THE EMBRYOLOGY OF A SCORPION.

105

The Embryology of a Scorpion (Euscorpius

italicus).
By

Malcolm Laurie, B.Sc,

Falconer Fellow of Edinburgh University.

With Plates XIII—XVIII.

SINCE 1870 there has been no detailed work on the de-

velopment of the Scorpion. As it seemed likely that with
modern methods of section-cutting and the great advance
which has been made of late years in the field of embryology,
a renewed examination might yield interesting results, I
have, at Professor Lankester's suggestion, examined and cut
sections of a large number of embryos of E u s c o r p i u s i t a l i c u s
preserved for him by the Zoological Station at Naples. I have
also examined- a number of embryos of Scorpio (Buthus)
fulvipes preserved and sent over from Madras by Professor
Bourne. These, however, chiefly owing to the small amount
of food-yolk, show such a great difference from E. i t a l i c u s
in their mode of development that it seems better to postpone
the description of them to a future paper.
The Scorpion is interesting not only as being the lowest, and,
as far as we know, the oldest type of air-breathing Arachnid,
but also as being exceptional among Arthropods in that the
whole development takes place within the body of the female—
in the ovarian tubes. The only other instances of this with
which I am acquainted are Phrynus, which is also viviparous,
VOL. XXXI,. PART I I . NEW SER. H
106 MALCOLM LAURIE.
and Sphoerogyna v e n t r i c o s a , one of the A.carina in which
the young are born sexually mature.
I may fitly here express my thanks to Professor Ray Lan-
kester not only for the suggestion that I should work at this
interesting subject, and for the generous way in which he has
provided me with material, but even more for his continual
and invaluable assistance and advice while the work has been
in progress.

HISTORICAL INTRODUCTION.

Johannes Miiller1 gave a short description, with five or six

figures, of the development of Buthus. Owing to its brevity
and the absence of any attempt to ascertain the internal
arrangement, his paper is of little value except from an histori-
cal point of view.
Duvernoy2 gives also only a few figures of Buthus and
of another form, probably Euscorpius. He describes at some
length a cord (baguette) which he says passes from the appendix
of the follicle in Buthus to the mouth of the embryo, and which
Miiller had compared to an umbilical cord. I hope to be able
in a future paper to give a detailed account of this and other
curious points in the development of Buthus. The chief value
of Duvernoy's paper was that he reconciled the contradictory
descriptions of the ovary which had been given by Miiller and
Rathke. 3 While doing this he makes a rather serious mistake
in describing the ovum of Buthus as occupying the whole of
the diverticulum of the ovarian tube, instead of only a small
space at the top.
The next writer on this subject is Leon Dufonr,* who gives
1
Joh. Muiler, " Beit. z. Anat. des Skorpions," ' Meckel's Arch. f. Anat.
u. Phys.,' Bd. xiii, 1828.
- Duvernoy, " Fragments sur les organes de la ge'ne'ration de divers Aui-
maux," 'Mem. de 1'Acad. des Soi. de l'lnstitut,' t. xxiii.
3
Rathke, "Zur Alorphologie," ' Reisbemerkungen aus Taurien,' 1837,
Riga, 4to.
* Dufour, " Hist. Anat. et Phys. de Scorpions," Mem. Presented a 1'Acad.
des Sciences,' t. xiv, 1856.
 THE EMBRYOLOGY OF A SCORPION. 107

an elaborate description with numerous figures of the anatomy

of the adult. His description of the embryo is, however, very
brief and his figures unsatisfactory.
Elias Metschnikoff1 is the only writer who has treated of the
development of the Scorpion with any degree of fulness. He
gives a detailed account of the whole development, and his
paper, which deals chiefly with the surface views and optical
sections, contains a large amount of accurate and laborious
observation. I t is the classic on this subject, and up to 1886
no attempt was made to add to it or supersede it.
In 1886 Kowalevsky and Schulgin3 published a short account
of the development of A n d r o c t o n u s o r n a t u s . Unfortu-
nately their paper has no figures, which detracts much from its
value. I find reason to differ from them on a few points, but
it is quite possible that this may be due to our having worked
on different genera.
The only other paper on this subject which I am acquainted
with is by G. H. Parker, 3 who treats at some length of the
development of the central and lateral eyes. I had worked at
this point before the appearance of his paper, and on the whole
agree with his conclusions. The.se are briefly that the lateral
eyes are monostichous, being formed from the hypodermis
without invagination. The median eyes, on the other hand,
are formed by invagination, and are therefore three-layered,
all the layers being derived from the hypodermis. The retina
is the second layer, the third being reduced to a post-retinal
membrane. The material at Mr. Parker's disposal did not
enable him to go back to the commencement of the formation
of the central eyes and their connection with the cerebral
invaginations.

1
Metsobnikoff, " Embryologie des Skorpions," ' Zeit. f. wiss. Zool.,' 1870.
a
Kowalevsky and Schulgin, " Entwickelungsgeschichte des Skorpions,"
' Biol. Centralblatt,' Bd. vi, 18S6.
3
Parker, "The Eyes in Scorpions," 'Bull. Mus. Comp. Zool., Harvard,'
vol. xiii.
108 MALCOLM LAUEIB.

T H E OVARY AND OVARIAN EGG.

The ovary consists, as is well known, of three longitudinal

tubes connected by transverse anastomoses, so as to form eight
quadrilateral meshes. The oviducts arise from the lateral
angles of the two foremost meshes and run forward to open
together on the genital operculum. The ovary appears to be
embedded in the liver, the chief mass of which lies dorsal to
i t ; this is not really the case, for, though lobes of the liver
pass through the meshes of the ovarian network they do not
unite on its ventral side. Both the longitudinal and transverse
tubes bear ova, which project from their outer surface as
oval bodies each attached by a short pedicle and measuring
when ripe about V% mm. in length and '83 mm. in breadth.
Ova in all stages of development are present on the ovarian
tubes at the same time, and there are in addition the corpora
lutea (v. p. 111).
The microscopic structure of the ovarian tubes i3 shown in
PI. X I I I , fig. 2. They are there seen to be formed of two
layers surrounding an irregular lumen. The outer layer, o. I.,
which is purely skeletal in function, consists of irregularly
polygonal cells, with circular nuclei and strongly marked cell
outlines. The contents of these cells are highly refractive.
Towards the inside of this layer the cells become flattened
so as to form a distinct, cellular, limiting1 layer. The inner
layer, which surrounds the lumen of the tube, is formed of
very long and thin columnar cells, with oval nuclei and clear,
faintly granular protoplasm. The nuclei are for the most part
confined to a central zone, leaving a large part of the outer
and a smaller part of the inner ends of the cells clear. It is
from this inner layer of cells that the ova and their follicles
are formed; it is, in fact, the germinal epithelium.
The first sign of the formation of an ovum is that one of
the cells of the inner layer of the ovarian tube begins to
increase in size (fig. 1, ov.). It contains finely granular
protoplasm, a large and distinct oval nucleus, and a darkly
 THE EMBRYOLOGY OP A SCORPION. 109

staining nucleolus. There is at first no sign of its presence

on the outside of the ovarian tube. As it increases in size,
however, it pushes its way, at the head of a column of cells,
towards the outside. The outer layer of the ovarian tube
becomes very thin, but remains as a membrane containing
few, if any, nuclei (fig. 2, fol'.). By the time the ovum is
about -04 mm. in length (fig. 2) it has passed completely
through the outer layer and is visible as a small protuberance
on the surface of the ovary. It remains connected to the inner
layer of the tube by a column of cells which is somewhat
expanded over the base of the ovum. The nucleus has not
increased in size in proportion to the growth of the cell.
The nuclei of the cells of the column which connects the
ovum to the inner layer of the ovarian tube next arrange
themselves so as to leave a clear space down the centre of the
column (PL XIII, fig. 3, mi.). They also grow round the ovum
so as to form a follicle (fig. 3, fol.) one cell thick. The
cells of this follicle rapidly become flattened and their nuclei
become smaller. The cells which remain clustered at the base
of the ovum (fig. 3, ffer'.) on the other hand increase iu size,
and shortly after the stage represented in fig. 3, which is a
drawing of an ovum of about *1 mm. in diameter, they begin
to secrete the yolk of which the greater part of the ripe egg is
composed. The outer layer of the ovarian tube can still
be traced as a thin and apparently structureless membrane
(fig. 3, fol'.) surrounding the egg outside the cellular follicle.
The nucleus has increased in size and is now a distinct oval
body with a round, granular nucleolus.
In fig. 4 is shown a longitudinal section of an egg of about
•4 mm. in length and -28 mm. in breadth. A considerable
quantity of yolk is now present in the form of spheres
ranging iu size from mere granules up to as much as "025 mm.
These spheres are clear, homogeneous, sharply defined bodies
showing no internal structure except that there is, in the
larger ones, a collection of granules at one point near
the outside. Round the nucleus the yolk-spheres are small,
and round the margin of the egg the protoplasm is coarsely
110 MALCOLM LAURIE.

granular, the rest of the space being filled up with the larger
spheres.
The nucleus (fig. 4, n.), which retains its central position,
is large ('05 mm.) but indistinct in outline and is probably
breaking down as I have been unable to find any trace of
it in eggs larger than that here figured. The nucleolus
(fig. 4, n'.)} which is situated towards one side of the nucleus, is
also large, staining darkly with carmine and showing a very
distinct circular outline. I t often contains one large, clear,
circular vesicle and a number of smaller ones.
The whole egg is surrounded by a distinct, rather thick
vitelline membrane (fig. 4, v. m.). No trace of pores or any
other structure was made out. Outside the vitelline membrane
the egg is surrounded, except at the base, by the follicle in
which the two layers (fol. and fol'.) of the ovarian tube can
still be traced. The cells of the inner layer of the follicle are
now flattened and small. The large yolk-forming cells at the
base of the egg {ger'.) have increased in size and arranged
themselves in a circle the centre of which is occupied by a
prolongation of the ovarian tube (mi.'). The egg is only
separated from this prolongation of the lumen by the vitelline
membrane. The spermatozoa are thus enabled to reach and
fertilize the egg while it is still in its follicle.
PI. XIII, fig. 5, shows the base of a ripe egg attached to the
ovarian tube. The pedicle has become shortened and its lumen
has increased very much in size. The yolk-forming cells have
degenerated, their flattened nuclei (ger'.) being, however, still
distinguishable, and the follicle has become much thinner owing
to the growth of the egg. The egg itself is a mass of tightly
compressed yolk-spheres, among which I have in vain sought
for the nucleus. It is probable, however, that the nucleus and
the greater part of the protoplasm migrate to the base of the
egg as segmentation commences there.
The yolk (PI. XIII, fig. 6) consists of spheres, ranging up to
•2 mm. in diameter. They are not homogeneous, but contain
spherical or prismatic bodies, which stain darkly with borax
carmine. These bodies are very large in the smaller yolk-
 THE EMBRYOLOGY OF A SCORPION. Ill

spheres, which contain one, two, or more of them. I n the

larger spheres they are much more numerous and much
smaller. Many of the spheres show round holes as if t h e
darkly staining bodies had dropped out. I t may be, however,
that these cavities contained a fatty or oily substance, which
has been dissolved out in the course of embedding and
mounting.
The only structures remaining to be described in connection
with the ovary are the corpora lutea mentioned above (fig. 7).
These are irregularly shaped bodies of about '12 mm. in
diameter, showing a slight tendency to radiate structure, and
containing a considerable number of nuclei, which are scattered
about without any definite arrangement. They project from
the surface of the ovarian tubes, and are evidently the collapsed
remains of the follicles after the egg has passed out. I was
confirmed in my idea that these were corpora lutea by their
resemblance to the structures described by v. Siebold 1 in the
ovary of Apus. They differ from these latter, however, in not
containing fluid.

FIRST PERIOD.—Formation of Blastoderm.

The egg is fertilized in the follicle, from which it does not
begin to pass out until the end of this period. I t then passes
into the ovarian tube in which it undergoes the rest of its
development, the young when born being exactly like the
parent in form. Kowalevsky and Schulgin 2 state that the egg
in Androctonus is not fertilized until it has entirely left the
follicle, and passed into the ovarian tube, or, as he calls it,
uterus. I can hardly believe this to be the case, but it is quite
possible that it leaves the follicle at an earlier stage in
Androctonus than in Euscorpius.
Stage A.—I have not, unfortunately, been able to observe
the processes of fertilization and the formation of the first
segmentation-spheres. I should think it probable that the
1
v. Siebold, 'Beitrage zur Parthenogenesis der Arthropoden,' Leipzig,
1871, p. 191.
2
Loc. cit., p. 526.
112 MALCOLM LAURIE.

greater part of the protoplasm with the nucleus collects at the

base of the egg. The youngest stage in my possession is shown
in surface view in PI. XIV, fig. 8, and insectiohinfig.9. The blas-
toderm forms a circular patch about "2 mm. in diameter, lying on
the surface of the yolk at the end of the egg nearest to the micro-
pyle, and consists of about twenty large cells, those in the centre
measuring about *03 mm. in diameter. In section (PI. XIV,
fig. 9) it is seen to be a single layer, the cells of which are
about -023 mm. thick in the centre. Round the margin the
cells are wedge-shaped so that the blastoderm lies flush with
the surface of the yolk. The cell-contents are coarsely granu-
lar, rather more so towards the lower side. The nuclei are
large, round and granular with distinct outlines.
The yolk-spheres under the blastoderm appear to be breaking
down. The blastoderm and yolk are closely surrounded by
the structureless vitelline membrane (v. m.). This stage seems
to be a little younger than that figured in Metschnikoff's paper
in PL XIV, fig. 6.
Stage B.—In the next stage (PI. XIV, fig. 10) the blastoderm
is somewhat larger, measuring '23 mm. in diameter. The
blastoderm is now almo.st twice as thick ("045 mm.). Some of
the cells are columnar, and occupy the whole depth of the
blastoderm, but the majority have divided in a plane parallel to
the surface, so that it is in places two or even three cells deep.
The nuclei vary in shape, those in the columnar cells being
oval.
Stage C.—In the next stage (PI. XIV, fig. 11) the blastoderm,
now - 3 mm. in diameter, is formed of an irregular mass of cells
showing as yet no trace of arrangement into layers. The cells
are comparatively small with well-marked outlines and large
nuclei. Round the margin of the blastoderm the cells form a
single layer on the surface of the yolk, but in the centre the
blastoderm is five or six cells thick, and the cells push their
way in between the yolk-spheres to which some of the cells
attach themselves. These cells, which attach themselves to
yolk-spheres, lose their definite outline and take, as far as I
have been able to ascertain, no part in the further growth of
 THE EMBRYOLOGY OF A SCORPION. 113

the embryo. There is no doubt that these yolk-cells are

derived from the blastoderm in this and the next stages, and
do not arise in the yolk by any process of free cell-formation.
Kowalevsky is also of this opinion. The yolk in the Scorpion's
egg shows no sign of segmenting as does that of the Spider. The
yolk of the Spider's egg seems1 to represent the hypoblast, and
takes an active part in the building up of the embryo; that of
the Scorpion, on the other hand, remains throughout develop-
ment an inert mass of food-material. This fundamental
difference in the segmentation makes any comparison of the
early stages of these two groups impossible, and would
seem to point to an independent origin for their abundance of
food-material. If the segmentation in Scorpions is a modifi-
cation of the centrolecithal type, as would seem probable from
the modes of segmentation in other groups of the Arachnida,
it is a very extreme one, and almost all trace of its origin has
been lost.

SECOND PERIOD.—Formation of the Three Layers and the Em-

bryonic Membranes.
Stage D.—It is difficult to get good sections at this stage as
the blastoderm is often humped up at the end of the egg and
compressed by the ovarian tube into which it is beginning to
pass. In one, and only one, series of sections I have seen what
appeared to be a longitudinal groove in the blastoderm. This
primitive groove is figured by Metschnikoff (PJ. XVII, figs. 2
and 3), but he may have been misled by the edges of the serous
membrane which is growing up and might easily give the
appearance of a groove in surface view. If the primitive groove
exists, which I am inclined to doubt, as the appearance in my
sections may have been due to shrinking, it is a very temporary
structure. Towards the posterior end of the blastoderm the
cells are proliferating and forming what I shall call the primi-
tive thickening1. From this primitive thickening is formed the
mass of hypoblast which is found later on in the tail-segment.
1
Locy, " Observations on the Development of Agelina nsevia," 'Bull.
Mus.j Harvard,' vol. xii.
114 MALCOLM LAURIE.

It would seem to represent a modified imagination, and is com-

parable to the primitive streak in the chick. I was at first in-
clined to call this the primitive cumulus, but considering the
fundamental differences between Scorpions and Spiders, and also
the fact that, while Balfour1 places what he calls the primitive
cumulus at the posterior end of the embyro, Locy 2 gives the
same name to a thickening at the anterior end, it seemed better
to avoid a term which might suggest erroneous homologies.
A layer of cells (fig. 12, pr. hy.) is seen to be forming under
the rest of the blastoderm, though not yet extending to its
edges. This is well marked in the next stage, and forms the
greater part of the primitive hypoblast or hypomesoblast. I t
would seem to be simply split off from the epiblast. I have
seen no appearance of a " down-sinking " of cells to form the
hypoblast, such as is described by Kowalevsky and Schulgin ; s
but, without the help of figures, it is not easy to be certain
of their exact meaning. Whether this " down-sinking" is
supposed to take place over the whole blastoderm or only
at the primitive thickening is not clear from their descrip-
tion.
Round the edges of the blastoderm a single layer of large
cells (fig. 12, s. m'.) is seen to be spreading a little way over the
surface of the yolk. These peripheral cells, which are at present
continuous with the epiblast, form later on the continuation of
the serous membrane. This serous membrane, or outer layer
of the amnion, is seen growing up as a single layer of cells
from the edges of the blastoderm (PI. XIV, fig. 12, s. m.).
I t spreads over the surface of the blastoderm from all sides,
and its edges ultimately meet and fuse in the middle line. At
this stage the edges have not yet come together, and the cells
of the layer are still small and similar in appearance to those
of the rest of the blastoderm.
The yolk is broken down to a considerable extent, and the
1
Balfour, " Notes on the Development of the Araneina," ' Quart. Journ.
Mior. Sci.,' vol. xx, 1880.
3
Locy, loc. cit.
3
Loc. cit., p. 526.
 THE EMBRYOLOGY OF A SOOEPION. 115
cells in it (fig. 12, y. c.) are numerous. Their nuclei are very
large and granular, and of irregular shapes. The cell-outlines
have entirely vanished, the cells being swollen up by an enor-
mous quantity of yolk-stuff. According to Kowalevsky and
Schulgin these cells are capable of amoeboid movements. Cells
continue to be added from the under surface of the blastoderm
to those already in the yolk up to the end of this stage. Their
function—of breaking down the yolk—is carried on at a later
period by the hypoblast.
Stage E.—In the next stage the blastoderm (PI. XIV,
fig. 13) has assumed an oval form, the thickened part or
ventral plate measuring "35 mm. in length and "25 mm. in
breadth, though the peripheral cells extend some way beyond
this. I have not been able, either in surface view or section, to
find any trace of the primitive groove, and imagine that, if ever
present, it has filled up. The primitive thickening (fig. 14, pr. t.)
is better developed than in the last stage, and the single layer
of primitive hypoblast (figs. 14 and 15, pr. hy.) is now quite
definite and extends a little way beyond the thick part of the
blastoderm, and forms a layer {hy'.) of cells under the peri-
pheral cells. These last (s. m'.) extend a good deal further than
in the last stage. The serous membrane (s. m.) is now com-
pleted over the surface of the ventral plate.
Stage F.—In the next stage the embryo, of which fig. 16
shows a longitudinal section, consists of two somites—those
which will afterwards bear the chelicerse and chelae—in addition
to the head- and tail-segments. The head- and tail-segments
are large, and a third somite is beginning to be formed from
the tail. The first somite is smaller than the second, and not
as yet very distinctly marked off from the head. It does not
become fully separated from the head until a much later stage
(eight somites). Except for this curious delay in the forma-
tion of the first, all the somites are formed and separated in
regular succession from the tail-segment.
The epiblast has undergone little change since the last stage,
except that it is somewhat thinner between the somites than in
them. It is beginning to grow up at the edges over the surface
116 MALCOLM LAURIE.

of the ventral plate as a single layer of flat cells to form the

inner embryonic membrane—the amnion proper (fig. 16, am.).
This amnion never loses its connection with the epiblast as-
the serous membrane has now done, but remains attached
to its edges and only extends round the egg as the epiblast
extends.
The most important change in this stage is the formation of
the mesoblast (mes.). This layer is formed under the whole
ventral plate by a multiplication of the cells of the primitive
hypoblast, from which it is in places not yet distinguishable.
The mesoblast extends across the whole ventral plate from
side to side, and is much thicker in the somites than between
them.
The serous membrane (s. m.) has, as mentioned above, now
lost all connection with the blastoderm, and is continued round
about two thirds of the egg by the " peripheral cells," which
are now beginning to separate from the egg and form a definite
membrane. The cells of the serous membrane are becoming
large and flat.
The hypoblast extends a little way beyond the ventral plate,
forming a single layer of cells (%.) in the periphery of the
yolk immediately under the serous membrane.
By the time the embryo has reached a stage with three somites
completely formed (PI. XIV, fig. 17) most of the changes which
were going on in the last stage are completed. The amnion has
entirely closed over the embryo (fig. 18, am.), though its cells
have not yet attained their characteristic form. The mesoblast
(mes.) is entirely separated from the hypoblast, and remains
henceforth a distinct and independent layer. The hypoblast
(hy.) is now a single layer, extending under the whole ventral
plate, except in the tail-segment, where it consists of a spheri-
cal mass. This hypoblastic mass in the tail-segment is the
direct product of the primitive thickening. The hypoblast
extends somewhat further round the egg than the other
layers, as is diagrammatically shown in fig. 19.
The description given above of the mode of formation of the
serous membrane and amnion differs very considerably from
 THE EMBRYOLOGY OF A SCORPION. 117
that of Kowalevsky and Schulgin. They describe it as a fold, the
outer layer of which forms the serous membrane while the
inner forms the amnion. This is probably the more primitive
mode of origin for these structures, and the mode described
above for E. i t a l i c u s is probably derived from it either by a
hastening of the formation of the serous membrane or a retarda-
tion of that of the amnion. I am unable to confirm their state-
ment that mesoderm cells are present between the two layers.

THIRD PERIOD.—Up to the Formation of Nine Somites.

This period covers the rest of the time before the appendages
begin to form. The egg has by this time entirely passed into
the ovarian tube. It has also increased considerably in size,
but I am unable to say whether this is due in any degree to
absorption of fluid or whether it is entirely due to internal
changes.
Stage G.—In the first stage belonging to this period which
I have examined (PI. XV, fig. 20) the embryo consists of
nine somites. The first of these—that which will bear the
chelicerse, is much smaller than the others, and is seen in section
to be not yet fully separated from the head. The second
somite, which will bear the chelae, is larger than those following
it. The next four are the ambulatory, and the seventh will
bear the genital operculum. A slight groove (n. g.) runs
down the middle line of the body; this is chiefly due to
the mesoblast having divided into two longitudinal bands
(figs. 21 and 22, me*.).
The epiblast is moderately thick in the somites, and is
beginning to grow as a single layer round the rest of the egg
(fig. 21, ep'.), carrying the amnion with it. By this stage it
has extended almost as far as has the hypoblast. The cells in the
middle line show a more definite arrangement than the rest of
the epiblast. This is preparatory to the formation of the
neural groove. The cells of the amnion (am.) have developed
their characteristic nuclei—spindle-shaped in sestien—'and
form a well-marked thin membrane lying close over the embryo.
The mesoblast (figs. 21 and 22, mes.) shows most iinpor-
118 MALCOLM LAURIE.

tant changes. As mentioned above, it has now separated into

two longitudinal bands. This separation does not extend into
the tail-segment (fig. 23, me*.), where the mesoblast remains
as a solid mass of cells somewhat thinner in the middle line.
The ccelomic spaces are now formed by a splitting of the meso-
blast in the somites. They are best seen in the posterior
somites (fig. 21, cm.), where the mesoblast is thin and forms
only a single layer on each side of the coelomic space. Further
forward (fig. 22) the mesoblast is thicker and the coelomic
space is not so well marked.
The hypoblast has undergone very little change. It is still
visible in the tail as a solid mass (fig. 23, hy. m.), and spreads
under the ventral plate and a little way beyond its margin as
a single layer (figs. 21—23, hy.). The cells of this single layer
have large oval nuclei which stain less darkly than those of
the epi- and meso-blast. These nuclei are somewhat widely
separated from each other, and the cells seem to contain a
considerable amount of food-stuff.
The serous membrane (figs. 21—23, s. m.) is by this time quite
separate from the egg all round. I t has attained its final
structure, the nuclei being enormously large ( - 05 mm.), flat,
and at a considerable distance from each other. As far as my
observations go I can confirm Blochmann's statement 1 that the
nuclei of the serous membrane divide directly without forming
any karyokinetic figures. As the serous membrane plays a
purely passive part in the future development it will not be
necessary to refer to it again.
Stage H.—In the next stage (PI. XV, fig. 24), which is
the last before the formation of the appendages, the embryo
consists of nine somites. The first is very much smaller than
the others, while on the second, which is the largest, a trace
of the appendages is just visible. The first six somites are
clearly distinguished from those further back, owing to their
sloping backwards and outwards, while the posterior ones are
at right angles to the axis of the embryo.
1
" Ueber direkte Kerntheilung in der Embryonalhiille der Skorpione,"
' Morph. Jahrb.,' vol. x.
 THE EMBRYOLOGY OP A SCOBPION. 119
A distinct groove, the neural groove (re. g.), runs down the
middle line and extends some distance into the head-segment.
It is due to a thinning of the epiblast in the middle line
(figs. 25 and 26, re. g.~). The ventral nervous system is
formed by a thickening of the epiblast along each side of
this groove.
The epiblast now spreads as a single layer beyond the
hypoblast {ep'.) and extends over nearly half the yolk, carrying
the amnion with it. This is diagrammatically shown in fig. 27.
In the head-segment (fig. 25) the epiblast is irregularly
grooved and thickened. This is the commencement of the
formation of the cerebral ganglion. In the thoracic somites
(fig. 26) the epiblast is very thick and solid at the corners (ap.)
where the appendages are about to appear. It is also some-
what solid just at each side of the neural groove (n. th.).
This is the commencement of the thickening which will form
the ventral nervous system.
The mesoblast is a thin layer in the head-segment (fig. 24,
mes.), but shows the ccelomic space (cce.) distinctly. This
development of a head coelom does not, of course, as Balfour
has pointed out, necessarily indicate that the head-segment is
equivalent to a body somite. In the body somites (fig.
26) the mesoblast is pretty thick and the coelomic space is
almost entirely closed up. The mesoblast does not extend
across the middle line or beyond the limits of the ventral
plate.
The hypoblast (figs. 25, 26, hy.) shows no change from the
last stage but remains as a single layer, except in the tail-
segment, where the hypoblastic mass is distinctly visible.
As the next stage shows the commencement of a large
number of new structures, the ventral nervous system, the
appendages, &c, it seems advisable to give a short summary
of what has taken place so far.
First Period.
(1) The blastoderm commences as a single saucer-shaped
layer of cells at one end of the egg (Stage A).
120 MALCOLM LAURIE.
(2) These multiply and form a thick mass (Stages B, C).

Second Period.
(3) The serous membrane grows up from the edges of the
blastoderm over its surface as a single layer of cells, and is
continued round the yolk by the peripheral cells (Stages D—F).
(4) The hypo-mesoblast is formed partly as a single layer of
cells split off from the under surface of the blastoderm and
partly, at the tail end, as-a thick mass, the primitive thickening,
which probably represents an invagination. Before and up
to this stage cells pass from the blastoderm into the yolk
(Stage D).
(5) The mesoblast is formed as a layer several cells thick,
extending right across the blastoderm. The hypoblast remains,
after the formation of the mesoblast, as a single layer, except
in the region of the primitive thickening, where it is a spherical
mass (Stage E).
(6) The amnion is formed as a single layer of cells growing
up from the edges of the epiblast, with which it retains its
connection. The serous membrane has by this time lost all
connection with the blastoderm, and spreads round the greater
part of the yolk (Stage P).
The embryo by this time consists of three somites and the
large head- and tail-segments. The somites are formed from
the tail in regular succession.
Third Period.
(7) The mesoblast divides into two longitudinal bands, and
ccelomic spaces are formed in the somites and in the head
(Stage G).
(8) The epiblast and amnion begin to spread round the egg
beyond the limits of the ventral plate (Stage G).
(9) The neural groove is formed by a thinning of the epiblast
in the middle line (Stage H).
(10) The epiblast in the head-segment begins to thicken to
form the cerebral nervous system (Stage H).
 THE EMBRYOLOGY OF A SCORPION. 121

FOURTH PERIOD.—From the Formation of the Appendages to the

Hatching of the Embryo.
Stage I.—The first stage of this third period shows—as
mentioned above—the commencement of some of the most im-
portant structures. The embryo, of which a surface view is
given in PI. XV, fig. 28, now consists of twelve somites in
addition to the head- and tail-segments. These somites are
no longer separate thickenings as. in the last stage, but have
grown close up to one another, and are marked off by narrow
grooves. The epiblast extends as a single layer all round the
egg. The longitudinal neural groove is well marked and
extends the whole length of the body with the exception of the
tail-segment.
The first six somites bear appendages, i . e . the chelicerse,
chelse, and four pairs of walking legs. These appendages are
simple outgrowths, and are, with the exception of the first two
pairs, of approximately equal size. The chelicerse are much
smaller, and the chelse somewhat larger than the other appen-
dages. The appendages are an outpushing of the epiblast and
the outer layer of mesoblast or somatopleure (PI. XVI, fig. 31).
They are hollow, the spaces being prolongations of the coelomic
pouches. There is at this stage no sign of appendages on the
somites behind those bearing the walking legs.
The embryo has a strong dorsal flexure so that the cephalic
segment curves round the end of the egg. This is best seen in
longitudinal section (PI. XVI, fig. 29). The anterior margin
of the cephalic segment is deeply cleft in the middle line, the
segment being thus divided into two lobes. The lobes are in
much the same state as in the last stage, and show no signs of
the cerebral invagination from which a greater part of the brain
is formed. In the middle line, and a very short way behind
the bottom of the cleft, is a circular raised area with a pit in
its centre (PI. XV, fig. 28, st.). This pit is the stomodaeum.
It is seen in section in PI. XVI, fig. 29, and is a simple inpush-
ing of the epiblast.
VOL. XXXI, PART II. NEW SER. 1
122 MALCOLM LAURIE.

The ventral nervous system consists of a pair of thickened

bands of epiblast running the whole length of the body on each
side of the neural groove (PI. XV, fig. 28). The bands are
cut up into blocks by the grooves which separate the somites.
The epiblast is not evenly thickened, but the nuclei are ar-
ranged so as to present a wavy outline. This is characteristic
of the formation of nerve-tissue in this animal, and was well
seen in the cerebral lobes in the last stage (PI. XV, fig. 25).
The small ganglia of the cheliceral somite are well seen at
this stage (fig. 28, g, I).
The tail-segment, from which the six caudal somites have yet
to be formed, has begun to be pushed out (PL XVI, fig. 29).
The epiblast in this region is very thick, and the cavity of the
outpushing is lined by a thick layer of hypoblast, which is the
" hypoblastic mass " of earlier stages (fig. 29, hy. m.).
Besides this mass in the tail-segment the hypoblast extends
as a single layer round the whole egg (PI. XVI, fig. 29, hy.).
Along the ventral side the cells of this layer are close together,
hut towards the sides and back they become more scattered,
and are to a great extent involved in the yolk. It is from the
mass in the tail-segment that the mesenteron is chiefly formed.
The hypoblast along the ventral surface also takes some part in
its formation, but that round the sides and back is not involved,
though it aids in the formation of the great digestive gland or
liver.
The mesoblastic bands (PI. XVI, fig. 31) are not yet united
across the middle line. The ccelomic spaces (PI. XVI, figs. 30
and 31) are well marked and quite separate for each segment.
Those in the first six somites are prolonged into the appendages.
The somatopleure is several cells thick ; the splanchnopleure, on
the contrary, consists of a single layer of cells. The mesoblast
in the cephalic segment is thinner than in the body somites,
and the coelomic space is narrower.
Stage K (PI. XVI, fig. 32).—The thoracic appendages have
increased very much in size, and the chelicera? and chelae are
both bifurcated at the extremity. A section through the base
of one of the ambulatory appendages (PI. XVI, fig. 33) shows
 THE EMBRYOLOGY OF A SCORPION. 123

a well-developed process extending inwards towards the middle

line. This is undoubtedly the sternocoxal process, which is
present on the second, third, and fourth appendages of the
adult. Lankester1 characterises the presence of this process
as a very important point of resemblance between the thoracic
appendages of Limulus and Scorpio. It is therefore interest-
ing to find it at this early stage present on all four pairs of
ambulatory appendages. A series of sections through the base
of the fifth appendage, i. e. third ambulatory (PI. XVI, fig. 34,
a—h), shows the first stage of another structure characteristic
of Limulus and the Arachnids—the coxal gland. This consists
of a simple tube opening to the exterior at the base of the fifth
appendage (fig. 34 a), and running forwards through the meso-
blast to open in fig. 34 h into the ccelomic space. There can be
no doubt that it is a nephridium. Grulland's researches3 on
the coxal gland in the young Limulus point to the same con-
clusion. I have been unable to find traces of nephridia in
any other somites, unless, indeed, the genital tubes are partly
nephridial. The six abdominal segments also bear appendages
(PI. XVI, figs. 32 and 35). These appear on surface view much
more prominent than they really are owing to their white
colour, which is due to the greater thickness of cells. In
section (PI. XVI, fig. 35) they are seen to project very slightly,
and to be formed by a thickening of the epiblast and somato-
pleure, but with no definite outpushing such as there is in the
thoracic appendages. The first pair of these appendages—the
genital opercula—is very small, and concealed by the last pair
of walking legs. The other five pairs—the pectines and four
pairs of lung-books—are all of approximately equal size and
structure. I have been unable to find the smallest trace of
appendages on the somites behind these, i . e . somites 13 —17,
and do not believe they exist.
The cephalic segment is not so deeply cleft as in the last
stage, and the mouth has shifted posteriorly so that now it lies
between the bases of the chelicerse. In the centre of each
1
' Limulus an Arachnid,' p. 20.
3
' Quart. Journ. Mier. Soi.,' vol. xxv.
124 MALCOLM LAURIE.

cephalic lobe is seen a dark spot (fig. 32, ce. in.). These spots
are the cerebral invaginations. They begin in a somewhat
earlier stage (PI. XVI, fig. 36) as a pair of small inpushings.
These extend rapidly backwards and meet in the middle line,
their two lumens becoming continuous. This is seen in PI.
XVI, fig. 37 A—D, in which four transverse sections through
this region are figured. Owing to the strong cephalic flexure in
this stage the stomodaeum (st.) is also shown in section. The
cells, both at the sides of the cephalic lobes and throughout the
greater part of the invaginations, are rapidly increasing in
number to form the cerebral ganglia. Those in the centre of
the cerebral lobes remain as a thin layer, and take no part in
the brain formation. The cells also on the dorsal side in the
middle, -where the two invaginations have united (PI. XVI, fig.
37 D, oc), are more closely packed than the others, and take no
part in the formation of the brain. They are the beginning of
the retinal layer of the central eyes.
The ventral nervous system is in much the same condition
histologically as it was in. the last stage. The commencement
of its separation from the hypodermis can, however, be seen
(PI. XVI, fig. 35) where the hypodermis is growing over it
from each side as a thin layer.
The tail segment is now divided into six somites, and extends
forward along the ventral surface of the body, reaching, at this
stage, to the third abdominal somite. The epiblast is thickened
on the ventral surface to form the nervous system. This is
not shown in fig. 35, as the section passes between two thicken-
ings. The cavity of the tail is occupied by a tubular extension
of the hypoblast (fig. 35, hy.) surrounded by mesoblast. There
is as yet no trace of the proctodseum.
The coelomic spaces in the thoracic somites have not
developed much. Those in the abdominal somites, however
(PI. XVI, fig. 35, coe.), have extended enormously, and now
reach round almost one third of the egg. The mesoblast, ex-
cept in abdominal appendages, consists of two single layers of
cells. In the tail the ccelomic spaces are not yet formed.
Stage L.—The embryo, of which fig. 38 (PI. XVII) shows a
 THE EMBRYOLOGY OF A SCORPION. 125

surface view, has by this time made considerable progress in

several important points. The thoracic appendages are slightly
segmented (PI. XVII, fig. 39, ap.), though this is not apparent
in a surface view. The chelicerse have moved in towards the
middle line, and the mouth is now concealed between their
bases. The chelae are very large, and have their pincers well deve-
loped. The coxal gland, which opens at the base of the fifth
pair of appendages, is no longer a straight tube, but has become
bent on itself, so that a section through it (PI. XVII, fig. 39,
cox.) shows the tube cut in three places. It can still, however,
be traced through a series of sections as a simple tube opening
into the ccelom. The abdominal appendages have undergone
great changes. The genital opercula are still simple thickenings
of the epi- and meso-blast, but the pectines (PI. XVII, fig.
40) have become folded in a direction parallel to the long axis
of the body, i . e . transverse to their own axis. The most im-
portant change is, however, that of the four following abdo-
minal appendages. These (PL XVII, fig. 41) are pushed in
so as to form shallow cup-shaped cavities. The inpushing is on
the posterior part of the appendage, and is directed slightly
forwards. This is the commencement of the formation of the
lung-book.
The cephalic segment, which is shown in PI. XVII, fig. 38,
extended in the same plane as the ventral surface of the
embryo, is no longer so distinctly bilobed as in the last stage.
The cerebral invaginations (PI. XVII, figs. 42, a and i, and 43)
are much shallower, and have entirely joined together, so that
there is now only a single inpushing. This lies just in front of
the chelicerse (PI. XVII, fig. 38). The brain is being formed
from the sides of the inpushing, and shows a very characteristic
structure'. The mass of cells is more or less grouped round
small circular clear spaces (fig. 43), which give to this part of
the brain the appearance of being composed of a number of
small vesicles. I have not succeeded in tracing the development
of the nerve-fibres, which occupy the centre of the cerebral
ganglion (fig. 43). This central portion appears at this stage
perfectly transparent and empty.
126 MALCOLM LAURIE.

The retina of the central eyes is still a thickening of the

dorsal layer of the cerebral invagination (PI. XVII, fig. 43,
rtn.). It is visible in surface view (fig. 38, oc.) as a white spot
on the margin of the invagination. The hypodermis imme-
diately outside it is somewhat thickened, and will in this
region form the vitreous layer (fig. 43, vit.).
The ventral nervous system is now completely separated from
the hypodermis (figs. 40 and 41, n. c ) . The cells are beginning
to congregate together to form the ganglia, though the nerve-
cord between the ganglia is still largely cellular. Nerves are
seen growing out from the ganglia as thick cords of cells
(fig. 40). The ganglia contain a clear space in their centre
which later is occupied by a mass of fibres.
The tail (PI. XVII, fig. 38) has now attained its full
number of segments but the sting is not yet formed. The
gut extends up almost the whole length of the tail. There is
no sign yet of the formation of the proctodseum. The hypo-
blast in the rest of the body remains as a scattered layer of
cells.
The mesoblast has now grown round the body as a double
layer, with the ccelomic space between. In the middle line of
the back, where the right and left folds of mesoblast meet,
there is a somewhat irregular thickening in which both soma-
topleure and splanchnopleure seem to be involved. From this
thickened band, which extends from close behind the brain to
the beginning of the tail, the heart is formed. On the ventral
side in the thoracic region the mesoblast of the outer layer is
broken up into long strings of cells—the muscles—so that the
coelomic space can no longer be very definitely made out.
The stomodseum reaches as far as the back of the cerebral
ganglion. This is the limit of its growth, and it remains a
closed tube until, at a much later stage, the gut has grown
forward and united with it.
Stage M.—The embryo (PI. XVII, figs. 44 and 45) does
not show very much change in surface view. The thoracic
appendages are longer and distinctly segmented. They overlap
across the middle line and conceal the pectines. The chelicerae
 THE EMBRYOLOGY OF A SCORPION. 127
are further forward in relation to the mouth, which can now
be seen lying between the bases of the chelse.
The genital opercula begin to grow out from the body wall
and the genital duct begins to be formed. This last (PI. XVII,
fig. 46) is developed in the mesoblast as a tubular portion of
the coelom, but does not open to the exterior up to the time of
hatching. It may be nephridial in its nature, but this very
late formation of the external aperture is not very favorable
to such an hypothesis. The pectines are separated at their
outer ends from the body wall. The inpushings for the lung-
books are much deeper, and the cavity, which extends forwards
from the opening, is divided up by lamellae which grow down
from its upper end (PL XVII, fig. 47). It is in close relation
to a space in the mesoblast which contains blood-corpuscles.
The cephalic segment (PI. XVII, fig. 45) is now rapidly
approximating to its final shape. The cerebral ganglion, which
is seen from the surface as a four-lobed white mass (fig. 45, ce.),
has now lost all connection with the epiblast. The invagina-
tion remains, but its sides no longer give rise to nerve-tissue
(PL XVII, figs. 48 and 49). The thickening for the central
eye (figs. 48 and 49, rtn.) is more largely developed, and
pigment is deposited in the ends of the cells furthest from
the invagination. The eye is plainly visible as a double
black spot on the surface. The upper edge of the invagina-
tion is growing down to close its orifice. The hypodermis
lying immediately above it is clearly marked off from the
rest as the vitreous layer (fig. 49, vit.). A considerable space
still separates the retina from the vitreous layer.
The lateral eyes now appear for the first time as black spots on
what Lankester terms the " optic area," i. e. the front margin
of the head (PL XVII, fig. 45, oc). Their development, as
Parker 1 has shown, is strikingly different from that of the
central eyes. Each eye, and in this species there are at first
three, is formed (fig. 50) by a slightly cup-shaped thickening
of the hypodermis. The nuclei of this thickened portion
become larger, and pigment soon begins to be deposited at the
1
Loo. cit.
128 MALCOLM LAURIE.

outer ends of the cells. Fig. 50 a shows a somewhat later

stage, in which the cupping of the hypodermis has become
flattened out. There is no invagination of any sort, and the
eyes are, as Lankester and Bourne1 described them, mono-
stichous. The ventral nervous system has not undergone much
development. It has sunk somewhat deeper and is separated
from the hypodermis by the mesoblast.
The tail has now developed its terminal segment—the sting.
The cavity of this last is partly occupied by the paired poison
gland, apparently formed by inpushing of the hypodermis (PI.
XVIII, fig. 51, p. gl.). Each mass is connected to the super-
ficial hypodermis by a short duct.
The gut extends down the whole length of the tail, and the
proctodseum is present in the form of a solid massof hypodermis
cells blocking up its end (PI. XVIII, fig. 51, prod.). The gut
has also begun to grow forward (PL XVIII, figs. 52 and 53). In
the last abdominal segment it is a complete tube surrounded
by a thin layer of mesoblast (fig. 52, int.). It gives rise to
two tubular outgrowths from its dorsal side, which are the
Malpighian tubes (fig. 52, mlph.). These run first towards the
back and then bend forward. There can be no doubt as to
their hypoblastic origin in this form, as the proctodseum is not
yet formed. They have been already shown to be outgrowths
of the mesenterou in some Spiders by Loman,2 and also in ter-
restrial Amphipoda by Spencer.3 Further forward (PI. XVIII,
fig. 53, int.) the gut is simply a semi-cylindrical layer of hypoblast
supported by a string of mesoblast and open to the yolk on its
dorsal side. In the thorax it has not yet begun to form.
The mesoblast is broken up into strings and bands. The
coelom is still pretty distinct in the abdominal region (PI. XVIII,
fig, 53, cm.), and the heart is a large thin-walled tube apparently
connected with both somatopleure and splanchnopleure. As
mentioned above, the genital tube is formed in the somato-
pleure in the seventh somite and is a portion of the ccelom.
1
' Quart. Journ. Micr. Sci.,' vol. xxiii.
3
' Tijdschrift der nederl. Dierk. Vereen,' i.
3
' Quart. Journ. Micr. Sci./ vol. xsv.
 THE EMBRYOLOGY OF A SCORPION. 129

Stage M,—The changes from the last stage up to the time

of hatching are not very numerous, though very important.
The body attains a structure almost exactly like that of the
adult, the appendages being segmented and the whole animal
covered by a thin, structureless, highly refracting cuticle. The
coxal gland still opens by a small aperture to the exterior at
the base of the fifth appendage (PI. XVIII, fig. 54). This
aperture, which is lined for a short distance by the cuticle,
leads to a straight duct (fig. 54) lined by cubical cells with
round nuclei, which closely resemble the cells of the gland.
The gland itself is distinguishable into medullary and cortical
portions as described by Professor Lankester 1 in. the adult. The
tubules have distinct lumens surrounded by a cubical epithelium.
The gland and its duct are surrounded by a thin capsule of flat
mesoblast cells.
The genital tubes have pushed their way some distance
between the lobes of the liver, but they are not yet connected
by transverse tubes nor do they open to the exterior. The two
layers of which the tube is composed in the adult (v. supra,
p. 108) are not yet distinguishable. The pectines approximate
very closely to their adult structure.
The ninth, tenth, eleventh, and twelfth appendages are also
very similar to those in the adult (PI. XVIII, fig. 55). The
number of lamellae is not so great, but their structure is very
well Bhown. Each lamella is covered by a thin cuticle, and its
cavity is in direct communication with a blood-sinus (fig. 55,
bl. s.). The cells which form the lamellae are very large, espe-
cially towards the base of the appendage. Towards the apex
they become smaller, and finally pass into a mass of different
cells from which more lamellae are formed as the animal grows.
The spaces between the lamellse (fig. 55, a. c'.) ai'e narrower and
in communication with the exterior through the stigma.
The head is now completely formed, the mouth having
shifted so as to lie behind the chelicerae. The invagination
which forms the central eyes has closed up. A stage immedi-
ately after its closure is shown in PI. XVIII, fig. 56. Here the
1
' Proc. Roy. Soc.,' vol. xxxiv, 1882-83,
130 MALCOLM LAURIE.

lips of the invaginations have come together, but not fused.

The posterior layer of the invagination is visible as a thin layer
of cells (fig. 56, rtn'.), separated from the retina by a narrow
space. The vitreous layer (vit.) is distinctly marked as a
thickening of the hypodermis on the top of the head, the nuclei
in that region being elongated, but there is still a small space
separating it from the retina. By the time the embryo is
hatched the eye (PL XVIII, fig. 57) has lost all connection
with the hypodermis at the point where it was invaginated.
The cells are long and deeply pigmented round their margins.
The pigment is not equally abundant throughout the whole
length of the cell, but five alternately more and less deeply
pigmented zones can be distinguished (fig. 58). The base of
the cells is most deeply pigmented, and their superficial ends
come next. The nuclei of the retinal cells lie in zone 4,
but I have been unable to find any trace of either rhabdomes
or phaospheres. I have also not been able to trace any migra-
tion of mesoderm cells among the retinal cells. The posterior
layer of the invagination can with difficulty be made out in
depigmented sections owing to the flatness of its nuclei, and it
is absolutely undistinguishable in sections from which the pig-
ment has not been removed. This posterior layer forms the
post-retinal membrane of the adult eye. The optic nerve is
beginning to grow out from the cerebral ganglion, but has not
yet come into connection with the eye. The hypodermis, im-
mediately in front of the eye, is formed of a single layer of
large transparent cells with faintly staining oval nuclei
(PI. XVIII, figs. 57 and 58, vit.). This vitreous layer is
covered by a thin cuticle exactly like that which covers the
rest of the surface of the body. The only sign of the formation
of the lens is a slight cupping of the vitreous layer at one point
(fig. 58). The hollow formed here is, however, not as yet filled
up by any cuticular substance, but the cuticle passes straight
over it. Round the area, where the lens will form the hypodermis
is deeply pigmented. The cells are much smaller than those
of the vitreous layer, and their nuclei are irregular in shape.
The cells of the lateral eyes (PI. XVIII, fig. 59) are about
 THE EMBRYOLOGY OP A SOOEPION. 131
the same size as those of the median ones. The pigment is not,
however, arranged in definite zones, though it is more abundant
at the base and at the outer ends of the cells than in the middle.
There is a small third lateral eye present, and the hypodermis
around and—the lateral eyes being so to to say on the edge of
the head—below the eyes is all pigmented. I have been
unable to find the nerve to these eyes and think it is probably
not yet formed. The cuticle over the lateral eyes is not
thickened to form the lens, and I have seen no sign of the
peculiar mode of lens-formation described by Parker, 1 i. e. the
ends of the perineural cells pushing in front of the retina. I t
is, of course, possible that this does not take place till later.
The brain and ventral cord have almost attained their
adult structure. In the nerve-cord there is a string of
cells in the middle line (PI. X V I I I , fig. 60) dorsal to the
cords proper, which seems to represent the centre of the neural
groove.
The tail is exactly similar to that of the adult, and is carried
in the same way curved over the back. The poison-glands are
fully formed and surrounded by muscles, but do not occupy so
much of the terminal segment as in the adult. The proctodseum
is lined by flat cells and has pushed its way almost to the
base of the tail. The mesenteron is fully formed only in the
hind segments of the body (PI. XVIII, fig. 60). From the
end of the stomodaeum to where the last hepatic caeca
join it the intestine (fig. 61) is surrounded by a definite
cylindrical layer of mesoblast which is continuous with that
surrounding the lobes of the liver, but the hypoblast cells
lining this cylinder (fig. 61, hy.) are not yet definitely arranged.
The nuclei are scattered about in groups for the most part
near the outside, and the cells are drawn out into irregular
more or less pyramidal form, the apex of the pyramid pointing
towards the centre of the tube. There is no definite lumen,
the space between the cells being filled up by small yolk-spheres
(fig. 61,2,*.).
The liver-follicles are much the same in structure as the
1
Loo. oit., p. 199.
 132 MALCOLM LAURTE.

intestine. They contain, however, a rather larger proportion of

yolk. The scattered layer of hypoblast cells, which in the
preceding stages surrounded the yolk, takes a large part in their
formation. They open into the intestine in pairs by wide
ducts.
The Malpighian tubes (PI. XVIII, fig. 60, mlph.) have not
undergone much development. They reach well forward in the
body, and open into the intestine in the first caudal segment.
It is evident from the structure of the intestine that the
young scorpion does not need food for some time after hatch-
ing. The large amount of yolk which still exists must last it
for some weeks, or most probably till the next spring. If this
is the case embryonic life practically lasts twelve months as the
eggs are fertilized in May.
The outer layer of the mesoblast has now for the most part
formed itself into muscles. The inner layer is very much com-
plicated, being folded in so as to surround the gut and the
lobes of the liver. The spaces between the lobes of the liver,
which are undoubtedly the true coelom, are filled up by a net-
work of trabecular tissue (PI. XVIII, fig. 60). The heart,
pericardium, and blood-vessels are fully formed and contain a
considerable number of large nucleated corpuscles.

Summary of the Changes during the Fourth Period.

(1) The thoracic appendages begin as simple outpushings of
the body wall containing a portion of the coelom (Stage I).
They rapidly increase in length and the chelicerse and chelae
become bifid at their extremities. Sternocoxal processes are
present on the third to sixth appendages (Stage K).
The chelicerse, which were at first behind the mouth, gradu-
ally move forward relatively to it till they come to lie in front
of it (Stage L).
(2) The coxal gland begins as a simple tube opening to the
outside at the base of the fifth pair of appendages, and opening
at the other end into the ccelom (Stage K). The tube soon
becomes coiled, but the external opening persists until after
hatching. It is undoubtely a nephridium,
 THE EMBRYOLOGY OF A SCORPION. 133
(3) The abdominal appendages appear as thickenings of the
epi- and meso-blast on the seventh to twelfth somites (Stage K).
The first pair (genital opercula) does not develop further till
a late stage (L).
The second pair (pectines) form a number of short longi-
tudinal ridges on the surface of the abdomen (Stage K). They
then separate from the body, the separation beginning at their
outer ends (Stage M).
The third to sixth pairs (gill-books) begin to be pushed in
(Stage L). The inpushing becomes deeper, and begins to be
divided up (Stage M), and by the time the embryo is hatched
they have attained their adult condition in every respect except
size and number of lamellae.
(4) The cerebral ganglion and central eyes begin as a pair of
invaginations on the cephalic lobes. These invaginations
meet in the middle (Stage K). The cerebral ganglion is formed
from the sides of the invaginations, which rapidly become
shallower and unite so as to open in the middle line. The
dorsal surface of the invagination becomes thickened to form
the retina of the central eyes (Stage L).
The brain becomes entirely separate from the hypodermis,
the invagination remaining to form the eyes (Stage M). The
invagination closes up and its lumen disappears. The cells of
its lower layer form the post-retinal membrane. Those of the
upper layer form the retina, and come in contact with the hy-
podermis on the top of the head, which is thickened in this
region to form the vitreous layer. The retinal cells become
deeply pigmented (Stage N).
(5) The lateral eyes form as cup-shaped thickenings of the
hypodermis in the " optic area," the cells of which become
pigmented. There is no invagination, and they consist of a
single layer (Stage M).
(6) The ventral nervous system forms as a pair of thickened
segmented bands, one on each side of the neural groove
(Stage I). The nerve-cords sink down, a thin layer of hypo-
dermis growing over them. There is at this time (Stage K)
a distinct postoral pair of ganglia for the chelicerse. The cells
134 MALCOLM LAURIE.
become aggregated to form ganglia, and the cheliceral ganglia
become fused with the cerebrum.
(7) The tail grows out, lying along the ventral surface
of the abdomen. The poison-gland in its terminal segment is
formed by a pair of invaginations of the epiblast.
(8) The hypoblast consists of an irregular layer under the
whole embryo and a solid mass at the tail end (Stage I). As
the tail grows the hypoblast grows into it as a tube reaching
down to the last somite (Stage K).
The hypoblast forms the gut in the abdominal portion of the
body, growing forward in a sling of mesoblast at first as a flat
layer, which soon becomes bent round into a cylinder. The
Malpighian tubes are formed as outgrowths from the mesenteron
in the first post-abdominal somite (Stage M).
The gut does not reach forward to the stomodaeum till
shortly before hatching, and at this period the portion of
it into which the liver-follicles open is not fully formed
(Stage N).
(9) The stomodseum is formed early. It lies at first in front
of the cbelicerse (Stage I), but soon shifts its position and
comes to lie behind them. It extends inwards as far as the
back of the brain.
(10) The proctodaeum is formed much later than the stomo-
dEBum. It is at first a solid plug of cells (Stage M). As it
increases in size it appears to replace the hypoblast in the
last four somites.
(11) The mesoblast consists at first of a pair of segmented
bands with a separate ccelomic space in each somite, and also
one in the cephalic segment (Stage I). The ccelomic spaces
soon unite, and the mesoblast bands join across the ventral
surface. Somewhat later they extend round—the ccelomic
space extending with them—and unite in the middle line on
the dorsal surface (Stage L). From the thickened band where
they have united on the dorsal surface the heart is formed. A
portion of the ccelom in the seventh segment becomes separated
off to form the genital tubes (Stage M). These do not open to
the exterior. The outer layer of the mesoblast forms chiefly the
 THE EMBRYOLOGY OF A SCORPION. 135

muscles of the body. The inner layer becomes folded so as to

surround the liver and intestine, and the ccelomic space becomes
partly filled up by trabecular tissue.

Conclusion.
The development cf this Scorpion, of which I have tried to
give an outline above, is interesting in many points. It does
not agree closely with any other Arachnid type as yet described,
and I have for the present given up all attempts at comparison.
The development of the central and lateral eyes entirely bears
out Lankester and Bourne's description of their structure. It
is true that the central eyes are three-layered, but as the retina
is the second layer from the surface—the third layer forming
only a post-retinal membrane—they may be called diplostichous.
The account given above of their development agrees in all
essential respects with that of Parker, but, having a larger
supply of embryos, I have been able to trace the earlier stages
and the connection of the eyes with the cerebral invagination.
Their mode of origin resembles very closely Locy's1 description
of the development of the eyes in Agelena nsevia, the chief
difference being that in Agelena the optic invaginations appear
to have no connection with the formation of the brain. Locy
does not, however, give a detailed description of the formation
of the latter.
The description given above of the development of the lateral
eyes also agrees pretty closely with that of Parker. In these,
as in the central eyes, Lankester and Bourne's conclusions are
confirmed, and Patten's 3 conclusions as to what the structure
of the eyes must be in order to fit in with his theories are
shown to be without foundation. The lateral eyes are mono-
stichous, being simply somewhat specialised hypodermis cells.
The mode of formation of the ventral nervous system is
exceptional among Invertebrates, resembling rather that of
1
'Bull. Mus. Comp. Zool., Harvard,' vol. xii, p. 85.
3
"Eyes of Molluscs and Arthropods," 'Mitth. Zool. Stat. Naples,'
Bd. vi.
136 MALCOLM LAUEIB.
Chordata. The nerve-coi'd instead of peeling off from the
superficial layer of epiblast sinks down bodily, and is covered
by a layer of epiblast which grows over it from each side.
The development of the coxal gland leaves, I think, no room
to doubt that it is a nephridium. That of the genital tubes is
less conclusive, but I should think it probable that they are
also, at least in part, nephridial.
The gill-books are undoubtedly appendages comparable to
the abdominal appendages of Limulus. Whether they are
really invaginated, i . e . whether the edge of each lamella in the
Limulus appendage corresponds to the bottom of the fold
between the lamellae in the Scorpion's gill-book, or whether the
whole appendage has become sunk in a hollow in the abdominal
surface without being invaginated, it is difficult to say. Un-
doubtedly, the surface now exposed to the air has always been
the external surface, but that would be the case with either of
the above modes of derivation. Although the second alterna-
tive has the advantage that it is easy to see how the change
could take place gradually, I am inclined to think the first is
probably the true way in which they have arisen. One argu-
ment in its favour is that if the second alternative were correct
one would expect the gill-book to commence as a distinct out-
growth, which would become sunk in a pit. Now, there is no
such outgrowth in the formation of the gill-book. The first
thing to appear on the thickened portion of the epiblast, from
which the gill-book is formed, is a pit (PL XVII, fig. 41). The
lamellae do not begin to form till a later stage. Again, the
abdominal appe adages of Limulus are directed towards the tail
as one would expect abdominal appendages to be. Now, if the
appendage had sunk in without invagination, one would expect
it to be still directed towards the tail unless there were some
very good reason for its having changed its direction. If, on
the contrary, it had become invaginated it would naturally be
directed in the opposite direction towards the head, and this is
what we find in the Scorpion. The inpushing is from the
beginning towards the head, and the aperture opens towards
the tail (PL XVII, fig. 47). I think it is quite conceivable
 THE EMBRYOLOGY OP A SOOEPION. 137

that the changed conditions of development, due to terrestrial

life, and the consequent pressure on the embryo, may have pro-
duced this change. A detailed account of the development of
these appendages in Limulus may throw more light on the
matter, but, unfortunately, though many authors have attacked
the problem, a complete and satisfactory account of the develop-
ment of Limulus is not yet in existence.

EXPLANATION OF PLATES XIII, XIV, XV, XVI,

XVII, & XVIII,

Illustrating Mr. Malcolm Laurie's paper on "The Em-

bryology of a Scorpion (Euscorpius italicus)."

Abbreviations.
a. c. Air-cavity in gill-book, ac . Air-spaces between the lamellse of gill-
book, ah. ap. Abdominal appendage, am. Aranion. am. c. Amniotic cavity.
ap. Appendage, bl. Blastoderm, bl. s. Blood-space, bl. c. Blood-corpuscle.
can. Caudal segment, ce. Cerebral ganglion, ce. in. Cerebral invagination.
ceph. Cephalic segment, ex. Ccelom. cox. Coxal gland, cox. d. Duct of
coxal gland, ep. Epiblast. ep'. Extension of epiblast beyond ventral plate.
fol. Eollicle. foil. Outer non-cellular layer of follicle, g. 1. Ganglion of
cheliceral somite, ger. Germinal epithelium or inner layer of ovarian tube.
ger'. Yolk-forming cells derived from germinal epithelium, ge. t. Genital
tube. hi. Heart. hy. Hypoblast. hy'. Extension of liypoblast beyond
ventral plate, hy. m. Mass of hypoblast in caudal segment, int. Intestine.
I. Gastric gland, mes. Mesoblast. mi. Prolongation of ovarian tube to egg.
mlph. Malpliigian tubes, n. g. Neural groove, n. n'. Nucleus, nucleolus.
n. c. Nerve.cord. n. gl. Nerve-ganglion. n. th. Neural thickening, oc.
Central eye. oc'. Lateral eye. o. I. Outer layer of ovarian tube. ov. Ovum.
p. gl. Poison-gland, pr. hy. Primitive hypoblast (hypomesoblast). proct.
Proctodseum. pr. t. Primitive thickening, rtn. Ketina of central eye.
rtn'. Third layer of central eye, post-retinal membrane, s. m. Serous mem-
brane, s. m'. " Peripheral cells." torn. mes. Somatic mesoblast. spl. mes.
Splanchnic mesoblast. si. Stomodseum. ste. p. Sternocoxal process, stg.
VOL. XXXI, PART II. NEW SEB. K.
138 MALCOLM

Stigmata, ie. Telson. tr. mes. Trabecular mesoblast occupying ccelom. vil.
Vitreous layer of central eye. y. c. Cells in yolk. yk. Yolk. The somites
are numbered i, n, m, &c.

PLATE XIII.
FIG. 1.—Transverse section of ovarian tube, showing the two layers; one
cell of the inner layer enlarging to form an ovum. X *-^.
FIG. 2.—Transverse section of ovum and ovarian tube. The egg has now
pushed its way through the outer layer, and appears as a small protuberance
on the ovarian tube. The follicle is beginning to form from the cells of the
inner layer, which have accompanied the ovum, x &^.
FIG. 3.—Longitudinal section of ovum of - 1 mm. diameter, showing the
two-layered follicle and the yolk-forming cells (ger',). x ^s•.
FIG. 4.—Longitudinal section of egg of '4 mm. in length, showing yolk-
spheres, indefinite nucleus, and strongly marked nucleolus. The egg is sur-
rounded by a vitelline membrane. The rest as in Fig. 3. X S-|3.
FIG. 5.—Section through the base of a ripe egg. x 1^s-.
FIG. 6.—Yolk-spheres from ripe egg, showing the darkly stained spherical
and prismatic bodies and the clear spaces. X i p .
FIG. 7.—Section through a corpus luteum and part of ovarian tube.

PLATE XIV.
FIG. 8.—Surface view of one-layered blastoderm, x -^-.
FIG. 9.—Section through one-layered blastoderm, same stage as Fig. 8.
x ^.
Fi&. 10.—Section .through blastoderm later than Pig. 9, showing the cells
multiplying to form a mass at one pole of the egg. x ?%&'
FIG. 11.—Section through more advanced blastoderm. X £4p.
FIG. 12.—Transverse section through blastoderm at time of formation of
primitive hypoblast and serous membrane. The yolk and yolk-cells are drawn
in detail in this figure to show the breaking down of the former, x • s -p.
FIG. 13.—Surface view of blastoderm now becoming oval, x -&p-.
FIG. 14.—Transverse section through posterior end of embryo figured in
Fig. 13, showing serous membrane, primitive thickening, primitive bypoblast,
and " peripheral cells." x &p.
FIG. 15.—Transverse section through anterior part of embryo about the
same stage, x l^-.
FIG. 16.—Longitudinal section through an embryo a little younger than
Fig. 17, showing two somites with a third forming. The mesoblast is
forming from the primitive hypoblast, the amnion is growing up from the
 THE EMBRYOLOGY OF A SCORPION. 139

edges of the hypoblast, and the primitive thickening is well seen in the caudal
segment. X -^p.
FIG. 17.—Surface view of embryo, with three somites fully formed.
x J£-.
PIG. 18.—Transverse section through posterior end of Fig. 17, showing
hypoblastic mass, mesoblast, &c. X i-^.
TIG. 19.—Diagrammatic representation of the relative extension of the
various layers in an embryo of the stage of Fig. 17.

PLATE XV.
FIG. 20.—Surface view of an embryo with seven somites, drawn as if
flattened out. a—6 and c—d are the planes of the sections figured in Figs.
21 and 23. X *£-.
FIG. 21.—Transverse section through one of the posterior somites of an
embryo with seven somites (a—b in Fig. 20), showing the three layers,
epiblast thinning in centre, and mesoblast thin; amnion, serous membrane,
and ccelomic spaces. X *\9-.
FIG. 22.—Transverse section through one of the anterior somites of Fig.
20. x -4*.
FIG. 23.—Transverse section through tail-segment (c—d) of Fig. 20,
showing the undivided mesoblast and the hypoblastic mass, x 4 ^ .
FIG. 24.—Surface view of embryo of nine somites, drawn as if extended.
X 3SL.
FIG. 25.—Transverse section through head-segment of Fig. 24, showing
epiblast thickening to form cerebral nervous system and spreading (ep'.), with
the amnion beyond the ventral plate, neural groove, thin mesoblast, with small
ccelomic space and hypoblast. X i-\Q-.
FIG. 26.—Transverse section through one of the anterior somites of Fig.
24, showing the epiblast very solid where the appendage will develop (ap.)
aud form the neural thickening {n. th.) at each side of the neural groove.
Mesoblast thick, and ccelom not very evident, x x\s-.
FIG. 27.—Diagrammatic representation of the relative extension of the
various layers in an embryo of the stage of Fig. 24.
FIG. 28.—Surface view of embryo at Stage I (ten somites) extended in a
plane, showing appendages, cheliceral ganglion, stomodseum, &c. X A&..

PLATE XVI.
FIG. 29.—Longitudinal section of Stage 1 in the middle line, showing
dorsal flexure of the embryo, commencement of tail outgrowth, stomodseum,
&c. x -^..
140 MALCOLM LAURIE.

PIG. 30.—Longitudinal section to one side of the middle line, showing the
appendages and coelomic spaces. X ££-.
FIG. 31.—Transverse section through the third somite of Stage I, showing
the formation of the appendage, the neural thickening, &c. X -e-£-.
FIG. 32.—Surface view of embryo at Stage K, showing the cerebral imagi-
nations, abdominal appendages, tail, &c. X -^-.
PIG. 33.—Transverse section through the base of a thoracic appendage,
showing the sternocoxal process.
PIG. 34, a—A.—Series of sections thiough base of fifth appendage, showing
the coxal gland.
PIG. 35.—Transverse section through one of the abdominal appendages and
the tail, showing the appendage, the neural thickening beginning to separate
from the epiblast, the gut forming in the tail, &c. X -715-
FIG. 36.—Transverse section through the cephalic segment of a somewhat
earlier embryo, showing the beginning of the cerebral invagination. x - ^ .
PIG. 37, A—D.—Sections through the head of an embryo of Stage K, show-
ing the cerebral-optic invaginations. X -^s..

PLATE XVII.
PIG 38.—Surface view of an embryo of Stage L extended in a plane,
showing the cerebral invagination, the central eyes, &c. X ~°-.
PIG. 39.—Transverse section through the base of the fifth appendage,
showing the coxal gland, x -2^-
PIG. 40.—Section through the pectines. X ^ .
PIG. 41.—Section through an abdominal appendage, showing the inpushing
to form the gill-book. X -',5-.
PIG. 42, a, b.—Transverse sections through the cerebro-optic invaginations.
x -^a-.
FIG. 43.—Longitudinal section through the cerebro-optic invaginations,
showing the formation of the brain and the central eye. X A-j-^.
PIG. 44.—Surface view of an embryo of Stage M, from the ventral surface.
X 4j2..

PIG. 45.—Surface view of the dorsal side of the head of the same embryo,
showing i he central and lateral eyes, x \ 0 -.
PIG. 46.—Section through the seventh somite, showing the formation of
the genital tube from part of the ccelom. X •sf-.
PIG. 47.—Longitudinal secion through a gill-book, showing the commence-
ment of the formation of the lamellee. x :L\^.
FIG. 48.—Longitudinal section through the head end, showing the stomo-
deeum and the cerebro-optic invagination from which the brain is now entirely
separate. X -*T*-.
PIG. 49.—Transverse section through the same region, x ^-.
 THE EMBRYOLOGY OF A SCORPION. 141

FIG. 50.—Longitudinal section through the lateral eye, showing its forma-
tion by a thickening of the hypodermis. X i-p.
PIG. 5O«.—Section through a somewhat older lateral eye in which the
inpushing of the hypodermis has disappeared, x ^s•.

PLATE XVIII.
FIG. 51.—Longitudinal section through the tail end of Stage M, showing
the poison-gland, proctodseum, intestine, &c. X 4A.
FIG. 52.—Transverse section through the posterior end of the body, showing
the intestine, with the Malpighian tubes, the heart, &c. X -^-.
FIG. 53.—Transverse section a little further foward than Fig. 52, showing
the intestine, which has not yet closed into a tube. X - ^ .
FIG. 54.—Section through the coxal gland of a newly-batched scorpion,
showing the opening to the exterior, &c. X ££•.
FIG. 55.—Longitudinal section through a gill-book of a newly-hatched
scorpion, x ^-jp.
FIG. 56.—Longitudinal section through the central eye of an embryo a short
time before hatching, showing the closure of the cerebro-optic invagination
and the three layers of the eye. X - ^ .
FIG. 57.—Longitudinal somewhat oblique section through the central eyes
of a newly-hatched scorpion. X *$&•.
FIG. 58.—A few cells of the same eye more highly magnified, and showing
the inpushing in the vitreous layer.
FIGS. 58, a, b, c.—Transverse sections through the same eye at different
levels.
FIG. 59.—Section through the lateral eyes of a newly-hatched scorpion.
X if*.
FIG. 60.—Transverse section through the posterior part of the body of a
newly-hatched scorpion, showing the fully-formed intestine, the Malpighian
tubes, the nerve-cord, and the trabecular tissue filling up the ccelomic space.
X ***.
FIG. 61.—Transverse section through the intestine further forward, where
it is not yet properly formed, showing the irregular hypoblast cells and yolk-
spheres. X if*.
Fccf.Z.
m

i • *

\ * *
 Fig.

\ ..-n

A^Sl-SV^-..

&O y-

/ • c
•^HtH.
y.k.

s.m.
 U

;
l
• •" i

.'
St^
.V.V.Y.V.W -
. ••>• nit
V, '.

:
m-.:••••
Ni-. ;
k-'.'r
- * . • :
/\ ^^ v
• • • • • ' t
: a | . - ••-

VIA.

•••«<&°*»i'ift,. ^
.Vlt.

»-i/r,'

Fiq.58c

vit.