remains which I describe later
(p. 07) under the name of " Pteronepionites " were found at the same horizon and
localities as Hrm/in priscotincta, and il is fairly certain that some of them are
immature forms of this species. It will be noted that these larval forms are broad-
bodied and well segmented, and with lateral outgrowths of a pleura-like character
upon the abdomen. They indicate that ]3rodiapriscotincta went through a progres-
sive metamorphosis, the rudimentary wings gradually developing as the insects
lived as ground-feeders among dank and rotting vegetation.

One of these wings (I'l. IV, tig. '•}) in the Madeley Collection in the British
.Museum (no. I. 2966) is an impression is mm. in Length by !• mm. in
maximum breadth, contained in a small grey ironstone nodule. A.> is usually
the case with these grey nodules, the details of the wing are much obscured
by the matrix, and the precise method of division of the veins is lar Iroin being
clear.

The outer margin is regularly convex and formed by the costa. Il gradually
merges into I lie expanded and well-rounded wing-apex. The >ubcosta extends

BRODTA PEISGOTINCTA. 63

the whole length of the wing, ending in the apex, and being parallel to the costal
margin. A wide area separates it from the next vein.

The course of the radius is similar to that of the sul>costa. The point at
which the radial sector arises cannot be determined, but it lies in the base of
the wing.

The radial sector is parallel with the radius over the greater part of its length,
and gives off three inward branches in the distal third of the wing, all of which
end on the inner side of the wing-apex.

The course of the inner third of the median is indistinguishable. The main
stein reaches the middle of the wing before it gives off the first outer branch,
which bends round and becomes parallel with the inner branch of the radial sector.
The second branch arises a little further out, and at a more acute angle than the
first, passing down to the inner margin of the wing midway between the first
branch and the main stem.

The cubitns is a long undivided vein which passes, first in a curve, and then
in a straight line to the inner margin.

Lying inward to the cubitus are traces of another undivided vein, which may
represent the anal. It bends inwards more rapidly than the cubitus, and the
interval between the two veins is very wide. The inner margin of the wing is
sigmoidal in outline.

The general characters of this immature wing are nnmistakeably those of the
genus Pn-odi<t, and only the absence of spinules on the costal margin and radius,
and the lack of a fourth branch to the radial sector, distinguish the specimen from
adult wings of II. priscotim'ln. These details are not of specific value, and may lie
due to the immature condition of the wing, which is lint one-third the length of
that of a normal T>. priscotlncta.

The general characters of the wing are much like those of UrnJin /_>rixcotiiieta
(/iu'enis), and the specimen may be a slightly older larva of that form. The
outline of the wing is much the same, but the branching of the principal veins is
better shown and their apical curvature less pronounced.

The wing also lends considerable support to the belief that these insects did
not possess a resting or pupal stage, but that the metamorphosis was regularly
progressive.

Brodia priscotincta, Scudder, fm-inn jitreiii*, Bolton. Plate IV, figs. 1,2; Text-
figures 17, IS.

1919. Timlin nijiiijiJta, Hamllirsch, Revision tier Palaozoischen lusekten, p. 76, fig. 90.

T</l>e. — A pair of wings, one almost complete, the other showing the apical half
only; British Museum (Johnson Collection, no. T. 1 •"><>:!).

(H FOSSIL INSECTS OF THE B1UTTSIT COAL MEASURES.

ll<ii'i::nii KHI] Lucnllfi/. — Middle.' Coal Pleasures (binds between the " Brooch "
and "Thick" coals) ; Coseley, Staffs.

Description. — The fossil is contained in a brown ironstone nodule, the whole
wing- having a length of 21 mm., and a maximum diameter of 4 mm. It is
membranous, and oblanceolate in shape. The outer wing-margin, of which only
the distal half is preserved, is flatly convex and curves inwards, meeting the inner
margin in a rounded apical angle. The inner margin is feebly concave. The wing-
has three longitudinal folds, and the extreme tenuity of the integument is accom-
panied bv a corresponding thinness of the veins. The greater portion of the costa
and the whole of the subcosta are missing. The outer portion of the radius is present.
It is parallel with the outer margin, and curves round into the wing-apex. The
radial sector is parallel with the subcosta, and curving round into the apex is lost,
by reason of its tenuity. Lying inward to the main stem of the radial sector in
the distal half of the wing are two long veins, and traces of two others, all
following the same course, and curving to the inner margin. The two long veins

Fi<i. 17. — TiroJln in Iscotincta, Seuilder. fin-mn juvenis, I!'>lton ; diagram of wing-neuration, t\vo-nml-a-
half times natural si/.c. — Middle Coal Measures (liimls lie) \\een the " llvoorh " ami "Thirl;"
roals): CVseley. Stall'i ii'dshi re. Johnson Collection, I'.rit. Mus. (no. I. 15(>:i) 1 lettering as in
Text-figure Hi', p. 62.

appear to arise from the radial sector, in which case the two of which traces onlv
are seen would do so also. The radial sector therefore seems to give origin to
four inwardly directed parallel brandies. The whole course of the next vein is
not clearly determin able. It passes to just beyond the middle of the inner margin,
in an almost straight oblique course, and gives off a single forked outer branch
parallel with the fourth branch of the radial sector. This vein can only be the
median. The next vein is the cubitus. Ft is undivided, and goes to the middle of
the inner margin. Anal veins are only indicated by feeble traces of a single

< . O

undivided vein, which apparently reached the margin midway between the cubitus
and the base of the wing.

Another similar wing from the same hori/.on and locality in the British Museum
(no. I. 1564) is 17 mm. long and "> mm. wide, and lies on the surface of a split
nodule of dark-brown ironstone. The wing-membrane is very thin, and forms a
slight gla/e on the otherwise granulated surface — a feature which has made the
details of structure difficult to determine. The costa is marginal, the outer margin
convex, and gradually curving into the rounded apex of the wing. The inner
margin is slightly convex distally, and straight proximally.

The Mibcosta is a feeble vein whose course cannot be traced with certainty

BROD1A PRISCOTINCTA. <>:•

beyond the middle of the wing. It is close to the margin and parallel \vith it For
its whole lengtli. The radius is well marked, sunken, and also parallel with the
margin except distally, where it is more inwardly curved. It ends on the apex in
a short fork. The radial sector arises near the base, and is parallel with the radius.
It appears to give off three to four branches, the first arising very near the base of
the wing. The whole course of this branch can be traced, but of the middle two
only faint traces are left in the region of the wing-apex. The fourth branch is
very short, and corresponds in position with the last branch of the radial sector in
BroiUci priscotincta. The median vein forks low down into two equal branches,
which reach the middle of the inner margin.

The cnbitus is a long undivided vein passing out almost to the middle of the
inner margin, and separated from the next vein, which seems to be the first anal.
The course of the latter is more oblique to the inner margin than that of the
cubitus.

In its general character this wing is distinctly similar to that of 13. priscotincta,
although it lacks a second branch to the median, and the first anal does not seem
to be forked.

FIG. 18. — Krodin priscotinctn, Seudder./nnmi /'nwnis, Bolton ; diagram of wing-neuration, twice natural
size. — Middle Coal Measures (binds} between the "Brooch" and "Thick" fals I ; Coseley,
Staffordshire. Madeley <'»ll.'eti.>n, Brit. JIus. (no. I. 1504). Lettering as in Text-fit^-. Hi. }< H-'

. I /////// (V.s-. — Handlirsch regards these as nymphal wing-sheaths, bat their
extreme tenuity militates against this view. He is probably wrong in referring
Ili-inliii to the Megasecopterida?, but right in regarding these wings as those of
heterometabolous insects, and not holometabolous as Lameere supposed.

Immature though the wings undoubtedly are, the more nearly perfect example
possesses an assemblage of characters which I believe points to its relationship.
The shape of the wing, the character and course of the subcosta, the number and
position of the branches of the radius, the simple forking of the median and the
anal veins, are all characters pertaining to the genus Ili-mlin. The differences in
detail between this wing and that of />'. priscotincta are such as may be looked for
between the nymph-stage and the adult. Among previously described larval wings,
the onlv type which seems comparable is J.nnu'ri-i'ifi'x r// /•/•/'/»•// ///x, Handlirsch, based
on four nymph-wings or wing-cases, as Handlirsch has them, found in the dial
Measures (Pennsylvanian) at Mazon Creek, Illinois, U.S.A. (I'.'ll, Handlirsch,
' Amer. Journ. Sci.' [•!•], vol. xxxi, p. :375).

In these wings, the outer or costal margin is more strongly curved, and the costa
and radius extend over the bluntly curved apex down to its junction with the
inner margin. The succeeding veins arise nearer the base of the radius, only one
9

C.r, FOSSIL TXSECTS OF THE BRITISH COAL MEASFRKS.

apparently being a branch of that vein, while the median and the cubital areas are
occupied by numerous veins whose origins are nut indicated.

The differences between Lameereites dtrvipennis and this specimen are consider-
able. The balance of evidence is greatly in favour of an affinity with llroiJin /irix-
i-ntinclii, and the specimen may represent a nymph or larval stage of that species.

It is undesirable to attach a specific name to immature wings agreeing so
closely with a known species. 1 propose to regard it as I!, prisnotincto,, j'nrinn
juvenis.

Brodia furcata, Haudlirscli. Plate III, figs. 7, S; Text-figure !'.».

1919. li rail iii J'urciita, Hanillirscli, Revision <ler Palao/oiseheii Insektt-n. p. 75, fig. xn.

Tt/{>r. — A left wing and impression showing the under-surface ; British Museum
(no. I. ;29G'2).

Horizon aiiJ Lix'nUtij. — Middle Coal Measures (above the '; Brooch " Coal) ;
Dudley and Coseley, Staffs.

FIG. 19. — Bfnilin J'ui'rutu, flandlirsch ; showing forking of the second branch of the median into two
equ;>l twi^s, natural size. — Coal Measures (clay ironstone nodule from binds between the " Brunch "
and "Thick" coals) ; Coseley, Staffordshire. Brit. Jlns. (no. I. :.'!>( iiM.

tijii'r/jir (.'Inifiifti'rx. — Radial sector reduced in area, and possibly with three
branches. Median having the second branch dividing into two equal twigs, both
of which reach the margin. Median area, enlarged.

Description. — This wing differs so much from the type-form as to be worthy of
specific distinction. Its total length is 44 mm. and the greatest width 1 Ll'5 mm.
The base of the wing is much more nearlv complete than usual, and very narrow
(•> mm.) for a short distance, bevond which it widens by the development of the
strong convex inner margin. The wing shows the usual plication, which fortu-
nately is not continued into its base, so that the course of the veins in the latter
is not obscured, as is so often the case in I!, firixfutiiirlti. The outer or COM a I
margin forms an almost straight line, and bears a double row of spmnles, those ai
the base of the wing pointing inwards. No basal hum]), as in //. /irixrotiiirlit, is
shown. The subcosta is widely spaced from the outer margin proximally. The
radius and radial sector present no special features, and but one inward branch is
present. Whether four branches arose from the mam stem, as in the t vpe-species,
cannot be determined owing to the loss of the apical portion of the \\IIIL;'. It is
doubtful if such was the case, as the portion missing is not great. The outer three
branches of the radial sector are usually 10 — 1 12 mm. apart, so that if the wing

" PTERONEPIONITES." 67

possessed the same number of branches of the radial sector as in B. priscotincta,
its total length would have been 70 mm. — an unusual length. The median vein is
the most powerful of the whole series, and occupies a middle diagonal area equal
in extent to the combined costal and radial areas. The first branch arises in line
with the radial sector, and much nearer the base of the wing than in B . priscotincta,
and also lies so much nearer the cubitus that the area separating the two is but
half the diameter of the area in the former species. This shortening of the main
stem and its movement inwards has been brought about by the division of the
second branch into two equal twigs, which pass out to the margin between the
first branch and the main stem. The median vein therefore takes a larger share
in the wing-structure than in B. j>r!sc.ottiH-tii, and the radial and cubital areas are
correspondingly reduced. The cubitus presents no special features. Notwith-
standing the shortening up of the area separating it from the median, the area
between the cubitus and the anal veins is very large. The cubitus has suffered no
displacement by the increased division of the median. The first anal vein is very
long, passing well beyond the first third of the wing and dividing just before
reaching the wing-margin. The second anal vein is two-thirds the length of the
first, and bends inwards more gradually to the margin, giving off two short oblique
branches in its basal half.

" PTERONEPIONITES."

Many larva.' of fossil blattoids have been recorded, but very few of other
groups. These larva? may eventually reveal the changes undergone up to the
adult stage, and the development of the neuration of the wings. In addition, the
occurrence of larva? in deposits is more likely to be indicative of habitat than the
presence of adult wings, as the inability of larva? to fly and their lesser power of
flotation would ensure inclusion in adjacent deposits. It is therefore necessary
that their occurrence, and as much as possible of their structural appearance, be
fully recorded. Any attempt to classify them under genera and species would
rather retard than accelerate progress, and it seems advisable to record them
under some term which will leave no doubt of their larval character. Handlirsch
(' Amer. Journ. Sci.' [4], vol. xxxi, p. 375, I'.Ul) has already described
larval "wing-cases" of a somewhat similar character under the generic name of
" Lameereites," and placed them under the order Megasecoptera.

Had not Handlirsch given a generic value to the name " Lameereites," it would
have been possible to extend the use of his term to all larval wings. Failing this
I would suggest the use of the word " Pteronepionites " for all larval wings which
cannot be referred to a known genus, adding a specific designation when any larval
wing presents features of a definitely recognisable character. Handlirsch restricts
the name " Lameereites " to the wings of larval Megasecopteridie, but recognises the

68 FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

close similarity between them and Pteronepionites (' Revision der Palaozoischen
Insekten,' p. 76).

"Pteronepionites" johnsoni, sp. nov. Plate IV, fig. 4; Text-figure 20.

T//JII'. — Immature wing, 12 mm. long and 4 mm. wide; British Museum
(.Madrley Collection, no. I. 2907).

Horizon mill Locality. — Middle Coal Measures (binds between the "Brooch"
and "Thick" coals); Coseley, Staffs.

I ti'.-ifi'ijilliiii. — The wing, like all these immature structures, is of extreme
temutv, and the finer details are masked by the coarse granular nature of the
matrix composing the nodule. The outer margin is straight, curving backwards
until it meets the inner margin in a bluntly-pointed apex. The inner margin is
strongly convex, a distal infolding indicating that the wing was apparently never
fullv extended. Very feeble traces occur of a short straight snbcosta, which

Kn, 20. — " I'ti'niiifjiiiinitft. " ./H/IHSMIII. sp. nov. ; iliu^i-iiin of win^-nrunitioi), t \v.,-;iinl-a-h:ilf times
natural si/.e . — JliiMIc Coal Mi MM ires (l.imls Iir1 wn-n tlic " Hn.ocli " and " Thick " ro;tls) : Coseley,
StaftYirtlsliiiv. .Ma.lclcy GVllivtinii, Brit. Mus. (no. I. ^'.H;TI

reached the margin near the apex of the wing. The radius is stout and straight
and reaches the wing-apex, giving off a well-marked radial sec-tor, which, diverging
from the radius, passes in a wide curve into the wing-apex. The median remains
undivided for nearly a third of its length, and then bifurcates into two eipial
branches which assume a parallel position, and reach the outer third of the inner
margin. The cubitus is represented by two veins, the first having a sigmoidal
sweep and the second a simple curve. Until reach the middle third of the inner
margin. No trace of anal veins can lie observed.

"Pteronepionites" ambigua, sp. nov. I'late IV, fig. 5.

/'///"'• — A pair of larval wings still attached to the crushed and a most
obliterated body; Uritish .Museum (Madeley Collection, no. I. 2'.Mi,S).

Horizon <m<l Locality. — Middle Cnal Measure's (binds between the "Brooch"
and "Thick " coals); Coseley, Staffs.

Description. — The insect lies in a fragment nf a liidit grey ironstone nodule,
and is not well preserved. One wing is whole, but mure than hall ol Ilie second
is niissiiiL!'. and it> ape.x is obscured by the matrix. The whole wing has a length
nl li'.i mm. and a width of 2 mm. Ivich is much thickened over the basal hall.

" PTERONEPIONITES " LKITS. li'.i

and strongly ridged or furrowed along the line of a powerful vein or tracheal
trunk (the wing is so immature as to render the latter possible), occupying the
position of the radius and median veins. Traces of a third vein occupying the
position of the cubitns are present on the inner half of the wing. The body-
segments are numerous, o — t mm. in depth, and seem to have had pleura-like
expansions. The region in front of the thorax is bent backwards and below the
abdominal segments. The thoracic segments are larger and more robust than
those of the abdomen.

The thinness of the body, and the difficulty of determining boundaries Avith
satisfactory accuracy, render all attempts at a more precise determination impos-
sible. We can, however, say with confidence that the remains are those of an
insect possessing a long segmented body, an elongated head-region, and wings
carried upright over the back.

" Pteronepionites " lepus, sp. nov. I'late IV. tig. <>.

e. — Remains of a larval insect, having a segmented body and two immature
wings, in a flattened nodule of light-grey ironstone ; British Museum (Madeley
Collection, no. I. 2%'.M.

Horizon and Lonilitt/. — Middle Coal Measures (binds between the "Brooch"
and " Thick " coals) ; Coseley, Staffs.

Description. — The impression of two wings is clearly discernible, the segments
of the abdomen are less so. With oblique lighting, six segments can be
made out behind the thorax. The wings are slightly unequal in si/.e, and the
anterior is longer and thicker than the posterior wing; it is also somewhat
infolded at the base. The differences between the two wino-s, and their relative

o 7

positions, would seem to indicate that they are the fore- and hind-wings of the
one side.

The fore-wing has a length of '.» mm. and a width of -^> mm., while the hind-
wing is 8 mm. long and 2 mm. \vide. The fore-wing has a strap-shaped
appearance, and the line of attachment to the body is its broadest part. A stout
ridge, swollen at its junction with the body, traverses the greater part of the
length of the wing, dying out before the apex is reached. The wing is too small
for definite determination of this swelling, but it appeal's due to incomplete
extension of the wing-membrane, rather than to the presence- <>f a vein. The distal
fourth is flat and thin, and the apex well rounded. The outer and inner margins
are parallel and undulated. A few faint and irregular lines may be indications
of veins.

The hind-wing has undergone greater expansion than the fore-wing, and lies
almost flat on the nodule. Both outer and inner margins are gently convex. th.e
convexity of the inner margin being the greater. Feeble traces of a subcosta. and

70 FOSSIL INSISTS OF THE BRITISH COAL MEASURES.

of a stout vein which forks before the middle of the wing is reached, are dis-
tinguishable. The latter vein occupies the position 1 should assign to the radius.
No further traces of veins are visible.

The two wings are separated by an interval of :>> nun. at their bases. Xo
definite details can be made out in the thoracic region. Behind the hind-wing
are faint impressions of a series of abdominal segments. These are about three
times as deep as wide, and appear to have borne lateral spiny processes. There
are also traces of tubercles.

The head-region is only indicated by faint discolorations. As in the case of
other examples of " Pteronepionites " we have met with, the remains are so filmy
in character, and merge so much into the matrix, that it is impossible to define the
outer limits of the various segments with absolute clearness, and no attempt can be
made at a systematic determination of characters.

The specimen is a larval form with wings not yet fitted for Might, but with a
degree of differentiation in the fore-wings which indicates that they were thicker
and less flexible than the hind-wings.

The abdomen is long, wide, and well segmented, the lateral expansions being
not unlike those of Kuphuleria J'ern.r. So closely does the abdomen resemble the
segmented body of a Diplopod, that in the absence of the wings we believe it
would be readily classed as belonging to that group, and as possessing nothing in
common with insects.

It is difficult to resist the belief that these larval insects were capable of
crawling about in decaying vegetation, and that their larval life was thus spent,
until by successive ecdysis the wings had acquired sufficient strength to lift the
body from the ground, -and enable the insect to fly.

Family .KxniMATonin.K, Handlirseh.

190IJ. Handlirseh. Prou. U.S. National Mils., vol. xxix, p. 680.
11)0(3. Haudlii'scli, J>i<' Kossilcii [nsekten, p. 116.

Wing stronglv arched, and broadly rounded at the apex. Anal area small, and
not marked off from the rest of the wing Subcosta reaching almost to the wing-
apex; radius simple, and radial sector with three divisions. .Median with lour
brandies. Costa represented bv an oblique vein with a terminal lork, followed by
three simple, stronglv curved anal veins. Interstitial neuration partly of regular
ei'oss-nervures, and partlv ol a polygonal network.

Handlirseh founded this family on an incomplete wing in which the greater
part of the outer border, sulieosta and radius is missing. From the upper Middle
Coal .Measures of .Ma/.oii Creek, 111.. I .S.A.

.ENIGMATODES (?) REGULAR IS. 71

Genus ^NIGMATODES, Ihmdlirsch.
1900. -^iiii/ii/iifiitlfft, Handlirsoli, Die Fossilcn Insekten. ]j. 111!.

Generic characters as above.
^Enigmatodes (?) regularis, sp. nov. Plate IV, fig. 7 ; Text-figure 21.

Ti/pi'. — Fragmentary wing; British Museum (no. Tu. IS, 604).

Hori::un <ii//1 Loi'iilltij. — Middle Coal Measures (over the Barnsley Thick Coal) ;
Monckton Main Colliery, Barnsley, Yorkshire.

^l»'i'i'fir I'/Kirnrfi'i-x. — Radial sector, median, and cubital areas occupying
most of the wing, and all with well-spaced branches united by straight nervures,
except in the marginal area between the median and cubital veins where they
form a slight mesh work. Inner margin well rounded.

l^'xiTijition. — Little more than half of this wing is preserved, and lies on a
fragment of hard grey bind associated with broken-up plant-remains. The
greatest length of the fragment is 42 mm., and the width 1-j mm. The length

Cu

FIG. 21. — .F.niijinntmli'* ('.') ir</ii'»ns, sp. nov. ; iliagram of neuv.it.ion of wing-fragment, one-aii'1-a-half
times natural size. — Coal Measures (over the Barnsley " Thick " coal) ; Monckton Main Colliery,
Bavnsley, Yorkshire. Brit. Mus. (no. In. 18,604). Lettering as in Text-figure Hi. p. <>2.

of the complete wing was probably 50 — 00 mm., and the width 18 — 20 mm. The
outer margin is missing, and of the subcosta and radius only portions of the basal
third are present. These are separated by a narrow area crossed by short trans-
verse nervures.

In the distal third of the wing is the innermost branch of the radial sector,
which ends on the margin in a small fork. The line of fracture of the wing has
closely followed the line of the radial sector, and a portion of another branch of
the radial sector may have been present along the line of the extreme broken edge
of the wing. The median vein sweeps in a convex curve to far out on the inner
margin, giving off two well-spaced branches. The next two veins, one of which
forks, are more oblique in their course, and may also belong to the median, but
the basal curve rather indicates that these veins are cubital, as is the succeeding
vein, of which only a small portion is left. A trace of an anal vein is present near
the wing-margin. The interstitial neuration consists of well-marked transvei-c
nervures, regularly spaced in the outer parts of the wing, and uniting in a loose

FOSSIL IXSFCTS OF THE BRITISH COAT, MEASURES.

mesh work in the wide area between the inner branch of the median and the
cubitus.

.(////////(.•.-•. — The nearest approach to a wing of this character is that of
jffinigmatodes ilmiiclxi, Ilandlirsch, in which the greater part of the outer margin
is also missing. The Yorkshire specimen, which may be a hind-wing, is three
times as long as .!•'. tliti/i<'lxi, and the veins have a stronger inward curvature.
In the absence of more definite details, it seems best provisionally to refer the
specimen to the genus /Enigmatodes.

Genus PSEUDOFOUQUEA, Handlirsch.
UM.iti. Pseudofouquea, Hancllirsch, l>ic Fossil™ Tnsoktcn. p. 125. «

Wings three times as long as wide. Cubitus with inner and outer branches.
Anal veins, so far as known, not united. Interstitial neuration of feeble cross-
nerviires except between cubitus and tirst anal, where it is irregularly reticulate.

Pseudofouquea cambrensis (Allen). Plate IV, tig. S ; Text-figure 22.

liMll. l<'i,in/in':i cambrensis, Allen, (icol. Alag. [4], vol. viii, p. (!5. text-fig, on p. I'll.
I'.MUI, P.-ifti/liit'in/i/iifi.! i^niiliffnaif. Iliunllii'Sc-li, l>ic Fossileu Insekten, p. 125, pi. xiii, fig. 5.
1!>1<;. ra,-iiiliifnn<ii<ni cambrensis, 1'mlion, (,>ii;ivi. Joiirn. Geol. Soc., vol. K\ii. p. 5!*, pi. iv, figs. 4 — 5,
text-fiff.

Ti/jx-. — A broken left fore-wing, of which the two parts are preserved on
fragments of blark shale: one fragment, bearing the basal part of the wing, in
the Museum of I'raclical Geology, .lermyn Street (no. 7-72). the other, containing
the impression of the distal 2S mm. of the wing, in the \Yelsh .National Museum.
Cardiff (no. 13,120).

//"/•/.v«// mill Lix-itlit//.- Lower Coal Measures (top of the Four Foot Seam);
Llanbradacli Colliery, near Cardiff.

i^pei-ipr ''l//i r<ir/i-i-x. --\Vings stout and obfuselv pointed. Costa marginal and
flatly convex. Snbcosta reaching margin in the outer third. Radius parallel with
subcosta. and ending in apex of winu'. Radial sector diverging widely trom radius,
and endiiiL:' on the win"'- apex in live divisions. Comstoclc regards it as a typical
dichotomies radial sector, with an accessory vein on the second branch (' \\ings
of Insects.' I'.'iS). Radius and radial seel or occupy ing all the wing-apex. Median
dieliotomonslv branched, with an accessor} vein in the fourth branch. ('ubitn-
widely divergent from median, ami giving off allernate twigs from its outer and
inner sides, those nf the inner side being much the Feeblest. Anal veins lour in
number. Interstitial nenralioii of feeble cross-nervures. except between the base
of tin- fii'sl anal vein and tin- cubitus, where it is irregularly ret iculal e.

Description. -The lo|;d len^lh of the wing is now :'>2 mm.. '.' mm. having been

PSEUDOFOUQUEA CAMBRENSTS. 73

lost from its tip since it was measured by Allen, who gives the total length as
-II mm. The greatest width is 1 •"> mm.

The subcosta is widely separated from the outer or costal margin in the base of
the wing, and gradually approaches and unites with it beyond the middle. The
radius is parallel with the subcosta throughout its length, and gives origin to the
radial sector basally. The radial sector diverges from the radius over the whole
of its course. It now shows but one inwardly directed branch, which forks at the
broken edge of the wing. Allen's figure indicates that two more branches were
given off, both being undivided. The radius and radial sector occupied almost the
whole of the apex of the wing.

The median vein forks in the basal fourth, the outer branch again forking
before the middle of the wing is reached. The inner branch of the median diverges
almost in a straight line from the outer branch, and also forks, and bears an
accessory twig upon the fourth branch. A feeble accessory twig appears to be
given off near the middle of the Aving, but dies out in the integument. The cubitus

Fia. ~2. — Pseudofovquen nniifuYiisis (Allen) ; restoration of whole win^, natural size. Lower Coal
Measures (top of the Four Foot Seam) ; Llanbradach Colliery, near Cardiff. Basal portion of
wing in Mus. Pract. Oreol. (no. 727-) Impression of apical portion of winy in the Welsh National
Museum (no. 13,120).

is a remarkable vein, unlike that of any other fossil wing (Comstock, ' Wings of
Insects,' 1918, p. 106). For nearly half its length it passes in a broad curve to
the inner margin, giving off a series of alternate twigs upon its outer and inner
sides, those of the inner beintr weaker than those of the outer side. The two

o

outer twigs are strongly developed, while those on the inner side of the cubitus,
four in number, are weaker and shorter. The feeble continuation of the main stem
reaches the* margin between the two sets of branches. Four anal veins are
distinguishable. The inner two arise from a common base, the outer two not
uniting. This is unlike the condition in Fnui/iimi. where the anal veins branch off
regularly from a single stem.

The area lying between the first anal vein and the main stem of the cubitus is
verv wide — much wider, indeed, than any other area.

The interstitial neuration consists of weak cross-nervures, except between the
base of the first anal vein and the main stem of the cubitus, where it is irregularly
reticulate.

Aljhiittffi. — The characters of the cubital and anal veins definitely remove the

species from the genus Finn/in-n, and the cubitus, with its strong, anteriorly

directed twigs, and its feebler inner series, is wholly unlike that of any other

insect, and would alone suffice to justify the generic rank given by Handlirsch.

10

74 FOSSIL INSECTS OF THE BEITISH COAL MEASUKES.

So far I am in agreement with Handlirsch, but 1 regard the enlarged areas
between the inner divisions of the radial sector and the cubitus, and between the
cnbitus and the anal veins, as more suggestive of the Protorthoptera, notably
Tln>i-uiii/xis iiif/l>t>rft'iix!x, Aminon. More than this cannot be said, and /V/i//,i-
/'iiiii/iK'it cambrensis must be regarded provisionally as Palreodictyopterid, with a
possibility of Protorthopterid or even Orthopterid affinities.

INCERT^E SEDTS.
Genus ARCHJEOPTILUS, Scndder.

1881. Arcli:n>]>til>is, Scudder, (leol. Mai,'. J2 ', vol. viii, p. 2'.'.V

A wing of unusually robust type. Only one is known, consisting of not more
than the basal fifth of a whole wing whose total length may have been 25~4 cm. to
o5'5 cm. The fragment is too small for a correct determination of its systematic
position, and has been referred to widely separated families by various workers.

Archaeoptilus ingens, Scudder. Plate IV, fig. (.»; Text -figure '2:>.

1881. Archn'optihis iiiijcim, Scudder, Geol. Mag. [2], vol. viii, pp. 295, 300.

1883. Ar<'Ji;>'<i1itiln.t /'//;/(•//.-•, Scudder, Mem. Bost. Soc. Nat. Hist., vol. iii, pp. 217, 22o, pi. xvii.

tigs. 10—12.

1885. Arclin'fuitihix ni//r//x, llnm-mail. Bull. Soc. Amis Sci. Nat. Rouen [3], turn, xxl, p. (JO.
1S85. An-li;i-'i/ililiif ///i/i //s. SiMiddi'i-, Zittcl's llaudlnicli der Pahi'ontoloi,'ic', vol. ii, p. 757.
lsc.i;3. Ai-<-li;i'tijit!lii* iiiiji'ii*, Iii-on^uiart, Faune Kutmn. Temps Prim., p. 1-08. j>l. xxxvii, fig. 6.
.l/-i7/,-n»////7//x Ingens, Handlirsrh, Die Fossilen Insekten, ]>. 117. pi. xii, fig. 18.

'/'///"'. — Basal fifth of wing, in counterpart ; British Museum (no. T. 3!»97).

llnrr.i'ii inn/ l,iicnli/i/. — Middle I'pper Coal Measures; between Shelton and
Clay Lane, near Chesterfield, Derbyshire.

N/)ir///Y Characters. — Wings very large; cosia. subcosta, and radius broad
and robust. The costal border spiny. Interstitial iienration of stout li-ansverse
nervnres.

I >i'xi-ri]ilimi. — Onlv the basal part of the wing and its counterpart are
preserved, having a total length of !-'> mm., and a greatest breadth ol ']•} mm.
Scudder's estimate of the length of the whole wing as o-V'i cm. is probably
excessive.

Scudder (/i/c. o'/., 1SS1) thus deserilies the specimen: "All the principal veins
are a millimetre or more thick, and the cross-veins of the upper interspaces are
tolerablv distant, stout, prominent, and generally simple. The marginal (costa)
vein, lormiiiL;' the I'ronl (outer) border of the wing is studded with short oblique
spines (r macro! richia). The other veins lie al very different levels on the stone,

AECHLEOPTILUS INGENS.

75

and below the interspaces mentioned, seem rather closely crowded, and much
more curved, sweeping downward, while the upper veins show little tendency to
turn from a longitudinal course."

The great apparent width of the costa is caused by the formation of an
expanded chitinous bur along its outer margin, the free edge bearing the spines
described by Scudder. The costn, with its chitinous bar, the subcosta and
the radius are so broad as to appear strap-like, are widely separated, and the
intervening areas are crossed by equally strong nervures. The costa can In1
distinguished on the cast from the frontal bar and appears as a narrow
rounded vein.

The subcosta is a very broad vein, crossed by oblique stria? which are directed
outward from the upper edge of the vein. A broad interval ('.> mm.) separates
it from the costa, the area being crossed by stout, slightly oblique, transverse
nervures. The general direction of the subcosta is such that it must have reached
the margin of -the wing near the apex. At the base it is much enlarged, the

FIG. 23. — Archaeoptilus imjens, Scudder ; diaynim of whole winy; from Sfiuldrr's on^-iiml restoration, one-quarter
f" natural size. — Upper Coal Measures ; near Chesterfield, Derbyshire. Brit .Mus. (no. I. 3997).

enlargement probably indicating the attachment to the body of the insect. The
expanded inner portion of the base is fused with the equally expanded base
of the radius.

The radius diverges from the subcosta in its outward course, and is even more
enlarged than the subcosta. It is connected with the latter by a series of thirteen
transverse nervures in the wing-fragment. Most of these are slightly convex,
and two are united by lateral branching.

The median vein was apparently closely apposed to, or united with the base
of the radius, and is much less robust than the latter, the intervening urea being
narrow, and crossed by short, thin, straight, transverse nervures which do not
appear to continue into the base of 'the wing. The cnbitus is not so readily
distinguishable, and diverges sharply inwards towards the inner margin. The
anal veins are 4 — 5 in number, and are strongly curved inwards, occupying
not more than one-fifth of the inner margin of the wing. The first three may
have been united at their base. The wing is marked by a series of folds along
the lines of the principal veins.

Afo'.iitiiex. — This remarkable wing-fragment has caused considerable conjecture
as to its true character and relationship. Scudder in his second note (Inf. fit.,

7C, FOSSIL INSECTS OF THE BRITISH COAL MEASURES.

188-3) somewhat vaguely placed it, " with a strong degree of probability, in the
same general group as some other I'ahwo/oic wings." This reference can only be
to the Palseodictyoptera. Later he published a restoration of the wing, and classed
it with the Protophasmidge. Brongniart atone time regarded it as a Dictyoneuron,
and later as a " Neurorthopteron " of the group Slhenaropterida. Still later
he placed it as a Nenropteron in the group Platypteridse. Handlirsch considers
that the costa is marginal, and that this character, with the sharp inward curve
of the anal veins, justifies its inclusion in the Palseodictyoptera, although lie does
not attempt to indicate its allies.

A second species was described by Brongniart as A. ln<-nxi (' Bull. Soc. Amis
Sci. Xat. Rouen' [o], ami. xxi, p. <><), 1885), but this throws no light on the
genus, nor does it appear to be generically referable to Archseoptilus.

The only details preserved which can be used in the determination of relation-
ship are the spiny outer margin, the great width of the principal veins, the
well-developed cross-nervures, and the strongly curved and numerous anal veins.
Even these are too fragmentary for safe conclusions to be drawn in the absence
of other material.

The general structure of the wing-fragment is to me more suggestive of the
Protodonata than of the Palseodictyoptera, but the presence of well-marked anal
veins discounts this view, unless we are prepared to accept the specimen as an
early and archaic prototype of the Protodonata.

Order M I XOTKRMITOIDKA, Handlirsch.

I'.MM;. Handlirsch, Proc. U.S. National Musc'Uiu. vol. xxix. p. <;!>.">. ami Die Fossil™ Inseklt'ii. \>. l^i',.
1919. Handlirsch, Ut-vision <ler Paliiozoisclieu Iiisekten, \>. -i>.

Subcosta much shortened; radial sector arising close to the base, \\itli - — :>
branches, of which only one forks. Median long, four-branched and suggestive of
the Palseodictyoptera. Cubitus with "2 — :.! inward branches. Anal vein :>, simple.
Cross-nervures strong, wide-spread and regular.

This is a provisional order established by Handlirsch to include two forms
only — Mixoter tries luganensis, Ster/el, from the Coal Measures of Saxony, and
Geroneura «•//>-•<>///, Matthew, from the Carboniferous of St. John, New Bruns\\ ick.
North America. l>oth wings show clearlv their Palseodictyopteroid ancesli-y, but
I landlirsch is uncertain whether thev should be brought near to the Protorthoptera
or to the I'ri'liihe.

(ienus GERONEURA, Matthew.
ls!-i;». lirfiiiii-iifii, Miiiihcw. '1'rans. li'ov. Sm- Canada, vul. \i. scrt. i\', \>. .".7.

fJeneric Characters. — \Vingtlirec times aslongas wide; apex obluselv rounded;

GERONEURA (?) OVATA. 77

subcosta short and joining outer margin before middle of wing. Radius and
radial sector occupying almost the whole of the wing-apex. Median, cubitus, and
anal divisions few. Inters! it i;il nenration of stout cross-nervures at wide intervals.

Geroneura (?) ovata, sp. nov. Plate V, fig. 1.

T//I>IJ. — Portion of left wing; British Museum (Madeley Collection, no. I. 29(>-">).

Hon'::nn <nnl Lacalllij. — Middle Coal Measures (binds between the "Brooch"
and " Thick " coals) ; Coseley, near Dudley, Staffs.

Nyirr///V Characters. — Radial sector rising in outer third of wing, with three
divisions. Median vein large, the two outer branches forking in line with origin
of the radial sector, the third branch undivided. Cubital veins few, undivided, and
reaching the distal part of the inner margin of the wing.

I >!'.-:, •njitinii. — The specimen consists of the impression of the upper surface
of the distal portion of a left wing, having a length of o2 nun., and a breadth of
'2(1 mm. The impression lies on the surface of a thin flattened half-nodule of fine
sandy grit, and is but faintly indicated. The total length of the wing Avas
probably from twice to three times the length of the portion preserved, and its
breadth may have been a little more than 2(i mm. The outer margin is gently
convex, and curves into the broadly rounded apex. Very little is left of the inner
margin, Avhich also seems to have been convex. The distal portion of the costal
margin is present for a length of 'JO mm. There is no trace of the subcosta, so
that this vein did not extend much, if at all, beyond the middle of the wing. The
radius gives off the radial sector about the distal third of the wing, the two veins
remaining almost parallel with the wing-apex. The radial sector gives off a single
inward forked vein. The next three veins seem to be divisions of the median.
The first two each give off an outer branch in line with the division of the radius
and radial sector, and the outer of the two also forks before reaching the edge of
the wing. The third vein is single for its whole length, but evidently united with
the second a short distance outside the line of*fracture of the nodule. The
remaining four veins appear to belong to the cubitus. \o anal veins are
distinguishable. All the veins, with the exception of the small forks of the
radial sector and first median, are parallel and Avidely spaced. They are united by
a series of strong, straight cross-nervnres placed widely apart. Xot withstanding
the strength of the veins and of the cross-nervures, the smooth impression of the
wing-fragment seems to indicate that the veins were not sunk below the general
surface of the Aving, as is usually the case.

Aljiiii/ii'x. — The determination of the relationship of so small a wing-fragment
would be difficult were it not for the unusual direction of the main veins, their
mode of branching, and the character of the cross-nervures. These characters are
_a special feature of the order Mixotermitoidea, Handl.

FOSSIL INSECTS OF THE BRITISH COAL MKASrUKS

The wing-fragment must lie referred to this provisional order in the absence of
knowledge of the whole wing-structure. The much-divided median vein is more
comparable with that of Gennn:in-(i iri/xnuL Matthew, than with that of Mixotermes
liigaitensis, Sterzel, and is also correlated with a shorter subcostal vein, although in
IT. irllxiuil that vein extends beyond the point at which the radial sector arises from
the radius. An open series of cross-nervures is present in both genera, as in this
specimen, and both have the same well-rounded apex. The wing-fragment is
suggestive of llfiiii'i-ixfitt orrii/fi/tnlix, Dana, but has a less branched radial sector.
I provisionally refer it to Geroiieura with the specific name of urnln.

Order PROTORTHOPTERA, Handlirsch.

I'.'IM;. Handlirsrh. 1'roc. U.S. National Museum, veil. xxix. p. >'<'.>'<. and l)ic tVissilt-n Iiisekteii, p. 1-3.
1919. Haudlirsch, Kevisiou der Palaozoischen lust'kteu, p. 28.

Head large, with strong mouth-parts, and bearing long slender antenna1;
prothorax large and elongated, and the body strongly built. Legs either uniform
in character and fitted for running, or the hind-legs modified for leaping. Wings
more specialised than those of the Palaeodictyoptera, and capable of folding on the
abdomen when at rest, with the enlarged anal areas of the hind-wings doubled
under, owing to the formation of a fold between the anal area and the rest of the
wing. The principal veins and their subdivisions not so strongly curved inwardly
as in the Palseodictyoptera.

I landlirsch established this order to include a series of insects intermediate in
character between true Orthoptera and I'aheodictyoptera, to which Scudder had
previously given the name of Palaeodictyoptera Neuropteroidea.

Genus JEDCEOPHASMA, Sendder.
1880. .Kilii'iiii/iiisiiin. Scudder, Greol. Mag. :> . vol. ii, p. -('>'>.

(! i' in1 i-'n- Clnii-iirfi'i-K. — Large wings two-and-a-half times as long as wide; inner
margin more convex than miter margin, and curving distallv into the latter.
Principal veins broad and Hat in the basal third, and diminishing in si/,e distallv.
Sulicosta and radius reaching the wing-apex. Median vein with two mam
branches, (lie outer with most subdivisions. Cultit us with two main branches, each
much subdivided. Anal veins numerous. Interstitial neuratioii of irregular
nervures, and a loose meshwork in the wider areas.

^Edoeophasma anglica, Scudder. Hale V, tig. L! ; Text-figure '_!!.

[885. .K<li <'<./< linaiii, i iiinjlii;i, Scinlilcr, Orol. MML;'. :! . vol. ii, p. L.Vi.">. and in /illcl's I landl'iirli ili-r
o^ic. vol. ii, p. 7-V\ tig. '.Ml

^BDCEOPHASMA ANGLIC A. 7'.)

linn;. . 7',Vii'i,y//(,is/mi tuii/lii-u, Haiidlirsrli, Die Fossilcn lusckU'ii, p. l-!-r>, ]>1. xiii, fi^. 4.
IJ'M. ./•;./.,',,/,//, />•///,; luiiilint. r>i.l1(in, Quart. Jniivn. Gcol. 8oc.. vol. Ixxii. p. 4:!, pis. ill, iv, ami
text-figure,

V//7"'. — Greater part of a left wing in an ironstone nodule ; Liverpool Museum
(presented by Major Chamber* in 1858).

\\tn-\-.ini inul I'jfii-filiti/. — j\liddle Coal .Measures; South Lancashire (locality
unknown, but the nodule so similar to those derived from the Ravenhead Railway
Cutting- that it may be from that section).

$jit',-ijii- ('liitrartrrx. — As generic characters.

])esci'ii>li<>n. — The specimen was partlv described and named by Scndcler in 1885,
and re-examined and figured by the present writer in ]'.»1G.

The wing lies in counterpart in a fine-grained ironstone nodule, and its total
length as now exposed is S7 mm., its greatest breadth (across the middle) •!•<> mm.
When whole, the wing was probably 100 mm. long.

Fir,. lil. — .Etlienjihitsma nnijllcii, Seudder ; restoration of whole wing, showing the general character of
the venation, natural size. — Middle Coal Measures; South Lancashire. Liverpool Museum.

The outer or costal margin is gently convex. The subcosta is a broad flat
vein, gradually diminishing in width towards the wing-apex, which it just fails
to reach.

The radius is an even broader vein than the subcosta, is also flattened in its
basal third, and reaches the outer angle of the wing-tip, keeping parallel with
the subcosta.

The median vein divides low down into two equal branches, the outer giving
off four inwardly directed twigs. The first of these remains undivided ; the second
forks twice, and the outer and inner divisions of the second bifurcation again
divide, so that the vein ends on the apical margin in six divisions. The remaining
two branches are undivided. The divisions of the outer branch of the median
occupy the greater part of the wing-apex.

The inner branch of the median does not divide until it has reached the apical
fourth of the wing, where it gives off four twigs which pass inwards to the
junction of the inner margin with the apex. Only the first of these twigs forks.

The cubitus has the same broad flattened basal portion which is so character-
istic of the veins we have already dealt with. The main stem lies somewhat near

so FOSSIL TXSECTS OF THE BRITISH COAL MEASURES.

the lirst branch of the iiit.'(li;iii, and remains parallel with it over almost the whole
of its length. Inwardly it is separated somewhat widely from a slighter vein
which, I believe, joined it near the base, and formed the first inward branch. The
main stem sends off at the middle of the wing a strongly curved lirancli which
bends first inwards and then outwards towards the apex, breaking up into five
twigs before reaching the inner margin. The second of these twigs forks, A
second undivided branch comes off a little further out, and a third verv small one
almost on the margin. The next two veins were probably united a little war out
from the base, and their direction is such that the single stem from which thev
arose may have arisen. a> suggested above, as the first inward branch of the
cubitus. The outer of the two veins is undivided, and reaches the inner margin
beyond the middle of the wing. The innermost vein runs fairlv parallel with the
first along its whole length, giving off, as it does so, four inwardly directed t \\ igs,
of winch the first and fourth fork. The whole vein ends on the margin in six
twigs. Four anal veins are shown, one only forking.

The interstitial neuration of the radial and median areas consists of straight
or slightly curved nervures, placed at nearly equal distances. The very wide
cubital and cubito-anal areas are filled by a loose meshwork, and a few irregular
wavy nervures. The anal area is crossed by simple straight nervures.

Affinities. — Scudder was originally of opinion that this wing was related to
Meganeura (Dicti/oneura) ntu/ti/i, Brong., representing a member of the group
Protophasmidse. Handlirsch removed the genus to the group of Palaeodictyoptera
incertaa sedis.

Scudder was undoubtedly mistaken in referring I he wing to the Protophasmidse,
as a glance at the figure of Protophasma iltinnixii, Brong., will at once show (' Die
h'ossilen Insekten,' pi. xvi, figs. 1 — 2). Handlirsch did not see the specimen, and
had to base his determination on a sketch of the wing which he considered "con-
fusedly drawn.'' The latter probably accounts for the interpretation which lie
placed on the various principal veins. More recently I have been able to expose
more of the structure, and diagnosed the wing accordingly. If Handlirsch's view
were correct, the radial sector would be of enormou> proportions, and occupy all
the wing-apex. The base of the radius, so f;ir as shown, is widelv divergent troin
the base of the median — more so, in fact, than at anv other part of the whole
course of the radius and supposed radial sector. These veins, therefore, have
come into union onlv at (lie actual point of origin of the wing. This may have
been the case, but in my opinion, the radius is whollv simple and undivided, and
no radial sector is present. The median and cubitus are large, much divided,
and take up the greater part of the wing-area, while the anal veins are lew.

If this view be correct, the wing is a verv primitive example ol the Protor-
thoptera, still retaining evidence in the cosla. Mibcosla and radius, ol its
I ';ihe' ii lid vi ipi eroid origin.

PLATE I.

Fiu. PAGE.

I**. Dictyoneura h'njij'nixii (Handlirsch) ; basal portion of left wing, x 1^.
Middle foal Measures (horizon not known) ; Ravenhead Railway
Cutting, nr. St. Helens, Lancashire. Liverpool Museum. 25.

\li. Ditto; impression of the wing. X H. 25.

'In. Orthocosta xjih'ixli'ii*, Bolton ; left whig. Natural size. .Middle Coal
Measures (below the Top Hard Coal) ; Shipley Manor Claypit,
Ilkeston, Derbyshire. Mns. Pract, Ceol. (Moysey Coll.), no. 30222. '27.

2/». Ditto; impression of the wing. Natural size. Mus. Pract. Geol. (Moysey

Coll.), no. 30223. 27.

3a. I'lrrn/iiiJi/i plicatula, Bolton; inner half of left wing, x H-. The
straight inner margin and the plication are well shown. Middle
Coal Measures (below the Top Hard Coal) ; Shipley Manor Claypit,
Ilkeston, Derbyshire. Mus. Pract, Geol. (Moysey Coll.), no. 30224. :!<».

3//. Ditto; impression of portion of the wing, showing the oblique cross-

nervures. X H. M"us. Pract. Geol. (Moysey Coll.), no. :)0225. 30.

4<i. Hypermegetkes ii<»-flriiiii/u-!;r, Bolton; basal half of a left wing. X 1^.
Coal Measures (shale above the Crow Coal); Phoenix l>rick\vorks,
Crawcrook, Durham. British Museum, no. In. 18524. 32.

4^. Ditto; counterpart of portion of basal half of the wing, the interstitial
neuration having been rendered visible by immersion of the nodule
in water during photography. X 1|. ;>2.

PAL/EONTOGRAPH ICALSOCIETYI9I9

Bolt on , Insects of Coal Measures.

PLATE I.

l.DICTYONEURA. 2 .ORTHOCOSTA . 3. PTERONIDIA.
4.HYPERMEGETHES.

W Tutcher, pKoto

London Stereoscopic Co. imp.

PLATE II.

FIG. PAGE.

!<?•. Cryptovenia iiioi/xfi/i, Bolton ; greater part of left wing, dorsal surface
uppermost. X •).!. Middle Coal Measures (below Top Hard Coal) ;
Shipley Manor Claypit, Ilkeston, Derbyshire. Mus. Pract. Geol.
(Moysey Coll.), no. 30^2(i. 35.

16. Ditto; impression of dorsal surface of left wing showing interstitial

neuration. x '-'>\. 35.

2fl. Mecyiioptera tn/ifrnil<if<i, sp. nov. ; portions of two fore-wings and cubitb-
anal portion of hind-wing superposed, x Iv,. Middle Coal Measures
(shales above the Royley or Arley Mine) ; Sparth Bottoms, Rochdale,
Lancashire. British Museum, no. In. 1S57G. 37.

2/<. Ditto; impression of the two fore-wings showing interstitial neuration.

x li 37.

3«. Palxomantis nidcro^tcru, Bolton; fragment of left wing, dorsal aspect,
the underside of the right wing showing beneath. Natural si/e.
Middle Coal Measures; Ravenhead railway cutting near St. Helens,
Lancashire. Liverpool Museum. 40.

3/;. Ditto; impression of greater part of dorsal surface of left wing. Natural

size. 40.

4. Liihomantis carbonarius, Woodward ; portions of fore- and hind-wing,
the prothorax with domed lateral lobes, and the anterior styliform
process. (The latter ends in a sharp point not shown in the photo-
graph. A little of the mesothorax is seen between the liases of the
fore-wings.) Natural size. Coal Measures; Scotland. British
Museum, no. I. 81 IS. |:).

Bolton . Insects of Coal Measures.

PLATE n.

", *-' „' *"jj\ « :.'fS' '"VWlSi .J-2-*/

'

1 CRYPTOVENIA. 2. MECYNOPTERA. 3. PAL^EOMANTIS.

4. LITHOMANTIS

PLATE III.

Fio. PAOK.

la. TAth o*iu Us bromjiiiarti (Mantell) ; left fore-wing-. Natural size. Coal
Measures ; near Coalbrookdale, Shropshire. British Museum
(Mautell Coll. ul'iin Parkinson Coll.), uo. 11 HI',). 46.

\b. Ditto; impression of greater part of left fore-wing. Natural size. 46.

In. Frtirostia x/>/'rt<ibilin, gen. et sp. nov. ; left fore-wing, x 1^. Middle
Coal Measures (ironstone nodules in the binds between the
"Brooch" and "Thick" coals); Coseley, near Dudley, Stafford-
shire. British Museum (Johnson Coll.), no. T. 15R9I. 48.

1I>. Ditto; impression of same, x H. 48.

3. fipilii/iti-rii xitfclffi'i, Bolton ; Itasal part of left wing. X 1^. Middle

Coal Measures (shales above the Royley or Arley Mine); Sparth
Bottoms, Rochdale, Lancashire. Manchester Museum, no. L. 8197. 54.

4. lloltoiiii'llii tenuitegminata (Bolton) ; right hind-wing. x 2. Coal

Measures (No. '2 Rhondda Seam, base of Pennant Series) ; Ij miles
north-east of Resolven Station, Glamorganshire. Mus. 1'ract. Geok,
no. 24509. 56.

5. 1.1 rod in in'isi-otiiicfii, Scudder; almost complete left wing, showing the

attenuated base. X 1^. Middle Coal Measures (clay ironstone
nodule from binds between the "Brooch" and "Thick" Coals);
Dudley, Staffordshire. British Museum, no. I. 20(51. 59.

6. Ditto; portion of right hind-wing, under surface uppermost. (The

original furrows of the wing have been flattened out, the veins more
widely separated.) X If. Middle Coal Measures (clay ironstone
nodule from binds between " Brooch" and "Thick" Coals) ; Coseley,
Staffordshire. British Museum, no. In. ]<S4ol. 59

7. llrniliii j'in-1-iitii, Handlirsch ; wing lacking about one-sixth of the apical

end. X i2. Middle Coal Measures (above the "Brooch" Coal);
Dudley, Staffordshire. British Museum, no. 1.2962. (56.

s. Ditto; impression of same, showing the narrow base of wing. X ' i- 66.

PAL/CONTOGRAPH I CAL SOCIETY, 1919.

Bolton . Insects of Coal Measures.

PLATE 111

l.LITHOSIALIS. 2.PRUVOSTIA. 3. SPILAPTERA.
4.BOLTONIELLA. 5-8.BRODIA.

J W Tutcher, photo

London Stereoscopic Co imp.

PLATK IV.

FIG. PAGE.

1. Brodia priscotinda, Scndder (jin-i-nix} ; wing. X H. Middle Coal

Measures (binds between the " Brooch " and " Thick " Coals) ;
Coseley, Staffordshire. British Museum, no. I. 15(33. 63.

2. Ditto; slightly older wing with a broad base. X 3. Middle Coal

.Measures (binds between "Brooch " and " Thick" Coals); Coseley,
Staffordshire. British Museum, no. I. 1564. 63.

3. Jiroilio priscotincta, Scudder ; immature wing. X 1 i-. Middle Coal

Measures (binds between " Brooch " and " Thick " Coals) ; Coseley,
Staffordshire. British Museum (Madeley Coll.), no. I. 2<><;<;. 59.

4(i. "Pteronepionites" joiiitKuiti, sp. nov. ; very immature wing. X 12.
Middle Coal Measures (binds between "Brooch" and "Thick"
Coals) ; Coseley. Staffordshire. British Museum (Madeley Coll.), no.
I. 2'.H;7. 68.

4li. Ditto; impression of same showing undeveloped apex of wing. X 2. 08.

5. "Pteronepionites" ambigua, sp. nov.; crushed larval insect with partially
developed wings. X 4. Middle Coal Measures (binds between
" Brooch " and " Thick " Coals) ; Coseley, Staffordshire. British
Museum (Madeley Coll.), no. I. 29G8. 68.

G. " Pteronepionites " />y*//x, sp. nov. ; remains of larval insect with immature
wings and long segmented abdomen, the photograph accidentally
placed upside down. X 2. (Obscure traces of what may have been
pleiiral spines can be distinguished.) Middle Coal Measures (hinds
between "Brooch" and "Thick" Coals); Coseley, Staffordshire.
British Museum (Madeley Coll.), no. 1. 29G9. 6i».

7. jSHnigmatodes (.') >-<-</n1iiri*, sp. nov.; portion of inner half of wing, x 2.
Middle Coal Measures (over the Barnsley Thick Coal) ; Monckton
Main Colliery, Barnsley, Yorkshire. British Museum, no. In. 1860 4. 71.

tin. Pseudofouquea <VM/I/>/V//.V/'.S (Allen) ; greater part of left wing. X -.
Lower Coal Measures (top of the Four Knot Seam); Uaubradach
Colliery. Mus. I'ract. Geol., no. 7272. 72.

Xl>. Ditto; impression of missing apex of same wing. X 2. 72.

9a. Arcks&optilus ing ens, Scudder ; basal fifth of (? right) wing. Xatural six.e.
Middle Upper Coal Measures ; between Shelton and Clay Lane,near
Chesterfield, Derbyshire. Uritish .Museum, no. I. 3'.)(.>7. 74.

'.)//. Ditto; impression of same. X 1^. Uritish Museum, no. I. 39(.>7. 74-.

Bolton . Insects of Coal Measures.

PLATE IV.

1-3.BRODIA. 4-6 PTERONEPIONITES. 7. ^ENIGMATODES.
8.PSEUDOFOUQUEA. 9 ARCH^EOPTILUS.

.9088 QQ765 6960