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COXAL GLANDS OF THE ARACHNIDS

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Coxal glands of the Arachnids.

B. H. Buxton. q^ '

(Cornell medical college, New York.)

With Plates 1—43 and 7 Figures in the text.

Part I, Pulmonate Arachnids.

Introduction.

Limtihis has probably in many respects the most primitive
arrangement of the coxal glands among the arachnids. According
to Patten each gland consists of four lobes (saccules), one at the
base of each appendage two to five. A short duct connects each
lobe with the common duct (labyrinth;, which after a series of com-
plicated twists and turns, finally ends in a straight dilated exit tubule
or end sac with its outlet just posterior to the base of the coxal
joint of the fifth appendage. In the embryo, lobes are also formed
on the first and sixth appendages, but degenerate and disappear in
the course of development.

The arachnids studied for the purpose of this article were the
Scorpions, Pedipalps, Spiders, Solifugae and more superficially the
Phalangids. In the coxal glands of each of these orders the same
fundamental arrangement as in Liniiihis can be traced, although the
number of the lobes, or saccules, to adopt Beuntz's nomenclature,

Zool. Jahrb., Supplement XIT. 16

232 B. H. Buxton,

lias been reduced to a single one in each case, with the exception
of the primitive Theraphosid spiders which have two saccules still
remaining-, one on the third and the other on the fifth appendage.
The coxal glands of the Theraphosids have also two outlets corre-
sponding to the position of the two saccules, i. e. on the third and
fifth appendages, but all the other arachnids have but a single
outlet.

In the scorpions and phalangids the outlet is at the base of
the fifth appendage, corresponding in position to that of Limulus.
In the true spiders (Araneae verae) and the Pedipalps the outlet
is on the third appendage and in the Solifugae on the second
appendage. My endeavor will be to demonstrate by means of
description and photographs that these conceptions of the arachnid
coxal gland are correct, and also to point out the various modi-
fications which the structure has undergone in the course of
specialisation.

Almost all of the animals studied were collected by myself,
killed with chloroform, and immediately fixed in Duboscq's picro-
formal- acetic solution. Before fixation, the legs and abdomen were
cut oft' so as to allow of penetration of the fixing fluid into the
cephalothorax. Imbedding first in celloidin and then in paraffin was
found to be the best method for obtaining good serial sections, as
in this way the use of xylol is avoided and the chitin, always a
difficulty, does not become so brittle. Moreover, when the more deli-
cate tissues are padded with celloidin they do not become distorted
in the hot paraffin.

For identification of the species I am indebted to Dr. Alex.
Petrunkevitch of Yale University to whom my best thanks are due.
A few specimens preserved in the ordinary way in alcohol were
also kindly provided by the late Prof. T. H. Montgomery, Prof.
Alex. Petrunkevitch and M. Eugene Simon of Paris, but naturally
sections from alcohol specimens are not often sufficiently well pre-
served for photography or for study of finer details. Where such
specimens are referred to they are indicated as such by (ale. spec.)
in brackets and the initial of the donor M., P., or S. The photo-
graphs were all taken direct from the sections and have not been
retouched.

Before beginning an account of the special work done, it may
be well to explain in a general way the characteristics of the organs

Coxal glands of the Arachnids. 233

and their cells ; the constituent parts in their order from the terminal
part to the exterior being- designated as the saccule (S), the col-
lecting tubule (CT), the labyrinth (CL), the bladder or vesicle (F)
and the exit tubule (ET) leading to the outlet (0),

The typical cells of the labyrinth coils have a striated base in
which the cell outline is lost, so that the striated layer of the cells
appears continuous (photo 5 b). Directly upon the striated layer or
immediately beneath its border, but never at its base, lie the nuclei,
generally very large and prominent with much chromatin (photo 1),
but the variations are considerable ; the cell nuclei of some scorpions,
and more particularly of the solifugae being extraordinarily minute
(photo la).

That part of the cell cytoplasm lying above the striated border
may be extensive, forming large well defined columnar or cubical
cells (photo 10), or the cell surface may be greatly worn down so
that the nuclei may appear to lie upon the striated border with
only fragments of cytoplasm attached to them (photos 15 a CL and 22).

The typical saccule which is commonly held to represent the
last remnants of the coelom, does not form definite coiled tubuleJi
like the labyrinth, but is actually of the nature of a sac, indented
in every direction by blood capillaries, which penetrate into the sac
and hang- in it; the very thin walls of the sac being lined intern-
ally with a delicate epithelium which may be flattened so as to
resemble an endothelium. The capillaries therefore would seem to
serve the same purpose as the glomeruli of the vertebrate kidney
and may be referred to as glomeruli (photo 30 GLR). The
capillaries however can hardly, strictly speaking, be considered as
such. They are simply lacunae of the hemolymph system without
any walls of their own, so that nothing lies between the body fluid
and the lumen of the saccule, except the thin wall of the saccule
itself with its lining epithelium.

At a certain point in the saccule the lumen, freeing itself from
the inhanging capillaries, becomes dilated, forming a central lumen
WSL) and from this central lumen of the saccule there runs a
short duct, or collecting tubule (CT) lined with epithelium (photo 9 b)
which then passes into the labyrinth (CL); the change from the
epithelium of the duct to the striated epithelium of the labyrinth
being very abrupt.

The exit tubule is very narrow, lined with chitin and evidently
formed by an involution of the external surface, the cells beneath

16*

234 B- ^- Buxton,

the chitin resembling those of the hypodermis. The arrangement
of muscles around the exit tubule seems to indicate that the outlet
can be closed and opened at will. Just before reaching the exit
tubule, the labyrinth tubule straightens out and sometimes becomes
dilated, forming a vesicle or bladder (Patten's "end-sac" in Limulus).

1. Scorpions.

The coxal glands of the scorpions were first described by
Lankester who recognized the tubules lined with striated epithelium
(the labyrinth), and a medullary portion, enclosed by the tubules.

The medullary portion was first recognized by Maechal (1892)
as the saccule, from which the collecting tubule passes into the
labyrinth. Maechal speaks of the lacunae of the saccule and the
lacunae of the hemolymph system, which together form the medullary
portion, but the saccule possesses definite walls lined with epithelium,
so it would hardly seem correct to describe its lumina as lacunae.

Maechal also drew attention to the homology between the
coxal glands of the scorpion, and the green glands of the decapods,
which also have a saccule and a labyrinth distinct from each other.

The coxal gland of the scorpion is very compact, situated oppo-
site the bases of the fifth and sixth appendages ; its posterior border
lying on the anterior surface of the diaphragm (D) which separates
the cephalothorax from the abdomen (photo 1, 3). The Saccule {S)
is surrounded and enclosed by the coils of the labyrinth tubules
(CL) except at one point where the blood supply enters the organ.
Opposite the entrance of the coxal artery the central lumen of the
saccule passes into a short, ill defined collecting tubule, connecting
with the labyrinth (Photos 1, la, for a general view, and lb, Ic,
for the details of the changes).

The tubules of the labyrinth form complicated coils and finally
end in a straight somewhat dilated tubule, which may be regarded
as the bladder. The bladder passes into a very narrow straight
exit tubule {ET, photo 2) lined with chitin, and obviously derived
from the external surface. The outlet appears to be functional even
in the adult in spite of much that has been written to the contrary.

Diagram Bl indicates roughly the position of the coxal gland
in cephalothorax and the plan on which it is constructed.

At the base of the sixth appendage there is a distinct dip of the
labyrinth tubule toward the surface, which in the embryo scorpion

Coxal glands of the Arachnids. 235

is very nearly reached (photo 3), although in no specimen has an
actual outlet been observed, but the dip may be an indication that
there was primarily an outlet at this point as well as the persisting
outlet on the fifth appendage. From the disposition of the coxal
gland, opposite the bases of both the fifth and sixth appendages, it
seems quite possible that it has been formed from a coalescence of
the originally distinct organs of the two appendages.

Bkauer (1894) in embryo scorpions observed the development
of nephridial "Anlagen" on segments two to eight. Those on seg-
ments five and eight persist and form the coxal gland and genital
system respectively, but the others degenerate and soon disappear.
He says nothing however, about a coalescence of the glands on the
fifth and sixth segments but remarks that the nephridium of the
sixth segment is the last of the transient ones to disappear.

The source of the blood supply for the saccule is interesting;
somewhat difterent from that found in other arachnids, and so far
as I can gather, has not hitherto been described. Accompanying
the nerve from the brain mass to each of the appendages is an
artery; a well defined thick walled vessel. The artery which ac-
companies the nerve of the fifth appendage (NV), on reaching the
base of the appendage, sends off a branch {CA) at right angles to
its course direct to the saccule of the coxal gland (photo 4. The
course of the artery can also be traced intermittently in photos 1,
and 1 a). On reaching the coxal gland at the point where the sac-
cule comes nearly to the surface of the gland, the artery breaks
up, and the fluid, passing through a mass of cellular lymphoid tissue
which may be referred to as the glomus {GL photos 5, 5 a), forms
numerous capillaries, or rather lacunae, which hang into the lumen
of the saccule; forming glomeruli in the typical manner already
described in the introduction.

So far the description of the coxal gland may be applied to
all scorpions but we now come to an accessory organ which is not
found in all of the families.

KowALEvsKY first described this organ in ButJms occitanus from
the South of France, but remarks that he could not find it in
Androdonus, an allied Buthid from Russia. Androdonus is simply
a synonym for Buthus and his species was probably B. caucasicus.
He called it the lymphatic organ to distinguish it from the lymphoid
organ which lies above the nerve cord along its whole length in
the abdomen of all scorpions. According to Kowalevsky the lym-

236 B. H. Buxton,

phatic org-au is a diverticulum from the diaphragm which becomes
filled with cells but remains open opposite the coxal glands.

It is obvious that the blood supply to the saccule of the coxal
gland must be very limited; merely a branch of a sing-le artery,
and the lymphatic org-an of Kowalevsky appears to have been
evolved as a means o^ bringing- waste products directly from the
abdomen to the coxal glands. It is probably a recent addition to
the execretory system, as there is nothing analogous to it to be
found in any of the other arachnids, nor is it present in all scor-
pions. The organ itself (LO) forms a kind of sac with a thick
hyaline wall lying in the abdomen, its interior being occupied by
small lymphoid cells, among which are also found the larger phago-
cytic connective tissue cells, often containing granules; the nephro-
cytes of Beuntz.^) Anteriorly there is an orifice which opens
through the diaphragm into the capsule of the coxal gland, but it
has no direct connection with the saccule (photos 6, 6 a, 6 b, 6 c,
at LO).

The walls of the lymphatic organ are certainly not muscular,
nor are any muscles attached to them, so there would not appear
to be any pumping action; the fluid either filtering in, or perhaps
excretory products being brought in by the large phagocytes of
the connective tissue, where they are worked over by the lympho-
cytes and passed on to the coxal gland.

Just ventral to the lymphatic organ and running parallel with
it is a large blood sinus (VS) from which, no doubt, the lymphatic
organ is supplied with waste products (photo 6).

Of the scorpions, I have examined representatives of four out
of the six recognized families.

1. Fam. Buthidae.

Subfam. ButJiinae.

Butlms australis, Biskra.

Buthus occitamis Banyuls, South France. All ages from the

advanced egg; just born; one year old
up to adult.

1) These cells help to eliminate carmine or other foreign substances
injected and therefore act as nephrocytes, but it can be shown by dif-
ferential staining that they also carry nutrient material from the intestinal
coeca, where it is stored up, to the various organs.

Coxal glands of the Arachnids. 237

Siibfam. Centrurinae.

Centrurus {?sp.), Arizona.

Tityns, Panama.

Centrums margaritakis, Jamaica and Mexico (ale. sp. P).

2. Fam. Scorpionidae.

Scorpio mmirus, Biskra, Algeria.

Biplocentrus scaber, Jamaica.

OpisthacantJms elatns, Panama.

3. Fam. Chadidae.

Euscorpius italicus, Lugano, Italy.

Ckactas {sp. ?), Colombia.

4. Fam. Vejovidae.
Vejovis flavies, Arizona.

The lymphatic organ is present and readily found in every
specimen from the families Scorpionidae, Chactidae and Vejovidae, but
no trace of it has been found in any one of the Buthid specimens
from the egg up to the adult. This fact, I think, tends to support
the hypothesis that it is a recently acquired organ, since if it had
been ancestral and discarded by the Buthids, there surely would
have been some evidence of it in the earlier stages of development.
If the organ can be regarded as a diverticulum or hernia from the
diaphragm, as suggested by Kowalevsky, its origin can be accounted
for in a very simple way.

There appears to be some contradiction here with Kowalevsky's
findings, since he describes the organ as occurring in Buthus occi-
tanus, but he himself says that he could not find it in the very
closely allied Buthid Androctonus, and it seems probable that the
specimens sent to him from the South of France were not Buthus,
but the equally common Euscorpius flavicauda (Chactid).

2. Pedipalps.

The conclusions here arrived at with regard to the coxal glands
of the pedipalps are based upon a study of 7 specimens of Tarantula
palmata from Colombia, and 4 of Tarantula fuscimana from Panama
and Costa Rica. The coxal glands of these two closely allied species
of the Tarantulidae {Phrynidae) are precisely alike in every particular

238 • B. H. Buxton,

SO that a description of one serves equally well for the other
species.

I have not been able to examine examples of the TJwJyphonidae
with the exception of a single specimen of Mastigoproctus giganteus
(ale. sp. P.) which was in such a bad state of preservation that
little could be made out of its details but so far as could be judged,
its coxal glands do not differ in general arrangement from those of
Tarantula.

Laukie found an outlet on the third appendage in Tlielijplionus
and BoRNER remarks that in the pedipalps the coxal gland is
extensive with an outlet on the third appendage; well marked in
Tlielyphonus but not so clear in the Tarantulidae.

The chief works on the development of the Pedipalps are those
of Sophie Pereyeslavzewa (1901), Gough (1905) and Schimkevitch
(1906). The two former observed in early stages the rudiment of
a second gland with indications of a lost outlet on the fourth appen-
dage, but the rudiment soon disappears and in the young pedipalp is
no longer traceable.

Borner (1906), in: Zoologica, describes the coxal glands of the
pedipalps at some length, but makes no mention of the saccule or
collecting tubule, and I cannot find in the literature that they have
been observed by any one who has recognised them as a part of the
glandular structure, although Pereyeslavzewa figures the saccule
in the embryo very clearly, but, mistaking its significance, calls it
an "agglomeration of connective tissue", forming tubules which will
give rise to additional coxal tubules. There is no evidence in her
description or figures that she found the collecting tubule, and since
the labyrinth consists of a single coiled tube it seems unlikely that
new tubules can be formed and added to it.

Schimkevitch finds in the young larva a communication with
the coelom, but in older stages the communication is lost, although
he notes the presence of "stroma cells" which he thinks may
correspond to the medullary portion of the scorpion's gland, but the
stroma cells lie outside the gland. The "stroma cells" can un-
doubtedly be referred to the saccule, but he missed the connecting
link between it and the labyrinth.

In Tarantula, the coxal glands are not compact as in the
scorpion, but elongated, stretching posteriorly from opposite the third
appendage to a point between the fifth and sixth appendages; the

Coxal glands of the Arachnids. 239

labyrinth tubules comprising by far the largest part of the gland
(photos 7, 12).

The outlet is on the posterior aspect of the base of the third
appendage (photos 7, 8) and immediately above the exit tubule {ET)
lies the saccule [S) (photos 7, 9) the details of which present much
the same appearance as in the scorpion, with its thin walls lined
with delicate epithelium, the inhanging glomeruli {GLJR) and central
lumen of the saccule {CSL) but the saccule in Tarantula is only
very partially enclosed by the labyrinth coils. The collecting tubule
(CT) which is short, but more clearly defined than in the scorpion,
forms the communicating link between the central lumen of the
saccule and the labyrinth with its striated cells (photos 9, 9a, 9b, 9c).

The labyrinth {CL) starting from the saccule opposite the
third appendage forms coils and twists running posteriorly as far
as the posterior aspect of the fifth appendage, where it turns a
little ventrally and then runs forward, underneath the coils, as a
straight tubule to the exit tubule on the third appendage. It may
be referred to as the ventral tubule {VTL) of the labyrinth
(photos 7, 7a).

In the coiled part of the labyrinth three divisions may be
recognised depending upon the character of the epithelium (photos 7,
12). Roughly speaking the anterior division {CLa) lies mainly opposite
the third appendage; the middle division (CLm) opposite the fourth
appendage, and the posterior division (CLp) opposite the fifth
appendage, but the coils of the three divisions overlap each other
and there is no abrupt change from one division to the other, such
as is found in the change from collecting tubule to labyrinth.

The cells of the labj'rinth throughout show the striated base,
but in the middle division the striated base is surmounted by well
defined large columnar cells with slightly basophil cytoplasm, the
nuclei lying at the base of the column, immediately upon the striated
base (photo 10).

In the anterior division the nuclei lie, almost bare of cytoplasm,
along the free border of the striated portion of the cell, while in
the posterior division the cells are for the most part fairly clearly
defined and cubical, although in this division there is a good deal
of variation and in some places the cytoplasm appears almost worn
down to the nuclei. These differences in the three divisions appear
to be constant; the middle division with its large basophil cells

240 -B- -S- Buxton,

«

being- always clearlj^ marked off between tlie other tw^o in both
species examined.

BoRNEE figures these cylindrical basophil cells and remarks that
they are different from the cells of the coxal glands in Thelyphonus;
and also from those of the palpigrades [Koenenia). He does not
however appear to have noticed that these cylindrical cells are
confined to one part of the labyrinth but speaks as if they were
the only kind of excretory cells found in the coxal gland.

With regard to the blood supply, there runs parallel with the
labyrinth a large blood sinus (VS) lying ventral to the labyrinth
coils (photos 12, 12a) alongside of and just internal to the ventral
tubule of the labyrinth. From the sinus small branches run in among
the coils of the labyrinth so that the tubules are all bathed in the
circulating fluid (photo 11). The sinus runs along the whole length
of the labyrinth and on reaching the saccule anteriorly, breaks up
to form the glomeruli which hang into the saccule. Photograph 11
sliows the relation of the ventral sinus to the labyrinth tubule in
transverse section.

3. Spiders.

Among the spiders there is great variation in the appearance
of the coxal glands. It has already been remarked that in the
Theraphosids there are two saccules and two different outlets, but
only one of each in the Araneae verae. The labyrinth of the
Theraphosids is very complicated but in the higher spiders becomes
more and more simplified as we pass from the Theraphosids to the
Dysderids and Sicariids, and from these through the hunting spiders,
Lycosids and Thomisids, to the highest of the web spinners like
the Epeirids and Theridiids, in which the labjTinth has become
reduced to such an extent that it is no longer recognizable as such.
The reduced condition of the coxal glands in the higher spiders
has been observed by Beetkau and Stueant, and referred by them
to degeneration, but it seems more probable that it is a case of
simplification with increased efficiency of the remainder, since the
saccule on which may be, the most important part of the work falls,
is still in good working order. The writers do not mention the
saccule and probably overlooked it. Even Beuntz, who is generally
very particular about the saccule, does not seem to recognise it in
the higher spiders, and in speaking of elimination of carmine by
them, mentions only the nephrocj^tes (amoeboid connective tissue
cells) and the malpighian tubules. Beenaed (1897) suggests that

Coxal glands of the Arachnids. 241

in the hig-her spiders the excretory products are utilized for silk
production, and the coxal glands therefore deg-enerate. The suggestion
is a good one so far as excretion hy the labyrinth is concerned,
but in all spiders excretion by the saccule seems as if it should
go on as usual, so far as one can judge by its histological appearance.
Both KowALEvsKT and Bruntz have demonstrated that in scorpions
and certain other Arthropods, some (colored) substances injected are
eliminated by the saccule and others by the labyrinth, but neither
of them appear to have recognised any excretion (of color) by the
spinning glands after injection into spiders. But the actual number
of spiders so treated up to the present time is very limited.

A. Theraphosids. Specimens sectioned and studied.

Genus ? Biskra. Immature.

„ Avicularia So. America. Adults and young just

leaving cocoon.
„ Eurypehnavagansa.ndL Panama. Adults and young with mother

rusticum and from cocoon.

„ Bugesiella Jientm Texas. (Peteunkevitch. Fixed in

sublimate.)
„ Hapdlopus pentaloris, Colombia. Immature.
„ Ischnothele Colombia. Immature.

Neodeniza mexicana Panama.

Ischnocolus Arizona.

Atyptis affinis Europe (ale. spec. S.).

The coxal gland is elongated as in the Pedipalps; the coils of
the labyrinth extending forward to the base of the second appen-
dage, and posteriorly almost to the posterior aspect of the sixth
appendage. The organ runs continuously throughout, just above and
external to the endosternite on either side. The labyrinth is so
complicated that it has not been possible to follow up the tubules
and determine if there are two distinct systems of labyrinth, one
for each saccule and outlet, but I am inclined to think not, and to
consider the labyrinth as a system of tubules common to the two
saccules and outlets.

The two saccules however are clearly distinct as can be best
determined by dorso-ventral (frontal) sections. In such sections,
tracing from the dorsum downwards, at about the middle of the
cephalothorax the labyrinth {CL) can be seen extending continu-

242 B. H. Buxton,

ousl}^ along its length from the third to the sixth appendage in-
clusive (photo 13), but no saccule is yet visible. On passing to
sections more and more ventral the labyrinth breaks up into four
parts, one of which is opposite the posterior aspect of each of the
appendages from the third to the sixth ; showing therefore, that there
are four distinct dips of the labyrinth toward the ventral surface
(photo 13 a). The two saccules on the third and fifth appendages
respectively now make their appearance, lying a little external to
the labyrinth tubules, but they are quite distinct from each other.
I have been unable to trace any saccule on the dip of the sixth
appendage, although there is some evidence of a saccule on the
fourth appendage, but no collecting tubule or change to labyrinth
in connection with it. It appears to be rather an outlying part
of the saccule of the third appendage, and not to have an indepen-
dent existence.

Examining sections still more ventral it is found that the dips
of the labyrinth tubules on the fourth and sixth appendages end
blindly not far from the ventral surface, while those on the third
and fifth pass on to their respective exit tubules and outlets, both
of which are functional in the adult as well as in the young. These
four dips with the two saccules and two exits can also be readily
traced in sagittal and transverse sections.

The dips of the labyrinth on the fourth and sixth appendages
seem to indicate that primarily there were outlets at these points
also. The dip on the posterior aspect of the sixth appendage in
particular shows in the young Avicularia from the cocoon, a very
marked string of labyrinth cells (the lumina of the tubules in these
specimens are barely apparent) extending right up to the external
surface (photo 14).

This extension is so clearly defined that for some time I thought
that there must be an outlet there also, but careful search through
all the specimens has failed to show any.

The saccules (S), as can be seen from the photographs (15, 15 a,
15 b, on third, and 16, 16 a, 16 b, on fifth appendages) present the
typical appearance as described for the scorpions and pedipalps
with the inhanging glomeruli (GLR) and the collecting tubule (CT),
passing abruptly into the labyrinth (CL). In the labyrinth the
cells have the usual striated base upon which rest enormous deeply
stained nuclei; the cytoplasm in places being worn down to the
nuclei and in other places forming large well outlined cubical cells.

Coxal glands of the Arachnids. 243

The exit tubules (ET) on the third appendage can be seen in
photographs 15, 15 a, 17, 17 a, and on the fifth appendage in 18
to 18 c. It has not been found possible to get sections on the fifth
appendage showing the saccule, collecting tubule and exit tubule
all in one field as is seen so clearly in 15 and 15a for the third
appendage.

In Atijpus, the two saccules and two outlets are present, but
the labjTinth appears to be less coiled than in the Theraphosids
proper. There is already a tendency toward simplification, but the
coxal glands of AUjpus approach much more closely to the Thera-
phosid character than to that of the Araneae verae. The finer details,
however, could not be determined as I only had at my disposal
alcohol specimens kindly provided by M. Eugene Simon.

It may be remarked that the aberrant IscJmothele, wiiich, unlike
the other members of the Theraphosid group, have become web
spinners, have lost the saccule and outlet of the fifth appendage,
retaining only those of the third appendage. The labyrinth also is
greatly reduced and simplified, so that the characters of the coxal
glands in this aberrant genus resemble rather those of the Dysderids
than of the Theraphosids in general. In the photograph (19) from
Ischnothele, the labyrinth tubule leading from the saccule is seen to
run posteriorly as a single elongated tubule without coiling, but the
walls are irregular in outline, indicating the probability that the
straightening out has been caused by the breaking through of ad-
jacent coiled tubules. There is evidence also in the other Thera-
phosids of this breaking through with consequent fusion of adjacent
coils, but it is only in IscJmothele that the process of straightening
out has gone far enough to enable one to follow up the course of
the tubule.

It is probable that but few well fixed specimens of the Thera-
phosid spiders have been available for studj', and there is some
apparent discrepancy in the reported observations. Bektkau (1885)
in AUjims found an outlet on the fifth appendage, and Sturany (1891)
confirmed this observation, but could find no outlet in a single alcohol
specimen of Mtjgale, and Pelseneer (1885) had also previously come
to the conclusion that in a large Theraphosid examined by him
there is no efferent duct either to the exterior or to any internal
organ.

Schimkevitch (1911) has recently studied the development of
a Theraphosid {Isclmocoliis) from Java, and in the embryo found an

244 ^- H. Buxton,

outlet on the third appendage, but denies the existence of one on
the fifth, although the cells dip down very close to the surface.
ScHiMKEviTCH figurcs Sagittal sections in which I have generally
found that the outlet on the third appendage is clearly defined, but
to observe the whole exit tubule and outlet of the fifth appendage
in one field, it is necessary to have transverse sections. In sagittal
sections the course of the exit tubule on the fifth appendage must
usually be followed up from section to section and it is often diffi-
cult to determine if there is actually an outlet or not. Photograph
18 b is taken from the only sagittal section 1 have which shows the
exit tubule of appendage V in its entirety. It may be observed
that 18, 18 a, 18 c are all taken from transverse sections.

ScHiMKEviTCH docs uot meutiou the saccule, and his specimens
were perhaps too young to show it clearly. I have myself been
unable to identify the saccules with certainty in specimens of Avi-
cularia from the cocoon, although in those of Eunjpelma (photo 15)
at about the same stage of development it is already clearly defined.

Beuntz (1904), so far as I can gather, is the only author who
mentions the existence of a saccule. He found it in Atypus and
demonstrated that the cells of the saccule eliminate carmine, but
not those of the labyrinth. Bruntz was unable to obtain specimens
of the Theraphosids, but in a living Bugesiella lienisii from Texas
(given to me by Dr. Peteunkevitch) inoculated with carmine and
killed three days later, the same results were noted as by Bruntz
for Atypus, namely that the cells of the saccule took up carmine,
but not those of the labyrinth (photo 20). Both of the saccules
were alike in this respect.

The fact, however, that there are two saccules and two outlets
in the Theraphosids does not appear to have been recognised by
anyone.

B. Araneae verae.

In all the Araneae verae there is a single saccule lying mainly
opposite the third appendage, but extending also posteriorly to about
opposite the fourth appendage. The thin walls of the saccule are
lined with delicate and sometimes fiattened epithelium, and glomeruli
hang into its lumen in the usual way. The saccule therefore preserves
its typical appearance, but the labyrinth has become greatly reduced
and simplified. According to the degree of simplification of the
labyrinth the Araneae verae may be divided into three groups. In

Coxal glands of the Arachnids. 245

all of them there is an exit tubule with an outlet on the third
appendage functional in the adult as well as in the immature. The
collecting tubule has been lost and the lumen of the saccule passes
directly into the labyrinth tubule.

Group 1. Sicariids and Dysderids.

The six-eyed Sicariids and Dysderids show the most primitive
condition of the labyrinth. Starting from the main body of the
saccule opposite the third appendage, it runs posteriorly as a single
straight tubule as far back as the sixth appendage where it turns
inwards, and then runs forward as a straight dilated tubule, inter-
nal to the first tubule, and along the endosternite to its outlet on
the third appendage (Diagram A).

Diagram A.

Coxal gland of group 1 of the Araueae verae.

The saccule {S) opposite appendage III, leads into the straight external
tuhule (LET) which runs posteriorly as far as appendage VI where it turns
and runs as the internal tubule [LIT) anteriorly to the outlet on appendage III.
Both (LET) and [LIT] are lined with striated epithelium. The vertical broken
lines indicate approximately the points through which transverse sections were
cut from different specimens and photographs taken (21 to 24).

Since the labyrinth tubule running posteriorly from the saccule
is, in the true spiders, always external to the other, it may be
referred to as the external tubule {LET) and the tubule running
anteriorly to the exit, as the internal tubule {LIT), the two to-
gether composing the labyrinth. In Scytodes (Sicariid) there are a
few loops at the posterior end of the labyrinth, but these are wanting
in Loxosceles (Sicariid) and Dijsdera, in which the labyrinth simply
runs backwards and then forwards again, so that in transverse sec-
tions one sees two tubules cut across, the external one leading from
the saccule posteriorly; the internal one close to the endosternite,
being the return tubule to the exit (photos 21, 22). The labyrinth
cells have the usual striated base on which the large nuclei rest,

246 S- H. Buxton,

but the cells are generally much worn down and the nuclei lie
almost bare of cytoplasm on the free border of the striation.

Photograph 23 shows the passage of the saccule into the exter-
nal tubule and photograph 24 the saccule and internal tubule. Dia-
gram A explains of itself how the internal tubule at this point
appears to be double. The photographs 21 to 24 are taken from
sections cut transversely approximately at the points indicated by
the broken vertical lines in the diagram.

Group 2. The hunting Lycosids, Thorn isids, Attids,

Drassids, Zodariids, Selenopids, Ctenids, and the

imperfect web spinners, Agelenids and Dictynids.

The labyrinth in the members of this group of spiders shows
very close resemblances, and one description will serve for all.

The walls of the external tubule leading posteriorly from the
saccule, have become very thin and are lined with a flattened epithe-
lium, the nuclei of which are not prominent so that the tubule
resembles the saccule, and until the tubule is closely followed up
from section to section it is often difficult or even impossible to
determine which is the saccule and which the external tubule.

The external tubule runs, as in the Dysderid group, posteriorly
to the sixtli appendage where it turns forwards again. At the turn,
or a little before the turn (photo 25, 25 a) the lining epithelium
changes its nature and becomes cubical and larger, remaining how-
ever relatively fine and delicate, and showing no evidence, or
mere traces, of the striated base, which is such a marked feature
in the typical labyrinth. The internal tubule is usually greatly
dilated and shows dips towards the surface between the fourth and
fifth, and the fifth and sixth appendages.

It has also dorsal extensions opposite the dips and also one
opposite the exit. It would seem to serve the purpose of an ex-
tensive bladder rather than of an excretory gland (photo 26, 27)
and probably the saccule alone has excretory functions. In some
members of this group, e. g. the Zodariids (photo 28) the external
tubule seems to have become actually fused with the saccule.
The process of fusion appears to be due to glomeruli pushing
in the thin wall of the tubule, so that its lumen becomes
irregular and broken up, glomeruli hanging into it, as into the
saccule itself.

Coxal glands of the Arachnids.

247

m

IV

Diagram B.

Comparison between

the coxal glands of

the scorpions, pedi-

palps and spiders.

1. Scorpions. The
saccule {S) surround-
ed by the labyrinth
coils, which termi-
nate in a vesicular
dilatation and an
outlet on appen-
dage V.

2.Pedipalps.From
the saccule the coiled
labyrinth tubule ex-
tends posteriorly as
far as appendage VI,
where it turns and
runs straight for-
wards to the outlet
on appendage III.

3. T h e r a ]) h s i d
spiders The two
saccules on appen-
dages III and V
communicate with
the coiled labyrinth
tubules and through
them with the two
outlets on III and V.
The exact relations
of the labyiinth with
the saccules and
outlets have not

been determined.

4. Aran eae verae.
Group 2. The Hun-
ting Spiders.

The saccule is exten-
sive but tlie tubule
leading from it poste-
riorly has become
straigliteued out and
the walls so thinned
down that they often CI

appear fused with

the saccule. The tubule leading anteriorly to the outlet on appendage III is
s-reatly dilated and appears to act only as a vesicle since the lining cells have
lost their striation.

5. Araneae verae. Web Spinners.

The saccule is well preserved but the labyrinth has become fused with it,
with the exception of a short vesicular portion leading to the exit tubule and
outlet on appendage III.

17

Zool. Jahrb., Supplement XIV.

IV

VI

248 B. H. Buxton,

On comparing the photographs of the labyrinth of group 1
(photos 21, 22) with those of group 2 (photos 26, 27), it will be
observed that although the tubules of group 1 are very much
shortened as compared with those of the Theraphosids, yet the
striation is very highly perfected ; more so in fact than in any of the
other Arachnids, except towards the anterior part of the internal
tubule in the neighborhood of the saccule (photo 24), where the striated
part appears somewhat reduced and there is a tendency towards
vesicular enlargement. It seems probable therefore that the ex-
cretory functions of the labyrinth of group 1 have not been in any
way impaired, whereas in group 2, they have been entirely or
almost entirely lost, and possibly, as suggested by Bernard, trans-
ferred to the production of silk.

Group 3. Epeirids, Theridiids, Pholcids,

F i 1 i s t a t i d s.

In this group, the members of which are all expert web spinners,
the labyrinth reaches its extreme limit of simplification. Practically
nothing is left of this part of the gland but a short bladder (F)
leading to the exit tubule (ET); the bladder opening above im-
mediately into the saccule (S) with its inhanging glomeruli (GLR)
(photos 29 to 32). The walls and epithelium of the labyrinth have
become thinned down throughout, and so fused with the saccule
that it is impossible to distinguish between them.

The well preserved condition of the saccule however and the
well marked opening on the external surface, show clearly that
the gland must be functional, although it would obviously be
impossible to homologise such an organ with that of the other
arachnids if it were not for the intermediate steps which can be
traced back to the Theraphosids through groups 2 and 1 of the
Araneae verae.

Diagrams A and B show roughly in a general way the main
points of the difference and similarity between the coxal glands of
the Scorpions, pedipalps and spiders, and the description attached to
the diagrams will serve also as a summary for Part I.

Coxal glands of the Arachnids. ' 249

Part II. Tracheate Arachnids and Peripatus.

Tracheate Arachnids. Solifugae and Plialangids.

Solifugae.

Specimens studied:

Galeodidae.

Galeodes arabs, Biskra, Algeria.
Galeodes (sp.?), Biskra, Algeria.
Paragcdeodes barbarus, Biskra, Algeria.
Paragaleodes scalaris, Biskra, Algeria.

Solpugidae.

Solpuginae. Solpuga flavescens, Biskra, Algeria.
Doesinae sp ?, Colombia.
Boesinae: Ammotrecha, Texas (ale. sp. M.}.
Eremobatinae \ Eremobaies, Texas (ale. sp. M.).

In the description of the coxal gland of the solifugae we shall
find so many new and remarkable features, that the inferences drawn
may very likely be erroneous in some respects. But in order to avoid,
as far as possible constant repetition of "I think" or "I believe",
I shall treat the subject dogmatically and appear to take it for
granted that my views are correct.

The coxal gland lies loosely in the hemocoele; the loops and
coils of its tubules extending posteriorly as far as the sixth append-
age. The main portion of the gland lies between the second and
third appendages and from this point a long narrow tubule runs
anteriorly with an outlet on the second appendage. The more
detailed description of the gland given below applies to Paragaleodes.
Variations from the type of Paragaleodes in other genera will be
commented upon later.

Beginning with the saccule [S) which lies in the main portion
of the gland we find thas it hat undergone considerable modification.
Since the solifugae, being tracheate animals, have no system of
blood vessels outside of the long narrow heart {H\ there is no means
of carrying blood directly to the gland as in the pulmonate scorpions,
pedipalps and spiders. To remedy this defect, the saccule appears
to have been developed into a pumping organ designed to force fluid
from the hemocoele into the coxal gland. All around the periphery
of the saccule are attached bands of finely striated muscle fibres
{MS) which radiate out on all sides (photo 33, 34, 34 a, 34 b) and

17*

250

B. H. Buxton,

at their distal ends attach themselves to anything which crosses
their path; walls of tracheae {TR), tubules of the coxal gland, or
to other muscle bundles.

Diagram C.

Oaleodes [^ Othoes). Frontal view.

Shows in a general way the disposition of the coxal glands.

The saccule (<S) and collecting tubule passing into the labyrinth sac {S.L)
(only partially drawn; the posterior extension is omitted) which changes to the
coiled tubule of the labyrinth {CL). The tubule [CL) runs posteriorly and then
anteriorly again to the exit tubule {ET), terminating in the nozzle (NZ). In
this species there is a definite tubule beyond che nozzle with its side outlet {SO)
and final outlet {FO) not far from the opening of the rostrum to the exterior.

In the diagram the tubule has been drawn pointing a little too much inwards.
It_ should have been drawn more obliquely. (Compare with photo 8.) The rostrum (E)
■with its sucking chamber lies in front, and the sucking chamber passes posteriorly
into the narrow oesophagus between the two lobes of the brain (dotted ovals).
The straight dotted line above indicates approximately the level at which the
chelicerae project from the body above appendage II.

Coxal glands of the Arachnids.

251

n

in

A^

It is obvious that when these muscles are contracted the walls
of the saccule would be pulled out, the lumen would become distended,
and filled with fluid filtering- in through the walls. On relaxation
of the muscles, the saccule would tend to collapse upon itself, and
press upon the contained fluid which would thus be forced through
the collecting tubule into the labyrinth.

To increase the
extensibility and also '

no doubt to strengthen
the walls of the sac-
cule, they are rein-
forced internally by
a thick layer of
spongy cellular tissue
(photo 33), between
the cells of which
run innumerable fine
fibres, which may be
extensions of the
muscle fibres, but the
fibres themselves do
not appear to be con-
tractile. Although the
cells of the spongy
tissue have lost all
appearance of epi-
thelium they are pro-
bably derived from
the original epithelial
lining cells of the
saccule.

There is a con-
siderable lumen (CSL)
in the center of the
saccule, and pouches
from the central lumen

TRS

V

VI

Galeodids

Solpugids

Diagram D.

Showing an outline of the coxal glands and the
way in which a second extension posteriorly of the
labyrinth sac in the Solpugids has led to the change
from SL to CL being removed for some distance
from the main part of the gland. In the Solpugids
also the coiling of the labyrinth tubule is much more
complicated than in the Galeodids.

extend into the spongy

tissue on all sides (photos 34, 34 a, 34 b).

Although this part of the organ has undergone such striking
modification that by itself it could no longer be recognised as the

252 S- H. Buxton,

saccule, there can remain little doubt that it does actually represent
the saccule when its relations with the rest of the gland are taken
into consideration.

At one point in the central lumen of the saccule a funnel like
collecting tubule with definite epithelial lining begins and runs as
a long narrow duct opening into the labyrinth (photo 35). The
labyrinth can be divided into two very distinct parts, the first of
which is not a system of coiled tubules, but is more of the nature
of a sac with innumerable pouches (photos 37, 37 a). The German
word "Schlauch" would best describe it, but it may be called the
"sac of the labyrinth", or "labyrinth sac" (SL) to distinguish it from
the true saccule.

In its simplest form (Paragaleodes), the long narrow multipouched
sac runs posteriorly to a little beyond the tracheal stigmata (TRS)
of the thorax which lie in a space between the fourth and fifth
appendages (Frontispice and diagram D), where it ends blindly.
The walls of the labyrinth sac are lined throughout with tall
columnar epithelium without any striation (photos 36, 37 and 45 (SL)
show this best); the closely packed nuclei lying at the base of the
cells immediately over the limiting membrane of the Avail.

The lumen of the sac is sometimes filled with a granular
detritus and when this is the case the cells appear worn down and
indistinct (photos 37 a, 42a). This part of the labyrinth appears
therefore to be a secreting rather than an excreting organ and may
be considered as such provisionally.

The structure and the cells of this part of the coxal gland are
something quite difierent from anything we have hitherto encountered
in the labyrinth and for some time I considered it to represent the
saccule in a modified form, and took the saccule to be a new
structure of the nature of a glomus, but this view proved to be
erroneous.

I find that Beenard appears to take this mistaken view of the
situation. He observed the two distinct parts of the labyrinth and
considers that the proximal part (labyrinth sac, of this article), can
be homologized with the proximal or medullary part (Lankestee)
of the coxal glands of the scorpions. But he overlooked the true
saccule and collecting tubule altogether and thought the outlet was
on the third appendage, so that his observations at any rate were
incomplete and in some respects actually erroneous. Bernard also
discusses the question whether these proximal parts of the glands

Coxal glands of the Arachuids. 253

in the solifugae and scorpions can be considered as tlie remnants
of the coelom or not, and comes to the conclusion that they cannot
be so considered, but are simply modifications of the ordinary laby-
rinth tubule; also a mistaken view in all probability.

At a point in close proximity to the saccule, the sac of the
labyrinth [SL) passes abruptly into the second part of the labyrinth
(CL) (photos 36, 36 a) which is typical in every respect. We find
here the coiled tubules lined with cells having the typical striated
base upon, or just w^ithin, which the nuclei are placed. From its
opening' from the secreting sac (diagram D, Galeodidae) the tubule
runs posteriorly as far as the sixth appendage, w^here, after forming
a few coils, it turns and runs anteriorly again to the main part of
the gland. Here, after coiling a little upon itself, it passes on
anteriorly as an almost straight tubule to the base of the second
appendage.

As the exit tubule {ET) it then extends forwards along the
dorsal side of the coxal joint of the second appendage just under-
neath the chitinous tegument its walls becoming muscular (photo 38),
to its outlet at a point about opposite the front of the* median eyes,
and just below the base of the chelicera. The tubule here turns a
little inwards and terminates in a strong chitinous ring with a
minute orifice in its center, which acts as a nozzle {NZ) (photos 38,
38 a, 38 b).

The nozzle lies in a groove, formed by two flaps of the chitin-
ous integument, which extend as a sheath {LE) for some distance
further forwards and provide a kind of conduit for the fluid ejected
from the nozzle (photo 40, 40 a).

The second appendage (palp) is always directed forward and
the dorsal aspect of its coxal joint works under, and rubs against
the ventral surface of the enormously developed first appendage
(chelicera), so that this extension of the exit tubule and the chitin-
ous sheath beyond its outlet run anteriorly and finally a little
inwards.

Since the natural position of the outlet would be at the base
of the coxal joint of the appendage it seems probable that this
anterior extension of the tubule along the coxa, with its muscular
w^alls and nozzle like outlet, must be a secondary development de-
signed to serve a special purpose, and in fact the outlet is situated
at a point from which fluid ejected with some force would be pro-

254 2- H. Buxton,

jected towards the outlet of the rostrum and the prey held hy the
chelicerae in front of the rostrum.

The apparatus for forcibly ejecting the fluid is there, and, the
chitinous sheath through which the stream passes w^ould keep it in
the right direction, since in the solifugae the mouth parts, instead
of being situated ventrally, as in the spiders and pedipalps, are
directed straight forward, the rostrum (R) forming a sucking chamber
(KS) in front of the mouth proper, and extending anteriorly between
the chelicerae (photo 39).

The solifugae have a bad reputation in many places as poi-
sonous animals, but no definite poison glands, such as exist in spiders
and scorpions, have ever been found. Pocock suggests that the
poison may exude from small setal glands situated in the teeth of
the chelicerae, but I do not know that this has ever been demon-
strated, and it seems more probable that the poison, if there is any,
is the product of the coxal gland.

When the coxal glands were first observed in the solifugae
they were taken for poison glands (in: Cambridge nat. History),
but this view was refuted and the glands are now considered to
be excretory, and homologous with the coxal glands of other arach-
nids. However it is possible that the older observers were not so
far wrong after all.

In any case the coxal gland would appear to act as a salivary
gland. In the spiders and scorpions special salivary glands have
been evolved on the coxal joints of those appendages which are
situated closest to the mouth and do most of the masticatory work.
Such salivary or buccal glands in the spiders are localised at the
base of the coxal joint of the second appendage, and in the scorpions
of the third and fourth appendages, but are quite independent of
the coxal or poison glands. There are no such separate buccal
glands in the solifugae w^hich affords an additional reason for sup-
posing that the coxal glands perform the duty of salivation.

Briefly then the spiders and scorpions have three distinct sets
of glands, poison, salivary and excretory, but in the solifugae all
these duties are performed by the coxal glands, parts of which are
specially modified for the purpose.

Before concluding, a few additional observations may be recorded.
With regard to the saccule; in some specimens of Paragaleodes, on
one side of the animal the collecting tubule is open, and its course
can be readily traced from the saccule to the labyrinth sac (photo 41,

Coxal glands of the Arachnids. 255

41a), but on the other side of the same specimen the colleetini^
tubule is closed by a bulge (B) from the wall of the labyrinth sac
(photo 42, 42 a) the bulge protruding also into the lumen of the
saccule (photo 46 is a compound of 41 and 42).

It appears that the muscle fibres of the saccule on this side
are contracted, and in pulling out the walls of the saccule a partial
vacuum is temporarily formed in its lumen, and the labyrinth sac,
which surrounds the collecting tubule, closes up over it; forming a
valve which prevents any back flow through the tubule from the
labyrinth sac, thus allowing the saccule to fill up again with fluid
from the hemocoele outside. As the muscles become relaxed the
tubule opens again and fluid is forced from the saccule into the
labyrinth.

In connection with the saccule, it may further be remarked that
the tissue composing it varies much in different genera (compare
photos 34, 34 a and 34 b). In Paragaleodes (34) it is very loose and
porous; less so in Solpuga (34a) and in Gakodes (34b) relatively
compact and dense. The tissues surrounding the saccule also show
the same progressive condensation. Finally in the specimen Galeodes
sp.? not only is the tissue of the saccule very compact, but the
special muscles (MS) appear to be wanting, so that it is doubtful
if there can be any pumping action.')

The collecting tubule is usually long and narrow, lined with
epithelium differing from that of the labyrinth sac on the one hand
and from the cells of the saccule on the other, but in the specimen
of a Galeodes just mentioned, the collecting tubule appears to be
lost as a separate entity, and the change from the cells of tiie
saccule to the columnar cells of the labyrinth is abrupt as is shown
in photograph 45, but the collecting tubule is in the usual place
between two pouches of the labyrinth sac. It may be that the loss
of the pump and the alteration in the character of the collecting
tubule have some connection with each other.

In this specimen also the columnar cells of the labyrinth sac
(SL) are much more regularly disposed than in any other examined,
and the columnar character is even retained in the cells of the

1) I am now inclined to think that this specimen belongs to the
genus Othoes of the Galeodidae (Nov. gen. Hirst 1910). In its internal
organisation it differs considerably both from Galeodes and Paragaleodes,
yet stands nearer to them than to the Solpugids.

256 B. H. Buxton,

labyrinth coils {CL), but in CL the nuclei are raised from the
base of the cells, and lie upon a striated border, the striation how-
ever being less conspicuous than is usual. A g-eneral view of this
specimen with the coxal glands and their relation to other parts of
the cephalothorax is shown in diagram C.

As to the labyrinth, the description given of it applies to the
Galeodids, in which the coils and twists are few and the tubules
can be followed up without difficulty throughout their course, but
the Solpugids, althougii fundamentally the arrangement is tlie same
as in the Galeodids, have a much more complicated labyrinth, but
it will be sufficient to refer to the diagram D, which shows how
the labyrinth sac has developed a second extension posteriorly,
dragging with it the point of change to the striated tubule, so that
the change in this case is far removed from the main part of the
gland. The striated tubules also become so greatly coiled in the
Solpugids that it is impossible to follow them accurately throughout
their whole length (photo 43).

There is much more coiling and twisting than is indicated in
the diagram of the Solpugids but that of the Galeodids shows
approximately all the coiling. In neither case however has any
attempt been made to show in the diagram D the pouching of the
labyrinth sac, a better idea of which can be obtained from the
photographs (37 and 37 a).

The pouching of the labyi-inth sac is most conspicuous in Fara-
galeodes. In the Solpugids and Galeodes the pouches are fewer but
individually longer.

The chitinous groove or sheath along which the stream from the
nozzle is directed towards the rostrum varies somewhat in appearance
in the diiferent genera. In Paragaleodes it is short and imperfect, but
in Solpuga much better developed, forming a long forwardly pro-
jecting sheath standing out from the coxa of the appendage (photos
40, 40 a). Finally in OiJwes (?) it is converted into a definite tubule
lined with epithelium which, starting from the nozzle, runs inwards
with a outlet just before reaching the rostrum (photo 39, LR and
diagram C). There is however a side outlet just beyond the nozzle, •
the object of which would appear to be to allow excretory products
to dribble out during a period of quiescence, and probably this side
outlet is closed and the fluid forcibly ejected through the tubule
beyond it for salivary purposes.

The side outlet is present also in other genera and in the photo-

Coxal glands of the Araclmids. 257

graph 44 of a transverse section from Eremohates can be seen on
the left of the rostrum at (^0). Just anterior to this point the
sheatli closes up again and proceeds to the final outlet {FO in
photo 39), a little further forwards. The photograph 44 and dia-
gram C show that the side outlet is directed away from the mouth,
whereas the final outlet is directed towards it.

It seems strange that there is no definite vesicle or reservoir
near the outlet for storing the products of secretion since these have
to travel so far before reaching the exterior, but the exit tubule
appears to be distensible. The photograph 38 b shows this condi-
tion of distension as observed in a single specimen of SoJpuga, but
in all other specimens examined the tubule is narrow and contracted
as in photographs (38, 38 a). Perhaps at the moment of death the
muscles usually contract and drive out any secretory products which
may be present in the exit tubule.

Phalangitis.

The Phalangids have not been studied at all closely, but in the
few specimens examined, the labyrinth is clearly marked out, and
there is an outlet on the fifth appendage. The saccule is small and
surrounds the exit tubule, but I have been unable to find the com-
munication between it and the labyrinth through the medium of a
collecting tubule. These observations simply confirm those of
Bruntz. Bruntz also mentions that there is always a large nerve
passing through the folds of the saccule. This observation, I can
also confirm.

Summarising the knowledge thus far gained we find that in
the coxal glaads of the scorpions, pedipalps and true spiders there
is a saccule, a collecting tubule, a labyrinth Avith striated cells, and
an outlet either on the third or fifth appendage, while the primitive
Theraphosid spiders have two saccules and two outlets; one cor-
responding to the third and the other to the fifth appendage.

In these groups then the coxal glands appear to be quite homo-
logous, but the solifugae possess in addition a secreting sac and
the outlet is on the second appendage not far from the mouth, so
that the gland acts as a salivary as well as an excretory organ
and also possibly as a poison gland.

On examining, without any special object in view, some sec-

258 S- H. BCXTON,

tions of Peripatus (specimens given by Dr. Alexander Petrunke-
viTCH and Dr. M. Grabham of Jamaica), I was astonished to find
that they threw a great deal of light upon the reasons for these
differences in the coxal glands of the Arachnids.

Perlpcttns,

Specimens examined :

P. juUformis. Male (Petrunkevitch, fixed in sublimate).

P. jamaicensis. Female (Petrunkevitch, fixed in sublimate).

P. juUformis and P. jamaicensis (Geabham, preserved in formol).

All of the above were collected in Jamaica and in addition I
have examined three adult females containing embryos, of P.
nicaraguensis var. isthmicola (Bouvier) collected by myself in Costa
Rica and fixed in Duboscq's solution.

Peripatus has a vai'iable number of legs, ranging from sixteen
in the Australian species to about forty in P. jamaicensis, and in
front of the legs are two segments; the jaws, corresponding to the
chelicerae, and the oral papillae, corresponding to the palps of the
Arachnids. The antennae, corresponding possibly to the rostrum of
the Arachnids arise from the pre-oral segment (Sedgwick) and are
not here considered to have any segmental value.

On each of the leg bearing segments of Peripatus there is a
nephridium; the first five and the last three of which, irrespective
of the number of legs, are modified from type. Thirty legs is the
usual number for the American species and in describing the
nephridia it will be convenient to take the number of legs as thirty,
and the segments therefore as thirty-two. The genital segment near
the posterior end has no nephridium, but the posterior segments
have no bearing on the present argument, so for the sake of brevity
the genital segment will be disregarded.

The typical nephridium found on segments eight to twenty-nine
possesses a saccule {8) from which a funnel bearing collecting tubule
[CT) leads into the labyrinth (CL) with its striated cells and
nuclei in the usual position (photos 47, 47a) but the striatiou is not
so clearly marked as in the Arachnids. The diagram E indicates
roughly the way in which the labyrinth coils and finally enters
the vesicle, but for a short distance at the loop (Lilf, of diagram
and photographs), the lining cells take on a totally difterent
character. The striation is lost and the nuclei lie at the base of

Coxal glands of the Arachnids.

259

granules; cells
47 to

therefore
47d L3I).

tall columnar cells, containing- basophil

which appear to have secretorj' functions (photos

The nephridia of segments six
and seven, on legs four and five,
have become modified from tj^pe in
the sense that the labyrinth is
very extensive; its tubules forming
complicated twists and turns, but
the modified secretory portion of the
labyrinth is entirely lost. The
vesicle has also disappeared, so that
from the collecting tubule, to the
exit tubule there is an unbroken
stretch of labyrinth with striated
cells, as in the scorpions, pedipalps
and theraphosid spiders.

In the modified nephridia of the
first three and last three legs on
the contrary it is the striated portion
of the labyrinth which is entirely
missing. The short collecting tubule
passes directl}' into a tubule lined
with cells resembling those of the
modified part of the typical labyrinth;
this tubule proceeding straight to
the exit tubule without any vesicular
enlargement (photo 49 NM).

The modified nephridium of the
second appendage, known as the
salivary gland, presents some very
remarkable features. The saccule
and collecting tubule can readily be
found and recognised in photos 48,
50, 50 a, 50 b, but the collecting
tubule opens into a diverticulum
(photo 51 DSL) from a long
narrow sac (SL), which extends

posteriorly almost to the end of the body (diagram F 1), occupying
a space in the lateral sinus, dorsal to the segmental nephridia.
The sac is almost straight, but shows a tendency towards pouch-

Diagram E.

Typical nephridium oi Peripatus.

The saccule (S) passes into the
funnel bearing collecting tubule (CT).
The striated part {CL) coils upon
itself, and at the bend in close
proximity to the collecting tubule,
there is a short stretch lined with
epithelium of a secretory nature (LM),
beyond which the striation begins
again, the tubule linally passing to
the vesicle and outlet at the base of
the appendage.

260

B. H. Buxton,

ing', and may or may not be filled with products of secretion
(photos 50 and 52 SL). The lining cells vary somewhat in different
parts of the sac, but all are of the fundamental type of a secreting
cell; those in the diverticulum leading to the collecting tubule being
filled with basophil granules (photos 50, 51 DSL), resembling very
closely those of the cells of the modified part of the labyrinth in
the typical nephridium (diagram E LM, and photo 47 Lili) of
which modified part the entire sac is probably a gigantic extension
and further modification. A comparison of the basophil granules in the
diverticulum (DSL) of the salivary gland, with those in the modified
part (LM) of the typical nephridium, and of the short tubule of
the small modified nephridia is made in photographs 54, 54 a, 54 b.
Not far from the collecting tubule and ventral to it (diagram F 1)
the sac opens into a tubule lined with striated cells (photo 53 CL)

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Diagram P.

Comparison between the salivary gland of 1. Peripatus and the coxal gland
of 2. the solifugae.

The figures refer to the parts corresponding to those of the typical nephridium
in diagram G 1.

1 Saccule. 2 Collecting tubule. 3 Missing. 4 Labyrinth sac. 5 Striated
part. 6 Missing 7 Exit tubule.

Coxal glands of the Arachnids. 261

which shortly passes into the exit tubule. The exit tubule joins
with that of the other side, and the two have a common outlet just
behind the mouth. There is a diverticulum from the striated divi-
sion, running- posteriorly (CR), called the "reservoir" by Bouviek,
the position of which can be gathered from the diagram (Fl).

Bouviek (1905), whose magnificent work is mainly systematic,
does not figure the saccule and collecting tubule of the salivary
gland, but they have been described and figured by v. Kennel (1886)
in embryos of the American species P. edwardsii and P. torquatus.
V. Kennel however says that in the adult the saccule disappears
and the collecting tubule is closed.

Sedgwick (1888) also describes the saccule and collecting tubule
of the salivary gland in embryos of P. capensis, remarking that they
persist in the adult, and this is certainly also the case in the spe-
cimens examined by myself, in all of which the saccule is clearly
present and the collecting tubule open. In P. nicaraguensis (photo 50)
the saccule and collecting tubule are somewhat reduced in size as
compared with those of the typical nephridium, but in P. jamai-
censis (photos 48, 50 a) they appear quite normal.

Peripatus also aft'ords an excellent example of the way in which
the extension of one organ, may lead to atrophy of others which
were originally its equal. In most species of Peripatus the striated
diverticulum (reservoir) of the salivary gland extends posteriorly to
the third leg, and the nephridia of the first three legs have lost
their striation. In P. janiaicensis however the striated part
of the salivary gland stretches back as far as the twelfth leg,
according to Bouvier, although in my specimens it does not
extend beyond the seventh or eighth leg. However this may
be the nephridia of legs six to twelve inclusive have all become
modified in the same manner as those of the first three legs, that
is, they are very small and have lost their striated part and vesicle.
But this modification does not affect the large nephridia of the
fourth and fifth legs, which in P. jamaicensis are relatively larger
and the striation of the cells more distinct than in the other two
species examined. It is difficult therefore to be sure whether the
salivary reservoir or the large nephridia of legs four and five have
had most eft'ect in reducing those which lie more posteriorly, but
the fact remains that in P. jamaicensis the excretory functions of
the first fourteen segments are performed by three large systems
of well defined striated tubules, whereas other species, in wiiich

262 B- H. BtjxTON,

these three systems are less developed, possess in addition seven
smaller systems of striated tubules in the iirst fourteen segments.

The similarity of the salivary gland of Perqmtus to the coxal
gland of the solifugae is most striking (diagram Fl and 2) and
there can be little doubt that they are homologous organs ; the long
narrow sac in each instance being an extension of the original
modified part of the labyrinth which can still be recognised in the
typical nephridia of Peripatus. Both have this extensive secreting
sac, which may or may not be filled with products of secretion
(photos 37, 37 a of solifugae and 51, 52 of Feripatus), and the outlet
on the second appendage close to the mouth opening.

The saccule in both cases lies anteriorly to the collecting tubule,
and the collecting tubule itself passes posteriorly to the labyrinth
sac; a point which has not been sufficiently indicated in the diagram (F).

With regard to the outlet in the solifugae we may infer that
as the mouth opening extended forwards the exit tubule of the
salivary or coxal gland had to follow it along the coxal joint of the
palp, and finally reaching a point where the joint begins to turn
outwards (Frontispice B of entire animal, ventral view), the exit tubule
itself could extend no further forwards so developed the squirting
arrangement which has been described, and the chitinous sheath to
keep the fluid in the right direction.

The coxal glands of the scorpions, pedipalps, and spiders on
the other hand appear to have lost the modified part of the laby-
rinth and resemble in a general way the large nephridia found in
two of the segments of Peripatus, except that the outlets are on the
third or fifth appendages with indications of lost outlets on the
fourth and sixth, and not on the sixth and seventh as in Feripatus.

In the scorpions and spiders, as we have seen, special poison and sali-
vary glands have been developed for these parts of the work, but such
special glands are absent in Tarantula. The outlet of the coxal gland of
Tarantula on the third appendage is not far from the ventrally placed
mouth, and we have seen that in the middle part of the labyrinth the cells
are columnar and basophil, so it is possible that they act as secreting
cells, and that the coxal gland in this case also plays the part of
salivary as well as of excreting gland. However these cells of the
middle part of the Tarantula labyrinth have the usual striated base
on which the nuclei rest, and do not resemble in this respect the
secretory cells of the labyrinth sac in Feripatus and Galeodes. U
then these cells are actually secretory it is probable that they have

Coxal glands of the Arachnids.

263

become so secondarily and do not represent the modified part of the
Peripatus labyrinth.

2.

3.

4.

6.

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Diagram G.

The typical and modified nephridia of Peripatus compared with each other
and also with the coxal glands of the Arachnids.

The relative sizes of the different parts are approximately shown in 1, but
in the other figures it has simply been attempted to show which of the original
parts have been preserved, and which are missing, without regard to the relative
sizes of the parts.

1 Saccule. 3 Collecting tubule. 3 First .striated excretory part. 4 Modified
secretory part. 5 Second striated excretory part. 6' Vesicle. 7 Exit tubule.

Those parts of 1 which are missing in the other figures are represented by
a single broken line, and doubtful parts by a double broken line.

In diagram G a comparison is drawn between the nephridia of
Peripatus and the coxal glands of the Arachnids. In the typical

Zool. Jahib., Supplement XiV. ^^

264 -B. H. Buxton,

nephridium of Feripatus (1) the relative sizes of the component
parts are approximately given, but for the other figures it has
merely been attempted to indicate which of the original parts have
been preserved or lost without regard to their relative size. In the
scorpions and higher spiders we find a vesicular enlargement just
before the exit tubule is reached, but it does not appear that this
vesicle can be homologized with that of the typical nephridium in
Peripatus. In neither scorpions nor spiders is the vesicular enlar-
gement lined with cells differing from those of the labyrinth, nor
has it any regularity of shape. The vesicle (6) therefore in the
last figure of the diagram is drawn as "doubtful".

If the hypotheses here suggested as to the origin of the coxal
glands are correct it would seem that the parent stock of the soli-
fugae must have been different from that of the other Arachnids
since they appear to have descended from a branch of the original
stock in which the salivary glands had become paramount, whereas
in the parent stock which gave rise to the other Arachnids, the
salivary gland of the second appendage had retrograded, and the
nephridia situated on the third to the sixth appendage had become
the most important; developing along lines similar to those of the
sixth and seventh appendages in Peripatus.

So far as I am aware the new observations are:

Scorpions. Blood supply to saccule from artery of appendage V.
Lymphatic organ wanting in Buthids but present in the other families.

Pedipalps. Saccule and collecting tubule opening into laby-
rinth. Columnar cells in labyrinth of Tarantula confined to the
middle division.

Spiders. T h e r a p h o s i d s. Two saccules, each with its col-
lecting tubule opening into the labyrinth, and two outlets.

Araneae verae. Saccule and its communication with the
disappearing labyrinth.

Solifugae. Modified saccule with pump and collecting tubule.
Outlet on appendage II with squirt and projecting sheath.

Peripatus. Modified (secretory) portion of the labyrinth of
the typical nephridium (?).

Modified nephridia of legs six to twelve in P. jamaicensis.

Homology of the salivary gland in Peripatus with the coxal
gland of the solifugae.

Homology of the coxal glands of the other Arachnids with the
large nephridia of Peripatus on legs four and five.

Coxal glands of the Arachnids.

265

Literature.

Beenaed, H. M., The comparative morphology of the
Galeodidae, in : Trans. Linn. Soc. London, Vol. 6,
1896, p. 377.

Beetkau, Ph., TJeber den Verdauungsapparat der Spinnen,
in: Arch, mikrosk. Anat., Vol. 24, 1885.

BoENEK, C, Beitrage zur Morphologic der Arthropoden.
I. Ein Beitrag zur Kenntnis der Pedipalpen, in:
Zoologica, Vol. 17, 1904, p. 95.

BouviEE, E. L., Monographie des Onychophores, in:
Ann. Sc. nat. (9), ZooL, Vol. 2, 1905, Vol. 5, 1907.

Beauee, a., Beitrage zur Kenntnis der Entwicklungs-

geschichte des Skorpions, in : Z. wiss. Zool., Vol. 57,

1894, Vol. 59, 1895.
Beuntz, L., Contribution a I'etude de I'excretion chez

les Arthropodes, in: Arch. Biol., Vol. 20, 1904.
GOUGH, L. H., The development of Admetus pumilio

Koch, in: Quart. Journ. microsc. Sc, Vol. 45, 1902.
V. Kennel, J., Entwicklungsgeschichte von Peripatus

Edwardsii Blanch, und Peripatus torquatus n. sp.,

in: Arb. zool.-zoot. Inst. "Wiirzburg, Vol. 7, 1884,

Vol. 8, 1886.
KOWALEVSKY, A., Una nouvelle glande lymphatique chez

le scorpion d'Europe, in : Mem. Acad. Sc. St. Peters-

bourg (8), Vol. 5, 1897.
Laueie, M., On the morphology of the Pedipalpi, in:

Journ. Linn. Soc. London, Zool., Vol. 25, 1894,

p. 29.
MaeCHAL, p.. La glande coxale du Scorpion, in: OR.

Acad. Sc. Paris, Vol. 115, 1892, p. 191.

Solifugae.

Spiders.

Pedipalps.
Peripatus.

Scorpions.

Arachnids.

Pedipalps.

Peripatus.
Scorpions.

Pedipalps.

Scorpions.
18*

255 B. H. Buxton, j

i

Patten, "W. and A. P. Hazen, The development of the j

coxal gland, branchial cartilages and genital ducts of j

Limulus polyphemus, in: Journ. Morphol., Vol. 16, j

1900. Limulus. \
Pelseneek, p., On the coxal glands of My gale, in: Proc. I

zool. Soc. London, 1885. Theraphosids. ;

Peeetaslawzewa, S., Developpement embryonnaire des \

Phrynes, in: Ann. So. nat. (8), Zool,, Vol. 13, i

1901, p. 232. Pedipalps. ]
Schimkewitsch, L. und W., TJeber die Entwicklung von i

Telyphonus caudatus (L.), in: Z. wiss. Zool., Vol. 81, '

1906. Tfielyphonus. |

— , Ein Beitrag zur Entwicklungsgeschichte der Tetra- |

pneumones, in : Bull. Acad. Sc. St. Petersbourg (6), j

Vol, 5, 1911. Theraphosids. I

Sedgwick, A., The development of the Cape species of ^

Peripatus, pt, 4, in: Quart. Journ. microsc. Sc. ;

(N. S.), Vol. 28, 1888, p. 387. Feripatus. .\

Sturany, E.,, Die Coxaldriisen der Arachnoideen, in: \

Arb, zool, Inst, Wien, Vol. 9, 1891, Arachnids. \

Coxal glands of the Arachnids. 267

Explanation of Plates.

Part I. Pulmouate Arachnids.

C Coeca of intestinal tract ON Ganglion

CA Coxal Artery LET External tubule of labyrinth

GL Coxal labyrinth LIT Internal tubule of labyrinth

CSL Central lumen of saccule LO Lymphatic organ

CT Collecting tubule M Muscle

D Diaphragm jV Nerve

END Endosternite Outlet

ET Exit tubule S Saccule

EXT External surface V Vesicle

G Genital duct VS Ventral sinus

GL Glomus VTL Ventral tubule

GLR Glomeruli

The photographs are so arranged that on reading the lettering , the
dorsal side of the specimen is uppermost in sagittal and transverse sections,
and the anterior end is uppermost in frontal sections.

Scorpions.

Plate 1.

Fig. 1. Vejovis flavies. Sagittal. 100:1.

Coxal gland showing the saccule surrounded by the labyrinth
coils. The coxal artery (CA) enters the saccule (S), and on
the right the lumen of the saccule passes into the labyrinth
tubule (CL). Below on the left is appendage V with its nerve
(N) and artery (AV).

From AV runs the coxal artery (GA) to the saccule.

The diaphragm (D) runs vertically across the field just posterior
to the coxal gland. On the right of D is the abdomen with its
intestinal coeca (C) filled with globules of nutrient matter. The
lymphatic organ lies more internally and does not appear in this
section.

268 B. H. Buxton,

rig. la. Buthus austrcdis. Immature. Sagittal. 100 : 1.

A view of the coxal gland corresponding to that of 1. The
difference in the size of nuclei and cells in the two different

species is noteworthy.

Plate 2.

Fig. lb. Vejovis flavies. Sagittal. 300 : 1.

Coxal gland. Passage of saccule into labyrinth.
A part of photograph 1, more highly magnified.

Fig. Ic. Bedims occitanus. Adult. 200 : 1.

Coxal gland. Passage of saccule into labyrinth. The striated
part of the labyrinth cells is stained black with iron hematoxylin.

Plate 3.

Fig. 2. Buthus occitanus. Young with mother. Sagittal. 300 : 1.

Vesicular part of coxal gland with exit tubule {ET) and

outlet (0), {EXT) is the exterior between appendages V and VI.

Fig. 3. Centruriis margaritatus. Egg just before birth. Frontal.
225 : 1.

Coxal gland with dip of labyrinth coils almost to surface (EXT)
on appendage VI. The saccule does not appear in this section.

Fig. 4. Buthus occitanus. Young with mother. Sagittal. 250 : 1.

Shows the whole course of the coxal artery (CA) from the
artery of appendage V to the saccule, where it breaks up into
the glomus (GL).

(0 in brackets) indicates the position of the outlet.

Plate 4.

Fig. 5. Buthus occitamis. Adult. 200 : 1.

The glomus {GL) formed at the entrance of the coxal artery
into the saccule (S). Compare with {GL) in the young scorpion,
photograph 4.
Fig. 5a. Vejovis flavies. Sagittal. 250 : 1.

A part of photograph 1 more highly magnified , showing the
coxal artery {GA) entering the saccule {S) and breaking up
into the glomus (GrL).
Fig. 5b. Buthus occitanus. Adult. 300 : 1 .

From the same specimen as photograph 5 to show the striation
of the labyrinth cells under a higher magnification.

Plate 5.

Fig. 6. Vejovis flavies. Sagittal. 100 : 1.

The lymphatic organ {LO) lying in the abdomen and opening
through the diaphragm (D) into the capsule of the coxal gland.

Coxal glands of the Arachnids. 269

Below the lymphatic organ is the large ventral sinus (VS).
The saccule lies more externally and is not visible.

Fig. 6a. Diplocentrus scaber (Scorpionid). Sagittal. 100 : 1.

Lymphatic organ and coxal labyrinth as in 6. The ventral
sinus is only partially seen.

Plate 6.

Fig. 6b. Euscorpms italiciis (Chactid). Sagittal. 100 : 1.

Lymphatic organ as in 6. In Euscorpius the lymphatic organ

is extensive and somewhat coiled upon itself. One of the large

binucleated nephrocytes is clearly seen in the ventral sinus. {BR)

is a part of the suboesophageal ganglion.

Fig. 6c. Scorpio inaurus (Scorpionid). Young with mother. 100 : 1.

Lymphatic organ and ventral sinus.

Pedipalps.

Plate 7.

Fig. 7. Taraniula palmata. Sagittal. 100 : 1.

Coxal gland. General view showing the saccule (5), the three
divisions of the labyrinth {CLa, CLm, CLp), and the straight
return ventral tubule (VTL) leading to the exit tubule (ET)
and outlet on appendage III.

Fig. 7a. Tarantula •pcihtaia. Sagittal. 100:1.

Coxal gland. Similar view to that of 7, taken from another
specimen. The central lumen of the saccule (CSL) narrowing
above to form the collecting tubule, is better shown than in 7.
C. in, C. IV, C. V, are the coeca corresponding to the appendages

below.

Plate 8.

Fig. 8. Tarantula palmata. Sagittal. 300 : 1.

The exit tubule, more highly magnified.
Fig. 9. Tarantula palmata. Transverse. 250 : 1.

The saccule with the inhanging glomeruli and central lumen
{CSL) with the collecting tubule {GT) leading into the
labyrinth {CL).

Below, on the left of ^the saccule, is the beginning of the exit
tubule. The glomeruli can be better distinguished in 9b. The
small spherical nuclei belong to the cells lining the lumina of
the saccule.

270 S- H. Buxton,

Plate 9.

Fig. 9a. Tarantula jyalmata. Transverse. 250 : 1.

Similar view to that of 9, but taken from another specimen.
Fig. 9b. Tarantula fuscimana. Transverse. 225 : 1.

Similar view to those of 9, and 9a. The cells lining the
collecting tubule are peculiar in that the nuclei lie on the free
border of the tall cylindrical cells.

This peculiarity is seen best in 9b. A part of the exit tubule
{ET) can be seen below the saccule.

Plate 10.

Fig. 9c. Tarantula fuscimana. Transverse. 75 : 1.

Compound of two photographs taken from the same specimen
eight sections apart, just posterior to appendage III.

The anterior part of the coxal gland with saccule, collecting
tubule, exit tubule and anterior division of the labyrinth lies on
either side of the enormously developed brain, pierced by the
narrow oesophagus.
Fig. 10. Tarantula palmata. Transverse. 400 : 1.

A transverse section across the middle division of the labyrinth
to show the cylindrical, slightly basophil cells, and the blood
channels running between the tubules.
Fig. 11. Tarantula palmata. Transverse. 100:1.

A transverse section taken near the middle of the coxal gland
to show the relation of the labyrinth {CL) to the ventral sinus
{VS) and the straight return ventral tubule (VTL). The
ventral sinus lies close to the endosternite, internal to the ventral
tubule. From the sinus blood channels run between the coils of
the labyrinth.

Plate 11.

Fig. 12. Tarantula fuscimana. Sagittal. 100 : 1.

The coxal labyrinth with its three divisions lying dorsal to
the large ventral sinus (VS). Below the sinus is the endo-
sternite (END). The basophil cylindrical cells of CLm can be
clearly distinguished from those of CLp and CLa.
Fig. 12a. Tarantula palmata. Sagittal. 100 : 1.

Similar view to that shown in 11.

The cells of CLm are clearly columnar but not so basophil
as is usual. Compare with 7a "from the same specimen. 1 1 and
11a are taken from sections lying more internally than 7 and 7a.
The saccule and ventral tubule lie externally and are not seen.

Coxal glands of the Arachuids. 271

Theraphosid spiders.

Plate 12.

Fig. 13. Hapalopus j^entcdoris. Immature. Frontal, 30 : 1.

The coxal labyrinth of one side, lying along the long axis of
the cephalothorax from appendage III, to the posterior aspect of
appendage VI. Section at about the middle of the cephalothorax.
On the right {CL') is a small part of the coxal gland of the
opposite side.

Fig. 13a. Hapalopus pentaloiis. Immature. Frontal. 30:1.

A section from the same specimen as 13, taken more ventrally.
The coxal labyrinth is now broken up into four parts, each of
which represents a dij) on the posterior aspect of an appendage.
(C) are the coeca of each appendage, and between the coeca and
the coxal labyrinth of appendages III and V, the saccules can be
seen, though not very clearly on account of the low magnification.

On the right of the endosternite (END) are (OE) the
position of the oesophagus (ST), a dilatation between the oeso-
phagus and sucking stomach (SCIt). The section is folded over
at (SC/i) so that the lumen of the sucking stomach is obscured,
but its lateral muscles and their relation with the endosternite
(END) can be seen.

Plate 13.

Fig. 14. Avicularia. Cocoon. Transverse. 250 : 1.

Showing the dip (DVI) of the coxal labyrinth almost to the
external surface (EXT) on appendage VI. There is however
no actual outlet. (CVl) is the coecum of appendage VI.
(BLS) is a blood sinus.

Fig. 15. Eurypelma vagans. Cocoon. Sagittal. 225 : 1.

Coxal gland on appendage III, showing the saccule (S), the
collecting tubule (CT) entering the labyrinth C'L) and finally
the exit tubule with its outlet behind appendage III. [The
glomeruli (GLR) can be distinguished from the lobular windings
of the saccule by the absence of the small spherical nuclei which
belong to the lining cells of the saccule.] At CL^ on the right
of the photograph the labyrinth extends posteriorly along the
length of the cephalothorax (compare photo 13).

Plate 14.

Fig. 15a. Enrypehna vagans. Young with mother. Sagittal. 250:1.

Coxal gland on appendage III. Similar view to that of 15.

A portion of the collecting tubule can be clearly distinguished,

but it is cut off from the main part of the saccule. At CL^

the labyrinth continues posteriorly as in 15.

272 B. H. Buxton,

rig. 15b. Hapalopus pentaloris. Immature. Frontal. 250 : 1.

Coxal gland on appendage III, A portion of the saccule witli
its central lumen (GSL), and collecting tubule (CT) leading
to the labyrinth (CL).

Plate 15.

Fig. 16. Ischnocolus. Transverse. 250 : 1.
Coxal gland on appendage V.

A portion of the saccule with its central lumen (CSL) and
the collecting tubule (CT) leading to the labyrinth (CL).
Fig. 16a. Hcqxdopus peiitaloris. Immature. Frontal. 250 : 1.

Coxal gland on appendage V. From the same specimen as
15b. The arrangement is precisely the same, but in 1 6a is on
appendage V, and in 15b on appendage III.

Plate 16.

Fig. 16b. Eurypelwa vagans. Young with mother. Frontal. 250:1.
Coxal gland on appendage V, with saccule, collecting tubule
and labyrinth all communicating.

Fig. 17. Avicularia. Adult. Sagittal. 100:1.
Exit tubule and outlet on appendage III.

Plate 17.

Fig. 17a. Avicularia. Cocoon. Sagittal. 250:1.

Exit tubule and outlet on appendage III. (CIH) Coecum of
appendage III.
Fig. 18. Avicularia. Cocoon. Transverse. 250 : 1.

Exit tubule and outlet on appendage V. (CV) Coecum of
appendage V.

Plate 18.

Fig. 18a. Hapalopus pentaloris. Immature. Transverse. 250:1.

Exit tubule on appendage V.
Fig. 18b. Eurypelma vagans. Cocoon. Sagittal. 250 : 1.

Exit tubule and outlet on appendage V.

Plate 19.

Fig. 18c. Theraphosid from Biskra. Young. Transverse. 250:1.
Exit tubule on appendage V, with portions of the labyrinth
(^CL) and saccule (»S'j. Just before the exit tubule is reached
there is a vesicular swelling (V).

Fig. 19. Isclinothele. Immature. Sagittal. 250:1.

Coxal gland. The saccule (S) changes at (CT) into the long
straight tubule of the labyrinth running posteriorly and corre-

Coxal glands of the Arachnids. 273

spending to the external tubule in the Dysderids. The internal
wall of the tubule is irregular in outline and the tubule seems
to have been formed by a breaking through of adjacent labyrinth
coils. The return tubule {RT) running forward to the exit lies
partly above and partly below, and corresponds to the internal
or return tubule of the Dysderids and hunting spiders. It also
is irregular in outline. The approximate position of the exit
tubule is indicated by {ET) in brackets.

Plate 20.

Fig. 20. Dugesiella hentzii. Adult. 400 : 1.

A portion of the coxal gland three days after inoculation
with carmine. The lining cells of the saccule are swollen, and
are filled with minute granules. The nuclei are stained with
hematoxylin and there is a counterstain of orange, but the granules
in the cells represent the red carmine granules. No such granules
can be seen in the labyrinth cells.

Araneae verae.

Group 1 .

Fig. 21. Loxosceles rufescens (Sicariid). Adult. Trans. 200 : 1.

Transverse section at about the middle of the coxal gland. LET
is the external tubule leading posteriorly from the saccule, and
LIT the internal tubule, close to the endosternite, leading an-
teriorly towards the exit tubule. The cell surfaces are worn down
to the nuclei but the striatiou shows no evidence of deterioration.

Plate 21.

Fig. 22. Dijsdera (sp.?) (Dysderid). Immature. Trans. 400:1.

Transverse section about the middle of the coxal gland.
Compare with 21.

Fig. 23. Dysdera crocata. Adult. Trans. 250: 1.

Transverse section across the coxal gland at the point where
the saccule (*S') passes into the external tubule {LET).

Fig. 24. Loxosceles rufescens. Adult. Trans 250 : 1.

Transverse section across the anterior part of the coxal gland,
showing the saccule and internal tubule (GLR) is a large
glomerulus with a mass of blood cells. The nuclei of the blood
cells are oval or flattened; those of the saccule are spherical.
The diagram explains better than a description how it comes
about that the internal tubule of the labyrinth appears to be
double.

274 B. H. Buxton,

Group 2.

Plate 22.

Fig. 25. Selenops mexicanns (Selenopid). Adult, Sagittal. 250 : 1.
Showing the saccule {S) passing into the external tubule
{LET) , the walls of which are thinned down , and the cells,
flattened so as to resemble those of the saccule. At Ch the
lining epithelium of the external tubule changes to that of the
internal tubule, the subsequent (internal) course of which is then
roughly indicated by the broken lines until it reaches LIT,
which then leads to ET, the beginning of the exit tubule. The
internal tubule retains some traces of striation, but in most
members of this group the striation is entirely lost.

Fig. 25a. Lycosa radiala. Adult. Sagittal. 100 : 1.

A view of the coxal gland comparable with that of photo 25.
There is an extension of the internal tubule {LIT) over appen-
dage III, which ends blindly a little further anteriorly. The
position of the exit tubule and outlet are indicated in brackets.

Plate 23.

Fig. 26. Lycosa radiata (Lycosid). Adult. Sagittal. 40 : 1.

The coxal labyrinth {CL) is here converted into a large
vesicle. The exit tubule and outlet are clearly shown, but the
saccule does not appear in this section.

{C) Coecum of appendage III. {BIS) Blood sinus.
Fig. 27. Tegenaria domestka (Agelinid). Adult. Sagittal. 100:1.
A view comparable to that of 26.

{CIIl) Coecum of appendage III. {NIII) Nerve of appen-
dage III.
Fig. 28. Zodariid Spider {S^.?). Sagittal. 250:1.

Showing the external tubule {LET) changing to the internal
{LIT). In this case the external tubule has become so much
fused with the saccule that it is impossible to distinguish between
them, except at the point of change.

Group 3.

Plate 24.

Fig. 29, 29a, 29b. Filistata hibernalis (Filistatid). Adult. Sagittal.

Three sagittal sections of the coxal gland, from the same
specimen, taken at different points.

The thin walled external and internal tubules of the labyrinth
have become so fused with the saccule that it is impossible to
distinguish between them. In these sections the glomeruli are

Coxal glands of the Arachnids. 275

filled with granular matter ; the lobes of the saccule appearing to
be empty.

Fig. 29. From a section through a more external part of the eland.
250:1. ^

The short vesicle (F) opens above into the saccule (S) with
its inhanging glomeruli {OLE). Below the exit tubule is
indicated by {ET in brackets).

Probably that part of the saccule into which the vesicle opens,
represents the remains of the internal tubule, the true saccule
being that part which lies on the right, immediately below the
coecum.

Fig. 29a. From a section a little more internal to 29. 100: 1.

A small portion of the vesicle can still be seen opening widely
into the saccule. The saccule itself now extends posteriorly for
a considerable distance, its posterior portion being probably
chiefly if not entirely composed of what were originally the
tubules of the labyrinth.

Plate 25.

Fig. 29b. From a section still more internal. 100: 1.

Showing the relation of the saccule to the endosternite.
Probably all of the saccule here visible represents the original
labyi-inth tubules, since the saccule itself in the Theraphosids and
in group 1 of the Araneae vei-ae does not extend so far inter-
nally.

{CUT, CIV) Coeca of appendages. Each coecum loops round
over and external to the coxal gland and enters the appendage.
(P) Loops of the poison gland which in Filistata is very extensive.
(BLS) Blood sinus from which glomeruli hang into the saccule.

Fig. 30. Lntrodedus madans (Theridiid). Adult. Sagittal. 250 : 1.
Views comparable with that of 29, but showing in addition
the exit tubule {ET) and outlet {()). The walls of the vesicle
are here surrounded by finely striated longitudinal muscles.

Plate 26.

Fig. 30a see Fig. 30.

Fig. 31. Epeira iyifoHum (Epeirid). Adult. Sagittal. 100:1.

May be compared with photograph 30. The vesicle is filled

with excretory products.

Fig. 32. Tkeridium tepidariorum (Theridiid). Adult. Sagittal. 100 : 1.

The details of the saccule are not well shown, but the vesicle,

opening widely into it and the exit tubule are clearly marked.

276 B. H. Buxton, *.

Part II. Tracheate Arachnids and Peripatus.

B Bulge into saccule MS Muscles of saccule

C Coeca of intestinal tract N Nerve

CL Coxal labyrinth NM Modified nephridium

CSL Central lumen of saccule NZ Nozzle

CT Collecting tubule Outlet

DSL Diverticulum of labyrinth sac i? Rostrum

EP Epidermis S Saccule

ET Exit tubule SAL Salivary gland

FO Final outlet SL Labyrinth sac

GN Ganglion SIG Slime gland

H Heart SO Side outlet

INT Intestine TR Trachea

KS Sucking chamber TRS Tracheal stigmata

LM Modified part of labyrinth V Vesicle

LR Sheath VNC Ventral nerve cord

31 Muscles

Solifugae.

Plate 27.

Fig, 33. Paragaleodes barbarus. Oblique. 250 : 1.

Showing the finely striated muscle fibres (MS) attached to
the periphery of the saccule {S).

Fig. 34. Paragaleodes scalar is. Oblique. 225 : 1.

Saccule with central lumen (CSL) and muscle fibres radiating
out from the periphery.

Plate 28.

Fig. 34a. Soljmga flaveseens. Sagittal. 250 : 1.

Saccule with central lumen and radiating muscle fibres, much
finer than the ordinary somatic muscles (M).

The tissue of the saccule is more compact than in 34, and the
saccule is more closely surrounded by somatic muscles.

Fig. 34b. Galeodes arabs. Transverse. 250:1.

Saccule with lumen. The tissues are more compact than in 34,
or 34a, and the short muscles radiating from the saccule are

arranged in bundles.

Plate 29.

Fig. 35. Ammotrecha (Ale. sp. M.). Transverse. 200: 1.

Showing the whole course of the collecting tubule (CT) from
the saccule (S) to the labyrinth sac (SL).

Coxal glands of the Arachnids. 277

Fig. 36. Daesina (sp.?). Transverse. 300:1.

To show the abrupt change from the labyrinth sac (SL) with
its columnar secretory epithelium, to the excretory cells of the
labyrinth tubule with striated border on which the minute
nuclei rest.

Fig. 36a. Paragakodes scalaris. Transverse. 300:1.

Change from labyrinth sac to striated tubules in another
specimen.

Plate 30.

Fig. 37. Paragakodes scalaris. Frontal. 100:1.

Labyrinth sac {SL) to show the pouching. The labyrinth
sac is lined with tall columnar epithelium, but there are no
products of secretion in the lumen.
Fig. 37a. Paragakodes harharus. Frontal. 100 : 1.

Labyrinth sac {SL) to show the pouching. The lumen of
the sac is filled with products of secretion, and the cells are
worn down almost to the nuclei. At {CT) is the entrance of
the collecting tubule.

Plate 31.

Fig. 38. Paragakodes harharus. Oblique. 250 : 1.

Exit tubule [ET) , with muscular walls, running anteriorly
just below the integument of appendage II, and leading to the
nozzle {NZ), which opens into {LP) the commencement of the
sheath. It seems as if the secretory products had just been
squirted out; some traces {Sp.?) still remaining in the expanded
orifice of the nozzle.

Fig. 38a. Paragakodes harharus. Sagittal. 250:1.

View similar to that of 38, in another specimen. The nozzle
is here cut through a little to one side of its orifice.

Plate 32.

Fig. 38b. Solpiiga flavescens. Frontal. 250 : 1.

View similar to that of 38 and 38a, but showing a dilation
the exit tubule.

Fig. 39. Oakodes {? Oihoes). Frontal. 100:1.

(7?) is near the opening of the rostrum to the exterior.
{KS) is the sucking chamber of the rostrum, passing posteriorly
into the oesophagus which lies between the two lobes of the
brain {BR). Above, on the right, the position of the nozzle
(V;t, in brackets) is indicated, anterior to which is the sheath {LP).

278 S- H- Buxton,

In this species the sheath is converted into a definite tubule
which runs a little further inwards than is shown in the photograph ;
the final outlet being approximately in the position indicated by
{FO in brackets) but the tubule does not project beyond the
appendage as in 40 and 40a. It can be readily seen that {LR)
is directed towards the opening of the rostrum.

Plate 33.

Fig. 40 and 40a. SoI^Juga flavescens. Sagittal. 250 : 1.

In 40 the exit tubule and nozzle are seen below, and above is
a part of the sheath (LR). 40a is taken from the same specimen
two sections further on. The position of the nozzle is indicated
by (NZ) in brackets, above which a longitudinal section of the
sheath can be seen, at first running along the appendage, and
then projecting beyond it from the point where the appendage
turns outwards, i. e. to the left in the photographs. The
approximate position of the final outlet is indicated by (FO) in
brackets, and that of the side outlet by ('S'O) in brackets.

Plate 34.

Fig. 41, Paragaleodes barharus. Frontal. 100 : 1.

Main part of the coxal gland. Above (anteriorly) is the
saccule (S) with its central lumen (GSL) passing into the
collecting tubule (CT), surrounded by pouches of the labyrinth
sac (SL). Below can be seen the abrupt change from the
labyrinth sac to the excretory tubule (CL) with its striated
cells. The course of the collecting tubule can be followed through
several sections to its entrance into the labyrinth sac, shown in
photo 37a, taken from the same specimen.

Fig. 41a. Paragaleodes barbarus. Frontal. 250 : 1.

A i^art of the same field as 41, more highly magnified, to show
the collecting tubule and its surroundings more in detail.

(B in brackets) corresponds to the bulge on the other side.
See photo 42 (B).

Plate 35.

Fig. 42. Paragaleodes barbarus. Frontal. 100:1.

Similar view to that of 41 taken from the other side of the
same specimen. The collecting tubule is closed by a bulge (B)
from the labyrinth sac, which protrudes also into the lumen of
the saccule. Just below the bulge is the change from the laby-
rinth sac to the excretory tubule of the labyrinth. By comparing
42 with 41 it can be seen how very closely the parts correspond
on either side.

Coxal glands of the Arachnids. 279

Fig. 42a. Paragaleodes harharus. Frontal. 250:1.

Taken from the same specimen as 42, two sections further on,
showing the bulge, more highly magnified, protruding into the
lumen of the saccule.

Plate 36.

Fig. 43. Solpuga flavescens. Transverse. 100:1.

Taken near the posterior end of the labyrinth coils, about
opposite appendage V, showing the complicated coils of the
tubules, some of which belong to the right side and some to the
left side, C is the coecum which runs from the intestine (AV7)
into appendage V {H) Heart. (.V) Ventral nerves.

Fig. 44. Eremobates (Ale. sp. M.). Transverse. 100:1.

E is the rostrum with its sucking chamber (KS), on the left
of which is the coxal joint of appendage II. On the dorsal
aspect of appendage II is the sheath (LIi) , cut transversely
just anterior to the nozzle, with its side outlet (SO) turned
away from the rostrum.

Just in front of this point the sheath closes up again, and
passes anteriorly as in photographs 40 and 40a,

/ is the chelicera which works above the coxa of appendage II.

Plate 37.

Fig. 45. Galeodes (? Othoes). Frontal. 250 : 1 .

A section showing the collecting tubule (CT) which is here
lined with columnar epithelium resembling that of the labyrinth
sac, so that the collecting tubule appears lost as a separate entity.
Its position however, between two pouches of the labyrinth sac
is the same as usual. In this specimen the columnar cells lining
the labyrinth sac are much more regular than is usual. In the
excretory part of the labyrinth (CL) , the columnar character of
the cells is maintained, but the nuclei are raised and rest upon
the striated base.

Fig. 46. A compound of photos 40 and 42. Frontal. 40 : 1.

The photographs are taken 12 sections apart. Each section
12 ^ thick. The two sections taken together show the relations of
the coxal glands on either side to each other and to the surrounding
tissues. On one side is seen the open collecting tubule, but on
the other side the tubule is clased by a bulge from the labyrinth
sac. For the finer details see photos 41, 41a, 42, 42a.

Zool. Jahi'b., Supplement XIV. 19

280 B. H. Buxton,

Plate 38.

Fig. 47. Peripatus nicaraguensis. Adult. Frontal, 250 : 1.

Typical nephridium near posterior end of body. The long
collecting tubule {CT) with ciliated cells leading out of the
saccule (S). The labyrinth {CL) changing to the modified part
(L3I) is seen on the right of the collecting tubule.

Fig. 47a. Peri]Kitiis jtdiformis. Adult. Transverse. 250 : 1.

Typical nephridium near anterior end of body with saccule,
collecting tubule, labyrinth changing to modified part, and the
vesicle (F). Above on the right is the labyrinth sac of the
salivary gland (SL). (EP) Epidermis. (GN) Ganglion.

Plate 39.

Fig. 47b. PerijMtus nicaraguensis. Adult. Transverse. 250 : 1.

Typical nephridium. The basophil granules in the cells of the
modified part of the labyrinth are clearly shown.

Fig. 47c and 47d. Peripahis jidiformis. Sagittal. 400:1.

Showing the change from the labyrinth to its modified part
under higher magnification.

Diagram 3 may be consulted in connection with photographs
47 to 47b, in order to understand how (CT) and (L3I) lie
closely apposed without however having any direct connection
with each other.

Fig. 48, Peripahis jamaicensis. Adult. Sagittal. 100 : 1.

Salivary gland showing the saccule , collecting tubule and
diverticulum which joins the labyrinth sac a little further poste-
riorly. {VI^C) is the ventral nerve chain. (CLR) the stri-
ated pai't (reservoir) of the gland which runs posteriorly under
the nerve chain to the eighth leg. The basophil granules of
(DSL) are stained very dark with hematoxylin , without reso-
lution of the individual granules.

Plate 40.

Fig. 49. Peripahis jidiformis. Transverse. 250 : 1.

Modified small nephridium of appendage III. (First leg),
which has lost all its striated, excretory cells. The whole of the
nephridium is shown (NMTII). Above the modified nephridium
is a part of the salivary gland, showing the saccule, diverticulum
(DSL) with basophil granules and labyrinth sac (SL) with
columnar epithelium, cut transversely. (CLR) is a part of the
reservoir (Bouvier) of the salivary gland.

Coxal glands of the Arachnids. 281

Fig. 50. PeripaUis nicaraguensit<. Sagittal. 250:1.

Salivary gland showing the saccule (S), the collecting tubule
{CT) leading into the diverticulum {DSL) of the labyrinth
sac. The diverticulum opens into the main part of the labyrinth
sac a little further back. (See photograph 51.) The lining cells
of the diverticulum are filled with coarse basophil granules (Com-
pare with those of photo 47b) and in the labyrinth sac itself
{SL) the nuclei lie at the base of the tall columnar cells con-
taining globules of secretion, but the lumen of the sac is free
from secretion and the cells are intact. Compare with photo-
graph 37 of Paragakodes. {CLE) is a part of the reservoir
which runs posteriorly as far as the third leg.

Plate 41.

Fig. 50a. Peripatus jamaicensis. Adult. Sagittal. 250 : 1.

Salivary gland. The specimen had simply been preserved in
forraol so that the tissues are somewhat swollen and distorted,
but the relations of the various parts can be clearly seen and
compared with 50 and 48.

The dark mass {SIG) is the slime gland of the oral papilla filled
with mucinous secretion.

Fig. 50b. Peripatus nicaraguensis. Embryo. Sagittal. 250:1.

Salivary gland of developing embryo in the uterus of the
specimen photograph 50. {SIG) is the slime gland.

Diagram F may be consulted in connection with photographs
50 to 50b.

Plate 42.

Fig. 51. Peripatus nicaraguensis. Adult. Sagittal. 100:1.

Salivary gland showing the passage of the diverticulum {DSL)
into the main body of the labyrinth sac of the salivai-y gland {SL).
Taken from the same specimen as photograph 50 a few sections
further on. The basophil granules of the diverticulum suddenly
cease where it enters the main sac. Along the dorsal margin of
the main eac the secretion appears to be beginning to pour out
into the lumen.
Fig. 52. Peripatus juliformis. Adult. Sagittal. 100 : 1.

Salivary gland. Labyrinth sac in sagittal section to show the
incipient pouching and the lumen filled with products of secretion.

Compare with photograph 37a of Paragaieodes.

Plate 43.

Fig. 53. Peripatus juliformis. Adult. Transverse. 250:1.

Salivary gland. Change from the secretory labyrinth sac {SL)
to the striated part of the gland. At {CLP) is the lieginuing
of the reservoir. {CL) runs anteriorly to the outlet.

19*

232 B. H. Buxton, Coxal glands of the Arachnids.

Fig. 54, 54a, 54b. Peripaius nicaraguensis. 400 : 1.

All takeu from the same specimen to show the similarity of
the basophil (hematoxylin) granules in the secretory cells.
Fif. 54. Diverticulum of salivary gland. DSL of previous photo-
graphs.

Fig. 54a. Modified part of typical nephridium. LM of previous
photographs.

Fig. 54b, Small modified nephridium of appendage 31.

The collecting tubule passing directly into a short secretory
tubule, corresponding to the part LM of the typical nephridium.
The short secretory tubule passes directly into the exit tubule.
Compare with photograph 49 (NM).

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