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Vor. II, No. 3. 


ON CA LIFORNIA EKUDRILID A. 


By GUSTAV EISEN. 


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SAN FRANCISCO, CAL., 
pas FEBRUARY, 1894. 


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ON CALIFORNIA EUDRILIDZ. 
BY GUSTAV EISEN. 


The California Oligochiets, to be described below, are all related to the well- 
known genera Microscolex, Pontodrilus, Plutellus, ete., referred by Benham, Rosa 
and others to the general family of Eudrilidee, and by Beddard to his family Crypto- 
drilide. I will not here discuss the relative merits of the respective families, as the 
number of species belonging to them is hardly sufficient to enable us to as yet come to 
any definite conclusion. For the present, at least, I retain these genera in the family 
of Eudrilidee. But I must concede that this family, as defined by the above inyesti- 
gators, contains two distinct main types—one represented by Eudrilus and the other 
by Microseolex—and a separation in families should be made in reference to them, 
though I am not prepared to make the separation as wide as Beddard has done. Our 
California species ally themselves to the Microscolex group, and especially to Micro- 
scolex and Plutellus. A species of Pontodrilus was found too late to be included. I 
believe this family may be conveniently divided into several subfamilies based upon 
the openings of the spermducts, prostates, penial sacs, as well as upon the location of 
the nephridio-pores. Our California forms may thus be grouped as follows: 


PONTODRILINI. 


The eight seta are separate though in four couples on every somite. 
Clitellum complete. 
Male pore in somite xviii. 
* Nephridia—mega-nephridia only—commence posteriorly to somite xiii; the pores are in line with sets 2’, 
not alternating in position. 
The prostate receives the spermduct previous to its entering the male pore. 
No gizzard, no typhlosole, no subneural yessel. Ponropritus E. P., 1881. 
PuHoropritus Girard, 1887. 
MICROSCOLECINI. 


Eight sets separate in four couples ; the set of the inner couples not always parallel. 
Clitellum complete. 
Male pore in somite xvii. 
The prostate receives the spermduct previous to its entering the male pore; or opens separately in the same 
somite, 
The nephridia commence anterior to somite y; the nephridia in line with one of the set 3 or4. The first 
few anterior pores may open in front of a different set from the posterior ones. 
No gizzard, no typhlosole, no subneural vessel. RuopopRitus Beddard, 1889. 
MicroscoLex Rosa, 1887. 
DeELTANTA Eisen, 1893. 


PLUTELLINI. 


Seta eight, in couples or equidistant. 

Clitellum perfect or imperfect. 

Male pores in xviii. 

The prostate either receives the spermduct or it opens separately, but in the same pore. 

The nephridio-pores show a systematic alternation in position either in front of or outside of the set. 

A gizzard; a small typhlosole may be present PLuvTELLUS Perrier, 1873. 
ARGILOPHILUS Hisen, 1893. 


bo 
bo 


CALIFORNIA ACADEMY OF SCIENCES. 


The genera related to Microscolex may be arranged as follows: 


I. Male pore in somite xviii. Nephridia commences posterior to somite xiii. 
PoNTODRILUS and PHOTODRILUS, etc. 
II. Male pore in somite xvii. Nephridia commences in the anterior somites ii to y. 
a. Spermducts open independently of the prostates. RwopopRiLvs. 


Sp.: minutus; nove-zelandiv. 
b. Spermducets open in the prostate, close to the body-wall. 


1, The sete of the inner couples (sets 1 and 2) in the vicinity of the clitellum are parallel with 
each other and do not converge toward any one of the genital pores. Nephridia commence 
in il to iv. MIcROSCOLEX. 

Sp.: modestus; algeriensis. 

2. The set of the inner couples (set 1 and 2) in the vicinity of the clitelluam converge toward 

one or both of the genital pores. Nephridia commence in somite ii to v. Deranta. 
Sp.: elegans; Troyeri; Benhami; Poultoni; dubia. 


DELTANIA. 
Deltania Eisen, Zoe, iv, 250, October, 1893. 


Prostomium encroaches on somite i. Eight sete in four couples ventral and 
lateral, beginning in somite 11. The sete of the inner couples in the genital region 
converging toward the male pore. The sete of the outer couples are further apart 
than those of the inner couples. There are a buceal cavity, pharynx, cesophagus and 
saeculated intestine, but no gizzard, thyphlosole, or cesophageal pouches. Clitel- 
lum, which is perfect, contains three large somites and two outside smaller ones, 
extending from xiii to xvii. No dorsal grooves. Testes in x and xi, free. Sperm-saes 
present, free. Spermatheca present or absent, variable in position and shape. 
Ovary one pair in xiii. Oviduet one pair in xiv. Two pair of ciliated rosettes in 
x and xi. Spermducts open in xvii together with a large prostate which is placed 
parallel to the segmental grooves. The spermducts join the muscular part of the 
prostate at the point where it enters the body-wall. The glandular part of the prostate 
consists of two layers of cells. Penial setee open in the same duct as the prostate. 

The dorsal vessel with three pair of lateral hearts in x, xi and xii. No sub- 
neural vessel. Only very few blood vessels on the nephridia. 

Nephridia are of two slightly different kinds. Those in the first few anterior 
somites, commencing in somite ii, open in front of and a little interior to the 4th seta, 
while those posterior to the former open in front of and a little lateral to the 3d seta. 
The nephridia begin generally in ii, rarelyin vy. All nephridia furnished with a large 
terminal bladder which in the posterior nephridia develop a ccecal prolongation. 

Small, transparent-glassy, more or less colorless worms with orange colored 
clitellum, and living in moist, especially sandy soil. 


Systematic position. Under the genus Deltania I group all species which 
would otherwise be referred to Microscolex Rosa, but which agree in having a deltoid 
arrangement of the ventral sete surrounding the generative and especially the male 
pores. With this character I believe a closer investigation of the species of Microscolex 
will join others, principally in regard to the nephridia which have not been sufficiently 
studied, probably on account of the scarcity of specimens for research. 

Among species which must be referred to the genus besides those described 


2. 


CALIFORNIA EUDRILID®. 23 


below are Microscolex dubius of Rosa and Eudrilus dubius of Fletcher. I will first 
refer to the latter. Fletcher’s description is sufficiently minute to allow us with cer- 
tainty to refer it to the genus, but the details are wanting to such an extent, that it is 
difficult to understand its further relationship. There are three points in the deserip- 
tion which are of special interest. 

1. Absence of spermathece. 

2. The beginning of the nephridia in vy. 

3. The junction of the spermduct and the prostate half-way between the 
glandular part and the body-wall. 

As to the first of these the spermathec may be really wanting or it may have 
a substitute similar to the one found in De/tania elegans as described below. At any 
rate this character brings the species Hudrilus dubius close to Deltania elegans as well 
as to Beddard’s Microscolex Poultont. 

The beginning of the nephridia in somite v brings 2. dubius close to Beddard’s 
species but separates it distinetly from Dedtania elegans in which the nephridia com- 
mence in ii, as will be shown below. 

The third character requires to be reaffirmed and described more in detail. 
The joining of the spermduct and the prostate is always of the utmost importance and 
interest and a mere general statement will not suffice for properly characterizing a 
species, especially when the group is little known. 

Rosa at first considered #. dubius to be identical with his Wicroscolex mod- 
estus, but a later investigation of new material convinced him of the distinct char- 
acter of the species and he then describes both as two different species of Microscolex. 
It must therefore be considered certain that the deltoid arrangement of the ventral 
setve does not oceur in JMicroscolex modestus. In regard to the respective species of 
E. dubius and M. dubius described by Rosa and Fletcher, I am not fully persuaded 
that both actually belong to the same species, and I believe that nothing short of an 
actual comparison of the specimens can decide if they do so. 

Beddard has at two different times described species of the genus Microscolex, 
but which differ from each other in several important points. J/icroscolex novw-zelan- 
dive resembles the old genus Rhododrilus in the independent opening of the spermduct. 
Instead of referring the above species to Microscolex, and merge Rhododrilus in the 
latter genus, I consider it more proper and convenient to retain Rhododrilus and re- 
fer M. nove-zelandic to it, as the independent opening of the spermduct appears to 
me of sufficient importance to be considered a generic character. Another species, 
Mieroscolex algeriensis, also described by Beddard, can, I believe, best be retained in 
the genus Microscolex, as it evidently possesses the set parallel throughout the 
ventral side of the clitellum. 


Microscolex Poultoni however is probably a true Deltania and Beddard’s excellent 
description leaves no important characters in doubt. 

In his description of MMicroscolex Poultoni, Beddard refers especially to the 
deltoid arrangement of the setee in the clitellial somites. He says: “ From segment 


24 CALIFORNIA ACADEMY OF SCIENCES. 


xix backward and from segment xiii forward, the distance between the two ventral 
sete of each side gradually increases.” 


Distribution and habitat. The species of the genus Deltania described below 
are undoubtedly natives of the Pacific Coast of North America, and more particularly 
of California. The species are found not only in the Golden Gate Park of San Fran- 
cisco, where they might have been introduced, but also in localities distant from 
gardens, and in which no cultivation has ever taken place. Thus I have species 
from Berkeley, Santa Rosa, Lake Chabot, Mount Diablo, ete. Moreover, since this 
paper was finished in MS8., I have found species of Deltania in localities so far from 
civilization that there can be no doubt as to the native habitat. I refer to species 
found in the high mountains of the Cape region (in the vicinity of Cape San Lueas) 
of Baja California, at an altitude of 4,000 feet, in almost inaccessible localities. A 
species has also been found in a gulch or river-bed at Ensenada, Baja California, 
thus proving the extensive territory over which the genus extends. The description 
of these species must be deferred to another paper. 

As regards Deltania dubia (? Budrilus dubius Fletcher) and Deltania Poultoni 
(Microscolex Poultoni, Beddard) it may be possible that the former, at least, is an 
introduction from abroad. The locality where the latter is found shows the genus to 
possess a wide neotropic extension. 

While it is thus certain that species of Deltania are natives of the new world 
and the Atlantic islands, and especially of the Pacific Coast of North America and 
Mexico, the distribution of Microscolex must yet remain undecided. Whether it 
was imported from the Argentine Republic to the Mediterranean region, or vice 
versa, must remain an open question, though the finding of two species in the Medi- 
terranean countries seems to point to the probability of that region being its real 
habitat. 


Species. Of the genus Deltania, California now possesses at least three species 
with the very great probability of a discovery of new species, as soon as a wider area may 
have been explored. All the species appear very sensitive to dryness and heat and 
disappear with the first warm and dry weather, hence the difficulty of finding them 
except at their proper season which appears to be limited to February to April in the 
vicinity of San Francisco. The species in Baja California were found in September 
and October. 


Deltania elegans. 
Figs. 1-20, 49-58. 
Deltania elegans Eisen, Zoe, iv, 248, October, 1893. 


Size about 2 to 4 inches by from ,'; to } inch wide. Septal glands very small, the 
posterior one the smallest. Spermatheci very pellucid, minute, and irregular in their 
position both as regards somite and the place in the somite, but generally opening be- 
tween vili and ix. Sperm-sacs comparatively small, deeply lobed, one pair in xi and 


bo 
Or 


CALIFORNIA EUDRILID-©. 


one in xii. Prostate is at the top helix-like folded. Penial papillee prominent, with 
two or more penial sete in each sac. 

The worm is very pale, glassy, semi-transparent with reddish-orange clitellum, 
and with the dorsal vessel showing prominently through the body-wall. It is very deli- 
cate, succumbs readily to heat and can only with difficulty be kept alive. It is the 
largest of the species belonging to this genus as far as known. 

Habitat.  Deltania elegans is so far only found around San Francisco, Califor- 
nia, and at Santa Rosa and Mount Diablo, especially under decaying manure in natura] 
hollows in the Golden Gate Park. It is common immediately north of Strawberry 
Hill in the sandy hollow where rain water collects and keeps the soil sufficiently moist. 
The worm is not found in the water, but at the water’s edge, the water, however, being 
only temporary during the rainy season. A few specimens also found in Berkeley 
along the creek. In the locality in Golden Gate Park the worms congregate in large 
masses, always in the sandy soil. At Santa Rosa and Mount Diablo I found only few 
specimens, the season being at my visit in May far advanced and the soil was drying 


_fast: Mature in April and May. 


DETAILED DESCRIPTION. 


Exterior characteristics. The worm yaries considerably in size, but averages 
about three inches in length, by a width of from two to three lines, tapering consid- 
erably towards the tail. The color is pale flesh, with a bluish cast, and quite trans- 
parent, glassy, with a yellowish clitellum. Altogether, the exterior appearance of the 
worm is one of great delicacy, greatly heightened by its semi-transparency. 

The cephalic prostomium is prominent, and divides the peristomium about ? 
of its width (fig. 5). This, the first somite, is wider than any of the following. 
Somites ii and ili are next in size, and about equal in width. Somites iv to xi are 
smaller, about equal in width, slightly decreasing backward. Somites xii and xiii are 
smaller than any of the other anterior somites, and of about the same respective width 
(fig. 2). No dorsal pores. 

The clitellum (figs. 2 and 15) commences with somite xiy, though xiii is gen- 
erally somewhat thickened. The clitellial somites xiv, xv and xvi are very wide, 
about as wide as somites 11 and iii, and of the same size. Somite xvii is much smaller. 
This somite carries the male pores and papille (fig. 15). These are situated in the 
posterior part of the somite, almost on the edge of the intersegmental groove (fig. 15). 
The papillee are slightly raised around the opening of the sac in the penial sete. The 
papillee are situate close together yery near the median line of the body, a short dis- 
tance only from the ventral ganglion. The papilla is oblong, sigmoid, with a pore 
for the sete at inner end. Between it and the intersegmental groove is seen the slit 
in which open the prostate and the spermduct. It is situated slightly to the outside 
of the penial papilla (fig. 14). 

The oviducal pores are in the anterior part of xiv, generally in a depression 
anterior to and more ventrally located than the inner couple of sete. Spermathecal 
pores are variable, not perceptible. The nephridio-pores are in front of the third sete, 


26 ; CALIFORNIA ACADEMY OF SCIENCES. 


while the three anterior pepto-nephridio-pores are in front of the fourth sete (fig 2). 
Clitellum is conspicious, occupying somites xiv to xvil, encroaching slightly on xiii 
(fig. 2 and 16). 

In viewing the caudal end of the live worm a number of irregular white spots 
are seen in mature specimens. These are rows of mature ova which agglomerate there 
sometimes in large quantities. How they finally find their way through the oviduct 
is not readily explained. 

Sete (figs. 2 and 4). As usual the sete begin in somite ii. All except 
those in the penial sacs, are sigmoid and arranged in couples of two, eight in each 
somite. The ventral sete are about 4 closer than the lateral ones. Thus if we con- 
sider the distance between the inner and outer couples to be 50, that between the 
setee of the outer couple is 30, that between the sete of the inner couple 22, that be- 
tween the inner couples 40. This being the distance in somite xxvii, where it may 
be said to be normal. In the region of the clitellum, and posterior to it, the inner 
sete are unequally distant in the respective somites. Thus we may consider the inner 
sete 1 and 2, to have the normal distance from each other in somites xii to xxvii. 
From these two somites the sete in the inner couples conyerge toward somite 
xviii in which the sete are about 4 as far apart as in xii and xxviii. In xvii the set 
of the inner couples are wanting. 

The innermost or row No. 1, forms a continuous line from one end of the body 
to the other, while the row No. 2, forms an angle with somites xvii and xyiii at the 
apex. The sete in rows 3 and 4 are parallel and normal. The seta 1 in xviii is pre- 
sent (pl. xii, fig. 4). 

This arrangement of the sete appears very constant, and is characteristic of 
the species, the details being somewhat different in the other species of the genus, 
while the general characteristics are the same. The normal sete (fig. 16) of the 
clitellum are not smaller than those of other parts of the body. The penial seti are, 
however, very much the largest. 

Penial sete (fig. 17). The two pairs of sacs containing the penial sete are situated 
in front of the spermidueal pore in somite xvii. They open immediately in front of 
that pore, in a slit, at either end of which is situated a pore, each pore being the 
outlet for the respective fork of the penial sac. The two sacs are connected at the 
upper margin as usual by arciform muscles. Each sac contains not less than two, and 
sometimes three or four sete, straight or slightly curved, but not sigmoid. The sete 
vary considerably as regards shape, but resemble each other in not being sculptured, 
and are only marked by rings. The penial sac reaches to the upper part of the mus- 
cular part of the prostate. 

INTERIOR CHARACTERS. 

The Septa begin between somites iv and y. Those between vii and viii, and 
following as far back as somite xiv, are slightly thickened, all, however, being of the 
ame general thickness, 

The body wall contains the usual layers. The hypodermis is slightly thinner 


i 


bo 
~I 


CALIFORNIA EUDRILID®. 


than the circular muscular layer, and this again is thinner than the longitudinal 
muscular layer. The clitellum is perfect, but somewhat thicker on the upper part. 
The glandular layer is very thick, being about ten times thicker than the two mus- 
cular layers combined. The longitudinal fibres are nowhere arranged in such 
feather-like bunches as in Lumbricus, but much more irregular, with a faint ten- 
dency to feather-like bunching. This is the rule both in the clitellum and elsewhere. 
This stratum of longitudinal fibres is, as usual, interrupted in four places by the setal 
grooves, though there are also fibres between the sete. In the center, between 
these grooves, the layer is thickest, from there gradually tapering towards the setie. 
The under side, or the ventral part, of the layer is somewhat thicker than the dorsal 
part. 

The clitellar glands are less regularly paired than in Lumbricus though 
the general arrangement is that of two or three rows of glandular cells together. The 
clitellum is very thick and developed all around the body. 

Alimentary canal (fig. 1). The bueeal eavity is eversible to a remarkable de- 
gree, so much so that it is often projected like a large sac or bladder, covering more 
or less perfectly the prostomium and part of the peristomium. Its walls are very 
thin and transparent. The pharynx commences with the prostomium and covers 
somites i, ii, ii, but is only dorsally developed. It is much and irregularly folded, the 
sinuses being sac-like and not parallel, the largest ones being in somites ii and iii. 
The muscles of the pharnyx are thickly covered with salivary glands. Superiorly 
these glands project along the under side of the muscular bands, which extend back- 
ward, thus forming three rows of parallel projections tapering from base to apex. The 
anterior of these salivary glandular masses is the largest, the third, or the most posterior 
one, the shortest (fig6). The posterior half of the salivary glandular mass forms one 
single projection equal to all the anterior ones together. This gland is connected with 
and partially rests on two muscular bands attached to the body-wall between somites 
vii and viii. 

(Hsophagus commences between iii and iv, extending backward to somite xvi, 
being slightly differentiated in xiv, xv, and part of xvi, as a tubular intestine (fig. 1). 
(Esophagus is much sacculated, first rising upward and forming a sigmoid plexus in 
somite vil, after which it lowers itself somewhat in viii and then extends gradually 
backward to the sacculated intestine, at the same time gradually diminishing in size. 
The tubular part in xv or between xv and xvi is the narrowest part of the cesophagus. 
The glandular epithelium of the cesophagus is very narrow in the anterior somites, or 
those in front of the clitellum, and the blood sinuses in them are narrow. In the 
anterior part of the clitellum the epithelial villi become greatly elongated with in- 
ereased blood supply, while in the central part of the clitellum these blood sinuses be- 
come very large, occupying the largest part of the epithelial lobes. The nuclei in those 
epethelial cells are everywhere round. 


The sacculated intestine, fig. 1, s. i., commences in xvi, is generally about four or 
five times wider than the tubular intestine. It does not inerease gradually, but at 


28 CALIFORNIA ACADEMY OF SCIENCES. 


once, in which it differs from the corresponding organ of’ the two other species de- 
scribed here. 


Septal glands (fig. 1). The septal glands of the esophagus are found paired in 
somites yi, vil, viii and ix. They are all comparatively small and very thin and 
flat. The posterior ones are smaller, but the difference between the glands is not 
so great as for instance in some species of Oenerodrilus, or as in Deltania Troyeri. 
They are more nearly of the same size, and they extend all around the cesophagus 
and are more or less divided in several separate lobes. We can, however, always 
distinguish one pair in each somite connected with the septum below the alimentary 
canal, extending upward with its free upper lobes, which do not connect. 


Nephridia (figs. 19 and 20). The nephridia commence in somite 11, and are 
found in all posterior somites. The three anterior nephridia—in ii, ii and iy—are 
slightly different as to size and outward form, and may be considered as a kind of 
pepto-nephridia. They open, also, differently from other nephridia, their pores 
being in front of and a little interior (nearer the ventral ganglion) to seta 4, while 
all other nephridia open in front of and interior to the third seta. The nephridio- 
pores of the clitellum are considerably larger than the pores in the other somites, and 
appear like large, transparent dises when viewed from the outside. All nephridia 
possess a vesicle or bladder next to the body-wall. A small collar of tubular cells 
surround the uephridio-pore. The vesicle is smaller in the anterior pepto-nephridia, 
gradually increasing in size backward, being largest in the post-clitellar nephridia, 
where it is several times larger than in the pepto-nephridia. In the three pairs of 
pepto-nephridia, the vesicles are almost circular, and of the same size in the three ne- 
phridia. That of the first common nephridium in somite v is of about the same size 
as the pepto-nephridial vesicles, but from this on the bladders increase gradually, but 
slowly, in size to the end of the clitellum. But in somite xviii and following to the 
end ‘of the body, the vesicles are much larger, about twice as large as those in the 
clitellum. Thus the nephridio-yesicles in the clitellum are about three to four times 
shorter than the tubular part, while the post-clitellar vesicles are half, or more than 
half, as long as the tubular part or duet, when ordinarily folded. In the pepto- 
nephridia, the vesicle is about five times shorter than the folded tube, and the tubular 
duct extends more backward than in the other nephridia—especially so in the first 
nephridium—encroaching on the next posterior somite, reaching diagonally across the 
somite, while all the other nephridia run parallel with the intersegmental grooves. 
This diagonal position of the nephridia is, however, not always constant, except in 
the most anterior nephridium. The vesicle in the posterior nephridia consists of two 
more or less distinct lobes, the posterior one (to the duct), which is more rounded 
and bladder-like, forming a caecum, and the anterior, which is elongated or deltoid. 
This difference is more pronounced in the posterior than in the anterior nephridia, 
most so in the nephridium in sonite xviii, which nephridium is generally the largest 
of all. From this somite the nephridia diminish somewhat in size, both anteriorly 
and posteriorly. The posterior margin of the vesicle is considerably lobed, and. in 


CALIFORNIA EUDRILIDE. 29 


general outline convex. The anterior margin is convex-deltoid, with the apex at the 
exterior pore. 

The single duct from the vesicle leaves the vesicle in a different manner in 
the respective nephridia. In the pepto-nephridia it ascends from the center of the 
vesicle; in the anteclitellar vesicles it leaves from the apex of the deltoid longer part 
of the vesicle, while in the postclitellar nephridia it leaves from the side of the vesicle 
below the apex. The inner structure of the nephridia corresponds almost exactly 
with that of the nephridium of Argilophilus, which genus, however, does not possess a 
vesicle. With this exception, the nephridia of the two genera might be considered as 
almost identical. The most characteristic feature is that the short single duct from 
the nephridio-stome after joining the nephridial body, does not enter it as a single tube, 
but as a spongy duct full of irregular and connecting ductules, which later on join 
into one larger branching canal, the ductules, or arms of which enclose the two reg- 
ular underneath-lying ducts. At the inner bend of the duct the ductules disappear and 
the returning lobe contains three main canals, one of which is ciliated. This arrange- 
ment reminds us greatly of the one observed in certain leeches and described by 
Bourne. ‘The vesicles consist of an outer muscular layer, which extends all around 
the bladder, and from it, along the duct through the clitellum or body-wall, to the 
exterior pore. Between it and the ccecal epithelium there is a continuous row of 
connecting chambers probably analogous with the ductules of the ducts. This epithe- 
lium is furnished with some few blood-vessels. The duct from the bladder to the 
nephridio-pore is not otherwise differentiated, the glandular cells of the clitellum joining 
directly on the muscular duct. Before reaching the pore but while in the body-wall, 
the duct is enlarged, forming a small pear-shaped urinary bladder, which again opens 
into a narrow duct surrounded by a row of long tubular cells, which open directly 
into the duct. These cells form a veritable collar, the upper cells lining the inner 
surface of the nephridio-pore (fig. 49). This in the clitellar nephridia. In the 
nephridia posterior to the clittellum, the long tube between the vesicle and the pore 
is entirely wanting, the bladder connecting directly with the collar of tubular cells 
(fig. 50). The tubules, or vacuols, in the vesicle collect into at least two tubes, which 
run downward between the muscular and glandular layers of the vesicle and appar- 
ently open on either side at the beginning of the collar at the nephridio-pore (figs. 49 
and 50). The secretion from this glandular layer of the bladder may be of such 
nature as to facilitate the ejection of coarse matter such as caleuli which are found 
often in enormous quantities in the vesicle or seen as just ejected through the pores. 
The single duct which leads from the bladder to the nephridial body proper or the 
folded canals, apparently does not connect directly with the ductules or vacuols of 
the bladder, but opens directly into the large central chamber of the bladder, which 
again connects directly with the collar of the nephridio-pore. For a more detailed 
account of the canals of the nephridia in these two genera see the description of next 
genus, Argilophilus. Below the nephridial collar in Dedtania elegans is found a large 
branching body, probably a ganglion. It sends out branches to the nephridial collar, 
though in my sections I have not seen their actual connection with the collar. In 


30 CALIFORNIA ACADEMY OF SCIENCES. 

fig. 50b., the section is made through the junction of the nephridio-vesicle and the 
collar, and the ganglion (n. g/.) below is small, not yet branched. In fig. 50c, the 
section is made close to the pore where the ganglion (. gi/.) is branched. These 
branches always extend towards the collar, never the other way. Both these sections 
from which the drawings were made are not quite in right angle to the longitudinal 
muscular layer, but slightly slanting. 


The nephridio-stome is rather large, but very transparent and quite difficult to 
detect. It is flat, rosette-like and placed half-way between setee 1 and 2. The duct 
descending from it is remarkably thin, straight and never winding. The nephridio- 
stome is normally situated, piercing the anterior septum as usual. There is no central 
cell and nucleus larger than the other, but several smaller cell nuclei are visible in 
the center below the large marginal cells, which are about 16 to 18 in number and as 
usual placed in a crescent. Between these cells and the real margin of the lip there 
is arow of smaller cells and nuclei, the latter being slightly lower than the inner 
large nuclei, but sufficiently situated in the same plane to be seen at the same time. 


Testes consist of two pairs of small bodies, as usually situated in somites x and 
x1, attached to the septum close to the body-wall in the anterior part of the somite. 
They are deeply lobed and the lobes are very narrow, parallel and generally three or 
four in number. The testes are free, not enclosed in the sperm-sacs. 


Sperm-sacs (figs. 6 and 6). There are two pair of sperm-sacs, one pair in 
somite Xi and one pair in xi, attached to the anterior septum in the somite, rather 
high up, but below the cesophagus. The four sperm-sacs are of about the same size, 
comparatively small, and deeply lobed very much like the posterior sperm-sac in 
Ocnerodrilus Beddardi, etc., none of the lobes being very much larger than the other. 
It must be remarked that the sperm-sacs are not situated in the same somites as the 
testes, somite x possessing testes, but no sperm-sacs, and somite xii possessing sperm- 
sacs, but no testes. The respective sperm-sacs are not connected with each other and 
there is no median sperm pouch. The trabecula or muscular divisions of the sperm-saes 
are numerous and the chambers enclosed by them are comparatively small. 


Spermducts and rosettes (figs. 7,8, 11). There are two pair of ciliated rosettes, 
one in xiand one in xii,as usual. They are very large and folded, deeply crumpled and 
very prominent, occupying about 5 of the somite. The funnels are wide, with a 
slight posterior swelling. The spermducts run backward and sideways, connect in 
xii, and continue to somite xvii, where they leave the body-wall, ascend slightly and 
connect into one single duct, which opens out jointly with the prostate, in the 
posterior part of the somite. The duct does not enter the prostate, but both open 
jointly in one slit, running parallel to the intersegmental furrow. The spermduct is 
throughout of the same size, only widening out just previous to entering the rosette. 
The lower descending part of the duct, between the spermiducal pore and the bend 
where it starts forward, is slightly narrower than the anterior part. 


CALIFORNIA EUDRILID®. 31 


Prostate (fig. 8). There is one pair of prostates of rather prominent size ex- 
tending parallel with the intersegmental groove reaching almost across the somite. 
The shape of the prostate resembles at the top somewhat a curved feather, the inner 
apex being helix-like, curving backwards. This form appears quite constant and while 
I found the length and the width of the prostate to vary, I never found one, which did 
not show the helix-like, convolution. The thickness of the prostate varies consider- 
ably. In some specimens it was almost twice as thick as in others, the increased 
thickness being caused by a gradual widening toward the inner apex. In some speci- 
mens the prostate was longer, more slender and its longitudinal sides almost parallel, 
but the convolution was generally always thickened. In most instances the convolu- 
tion could be considerably straightened out by a pushing with a needle, but it would 
when released assume its natural helix-like form. The spermduct connects with the 
muscular part of the prostate in the muscular layer of the body-wall. The sae con- 
taining the penial sete is situated immediately anterior to the prostate and somewhat 
closer to the ventral ganglion in line with the regular setze, but opens in the same pore 
with the prostate and spermduct. (Figs. 8, 51, 52, 53.) 

A cross section of the glandular part of the prostate shows that it is composed 
of two layers of cells, the outside one containing large cells of flask-like shape, the 
inner are narrower rectangular cells. The contents of both layers resemble each 
other greatly and are difficult to discern. Both layers of the prostate contain nu- 
merous blood vessels arranged like radii in a circle, penetrating both of the cellular 
layers. But the inner layer is seen to also possess a vascular system of its own with 
many smaller vessels similarly arranged. These vessels are generally wider at the 
periphery of the prostate and narrow toward the center, many if not all collecting in a 
network of capillaries spreading on the inner surface of the prostate (figs. 55 and 56). 
Otherwise these vessels do not anastomose. All these vessels are fed by a branch from 
the ventral vessel of the body, which divides on the prostate into two or three large 
branches which again fork toward the apex of the prostate in many smaller ones, as 
in Deltania Troyeri (fig. 45). 

This junction of the various male organs is affected in this manner. The two 
spermducets run jointly on the top of the inner longitudinal muscular layer of the 
body-wall. When reaching the lower or muscular part of the prostate they turn and 
run parallel to it. Immediately before reaching the place where the prostate enters 
the wall, the two spermduets fuse into one duct, the lumen of which then is wider 
than the adjoining part of the prostate. This duct joins the prostate in the iongitudi- 
nal muscular layer of the body-wall. After reaching the transverse muscular layer 

this duct joins the pore of the penial sete (figs. 51, 52, 53). 


Spermatheca (fig. 13). The spermathecze consist of very minute bodies, pear- 
shaped in outline, and of extremely delicate structure, without any differentiation of 
the wall in a muscular and glandular layer. In size, the spermatheca is not much, 
if any, larger than one-half the width of the somite, when contracted in alcohol. But 
the most peculiar feature of this organ is that it is variable in number and_ position. 


32 CALIFORNIA ACADEMY OF SCIENCES. . 


In one specimen I found only one spermatheea, situated in the center of the ventral 
line, between somites ix and x, but in x. Another specimen had two spermathece, 
one in ix /x, another on the left side, between x and xi. Another had two sperma- 
thecee betweeen x and xi, one on each side. Another had two spermathece between 
ix and x, one in center of median line, and one on the right side, always attached to 
the intersegmental wall. This arrangement and variability of the spermathece 
reminds us of the spermathecze in Microchieta, where the number varies on either 
side; but in other respects there is no similarity between the two. The exterior 
spermathecal pores are not conspicuous, and not perceptible when viewing the out- 
side of the body, mounted, for instance, in glycerine. As will be seen, the sperma- 
theea in this species differs very much from those in Deltania Troyeri and Benhami, 
in which two species this organ is constant, and furnished with two diverticula each. 
The spermatheeca in De/tania elegans resembles greatly in structure a sperm-sac, from 
which it only differs in size and in position. It reminds me greatly of the peculiar 
organ described by me in Qenerodrilus occidentalis, where, apparently, the posterior 
testes have become modified, and assumed the function of spermathecze, with a dis- 
tinet and ciliated duct perforating the body-wall. 


The spermatozoa are always found agglomerated in sphzerical masses in the 
spermatheca, hardly regular enough to be designated as spermatophores. The tails are 
long, either extending straight out, or arranged screw-like in the same direction 
around the sperm-ball. These balls vary greatly in size, some being twice as large as 
others, but they are always round and apparently globular (fig. 15). 


Ovary and Oviduct (fig. 9). As usual the ovary is found in xiii. It offers no 
great peculiarities. It is rather deeply lobed and very large. The oviduct opens in 
xiv, with its funnel in xii. The ovidueal funnel is very thick, substantial and round, 
with a circular and very regular outline. The figures (5 and 9) give correctly its 
outline, but the depressed folds have been too distinctly marked. There is no ovisac, 
and the ovary and oviduets are entirely free. 


Llood vessels. The dorsal vessel emits three pair of hearts in x, xi, xii, and the 
ventral vessel is forked between somites ix and x. The blood is yellowish-red, more 
decidedly yellow than red. There are but few blood vessels on the nephridio-tubes and 
none on the nephridio-vesicle. 


The ventral nerve-cord is considerably wider in the posterior part of the 
somites where it emits the customary pair of nerve fibers. In the anterior part 
where the single septal nerve pair is emitted, the nerve-cord is quite narrow. In 
Deltania Troyert the nerye-cord is quite uniform without any nodular enlargements, 
as wide at the anterior as at the posterior end of the somite. The brain is narrow, 
slightly curved and the posterior sinus shallow. It is situated in somite ii (fig. 6). 


CALIFORNIA EUDRILID®. D0 


Deltania Troyeri. 


Figs. 21 to 39. 


Deltania Troyeri Eisen, Zoe, iv, 251, October, 1893. 

Size about 1} inch by } line. Septal glands comparatively large, the one in 
vi the largest. One pair of large, opaque spermatheca, furnished with one pair of 
diverticula, which are about 5 or more longer than the spermatheca proper. Sperm- 
sacs in x and xi, not lobed. One developed seta in each sac of penial sete. Pros- 
tate is tubular, not helix-like, with the top either straight or bent at right angle, pro- 
jecting backward. The exterior penial papille not as prominent as in the preced- 
ing species. The inner couples of sete are further apart than in the following species. 


Habitat. This species was first brought to my attention by Professor Carlos 
Troyer, of San Francisco. who found it, together with the preceding species, in the 
Golden Gate Park, in San Francisco, immediately north of Strawberry Hill. It 
occurred there in sandy depressions, where the rain and drainage water had moist- 
ened the soil in March and April. As the soil dried up the worms disappeared. 
The worm is very scarce at any time, and not one specimen is found to every hun- 
dred of Deltania elegans. 


Euvterior characteristics. Eixteriorly this species is characterized at once from 
Deltania elegans by being very much smaller, as much so as an Enchytreeus is 
smaller than an average medium-sized Lumbricus. The length in the largest speci- 
mens is about two inches, when stretched to its full capacity, though the average ones 
hardly reach one inch. The width is less than one line at the clitellum and less than 
+ line at the tail end. The first somite is much longer than any of the following. 
The second somite is next in size, while all the others are smaller and of very much 
the same proportions as in Deltania elegans. ‘Thus iii, iv and y, are larger than the 
following, and those between v and xiii are smaller and of about the same size. 

The clitellum occupies the same somites as in Deltania elegans, or from xiv 
to xvii, with the two outside somites smaller than the central ones. The body tapers 
towards the tail end, the last somites being somewhat larger than the others and 
‘ather obtuse. 

The color is pale flesh, with a darker, yellowish clitellum. The whole body 
is very transparent, just as the former species, but much less so than in the following. 
It is a very tender worm indeed, and can only be brought home alive with great 
care, as the least increase of temperature is apt to kill it. In no instance did I sue- 
ceed in keeping it alive more than a couple of days. In this respect, however, all 
the species of the genus are yery much alike, and if there is any difference the larger 
species Is the most tender. 


Setw (figs. 21, 24 and 39). The general arrangement of the setz is similar to 
that of Deltania elegans, but the two inner setee are much less close together, in compari- 
son with the two outer ones, than in the latter species, but not as close by one-half as in 


34 CALIFORNIA ACADEMY OF SCIENCES. 


Deltania Benhani. In the genital region, the distance between the two inner setz 
diminishes toward the male pore, almost in the same way as in Deltania elegans, with, 
however, a slight but characteristic difference. The inner or first seta in xyiii is 
often, but not always, wanting, probably falling out when young, before its full devel- 
opment, as more frequently a rudimentary seta is seen in place of a fully developed 
one. 

The first and second sete in xix and xx are closer together than normally. but 
already in xxi the setee have regained their proper distance. Again, anterior to the 
male pore, the sete 1 and 2 in xi to xvi are closer than normally, those in xiv to xvi 
are equidistant, while those in x to xiv rapidly approach. If we thus compare with 
Deltania elegans, we find that the arrangement relatively in front and behind the 
male pore is reversed. While in Deltania elegans, the anterior sete quickly con- 
verge, the posterior ones approach slowly. In Deltania Troyeri, the opposite is the 
case. 

The shape of the sets in the two species is very similar. Compared again 
with the arrangement of the setee in Deltania Benhami, we find that in the present 
species the sete in the ventral couples, as well as those couples themselves, are 
much further apart than in Deltania Benhami. The deltoid arrangement, also, is 
different in the two species, of which the figures give a better idea than any lengthy 
description (figs. 24, 39, 40). 


The saes of penial sete (fig. 55) are found as usual in the vicinity of the male 
pore in xvil. There are seldom more than one seta in each sac. This seta is long, 
slender, almost straight, occupying the whole length of the sac. Now and then there 
is a rudimentary seta in the same sac, but neyer more than one deyeloped seta. In 
Deltania elegans there are three or four sete in each sac. 


Alimentary canal (figs. 26 and 27). The bueeal cavity extends superiorly to 
ii, inferiorly to y. The pharynx ends in y, and is much less developed than in 
Deltania elegans. The upper fold is, however, very large. There are one pair of 
long and narrow salivary glands in each of iii and iy, and one pair very large com- 
pact ones in iv. The cesophagus commences in y, and rises to a sigmoid plexus in 
vill. It is greatly contracted at the septa. In xv and xvi it narrows down to a tubu- 
lar intestine. 

The sacculated intestine commences in xvii, but attains its full width first in 
xix or xx. There is no gizzard, no ealciferous glands nor pouches of any kind at- 
tached to the alimentary canal. 


Nervous System (fig. 28). The characteristic feature of the nervous system is 
the even width of the ventral ganglion, the two sides being nearly parallel throughout, 
with almost imperceptible contractions at the septa. In Deltania elegans this contrac- 
tion is very prominent, and the ganglion is almost twice as wide in the posterior part 
of the segment as in the anterior one. This characteristic appears constant. In 
Deltania Benhami the ganglion is narrowed somewhat at the septa, but the posterior 
part in each segment is not any wider than the anterior part. 


CALIFORNIA EUDRILID®. Ta) 


Septal Glands (figs. 26, 27, 29). When the worm is laid open and the cavity 
viewed from above, it is seen that there are 4 pair of septal glands surrounding the 
cesophagus in somites v, vi, vii and viii. In this view the anterior gland appears the 
largest, and the posterior one in viii the smallest. This is, however, only an illusionary 
appearance, caused by the position of the glands. There is in reality not any very 
considerable difference in their size, as may be seen when separated and spread out. 
Seen in a slightly eccentric longitudinal section, the gland in vi appears the largest both 
above and below the cesophagus, though in some sections the lower part is not as large 
as the lower part of the gland in vii. The anterior gland in vy is short but broad. The 
one in vii is larger than those in v and viii, but smaller than the one in vi. The 
upper part of the gland in viii is larger than the corresponding part of v, the lower 
part of the latter being the smallest. As will be seen, all the glands are developed, 
both superiorly and inferiorly, as regards the cesophagus, but the glands on either side 
in the somite do not connect, but only touch. There is a slight, but irregular lobing 
of the glands, frequently unequal on either side, as one gland may be almost entire, 
while the other again is furnished with three indentations. On the under side of 
each main gland there is a smaller lobe, almost entirely separated from the rest (fig. 
29). The glands are furnished with blood from the subcesophageal longitudinal blood- 
vessel which projects a branch to each gland, on which it again divides in two or 
three parts (fig. 29). I may add that the septal glands are very large, almost filling 
the respective somites, and as compared with those of Dedtania elegans, about three 
times as large, considering however the relative size of the two species. The glands 
are nearly similar to those of De/tania Benhami, but with more unequality as to size. 


Salivary or Pharyngeal Glands (fig. 26). ‘These glands resemble those of the 
preceding species, De/tania elegans, in general appearance. There are two very long 
glands behind the brain, attached on the underside of the two long muscular bands 
which stretch upward. The anterior one of these is the smallest and rather short. the 
second in order from the brain is the longest. The posterior gland, which forms the 
posterior projection of the pharynx, is much shorter, more compact and rounded 
than the corresponding gland in Deltania elegans. The whole mass of glands projects 
much less posteriorly than the glandular mass of the pharynx in that species. 


Spermatheca (figs. 30, 31, 32 and 39). These organs are prominent and char- 
acteristic of the species. There is one pair in somite ix opening in the inter- 
segmental groove between that somite and yiii. The external pore is in front of the 
inner couple of sete, but not interior to the sete. The organs are thick, opaque and 
of the form of pointed sacs, each one with two diverticula, one on each side, which 
connect with the main sac close to the external pore. The outline of the sac is irreg- 
ular in some places, toward the inner apex assuming the appearance of one or more 
warty diverticula, which, however, never assume the size of the diverticula. Of these 
latter there are one pair which are slender, cylindrical, of more or less irregular out- 
line with the apex sometimes slightly wider, sometimes helix-like, turned on itself. 
The lower part of the spermatheca is muscular, but this muscular part is quite small, 


36 CALIFORNIA ACADEMY OF SCIENCES. 


not extending above the junction with the diverticula. In general form the sperma- 
theea resembles that of the following species, D. Benhami, but the outline of the main 
sac is less regular and the diverticula are larger, being about one-half as long as the 
main sae (figs. 50 to 32). 


Testes. As usual there are two pair of small testes occupying somites x and 
xi. They differ in nothing particular from those of the other two species of the 


genus. 


Sperm-sacs. There are two pair of sperm-sacs, one each in somites x and xi. 
They are long, not very opaque bodies, occupying a large part of the somite, though 
not filling it closely. The spermatophoric spherules are comparatively large, 
globular. The sperm-saes are not greatly lobed, extend considerably in length from 
the dorsal to the ventral side, and are of more undecided shape than the sperm-saes 
of Deltania elegans and Benhami. They are also, comparatively, much larger (prin- 
cipally higher) than in that species. It will be noticed that the sperm-sacs occupy 
somites x and xi, while in Detania elegans, Benhami and dubia they occupy somites 
xi and xii. 


Ovary and Oviduct (figs. 35, 36, 37, 38). As might be expected the ovary is 
situated in xii. It offers no characteristics of interest. There is no ovisac. The 
oviduct is as usual funnel-shaped, either deeply cut or folded on one side, the inner 
funnel in xiii, the ovipore in xiv. Close to the oviduct in xiy there is a yery peculiar 
sac (figs. 35 et seq.), of the size of the oviduct or smaller. It does not open directly in the 
oviduct, and it has the general shape of aseptal gland with many rounded lobes arranged 
as the petals in a rose. The epidermal covering does not closely cover the inner cells 
which are irregular, apparently not closely packed, of uneven size and shape, round 
or conical, each with a round nucleus, and grainy cell contents. This gland does not 
connect with the opening of the oviduct funnel, from which it is separated by the 
septum between xiii and xiy. There is one pair of those glands, one behind each 
oviduct. The gland is affixed to the posterior part of the anterior septum in xiy. A 
similar gland does not exist in De/tania elegans. As to the nature of this organ I can 
say nothing definite, and I hesitate to consider it as an ovisac, until a more extensive 
material will allow other investigations. 


Spermduct and Prostate (fig. 35). As in the preceding and following species, 
the spermduct and prostate open in the same pore in somite xvii. The spermduct 
opens slightly posterior to the prostate and also more outwardly, though both organs 
closely join at the pore. The spermduct is quite wavy throughout its length to the 
ciliated rosettes, which as usual are found in x and xi, The rosettes are less folded 
and crenate than those in Deltania elegans. The prostate differs somewhat from the 
one in the latter species. The muscular duct is not helix-like at its upper end, either 
straight or bent at right angle to itself, with the distal end pointing backward, being 
parallel to the longitudinal axis of the body. The relative size of the prostate is in 


CALIFORNIA EUDRILID®. 37 


this species much larger than in De/tania elegans, which appears to be characteristic 
also with nearly all the other organs. The upper glandular part of the prostate is 
about three times wider than the lower muscular part. The latter is about equal in 
length to the penial sete and their sacs. 


Genital or exterior male zone (fig. 25). In somite xvii there is a pair of ven- 
tral papillee close to the ventral ganglion, and situated in the transverse median line 
of the somite. In these papille open the penial sete, in the place which otherwise 
would be oceupied by the regular sete. Between these papille and the ventral 
median line of the body, somewhat nearer to the posterior margin of the somite, are 
seen on either side a circular cup-shaped depression, from the center of which is spread 
backward a large fan-shaped branch of muscles connecting with the posterior inter- 
segmental groove. In the center of this suctorial organ, and at the very point from 
which the fan-shaped muscular fascicle starts, is situated the exterior opening of the 
spermduct and the prostate. In the median line between the suctorial cups is a smaller 
triangular depression. The anterior part of the somite is raised and thicker than the 
posterior part, or rather there are two large anterior folds, and seyeral smaller posterior 
ones in the vicinity of the male pores (fig. 25). 


Deltania Benhami. 
Plates xv and xvi, figs. 40-48. 
Deltania Benhami Eisen, Zoe, iv, 212, October, 1895. 


Size about 1 inch by ;;.. The inner couples of setee as well as the sete in the 
inner couples are much closer together than in any other species. The spermatheca 
are large, opaque, situated in ix, and opening between ix and viii, with two diverticula, 
which are less than 5 as large as the main spermathecal sac. A small species, in 
many respects resembling Deltania Troyeri, bat very distinct by the above character- 
istics. 

Habitat. 1 haye found this worm only in a guleh or cafion at the outlet of the 
waterworks and dam, known as Lake Chabot, east of Alameda and San Leandro, in 
Alameda Co., California. The worm is very scarce and lives under damp leaves in 
the very top layer of the soil around the roots of trees. The exact locality is to the 
right of the gate which closes the reservation, down by the creek, not far from the 
wire fence. It occurs here alone, not mixed up with any other species, and to all ap- 
pearances this species is a true native and not introduced. It is an exceedingly deli- 
vate worm, almost transparent, white, with yellowish clitellum, very impatient of 
being handled and can only be kept alive with great care. It is much more trans- 
parent than any of the other species. 


Exterror characters. In general appearance, the worm resembles De/tania 
Troyeri, but is slightly larger in size. The second somite is much narrower than in 
that species, being larger than the third somite. But it is especially as regards 
position of the sete that the greatest external difference exists (fig. 40). The yven- 
tral setee in one species are much closer together than in the other species of the 


38 CALIFORNIA ACADEMY OF SCIENCES. 

genus, and average about twice or three times as close as in Deltania Troyeri. 
This is a prominent and constant characteristic. The characteristic feature of the 
genus, which consists in the narrowing together of the ventral setze on both sides 
and towards the male pore in xvii, is readily seen in this species. On account of 
the already very close approach of the ventral setee of each couple, this narrowing 
together is, however, less perceptible in our present species, but it is still consider- 
able and readily seen. The figure (fig. 40) will illustrate this better than any 
lengthy description. Not only does this approach of the setee exist on both sides of 
the male pore, but it is also seen towards the spermathecal pore, though here in a 
smaller degree. In Deltania Troyeri, there is not a trace of such an increased ap- 
proach of the ventral setee towards the spermathecal pore. Another characteristic as 
regards the setee is that the ventral or inner couples are much closer together than 
in the other two species described here, so close that the main bodies of the sperma- 
thecze almost touch. 

The spermathecal pore is situated in the intersegmental groove between viii 
and ix, in front of seta 1. In Deltania Troyeri, the spermatheeal pore is in front of 
the corresponding seta 1, but considerably more lateral, and on account of the 
greater distance of the ventral couples of sete, the spermathece in that species are 
much further apart, and do not crowd the ventral ganglion. Ovidueal pore as usual 
in xiy. Penial setze on papille in xvii, opening in the male pore. The clitellum 
comprises three full and wide somites, xiv, xv, xvi, and two smaller ones, xiii and 
Xvll, as in other species. 

The genital region (fig. 48) around the male papille is much simpler than in 
Deltania Troyeri. The two papille are situated much closer together, and there are 
no suctorial depressions with their fan-shaped arrangement of muscles. The penial 
setee are very slender, sickle-like, bent at the free apex, with a narrowed and sharp 
point. There are a dozen or more external fibres of exceeding thinness stretching 
from the anterior part of somite xvii to the posterior part of xviii. These 
threads, however, exist only in the longitudinal groove between the papille, all run 
parallel with each other and with the long axis of the body (fig. 48). As to their 
nature Tam not at all certain. They are apparently too thin to be muscles. The 
clitellum proper in this as in the preceding species ends at the papillee or male pore. 
Anterior to it the clitellum consists of regular flask-like cells; posterior to it again 
the body-wall has assumed its regular appearance. In De/tania Troyeri the clitellar 
thickening stops just before it reaches the papillee, but in De/teinia Benhami it stops 
just as it reaches the male pore. ‘The papillee, however, continue to the posterior end 
of the somite. 

The length of the species is about 1 and 14 inch, by ; to § inch wide, taper- 
ing towards the caudal end. In fact the general form and size does not differ from 
the preceding species. In all I found some fifteen specimens, but they were, un- 
fortunately, in poor condition when I arrived home, and the part posterier to the 
clitellum had already begun to decompose. The following account of the anatomy 
is, therefore, not as full as desirable, but enough is known to perfectly characterize 


CALIFORNIA EUDRILID#&. 39 


the species. There are four pair of septal glands in y, vi, vii and vii. The glands, 
as far as I can judge from dissections only, are of almost equal size, somewhat longer 
there than in Deltania Troyeri. They are equally developed on the lower and upper 
side of the csophagus. The alimentary canal offers no great peculiarities. The 
specimens were all very much stretched, and I am not certain if the form of the 
cesophagus will prove constant. However, the contractions at the septa were much 
smaller than in any other species. The cesophagus from somite ix narrowed down 
toward somite xii, here it began to gradually increase in width, but the sacculated 
intestine begins evidently first in xvii, increasing in width gradually backwards until 
it reaches the region between xxvii and xxxy, where it suddenly narrows and con- 
tinues as narrow to the end of the body. 

Spermatheca (figs. 42 and 43). There is one pair in ix, opening between viii 
and ix, of the same general appearance as in Deltania Troyeri with minor character- 
istic details. The main sac is ovoid and somewhat lunate, pointed, with very smooth 
outline and with no trace of warty excrescences. There are two diverticula affixed 
halfway between the base and the glandular part. They are much smaller than 
those of Deltania Troyeri, being less than one-third as long as the main spermatheca, 
while in Deltania Troyeri the diverticula are one-half or more as long as the main 
spermatheca, and affixed to the muscular part close to the base. The spermathecze 
open in front of the 1st setae and are situated much closer together than those in 
D. Troyeri. The opaqueness of the spermatheca is the same as in the latter species. 
There is, however, a decided difference in the location of the spermatheca. In 
D. Troyeri they are situated so far apart that they do not touch the ventral ganglion. 
In D. Benhami, however, they crowd it, this approach being caused partly by the 
closer proximity of the ventral or inner couple of sete, partly by the situation of the 
spermathecal pores which in our present form are more ventral to the sete. 

The sperm-sacs (fig. 41) are of a very characteristic form. They are larger in 
proportion than those of D. e/egans, but not quite as large as in D. Troyeri. There are 
two pair, one each in ix and xii, and they do not connect with each other. 
Each one consists of a very large flesh-like lobe, at the base of which are seen half a 
dozen smaller and globular saes, all connected at the place of adherence to the an- 
terior septum. These different lobes, the large one as well as the small ones, are 
full of rounded, oblong or irregular spermatophoric spherules. The sperm-saes do not 
by far fill the somites. The form of the sperm-sacs varies to some extent, but the 
main character is the same in all, a large, rounded or flask-like lobe, at the base of 
which are several smaller ones. 

Spermduct and ciliated rosettes as in D. Troyeri. The prostate offers the char- 
acteristic of having the lower part of the glandular sac considerably swollen, conical 
and gradually diminishing in size towards the apex, which again is slightly enlarged 
like a knob. There is only one long slightly bent or almost straight seta in each of 
the penial seta-sacs, both setze opening on a smali papilla, and which as far as I can 
see more resembles that of D. Troyeri than D. elegans. The blood is very pale yellow, 
paler even than in the other species. There are three strongly pulsating hearts in 


40 ; CALIFORNIA ACADEMY OF SCIENCES. 


somites x, xi and xii, connecting the dorsal and ventral vessels. There are also 
connections between those two vessels in ix, x and xiii as in the other species, but 
those connections consist of regular secondary vessels narrow and tubular, and in no 
way resembling hearts. There is no subneural vessel. The ventral vessel is divided 
in somite vi, and in the anterior six somites there is a heavy capillary system connect- 
ing the ventral and dorsal vessels. 

In the following I have endeayored to give a comparative table of the species 
which I refer to De/tania. As far as concerns those species which I have not myself 
investigated the table must be considered tentative, especially so in regard to the char- 
acter of D. dubia, the description of which by Fletcher is somewhat insufficient. I need 
here hardly add that Deltania Poultoni is the species described from Maderia by Bed- 
dard as Microscolea poultoni. The description given for D. dubia is taken from Rosa’s 
description of Jicroscolex dubius Fletcher, and not from Fletcher’s own. 

DELTANIA. 


—s « = es 


D. elegans. sp. D. Troyerin. sp. D. Benhamiu.sp. D. Poultoni Bedd. D. dubia Rosa. 


Septal glands in vi, Vili, viii, ix. Vv, Vi, vii, viii. Vv, Vi, vii, vill. Not present. 
Sperm-sacs in aE aah xxi xi, xii. Xi, Xi xi, xii. 
Spermatheca Very small and Large, apart, with Large, close togeth- Not present. Not present. 
pellucid. No di- 2 diverticula each. er, with 2 diverti- 
verticula. Opaque. culaeach. Opaque 


The ventral or inner Only a little closer Only a little closer Several times closer Several times clos- Several times clos- 
sete in each couple! together than the together than together than er together than er than the outer 
exclusive of those sets of the outer those of the outer those of the outer the outer couple.| couple. 


in the genital re- couple. couple. couple. 
gion, are: 
Prostate, the gland- Helix-like at the Not helix-like; tub- Not helix-like; tub- Helix-like at the 
ular part top. Muscular ular, straight or ular and broad at top. Muscular 
part not very bent. the base. Top en- part very short. 
short. larged kuob-like. 
Gizzard None. None. None. None. Rudimentary. 
Diverticula of the Absent. More than }as large About $ as large as 
spermatheca as the spermathe- the spermathecal 
cal pouch. pouch. 
Penial seta Two on each side. Two on each side. Two on each side. One on each side, Two on each side. 
‘ | | 
= oe | ox 
Nephridia com- il. li. | il. il. 5 
mence in 
Nephridio-pores in 4th and 3d sete. 4th and 3d sete. | 4th and 3d sete. 


| 3d sete, 3d seta. 


front of and little Prominent in the Not prominent in 


interior to | clitellum. the clitellum. 
Spermducts Remain separate|Remain separate up|Remain separate up Remain separate Unite to form one 
up to the atria. | to the atria. | to the atria. up to the atria. tube on each side 
| which opens in 
} the muscular part 
of the atrium. 
Posteriorly to the xxviand xxyii. xxl. xxl. XXxili and xxiv. 


clitellum the setz 
have regained their 
normal distance in 
somite 


CALIFORNIA EUDRILID®. 4] 


ARGILOPHILUS. 
Argilophilus Eisen, Zoe iy, 252, October, 1893. 


Prostomium encroaches on somite i. Eight setee in four couples, ventral, 
lateral and dorsal, commencing in somite ii. The sete of the inner couple not con- 
verging towards the male pore, but closer set than the sete of the outer couple. The 
alimentary canal consists of an eversible buccal cavity, a pharynx, cesophagus, gizzard, 
tubular intestine, sacculated intestine, typhlosole, but no cesophageal glands or pouches. 
Clitellum not developed ventrally, occupies somites xiii to xviii. 

Spermathecal pores, one pair vii /viii and one pair vili/ix. Ovipores in xiy- 
Male pore in xviii. One or two rows of ventral intersegmental papille. Two pair 
of spermathece. Testes in x, xi. Sperm-sacs paired in x, xi, xii, generally enclos- 
ing the ciliated funnels and testes. Two pair of ciliated funnels. Two pair of sperm- 
duets, which join a pair of very large, tubular-coiled prostates in xviii, at the upper end 
of the muscular duct. Two penial sete open iu the same pore, but not in the same 
duct as the prostate. 

Dorsal vessel and ventral vessels connected by 5 pair of hearts in xiv to x. No. 
subneural yessel. Blood red. Many blood vessels on the nephridia. No pepto-nephri- 
dia. The nephridio-pores variable as to location, the majority open in front of or lateral 
to the 4th sete, though many open interior to the 4th sete. No coecal bladder at the 
exterior pore. Large earthworms with thick round bodies, of pale flesh color, mar- 
bled bluish. 


Distribution and habitat. The genus Argilophilus appears to be an undoubted 
native of the Pacific Coast. Specimens have been found in the San Joaquin Valley 
in California, and as far north as British Columbia (Vancouver Island). In California 
these worms are our most common earth worms, appearing close to the surface with 
the adyent of the rains in the autumn and disappearing deep in the soil with the dry 
weather in May, after which time they are not any more found in even locally moist 
places. During the summer months I haye sometimes dug up these worms from a 
depth of 5 to 6 feet or more, each worm tightly rolled up as a little ball and appear- 
ently eneysted in a chamber of clay, the inner surface of which is smooth and hard. 
In these cysts the worms pass the dry season. These worms are hardly ever found 
outside of heavy adobe or clayey soil; the more clayey the soil, the better the worms 
appear to thrive, provided also the soil is rich and fertile. In poor soil the worms are 
seldom seen, and the best indorsement for a soil is that it contains worms of this genus: 

The color of the worms of this genus is fleshy pink, thickly marbled, with steel 
or slate gray, (fig. 132). The clitellum is yellowish red, and the whole anterior part 
is pinkish. The color of these worms is very handsome and distinguishes them from 
the deep brown Allolobophora so common in moist or swampy places in this State. 


Exterior characteristics (figs. 125-131). The most prominent exterior feature 
of this genus is the color which has just been described. Another is the frequent 
eversion of the lining of the buccal sac (fig. 150). As to size the worm must be 


42 CALIFORNIA ACADEMY OF SCIENCES. 


considered as our largest earthworm, though very variable, as might be expected. 
The smallest adult worms measure about 15 inches by .5 lines, the largest again 6 
inches by 4 to 43 lines, the size appears to depend greatly on the locality and richness 
of the soil. The prostomium divides somite i to about $ or 3 (fig. 150). Somites vii, 
vill, ix, are larger than the other anterior somites. The clitellar somites (figs. 125 to 
151) are large, the post-clitellar ones are very much smaller. The spermathecal 
pores are more or less conspicuous (fig. 129 spth.), sometimes hardly visible, at times 
again elevated and appearing as small round rings. ‘They are situated just lateral to 
setee 2, but of course in the intersegmental grooves between vil /villand viii /ix. The 
ovipores are closer together, situated a little more ventrally than sete 1, sometimes, if 
not generally, connected by a depression. The clitellum is only developed dorsally 
and laterally, the ventral part between the male pores being normal and appearing as 
considerably depressed, in hardened specimens the depression reaching as far forward 
as to the center of somite xiv or the ovipores. 

The male pores which open in line with sete 2, are situated on either side on an 
elongated papilla, which again is more or less surrounded by a circular depression, 
outside of which is seen a high semi-circular ridge, which is thicker anteriorly and 
posteriorly than laterally (fig. 25, 129). The penial setee are one pair in each 
pore, and are seen protruding through the male-pore. 

The regular sete are sigmoid, not greatly bent. The setee of the inner couples 
are closer than those of the outer couples; all the setee are in parallel rows (fig. 24). 
The nephridio-pores are difficult to view from the outside. Their arrangement is 
variable, but the majority are found outside of, or more lateral than the fourth row of 
sete. The three anterior nephridio-pores are seen in front of sete 4 (fig. 24). A 
more detailed description will be given further on. 


Setw. The ordinary sete have been already described as sigmoid. There 
are two sacs of penial setze attendant to each prostate, and opening in the same pore, 
but not in the same duet, as that organ. There is only one seta in each sac. This 
seta is sickle-like, much more curved than those in Deltania. The point is needle- 
like, and curved (figs. 122, 123). The very point is void of sculpture, but the part 
back of the point and up to the sae is sculptured as in the figure 123. The largest 
part of the seta is smooth, only showing the rings for the attachment of the muscles. 
The inner couples of sete of the clitellum are somewhat raised, though otherwise not 
differentiated. 

ANATOMICAL STRUCTURE. 


The body-wall (figs. 114, 115, 116, 117, 118). The body-wall outside of the 
clitellum shows the usual sets of layers. The innermost vascular layer, which covers 
the longitudinal muscular layer, is very thick and prominent, though not very 
crowded with blood vessels (tig. 114). Under this layer, and between it and the 
longitudinal muscles, passes the spermduct, almost throughout its length, from the 
place where the two ducts unite to the one where they rise to join the spermduct. 
This layer is less pronounced anterior to clitellum, and appears absent in the vicinity 


CALIFORNIA EUDRILIDA. 45 


of the spermatheca and anterior to them. In the elitellar somites this vascular layer 
is especially prominent. The muscles of the longitudinal muscular layer in this 
genus, as well as in Deltania, are arranged in groups or projecting lobes, between 
which pass projections of the vascular layer, as well as transverse muscles in certain 
places. These lobes vary in width, and on the ventral side below the ganglion are 
arranged fan-like (fig. 105, v. p.), diverging from the median line. The longitudinal 
muscles are never arranged around a central axis, as is the case in so many lum- 
bricides, though they show a faint trace of symmetrical arrangement. ‘The zone of 
the transverse muscles is much thinner, five or six times narrower than the longi- 
tudinal zone (117). 

The hypodermis is thick, with large, glandular cells of a flask-like or spindle- 
like shape. In the clitellar somites these glands become irregular, club-like, and 
project as far inside the layer of clitellar glands as the hypodermis is thick, or more 


(fig. 116). 


The clitellum is developed only dorsally. The glandular layer is much 
thicker laterally and dorsally, tapering towards the ventral side, and ceasing entirely 
at a line drawn outside of the male papilla and parallel with the ventral ganglion. 
The glandular layer of the clitellum is very thin, and as compared with that of 
Deltania, about 4 narrower. The cells of this layer are, however, very wide and 
long, there being generally eight or nine in the row. They are irregularly grouped 
in twos or threes, separated by narrow blood vessels, which, at rather regular inter- 
vals, are thicker. They are supplied with blood from sinuses situated between the 
transverse muscles, and which connect through these branches with a capillary net- 
work on the hypodermis (figs. 115 and 116). 


Transverse muscles (figs. 118 and 119). There are numerous sets of trans- 
verse muscular bands in the clitellar somites, quite similar to those described by 
Benham in JJoniligaster indicus. They are more numerous and prominent in the 
somite of the oviduct than elsewhere, and form there three distinet muscular bands 
the ventral ends of which terminate at the inner couple of sete (fig. 119); the lateral 
ends again terminate on the lateral side of the body wall. 

The posterior band is the smallest of the three and begins in the posterior and 
ventral part of the somite, in line with, but slightly posterior to the inner sete, stretch- 
ing from there diagonally across the somite, ending laterally at the seta 3 (fig. 
1S) ie WN 
The next band is much larger and begins in the anterior part of the somite 
also in line with the inner setie and stretches diagonally backwards ending posterior to 
but in line with seta 4 (fig. 119, m 2.) The third muscular band is of the same 
size and runs in the same general direction as the last, begins and ends in front of it, 
ending in front of the fourth seta (tig. 119, m3.) A fourth muscular band of a 
somewhat similar character connects the posterior part of the oyiduet with the body- 
wall terminating in front of the muscular band just described as m 1, on the figure 
(fig. 119, m 4.) Similar museles as the oyidueal one, are common in all earthworms, 


44 CALIFORNIA ACADEMY OF SCIENCES. 


connecting the various organs with the body-wall and require here no particular men- 
tion. ; 

In the other clitellar somites we find bands of transverse muscles which 
stretch diagonally (fig. 118) across the body wall from the ventral part, almost im- 
mediately below the ventral ganglion, to the vicinity of the outer sete. The ends of 
these muscles penetrate between the lobes of the longitudinal muscular layer (fig. 
118, 7. m.) These muscles are not continuous through the somite but are grouped in 
bands. 


Ventral papille (figs. 120, 125 to 151). Argilophilus is readily distinguished 
from other California Eudrilidee so far known, by its ventral papillee, occupying the 
ventral side of the intersegmental grooves from the spermatheca to the segments next 
posterior to the clitellum. In Argilophilus marmoratus ornatus we meet sometimes with 
as many as 7 or 8 pairs, while in A. m. papillifer we find as high a number in a single 
median row, under the ventral ganglion. In no instance did I find a papilla on the 
segment itself, all invariably occurred rising from the grooye between the somites, 
being in other words intersegmental. From cross-sections it will be seen that the 
papilla consists of two distinct parts, one exterior and lateral, consisting of elongated 
hypodermic supporting cells, which more or less fully enclose the interior main body 
of the papilla. This central part consists of larger or smaller gland-like bodies, vary- 
ing in size and number according to the size and age of the papilla, ete. In some 
papille these bodies fill the larger part of the papilla, in others they are confined to 
the bottom or very center of the organ. As to the nature of these organs I am, as yet, 
somewhat in doubt, though they certainly must be considered as sensory organs 
of some kind. Stained with osmic acid the bodies present in sections a darker center, 
which appears of ganglionic nature, around which are grooped larger sacs which 
again are composed of smaller, light-refractive granules, giving the idea of a reticu- 
lated protoplasm. Numerous nerve fibres connect with those bodies, and evidently 
penetrate to the gaglionic center. Long tube-like cells butt directly on these gland- 
ulous bodies, while others grooped in bundles penetrate between them connecting with 
nerve ganglia. The transition between the supporting cells and the drainpipe cells 
of the papilla is sudden and not gradual. In fact the line of demarcation between the 
two is quite prominent. The supporting cells in the young papilla, entirely enclose 
the papilla, while in the larger and full grown papilla, they are pushed towards the 
side leaving the whole center to the drainpipe cells and the glands. 

When stained with safranine the nuclei of the glandular cells become prominent, 
less so when stained with heematoxylon. The nuclei vary greatly in size and number. 
In some of the glandular bodies there are from one to three nuclei, in others none. In 
sections there are always less nuclei than divisions (cells?) apparently lying on the top of 
the reticulated mass. The nuclei may be seen directly above the nerve center, but 
generally they are found outside of it. It seems as if these glandulous bodies are 
modified agglomerations of hypodermic glands. 

The arrangement of the glandulous bodies and the tube-like cells is very 


CALIFORNIA EUDRILID&, 45 


regular, the former resembling small sugar-loayes standing in a row, with the fan-like 
arranged tubular cells between them (fig. 120). In some sections a bunch of nerve 
fibers is seen on either side touching the papilla, connecting on the ether hand with 
the ventral ganglion. As to the nature of the papilla Dr. Michelsen suggests that a 
somewhat similar organ in Acanthodrilus georgianus is a taste or “ Wollust”’? organ. 
Tt is, however, not unlikely that the minute light-refractive bodies are glandulous. 

Many of the tubular cells contain a fine granulated secretion in varying quan- 
tities, which stain dark red with eosine. The fact that the whole papilla is concave in 
the center speaks also strongly for the glandulous nature of the organ. The figure of 
the “Augenapfela’-like organ of Acanthodrilus, given by Michelsen, may possibly have 
been taken from a young papilla in which the glandulous bodies had not yet developed. 
Organs of a somewhat similar nature have also been described by Horst from Ponto- 
scolex: corethrurus which, however, he does not figure in connection with nerve fibres, 
It is not improbable that all the sensory organs in the genital somites of the higher Oligo- 
cheeta are of an analogous nature. Among such organs would be included the tuber- 
cula pubertatis, the puberty grooves, as well as some other epidermal structures in 
Heliodrilus, Hyperiodrilus and Eudrilus described more in detail by Beddard. How- 
ever all the organs require re-examination by the aid of other methods, as one single 
method of staining will not suffice to reveal their true nature. 


Septa (fig. 86). The anterior septa are greatly pouched, generally to such 
an extent that in cross-sections the various organs appear to lie several somites further 
back than they really do. Thus the gizzard and the posterior spermatheca may be 
seen in the same cross-section and this is also the case with the oviduct and the sperm- 
sacs. This pouching is principally restricted to the septa 5 to 15. These septa are 
also slightly thickened especially those bounding somites viii to xi. 


Nephridia (figs. 59 to 77). The position of the nephridio-pores places Argilo- 
philus very close to Plutellus. There are one pair of nephridia in each somite as usual. 
The first pair of nephridia are found in somite ii and others follow in all the posterior 
somites. The first five or six nephridia are somewhat larger than the others and open 
in front of the fourth sete. All the following open irregularly in front either of the 
third or fourth setee, or in the space between and anterior to them, or even outside of, 
or more lateral than seta four. Those which open laterally of seta four do not even 
open in the same row, as we find one nephridio-pore say as far out laterally from 
the fourth seta, as that seta is distant from the third, while others are half-way or one- 
third of the way between the fourth seta and the most lateral nephridio-pore. There 
is no regularity as regards this succession, though rarely two successive nephridia open 
in line behind each other. For instance, one nephridium opens in front of seta 
3; the second as far outside of 4.as 4 is distant from 3; the third nephridio-pore is 
in front of seta 4, the following half-way between seta 4 and the most lateral pore, 
the following % of the distance from seta 4 to the most lateral pore, the following in 
front of seta 4, the following again % of the distance from seta 4 to the most lateral 
nephidio-pore, etc. (Fig. 124 np. p.) 

Memoirs, Vo, II, 3. January, 1894. 


46 CALIFORNIA ACADEMY OF SCIENCES. 


The structure of the nephridium is considerably complicated. The nephridio- 
stome, found in front of the ventral sete, is unusually minute, and leads to a com- 
partively short duet, which connects with the body of the nepridium proper, at the 
point where the single-tubed outlet leaves the folds. This tube is not convoluted (fig. 
59 @)), but almost straight, very hard and solid, widening toward the base. When 
it reaches the folds it does not at once enter a tube, but forms a long, cylindrical, 
spongy mass (fig. 63), which extends the whole side or one-third of the length of the 
nephridium, before it assumes the proper shape of a clear canal (fig. 59, @) to @), 
and fig 64). In the beginning at @), this mass shows no regular lumen, but a num- 
ber of irregular pores and tubes, which might best be compared to the inner canals 
of a common washing sponge (fig. 64, @), @),(@)). In the center of this mass are 
imbedded the two parallel folds of the main nephridial canal (fig. 64, @ and b). At 
first the spongy tube is located principally above the two canals, but soon the mass is 
shifted and the canals become imbedded in the center of the mass, or very nearly so 
(fig. 64, (@) toG)). The small connecting tubules at @), which were at first so ir- 
regular, soon assume a greater degree of regularity at (8), while at the same time two 
longitudinal lumens are formed—one on the upper () and one on the under side 
(@),, fig. 64) of the two central canals (a and b). At first these canals @) and @) 
are indistinct and irregular, but soon they assume the character of regular longitudinal 
canals (from (4) to@), @ and @)). From @) to @) these canals (@) and G) are 
connected by the transyerse tubules, which completely surround the two central 
canals (2) and @). At G) the lower canal G) becomes narrower, and the trans- 
verse ducts drop into the main canal (©), while the former lower canal @) assumes 
the character of a rather thick, epithelial lining. The general effect of this arrange- 
ment is, that, seen with a lower power, the spur between () and () appears to con- 
sist of four distinet parallel canals, while from («) to (3) the fold contains only three 
parallel canals. At @) these canals become very crowded, the whole fold being nar- 
rower. The length of this narrower part varies, but generally already at (), or 
shortly before entering on the bent plexus at (), the fold has regained its original 
width. At @) the canals turn; that is, @) connects with (@) and the central canal 
folds upon itself, and for a short distance we have four almost parallel canals. At 
(8) the formerly central canal (from (8) to @), ©) and @)) leaves the lobe and crosses 
over to the other fold, and at (@) becomes the original canal @), which runs its 
course all through the two folds, at @) turning downward, becoming the lower canal, 
which again at @2) becomes canal (©), from there on pressing backward and again 
forward from (@) to @), at which point it separates itself from the fold, and, running 
along the inner body-wall, forms the outlet duct opening at the exterior pore at (#). 
The connecting bridge between the two main folds, or rather between one fold and 
the free lobe of the other, between (@) and (®), is very narrow at (2), suddenly 
increasing at (@), gradually tapering toward @). As regards the ciliation of the 
canal, it may be stated that it does not extend all through the tube. It appears that the 
canal is ciliated at places where the passage of the excretions is difficult. The narrow 
duct from the funnel to () is ciliated as well as the funnel itself. Again, the ciliation 


CALIFORNIA EUDRILIDA. 47 


begins at the point of recurrence at (@), but ceases after the duct has assumed its straight 
course at (6); it begins again at (@), and is with a certainty found between () and 
@), and possibly between (@) and (2), where, however, I am not quite certain of 
its presence. The whole nephridial fold is imbedded in a wing-like cellular and 
fibrous mass, which at either end becomes thicker, supporting large pellucid perito- 
neal cells, such as found in the nephridia of many Oligocheta (fig. 64, per. c.), and a 
complex system of blood-vessels, which branch and form capillary ioops partly on the 
folds, but principally between the spongy tubes and the two central canals (figs. 59, 
63, 64, 67, b/.). The blood-vessels originate from a branch of the ventral vessel as in 
Lumbricus. 

Scattered over the peritoneal cells are masses of free cells, generally agglom- 
erated in separate heaps. Each cell is deeply crenate, as if it consisted of several in- 
dividual cells, but in each such small agglomeration there is only one round nucleus 
(fig. 80). 

If we recapitulate, we find that the nephridum of Argilophilus consists of the 
following distinet part from the nephridio-stome to the nepridio-pore: 

1. Nephridio-stome, engaged in the anterior septum. 

2. Narrow duct, which connects the nephridio-stome with the body of the ne- 
phridium proper, especially with the spongy tubules. 

3. Spongy tubules, which are at first irregular, but which soon fuse into one 
main-tube with many branching tubules. 

4. Main nephridial canal, which, recurring on itself several times, forms two 
distinct folds and one spur. The posterior fold contains besides the tube and tubules 
two turns of the canal, one of which is recurring. The anterior fold contains three 
turns of the canal, two of which are recurring. Part of this anterior fold disengages 
itself from the main nephridial mass and forms the : 

5. Spur. This spur contains four canals, two of which are recurring, the 
point of recurrence for all four being at the distal end. One of the recurring canals 
of the spur is connected with the posterior fold by the 

6. Bridge, a part of the canal much narrower than the other, spanning the 
distance between the two folds. 

7. The wide duct leading to the nephridio-pore, directly connected with the 
recurring canal of the posterior fold. In different nephridia this wide duct is of 
varying length. 

8, Nephridio-pore, apparently without urinary bladder or collar. 

9. <A large wing-like, but rather thin, mass of peritoneal cells. 

10. <A complicated and extensive system of capillary blood vessels on the va- 
rious parts of the nephridium. 

Compared to the nephridium of Lumbricus as described by Benham we find 
a few more important differences, especially in the post-septal part. Thus the post- 
septal part of the narrow tube after reaching the first loop assumes the shape of a 


48 CALIFORNIA ACADEMY OF SCIENCES. 


tubular network, which only gradually fuses itself into one duct with numerous 
branching duetules, while in Lumbricus the single duct continues through the differ- 
ent lobes and in recurring winds around itself. The tubules in Argilophilus em- 
brace the two tubes but cease after leaving the first fold. The muscular duct which 
is so prominent in Lumbricus is not represented in Argilophilus, but is replaced by an 
elongation of the single wide tube corresponding to the “‘ wide tube of the 3d lobe ” in 
Lumbricus. 

Compared again with the nephridium of Deltania, we find that the principal 
difference consists in the absence of the urinary bladder and the collar at the ne- 
phridio-pore as well as in the absence of a coecal bladder. The irregular or alternate 
locations of the nephridio-pores distinguish Argilophilus and Plutellus from all other 
earthworms, as far as known. 


ee 


Alimentary canal (fig. 87 to 92). The buccal cavity is, as has been stated, 
greatly eversible, and generally remains everted after the worm is dead. The 
pharynx is only developed superiorly though there is a slight thickening of the lower 
wall of the buceal cavity at the junction with the cesophagus, at which place numer- 
ous muscles are seen to connect with the lower part of the body-wall. The pharynx 
is as usual furnished with numerous salivary glands (fig. 86, s/. g/.), extending from 
the vicinity of the brain to the posterior part of somite iv, the most posterior glandular 
mass being the largest (fig. 86). The various glandular lobes offer some characteristics, 
which if carefully noted and compared may be found to be constant enough to serve 
as species characteristics. 


The @sophagus begins in ili and oceupies somites iii, iv and y, (fig. 86, A. @.) 
forming first a narrow tube, which widens out, and rising upwards connects with a 
very large gizzard (gz.). This gizzard occupies in reality only somite vi, but its great 
length causes it to push far backward to such an extent that it actually occupies the 
space covered by vill and sometimes by ix. The gizzard is compressed from above, 
but widened laterally which makes it appear very much larger when viewed from 
above than when seen in vertical section. It connects posteriorly, in vii, with a very 
long narrow tubular intestine, which extends to somite xii, but which in somites xiii 
to xv is strongly nipped by the septa and considerably enlarged (fig. 86, s,) without, 
however, being strictly sacculated. 


The saceulated intestine proper, however, begins first in xvi, and is much wider 
than the other part of the alimentary canal. The structure of the pharynx offers 
nothing of unusual interest. The wall of the gizzard contains the usual layers, but they 
are poor in blood vessels. The longitudinal muscular layer is thickest, and on the 
widest part of the gizzard it is five or six times as thick as the epithelium, gradually 
diminishing in size anteriorly and posteriorly (figs. 86, 87, 88, 89). 


Pear-shaped or Chylus Chambers. Beginning immediately behind the gizzard, 
and extending throughout the tubular intestine, we find imbedded between the 
epithelial folds numerous pear-shaped organs of doubtful function. In pronouncing 


CALIFORNIA EUDRILID&, 49 


these as chylus chambers, I do so with much hesitation, as I am not at all satisfied but 
that instead of being organs of absorption, they may not in reality be organs of dis- 
charge, or in other words, glands. I found these chambers always empty, appearing 
entirely transparent in sections of the intestine, and they never stain, apparently 
showing a want of contents, though this may be a temporary condition, owing to 
the state of the worms when killed, at which time these organs may have happened 
to have been temporarily empty. As regards location, they are found principally in 
somites vii to xii, und seldom extend further back than that somite. It is to be re- 
membered that in somite xiii the tubular intestine changes its form and becomes 
considerably sacculated. In somites vii to xii this tubular part is very poor in blood 
vessels, or rather in large blood sinuses, while in xiii these sinuses begin to appear in 
large number and of large size. Thus, with no blood sinus in the epithelial folds, there 
are many pear-shaped chambers, while on the contrary, the cessation of pear-shaped 
chambers is accompanied by numerous and large blood sinuses (fig. 93). The pear- 
shaped chambers are imbedded between the epithelial cells, and probably all of them 
reach the inner cavity of the alimentary canal, though, from the sections made, this is 
not quite evident. Some of them, however, do, as will be seen from the figures 90 and 
91. They are unequally distributed; in some places they fairly crowd out the epithelial 
cells, as in fig. 91, which is drawn from a longitudinal section, showing an unusually 
large number. Fig. 92 represents a surface view where they are less numerous, 
and fig. 90 represents a camera drawing from a transverse section of the gut. Each 
chamber consists of six, seven, or, in some instances, of only two or three cells, arranged 
as the clefts in an orange, around a central pore or short tube, which, however, does not 
extend all through the papilla, but ends blindly. At the lower end these individual 
chambers connect with numerous smaller and more irregular cells, which join 
each other close to the transverse muscular layer. The small central pore stands 
frequently in connection with the alimentary cavity. In the smaller and inner 
chambers this pore is less pronounced or entirely wanting. At the base of the 
chambers are seen a number of small glandular masses, with grainy and opaque 
contents (fig. 90 g/.). As regards the distribution of the chambers among the epithelial 
folds of the same somite, it may be noted that they are principally numerous in the 
dorsal and ventral regions, disappearing in the lateral regions (fig. 133, ep. and ec. 7.) 
This is the case in all the somites where these chambers appear in the tubular intestine. 
Chylus canals have been described by Michaelsen as present in the intestine of En- 
chytreeus, but the difference between their structure and those of Argilophilus is 
quite great. In Enchytreeus these canals are principally intercellular, and connect 
with bloodvessels or sinuses, while the pear-shaped organs in Argilophilus are extra 
cellular, and occur in parts of the intestine especially poor in blood vessels. The im- 
possibility to procure fresh worms will necessitate a closer study of these organs, to 
be deferred to a more opportune time. . 


7 esl 5 =< . F Sy 5 - . le < 
Typhlosole. The typhlosole and typhlosolax region is small, and not especially 
pronounced in front of and in the clitellar somites. Anterior to somite xix the 


50 CALIFORNIA ACADEMY OF SCIENCES. 


typhlosolar vessel, which is entirely dorsal, is many times smaller than the longitudi- 
nal dorsal vessel, but posterior to that somite the typhlosolar vessel assumes a larger 
size, almost equalling the main dorsal vessel. In the anterior somites the typhlosolar 
region does not project down in the intestine, but is only somewhat wider than the 
balance of the intestinal wall. The typhlosole in this part appears to be filled with a 
fibrous and spongy mass, in which, however, I have seen no distinct nuclei (fig. 154, 
c.”.). These cavities are more oblong at the walls, more round or angular at the 
center, where several larger cavities are seen. At the upper margin of the typhlo- 
sole this fibrous body gives room to two large and several smaller longitudinal canals, 
one at each extremity of the typhlosole, all separated by muscular fibers or cell-like 
chambers. In fig. 136 the fibrous nature of the interior of the typhlosole is somewhat 
more pronounced than it should be, the fibers appearing rather more regular than in 
reality, though the appearance is always as if the majority of the fibers radiated in a 
fan-like way from the central spongy mass. This part of the posterior typhlosole is 
in cross-section deltoid, with the point projecting into the cavity of the intestine 
(136, ty). The size of the typhlosole varies with the individual specimen. 


Spermatheca (figs. 81 to 86). There are two pair of spermathece in viii and 
ix Opening in the intersegmental grooves between vii and viii and yili and ix. The 
spermthecal pores are found in front of and slightly outside of the second sete. As 
regards the size the spermathecze may be said to be very large, but unequal, as we 
seldom find two of the same size. One or two are generally developed at the 
expense of the others and fill all the available space in the somites, frequently pushing 
the septa into the nearest somites. Seen in a transverse section of a segment the 
larger spermatheca may be occupying as much as three-fourths of the cavity (fig. 86). 
The spermatheca consists of two distinct parts, of which the upper is by far the largest, 
rounded in outline or potato-like with comparatively thin walls (fig. 82). This part, 
though somewhat warty, carries no diverticula and there are no smaller cavities for 
the storing of the spermatozoa. The lower muscular part is twisted, and set obliquely 
to the former, but can in no way be said to form any kind of a diverticulum (figs. 81 
and 82). The muscular layer, which is a direct continuation of the longitudinal layer 
of the body-wall, is only arranged in one way forming a circular muscular stratum of 
the spermatheca much thicker at the base than at the top (fig. 82 /.m). This muscular 
layer extends to the upper part of the spermatheea (82 a), but is here quite narrow. 
The inner glandular layer is singularly well developed (figs. 82, g/. e.; 84 gl. ep. and 
85), projecting inwards in large folds like the epithelial folicles of the intestine. 

Secretions accumulate as a large whitish mass in the upper part of the sperm- 
atheca, and are seen to be sparingly mixed or streaked with spermatozoa. But 
the most characteristic part of the spermatheca is the interlacunary system for the 
storage of the spermatozoa in the lower or muscular part. A section of this part shows 
(figs. 82, 83 and 84, /. s.) a row of chambers imbedded between the epithelial cells, 
or between them and the muscular layer, and which connect more or less directly 
with the cavity of the spermatheca by means of narrow passages (fig. 83 p.) Some 


CALIFORNIA EUDRILID®. al 


of the chambers stand also in connection with each other as at 84 sp. b. and 83. 
In these chambers are stored the spermballs proper, one in each, rarely oceupy- 
ing the whole space of the chamber, but leaving considerable of the lacunary room 
empty, it never containing any free spermatozoa, only agglomerations or spermballs. 
These spermballs may also be seen as white opaque globules from the outside of the 
spermatheca as represented in figs. 81 A and L, sp. b. 

Testes (figs. 94, 95, 96.) There are two pair of testes, one in somite x, one in 
xi, as usual post-septal. The anterior testes are enclosed in the sperm-sac, the pos- 
terior ones are generally free. Cross-sections show the testes to be deeply multilobed, 
with the lobes spreading. As regard the enclosing of the anterior testes it may be 
remarked that it is more or less complete, evidently depending on the size of the 
sperm-sac. In some specimens the testes were entirely enclosed, in others only the 
posterior apex was invested in the sperm-sac. In most specimens the testes were 
found pressed close to the body-wall and projecting backwards, the point of adherence 
to the anterior septum being immediately above and adjoining the ciliated funnel of 
the nephridium in line with the ventral sete. 


Sperm-sacs. ‘There are three sperm-sacs, more or less but generally paired in 
somites x, xi and xii, the one in the latter somite being much more lobed than the 
anterior ones, and more frequently paired. The two anterior sperm-sacs are much 
the largest (fig. 96 and 97). They fill the whole somites, are not always paired, but 
the lobes connect all along the dorsal body-wall. The anterior sac closely invests the 
testes and ciliated rosettes in x and furthermore often encloses the ventral ganglion (fig. 
96 and 97). The sperm-sae in xi is also often closed on the dorsal side, but it does not 
invest the testes nor the sperm funnels of that somite (fig. 98). The sperm-sae in xii 
differs from the other by being deeply lobed in the plane of the trabecule which are 
arranged ina fan-like shape from the alimentary canal. This sperm-sac which is 
seldom paired, does not extend below the alimentary canal as the anterior ones do, 
but so to say rests entirely on the intestine (figs. 86 and 99). The anterior sperm-sacs 
are also traversed by numerous trabeculee, which however are irregularly arranged 
(fig. 100). The sperm-sacs are frequently infested with parasitic coceidixe of round 
shape and with from one to three germ cells (fig. 100, coc.) In size the sperm-sacs 
are so large that they push far backwards, the one pair covering the other, the 
whole mass often reaching as far back as to touch the prostate, thus entirely covering 
the ovary and oyiducts. This is especially evident in transverse sections when the 
ovary, oviduct and the three sperm-sacs may be seen at once. 

The ovary is, as usual, found in xiii and offers no great peculiarities. It is 
generally deeply lobed, and in cross-section shows the same projecting lobes as the 
testes. 


The oviduct is rather thin and characterized by a thick and long upper lip 
(fig. 103 uw. /.), which generally is bent to one side. In one instance the upper lip 
was forked (as in fig. 103, c.). The muscle attaching the upper part of the oviduct to 
the body-wall is unusually strong (fig. 103, m. s.). 


52 CALIFORNIA ACADEMY OF SCIENCES. 


Spermducts and prostates (figs. 105 to 115). There are two pair of sperm funnels 
and duets, the ducts joined together. The ciliated rosettes or sperm funnels are 
found in x and xi. The anterior pair is engaged in the sperm-sac of that somite 
(fig. 97), while the posterior pair is generally free (fig. 98). I have, however, seen 
the sperm-sacs attached to the funnels in these somites in some specimens, and there 
appears in this respect to be considerable variation; generally, however, the rosettes 
are free (lig. 98) in somite xi. Also with the anterior funnel there is some variation. 
In some specimens the funnels were entirely enclosed in the sperm-saes, in others the 
sperm-sac was merely attached to the free surface of the funnel, while in others again 
one-half of the funnel was imbedded in the sperm-sac, while the other half was free. 
Figs. 97 and 98 show cross-sections with the sperm funnels or ciliated rosettes free or 
imbedded. 


The spermducts join in xii and continue in a direct line to the prostate, which 
they enter in somite xviii. The point of junction is in the lower part of the glandular 
part close to the muscular duct (fig. 86, e.). The duct runs between the very thick 
vascular layer and the longitudinal muscular layer of the body (figs. 117 and 118, sp.). 
The two ducts are never fused together, but continue distinct and separate, though out- 
wardly joined, as to the very entrance in the muscular part of the prostate (figs. 112 
and 113). While the junction of the spermduct and the prostate is, to all appearances, 
in the glandular part of the prostate (figs. 110 and 113), the real point of entrance is 
in the muscular duct (fig. 115, spd.). After having touched the glandular part, the 
spermducts bend and eross the intervening space to the muscular prostate which they 
enter in a slanting direction, then passing considerably downward enclosed in the 
muscular part of the prostate, before entering the lumen proper (figs. 106, 107, 110, 
111, 112,113). Fig. 113 represents a cross-section in which the spermducts haye 
been cut twice. Part of the spermduct is seen free close to the glandular part of the 
prostate, part again is seen just at the fusion of the ducts with the lumen of the 
muscular part. The duct closest to the lumen has been partly differentiated, the cells 
having lost their nuclei. 


The prostate consists of a very large cylindrical, but greatly coiled, duct (figs. 
106 and 107 and 1), which generally lies pressed flat to the body wall of xviii (fig. 386, 
pr.) Tt opens outwardly in xviii in the posterior part of the somite in the same pore 
as the penial setze. 

There are two layers of cells in the glandular part, but apparently no muscular 
layer between them. The outermost layer consists of large glandular lobes containing 
glandular cells which pass between the inner cell layer, and discharge in the lumen of 
the prostate (see figs. 108 and 109). There is a large system of blood vessels which 
penetrate both of the cellular layers, but which is not developed to the same extent as 
in Deltania. 


In cross-section of the body-wall (fig. 103 A) the prostate is seen to open 
laterally to the penial setee, though in the same pore, situated at the very junction of 
the glandular clitellum and the ventral zone (v. p.) of the body (fig. 103 A.¢ ). 


1 
eX) 


CALIFORNIA EUDRILID#E. vd 


Vascular system (fig. 86 6 and 119). The following remarks on the vascular 
system can only be considered as preliminary, a more detailed report being reserved 
for a future study of living specimens. The main system consists of two longitudinal 
vessels, one dorsal and one ventral. There is no subneural vessel. Three pair of 
stout, oblong, thrice-contracted and sac-like hearts connect the ventral and dorsal ves- 
sels in x, xi and xii. The typhlosole has already been described. 

The secondary vessels are characterized by numerous bead-like constrictions 
and swellings of the smaller branches, especially of those surrounding the funnels of 
the nephridia, and of the secondary vessels of the ventral longitudinal vessel. 

Of the genus Argilophilus there are two rather distinct forms in California, which 
however, only differ externally, and, strange enough, have not, as a rule, been found in 
the same locality. Through careful dissection and sectioning I have not been able to 
distinguish any anatomical differences between the two species, though I have never 
seen any transitory forms as regards to the external markings. According to our 
present knowledge of earthworms we always expect to find internal specific charac- 
teristics of the species, and if such are not found we must hesitate to consider the re- 
spective forms as different species. Under such circumstances I will here refer to 
the two forms of Argilophilus only as varieties or subspecies, leaving to further inyes- 
tigations, if possible, to detect any internal differences. 


Argilophilus marmoratus ornatus. 
Figs. 125 to 129. 


Argilophilus marmoratus ornatus Eisen, Zoe, iv, 253, October, 1893. 


There are two rows of ventral papillae between some of the somites in the vi- 
cinity of the clitellum, interior or posterior to it, or both. These papille are always 
more or less in line with sete i, of rounded or slightly oblong form, and generally 
more or less of the same size. The number of papille varies, and frequently one 
papilla in a pair is wanting. 


Habitat. I have found this worm only in the vicinity of Santa Rosa, Sebasto- 
pol, etc., north of San Francisco Bay. It occurs there in rich heavy soils, and con- 
stitutes the most common earthworm of the district. In the vicinity of Sebastopol I 
found this subspecies both in manure piles and in the wet, soggy places near the la- 
goon, places which part of the year must be covered with water from the slough or 
lake. Among several hundred worms collected in the month of May during one day, 
I found only one single specimen with median papille; all the others possessed lateral 
papille (fig. 125 to 128). 


The body is thick, cylindrical, and only slightly tapering toward either end 
; y sugntly g 
fig. 152). As usual, the anterior segments are much wider than the post-clitellar 
, g l 
ones. The prostomium frequently protrudes like an inverted sac hanging over the 
peristomium (fig. 130), as is probably the case with most genera of this family. 
While the anterior somites are well set off, the clitellar somites are less distinct, 


54 CALIFORNIA ACADEMY OF SCIENCES. 


though they can always be made out. The clitellum comprises somites xiii to xviii, 
though the posterior + of xii is generally also somewhat thickened. Of the anti- 
clitellar somites vii, vill, ix are thicker than the others, and as thick as the somites 
of the clitellum. All the somites are deeply segmented, except the clitellar somites 
and those anterior to iii. The external pores are situated as follows: Spermathecal 
pores, two pairs, between yii/yili and yiii/ix, a little lateral of the seta 2. Oviducal 
pores in xiv in front of seta 1, generally in a slight depression, bordered by a swelling 
of the body-wall. The male pore and pore for penial setee combined in one, open in 
line with seta 2 in xviii. The male pore is characterized by being situated on an ele- 
vated papilla more or less oblong. This papilla is surrounded by a circular depression 
which encroaches on the two adjoining somites (xvii and xix), and this depression 
is again surrounded by a lunar ridge, open ventrally, and extending from the center 
of xvii to the center of xix, thus reaching slightly outside of the clitellum (fig. 129). 
The clitellar glands do not extend further ventrally than toa line running from the 
center of the lunar ridge parallel to the sete. The ventral part of the clitellum is 
depressed. 


Ventral papilla. One of the most interesting features of this genus are the inter- 
segmental glands occurring on the ventral side and which in our present species are 
paired. They have possibly the same general function as the tubercula pupertatis of 
the true Lumbricids, or the ventral papillae of Pericheeta, ete., which latter they 
greatly resemble, but unlike the former they are variable in position, sometimes oc- 
curring to the number of as many as seven pairs. In other specimens again they are 
reduced to one solitary tubercle, situated on one side of the ventral median longitud- 
inal line. The most common form is as follows: 

One pair between vili/ix. 
One pair between ix/x. 
One pair between xv/xvi. 
One pair between xvi/xvii. 
One pair between xix/xx. 


Other specimens possessed in addition to these pairs: 
One between xx/xxi. 
One between xxi/xxii. 


To show the variability of these papillee I give their places in a few individuals 
picked out at random, and I only add that other variations occur, as may be expected, 
from what is shown here. All these specimens were taken the same day and at the 
same locality : 

No.1. One pair, viii/ix. 
One pair, ix/x. 
One pair, xv/xvi. 
One pair, xvi/xvii. 
One pair, xix/xx. 


CALIFORNIA EUDRILID®. v0 


No. 2. One pair, ix/x. 
One single, xv/xvi, left side. 
No. 38. One pair, ix/x. 
One pair, x/xi. 
One single, xiv/xv, right side. 
One pair, xv/xvi. 
One pair, xvi/xvii. 
One pair, xx/xxi. 
One pair, xxi/xxii. 
No. 4. One pair, x/xi. 
One pair, xv/xvi. 
One pair, xix/xx. 
No. 5. One pair, x/xi. 
One pair, xix/xx. 
The structure of these papillze has been referred to in connection with the 
body-wall, and apparently does not differ in the two forms. 


Argilophilus marmoratus papillifer. 
Fig. 131, A and B. 
Argilophilus marmoratus papillifer Eisen, Zoe, iy, 253, October, 1893. 

The ventral side of the somites with one single median row of ventral papille, 
which are generally largest between the clitellar somites, diminishing gradually in 
size toward the anterior somites. The papill of this form are generally, but not al- 
ways, more oblong than in the preceding form, where they are much more rounded. 
The clitellar papille are frequently diamond-shaped (fig. 131). The papille vary 
greatly in number and size in various individuals, but they are always median, never 
paired. Sometimes there are 6 to 7 papille posterior of the clitelium. 


Habitat. This worm is very common in the vicinity of San Francisco Bay, 
south of Santa Rosa, where the former form begins. I have also this form from Santa 
Clara County, Monterey County, Fresno County, ete. It is common in the foothills 
of the Sierra Nevada, in Nevada County. Among many hundred specimens col- 
lected there was only one which possessed the paired papille of the former form. 


Systematic position. The peculiar variation of the nephridio-pores places 
Argilophilus in undoubted proximity to Plutellus, both as described by Perrier and 
Benham. ‘The extra chetal pores in Argilophilus warrants however the forma- 
tion of a new genus, even if no other important characteristics would help to make it 
yet more distinct. As is well-known, Perrier’s description of the ovaries, ete., in 
Plutellus have always been considered doubtful, and by Benham have been shown to 
be incorrect. This of course only in the case that Benham’s worm really belongs to 
the same genus as the one described by Perrier. I do not doubt that this is so, be- 
cause the positions assigned to the ovaries by Perrier is so abnormal that it is more 


56 CALIFORNIA ACADEMY OF SCIENCES. 


than likely that a mistake was made, as Benham has already pointed out. But on the 
other hand Benham’s worm came from Queen Charlotte’s Islands in the Pacific, while 
Perrier’s worm came from the Atlantic Coast, many thousand miles away, and it will 
always be impossible to with certainty decide upon the position of the ovaries in 
Perrier’s Plutellus until the original worm has again been investigated. That it will 
be found to entirely agree with the normal type I do not doubt. 

From Plutellus our Argilophilus differs in several important points besides the 
location of the nephridio-pores. First, in regard to the number of the ciliated rosettes— 
in Plutellus only one pair, in Argilophilus two pair. One pair of sperm-saes in Plu- 
tellus, three pair in Argilophilus. One pair of testes in Plutellus, two pair in Argilo- 
philus. Four pair of small spermatheca in Plutellus, two very large ones in Argilophilus. 
Perfect clitellum in Plutellus in at least two somites, ventrally imperfect clitellum in 
Argilophilus. In other respects the two genera resemble each other greatly, and 
through the respective arrangement of the nephridio-pores, they form a distinct tribe 
or sub-family for which I propose the name Plute//ini. 


The following table will show the principal characteristics of the two genera: 


Argilophilus. Plutellus. 
Prostomium dovyetailing the peri- 
BEONUUU oie hate ee ieee etre Partly. Completely. 
(UNA DeS eo rabbit dGonDonenS pees Ventrally imperfect. Ventrally perfect in at least two 
somites. 
Nephridio-Pores.. 2a. e/ ein ...| In front of sets 3, 4, and outside of 4; gen-| In front of 3 and 4, alternating 


erally the latter, the arrangement being) rather regularly. 
irregular, 


GigzOG 1M yin: sede esas oR aoe v. vi. 
SDENMONLECE: pee giracn she mene a hete Two very large pair in yii and viii. Four or five pair in vi, vii, viii 
and ix. 
Sperm-funnel . .......-....-.-+-«.| LWO pair in x and x1. One pair in x. 
LeBLES tral toranisy ie stern ese A Two pair in x and xi. One pair in x. 
ISDORIM-SACES che ee-oncesieisienel eda cia Three pair in x, xi, xii. One pair in xi. 
Ovary and oviduct im ........... xiii. xiii. 
Onipore its Wye ein tee tics ert xiv. | xiv. 
Male pore oh a ee eed Fees SPR 5 | xvlil. xviii. 
lea . 
POSLGER ce eis Pe cals ee Paceline Very large, coiled, extending through several) Straight, tubular, not very large, 
somites. confined to one somite. 
| 
r 30 = ee Pee3 
Nephridia commence in.......... rin ili. 


Nephridral bladder at the entrance) Absent, the duct being single and straight. | Large, but of varying size in the 
toy the bodiy-wall < ose.) eee aye two series of nephridia. 
Ventral papille on the body-wall. | Present in pairs or in a single one. Absent. 


Copulatony, S0be. 6.6.5 2. ncn cons | Present. Absent. 


Ty RLOS G08. war stal See eaten eae cont ee | A small typhlosole present. | No typhlosole. 


CALIFORNIA EUDRILID®. Dia 


PAPERS REFERRED TO. 


Benuam, W. Biaxtanp. An Attempt to Classify Earthworms. Quarterly Journal of Microscopical Science, vol. 
xxxi, pt. ii. 

Benuam, W. Briaxtanp. The Nephridium of Lumbricus, ete. Quarterly Journal of Microscopical Science, xxx, 
ii, 1891, 293. 

Bennam, W. Braxianp. Description of Three New Species of Earthworms. Proceedings of the Zoological Society of 
London, Feb. 16, 1892. 

Bepparp, Frank HE. The Classification and Distribution of Earthworms. Proceedings of the Royal Physical 
Society, Edinb., 1891. 

Bepparp, Frank E. On the Earthworms collected in Algeria and Tunisia. Proceedings of the Zoological Society 
of London, January 5, 1892. 

Brpparp, Frank E. Some New or Little Known Oligochewta. Proceedings of the Royal Physical Society, Edinb., 
June 13. 1893. 

Bourn, A.G. The Nephridia of Leaches. Quarterly Journal of Microscopical Science, vol. xxxiy, 592. 


Eisen Gustav. California Earthworms of the family of Eudrilidw. Zoe, a Biological Journal, San Francisco, Cali- 
fornia, vol. iv, No. 3, 1893, pp. 248-254. 
Fretcner, J.J. Notes on Australian Earthworms. Proceedings Linnean Society of New South Wales, vii, 374. 


MICHAELSEN, W. Oligochwten von Stid Georgien, Jahrbuch der Wissenschaftlichen Anstalten zu Hamburg, 1887. 
Hamburg, 1888. 

Rosa, Dr. DanteLe. I Terricoli Argentini, ete. Annali del Museo Civico di Storia Naturale de Genoa, ser. 2, 
vol, ix, 509, 1890. 

Rosa, Dr. DANIELE. Sui Generi Pontodrilus, Microscolex, Photodrilus. Bol. Musei, ete., di Torino, No. 39, vol. 
iii, 1888. 


Rosa, Dr. DANTELE. Microscolex Modestus. Bol. Musei, ete., di Torino, No. 19, vol. ii, 1887. 


EXPLANATION OF THE FIGURES. 


DELTANIA ELEGANS, Figs. 1-20. 


1. A medium sized worm, natural size. 
2. The anterior part magnified, seen from the yentral side, to show arrangement of sets and the exterior pores of 


various organs. p.n.p. the first three pepto-nephridio-pores in somites ii, iii and iy. mn. p. nephridio-pores 
of the following somites, opening in front of the 3d seta, the pepto n. p. opening in front of the 4th sete. 
0.p. ovipore. s. pd. p. spermiducal pore. 
In order to show the outer sets, the body has been slightly flattened out, In reality the outer setw should be 
slightly more dorsal than what appears. 


3. The prostomium with somites i and ii. 
4. Schematic view of the arrangement of the sets on both sides of the male pore. The italic numerals indicating 


the order of the sete, the roman numerals indicating the number of the somites, 

5. The interior of the yentral surface of the genital somites viewed from above, showing the arrangement of 
the generative organs, etc. spth. spermathecsw. c. 7. ciliated rosettes. sp.s. sperm-sacs. sp.d. spermducts. 
ov. ovary. od. oviduct. p.s. penial sete and sacs. pr. prostate. v.gl. ventralganglion, ¢. testes. 

6. Longitudinal section of the anterior part of the body. The posterior section is a little more eccentric than 
the anterior. br. brain. si. gl. salivary glands. s.gl. septal glands. pha. pharynx. s. pla. sigmoid plexus 
of the esophagus. sp.s. sperm-sacs. h. hearts. dv. dorsal vessel. ¢.7. tubular intestine. s. 7. sacculated 
intestine. pr. prostate. s. ps. sacs with penial sete. v.gl. ventral ganglion. «s. esophagus.  spth. 
spermatheca. ¢. testes. c. 7. ciliated rosettes. ov. ovary. od. oviduct. 

8. Prostate and male apparatus. pr. prostate. sp.d.spermduct. p.s. penial setw and sacs. ms. longitu- 
dinal muscular layer of the body wall. 

9. Oviduet in xiy and xiii. 


26: 


30. 


35. 


42. 


47. 
48. 


CALIFORNIA ACADEMY OF SCIENCES. 


One of the septal glands in somites vi, vil, viii and ix. 

Ciliated rosettes. 

Ovary. 

Spermatheca. - 

Exterior male-pore and penial papilla. 4 male-pore. p.s. penial sete and papilla. 
Exterior view of the clitellum seen from the ventral side. op. ovipore. ¢ male and prostate pore. 
One of the regular sete. 

Penial set from somite xvii. 

Testis from somite x. 

Nephridium from somite xv. 

Anterior nephridium from somite ii. 

A nephridio-stome from one of the posterior nephridia. 


DELTANIA TROYERI, Figs. 21-38. 


Diagramatic view of the regular arrangement of the sets in several somites posterior to the clitellum. 

A medium size worm. 

The largest specimen. 

The anterior part seen from above, showing prostomium and somites i, ii, iii. 

Schematic view of the arrangement of the inner sete in the region on both sides of the male-pore. ovp. ovi- 
pore. p.s. penial sete. ¢ male-pore. 

The outside of the male genital region. f. folds inthe body-wall. p.s. penial seta. p. p. penial sete papilla. 
pr. and ¢ prostate and male-pore. c.d. cupshaped concave region or sucker near the male-pore. c. c. 
central cavity between the male-pores. m. st. place for missing set# No.1. s. st. seta No.2. p.m. penial 
muscles, arranged fanshaped, connecting the male-pore with the intersegmental groove. 

Longitudinal section of the anterior part of the body. 6b. c. buccal cavity. phnx. pharynx. sl. gl. salivary or 
pharyngeal glands. br. brain. s. gl. septal glands. ss. ple. sigmoid plexus of the cesophagus. sp. s. sperm- 
sacs. «es. cesophagus. ¢. testes. spih. spermatheca. 

Somewhat schematic view of the alimentary canal seen from above. es. cesophagus and pharynx. sl. gl. 
salivary gland. s. gl. septal gland. d.v. dorsal vessel. A. hearts. ¢. 7. tubular intestine. s.7. sacculated 
intestine. 

Brain and ventral ganglion. 

One pair of the septal glands (vi). 

Spermatheca from ix. Side view. 

Spermatheca seen from below. 

Spermatheca seen from above. 

Male apparatus. pr. prostate. p.s. penial sacs andsets#. sp. d. spermduct. ms. muscles of the body-wall. 

Nephridium from somite xv. 

Oviduct and oyisae, left side. 

The same from the same individual, but from the right hand side of the ventral ganglion. 

The same sac seen from above. 

Detail of the same. 

Arrangement of the set surrounding the spermathecal pores. 


DELTANIA BENHAMI, Figs. 40-48. 


Arrangement of the sete surrounding the spermathecal and male-pores. The same proportion and scale as in 
the preceding figure. 

Sperm-sac. 

Spermatheea, left side. 

Spermatheca, right side of the same individual. 

Oviduct. 

Prostate gland and penial sete. v.v. blood vessel covering the prostate. 

Alimentary canal. pha. pharynx. sil. gl. salivary glands. ss. gl. septal gland. as. esophagus. s. 7. saceu- 
lated intestine. c.i. contracted part of the intestine generally commencing somewhere between somites xxvii 
to xxxvi and continuing towards the tail. In the first six somites the contraction is generally much narrower 
than in the following. 

The worm natural size. 

The male papille on somite xvii, showing the projecting penial set. The upper shaded part is the clitellum 
proper which ends at the center of the somite in line with the male-pore. 


CALIFORNIA EUDRILID®. 59 


DELTANIA ELEGANS, Figs. 49-56. 


49. The lumen and cecal part of the nephridium in transverse section to the body. fl. c. e. flask-like glandular 
cells of clitellum. 5/. v. blood-vessels in the transverse muscular layer. ¢. m. transverse or circular museu- 
lar layer. /. m. longitudinal muscular layer. J. x. lumen of ceecal part of the nephridium. i. /. interior 
lining of the nephridium. g/. glandular outer layer of nephridium. cal. calculi projected by the nephridium, 
gl.c. glandular cells of the clitellum. atr. urinary bladder or large contractile (?) chamber near the exterior 
pore of the nephridium. ms. muscular layer of the nephidial duct joining the flask-like cells of the clitel- 
lum. col. collar of the nephridium forming the exterior pore and consisting of regular layers of large, almost 
rectangular cells. 

50A. Section through the body-wall and the lower part of a nephridium in somite xviii, but posterior to the clitellum. 
n.d. nephridio-duct. ¢.n. cocal nephridio-bladder. /. m. longitudinal muscles. ¢. m. circular muscles. 
ep. epidermis and cuticle. 

50B. Section through the body-wall and the nephridial collar in a somite posterior to the clitellum, illustrating the 
large ganglion (?), below the nephridial collar. The section shows the beginning of the nephridial collar and 
the place where it joins the cacal vesicle. 

50C. Section closely following the former nearer the nephridio-pore, showing a larger part of the ganglion. i. c. 
interior chamber or tubule, probably opening into the nephridial cecum. e.). inner lining of the cecal 
bladder. m. 6. muscular layer of the bladder. c. np. collar of nephridium. xn. g/l. nerve ganglion. /. m. 
longitudinal muscular layer of the body-wall. 4. m. transyerse layer of the body-wall. 4. hypodermis. 

51. Lower part of the prostate and penial sac, section through the male-pore, showing the junction of one of the 
penial sacs with the prostate. p.s.s. penis sac with sete. p.s. one of the penial setw broken. pr. 
and ps. prostate and male-pore. pr. d. prostate duct. 

52. Section through the male-pore, cross-section through the body-wall, showing junction of the two spermducts, 
and prostate, and one of the penis sacs. sp. d. spermducts. j. sp. junction of spermducts. pr. prostate. 
j. sp. and pr. junction spermduct and prostate. s. ps. sac of penial setw. j. pr. and p.s. junction of prostate 
and penal set. 4 male-pore. ep. epidermis. c. m. layer of circular muscles, body-wall. Jl. m. layer of 
longitudinal muscles, body-wall. x. c. nerve cord from ventral ganglion. p. p. penial papilla. 

53. Section through the male-pore showing junction of the two penis sacs with the prostate. ps. the two 
branches of the penis sac with the penis sheaths and penial setw. pr. prostate. c. m. circular muscles. 
l,m. longitudinal muscles. 4 male-pore. atr. atrium, or junction between the penis sacs and the prostate. 

54. Section through the muscular part of the prostate. gl. inner glandular layer. m. outer muscular layer. 
blw. blood-vessels and cuticle. 

55. Section through upper end of prostate. b/v. bloodvessels. c.v. vessels between the inner and outer layer of 
cells. i. v. inner capillary vessels, lining the inner surface of the prostate. 

56. Section through prostate at the junction of the muscular and glandular part. m. pr. muscular prostate; other 
letters same as in Fig. 54. 

57A. Section through intestine in somite xy. ch./. chloragogic layer. /. m./. longitudinal muscular layer. ». /. 
vascular layer. lv. blood-vessel of the epithelium. ep. epithelial layer. 

57B. Section through intestine, somite xiv. 

57C. Section through intestine, somite xvi. 

58. Cross-section through the clitellum, showing the relative size of the various layers. 


ARGILOPHILUS MARMORATUS ORNATUS, Figs. 59-130. 


59, A nephridium from one of the somites posterior to the clitellum. ~* 0, inner orifice or nephridio-stome engaged 
in the muscular dissepiment d. 1, narrow duct connecting the funnel with the fold. p. f. posterior 
fold. a. /. anterior fold. wu. upper part of the spongy duct, which at 15 stands in direct connection with 
the narrow tube 1. /. lower part of the spongy duct gradually increasing in size towards 5, a. upper main 
canal. 6. lower main canal, both the upper and the lower being different ends of the same canal. o. c.f. 
point of recurrence. jp.c. peritoneal cell covering or support of the nephridium. n. p. nephridio-pore. 
c. t. capillary tubules of the spongy duct. 6/. blood-vessels. cap. capillary blood-vessels surrounding the 
canals aand b. 1 to 17 mark places from which the detail drawings are taken. The nephridium is longer in 
proportion to its thickness than what is shown in the figure. If the part between the two parallel lines at 
18 was increased to about one-third or one-fourth the entire length of the nephridium the proportion would 
be correct. If the whole nephridium had been drawn the figure would have been too long for the plate. 

60. Nephridio-stome, front view. 

61. Nephridio-stome, side view. 

62. Nephridio-stome, end view. 


* Letters and figures in full-face type refer to those in rings on the plates. 


60 


63. 


64. 


66. 
67. 


69. 


71. 


83. 


84. 


CALIFORNIA ACADEMY OF SCIENCES. 


Inner part of nephridium more highly magnified. The numerals indicate the same places as in Fig. 59. The 
letters are the same as in the above figure. 

Part of the same more highly magnified in order to show the capillaries and their connection with the inner 
duct of the tube. ¢./. tube leading to nephridio-stome. a. upper canal. b. lowercanal. 7. /. innerlumen 
of the duct which connects at c. with the capillaries. c. ¢. capillary tubes, longitudinal view. ¢.¢.¢. capil- 
lary tubes seen in cross-section. sp.c. spongy canal or tube directly connecting with the duct 1 and con- 
taining the capillaries. o. c. outer canal leading to the nephridio-pore. c. br. the narrow bridge connecting 
the two main folds. p.c. point where ciliation begins. b/. v. blood-vessels, connecting with capillary 
blood-vessels surrounding the two inner canals a and}; only a part of the blood-vessels are drawn. This 
figure was carefully drawn with camera lucida, though, on account of the minuteness and numbers of the 
ductules c. tc. and c. ¢., it was impossible to delineate all, almost one-fifth having been left out in places where 
they crowded each other. 

View of the spongy duct between 3 and 4 showing the inner canals a and 6 and the connecting ductules; 
letters indicate the same as in previous figures. 

The same spongy duct near 4, end view, and partly cross section. 

The posterior fold as seen at 5, the crosses + and X indicate the respective point corresponding on fig. 59, 
in order to show the direction of the tube, which is contrary to the one shown on fig. 59. sp. ¢. the spongy 
tube with the upper and lower ducts, connected by ductules c.¢. 6/. bloodvessels. cap. capillary blood - 
vessels, some of which surround the inner canals, others the outer fold. At the place marked with a x the 
ductules begin to connect the upper spongy tube with the lower main canal instead of with the lower part 
of the spongy tube, as atc. ¢. 

A somewhat larger figure of the last taken at the end x, where the lower part of the spongy tube has almost 
vanished and become bereft of its tubules c.¢., capillary tubes entering canal b. c. chambers on the central 
canal in the spongy tube, which send off tubules on the other side of the two main canals, and which also 
enter the canal. 6.1. lumenof the spongy tube. In this and the previous figure will be seen how the tubules 
of the spongy tube shown in figs. 65and 66 have been found forming a more regular lumen /, which in the 
next figure will be seen to emerge into or change to a regular canal. ; 

Part of the upper fold at 7, showing the tubules of the spongy tube to cease, the tube itself to become more 
regular, fusing itself in canalb. The arrows show the direction of the fluid or excretions from the nepridio- 
stome to the pore. 

Diagramatic section of the main fold at the place where the capillaries of the spongy mass have ceased, and at 
the very point of the outer band marked fig. 59 0. c.f. The spongy mass surrounding the canals is now void 
of tubules. 

View of the anterior fold at 10, showing the inner ciliated canal—formerly the lumen of the spongy tube— 
and the two canals a and b, which are enclosed by a common glandular mass, a continuation of the spongy 
tube. The lower fan-shaped lines are possibly openings or valves in the tube, connecting with the glandular 
mass. As will be seen, the glandular mass is not continuous, but seemingly separated by lumens—/—which 
may prove to be canals connecting with the tube at the slits above referred to. 

A part of same fold between 9 and 10, The lumen is here void of cilia, and the glandulous mass is smaller 

The main fold as seen at the narrow place 9, /. lumen yoid of cilia. a. one of the canals @ or b seen on the 
other side of the lumen. O6/. v. bloodvessels surrounding the lumen and its glandular walls. 

Cross section of the posterior main fold taken at 3, before the main central lumen in the spongy duct was 
formed. 
Cross section of the posterior duct at 7, showing the blind ending of the tubules. 

A longitudinal section of the fold at 5, showing only one of the central canals. 

The same, the under side of the lumen being in focus. 

A longitudinal section near 3; here also one of the canals has been cut away. 

Peritoneal cells of the nephridial supporting covering from p. c. fig. 59. 

The same, end view. p.c. masses of perigastric cells.. 

One of the perigastric cells more highly magnified. 

One of the spermathece, exterior view. 

Cross section of a spermatheca. a. upper sac-like part. 6. lower muscular part. m. muscular layer of the 
sac-like part. /. m. longitudinal muscular layer of the body-wall. c. m. circular muscular layer of the 
spermatheca. gl. ep. glandular epithelium. 7. inner cavity of the spermatheca. sp. hb. sperm balls. /. s. 
lacunary system, or chambers for the storage of the spermatozoa. s. m. secreted mass with streaks of sperm- 
atozoa. spz. spermatozoa. ¢.m. transverse muscular layer of the body-wall. spth. p. spermathecal-pore. 

A longitudinal section of the spermatheca more highly magnified, showing the lacunary system and its position 
to the epithelial cells. Letters as in the last figure. 

Cross section of the muscular part of the spermatheca, showing the arrangement of the storage chambers oc- 
cupied by sperm balls, Letters as in fig. 83. 


85. 
SGA. 


S6B. 


100. 
101. 
102. 


CALIFORNIA EUDRILID®. 61 


One of the sperm balls more highly magnified. Of these never more than one occupies the same chamber. 

A somewhat diagramatie view of the anterior somite, composed from several sections, in order to show the lo- 
cation of the organs, theirrelative size, ete. c/. clitellum. br. brain. c. lower commissure of ventral ganglion. 
phx. pharynx. si. gl. salivary glands. w. esophagus. gz. gizzard. ¢.7. tubular intestine. s.7. sacculated 
intestine. nph. cross section of the anterior nephridia in somites ii, iii, iv. The other nephridia are not 
shown. sph. spermathece. sps. sperm-sacs. /. sps. lobed sperm-sacs in xii. ov. ovary. ovd. oviduct. 
c. r. ciliated rosettes. sp. d. spermduct; the connection with the ciliated rosettes not shown. ¢ male 
pore. pr. prostate. p.s. sacs with penial sete opening in the same pore, but more ventrally than the pros- 
tate. e. the place where the spermduct enters the prostate. 

The body opened from above and the interior organs exposed to view. A partially diagramatic view. The 
dotted lines indicate the organs covered by others, and which could not be seen except by remoyal of overly- 
ing organs. Letters indicate as in the preceding figure. 

Diagramatic longitudinal section of gizzard. 

Diagramatic transverse section of gizzard, to show its compressed shape. 

A part of the epithelium and muscular layer of the gizzard seen in longitudinal section. ep. epidermis. cpt. 
epithelium. /. m. longitudinal muscles. 

A transverse section of the gizzard. (Letters as in fig. 88.) 

A transverse section of the alimentary canal adjoining the gizzard posteriorly. ¢. m. transverse muscles. 
1. m. longitudinal muscles. ep. epithelium cells. chy. supposed chylus chambers. c.c. chloragogie cells. 
gl. glands. 6/. bloodvessels. 

The same in longitudinal section. (Letters as in fig. 90.) 

Surface view of the same, showing the arrangement of the chylus chambers. (Camera drawing.) 

Section of a part of the dorsal vessel in somite xvi. m. muscular layer. b/. blood clot. c.c¢. chloragogic 
cells. 

Section through the alimentary canal in somite xiv. 

One of the testes. 

Right testis in cross section. 

Left testis in the same somite xi in cross section. 

Ovary. 

Section of the body in somite x, through testes. 

Section of the body in somite x through the sperm funuels, very close to the septum. 

Section through somite xi. 

Section through somite xii. sps. a. sperm sac in x. sps. a. sperm sac in xi. sps. ai. sperm sac in xii. 
m. longitudinal muscles connecting various organs with the body-wall. 7. intestine. d.v. dorsal vessel. 
v.v. ventral vessel. A. hearts. ¢. testes. v. gl. ventral ganglion. s. septum. 

Part of the sperm-sac. coc. parasitic coccidie. 

One of the sperm-sacs separated. 

Oviduct a and b, front view. ¢. side view of an abnormal oviduct with two, instead of one, upper lips. w./. 
upper lip. s. septum. m. muscular band. 


103A. Cross section of the body-wall through the male-pore, showing sections of prostate and penial sete. pr. gl. 


sections of glandular part of prostate. pr. m. muscular part of prostate. p. s. penial sete and their sacs. 
v. vascular layer of the body-wall. J. m. longitudinal layer of muscles. ¢. m. transverse layer of muscles. 
c. gl. glandular layer of clitellum. fh. hypodermis. yg. ganglion. v. p. ventral part of the worm. 


103B. Detail of former, nuclei in the muscular part of the prostate. 


104. 
105. 
106. 


107. 
108. 


109. 
110, 


114. 


One of the ciliated funnels freed from the attached sperm-sacs. 

Diagramatic section through prostate, showing relative size of spermducts where they join the prostate. 

Diagramatic representation of the prostate, showing its natural position when the body-wall is spread out. 
pr. glandular prostate. spd. spermduct. p.s. penis-sacs. m. pr. muscular prostate. v./. vascular 
layer. v. ventral side. 

Detail of the above, to show junction of spermduct and prostate. 

Cross section of prostate. 61. v. blood vessels. o. gl. outer glands or prostate proper. i. gl. inner gland 
or atrium. 

Detail of the former, showing the way the outer glands open into the atrium between the inner cell layer. 

111, 112, 113. Successive sections through prostate and spermduet, illustrating the junction of the two. In 
the first figure the spermducts have not yet entered the muscular part of the prostate. In the three follow- 
ing figures this junction is progressing, and finally almost accomplished. 7. m. /. inner epithelium of the 
muscular part of the prostate. pr. glandular prostate. spd. spermduct. 0/. v. blood yssels. m. /. 
muscular layer of the spermduct. In 110 the spermduct has been sectioned twice, as it is here convoluted. 

Cross section through the body-wall in somite xix. v./. vascularlayer. J. m. longitudinal layer of muscles. 
t. m. transverse muscles. h. hypodermis. 


62 CALIFORNIA ACADEMY OF SCIENCES. 


115. Cross section through clitellum. gl. glandular cells with fine secretions. c. gl. glands from the hypodermis 
projecting into the former layer. /. v. blood vessels. 

116. Detail of the former, illustrating the hypodermal glands, ete. 

117. Cross section through body-wall, somite xii. Letters same as above, illustrating the arrangement of the 
longitudinal muscles. spd. spermduct. 

118. Cross section through body-wall somite xvii, illustrating the transverse muscles. 

119. Somite xiv laid open, illustrating the diagonal muscular band. ovd. oviduct. s. septum xiii /xiv. ovp. ovi- 
pore. 1.2.3.4. sete. m. 1.2. 3. diagonal muscular band, beginning and ending on the body-wall. 
m. 4. muscular band connecting the oviduct and septum with the body-wall. 2x. /. nephridio-funnel. 
is. g. intersegmental groove. v. v. ventral blood-vessel. . ventral part of the hearts. /. v. lateral vessel 
spreading on the body-wall and supplying the nephridium. vy. g. ventral ganglion. 

120A. Cross section through one of the ventral papilla of Argilophilus marmoratus papillifer. ¢. ¢. tubular cells 
butting on the central glandular masses. c. gl. supporting and protecting cells, with coarse grainy con- 
tents. A. common supporting cells of the hypodermal layer. J/. m. longitudinal muscles. ¢. m. trans- 
verse muscles. v./. inner vascular layer. nf. nerve ganglions, connecting the glandular masses with the 
ventral nerve ganglion. o. glandular papillw with central nerve ganglia. 

120B. The corresponding organ, or ventral papilla in Argilophilus marmoratus ornatus; letters indicate the same. 

121A. Detail of the last figure showing the most lateral of the glandular masses. o. glandular masses with a 

central nerve ganglion. 

121B. Detail of the same figure showing the most centrally located glands. 

122. One of the common set. 

123A. One of the penial sete. 

123B. The tips of the former more highly magnified. 

124. A part of the body-wall viewed from the inner side. A semi-diagramatic figure illustrating the arrangement 
of the setze and the nephridio-pores over a certain space. J, 2, 3, 4, the various rows of sete. np. p. 
nephridio-pores, indicated by round circles. Two of the nephridiaare drawn in order to show their position, 
as well as the relative size of the ducts. 

125, 126, 127, 128. The anterior part of the body seen from the ventral side. Various specimens to show the 
yariation in position and number of the ventral papille. spth. spermathecal-pores. ovd. oviducts. 
¢ male-pores. The ovals indicate the ventral papille. 

129A. The clitellar and some of the anterior somites, viewed from the ventral side, showing the various pores and 
the genital zone. Letters indicate the same as in the preceding figures. p.s. penial sete protruding 
from the male-pores. 

129B. The twenty-three anterior somites of a specimen killed in alcohol gradually increased in strength and hardened 
in strong alcohol, showing the spermathecal, 9 and 4 pores, the ventral papille, etc. 

129C. The most anterior somites seen from above. 

130. The two and three anterior somites of two different specimens, showing the inverted buccalic sac, the prosto- 
mium, ete. a. and c. ventral; c. d. and f. dorsal; ¢. lateral view. 


ARGILOPHILUS MARMORATUS PAPILLIFER, Figs. 131-136. 


131. The anterior somites viewed from below showing the single row of ventral papillew, diminishing in size an- 
teriorly. The outlines at the side represent the shape of the papillw as found in the figured specimen. 

132A. A medium size specimen preserved in strong alcohol. The specimen was first killed in an increasing solu- 
tion of corrosive sublimate. 

132B. A large specimen of the former figured while alive. 

133. Section of the intestine and dorsal vessel in somite ix, in order to show the typhlosolar vessel. t.v. typhlo- 
solar vessel in the intestine. d.v. dorsal longitudinal vessel. ch. c. chloragogic cells of the intestine. 
€.p. epithelial folds of the inner lining, void of chylus glands. c¢.v. the chylus region of the inner 
epethelial lining. 

134. The typhlosolar region of the intestine, detail from the last figure more highly magnified. ¢. v. typhlosolar 
lining. /. m. longitudinal muscles of the intestine. c. m. circular muscular layer of the same. Other 
letters as in the last figure. 

135. Cross section of the intestine and dorsal vessel in somite xxiv illustrating the typhlosole. ss. 7. sacculated 
intestine. O/.s, blood-sinuses. ty. typhlosole. 

136. Detail of the former, showing the typhlosole proper, greatly magnified. d.v. dorsal vessel. ty. typhlosole. 
bl. v. blood sinuses in the intestine. /.m. longitudinal muscles. c¢.m.and e.p. circular muscles and 
epithelial lining of the intestine. 


MEM. CAL. ACAD. Il. — PLATE XI 


tS a ee 


: 7K , ’ q , , 
A et a | & 


a, 


i Doe 


; ETH BASTTONS REY 5. 
DELTANIA ELEGANS Fic’s. 1 rol. 

2: 7 

Th 


vA > 
ss ae 
ba kd 


-MEM. CAL. ACAD. Il.. PLATE ir 


‘UTH, BRITTO GREY SF. 


IA ELEGANS FG's. _12 To 20. 
- % & . 
ee i : a ° 
LS eT, _ 


Aaa 
vy \ 
< al 


iN) 
ed 


We NT 


UTM. BRITTON SRET S.F. 


DELUTANIA TROYERI Fie’S.-25 ro 34. J d 


PLATE (XV 


ee a 


-MEM. CAL. ACAD. II. 


Vil 


1x 


39. 


Oe fae be 
| 
= 
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ea lS qi 
} 
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AS le 


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of 


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hrex 


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OTH. BRITTONS REY SF, 


149 TO 48.. 


7 


a 
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Leal 
fy 
a 
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SI 
t 7 
a 
be 
a | 
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ITH. BAITTONS REY 3.F. 


DELTANIA ELEGANS FiIe’s.49 & 50. 
& = 


UTM .BAITTON SREY &.F 
GUSTAV EVSEN, DEL.. 


DELTANIA ELEGANS: FiG’s. _51 to 58 


e 


as ei ae 


-MEM.CAL.ACAD. Il.. PLATE XVI 


.. GUSTAV EISEN. DEL: : s LTH. BRITTONEREY 5.7. 


; ARGILOPHILUS MARMORATUS ' ORNATUS “Fia's. _59 ro 63 


es E 
s : 
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ARGILOPHILUS MARMORATUS ORNATUS 


ne 


GUSTAV EISEN, DEL. - 


. MEM. CAL.ACAD. II. - PLATE XX 


BIN BAITTONS MEY =F, 


ARGILOPHILUS ~ MARMORATUS ORNATUS FIGs. 78 Tro 85 
- 7 Q d a 


‘ < 
© 
© 4 
3 S : 
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f tim a = > ~ 4 “gp 
shige ues Dt pee = ted 
~~ z f f, } 
Ne — Seno a ; 
A a / | an) 
= \ A f 
i jj } 
lat / 
) Ag Ve { 
re ra 4 TE 
Y oe Ae yer pi 
* z -. ta . 


87.A. 
88. 


yh 


89. 


PS = 
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A Ko 


pr 


Lees 


S 


q 


a 


- 
ae 


CH, 


ate Cava 91. 


: S4A. 


94B. E 


ra g a = 
=) 2 : 2 
led \ 
- . wh 
et ‘ nw 95B 
\ Y 


GUSTAV EISEN, DEL. LTN.BAITTONS REY SF 


ARGILOPHILUS MARMORATUS ,ORNATUS Fia@'s. 90 To 95. 


spsXl ----- 


3 : 
ee, , st 97. 
-s 100. 
% Gy 
= 5 
wot 
98 
| @ 
\) | 
\ 
ssh) 4: \)\ y] Hf} 
{ — Sg 101. 
\ \ Vis Pr: } / 
\ ‘ ) =| / ; 
\ AN \ ( j y, \ 
oN . y SA | 
Sah SZ 
Ce ae ae J nga XI 
~ \ as 
Se ) } pee 
} eee od See ee k 
Fa =>) XI a eS 
3 } tad Ss : 
BUSTAVY EVSEM. DEL. 
~= - Tre > = +) 
ARGILOPHILUS Cc FIG TO 101, 


UTH.SAITTONS REY 5.P, 


_ ARGILOPHIRUS MARMORATUS ORNATUS Fra's. .102 TO 107. 


aay ee 


aTE a. ; ~ s3 


au 


ra) 


14. 


XIX 


- GUSTAV ELS 


115. 
>» w&S 
117. 
a ‘F / 


Ay 118. 


z ' ¥ LTM.BAITTON & 
: : : Ze oni mt 

. a. Vg he j 7 f 

GUSTAV EISEN. DEL, - - 


ARGILOPHILUS 


PLATE XXVII 


-MEM. CAL. ACAD. II. - 


UT BAITTOM Ma REY 3.f. 


_ ARGILOFHILUS MARMORATUS ORNATUS Fie's. 12] To 124 


J 
; 5 5 x x 28 5 z& 2 F $ Po 5 ; | 
2 EB o bie ty ae o. 98 : 
ae ET vieheai rT : 
RB | ifr oao DDD — ii ) 
& 3.4 coy } a f 4 
{ Ee ; a 
4 Sng ee eset al a Q } 
A, aa a a Sa) ee 
a 
a | 
3 
q 
i 4 i. 
: 3 °° 
, r S32) ; 
} = 


Fia's. _125 ro 130 


ARMORATUS PAPILLIFER Ftq@’s. 131 & 132 


& 


GILOPHILUS M 


ARGILOPHILUS MARMORATUS ORNATUS 


AR 


MEM. CAL. ACAD. II. PLATE XxXIxX 


: t 
nmdep, | 


blo, 


UTH.BRITTORSREY oF 


ARGILOPHILUS MARMORATUS PAPILLIFER FIGs. 133 70 136. 


OF 


THE 


~— Cauirornia Acanemy or Scrsnens 


L. 


nee 


By GUSTAV EISEN. 


SAN FRANCISCO, CAL. y 


MARCH, 1895. 


| ay ZAMASON 4x 
e \ 
NOV 8- 1956 
® 


Inv. 


PACIFIC COAST OLIGOCH ATA. 
I: 


PHGNICODRILUS TASTE: PONTODRILUS MICH -ELSENI; ECLIPIDRILUS FRIGIDUS. 
BY GUSTAV EISEN. 
PHCENICODRILUS owov. gen. 


Small slender terrrestrial oligochzt closely allied to Ocnerodrilus from which 
genus they differ only in the absence of a prostate. In Ocnerodrilus the organ 
known as a prostate, or by some as atrium, is a prominent and important characteristic 
also shared with such genera as Kerria and Gordiodrilus. These genera which form 
a natural group have been chiefly characterized by one or more prostates, opening 
either with or independently of the spermducts. Our present form resembles 
Oecnerodrilus in all principal characteristics which distinguish that genus from Ker- 
ria and Gordiodrilus, with the exception that it has not even a trace of a prostate. 
There are also some other minor characteristics as will be seen from the description, 
but with the knowledge of only one species it is yet impossible to know if these are 
genus or species characteristics. For the present the following genus diagnosis may 
suffice: 

Small, terrestrial oligochste inhabiting damp soil. 

Clitellum imperfect; comprises the oviduct and the male pore. 

Spermathec with rudimentary diverticula at the inner free end—in ix. The 
spermathecal pore between vill and ix. 

No differentiated penial sete. The common sete sigmoid, 8 in each somite in 
4 couples. 

Nephridia paired, those posterior of the clitellum surrounded by large peri- 
toneal cells. 

Alimentary canal without gizzard and typhlosole, but with one pair of large 
diverticula in ix, connecting with the tubular intestine in the posterior part of the 
somite. Four pair of septal glands in y, vi, vii and viii. 

No subneural vessel. Dorsal and ventral vessel connected by hearts in x and 
xi. Lateral vessel projected anteriorly from each of the diverticula. Blood yellowish- 
red. 

Testes two pair, in x and ix. Large sperm sacs in ix, x, xi, xii. Ovaries in 
xiii. Oviduct in xiv. Two pair of ciliated rosettes in x and xi. Sperm ducts not 
fused, open in the male pore in somite xvii. No prostate. 


64 CALIFORNIA ACADEMY OF SCIENCES. 


Pheenicodrilus taste n. sp. 


The size is that of a very large Oenerodrilus though even fully matured speci- 
mens varied greatly as to length. My largest specimens, which first had been slowly 
killed by dropping solution of corrosive sublimate in the water dish and then extended 
before being hardened with alcohol, reached 2? inches by 1} line in thickness at the 
clitellum. Average-sized specimens were considerably over 2inches long. This refers 
to the mountain specimens collected in the Sierra El Taste; the lowland specimens 
from Pescadero were much smaller not reaching the 2-inch mark. The body is 
slightly tapering towards the tail end. The somites are well set, those of the clitellum 
are hardly distinct. The prostomium is not long, but broad, dovetailing the peristo- 
mium about one-half. From here on the somites gradually, though slightly, increase 
in size until somite x which is a little the largest, xi, xii and xiii are smaller. The 


somites posterior to the clitellum are slightly larger than somite x, except the last few - 


posterior ones. 

Clitellum comprises somites xiv to viii. Vertically it ends at a line drawn 
halfway between sete 2 and 3, slightly receding in somite xiv. Strictly speaking the 
clitellum does not enclose the male pore, as the pore is situated more ventrally than the 
thick clitellar layer, and between that and the pore there is no connecting ridge or 
papilla. 

An accessory copulatory swelling is seen around the outer couple of sete in 
somite xiv (fig. 26, c. c.), the body-wall here being raised like a small mound, with 
the sete slightly outside of its center, from which the cells are arranged as radii in a 
circle. 

The male pore is surrounded only by a very small ridge or papilla, not high 
enough to be seen with a magnifying glass sufficiently strong to reveal the elevated 
papillee of the oviducts. But the whole zone around the male pore is often considerably 
elevated, turned inwards or towards the median line of the body and rounded 
forming a longitudinal groove. 


Exterior pores. The spermathecal pore is situated in the intersegmental groove 
between somites vill and ix, in front of and slightly outside of seta 2, the inner angle 
of the pore being in line with that seta, while the body of the papilla is situated more 
dorsally. The ovipore is situated close by, in front of, but not outside of seta 2. The 
male pore is situated in xvii exactly in a line with sete 1 and 2 according to the lon- 
gitudinal muscular fibres, but as the body-wall is slightly contracted in this somite the 
pore appears as if situated slightly more ventrally; the exact location is, however, the 
place left vacant by the absence of the ventral couple of sete. This couple (sete 1 
and 2) are never, at least not in adult specimens, found developed in this somite, 
though the tips of the young reserve sete are sometimes seen in their sac close to the 
pore. 

Nephropores open in line with seta 2, and are situated in the anterior one- 
third of the space between the sete and the anterior septum. The pores are large, 
round and easily distinguished. 


a 


PACIFIC COAST OLIGOCHETA. 


Body-wall. The various layers offer nothing of great interest. The longitud- 
inal muscular layer is slightly thicker at the dorsal part, and so is the part situated 
between the 3d and 4th sete. The muscles in this layer do not show any feathery 
arrangement around a central fiber, but are irregularly distributed. There are no 
dorsal pores. The body-wall in somites xi and xii is thinner than the anterior somites. 


Arciform muscles. The inner surface of the body-wall in somites xvii and 
xviii, and partially also in xvi, are covered with numerous arciform muscles which 
stretch transversely or diagonally across the body cavity, connecting the region run- 
ning through the outer couple of setee with that of the male pore and the inner set. 
The number of muscular bands in these somites is very great and they vary in length 
and thickness. There is also a slight variation in different specimens, the main fasci- 
cles being, however, always in the main the same. ‘These muscles are best viewed 
when seen from above, the body wall being spread open and the sacculated intestine 
removed. As will be seen from the drawing, which is a careful representation of the 
principal muscular bands, most of the bands begin or are attached tothe body-wall on 
a line running through sete 1 and 2 and ending in a line running through sete 
3 and 4. But there are some which begin more ventrally and end more dorsally 
than either couple of sete. In somite xviii most of the fascicles run in right angle 
to the median line, but the most posterior one as well as one or two more run diag- 
onally backwards. In somite xvii there is one large group of fascicles beginning 
around the male pore and running transversely sideways, while another group of fas- 
cicles run diagonally forward connecting the male region with the anterior part of 
the somite. There are also longer fascicles anterior of the male pore which run much 
further sideways than the outer sete. In somite xvi there are but few fascicles, 
much smaller and shorter. The male pore and spermducts are so entirely covered 
up by these muscular fascicles that they can only be seen with the greatest difficulty 
when viewed on the spread body-wall. In the figure (fig. 7), only the principal mus- 
cles of one side are drawn. ‘The shaded bands which are seen crossing the median 
line are slight ridges in the body-wall, connecting the principal muscle fascicles. The 
objects of these muscles are of course to eleyate and depress the male zone. Similar 
muscles were first described by Benham in Moniligaster, and later by myself in Argi- 
lophilus. They no doubt exist in most species of Oligocheta but are of great interest 
and value as species characteristics. 


The septa begin between somites vy and vi, and gradually increase in size 
toward somite ix. The most anterior septum, however, is not unusually thick. The 
following three septa are very thick, that between viii and ix being the thickest. 


Alimentary canal. 'The buccal cavity is very large and oceupies somites i and 
ii, Pharynx begins in ili or posterior part of ii and extends to the posterior part of 
iii. It is only developed superiorly and merely attached to the cesophagus in the pos- 
terior part of ii. Qsophagus consists of one in the beginning narrow and compara- 
tively even duct, which in iy rises upwards and then becomes considerably sacculated 


66 CALIFORNIA ACADEMY OF SCIENCES. 


in somites y, vi, vii and viii, again to assume a more tubular form in ix. In the pos- 
terior part of this somite it is joined by the two diverticula. The tubular intestine 
proper occupies the twosomites x and xi. In xii begins the sacculated intestine. The 
diverticula of the cesophagus are very long, narrow and slender, more so than in any 
species of Ocnerodrilus which I have seen. These exterior features offer nothing 
peculiar. The lateral blood vessels issue, as usual, from the anterior points of the 
diverticula. 

These diverticula of the cesophagus originate in the posterior part of the 
somite, and not in the anterior part as in Kerria. The structure of these diverticula 
corresponds with that of the same organ in Ocnerodrilus. Only in that genus the 
rule appears to be that the interior of the diverticulum consists of one single chamber 
encroached on by a few parallel ridges. The diverticulum of Phwnicodrilus taste is 
in the central part four to eight chambered, divided off longitudinally, presenting the 
same appearance as an orange when cut through crosswise (figs. 14-16). The number 
of chambers yaries, as in one specimen I found the right hand diverticulum to have 
four chambers, while the left one had five chambers. In sections near to either end 
these chambers fuse further and finally only two and one chambers are left. 

But this fusion is unequal at the two respective ends. In the end nearest the 
junction with the alimentary canal there is only one chamber found, and a little 
further forward there are two chambers, the number increasing until generally eight 
chambers are found in the center. From there on no increase is made, but the cham- 
bers decrease in width, and at the anterior end suddenly fuse into one which is small 
and narrow, not thicker than a blood vessel. The longitudinal blood vesssels are very 
much the same as in Ocnerodrilus, but generally more in number. 

The number of blood vessels in each diverticulum varies with the place where 
located in the diverticulum. The section nearest to the posterior septum of the ali- 
mentary canal shows the two laterals from the dorsal vessel entering the diverticula. 
To begin with, they are seen on the outside of the cellular mass of the diverticulum, 
while in succeeding sections they appear like a few blood vessels scattered on the out- 
side of the epithelial cells. In succeeding sections these vessels are seen to increase 
in number until in the center of the diverticulum they number about one hundred. 

The blood supply for these organs come from branches of the dorsal vessel and 
not from collective vessels from the alimentary canal. 


Salivary glands are as usual found attached to the muscles of the pharynx, but 
they are smaller and less numerous than in Ocnerodrilus, the pharynx being less 
lobed and more compact than in that genus. These glands open through ducts, which 
follow muscular strands into the pharyngeal cavity in a similar way as will be more 
in detail described in Pontodrilus. In fact it is probable that all the suprapharyngeal 
glands in the respective genera of Lumbricids open similarly and without any great 
variation as to detail (fig. 18). The narrow ducts from the gland penetrate the pha- 
ryngeal epithelium and form at its outer edge small ovoid pockets for temporarily 
storing a small amount of the salivary secretion. These ducts end with the pharynx, 


PACIFIC COAST OLIGOCHETA. 67 


the cesophagal epithelium neither being furnished with ducts nor storage pockets (fig. 
18). 

The suprapharyngeal glands are posteriorly connected with the septal glands, 
not only with the nearest pair, but with all the pairs in the respective somites. In 
Ocnerodrilus Beddardi 1 ealled the attention to this fact, but I was not able to point 
out the connection between all the glands, which connection, however, I do not doubt 
really exists in all genera of this family, and probably in most other Oligocheeta. 


Septal glands(fig.2). As regards these glands our present species offers no great 
peculiarities different from species of Ocnerodrilus generally. There are four pair which 
surround the cesophagus in the usual way in somites y, vi, vil and vill. The glands are 
considerably lobed (fig. 1-2), and decrease in size posteriorly. That is, the pair in v 
is by far the largest, the one in vi is smaller and so on, the one in viii being much the 
smallest. This gradual decrease in size posteriorly, though the most common one in 
this class of Oligochxta, does, however, not always exist in all species. I have one 
species yet undescribed from Guatemala in which all the glands are of the same size. 
In our present species the glands are distinctly paired, but they lie so close together that 
that they appear in each somite as one single gland surrounding the intestine. In 
sections the glands are seen to be abundantly supplied with blood sinsues or larger 
vessels. 

These septal glands (fig. 2), connect one and all with the suprapharyngeal 
gland, being, so to say, superposed on several main longitudinal muscular bands con- 
necting the pharyngeal glands with the body-wall in somite ix. Wide and narrow 
duets follow these muscles, causing the secretions of the septal glands not to empty in 
the alimentary canal in the respective somites in the glands, but in the pharyngeal 
cavity as shown in fig. 18. 

Fig. 2 is, as far as outlines are concerned, a correct representation of these 
glands from a section lateral to the cesophagus. Most details, however, are not filled 
out. The glands on the upper side are those above the cesophagus, the lower row 
again those below the cesophagus, both opening on the upper side of the pharynx. 


The sete occur in couples of two, as usual, the distance between sete 1 and 1 
being about the same as the distance between 2 and 2. The shape is the usual sigmoid 
one found in Ocnerodrilus, and the size is rather large. The free points are slightly 
corrugated. The sete occur in all somites after the first, but there are never any devel- 
oped sete where should be the inner couples in xvii. Very small undeveloped tips 
may sometimes be seen close to and lateral to the male pore enclosed in the reserve 
bag, but even they are not always present. 

The blood vessels agree in all respects with those of Ocnerodrilus. There are 
two pair of hearts, one each in x and xi and one pair of connecting vessels in ix. 


Nephridia are found in all the somites except the first few anterior ones. In 
somites ili to v the nephridia are very small and dwarfed, but from there 
on posteriorly they increase in size. Those in front of the clitellum are not fur- 


658 CALIFORNIA ACADEMY OF SCIENCES. 


nished with a glandular covering of peritoneal cells; those in the clitellum show a few 
of those cells, while the nephridia posterior to the clitellum show a highly developed 
envelop of peritoneal cells, similarly as is the case in some species of Ocnerodrilus. 
In our present form the difference between the anterior and _ postclitellar nephridia is 
very marked, the latter ones being prominently visible through the body-wall both in 
alcoholic specimens as well as in alive ones. As regards the form of the nephridia, 
it agrees in general with that of the various species of Ocnerodrilus, some of which I 
have re-examined. The windings of the canals as well as the general arrangement 
of the ducts is much the same in the genera which I haye examined more in detail, 
such as Ocnerodrilus, Kerria, Deltania, Argilophilus, Pontodrilus. Especially is this 
the case with Ocnerodrilus and Kerria, the nephridia of which have been misunder- 
stood, in several particulars, especially so as regards the windings of the canals and 
the presence of blood vessels. 

In Ocnerodrilus as well as in Pheenicodrilus the nephridium consists of two 
distinct parts, A, the folds, with the winding canals, and B, the peritoneal covering, 
with more or less numerous blood vessels. The peritoneal covering again is also divided 
in two parts, one upper almost free, and one lower surrounding or at least adjoining 
the canals, about which more later on. 

In Pheenicodrilus the nephrostome leads to the narrow duct which connects 
with the folds of the main nephridial body. The folds of these canals are placed on 
the outside of or rather on the upper edge of the peritoneal covering. The narrow 
duct when it enters the fold is very narrow, in fact conspicuously so. In the neck of 
the nephridium it coils itself several times around the part of the wide duct that is 
enclosed in the neck. Retaining its narrow size it enters the anterior fold, in which 
it is the most anterior and exterior canal, but nowhere does it appear to ramify as in 
Pontodrilus, Deltania, Argilophilus, ete. It retains its narrow size all through the 
windings (fig. 12), but increases in size in the posterior fold, in which the three canals 
are of equal size. 

In the apex of the spur, which as usual is thicker than the fold, the continua- 
tion of the narrow duct connects with the very wide canal which later on forms the 
bridge. After passing the bridge this wide canal becomes much narrower but still 
continues as thicker than the other canals until it enters the posterior fold. It is also 
less coiled than any of the other canals, in fact it is most conspicuous by being very 
straight—it always occupies the under and inner side of the folds. Even in the 
‘ windings,”’ which, in this species, occupy a very large part of the folds, this canal is 
straight, while the two other canals are coiled and bent. We thus find in this genus all 
the principal parts of the nephridium of the much larger Argilophilus, and it may 
be safely stated in all highly developed Oligochzta the structure of the nephridia 
are in the main the same. As far as Iam aware we may distinguish the following 
divisions of a perfect nephridium: 


PACIFIC COAST OLIGOCHETA. 69 


a. Nephrostomal part. 
Nephrostome, or funnel. 
Neck of nephrostome, consisting of glandular cells. 
Narrow, or nephrostomal duct. 
b. Main glandular part. 
Neck of glandular lobe, connected with the nephrostomal duet. 
Anterior fold. 
The windings, or the coiled part of the two folds. 
Posterior fold, in which the three canals are of the same size. 
Spur, with four canals. 
Bridge, with only one canal. 
c. Efferent part. 
Wide terminal-duct or outlet. The latter is frequently furnished with a 
ceelomic bladder. 
Nephropore. 


When the nephridium is spread out and mounted in glycerine it is seen that 
the canals are of different transparency. Thus the wide canal coming from the bridge, 
together with its continuation in the spur, is much darker than the two .other narrow 
canals, which are conspicuous through their brilliancy. These two white canals meet 
in the very apex of the spur, while the two darker canals meet a little further in. 
The greater obscurity of the wider canal is partly due to ciliation. 

The nephrostome of Phanicodrilus taste is furnished with a large neck, almost 
as large as the rosette, containing several very large cells with conspicuous nuclei. 
The nephrostome is about as Jong as the narrow tube. 

In size the nephrostome ranges with the very largest, it being conspicuous 
and readily dissected. 

The marginal cells vary in number between eight and fourteen. There is a 
larger central nucleus as usual. The glandular neck of the nephrostome contains 
two of these nuclei much larger than any of those found in the rosette, though it also 
contains several smaller ones. When the body of the annelid is laid open and viewed 
from above, the nephrostomes are seen to lie on their side as in fig. 9, the side of 
the thick rim of the rosette being uppermost. In order to see the rosette flattened 
out it is necessary to dissect out the nephrostome, which operation in this species is 
not particularly difficult, as the muscular tissues easily separate. 

The peritoneal covering of the posterior nephridia separates itself in two dis- 
tinct masses, one dorsal and one ventral, the latter being somewhat the smallest. At 
the point where they join the peritoneal covering has narrowed down to « narrow 
band connecting the two main parts. 

The upper peritoneal mass does not surround or contain any of the canals or 
ducts, but appears to be merely an appendage, a gland, as it were, engrafted on the 
lower part. The canals and tubes are entirely confined to this part, as is also the case 
in the nephridia of Ocnerodrilus and Kerria. 


70 CALIFORNIA ACADEMY OF SCIENCES. 


The peritoneal sae consists of very large cells with small, sharply defined and 
very round nuclei. When these cells are empty they are very transparent and 
their walls are very plain. There are a few blood vessels on the peritoneal sac, and 
few of these cover also the folds. 


Nephridia of Kerria (fig. 23). Having recollected Kerria McDonaldi in the 
pond near Miraflores in the Cape Region of Baja California, the only place where it is 
found to date, I have taken the opportunity to re-examine the nephridia in order to 
ascertain their resemblance to those of our present form as well as Oecnerodrilus. As 
a consequence I am able to correct some errors and to add several details. The gen- 
eral structure is the same as in the Oenerodrilus and Pheenicodrilus and the windings 
of the canals the same as in the Argilophilus, ete. The two folds make a large 
rounded loop upwards, and join posteriorly in a very long spur partly free from 
peritoneal covering. The bridge connecting the junction of the spur and the 
posterior fold with the junction of the anterior fold, wide duct and narrow duct, 
is wide and ciliated, in fact it is the widest part of the canal. Posteriorly the 
canal of the bridge projects into the spur forming the widest of the four canals in 
this part. At the apex of the spur two and two of these canals are seen to join as 
usual, forming two loops, one outside of and above the other. The inner and lower 
one of these is the bend of the ciliated canal from the bridge, but it does not ap- 
pear to be ciliated at this point. 

The posterior fold contains as usual three canals, and so does the anterior fold. 
The rounded stretch where the two folds meet is more irregular in outline, and con- 
tains more windings than any other part of the fold, though not as many as in Ponto- 
drilus. The nephrostome is connected with the main body by a slender and nar- 
row tube, the connection being a little in front of the one between the wide duct and 
the main body. This wide duct is almost straight with only a slight curve away from 
the nephridium. It becomes slightly wider towards the nephropore, just before 
reaching which it turns sharply upon itself. 

Another point. of considerable interest is the presence of numerous blood 
capillaries on the nephridia, especially in the peritoneal sac, which they permi- 
ate. The blood has its origin from two vessels, one from the branch from the ven- 
tral main vessel and one from the branch from the dorsal main vessels, the two con- 
necting by capillaries on the nephridial folds. Until now it has been supposed that 
Ocnerodrilus and Kerria did not possess blood vessels on their nephridia, but this is evi- 
dently an error as far at least as some species are concerned. Ordinarily these vessels 
‘are not visible and not distinct from the peritoneal cell-walls but a staining with 
orange G. will bring them out at once. 


The generative organs, with one or two exceptions, resemble those in Oenero- 
drilus in form and general arrangement, and if it were not for the regular absence of 
a prostate our species would be considered as a true Ocnerodrilus. The Spermatheca 
is very large and resembles in general outline that of Ocnerodrilus Beddardi. In 
species of Oenerodrilus the spermatheca always stands up and is pressed close to the 


PACIFIC COAST OLIGOCHETA. Vi 


anterior septum. In Phenicodrilus taste, however, the spermatheca lies always flat 
pressed against the ventral side of the body-wall, and is of sufficiently large size to reach 
as far backwards as to the posterior septum between ix and x, which makes it about 
equal in length to the diverticula of the cesophagus (fig. 1, spth.) The lower part 
of the spermatheca is as usual more muscular than the free end, which in this species 
is more or less, though always shallowly, lobed, showing a large number of incipient 
diverticula irregularly formed and arranged. The spermatozoa are found principally 
in these warty diverticula. 


Testes are very long and situated as usual in x and xi. The posterior one at 
least, and propably both pairs, connect with the sperm-sacs in the same somite. 


Sperm-sacs are arranged as in some species of Ocnerodrilus. They are all 
paired. The anterior pair in ix is attached to the posterior septum of that somite. 
This pair is very much lobed, the lobes being more in number and in shape more 
round than in any species of Ocnerodrilus, the pair resembling two bunches of large 
grapes, completely filling the whole available space in the somite, especially above the 
diverticula and the cesophagus. The sperm-sacs in x and xi are less or hardly lobed, 
connecting with the the testes below, the latter, being long, slender and not branched, 
reach across the somite and joining the sperm-saes in the posterior part near the sep- 
tum. The sperm-sacs in xii are connected directly with those in xi, but otherwise 
attached to the anterior septum of somite xii. This pair of sperm-sacs are lobed but 
not as much so as those in ix. 

There are two pair of ciliated rosettes in x and xi, and two pair of spermducts 
as usual leading from them. The spermducts join, as is generally the case, forming a 
single strand which runs close to, but a trifle more dorsally, than setze 2, until somite 
xvii is reached. In this somite each spermduct enters a small muscular atrium devoid 
of prostate, and entirely confined to the longitudinal layer of the body-wall. As soon 
as entering the body-wall and this muscular chamber the two lumens of the spermduct, 
which until now had been separate, fuse together and enter the muscular chamber as 
one single duct. 

As atrium I consider a small muscular chamber entirely confined to the body- 
wall, in which the spermducts open. This chamber is, however, devoid of any 
glandular cell prolongation such as we are accustomed to find in Ocnerodrilus, and 
ordinarily it ends where the spermducts enter, which is at the upper end of the layer 
of the body-wall. The atrium itself consists of an inner layer of epithelial cells, 
which at the very pore are much larger and furnished with larger nuclei, but which 
gradually decrease in size as they approach the place where the spermducts enter. 
This layer is a direct continuation of the hypodermal layer of the body-wall., An 
outside layer again consists of fine muscular fibres with smaller nuclei directly con- 
tinued from the transverse muscular layer of the body-wall. We see thus that this 
short chamber might in reality be nothing but the remnants of a degenerated atrium, 
or rather remnants of the lower muscular part of a degenerated prostate, which gland- 


ular prolongation has disappeared. That this is the case I judge from the structure 
Memorrs, Vou. II, 4. February, 1895. 


72 CALIFORNIA ACADEMY OF SCIENCES. 


found in one specimen differing from any other which I investigated. Out of six 

specimens which I sectioned off five agreed in all particulars as regards the absence 

of a prostrate, as generally understood in Oenerodrilus, neither did seven specimens 

which I dissected show any trace of such a prostate. One specimen which I sec- 

tioned, however, showed an abnormal prolongation of the muscular chamber, in every 

particular resembling the lower muscular part or the atrium proper of the prostate as 

characteristic of Ocnerodrilus. It ascended inwardly in the cavity of the body as 

high as to sete 8 and 4, ended here blindly without any differential glandular part 

or prostate proper, as is always found in Oenerodrilus. Such “returns” to original 

characters and ancestors must, of course, be expected, and are the more interesting” 
when encountered. We might on the other hand consider Phwnicodrilus taste as- 
standing on a lower grade than Oenerodrilus, the prostate not having yet appeared. 

But against such a view speaks the fact that her organs are as highly developed as 

in Oenerodrilus, which would hardly be expected of a lower form, in which we would 

naturally look for a lower degree of development in several organs at the same time. 

A degeneration of a certain organ, however, such as the prostate, would not necessitate 

a similar degeneration of several other organs at the same time. 

The absence of a prostate in Pheenicodrilus is of considerable interest, and I 
think it clearly demonstrates that the absence or presence of this organ cannot be laid 
at the foundation of families. Such absence of the prostate in an Ocnerodrilide is not 
unexpected. A perusal of the various species of Ocnerodrilus shows us how. these 
species may be arranged in a series according to the size of the prostate, the list being 
headed by Ocnerodrilus occidentalis, with a very extended prostate, while at the other 
extremity we find Ocnerodrilus Hendriei with the most diminutive prostate, not extend 
ing outside of the somite. There is thus only a step further to Phcenicodrilus where 
this organ is entirely absent. That this fact will have some influence upon our views 
of the classification of Oligocheta is evident, and several genera or even families 
which hitherto have been considered far apart solely on account of the presence or 
absence of a glandular prostate, must now be brought closer together. 


Ovary and oviduct occupy the same somites as in Oecnerodrilus and offer no 
characteristics of interest. 

Halitat. 1 found this species in the Sierra el Taste, in the Cape region of Baja 
California, some 40 or 50 miles north of Cabo San Lucas, at an altitude of 4,000 feet. 
Later on I found specimens in great number in a garden at Pescadero on the Pacific 
coast, on the western slope of the same sierra, but at an altitude of only a few feet above 
the ocean. The water used for irrigation was taken from a creek coming from the 
sierra. 

The species lives in damp soil and occur in great numbers, not mixed with 
any other form as far as my experience goes. The distribution of the species is most 
interesting as on the eastern side of the Sierra in the valley of San José, Inever found 
any other Oenerodrilid than Ocnerodrilus Beddardi. The question if the Sierra really 
absolutely divides the habitats of the two only forms of this family found in the Cape 
Region, further explorations are necessary to decide. 


PACIFIC COAST OLIGOCH®TA. 


~J 


oo 


PONTODRILUS Grube. 


The genus Pontodrilus, together with Photodrilus, is distinguished from all re- 
lated genera by the absence of nephridia in the twelve anterior somites. In the 
majority of species the nephridia commence in somite xiii, but in two species they 
commence respectively in xivand xy. A common character to all the species appears 
to be the very great thickening of the septa between somite v and xiii. 

I have so far on the Pacific Coast only found one species of the genus Ponto- 
drilus. Examinations were made from alcoholic specimens, I having no opportunity 
to examine them when I found them in their native habitat—Mexico. 


Pontodrilus Michaelseni n. sp. 


This species differs from all other species, except P. Maronis,* which have been 
referred to this genus in possessing a glandular crop occupying somites xiv, xv and 
xvi, as well as in other minor details. The habitat of the species is the very narrow 
moist line between high tide and dry soil on the shores of the Gulf of California 
around Guaymas, Mexico. The soil in which it occurs is very sandy and thoroughly 
soaked or moistened with the strongly saline water of the gulf. It occurred here in 
large numbers, but unfortunately at my visit most of the specimens were immature, 
only two possessing clitellum in the end of November, 1895. I dedicate this species 
to Dr. W. Michaelsen of Hamburg, whose labors in our common field are among the 
the most thorough and best. 


EXTERIOR CHARACTERS. 


The body of this species reaches a length of 3} inches with a width at the 
clitellum of less than $ of an inch, but the majority of specimens are somewhat 
smaller. The above measurement refers to specimens slowly killed and then hard- 
ened in alcohol. The body is tapering towards the tail end, the latter however being 
slightly swollen (fig. 24). 

The prostomium encroaches on somite i, dividing it about one-half. Somite i 
is slightly larger than any of the following somites (fig. 25). 

The clitellum commences in xiii and in full-grown specimens includes part of 
xix. It is incomplete in a peculiar manner. In xiii to xvii inclusive it is only 
developed on the dorsal side of the body. But in xviii and in part of xix the clitellum 
is only developed on the ventral side of the body, though this fact cannot be ascer- 
tained from exterior inspection. Viewed from the underside the clitellum appears to 
be on a line drawn through seta one. Between xvii and xviii the clitellar swelling 
recedes slightly, again to widen out in xviii, and here joining to a pair of ventral 
cushions, between which and the clitellum proper are situated the male pores. 

Another pair of swellings are noticed around the spermathecal pores, covering 
on either side parts of somites vi, vii and viii (fig. 26). 


“TI received M. Perrier’s memoir on Pontodrilus only after this paper was partly in print. 


74 CALIFORNIA ACADEMY OF SCIENCES. 


External pores. There are no dorsal pores. The spermathecal pores are 
found between somites vii/vili and viii/ix in front of seta 2, each one situated ona 
slightly elevated cushion. The ovipore in xiy in front of seta 1. The male pores are 
in xvili in front of and in line with seta 2 (fig. 2646). The nephropores are in front 
of seta 2. 


Sete. The sete commence in somite 11, eight in each segment and in couples. 
The distance between 3 and 4 is only slightly larger than that between 1 and 2. The 
distance between 1 and 1 is nearly twice as large as that between 1 and 2, and the 
distance between 2 and 3is a little smaller than that between 1 and 1 (fig. 27). 


Color of body pale flesh, rather transparent and marbled very much like 
Deltania. Clitellum yellowish. 


INTERNAL ANATOMY. 


Body-wall. The body-wall appears to me to be of unusual thinness, throughout 
the length of the body. The dorsal side is slightly thinner than the ventral side, at 
least anterior to the clitellum (fig. 29). Dorsally the longitudinal muscular layer is 
of about the same thickness as the transverse layer while on the ventral side the 
longitudinal muscular layer is about twice as thick as the tranyerse muscular layer. 
This refers to the anterior somites. To this there is however an exception in somites 
vill and ix where on the ventral side in the vicinity of the spermathecal pores the 
transverse layer is thicker than the longitudinal layer. The transverse layer tapers 
down towards the spermathecal pores, but this thickening is found only in the imme- 
diate vicinity of the spermatheca. In the clitellar somites the relative development of 
the muscular layer is very different. Here the inner or longitudinal muscular layer 
is enormously thickened laterally in somites xvil and xviii or in the vicinity of the 
male pores (figs. 37, 38, 40, 41), while in the anterior part of the clitellum the longi- 
tudinal layer is only thickened ventrally, between the inner couples of sete it here 
being at least twice as wide as it is dorsally (fig. 59). 


Clitellum offers many points of interest. It has already been stated that this 
organ is incomplete, that is, not simultaneously developed on the dorsal and ventral 
sides. A section through an immature specimen shows (fig. 38) that the clitellar 
glandular layer is developed only between the seta, that is, from seta 4 ventrally to 
seta 4, while dorsally there is no trace of such cells. As regards the nature of these 
cells it is to be remarked that they are unusually small or rather thin compared to the 
larger and thicker cells of the dorsal part of the clitellum in the anterior somites of 
that organ. These latter cells offer nothing in particular of interest, resembling those 
of other genera of the family as far as I can make out. Unfortunately most of the 
specimens were immature and only two possessed clitellum, but these two had unavoid- 
ably not been treated, and had contracted to such very great extent that the finer 
structure of the clitellum had been hopelessly lost. From cross and longitudinal sec- 
tions made it was, however, evident that the clitellar glandular cells, which constitute the 
clitellum, do not extend ventrally further than to sete 1, thus leaving the ventral space 


=—I 


iy | 


PACIFIC COAST OLIGOCHMTA. 


between sete 1 and 1 with the usual layers less the glandular cells (fig. 41, 6. c/.) 
marking the points where the clitellum ceases. We have thus before us a species 
with a dorsal and lateral clitellum in some somites, and with a ventral and lateral in 
one somite. 

In all of the clitellar somites the inner or longitudinal muscular layer possesses 
an enormous development laterally, especially so in the region of the prostate, where 
it reaches a width four times that at the dorsal and ventral part. The reason of this 
enormous thickening of this layer is readily understood when we view one of the 
sections of this region. The part of the body-wall, on which rests the prostate, is 
attached by numerous arciform muscular bundles, arranged in a fan-shaped manner, 
to the dorsal and lateral sides of the body. These muscles are more numerous and 
stronger than any I have seen described in other species, making it possible for this 
region of the body to contract in a most characteristic manner. 

When the body is opened and flattened out these muscles are seen to spread out 
from the male pore laterally, connecting the center of somite xviii with the two 
nearest somites xvii and xix as I have endeavored to show in fig. 42. As may be 
judged from the figure the muscles are arranged in regular fascicles of which we may 
count twenty-two to twenty-four as being more prominent and overlapping the other— 
a smaller number of fascicles situated below. The wider periphery of the attachment 
is situated on the body-wall above the prostate, but the lower and narrower part of 
the attachment of the fan is situated on a peculiar swelling which I have designated 
as the copulatory cushion (fig. 42, ¢. ¢.; fig. 41, ¢. ¢.; fig. 38, ¢. ¢.; figs. 44, 45), 
apparently a thickening of the longitudinal muscular layer, transversed by innumer- 
able other muscles in every direction, all connected by what looks like connective 
tissue. The size of this copulatory cushion is very great; not only do these swellings 
project considerably outward but inwardly they encroach to a great extent on the 
internal cavity, especially so in mature specimens. This peculiar organ is entirely 
muscular, there being no sign of any outwardly opening papille. The prostate pore 
is situated between this cushion and the projection caused by the increased thickness 
of the longitudinal layer and the contraction of the fan-shaped muscles. In fig. 43 
the section, a longitudinal one, is somewhat oblique, having followed the large muscular 
strands which surround the outlet duct of the prostate as well as the muscles of the 
copulatory cushion. Fig. 44 represents a more vertical, longitudinal section, more 
interior to the large fan-muscles, which do not show in the section except their inner 
attachments (a. m.) The large arciform strands vary considerably in size, those 
nearest the body-wall being the smallest, those nearest the intestine the largest. 
When a cross-section through somite xviii is viewed it will be seen that the smaller 
strands are filled between with connective tissue (fig. 41, c. ¢.), while a concentric 
transverse muscular strand is crossing them near the inner angle of the prostate pore. 
Other muscles connect the glandular prostate with the body-wall (figs. 37, 40 m and 
42 sp.m.) The large strand sp. m in fig. 42, connects the bend of the prostate, where 
enters the sperm duct, with the ventral part of the body-wall under the ventral nerve- 
cord, 


76 CALIFORNIA ACADEMY OF SCIENCES. 


Septa. The first very pronounced septum is found between somites iv and vy. 
Beddard has already remarked that the septa in Pontodrilus hesperidum are unusu- 
ally thickened, some of them being thicker even than the body-wall at the point 
where they are attached to it. The septa in our present form are almost gradually 
increasing in ‘thickness towards the one between xi and xii, which septum is the 
thickest, being almost as thick as the ventral body-wall. The septum dividing xii 
and xiii, while thickened, is much thinner, and those bounding the somites down to 
xix are almost normal in thickness, that is, equal those posterior to the clitellum. As 
a rule the attachments of the septa correspond with the intersegmental furrows. The 
septa bounding vii/viii and viii/ix are, however, exceptions as far as the place of their 
ventral attachment is concerned. These two septa are here affixed to the body-wall 
half-way between the sete of the anterior somites, respectively viiand viii. This makes 
it appear as if the spermathec:e opened in the centre of the somite, when in reality 
they open as usual in the intersegmental groove. 


Alimentary canal. The alimentary canal takes the shape of a long, narrow 
duct, singularly straight and without any prominent characteristics until it reaches 
somite xiv, in which somite commences a kind of gizzard of peculiar construction 
(fig. 29, giz.) Though this organ resembles a gizzard in outward form it is in reality 
no gizzard at all, but rather a glandular modification of the alimentary wall. With 
a gizzard we must of course mean an enlargement of the alimentary canal in which 
the muscular part has reached an enormous development in order to grind the food 
properly. In the organ referred to in our present species the muscular layers are on 
the contrary not increased in size, the thickening of the wall being caused exclusively 
by a new layer composed of glandular cells, which has been interposed between the 
transverse muscles and the inner epithelium, thus forming a glandular crop between 
the cesophagus and the tubular intestine. This organ occupies three somites, xiv, xv, 
xvi, or very much the same place as is so frequently the location for gizzard in other 
Oligochete. If we view a longitudinal section of the body through this crop (fig. 47) 
we find it to be more or less tapering towards either end. The large longitudinal 
blood vessel lies almost immediately on the top of the outer or chloragogie cells, in 
places penetrating them with connecting vessels which supply the underlining sinus 
with blood (fig. 47, d. v. ¢.). 

The chloragogie layer of cells vary considerably in size. Sometimes there is 
more than one row of cells, one projecting above the other. The nuclei are oyal and 
situated at the place where the cells become narrower. The longitudinal muscular 
layer is narrow, about two strands thick, immediately superposed the transverse layer, 
which consists only of one single thickness of strands (fig. 48 to 54 ¢. m.) The case gen- 
erally observed in gizzards is that this layer is composed of a great number of strands 
more or less regularly arranged around a central plate. Below this transverse layer, 
commences the very thick layer of glandular cells, about 12 cells wide in centre. 
In the upper part of this layer are seen numerous blood lacunes, which in 
places join the muscular layers (fig. 49 to 53 02), at other times are more or less 


PACIFIC COAST OLIGOCHETA. bre 


perfectly enclosed in the glandular layer, forming generally a row of vascular 
lacunes near its outer margins (fig. 48 (/. s.) apparently without touching the mus- 
cular layer, or doing this only at certain places. Figs. 49 to 53 represent cross-section 
from different parts of the crop, from the posterior part, near the boundary of somite 
xvii (fig. 49), to the anterior part of xiv (fig. 52), illustrating the variations in thick- 
ness of the glandular layer. In the posterior part this layer is very thin (fig. 49 g/.) 
consisting only of a few cells, from one to three cells wide. In figs. 50 to 52 the 
glandular layer is seen to have increased in thickness and so has the inner epi- 
thelial cells. Fig. 55 is a portion of the glandular layer, in which the inner lumen 
is more plainly represented. The outer part of this glandular layer is divided up in 
lobes by the numerous blood sinuses, and in each such lobe there is a wider or nar- 
rower, generally, branched lumen, which, however, I have not been able to follow 
down to the epithelial cells. 
The sacculated intestine commences at the posterior end of the crop, and 
offers nothing of great interest or characteristic. 
There is no thyphlosole, but the intestine is otherwise very rich in blood 
lacunes. 


Pharyngeal or Salivary Gland. Pharynx which occupies somites ii and iii is 
only developed dorsally. It is superposed by a large mass of glands and muscles, as is 
usual in a large number of Oligochetee. In outline this glandular mass is remarkably 
even, especially so at its posterior end. In a longitudinal section we see customarily 
three lobes (fig. 29), supported by long strands of muscles, running back to the pos- 
terior boundaries of somites vi, vii and viii. 

On the ventral side there are seen three of those muscular strands, similarly 
running back to vi, vil and viii, indicating that there is a row of similar strands corres- 
ponding with somites ii, ii and iv. In somite vy there is a pair of smaller glands of 
similar nature attached to muscles which connect with the larger strands of the main 
gland (fig. 29, s.s. g/.) 

A cross-section of this glandular mass (fig. 80), shows us that the glands are situated 
principally on the periphery, supported by muscles (7s.), while the inner and posterior 
part is principally taken up by strands and ducts. A division of the mass in three more or 
less distinct layers is discernible, probably corresponding to somites li, iiiand iv. These 
glands communicate directly by means of ducts with the epithelium of the pharynx. 
In fig. 29 these ducts are roughly represented as dark violet. In the cross-section 
(fig. 30), the darkest blotches are intended to represent the ducts, while the lighter 
colored violet ones are the glands. In fig. 31 a lobe of the longitudinal section is seen 
in a larger magnification, and in fig. 52 a smaller lobe, yet more highly magnified. 
The glandular cells are rather of varying size, and arranged around the margin of 
glandular sack, leaving the inner space open. The cell cytoplasm is massed in places, 
leaving in other places larger or smaller, generally roundish, vacuoles (fig. 35, va.) 

In figs. 83 and 34 I have endeayored to show the relative arrangement of the 
muscular strands, the glandular ducts and the glandular cells. As will be seen these 


78 CALIFORNIA ACADEMY OF SCIENCES. 


ducts are in places entirely closed in by muscles, while the glands themselves are 
only supported by them. 

The ducts lead directly to the pharyngeal epithelium; arrived here they branch 
out sending numerous discharge-tubes between the epithelial cells (fig. 36 g/. dt.), dis- 
charging the salivary mucous in the pharyngeal cavity. These ductules are fre- 
quently, though not generally, branched while in the epithelial layer. Each ductule 
is furnished at the distal end with a small storage chamber (36a) of oblong form and 
considerably smaller than the nucleus of the epithelial cells. 


Septal glands. As hasalready been stated there are five pair of very small glands, 
which are pricipally attached to the connecting vessels in somites v to ix, and sit- 
uated on the ventral side of the cesophagus. These glands do not hang on closely to 
the septum, but are apparently suspended from it only by a few tiny mesenteric tis- 
sues and by a muscle or two. In longitudinal section they appear as suspend entirely 
between the two septa (fig. 29), while in cross-section they are seen to be affixed to 
the connecting vessels and from them project laterally, the point of affixion being 
close to the ventral vessels (fig. 70 g/.) The general outline of these glands resemble 
the so-called liver cells attached to the connecting and other vessels in some Lumbri- 
cides, but the structure is similar to the salivary glands. The gland in v resembles 
exactly the structure in the salivary glands which open in the pharynx, it being 
transversed by blood capillaries, infested with the same parasites, supported by mus- 
cles, and finally is only sparcely surrounded by floating, globular, ecelomie cells. The 
other gland in vi to ix are all surrounded by a thick coating of these floating coelomic 
cells. These glands stain in the same way as the salivary glands, their secretions being 
stained deep violet with hematoxylon-orange, while the ccelomic cells stain pale yellow. 
A fine and very thin duct runs backwards and upwards from the far upper end of 
each gland towards the alimentary canal to its junction with the septum, but I 
have some doubt about it emptying into the intestine, and it is much more 
probable that in Pontodrilus, as well as in Pheenicodrilus and Oecnerodrilus, these 
septal glands empty into the pharynx. None of my sections, however, show 
this to be the case. Certain it is that in Pontodrilus the various septal glands 
are not as closely connected with the salivary glands as in the just mentioned 
genera, in which the respective glands are actually not only suspended from the same 
longitudinal muscular band, but along and resting on the latter run also the collective 
ducts of the glands. Among the salivary and septal glands are seen numerous ir- 
regular, generally oval or oblong bodies full of nuclei. These are the terminal pock- 
ets of the capilaries, generally termed blood glands. 


Blood glands (fig. 754, t.). Ed. Perrier was the first to describe blood glands 
in Pontodrilus, but he found them exclusively in the blood vessels or at the end of 
the capilaries investing the nephridia. In our present species, P. Michaelseni, I have 
found these glands only in the capillaries of the salivary and septal glands. They 
here occur in very large numbers, especially in the former, being massed at 
or near the posterior edge of the gland in varying numbers. Some specimens con- 


PACIFIC COAST OLIGOCHETA., 79 


tain comparatively few blood glands, others three or four times as many. They 
are of all sizes and shapes, as Perrier has shown. Some contain only one single 
nucleus, then frequently surrounded by a blood clot; others again contain a very great 
number of nuclei, which are then situated in a sac-like pocket at the end of the blood 
capillary. In some of the larger blood vessels in the salivary gland the blood gland 
takes the form of a ‘“hertzkérper.”’ The smaller ones situated on the capillaries 
may be named terminal blood glands, while those situated inside the larger vessels 
may be designated interior blood glands. ‘The structure of the two are at least in this 
species very similar. 

In fig. 75, a. to s., [ have endeavored to illustrate the structure of these blood 
glands from sections. In g.a large blood vessel with a blood clot, at the base of which 
is an inner blood gland. On one side of the blood vessel is a part of a salivary gland 
with brown secretions. In a. a small terminal blood gland is shown, and in 4. and e. 
some of a greater development. The nuclei are not always surrounded by a distinct 
cell membrane, in fact in almost every gland are found some nuclei with distinct cell 
membranes while others lie loose in the granular serum. The exterior line in all 
the figures represents the wall of the blood vessel, and the difference between the 
terminal and inner blood glands consist in reality only in the absence or presence of 
blood surrounding the glands. As far as the granular protoplasm concerns it is always 
differentiated in two parts. The one at the distal extremity is more evenly dif- 
fused and finer grained than the one next the capillary, which again is coarser, streaky 
and which, besides, stains differently or at least more intensely than the other. Many 
of the glands contain larger or smaller bodies (p., ¢. and 0.) equally of round form 
and lighter in color than the cytoplasm, but sometimes they are very opaque, stain- 
ing deeply as at r., the two classes probably being of entirely different character. 
The former resembles a pale nucleus, while the latter opaque bodies appear only to be 
secreted matter. The paler ones may possibly be parasitic protozoa. 

The blood glands described by Claparede, Lankester and others in Lumbricus, 
ete., are probably of a similar construction, and judging from the figure given by 
Michaelsen of the “ hertzkérper” in Enchytreeus, we may conclude that it, too, is 
identical with the blood gland in Pontodrilus. 


Spermathece (figs. 30, 55, 56, 57, 58). There are two pair of spermathece 
found in somites viii and ix, the exterior pores being as usual in the intersegmental 
grooves between vii/vili and viii/ix in line with setee 2. Each spermatheca possesses 
a tubular diverticulum, the junction of the two being in the body-wall. The position 
of the diverticulum is always ventral to the spermathece proper. This is cylindrical, 
quite narrow, with a larger globular chamber at the free inner end, in which the wall 
is much thinner than in the cylindrical part. At the junction with the body-wall 
is a much larger swelling, the lower and more strongly muscular part of the main 
cylinder being greatly widened, presenting a muscular cushion partly projecting above 
the body-wall, partly again being immerged in it. The spermatozoa are principally 
massed in the inner globular chamber, though they are seen also in the diverticulum. 


80 CALIFORNIA ACADEMY OF SCIENCES. 


The main spermatheca is strongly muscular, especially at its lower end where 
the muscular layer is much thicker than the inner epithelial cells. These latter are 
large with oval nuclei. Outside of these cells and between them and the muscular 
layer are seen a row of interstitial nuclei of much smaller size than the nuclei of the 
epithelial cells (fig. 56 intes. n.) The structure of the diverticulum is somewhat 
different. Interiorly we find large epithelial cells with large round nuclei. Outside 
of them is a single row of interstitial nuclei. Surrounding them we find a circular 
muscular layer varying in thickness (fig. 57, ¢. m.) and with few, small nuclei. 
Exterior to this layer are seen numerous blood vessels, and outside of them a two or 
three cells thick layer of glandular cells (g/. c.) 

When the body of the worm is viewed from the interior, with the alimentary 
canal removed, the spermatheca as well as their diverticula are seen to extend back- 
ward, parallel with the ventral ganglion. The diverticulum is always much shorter 
and narrower than the spermatheca proper. 


Testes and Ovaries. The former organs consist of two pair of minute narrow- 
lobed bodies (fig. 29, t. 59, 60), the lobes being all in one plane, parallel to the body- 
wall. One pair are in x and one in xi as usual. 

The ovaries consist of one pair of flat bodies with wavy margin and wide and 
shallow lobes, distributed in both a horizontal and vertical plane (figs. 61 and 62). 
As usual the ovaries are in xiii. 


The oviduct is placed as usual with the funnel in xiliand the pore in xiv. The 
funnel part is very thick, fig. 63 drawn from a longitudinal section. 


Oihiated rosettes, Spermducts and Prostates. There are as usual two pair of 
ciliated rosettes in x and xi opposite the testes. The funnels are very thick and not 
much crimpled or hardly crimpled with one flare on either side. The epithelial nuclei 
are quite long, and their cells are superposed a thick layer of very distinct blood 
vessels (fig. 65, cr. 66 and 67). The spermducts unite to a single duct which passes 
immediately outside of the second setee. The duct is unusually narrow, the narrowest 
I have seen in any species of this size. The spermduct enters the glandular part of 
the prostate just above the intersegmental groove separating xvili/xvil (fig. 42, spd.; 
68, spd.) 

The prostate is tubular, very large, bent upon itself once. It starts from the 
male pore, which is situated in the center of xviii, forwards, running parallel with 
ventral nerve cord. “When it reaches xvii it turns backward, its apex being in the 
center of xviii. The prostate consists as usual of two distinct parts, connected in the 
center of xvii. The advancing part is strongly muscular, the returning part again is 
glandular. The prostate is cylindrical, the two halves being almost equal in thickness. 
The part which penetrates the muscular body-wall is several times thinner than the 
other part. The muscular part consists of two layers, the inner one consisting of a 
row of epithelial cells with oblong nuclei. The outer layer, which is very thick, con- 
sists entirely of circular muscles with a few small nuclei. 


PACIFIC COAST OLIGOCHETA. 81 


The glandular part of the prostate which commences at the anterior bend of 
the organ consists of two or possibly of three layers. The inner lining consists of only 
one layer of epithelial cells with ovoid nuclei. These long cells appear to be sur- 
rounded by a narrow zone of fibrous or perhaps muscular tissue with few nuclei. 
But by far the greatest part of the prostate consists of fibrous tissue with numerous 
small roundish nuclei, with here and there a cell being visible, and with a few cells 
of a glandular appearance (fig. 68), especially toward the circumference. In cross- 
section the anterior part of this prostate is triangular in outline, while the muscular 
part is always circular, and in size always thicker. The very narrow part of the 
prostate which penetrates the body-wall is strongly muscular of the same general 
structure as the free muscular part. The general structure of the prostate appears 
similar to the one of Pontodrilus hesperidum as described by Beddard. He has 
pointed out the absence of a regular layer of glandular cells in the prostate of that 
species, and it is possible that this construction of this organ which thus ap- 
proaches that of Ocnerodrilid, is not a species but a generic character, if it does not 
prove to be of even greater value. 


Vascular system. There are a dorsal and ventral vessel, but no subneural, nor 
any subintestinal vessel, and no thyphlosole. The two main vessels are connected 
with hearts in x, xi, xii, xiii, the most posterior one of which is found in somite xiii, 
immediately in front of the sacculated intestine. This heart is the largest, the others 
gradually decrease in size forward. The posterior part of these vessels are entirely 
free of brown cells. The ventral vessel is forked in somite ix in two parallel 
lateral vessels, there being no central vessel left. These two branches are always of 
unequal size, both being situated immediately under the cesophagus (fig. 64 v. v.) In 
the somites anterior to x, these branches of the ventral are connected by laterals with 
the dorsal vessel. In one specimen the ventral fork commenced in xii (fig. 64). 

Between the dorsal vessel and the ventral forks there are connecting vessels, 
one pair in each of the somites y, vi, vii, viliandix. To the ventral parts of these 
connecting vessels are attached oblong glands, which again are surrounded by a coat- 
ing of globular brown cells. These glands do not extend clear to the dorsal vessel, 
but end laterally before reaching it. The nature of the glandular cells appear the 
same as those of the pharynx, staining in exactly the same way. ‘The cells of these 
septal glands are more numerous in the anterior somites, gradually diminishing 
posteriorly, while the opposite is the case with the free round cells which are more 
numerous in the posterior glands (figs. 29 g/. and 71 g/.) . 


Nephridia (figs. 71 and 72). These organs commence in somite xiii, or in the 
same somite as the ovaries. The first two anterior nephridia are furnished with a 
smaller covering of peritoneal cells, but already in xvi do the nephridia attain their 
full size, as in all posterior somites. 

The nephridia are built upon the same general principles as those of Argilo- 
philus, Deltania, Ocnerodrilus and Pheenicodrilus, as well as of Lumbricus as shown 
by Benham. We find here the corresponding duets, canals, lobes, ete., and a gen- 


82 CALIFORNIA ACADEMY OF SCIENCKES. 


eral description of them will be superfluous. The spur is directed backwards and the 
two folds are directed upwards in about right angle to the spur and narrow duct as 
well. The spur and the folds rest on the large lobes of peritoneal cells, one of which 
is posterior and one anterior. The posterior one surrounds the spur, upon the ante- 
rior one, which is the smallest, rests the two folds. The narrow duct is not unusually 
narrow, while the wide duct or outlet duct is very narrow, not any wider than the 
narrow duct. The neck of the anterior fold is, where it connects with the nar- 
row duct, very wide, enlarged, irregular and sigmoid, gradually increasing in size to 
the anterior fold. Where the two folds join, the fold is always very coiled. The tube 
forming the bridge is not any wider than the clear canals, but it is less clear or trans- 
parent, just asin Lumbricus. The canal leading from this bridge into the anterior 
fold, is straighter, darker, and slightly wider than the two bright tubes which are 
much coiled and situated more anteriorly and superiorly to the straighter canal. This 
coiling ceases as soon as the big bend and windings are passed and the posterior fold 
is reached. 

The nephrostome is large. The marginal cells in the rosette are only slightly 
decreasing in size toward the extremities or centripetal marginals. There is a large 
centripetal protuberance surrounding the inner opening of the duct, as in Lumbricus. 
as described by Benham, but the centrifugal cells are less regular and more scattered, 
The centrifugal cells are never hidden by the centripetals as in Lumbricus, and the 
whole centripetal protuberance is most prominent seen in whatever direction. The 
outlet duct enters the fold much closer to the narrow or nephrostomal duct than is 
usual in Oligocheeta, in fact it connects with the free neck of the anterior fold, close 
behind the septum. 

The relationship of Pontodrilus Michaelseni to the other species of the genus 
is not as clear as we might wish. Beddard’s description and notes in his paper, 
“V. Some new or little known Oligocheta,” are the only comparative remarks yet 
made on the few worms which are grouped under this genus, an arrangement which 
must be considered as entirely preliminary. The only very characteristic features 
which connect the six species of the genus is the commencement of the nephridia 
posterior to somite xii, and the opening of the spermduct into the prostate, absence 
of typhlosole, grape-like sperm-sacs, and no penial sete. None of these species 
have been sufficiently described, an unavoidable fault attendant all species im- 
mersed in alcohol without previous careful preparation and evacuation. In the 
following table I have endeavored to compile the characters of the various species as 
far as I can make out fromthe descriptions, no specimens for comparison being in my 
possession. I include here, as suggested by Beddard, the genus Photodrilus Giard. 
I have had no access to Grube’s description of P. Jittoralis, and haye therefore 
excluded it from this table. 


PACIFIC COAST OLIGOCHEHTA. 


83 


SPECIES OF PONTODRILUS AND PHOTODRILUS. 


P. Michaelseni| P. arene P. hesperidum' P. insularis P. Marionis | P. phosphoreus 
n. sp. Fr. Miller & Bedd. Rosa. Perrier. Dugés. 
ose Michaelsen. 
Nephridia commence 
it SOMES. 3.5.00 xiii. xiii. xiii ? xiii. XV; xiv. 
Nephropores in line 
with seta. .......... 2. ? ? 4. 2. 2 
| 
Gizzam de tachetie seats = Rudimentary. | Absent. Absent. Rudimentary. Rudimentary?) Absent. 
esophageal swellings..| None. None. | None. None. None. Four in x-xiil. 
Glandular crop....... Large in xiv, | None. None. None. Present. | None. 
vt : 
XV, Xvi. } | 
| 
‘ 2 
Septal glands......... In v to ix. ? None. | None, Present. | In y-ix. 
L g 
Penial ste oo. nce ete Absent. Absent ? | Absent. Absent. Absent. Tn xvili and xii 
Common sete. ........ Plain, regular.. Ornamented, | Plain, regular. Plain,irregular, Plain, regular. Plain, regular. 
regular. | behind. | 
Spermatheca.........+ | Two pair in| Two pair in|; Two pair in| Two pair in| Two pair in | One pair in ix; 
| vill and ix;| viii and ix;, viii and ix;| viii and ix; viii and ix; one diverticle. 
| one diverti-| one diverti- one diverti- no diverti- | one diverti- | 
cle. cle, | cle. cle, | cle. | 
| | 
Sperm-8cs........+-5 Two pair,| Two pair, | Twopair, rac-| Two pair, | Two pairin xi Two pair in xi 
| grape-like, grape-like,| emose,in xi| grape-like, |) and xii. and xii. 
in xiand xii.} in xiand xii. and xii. | in xiand xii. | | 
Clitoris a clncace:- = dsc | xili—xvili, xili-xvii, hee vg xiii to xvii xili-xvii ? 
: : | | 
| incomplete. incomplete. | | | complete (?). 
Prostomium . .....0.- Encroaches on| Encroaches } | ? | Does not en- | Encroaches on Does not en- 
somite 1. on somite 1. | croach on somite 1. croach on so- 
: ; 
| somite 1. | | mite 1. 
| | 
Ventral papille....... In xviiand xix) | One central, | 
| xix-xx, and one) 
| | ditto, xx-xi. 
ELD Satis cahetctetcl 32 stele Guaymas, Desterro Jamaica. Arn Island. | Marseilles. France. 
é z = | 
Mexico. Brasil.| 


In the above table I have principally followed Beddard, but I cannot agree 
with his opinion as regards the identity of Grube’s P. /ittoralis, with that of P. Marionis 
of Perrier. As Perrier remarks, the ventral median papills described by him in P. 
Marionis are not found in P. littoralis, in which species these papillae were paired. 
Until Grube’s species has been found and redescribed we must therefore accept it as 
a separate one. It is of course not by any means impossible that one or more species 
of Pontodrilus are to be found in the same locality. 

In reference to the gizzard I have given it as rudimentary in two species. This 
refers only to an anterior gizzard close behind the pharynx, in which the muscular 
layer of the cesophagus is simply thickened as shown in fig. 46 Aands. Beddard says 
that P. littoralis Grube has also a rudimentary gizzard, but I think that he judges by 
M. Perrier’s description of what I call a glandular crop in the somites posterior to the 
testes. It may not be impossible that the muscular thickening of the csophagus 
really exists in all the species more or less prominently, as it is easily overlooked. 


84 CALIFORNIA ACADEMY OF SCIENCES. 


That the glandular crop really exists in P. Murionis is evident from Perrier’s drawing 
(fig. 22, pl. xvi), though I cannot accept the epithelial nature of the cells. Perrier 
says nothing of the place from which this drawing is taken, bunt I suspect that it was 
from a section of the intestine between the tubular and sacculated parts, similar as in 
P. Michaelsent. 

LITERATURE. 


Bepparp, Fr. E. Abstract on Some Investigations into the Structure of the Oligochwta. Ann. Mag. Nat. Hist. 
Jan., 1891, p. 96. 

Bepparp, Fr. E. Some New or Little Known Oligochswta. Proceedings of the Royal Physical Society, Edinburgh, 
June 13, 1893. 

Bepparp, Fr. E. On the Anatomy of Ocnerodrilus. Proc. Royal Society of Edinburgh, vol. xxxvi, p. 563. 

Micwagtsen, W. Terricolen der Berliner Zoologischen Sammlung II, page 14. 

Brenuam, W. B. An Attempt to Classify Earthworms. Quarterly Journal of Microscopical Science, vol. xxxi, 
part li, page 243, ete. 

Rosa, DanieLE. Sui Generi Pontodrilus, Microscolex, e Photodrilus. Boll. Mus. Torino, vol. iii. 

Rosa, DanieLE. Die Exotischen Terricolen des K. K. Naturhist. Wien. Annal. K. K. Nat. Hofmus. Wien, Bd. vi, 
p. 387. 

Eisen, G. Anatomical Studies on New Species of Ocnerodrilus. Proc. Cal. Acad. Sci., ser. 2, vol. iii. 

E1sen, G. Anatomical Structure of Two Species of Kerria. Proc. Cal. Acad. Sci., ser. 2, vol. iii. 

PERRIER, EpMoND. Etudes sur Organisation des Lombriciens Terrestres. IV Pontodrilus. Archives Zoologie 
Experimental, vol. ix, pages 175 to 248, pl. xiv. to xviii. 


Eclipidrilus frigidus Eisen. 


Through the kindness of a friend traveling in Sierra Nevada I have received a 
small number of specimens of this interesting oligocheta, but unfortunately all the 
specimens were in a poor state of preservation and much macerated. However 
I was enabled to make several very nearly continuous series of sections and thus settle 
several very important points in the anatomy of this worm. My former study of the 
species was entirely dependent on dissection, which could not possibly reveal all the 
details of this minute species, especially as regards the spermducts, the presence of 
which I am now able to demonstrate. My present researches show that the species 
is less erratic in its anatomy than I first supposed, while again in many respects it 
differs strangely from its nearest allies, the various genera of Lumbriculide. For 
the present I retain the family of Eclipidrilide, but not on the same grounds as 
formerly, and I now consider it rather as a subfamily to Lumbriculidz than one 
standing isolated, however with strong leaning towards Moniligaster. 

The generative organs are situated as follows: 

Testes, two pair. The anterior pair attached to the anterior septum of somite 
ix. The posterior pair similarly to the anterior septum in somite x. 

Ovary, one pair attached to the anterior septum of somite xi, 

Oviduct in xii, opening in front of the inner pair of sete. 

Spermathece, one pair in ix opening posterior to the sete and near the posterior 
septum. 

Atrium and prostate, one pair opening in x, posterior to the inner pair of sete. 
This organ, which is very long, occupying seven to eight somites, cousists of three 
parts, first, one anterior atrium and prostate proper, second, a thin and narrow part 


PACIFIC COAST OLIGOCHETA. 85 


connecting the former with, third, the posterior one, a storage chamber for sperm- 
atozoa. 

Spermducts are two pairs, exceedingly narrow, opening close together in the 
posterior part of the storage chamber. 

Ciliated rosettes, two pairs, the anterior pair opening in ix. The posterior pair 
in x, both in front of the septum. They are exceedingly thin and flat. The posterior 
wall is attached to the septum. 

Sperm-sacs consist of a pair of very long continuous sacs, which cover the 
generative organs, including the spermatheca, and extending from somites x to xvii 
or xvill, generally several somites posterior to the caudal part of the storage chamber. 

Setw are 8 in 4 pairs in each somite, commencing with the second. 

The vascular system is characterized by blind forked vessels in the posterior 
30 odd somites, thus bringing the genus in close relationship with Lumbriculide. 
Another characteristic is the two loops of lateral vessels which branch out from the 
main vascular trunks in somites ix and x, and which run backwards as far as the end 
of the storage chamber. The anterior eight somites contain winding lateral vessels 
connecting the two main dorsal and ventral trunks. Hertzkérper in the dorsal vessel 
as well as in the branches in the intestine. After these preliminary references to 
the main anatomical points I will enter more fully upon the anatomical and _histo- 
logical structure of the various organs. 


Body-wall and Clitellum. The finer structure must be left for future study. 
The inner longitudinal muscular layer is considerably thicker than the transverse 
layer and hypodermis together. The longitudinal strands are very thin and ribbon- 
like, some being much longer than others and reaching through the width of the 
layer, others being very much smaller, situated principally close to the transverse layer. 
Figs. 77, 79, 99, ete. 

The clitellum comprises about 6 somites, commencing in the posterior part of 
ix and extending to the center or posterior part of xiv. The clitellar glandular 
cells, one layer thick, are oblong, irregular, flask-like, containing very coarse, angular, 
grains (fig. 79). They are separated or interspersed by large non-staining cells. 
The peritoneum is rather poor in blood vessels, but the layer is very thick, in places 
almost as thick as the longitudinal muscular layer (fig. 82, pr). 


Septa. None of the septa are abnormally thickened. The first distinct sep- 
tum is seen between somites v andvi. The septa are straight, not cup-shaped. Those 
surrounding the various divisions of the prostate are much firmer than the others. 
They constrict the prostate, in fact the latter appears notched at every septum 
(figs. 78, 92, 94). 


Alimentary canal. The alimentary canal can properly be only divided in two 
parts—pharynx and intestine. The pharynx which ends in somite vy is developed 
latterally and dorsally, but not ventrally (figs. 83, 77). The thickened part is very 
thick, consisting of the usual narrow and almost filiform cells. The ventral part is 


86 CALIFORNIA ACADEMY OF SCIENCES. 


very thin, thinner in fact than any other part of the alimentary canal. Of the balance 
of this canal there is no distinction between cesophagus and sacculated intestine. The 
gut is every where sacculated, only increasing in thickness towards the genital somites, 
where it is thickest. The alimentary canal throughout its length is lined by a 
columnar, ciliated epithelium, outside of which is a very thick vascular layer, with 
large blood lacunes, directly connected with the dorsal and ventral vessels, which are 
closely attached to, or almost imbedded in the intestine. The latter as well as the 
vessels are covered with chloragogic cells, which, especially in the region of the 
dorsal vessel, are very large (figs. 85 to 91), the layer being thickest close to the 
strands of mesenteric tissue connecting the intestine with body-wall. 

The free coelomic lateral vessels in the eight anterior somites are similarly 
surrounded by a thick mass of glandular cells, arranged around muscular strands, and 
which are quite distinct from the chloragogic cells and more resemble real glands. 
Their reaction to stains is entirely distinct from that of the chloragogic cells of the 
main vessels and of the alimentary canal, staining very deeply with ammoniated 
hzematoxylon (fig. 84), and showing a coarser grainy secretion, while the real chlora- 
gogic cells remain much more pellucid and contain much finer grains. Cells similar 
to the former are also seen attached to the cceelomie covering of the prostate (fig. 
107, etc.) They also greatly resemble the glandular cells, or multicellular glands 
from the pharynx of Pontodrilus and other oligocheta, possessing pharyngeal glands. 

The vascular layer of the intestine is very much developed, especially on the 
ventral side, where it connects with the ventral vessel, through a thick band of mesen- 
teric and connective tissue. This as well as the walls of the blood vessels were so 
thickly studded with a protozoa (Hemagregarina) that the structure of the layer 
could not even in a single instance be properly made out. 

There are no pharyngeal glands, though a few glandular cells are seen scat- 
tered about between the muscular strands connnecting the pharynx with the body- 
wall. But these cells resemble more chloragogen cells than true pharyngeal glands. 

The ¢estes are of no unusual structure. The anterior pair, in the specimens I - 
opened, are smaller than the posterior pair, which were always forked, while the ante- 
rior pair was undivided. 

The ovary in xi isalways sigmoid of irregular shape and present the peculiarity 
that seldom more than one ovum is developed at a time, this one being situated not at 
the periphery or at the free end of the ovary, but in the inner angle of the sinus. The 
ovum is unusually small in size and readily detached from the gonad (figs. 78, 108). 
It grows large after separation, and is found in numbers in the posterior somites. 

The ovary is of large size reaching far back to the posterior septum. Its lower 
end is not only attached to the septum and body-wall, but also to the narrow end of 
the outer sperm funnel (or ciliated rosette) fig. 96. In one of the specimens sectioned 
the ovaries either extended past the oviduct through somite xii, or there was a second 
pair of ovaries in xii. Beddard has similarly remarked that the ovary in Sutroa is 
attached to the cells of the spermiducal funnel. 


PACIFIC COAST OLIGOCHETA. 87 


Spermatheca. As stated there is one pair situated in ix, the spermathecal pore 
being posterior to the sete, and in line with the inner couples. The spermathec: 
consists of two distinct parts, as is usual in this group, the lower muscular part, and 
an upper rounded part consisting of an unicellular layer of dice-shaped cells. This 
rounded chamber was sometimes situated in the same somite, ix, as the muscular 
body, but frequently it projected backwards into somite x. 

The structure of this muscular part is represented in figure 82. The inner epithe- 
lial layer, consists of narrow columnar cells, with rounded nuclei and striated proto- 
plasm. Exterior to this layer, there is a thin one of transverse or circular muscles, 
outside of which again is an epithelial stratum one or two cells deep, with slightly 
oblong nuclei, the cells themselves being irregularly dice-shaped (fig. 82). 


Prostate and spermducts. The most interesting, as well as the most complicated 
structure of this species, is the prostate, of which I am now able to give a fuller ac- 
count, which I believe will not leave any of the points of its structure in doubt. 

I have already referred to the three main divisions of the organ, the proximal 
one consisting of a long cylindrical tube, containing penis, atrium and _ prostate. 
Second, a very narrow tube of almost the same structure as the prostate part of the 
former, connecting with a long cylindrical chamber of somewhat modified structure, 
into which the two spermducts open, quite near its junction with the narrow part or 
bridge. If we, however, disregard the difference in size of these various parts and 
only consider the structure, we find that the whole organ may also be divided in 
three parts: 

First, the proximal part, which is entirely confined to somite x. This part is 
upright, so to say, does not run backwards; it is also somewhat bent, forming a right 
angle with the balance of the organ. This part consists of penis, and a long tubular 
part which, in want of a better name, I designate as atrium (fig. 100 qtr. and p.). 
With somite xi commences a change of structure of this organ, common both to the 
wide part and to the very narrow posterior part. I will refer to it as the prostate 
proper, as it contains the thick layer of regular prostate cells so common in all higher 
oligocheta where this organ occurs; it is the “two layer”—prostate of Beddard. The 
third part or storage chamber is characterized by the absence of this layer. 

I will now refer to each one of these three or four parts in succession and more 
in detail, beginning with the penis and atrium. 

Next to the body-wall, ending at the transverse muscular layer and from there 
stretching inwards, is a reversible sae—a preputium—consisting of epithelial cells, 
with very large, round, compressed nuclei and striated contents. This epithelium is 
surrounded by a thin muscular layer (fig. 100, pre., 101). 

Into this preputium opens a penial glans (fig. 100 p. gi/s., 101, ete.), consisting of 
two separate covers, posteriorly attached to a collar of larger rounded or pear-shaped 
cells, at the base of which are seen a number of muscular plates. This collar is 
folded on itself, one part of the fold connecting with outer and one part with the 
inner cover of the glans. Through the median line of these parts runs a long, very 

Memorrs, Vou. II, 4. February, 1595. 


88 CALIFORNIA ACADEMY OF SCIENCES. 


narrow excretory tube with thick walls. Nearest the male pore this tube is quite 
straight, or only slightly folded, but at its distal end the windings are greatly packed 
or irregularly coiled (fig. 99, tube). 

The part posterior to the penial collar I have designated the atrium proper, 
simply because it does not contain any inner layer of glandular cells, and because in its 
distal end it connects with the true prostate. The excretory tube continues all through 
this part and is, so to say, suspended in a more or less dense mass of fibrous tissue. 
Nearer the collar this mass is reduced to a few strands only, but towards the distal end 
it becomes quite dense, especially so nearest the tube, while towards the periphery it is 
much less dense (fig. 92, f. ¢., fibrous tissue; /. st., fibrous strands). Fig. 100 repre- 
sents a cross-section taken near the distal end; the tube is seen coiled in the center, 
while fibrous tissue connects it with the walls of the atrium. 

The exterior layers of the atrium are constructed very much as the same lay- 
ers of the prostate proper and the storage chamber. There are three layers which 
are common to the two different parts of the prostate. Interiorly a thin layer of cir- 
cular muscles somewhat variable in thickness in different parts, but generally only 
three or four strands thick. Exterior to this layer runs a spirally wound layer of lon- 
gitudinal muscles, arranged in band-like plates, and when seen in cross-section re- 
sembling a row of fringes (figs. 100, 101, 102, 103, 106, 107,7.m). Fig. 107 shows these 
plates, highly magnified, to consist of rather rectangular muscular strands. Exterior 
to this layer of /. m. muscles we find everywhere a broken row of prostate glands of 
very minute size, appearing to penetrate in between the muscular plates, though on 
account of the macerated condition of the tissues I could not follow their tubes. These 
small glands, which barely project outside of the muscles, are found everywhere from 
the distal end of the storage chamber to the penial collar, wherever this muscular 
layer isfound. In places they are continuous, in others scattered about, seldom more 
than one row thick (fig. 101, g/s). But as we reach the region of the narrow part of 
the prostate and especially the part of the storage chamber where the spermducts 
enter, we find another thicker layer of prostate glands similarly scattered over the 
longitudinal muscular fringe (figs. 102, 103, 106, 107, pr. g/. s, small prostate giands; 
pr. gl. l., large prostate glands). In the region where the spermduets enter the stor- 
age chamber this layer of large prostate cells is almost continuous and considerably 
thicker than the muscular layers combined (fig. 103). 

The prostate proper contains besides these jiayers of muscles and exterior 
prostate glands two inner layers, which resemble and propably correspond to the two 
cell-layers found in the prostate of the higher oligocheta, viz.: one layer of lining 
epithelium (fig. 101) and one layer of glandular prostate cells, with very large nuclei, 
and separated one from the other by transparent spaces, through which possibly enter 
projections of the exterior prostate glands. These two characteristic layers do not 
extend to the narrowest part of the prostate. Hence the muscular layer is covered 
by a single row of inner lining epithelium with large, slightly oblong nuclei (fig. 
102), which in the very narrowest part are about 12 to 14 in the row. 

The proximal end of the storage chamber is lined by a very thick epithelium 


PACIFIC COAST OLIGOCHATA, 89 


with very oblong nuclei (fig. 105 ep.). This epithelium is thickened only at the 
anterior side, nearest the narrow tube (fig. 104), but narrows quickly both superiorly 
and posteriorly. The inner transverse muscular layer is similarly thickened in this 
region (fig. 105, ¢. im.). 

The storage chamber contains the same general layers as the narrow bridge. 
The inner lining epithelium is narrow, with very compressed nuclei, with the flat side 
lying against the muscular wall (fig. 107). 

The nature of the prostate differs apparently much from that of the prostates of 
some members of Lumbriculidee, especially Sutroa, as lately more minutely described 
by Beddard, but more resembles in structure Lumbriculus. But I am more inclined 
to compare its makeup with the prostate of Moniligaster. In Eclipidrilus the pros- 
tate contains the characteristics of both Limicole and Tericol, if I may yet use the 
expression. ‘The inner two-celled layer of the prostate which is characterizing most 
higher earth worms, possessing a prostate, is superposed by the layers, muscular and 
glandular which characterize Tubifex, Moniligaster, ete. 


Spermducts. IT am now able to describe the spermducts for the first time. 
They enter the storage chamber of the prostate near to the bridge and close to- 
gether, but still entirely separate (fig. 95), and when the inner surface of the 
chamber is viewed from above the entrance pores are seen as two small slightly ele- 
vated papilla (fig. 95 sp.) From these the spermducts, which are of very minute 
size, run forward parallel to the prostate, one on either side, except alongside of the 
bridge where they run close together. The outer pair leave the prostate in xi, dip 
down to the ovaries in which they are partially engaged and push their ciliated 
rosettes through septum x/xi, the rosettes opening in x. The inner spermduets are 
similarly engaged in the gonads in x, push through septum ix/x and open their 
rosettes in ix, all very close to the body-wall. 

The rosettes are only one cell thick, very thin and flat, with the posterior sur- 
face attached to the septum, the anterior lip only being free. Only the inner deeper 
surface of the rosettes is ciliated. 


Sperm-sacs, one on each side and continuous from the beginning of the sperm- 
atheca, reach as far back or further than the posterior end of the storage chamber of 
the prostate. Generally the sperm-sacs reach two or three somites further back. 
Each sac is separate from the other and consists of one continuous bag contracted 
somewhat at the septa. It is not racemose and does not connect with the septa as in 
so many of the higher forms, but greatly resembles those of Sutroa. The sperm-sac 
only covers the prostate, but does not properly enclose it, as it does not extend to the 
space between the prostate and the intestine. With the latter, however, it is con- 
nected by two continuous walls of connective tissue, one on the dorsal and one on 
the ventral side of the intestine. In the enclosure thus formed the prostate, as well as 
the upper part of the spermatheca, lies free. 

In the posterior parts of the sperm-sacs are always seen very large sacs of yolk 
granules. In one specimen I found a mass of these yolk granules surrounding the 


vO CALIFORNIA ACADEMY OF SCIENCES. 


spermatozoa in the prostate storage chamber (figs. 120 to 123). The yolk sacs are 
frequently so large that they fill the larger part of the caelomic cavity pressing 
the intestine close up against the wall of body. 


Nephridia. The most anterior nephrostome is found in iii in front of septum 
iii/iv. The most anterior nephropore is found in iy in front of the inner couple of 
setee. There are nephridia (pores) in iv, v, vi, vii and viii, while somites x, xi, xii 
have no nephridia. The neck of the nephrostome contains a large glandular swell- 
ing, to which is attached museles connecting with the septum. This neck is perforated 
by several narrow ductules, which occasionally branch. The duct appears to con- 
sist of a single tube not covered by peritoneal glands. On account of the rather mac- 
erated condition of the specimens I could not investigate the nepridial structure any 
closer. 

Some of the specimens collected by my friend were from a new locality and 
somewhat larger in size. This locality is Three Spring Meadow, on the east 
side of the North Fork of Kings River, opposite the natural bridge, at an altitude of 
about 8,000 or 9,000 feet, and several thousand feet above the river bottom at that 
point. They were there found under some old logs lying across the meadow, and over 
which the water was flowing, the worms being attached to the surface of the decayed 
wood. These specimens were about 4 larger than those from the springs at Alpine 
Meadows on the South Fork of Kings River several thousand feet higher up, but I 
find no distinct characteristics, though the spermatheca appeared more twisted. 
Among the specimens were a few of Ze/matodrilus Vejdovskyi, for which I am thus 
able to note a new locality. 


. 


>) 


PLATE XXX. 


Fi 


, 


92 


1A. 


i) 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXX. 
PH@NICODRILUS TASTE. 


Semi-diagrammatic view of the body in a longitudinal vertical section, showing the position and proportion of 
the various organs. pha. pharynx. sl. gl. salivary glands. sp.gl. septal glands. sp.s. sperm-sacs. cl. cli- 
tellum. h. hearts and dorsal vessel. @. cesophagus. spth. spermatheca. divi. diverticulum of cesophagus. 


t. testes. si. sacculated intestine. @. ovipore. ov. ovary. 4 male pore. 

View of the interior organs, the body-wall being laid open from above. Onthe right hand side the sperm- 
sacs haye been removed in order to show the underlying organs. The letters indicate the same as in figure 
one. The shaded part of somites xiv to xviii indicate the clitellum proper. ac. accessory copulatory papillz 
around the outer set# in somite xiv. 

Longitudinal section through some of the anterior somites, from a section lateral to the esophagus, illustrating 
the connection between the septal and supra cesophagal glands and their ducts, which all open into the upper 
wall of the pharynx. s. septum. 6d. w. body-wall. si. gl. salivary or supra pharyngeal glands. ss. gl. sep- 
tal glands. 


Spermatheca in outline. 


The spermduct and male pore viewed from above or from the interior of the body. Both are covered by the 
longitudinal muscles, as well as by numerous arciform muscles. 


sr 


LTH BRITIONS REY, 


RS 


» \ 
~~ 3 
=A 


DH, Pare AK 


4 


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= 

A = “st 
sae SS 


sa a a Aled 


/ 704. 


HGS. 


JE 


ae 
SS ee 


t 
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o 
S 


aa 


> 


i eee eer a 


PROEMICODRILUS TAS 


PLATE 36xX1 


94 


5. 


GA. 


6B. 


ifs 


8. 


9A. 


9B. 


10. 


aie 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXI. 
PH@NICODRILUS TASTE. 


A transverse section of the body-wall in somite xvii, through the male pore showing the degenerated atrium 
and the wider upper part where enters the spermduct. These latter have just been fused and enter the pore 
as one single duct. cl. clitellar cells. Ay. hypodermal layer. cl. collar of epithelium of the atrial chamber. 
ac. m. arciform muscles. f. sp. fused spermduct just entering the atrial chamber. /. m. longitudinal mus- 
cular layer of body-wall. ¢r. m. transverse muscular layer of body-wall. ¢ male pore. 


The section nearest anterior to the former, showing the spermducts just as they enter the atrial chamber. 
The spermducts are not yet fused. Figures indicate the same as in the preceding figure. sp. spermducts just 
before being fused together. 


Section nearest anterior to the former showing the two spermduets, etc. 


The three somites xvi, xvii and xviii viewed from the interior, the alimentary canal and nerve-cord having been 
removed, as well as nephridia. 


One of the sete» magnified, the same relative proportion as that used in my paper on Ocnerodrilus. 


One of the nephrostomes seen in front views. When in proper position the face of the rosette stands 
parallel to the septum. 7. rosette proper. 7. neck with very large glandular cells. m». d. narrow duet. 
s. septum. f. fold of the main nephridial body. 


Nephrostome seen from the side, the posilion when the body is opened and spread out. Letters indicate the 
same as in the last figure. 


One of the museles connecting septal glands with body-wall, in cross-section. s. septum. m. muscular 
fascicle. 


One of the posterior nephridia seen under a low magnifying power in order to show the peritoneal covering 
and the general outline of the canals in the lower peritoneal lobe. a. f. anterior fold of canals. p.f. posterior 
fold of canals. spu. the partially free spur. s. septum. sf. nephrostome. np. nephropore. w.t. the 
wide tube leading to the nephropore. br. bridge connecting the spur with one of the folds. wu. /. upper 
lobe of peritoneal cells which does not contain any canals. c¢. 1. connecting lobe. J. 1. lower lobe. The two 
peritoneal lobes vary greatly in shape and hardly two are found exactly alike, though the |general form is the 
same in all. 


AL ACAD. Plate 0, 


Meu. 


all 


vf 


pris. laste Figs. Soll. 


Pren 


be 


ATE 


16A. 


16B. 
16C, 
16D. 


16E. 
16F. 


CALIFORNIA ACADEMY OF SCIENCES, 


PLATE XXXII. 
PHG@NICODRILUS TASTE. 
One of the posterior nephridia more highly magnified, to show the course of the canals. 


Phoenicodrilus taste, natural size, the worm having first been killed gradually with increasing strength of 
alcohol. Largest specimen. 


Section of the intestine at the point of connection with the diverticula, in order to show the arrangement 
of the blood vessels and the subdivisions of the diverticula. This figure is taken at the junction and shows 
the diverticulum as one chambered, and with a large blood vessel penetrating the diverticulum from the 
outside. This blood vessel comes from the dorsal vessel. This and the following five figures are outline 
drawings from slides, but the details of less importance are not filled in. 6. small vessels in the vascular 
layer of the alimentary canal. dv. diverticulum. Dl. v. large blood vessel penetrating the diverticulum 
from the exterior. ep. epithelial lining of the intestine. m./. muscular layer of the intestine. 


Section elosely following, or anterior to the former, showing the diverticulum as one chambered and with a 
division of the blood vessels into smaller ones. Letters indicate the same as in the former figure. 


A section of one of the diverticula a little anterior to the former, the organ is yet one chambered, and a large 
blood vessel is seen on the left side or the side nearest the intestine. J/. b/. v. very large vessel. b/. v. small 
vessels in the wall of the diyerticulum. 


A section anterior to the former, showing the beginning of the subdivision of the diverticulum. 
A section next anterior to the former, showing the diverticulum two chambered. 


A section near the center of the diverticulum, showing six interior chambered and a multitude of blood 
vessels. 


A part of diverticulum more highly magnified, showing nuclei and striated cycloplasm. 


Section of alimentary canal, 


PACIFIC COAST OLIGOCHATA. 


dS 


19. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXIII. 
PHC@NICODRILUS TASTE. 


Longitudinal section of the alimentary canal (cesophagus), in vi. ep. inner lining epithelium. gl. glandular 
cells. 61. blood sinus. m. the two muscular layers. pt. peritoneal epithelinm. 

Longitudinal section of the pharynx, showing the glandular ducts and ductules penetrating the pharyngeal 
epithelium. cl. ciliated surface. mn. ep. nuclei of epithelium. dus. ductules. duct. large duct from the 
pharyngeal glands aboye the cesophagus. @. ep. cesophageal epithelium joining the pharyngeal ciliated 
epithelium. 


Section of the body through one of the clitellar somites, a diagrammatic view, to show the relative size of the 
different layers of the body-wall. d.v. dorsal yessel. s. i. saculated intestine. m. /. the muscular layers. 
cl. c. clitellar cells. sp. d. spermducts. v.v and. c. ventral vessel and nerve cord. 


A diagrammatic section of the body in somite ix showing the relative size, etc., of the diverticula and the 
layers of the body-wall. d.v. dorsal vessel. sp. s. sperm-sacs in somites ix and x. div. diverticula of the 
intestine. v. v. ventral vessel. mn. c. nerve cord. 


Section of body-wall of a non-clitellar somite. g.c. unicellular glands. s. c. supporting cells. ¢. m. trans- 
verse muscular layer. /. m. longitudinal muscular layer. pr. peritoneum. 


KERRIA McDONALDI. 


Nephridium of Kerria McDonaldi from one of the somites closely posterior to the clitellum. pr. s. peritoneal 
sac with nuclei. The posterior end of this sac is not furnished with blood capillaries. c¢. 6/. capillary blood 
vessels, spreading principally through the peritoneal sac. a. 1. anterior lobe. p./. posterior lobe. nephr. st. 
nephrostome. nephr. pr. nephropore. w.d. wide duct outlet. x. d. narrow duct. nec. neck of the 
nephridium. 67. bridge. spr. spur. s. septum. 


PONTODRILUS MICHAELSENT. 


A specimen, natural size, having been slowly killed and extended. One of the largest specimens, the majority 
being only one-half as large. 


The three anterior somites seen from the side. pr. prostomium. per. peristomium. 


The anterior part of the worm more highly magnified, ventral view. cl. clitellum. ¢ male pore. spth. p. 
spermathecal pores. ov. p. ovipore. c.pr.m. cushion where ends the fan-shaped fascicle of prostate or arci- 
form muscles. ec. ¢. exterior copulatory cushion. 


Part of the body-wall iaid open and spread out to show the relative distances of the set, the numerals 
indicating the number of the seta. The ventral nerve cord is indicated. 


One of the setw highly magnified. 


Meu CAL Aso 1 Pare YO 


‘ 

. 
Ly 

‘ 

Fonan-cileeslort court. \ 
7 
. f 
-— eka 

bs = 
SusvEen.Des, ITH BRITTON SREY SE “I 


> Piemcoormus: Mase “Ties. 7c. = Heme M'Dowetal Hie &3. 
= —  — Paomius Mowetsem Fos 241028. ue 


ey ee " 


ie en ay soa us y ; a 
’ he yo ke? os a a a at i : dar: r ater 


PACIFIC COAST OLIGOCHMTA. 


PEAR SOOCLV: 


00 


y 
4 


100 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXIV. 
PONTODRILUS MICHAELSENI. 


29. A somewhat diagrammatic view of the anterior somites seen in longitudinal section. br. brain. mo, mouth. 
ph.c. pharyngeal cavity. pha. pharynx. m. muscular strand connecting pharynx with the body-wall. sl. gl. sal- 
ivary or pharyngeal glands opening by ducts in the pharynx, the ducts being supported by muscular strands 
(stained yellow). ss. gl. smaller gland; connected with blood vessel. g/. glands connected with the lateral vessels. 
w. esophagus. d.v. dorsalblood vessel. s.septa. spth. spermathecw, and their diverticula. The lower end 
of the spermatheca is greatly enlarged, forming propulsory bursa. spth. p. spermathecal pore. ¢. testes. 
ov. ovaries. ovd. oviduct. c. 7. ciliated rosettes. h: hearts. sp.s. sperm-sacs. These sacs are principally 
ventral to the esophagus. nephr. nephridia, the most anterior one is found in somite xiii. cl. clitellum. gl. cr. 
glandular crop in shape resembling a gizzard. s. 7. sacculated intestine. pr. prostate. ¢ the muscular duct 
of the prostate eut through. c.c. copulatory cushion. The dotted line indicates the outlines of the duct as 
they appear in other sections. 

30. Section through the posterior part of the pharyngeal glands, showing the arrangement of the glands, ducts, 
muscles and blood vessels, all in vertical section. «@. esophagus. ep. inner epithelial lining. m. @. circular 
cesophageal muscles. m. muscular strands. phx. gl. pharyngeal gland. d. v. dorsal vessel. 

In this figure the glands are stained violet, the muscles appear as yellow and the ducts are dark violet. The 
blood vessels are black (Corosive sublimate, abs. alcohol, orange G. amm. hematoxylon. Thus in xylol. 
This and the following five figures are all drawn from sections treated in the same way.) 

31. A longitudinal section of a lobe, showing the general arrangement of the glands and ducts. duc. ducts from 
the gland, leading anteriorly to the pharynx. gl. c. glandular cells. _6/. gi. blood glands. m. supporting 
muscular strands. 6l. v. blood vessels. 

32. A small glandular lobe more highly magnified. The letters indicate the same as in the preceding figure. The 
nuclei are stained yellowish with orange G. The secretion of the cells is precipitated? and stained dark violet 
with amm. hwmatoxylon, 

33. <A part of a gland, with supporting muscles. 

34. Two sections of muscular strands, showing the connection with the ducts in which the secretion has been 
stained deep blue. 

35. One of the glandular cells with nucleus stained yellow. ‘The vacuoles (va.) are white, the protoplasm grayish, 
while the secretions are stained deep bluish-violet. 

36A. The proximal end of two glandular ducts where they penetrate the epithelial lining of the pharynx, near its 
dorsal side, showing the ductules or end ducts leading between the epithelial cells. 67. blood vessels. m. ph. 
supporting muscles for the ducts connecting with pharynx. ¢. m. transyersal muscles cut across. gl. d. 
glandular ducts leading from the gland at the distal end beyond the pharynx. These ducts are the largest 
and collective ones, having received the secretion from many smaller ducts. ep. epithelial cells lining the 
pharyngeal cavity. gl. dt. glandular ductules passing between the former into the pharyngeal cavity. ss. gl. 
two of the smallest of the salivary glands, a few of which are found at the periphery of the pharynx. 

36B. A longitudinal section of one of the small septal glands, one pair of which is found in vy, vi, vii, vili, ix. 
c. bl. v. connecting blood vessel between the ventral vessel c. b/. yi and the dorsal vessel. This vessel is 
cut in two places, the one marked ec. b/. vii being nearest to and just below the esophagus (a. v.) the ventral 
outline of which only is figured diagrammatically. s. septum, outline of, between vi/vii. g/. small glands, a 
few of which surround the somewhat larger septal gland. s. gl. the largest septal gland. w. ¢. wandering or 
perigastrie corpuscles. d. ¢. discharge ducts from glands, probably connecting with the pharynx. 

37. Cross-section of an immature specimen, through the intersegmental groove between somites xvii/xvili, just 
anterior to the copulatory cushions. In this section no clitellar cells haye yet developed on the ventral side 
of the body. hy. hypodermis. ¢. m. transverse muscular layer. 7. m. longitudinal muscular layer. neph. 
nephridium. s. a. septum between xvii/xviii. m. muscles connecting the glandular part of the sperma- 
theca with the body-wall. gl. pr. glandular part of the prostate. spd. the sperm-duct just entering the 
glandular prostate. m.pr. muscular part of the prostate. v. ne. ventral nerve cord. ss. i. sacculated intes- 
tine. 

38. Cross-section of the body of an immature specimen, through somite xviii, the section passing through the 
copulatory cushions and through a part in which the clitellar cells on the ventral side of the body are already 
developed. «a. m. arciform muscles connecting the copulatory region with the dorsal part of the body-wall. 
el. c. clitellay cells developed in this somite only on the ventral side, never extending to the dorsal side even 
in fully mature specimens. Other letters indicate the same as in the preceding figure. 

39. Cross-section through somite xiii close to the anterior septum. ‘The clitellar cells are here fully developed. 
s. 8. part of septum between xii/xiii. cl. ¢. clitellar cells, only developed dorsally. v. ¢/. ventral termini of 
the clitellar cells. ov. ovaries. spd. spermduct. ¢. 7%. tubular intestine. dv. dorsal vessel. h. hearts. 


Tras sy, 
ere a 


‘ ay ; \ Vase 
<9) Se. m oes oe 


: 
[ 


{ 


DRILLS MICHAEL SRM FIGS. 29.19 39 


H) 
1 
t 


PACIFIC COAST OLIGOCHETA. 101 


PLATE XXXV. | 


. ‘ 
‘ 
; 
G 
, j 
s 2 * 3 
; 
= A , 
ae 
: 
“ -_ 
® S = 
; 
ee: : : 
1 eel od : 
f Y = = 


102 


40. 


41. 


42. 


43. 


44. 


CALIFORNIA ACADEMY OF SCIENCES, 


PLATE XXXYV. 
PONTODRILUS MICHAELSENI. 


Cross-section through somite xvii of a mature specimen showing the development of the clitellar cells on the 
dorsal and lateral sides of the clitellum. vv. cl. ventral termini of the clitellar cells. d. v. dorsal vessel. 
s. i. sacculated intestine. neph. nephridium. gl. pr. glandular part of prostate. pr. m. muscular part of 
prostate. m. muscles connecting the prostate with the body-wall. 4. m. transverse muscles. /. m. longi- 
tudinal muscles. cl. clitellar cells. v. cl. ventral termini of the clitellar region. 


Cross-section through a strongly contracted, fully matured specimen, through somite xviii, showing the copula- 
tory cushions as fully developed, as wellas the clitellar cells on the ventral side. Unfortunately the only fully 
developed specimen at my disposal had been badly contracted, hence the cells appear smaller than they would 
have done in properly prepared specimens. c.c. copulatory cushions. a.m. arciform or fan-shaped muscu- 
lar strands, which contract the copulatory region. i. ¢. m. interior transverse muscular strand, ending 
respectively in the copulatory cushions and in the projecting ventral lobes of the body. Other letters as in 
the last four. figures. 


The body-wall viewed from the inner side of somites xvii, xviii and xix, showing the genital region of the left 
side. pr. m. prostate muscles connecting this organ with the body-wall. «a. m. arciform muscles connecting 
the copulatory cushion with the upper body-wall. c.c. copulatory cushion. sp. m. muscles connecting the 
body-wall with the prostate at the junction with the spermduct. sp.d. spermduct. 7 and 2s. first and 
second seta. v.c. ventral nervecord. gl. pr. glandular prostate. m. pr. muscular prostate. s. m. smaller 
muscles, fan-like arranged, confined entirely to the copulatory cushion. sp. d. p. spermidueal pore overlaid 
by muscles. s. septa. 


A somewhat diagonal longitudinal section through the body-wall and copulatory cushion in somite xviii. The 
section is parallel with the muscular fan of arciform muscles, passing through the copulatory cushion near 
but not through the body-wall. Illustrating the arrangement of the muscular strands around the prostate in 
the vicinity of the male pore. The letters indicate the same as in the preceding figure. 


A longitudinal section through the copulatory cushion aud the body-wall, the section being almost vertical. 
pr. prostate, just leaving the arciform muscles, and immediately before bending down towards the prostate 
pore. m. muscles connecting the prostate with the body-wall. «a. m. the entrance of the arciform muscles 
into the copulatory cushion. 


A horizontal section of the body through somite xviii and the copulatory cushions, showing their size and 
position, ete. 


Mew (AL Acab HT. “Ptate MAY 


J 
. 
LTH SRTIOREREY SE 
is 


ae @ 1? _—_ 


e : 
PACIFIC COAST OLIGOCHATA. 1038 
i es 

= mete. ak tet : ibs r 
PLATE XXXVI. ; | 
Me 
Fi 
, e 
« , Le, 
4 os ; 
' . - = °5 


t 
= 
« 
Pe 
~ aJ 
7 
- . A 


a 


104 


46A. 
46B. 
46C. 
47. 


48. 


49. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXVI. 
PONTODRILUS MICHAELSENT. 
Section through the rudimentary gizzard or thickening of the intestinal wall in somite v. 
One of the circular muscular strands of the former figure. 
Section of the alimentary wall adjoining posteriorly to the rudimentary gizzard in somite vi. 


A longitudinal section through the intestine showing the glandular-crop in somites xiv, xv, xvi, and the be- 
ginning of the sacculated intestine in xvii. d.v. dorsal vessel. d.v.c. connection between the dorsal vessel 
and the vascular system of the crop. gl. cr. glandular crop. bl. s. blood sinus in the outer layer of the 
glandular layer. gl. c. glandular cells constituting the bulk of the crop. clo. chloragogen cells. m./. the 
two muscularlayers. ¢.7. the end of the tubular intestine or esophagus. 


A section of the former more highly magnified, showing the glandular cells which constitute the crop and the 
branched lumens which are especially pronounced in the outer part of the layer. /.m., t. m. the two mus- 
cular layers, which here are not more developed than in any other part of the intestine. 


Cross-section through the glandular crop in the posterior part of the organ in somite xvi. In this section only 
few of the glandular cells are found. 


A section anterior to the former, but inthe same somite. Here many more of the glandular cells are seen. In 
these two sections the chloragogen layer is thick and the cells crowded, similarly the ridges of the epithelial 
cells are high and separated by deep grooves. 


A similar cross-section through the crop in somite xv. Both the chloragogen cells and the epithelial cells are 
lower. 


Another cross-section showing the entrance of a blood vessel from the dorsal vessel into the vascular system 
of the glandular crop. 


A part of the glandular cells of the crop more highly magnified showing the structure of the cells and the 
branched lumens between them. g/l. c. glandular cells. /. lumen. 6/. blood sinus. ep. inner epithelial 
cells. 


An oblique section through the surface of the crop showing the two muscular layers, a blood sinus and a few 
chloragogie cells cut through. The glandular cells immediately below are not delineated. 


105 


PEATE. <x vil. 


< 


or 
or 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXVII. 
PONTODRILUS MICHAELSENI. 


A spermatheca with its diverticulum seen from interior of the body, whenthe body-wall is spread out. spth. p. 
spermathecal pore. d. diverticulum. spz. inner sac-like enlargement where the spermatozoa are massed. 


A cross-section of the body-wall through the spermatheca near its pore. v. /. thin vascular layer or perito- 
neum. /. m. longitudinal muscles. ¢. m. transverse muscles. hy. hypodermis. spz. spermatozoa. int. n. 
interstitial nuclei, en. epithelial nuclei and cells. c. m. circular muscles of the spermatheca. div. diver- 
ticulum. 


A cross-section of the diverticulum of the spermatheca. in. inner nuclei of the epithelium. ints. n. inter- 
stitial nuclei. c.m. circular muscles. 0b/. vascular layer. gl. c. exterior glandular layer consisting of a 
double or triple row of cells. 


A longitudinal section of the body-wall, somites vii, viii and ix made through the spermathecal pore. spth. p. 
spermathecal pore. spth. the lower, enlarged part of the spermatheca, which forms a strongly muscular 
papilla. dv. the junction with the diverticulum. set. seta. J. m. longitudinal muscles. ¢. m. transverse 
muscles. a. s. anterior septum of somite yili. p.s. posterior septum of somite viii. 


One of the testes. 

A longitudinal section of one of the testes. 
An ovary. 

Longitudinal section of an ovary. 


Longitudinal section of the oviducalfunnel. s. septum. 


FyvronrLus Mictuznsenr Figs. 55 1063 
: oe ta GS. 306) 


- 


PACIFIC COAST OLIGOCHMTA. 


PLATE XXXVIIL._ 


107 


:, 


108 


67. 


68. 


69. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXVIII. 
PONTODRILUS MICHAELSENI. 


Section through the body in somite xi, showing the sperm-sacs and their relative position. s. anterior septum. 
d.v. dorsal vessel. A. hearts. @. esophagus. v.v. the branched ventral vessel. sp.s. the racemose 
sperm-sacs situated much ventrally of the intestine. n. c. ventral nerve cord. /. m. and ¢. m. longitudinal 
and transverse muscles of the body. 


A cross-section of the same region more magnified. s. x/xi. septum between x/xi. s. xi/xii. septum between 
xi/xii. sp.s. one of the sperm-sacs. spz. spermatozoa. m.c. mother cells. c. 7. ciliated rosette. m. m. 
muscles connecting the septum below the sperm-sac with the cesophagus. 


A more longitudinal section of the ciliated rosette, showing the muscles and the blood vessels in the thick 
funnel. 


Another section through the ciliated rosette, showing the thick layer of blood vessels below the ciliated 
epithelium. 


A cross-section through the glandular part of the prostate, near the entrance of the spermduct. ep. epithelial 
cells with round nuclei. c.m. circular muscles. c. glandular cells. m. muscles connecting with the body- 
wall. sp. d. spermduct. 


Cross-section through the muscular part of the prostate. 


LITRERITIONR REY SE 


rm nies Py PYRE 


Meu Ca Aco ll. Pare AOU 


G40 62. 


GS 


Fowroorius Micwensen Fi 


ee | . . 
e | PACIFIC COAST OLIGOCHETA. 109 


Pi 


PIRATE SKK: 


‘5 
; 
@ 


110 


70. 


~1 
tw 


73. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XXXIX. 
PONTODRILUS MICHAELSENI. 


Cross-section of the body in somite xi, showing the relative thickness of the septa and body-wall, the septa 
being very much cupped, showing the cesophagus, ete., of the somite ix, as well as the septum and ciliated 
rosettes of x. v.v. x. the ventral vessel in x where it connects with the hearts. v.v.ix. the two forms of 
the ventral vessel. gl. septal gland surrounding the connecting vessels in somite ix. There are five pair of 
these glands, one each in vy, vi, vii, viii, ix. 


One of the nephridia isolated. p./. posterior lobe of the peritoneal sac of the nephridium. a./. anterior 
lobe of the same. w. windings of the canals where the two folds meet. p. /. and a. f. posterior and anterior 
folds of the canal. x. p. nephropore. x. st. nephrostome. spr. spur with four canals. br. bridge. 
bl. v. blood vessels on the nephridium. 


Nephrostome isolated. m. marginal cells. w.d. wide duct. nec. neck in which the narrow duct is seen to 
be branched. > 


An enlarged and somewhat diagrammatic drawing of the course of the ducts, etc., of the nephridium. The 
letters indicate the same as in the preceding figure. The canals have been represented as further apart 
than they are in reality, otherwise their course could not have been clearly delineated. When the nephridium 
is viewed from above mounted in glycerine, the outlines of the canals are only dimly discernible, being greatly 
obscured by peritoneal cells and blood vessels, neither of which have been delineated. The shape of the 
neck varies tosome extent, in some specimens being much wider than in others. 


Various forms and sizes of blood-glands from the supra pharyngeal and septal salivary glands. Drawn from 
paratine sections, hardened in Formaline, and stained: a. orange G. alcohol, Ehrlich’s hematoxylon amm. 
b. rose aniline in hydrochloric alcohol, Bismark brown, and Ehrlich’s hamatoxylon. The latter combination 
gives by far the finest results, clearly differentiating the blood from the gland-secretion, this being very 
imperfectly done by the orange G. 


ESTHBRITIONS REY, SF 


<x 


PACIFIC COAST OLIGOCHATA. 


PLATE XL 


dit 


112 


76. 


fle 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XL. 
PONTODRILUS MICHAELSENI. 


Cross-section through the body-wall and through the nephropore and wide duct. The latter is seen to enter 
the glandular neck of the nephridium (vec), its inner duct haying been cut twice, these cross-sections 
being always of unequal width. The section a passes through the distal windings of the ducts, the letters 
in the duct indicating that they belong to the same duct. c. c. ciliated canal, always wider than the other. 
The canal c. is a branch or forking of one of the other ducts. At d. about an inch of the drawing is left out 
in order to economize room. 


Longitudinal section of the nephridium near the windings showing the forkings of one of the canals. c. c. is 
the ciliated larger canal. 6. is the body of the nephridium, in which are seen the branched canals and lacunes, 
as well as the branching blood vessels which are held back. 


Part of the neck in cross-section, a larger magnification of a section similar to a 74, showing the branching of 
the narrow duct n.d. The wide duct w. d. ¢. is cut twice; the part marked w. d. is the part of the wide duct 
as it leaves the neck for the exterior pore of the body-wall. 


ECLIPIDRILUS FRIGIDUS. 


A diagrammatic view of the anterior somites of the body, composed from several longitudinal sections. Only 
part of the vascular system is shown, and none of the nephridia, the principal object being to 
show the arrangement of the reproductive organs and the shape of the alimentary canal. Sperm-sacs not 
shown. br. brain. bd. body-wall. pha. pharynx. c.v. connecting vessels. dd. v. dorsal vessel. v. v. 
ventral vessel. c/. clitellum. v.c. ventral nerye cord. 1+. first and second pair of testes. spth. spermatheca. 
ov. ovary. ovd. oviduct. c. 7. inner and outer ciliated rosette. ci. clitellar layer of glands.  sps. 
spermatozoa. ¢. testes. chlo. chloragogen cells. ¢ male pore. jp. penis. atr. & pr. prostate and 
atrium. pr. m. muscular layer of prostate. pr. g. glandular part of prostate. aér. atrium. sér. storage 
chamber for protozoa, spd. spermduct. int. intestine. br. bridge. 


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PACIFIC COAST OLIGOCHATA, 113 


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114 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XL 
ECLIPIDRILUS FRIGIDUS. 


78. A diagrammatic figure of the region of the reproductive organs, including somites ix to xvii, showing the arrange- 
meuts of the reproductive organs. The body-wall is represented as being cut open dorsally and spread out. 
br. the bridge or narrow part of the prostate connecting the wide and lower part with the storage chamber of 
the spermatozoa. str. storage chamber of spermatozoa, The prostate and ciliated rosettes are somewhat 
stretched, as otherwise they could not be seen both at the same time. 


79. Part of the clitellum in transverse section, showing the clitellar glands, the longitudinal and transverse muscu- 
lar layers. 


80A. One of the blood glands or herzkérper. 
80B. A single gland cell of the former. 
81. A transverse section of the dorsal vessel, showing the blood gland occupying the ventral part of the vessel. 


82A. Cross-section of the lower part of the spermatheca, at the spermathecal pore. bd. body-wall. ep. inner lining 
epithelium. m. muscular transyerse layer. ex. exterior layer. /. m. longitudinal muscular layer of the 
body-wall. ¢. m. transverse muscular layer of the body-wall. 


82B. Cross-section of the distal end of the spermatheca. 


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116 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLII. 
ECLIPIDRILUS FRIGIDUS. 


83-91. Represent somewhat diagrammatic views of transverse section of the body, respectively through somites iii, 


$4. 


85. 
86. 


89. 


90. 


91. 


93. 


94. 


iv, ix, x, x, xi, xi, xiv andxy. Of somites x and xi sections from the anterior and posterior parts are shown. 
The following letters are common and indicate the same in each one of the figures: v. v. ventral vessel. 
d.v. dorsal vessel. int. intestine. J. m. longitudinal muscular layer. 1. m. transverse muscular layer. 
v.c. ventral nerye cord. m. muscles and connective tissue uniting the intestine and body-wall.  sp.s. 
sperm-sacs. c. v. connecting yessels, cut in various places. pha. pharynx, shown to be developed only on 
the upper side of the alimentary canal. 


The intestine is here seen only to be connected with two superior muscular fascicles with the body-wall. The 
connecting vessels between the ventral and dorsal vessels are covered with glandular cells, the contents of 
which stains deeply with hematoxylon, and are of a different nature from the chloragogic cells. 


Section through the first clitellar somite ix. cl. clitellar layer. spth. spermatheca. spz. spermatoza. 


Section through somite x. Inthis specimen both the spermatheca project into somite x. The blood vessels 
which are seen in the sperm-sacs and surrounding the spermatheca are the anterior part of the anterior pair 
of lateral vessels, which project no further than here. 


Section through the posterior part of x cutting through the atrium and penis. The ciliated rosettes were not 
seen in this section, only some parts of the spermduct. The blood vessels seen surrounding the penis and 
atrium are the anterior parts of the posterior pair of lateral longitudinal vessels. 


Section through somite xi, close behind the anterior septum, showing the middle part of the outer ciliated 
rosette cut through. The other rosettes having been torn were not seen on the section. The spermducts 
are seen as a small circle, one on either side of the prostate. 


Section through the posterior part of somite xi. The outer spermducts are seen as situated further down, 
approaching each other, the approach being yet closer in the next section. 


Section through somite xiv. The prostate being bent zigzag is seen as cut in three different places. At athe 
part anterior to the very narrow pitt or bridge is seen cut. At} this very narrow part and at ¢ the posterior 
part or storage chamber where the spermducts are seen entering the chamber. 


Section through somite xy. The storage chamber is cut close through its anterior end and is seen surrounded 
by a thick layer of prostate glands. 


Cross-section of the atrium below its junction with the prostate. gl. small prostate glands penetrating the outer 
muscular layer. m.s. muscular layer. tube tube winding in the inner layer of fibrous tissue. n. nuclei 
of lining layer, boundaries of the cells not distinct probably on account of maceration. A few nuclei only are 
seen in the fibrous mass, which is much thicker in the center surrounding the tube. 


Cross-section of the prostate proper in somite xii. g/l. s. smali surface glands penetrating between the longi- 
tudinal muscular layer. m.s. longitudinal muscular layer composed of fascicles or plates of strands. ¢. m. 
transverse muscular layer. gl. c. thick glandular layer of one row of cells. J. ep. lining ep. of cells. spz. 
spermatozoa. 


Cross-section of the narrow part or bridge of the prostate. The layers are the same as in the preceding figure, 
but of different relative thicknesses. Letters as in preceding. sp. sperm ducts. out. m. outer layer of 
muscles, running in different directions. 


Cross-section of the storage chamber in somite xy, posterior to the entrance of the spermducts, but very close 
to it. sp.d. spermducts. gl. s./. large surface or prostate glands. gl. s.s. small surface or prostate glands. 
m.s. longitudinal layer of muscles. ¢. m. transversemuscles. ep. lining epithelium with large nuclei. 


A somewhat oblique section of the posterior part of the bridge, near its junction with the storage chamber, 
but anterior to the entrance of the spermducts, showing the relative size of the layers and the increased 
epithelium in the anterior part. br. posterior part of bridge, near its widening. ep. lining epithelium. pr. 
gl. prostate glands. Jl. m., t. m. longitudinal and transverse muscular layers, both cut obliquely. Both the 
transverse muscles and the epithelium are thickened towards one end. 


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| PACIFIC COAST OLIGOCHETA. ii 


PLATE XLIII 


97A. 
975. 
97C. 
99A. 


99B. 
100. 


101. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLIII. 

ECLIPIDRILUS FRIGIDUS. 
Cross-section of the neck of a nephridium. 
Section of the long duct. 
Nephrostome and part of duct. 
Section of the lower part of the male organ, slightly oblique to the longitudinal axis of the penis, but parallel 
to the atrium. aftr. atrium. /f.¢. fibrous tissue. im. inner lining of the atrium with long oval nuclei. 
1. ms. longitudinal muscular layer of atrium. ¢. ms. transverse muscular layer. gl. small glands scattered 
over the surface of atrium and prostate. spd. spermduct. sep. part of the septum between x/xi. spAzt. cir- 
cular muscles or sphinxter. g/. c. large glandular cells composing the collar. m. p. muscular plates, to which 
the longitudinal muscles of the atrium are joined. ube. penis tube. ps. penis sheath. pre. preputium. 
bd. body-wall. 


Section of inner penis tube of the former figure. 


The lower end of the male organ, atrium and prostate in somite x, xi, somewhat diagrammatically figured, in 
order to show the different layers, penis, ete. pr. prostate proper, projecting somewhat into x. s. septum. 
t. pr. terminus of prostate. pr. m. muscular layer surrounding the prostate. This layer contains numer- 
ous glands. pr. gl. glandular layer of long flask-like cells. ep. inner lining epithelium. tube tube running 
from prostate through penis and atrium—a penis lumen, with thick walls connected with the atrial wall by 
numerous strands, which on the upper part form a thick layer surrounding the tube, but lower down becomes 
thin, at least consisting of only few strands. col. collar surrounding posterior part of penis. jp. penis, 
projective part. pre. preputium, eversible. bd. body-wall. /f. t. fibrous tissue. 


A somewhat larger view of the lower end of the penis. Both this and the former drawing are made from 
dissections mounted in gum-thus and yiewed from above. Letters indicate the same as in fig. 92. 


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102. 


103. 


104. 
105. 


106. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLIV. 
ECLIPIDRILUS FRIGIDUS. 


The central part of the dissected prostate and male organ, as seen when mounted in gum-thus. This figure 
represents the narrow part of the prostate or tube, connecting the lower or glandular part with the upper or 
storage part. The two spermducts are marked much more strongly than they appear and are more spread 
or separated. The drawing represents views at two different foci. Zeiss C. Eyep. 3, ¢. 165. a. the posterior 
part or storage chamber in which the spermducts open at j. sp. The anterior end of this chamber is lined 
by a very thick epithelium, which however rapidly diminishes in thickness backwards. b. the very narrow 
part connecting the posterior storage chamber with the anterior prostate. c. the anterior prostate. out. 
outer layers of the storage chamber and prostate, surface views. du. lumen of storage chamber. /um. lumen 
of prostate. ep. very thick epithelium at the anterior end of the storage chamber. j. sp. junction of sperm- 
ducts and storage chamber. /.v. lateral longitudinal blood vessels. se. septum between xiv/xy. spt. sep- 
tum between xv/xvi. spd. spermducts, br. narrow part of prostate or bridge. gl. glandular layer of 
prostate. ept. lining-epithelium of prostate, 


The anterior end of the storage chamber, dissected, mounted in gum-thus. The focus is set on the inner 
concave surface, showing the junction of spermducts and bridge with the storage chamber proper. sp. the 
pores where enter the two spermducts. outer. coats of muscles and small prostate glands of the storage 
chamber. br. bridge or narrow part of the prostate. 


Longitudinal section of the body-wall, showing posterior ciliated rosette, etc., behind the male pore. 


The inner end of the former more highly magnified. The letters indicate the same as in the last figure. 
Only some of the larger prostate glands are figured. Zeiss C. Eyep. 3. 


A part of the former more highly magnified. Zeiss Hom. Im. 1-12, Eyep. 3-155. The letters indicate the same 
as in fig. 104. This section is more parallel with the longitudinal axis of ihe organ. The cell walls of the 
epithelial cells were not distinct, probably on account of maceration. 


PACIFIC COAST OLIGOCHETA. 121 
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122 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLV. 
ECLIPIDRILUS FRIGIDUS. 


107. A transverse section, the storage chamber more highly magnified. Zeiss Hom. Im. 1-12 Eyp. 3. 155. pr. gl. 1. 
large prostate glands. pr. gl. s. small prostate glands. J. m. longitudinal muscles, arranged in strands. 
t. m. transverse muscles. x. nuclei of inner lining epithelium. 


108A. A dissected ovary. 
108B. A ripe ovum. 
109A. A dissected oviduct. . 


109B. Longitudinal section of the body-wall, at the ovipore, showing the latter to be situated in the intersegmental 
groove. 


110 to 124. Various and successive stages of development of spermatozoa. 
110. A resting spermatogonium (spermatospore) from the testes. 


111. A spermatogonium from the outer edge of the testes ready to fall offinto the sperm-sac. The chromosomes of 
the nucleus have begun to develop into winding rods. 


112. Nucleus from a spermatogonium from the sperm-sac. 


113. Spermatogemme (spermatosphere or spermpolyblast). The spermatocytes (spermatoblasts) are surrounding 
and attached to a central, non-nucleated cytophore (or spermblastophore). 


114. The same shortly before division of the spermatocytes. 


117. A spermatogemme in the next last stage of development. The spermatocytes haye through division reached 
their final nomber. The cell divisions are indistinct. The nuclei are globular with scattered chromosomes. 


118. A further developed or next last stage of thespermatogemme. The nuclei have assumed an ovoid shape, stain- 
ing very dark with saffranin. 


119. A part of a spermatogemme, last stage. The nuclei have again diminished in size and become perfectly round, 
previous to growing out into spermatozoa. 


120. Part of spermatogemme in which the nuclei of 119 have begun to grow out into spermatozoa. 

121. A fully developed spermatogemme with grown spermatozoa attached to the cytophore. 

22. Thesame in a state of dissolution, the spermatozoa detaching themselves. 

123. Spermatozoa fully developed, the nuclear end being only slightly thicker than the other part of the body. 


124-125. Sections of fully developed spermatogemmes, showing spermatozoa in cross-section, they having gradually 


diminished in diameter since the nuclei were first concentrated (fig. 119), but having proportionately grown 
in length. 


126. Nuclei from 113, 117, 118, show their relative size. 
127-129. Yolk sae with yolk cells. 

127. The whole sac in section. Zeiss C. 

128. Yolk cells without granulation. 

129. Further developed yolk cells with granulation. 


KERRIA McDONALDI. 


130A. Longitudinal section of the alimentary canal of Kerria Mc Donaldi in the somite next posterior to the male 
aperture, showing unicellular glands alternating with ciliated epithelial cells. gl. unicellular glands. 


130B. A more highly magnified part of the former, showing one of the unicellular glands, surrounded by common 
epithelial cells, also rudimentary glands. r. gl. rudimentary glands. gl. large gland. p. its poe. ep. 
epithelial cells. /. v. blood-lacune. /. m. longitudinal muscles. ¢. m. transverse muscles. 


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VoL. ll, No. 5. 


PACIFIC. COAST: OLIGOCH ATA. a 


aad | II. fs 


By GUSTAV EISEN. 


eit ed SAN RRANCISCO, CAL, 


FEBRUARY, 1896. ; ie 


PACIFIC COAST OLIGOCH ATA. 
ER 


BrEnHAMIA, ACANTHODRILUS, ALEODRILUS, SPARGANOPHILUS, DELTANTA, 


PH@NICODRILUS. 
BY GUSTAV EISEN, PH. D. 


The following genera and species are treated of in this paper: 


Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 
Benhamia 


nana n. sp. 
mexicana Rosa. 
Bolavi Michaelsen. 
palmicola n. subsp. 
papillata n. sp. 
rugosa n. sp. 
octonephra Rosa. 
Godeffroyi Michaelsen. 
malayana Horst. 
floresiana Horst. 
Annz Horst. 


Acanthodrilus tamajusi n. sp. 
Acanthodrilus Vasliti n. sp. 
Aleodrilus Keyesi n. sp. 
Sparganophilus 
Sparganophilus 
Sparganophilus 
Sparganophilus 
Sparganophilus 
Sparganophilus 
Sparganophilus 
Deltania Troyeri var. crassa n. var. 


Benhamii n. sp. 

Smithi n. sp. 

sonome n. subsp. 
Eiseni Smith. 
guatemalensis n. subsp. 
carneus n. sp. 

tamesis Benham. 


Deltania Troyeri var. lagunz n. var. 
Pheenicodrilus taste Eisen, 


Pheenicodrilus tepicensis n. sp. 


Memoirs, Vou. II, 5. December 14, 1895. 


124 CALIFORNIA ACADEMY OF SCIENCES. 


BENHAMIA Michaelsen. 


The genus Benhamia was for some time considered a typical African genus, 
and when later on a few extra-African species were found it was supposed that these 
were recent emigrants, which had become more or less cosmopolitan. Of American 
species two are known from Mexico and Venezuela, viz.: 6. Bolavi and b. mex- 
icana, and one, Bb. octonephra Rosa, has been described from Paraguay since this 
paper was presented for publication. This latter species appears very nearly related 
to B. rugosa, described below, but differs through the absence of penial sete. 

The discovery of species of Benhamia at Miraflores, in the Cape Region of 
Baja California, in a locality to which plants of any kind have rarely if ever been 
introduced directly from foreign countries, would indicate that this genus has possessed 
representatives on American soil for ages past, and that we really must consider these 
American species as truly endemic. A final answer to this question of habitat must 
be deferred to a future time, when more researches will have been made as regards 
the distribution of these and other Benhamia species, as it is probable many more will 
be found on this continent. A difficulty, which besets us from the beginning, is, that 
so few species have been properly delineated, all descriptions having been made chiefly 
with a view to distinguish the species from others already known, while with proper 
delineations of the various organs, we would in all probability be able to make a satis- 
factory comparison between species new and old. I have received much aid from 
Dr. W. Michaelsen, of Hamburg, who has described more Benhamia species than any 
other investigator, and who has written extensively upon this genus. He has kindly 
placed at my disposal several species of African Benhamia, as well as of B. Bolavi, for 
comparison with forms found by me. This has enabled me to point out several im- 
portant differences between LG. Bolavi and B. palmicola, which are sufficient to dis- 
tinguish these as subspecies from each other, as well as from others previously known. 
At the end of the descriptions of the various new species I append a table of charac- 
teristics, etc., between species which may be confounded with our present ones, either 
on account of similarity of some characters or because of their geographical distribu- 
tion in the Malay archipelago, or in America. 

As we now understand the distribution of this genus, the species are divided as 
follows: America, 7 species; Malay archipelago, 3 species; Africa, 25 species; West 
Indies, 2 species. 

It may, however, be remarked that B. rugosa described below is of uncertain 
habitat having only been found in a hot-house, to which it had been imported from 
unknown country. 

Considering our various new species of Benhamia this genus may be charac- 
terized as below: 

Benhamia Michaelsen. 

Acanthodrilid oligocheta. Sete strictly paired, ventral and lateral. Cli- 
tellum generally incomplete, but in some species complete in some somites. Two 
gizzards in succeeding somites. Calciferous glands generally three pairs, but some- 
times only two pairs, very distinctly set off from the tubular intestine. Nephridia in 


PACIFIC COAST OLIGOCHAETA. 125 


respective species either diffuse or micronephridia arranged in three, four or more 
rows on either side, some species showing several gradations. Spermatheca two pairs, 
contracted at center with one or more small diverticula, Penial sete nearly always 
present. Generally small, tropical worms. 


DEFINITIONS OF AMERICAN SPECIES OF BENHAMIA. 


Benhamia nana n. sp. 


Derinition. Size—Length 20 to 30 mm.; number of somites 110; skin strongly 
pigmented. First dorsal pore between iii/iv. Prostomium very swollen and overlapping. 
Clitellum complete in central somites. Penial sete, the largest with 12 or more bristles. 
Common sete, distance between the dorsal couples about equal to that between the ventral 
couples. Oviducts, two pores, open in front of 1 and 2. Prostate pores not on papille. 
Pharynux situated very far forward. Gizzards in viii and ix. Calciferous diverticula, 
two pairs in xv and «xvi. Sperm-sacs in x and ai. Spermducts thicker at the base in 
xviii, evit and xvi. Nephridia in three rows on either side, the posterior with calomic 
mantle. Spermatheca, basal part with one diverticulum, apical part warty and irreg- 
ular.  Typhlosole large, begins in xviii. Color brownish red. Habitat, San Blas, 
Mexico, at sea level. 

Benhamia mexicana Rosa. 

Derinition. Length 30 mm.; number of somites 120. Skin not pigmented. 
First dorsal pore between wi/iv. Prostomium divides the first somite completely. Cli- 
tellum complete in aiti to wai. Penial sete not ornamented (?); common sete, 2 and 4 
further apart than Land 2. Oviducts, two separate pores in front of 1and 2, on small 
papille. Gizzards in viii and iv. Caleiferous gland 3 pairs in xv, xvi and xvii. Ne- 
phridia in 3 rows on either side. Spermatheca with a single short diverticulum. Alco- 
holic specimen colorless. Habitat, Durango, Mexico, at 2500 meters altitude. 


Benhamia Bolavi Michaelsen. 


Derinition. Length 40 to GO mm.; number of somites 97. First dorsal pore 
between v/vi. Clitellum incomplete in all somites, viii to wx. Penial sete, the largest 
with five to eight notches, the smaller spoonlike and slightly forked; common sete couples 
equidistant. Oviducts open in one single pore on a median papilla in xiv. Gizzard two 
in vii. Caleiferous gland three pairs in xv, xvi and xvii. Sperm-sacs one pair in xi. 
Nephridia in three rows on either side. Spermatheca, the basal part with a single small 
diverticulum. Saculated intestine begins in wxi. Color dingy flesh. Habitat, Venezuela 
(and Hamburg). 

Benhamia palmicola n. subsp. 

Derinition. Length 50 to GO mm.; number of somites 90. Skin not pigmented. 
First dorsal pore between iv/v. Clitellum incomplete in all somites. Penial sete, largest 
with four notches, the smaller one spoonlike and forked; common sete about equal dis- 
tance between the dorsal and ventral as between the ventral couples. Oviducts open in one 
single pore on a median papilla. Gizzards both in viii. Calciferous gland 3 pairs in 


126 CALIFORNIA ACADEMY OF SCIENCES. 


av, aviand avii. Sperm-sacs in xi and xii. Spermducts of even width. Nephidia 
in 3 distinct rows on either side. Spermatheca basal division globular, equal in size and 
shape to the apical part, single small diverticulum. A typhlosole. Color reddish flesh. 
Habitat, Baja California, Cape Region, 1000 feet; Tepic, Mexico, 4000 feet. 


Benhamia papillata n. sp. 


Derrinition. Size 50 to 70 mm. by 2 mm. Number of somites 125. First 
dorsal pore v/vi. Prostomium divides somite I more than one-half, and is much swollen. 
Clitellum complete, xiii-vx. No ventral sete in xviii, but sete 3 and 4 present. Prostate 
pores on large papilla, the four papille being close together in the sunk clitellar pit. 
Oviducal pores on a large median papilla, but the pores are separate and on a line drawn 
between the sete. Penial sete, largest slightly hooked with 8 indistinct notches, smallest 
with a hair-like sigmoid tip. Spermathece, the apical-sac not globular, basal part 
“with a swell diverticulum. Sacculated intestine commences in xix. Typhlosole in xx—win. 
Calciferous diverticula 3 pairs in xn, xvii, xvii; the anterior one is much smaller. 
Sperm-sacs inx and wi. Nepridia a consists of three distinct lobes, each one with a 
caelomie mantle. Color pale flesh, no pigment. Habitat, Tepic, Mexico, 4000 feet. 


Benhamia octonephra Rosa. 


Derinition. Compiled after Rosa. Length 20 to 40 mm.; number of somites 
85 to 95. First dorsal pore between » and vi. Clitellum incomplete, in xiii to va. Pe- 
nial seti, the larger with six groups of two blunt tubercles; the smaller seta like a scalpel. 
Common sete, couples equidistant. Oviducts open into a single median pore on a slight 
swelling. Calciferous diverticula three pairs in xv, xviand xvii. Sperm-sacs in xi and 
wii. Nephridia micronephric in four rows on either side. Spermatheca, basal part 
pear-haped with a globular diverticulum. Habitat, Paraguay. 


Benhamia rugosa pn. sp. 


Derinition. Length 30 mm.; number of somites 118. Skin slightly pigmented 
and corrugated. Clitellum incomplete, in wiii to wu. Penial sete, largest with five 
notches; smaller seta forked with prongs of equal size. Common sete pointed and ven- 
tral. Oviducts open jointly in a median pore. Gizzard in vii and vin. Calerferous di- 
verticula in wv, avi and «vii. Nephridia micronephric in four rows on either side. 
Typhlosole present. Spermatheca much flattened. Color reddish flesh. Habitat, Cal- 
ifornia, in hothouse (imported). 


Benhamia Godeffroyi Michaelsen. 


Derimition. Length 90 mm. by 4 mm.; number of somites 174. Skin pig- 
mented anteriorly. Clitellum incomplete (xiii) wiv to wx. Penial sete slender with nu- 
merous irregular notches; many sete in each sac. The curved fossa between the pros- 
tate pores with the convex side toward the median line. The anterior prostate largest. 
Calciferous diverticula three pairs. Nephridia plectonephric. Spermatheca without diver- 
ticulum, but with warty protuberance. Color, anterior pale reddish, posterior part gray. 


Habitat, Hayti (?/). 


a yreereseeneres 


see eee 


: 
nana D. sp. 


pic, Mexico, at sea level. 
Strictly tropical form. Temperate form. 


20 to 30 mm. 30 mm. 
110. Skin strongly pig- 
mented in anterior so- 
mites. 


lil-iv. iii-iv. 


Very swollen and overlap- 
ping. 


Complete in the central | Complete, xiii-xxi. 
somites, xiii-xx. 


completely (?) 


The largest with 12 or 
more strong bristles. 


enings. 


Distance between the dor- 
sal couples about equal 
to that between the ven- 
tral couples. 


than 1 and 2. 


Two pores, open in front 
of sete | and 2. 


their bases. 


Not on papille. 


| 
Not on papille. 


Situated very far forward. 


In viii and ix. In viii and ix. 


Two pairs in xy and xvi. 

xvii. 

In x and xi. In x and xi. 

Thicker in xviii, xvii and | Not enlarged (?) 
xvi. 

In three rows on each side 
of the median line. The 
posterior ones with ccelo- 
mie cell- mantle. The 
nephridia in each somite 
of about the same size. 
The nephridia on each 
side about equidistant, 
but the dorsal nephridia 
are much further apart 
from the dorsal median | 
line than from the lateral 
nephridia. Nephridia a 
open in front of 3 and 4. 


same size. 


Basal part with one plain 
diverticulum. Apical 
part warty and irregular. 

diverticulum. 


Very large; begins in xviii. 


Reddish-brown, even in | Alcoholic specimens color- 
less. / 


alcohol. 


mexicana Rosa. 


San Blas, Territory of Te- Durango, Mexico, 2565 | Hamburg, at Bergedorf, 
metres above the sea level. 


120. Skin not pigmented. 


Divides the buccal somite 


Not ornamented (Rosa). 
According to Ude, faintly | 
ornamented with 6 thick- 


Sete 3 and 4 further apart 


Two pores in front of 
sete 1 and 2 on small 
papille, which touch with 


Three pairs in xy, xviand 


In three rows on either 
side, probably all of the 
Nephridia a | 
correspond to sets 1 and 
2. The nephridia on each 
side are equidistant, and 
the dorsal nephridia ¢ are 
as equidistant from the 
dorsal median line as — 
from the lateral nephridia 
b. With ecelomic mantle. 


The basal part wide, grad- 
ually changing into the 
apical part. Single short 


¥e fj 
a) 


TABLE 
Bolavi Michaelsen. 


imported, and Venezuela 


(Ude). 


40 to 60 mm. 50 mm. 


07. 90. Not pigmented. 


v-vi. iv-v. 


Does not divide buccal 
segment. 


Incomplete in all seg- 
ments, xiii-xx. 


| 
The larger seta with five 
(or eight B) notches on 
apex; the smaller is spoon- 
like and slightly forked, — 
with no denticulation. 


Those on posterior seg- 
ments larger. The couples 
equidistant. 

tral couples. 


One single pore on a large 


median papilla in xiv. media papilla. 


Never elevated. 


Not forward. Not forward. 


Two in somite vii. 


Three pairs in xv, xviand 
xvii. xvil. 


One pair only in xi. In xi and xii. 


mites. 


In three rows on either 
side; the anterior ones 
have no ccelomic mantle 
like those posterior to 
the male pore. The two 
lateral nephridia of equal 
size; the ventral nephrid- | 
ium is much narrower, | 
but furnished with two | 
separate celomic man- | 
tles, the inner of which is 
smallest. The nephridia 
are more separated than 
in B. palmicola. 


median line. 


of 1 and 2, 


Basal part with a single 
small diverticulum. 
Apical division not glob- 


ular. lum. 


not very large. 


Dingy flesh-colored alive; 
clitellum yellowish or pink. 


palmicola n. subsp. 


Baja eels, Cape Re- 
gion an epic, Mexico, | 
1000 and 4000 feet. 


Not very pointed. 


Incomplete in all somites, 
xiii-xx. 


The larger seta with four 
notches, the smaller one 
spoon-like at apex and 
forked. 


About equal distance be- 
tween the dorsal and yen- 
tral as between the ven- 


One pore on a central 


Not on papille. 


Two; both in viii. 


Three pairs in xy, xvi and 


Stones ce minG eel haeips | Not thicker in 


Arranged in three distinct 
rows on either side of the 


ium a consists of two 
unequal parts, the most 
ventral of which is the 
smallest. Nephridium a 
is much larger than the 
other nephridia. 
dorsal nephridia far apart. 
| Nephridiaa open in front 


Basal and apical divisions 
globular, equal in size. 
Single small diverticu- 


Typhlosole present, but 


Reddish flesh when alive. 


ap 


60 to 70 mm. 


> 
125 (largest specimen). 


y-yi. 


Divides partly buccal 
segment; swollen. 


Complete, but ventrally 
thin, xili-xx. 


Larger with eight faint 
notches; smaller with 
a hair-like sigmoid 
point. 


Inner couples wanting 
in xviii, equidistant. 


Two closely approxi- 
mated pores on a me- 
dian papilla. 


Each on a large papilla. 


Not forward. 

Two. 

Three pairs in xy, xvi 
and xvii. 


In x and xi. 


In three rows, the pos- 
terior ones with ccelo- 
micmantle. Nephrid- 
ium a consists of three 
lobes, each with a sep- 
arate coelomic mantle. 
The inner lobe is much 
the smallest. The ne- 
phridia are closer, over- 


lapping. 


Apical division not glob- 


ular; dorsal part with 
a single diverticulum. 


Present in xix-xxiy. 


Pale reddish; no pigment. 


- . ; ad 


— pee 


octonephra Rosa. 


| Tepic, Mexico, 4000 feet. | Paraguay. 


20 to 40 mm, 
85 to 95. 


xiii to xx, incomplete be- 
tween the ventral seta. 


The larger curved, strong- 
ly curved at apex, and 
with 6 groups of 2 blunt 
tubercles, the smaller 
expanded at apex like 


a scalpel. 


Strictly paired, all ven- 
tral; couples  equi- 
distant. 


A single pore on median 
line in xiy, on a slight 
swelling. 


In line with ventral sete. 


Three pairs in xv, xvi 
and xvii. 


In xi and xii, two pairs. 


Arranged in four series 
on either side of the 
median line; the ven- 
{ral one corresponds to 
the dorsal sete. 


Basal part pear-shaped; 
apical part sac- like, 
basal part with a glob- 
ular diverticulum di- 
rected forward. 


¥ 


en nal 


malayana Horst. 


Malay Archipelago. 


20 to 30 mm. 


95. Skin without pig- 


mentation. 
v-vi or vi-vii. » 


Continued into the bue- 
cal segment, 


Complete in the anterior, 


incomplete in the pos- 
terior somites, xiii to xx. 


The larger seta with 4 
broad bristles; the 
smaller spoon - like, 
but not forked. 


Distance between dorsal 
and ventral couples 
about equal to that be- 
tween ventral couples. 


Two distinct pores. 


The depression connect- 
ing the prostate pores 
concave, with the con- 
vex part turned toward 
the median line. An 
elevated area around 
the depression. 


. =3: | . . 
The posterior nepbridia | Arranged in four longi- 
tudinal series on each 
median 


arranged in 3 groups 
of delicate tubuli on 
either side, the inter- 
nal ones situated near 
the dorsal sete. 


Basal part longest and Basal part the longest, 
single pear- 
shaped diverticulum, 


as broad as the apical 
part, with a narrow di- 
verticulum. 


 walebe a date St vs 
TRA-AFRICAN SPECIES OF THE GENUS BENHAMI 


floresiana Horst. 
Malay Archipelago. 
35 to 40 mm, 
mentation. 
vi-vii. 


one-half. 


Incomplete, xiii to xxi. 


dulated, the 


bristles at 
largest seta. 


ventral couples. 


sets 1 and 2. 


square area, 


| 
| side of the 
| line. 


narrow; 


30 mm, 


125. Skin without pig- | 85. Skin without pig- 


iy-v. 


Divides somite I about | Divides partly the bue- 


Anterior part much un- 
smaller 
seta less so; three small 
apex of 


Distance between a dor- 
sal and ventral couple 
about three-fourths of 
that between the two 


Open in an oval gland- 
ular area, pores open 
separately on line with 


Not elevated, situated 
at the corners of a 


A. 


Anne Horst. 


Java. 


ment. 


cal segment. | 


Incomplete in xiii to xxi. 


Undulated, with 12 
(about?) slight dentic- 
ulations or notches. 
The form of smaller 
seta doubtful. 


The four couples are 


| 


rugosa n. sp. Godeffroyi Michaelsen, 
San Francisco, Califor- Hayti (or New Zealand ” 
nia, imported. : 
30 mm. 90 mm. by 4 mm. 


118. Skin slightly pig- | 174. 
mented and anteriorly 
corrugated. 


Anterior part of 
body pigmented. 


v=vigee > he yo ee 
Divides buccal somite Divides buccal segment 
about one-half. | Somewhat less than }. 


Incomplete in xiii to xx. Incomplete (xiii), xiy-xix 


Largest seta with five Slender, numerous irreg- 
notches; smaller seta ular notches, giving 
forked, with prongs of — knotty appearance, but 

| equal size. ' no teeth or bristles. 

| | Many sete in each sac. 


| Strictly ventral. In 4 couples, all ventral. 


ventral and equidis- 
tant. 


Unknown. 


Depressions connecting | 


pores lunate, with the 
convex side turned to- 
wards median line, 
Area oval, not square; 
a ridge borders the 
grooves. 


I royce SA a | Not forward. 


Two in vi and vii. 


XV, xvi and xvii. 


| In ix-xii. 


A network of delicate | 


tubules anteriorly 
spread over the body 
area, except for a nar- 
row dorsal and ventral 
median line. Posteri- 
orly arranged in four 
rows on either side of 
median line, and fur- 


nished with ccelomic 


mantle. 


Basal part longest and 
very narrow, with 
narrow diverticulum 
situated near the cou 
traction. 


Alcoholic specimens col- | Alcoholicspecimens dis- | Colorless. 


orless. 


| colored. 


Open jointly ina median 
| pore in xiv. 
| | 
Anterior prostate pores | [he curved fosse between 
elevated on papilla. | the pores, with the con- 
| vex side turned towards 
) the ventral median line. 
The anterior prostate 
the largest. 


| 
| In vii and viii. 
} 
| Three pairs in xy, xvi 
| 


| Three pairs. 
| and xvii. 


Arranged in four rows | Plectorephrie condition, 
on either side of me- | but not known in detail, 
dian line. Posterior | except that they cover 
nephridia with cclo- | the whole of the body- 
mic mantles. Neph- wall except a narrow 
ridia of nearly thesame | strip dorsally and yen- 
size. Neph. d and d trally. Stronger in the 
about equidistant as | clitellar somite, where 
a and @ The most | they form a thick cov- 
ventral part of nephrid- | ering. 

ium @ not covered by 
mantle, Clitellar ne- 
phridia much larger 
than those anterior of 
clitellum, 


Spermathece with no 
distinct diverticula, but 
tened. Very small di- | the smaller basal part 
verticulum. with irregular warty 
| protuberances. Anterior 
spermathecs the larg- 
est. The apical part sac- 
like, larger and thin- 
walled. The basal part 
shorter. 


Apical part smallest. 
Both parts much flat- 


yg ER ermine « 


Reddish flesh. 


| Anterior part reddish; 
posterior part gray. 


bo 
-~J 


PACIFIC COAST OLIGOCHAHTA. 1 


DETAILED DESCRIPTION OF NEW SPECIES. 


Benhamia nana np. sp. 
Figs. 1-42. 


Habitat. San Blas, territory of Tepic, Mexico. I found a small number of 
specimens, of which only two possessed a perfectly developed clitellum, October, 1894. 
_ A real tropical form. 

Color. Deep reddish-brown and very opaque. 


EXTERIOR CHARACTERS. 


Size small, 20 to 80 mm. 

Somites 110. 

Clitellum comprises somites xiii to xx, being complete in the central somites, 
but incomplete in xiii, xvii, xvili and xix. 

First dorsal pore iii-iv. 

Spermathecal pores, one pair vii-viii, and one pair between viii-ix, rather close 
together in a ventral grove. 

Oviducal pores open separately in xiy in front of sete 1 and 2. 

Prostate pores, one pair in xvii, one pair in xix, open with penial sete. 

Penial set, two in each pocket, the largest of which is furnished with 12 or 
more bristles. 

Common set, distance between the dorsal couples equal to that of the ventral 
couples. 

INTERNAL CHARACTERS. 


Buccal cavity with a dorsal pocket. 

Suprapharyngeal glands with pharynx opening far forwards. 

Gizzards in vill and ix. 

Oalciferous glands two pairs, one in Xiy, one in Xv. 

Septal glands very minute in ix, x, Xi. 

Sacculated intestine commences in Xiv. 

Typhlosole very strongly developed in xviii to xxii. 

Sperm-sacs two pairs in xX, X1. 

Ciliated rosettes in x, xi. The lower part of the spermducts in xvi, xvii, xviii 
is thickened and twice the diameter of the anterior part. 

Prostates are straight and tubular, each one confined to one or two somites. 

Spermathece in vii-viii, vili-ix. The apical part much smaller than the 
basal part, the latter with a small diverticulum placed high up near the constriction. 

Nephridia arranged in three rows on either side of the median line. Pair a 
open in front of the outer couple of sete, pair 6 and ¢ open laterally and dorsally as 
regards the sete. The posterior nephridia furnished with a large coelomic cellular 
mantle. The nephridia in each somite of about equal size. The nephridia on each 
side about equidistant, but the dorsal nephridia ¢ are much further apart from the 
median line than from nephridia 0. 


128 CALIFORNIA ACADEMY OF SCIENCES. 


DETAILED DESCRIPTION. 


Body-wall. The prostomium with buccal cavity is strongly eversible, forming 
a bladder-like apex to the body, as is so frequent in oligocheeta. Somite i is very 
narrow, especially laterally, and may readily, when viewed from the exterior, be taken 
for part of the prostomium. 

The body-wall contains the usual layers of which the muscular fibers show the 
same bipinnate arrangement as in Lumbricus, ete. This arrangement is less regular 
and pronounced in the anterior somites, but quite plain in the genital and clitellar 
ones, especially so on the ventral side, immediately below the nerve-cord. 

Sense organs of the epidermis. All the species of Benhamia described here 
possess a continuous row of sense organs, in the equatorial plane of each somite, be- 
tween the sets. Outside of this equatorial circle I have not found them anywhere 
in the epidermis; there, however, they are very plain and prominent, appearing under 
low power and in longitudinal sections, as a large pellucid spot in the center of each 
somite. The organ consists of two distinct kinds of cells, a double line of large lunate 
cells, surrounding a row of sense cells, several layers thick. These lunate cells are 
generally three or more thick in the row, evidently modifications of the common 
goblet cells of the epidermis. They do not stain with the ordinary aniline colors, or 
only so with difficulty, and generally remain transparent and white. These lunate 
cells run continuously around the somite, and enclose between them bunches of sense 
cells, which may now and then be seen to penetrate the cuticle (fig. 20). Somewhat 
similar sense organs have been known in Lumbricus, ete., for considerable time, but 
have lately been described more in detail by Richard Hesse and F. E. Langdon. 
The sense organs of Benhamia differ from those of Lumbricus agricola (probably a 
collective name used for some East American Allolobophora) in two prominent 
points. Presence of the large lunate cells in Benhamia, which are not seen in 
Lumbricus. The continuous and broad circle of these organs in Benhamia, while in 
Lumbricus they appear to be much further apart. Unfortunately my Benhamias 
were collected at a time when no special preparation for nerve structure was feasible, 
and this made it impossible to work out the details as minutely as desirable. The 
work had already been finished when Langdon’s beautiful paper reached me. 
Fig. 20 represents a section of the body-wall in somite iv. Letters org. signify 
the sense organ. 

Besides these epidermal sense organs I find in all the Benhamias, observed by 
me, a large zone in the buccal cavity characterized by almost cubical transparent 
cells arranged in one single row deep, just as the epithelial cells in the pharynx, but of 
the same nature as the transparent cells in the sense organs of the epidermis. This zone 
is nearly always folded against itself like a sac, and is of considerable extent, as long 
or longer even than the pharynx. In certain places apparently scattered about, but 
principally near the opening of this sac, I find clusters of sense cells of the same 
structure as those of the epidermis. They are all narrow, do not reach below 
in the coelomic cavity, but end in this direction in line with the pellucid cells and 
connect at the end with nerve fibres. The free ends penetrate above in what ap- 


PACIFIC COAST OLIGOCH2/TA. 129 


pears as a veritable cuticle, though I can find none above the pellucid cells. In 
fact the whole structure of these organs is very much the same as the sense organs of 
the epidermis of the body-wall. In connection with them I may point out the simi- 
larity of structure of these organs with those described below in Acanthodrilus Vasliti, 
in the vicinity of the prostates; and with those in the tubercula pubertatis of Sparga- 
nophilus Smithi and tepicensis, in all of which I find the same sense cells. 

It remains to add that this pouch-like area of sense cells and glandular cells has 
previously been observed by both Michaelsen and Horst and designated as a diver- 
ticulum of the buccal cavity, but the true nature of the pouch as a sense-area has, I 
believe, not been previously recognized. Bb. Bolavi, malayana and probably most, if 
not all other, Benhamias possess this organ of the buecal cavity. 

Septa. These are generally very thin, only two being thicker than the rest, 
viz.: Xii—xili and xili-xiy. These septa do not strictly correspond with the inter- 
segmental grooves, but are affixed much further back in the somites. 

Pharyngeal and septal glands. The usual supra-pharyngeal glands are present. 
They are evidently unicellular and arranged ribbon-like as will be directly described 
below. The septal glands are found in ix, x and xi, are very narrow and only one cell 
thick in the row as the former. These glandular masses are much thicker in B. 
palmicola, but not any longer. 

Intestine. In all the specimens which I could examine I found the pharyngeal 
parts strongly everted and protruded to such an extent that it formed the lip or front 
margin of the body, taking the usual place of prostomium proper. I cannot ascribe 
this entirely to a simple protrusion of the pharynx, but believe that this part is actu- 
ally situated much more forward than in other species, as I found it to be the case in 
every specimen observed. ‘The pharynx was found actually on the outside of the body 
(figs. 15, 16, 17, 18). The real pharynx, of course, is the zone in which open the 
supra-pharyngeal glands. The wall of this part is in our present species hardly 
thicker than those of the buccal cavity and the cesophagus, but it contains the usual 
arrangement of narrow epithelial cells, between which penetrate the fine ducts from 
the supra-pharyngeal glands. The ends or discharge pockets of these glands are 
almost globular or rounded flask-like (fig. 18). The supra-pharyngeal glands are 
arranged as ribbons, running singly along muscular strands. They differ from similar 
glands in Pontodrilus, Phoenicodrilus, Oenerodrilus etc., by their arrangement in 
single rows, and here and there the duct of a single glandular cell may be followed 
clear to the discharge pocket (fig. 16). These glands appear to consist of a single 
cell with a long duct, just as the corresponding glands in Euchytreus, described by 
Hesse. But to draw the conclusion from this fact, that all the pharyngeal and septal 
glands are unicellular is, I think, premature. In Pontodrilus, at least, there 
may be seen plainly numerous nuclei on the gland ducts, which, of course, indi- 
cates that we here have a fusion of several cells. The pharyngeal glands in that 
genus do not show this ribbon-like arrangement as in Benhamia. I could, however, 
see that some of the smaller glands nearest the pharynx consisted of only one cell, 
but the majority, and all the large glands, consisted of several cells, the respective 


130 CALIFORNIA ACADEMY OF SCIENCES. 


ducts of which finally united into one. In Benhamia I could see no such union, and 
the single ducts could be followed with great facility to the outlets. Fig. 17 repre- 
sents some of these cells with three narrow ducts and secreted matter. 

(Esophagus, following pharyngeal division, is very long. The upper part, im- 
mediately below the pharyngeal gland, is very thin-walled, consisting of only one 
strand each of transverse and longitudinal muscles, and lined by a very narrow epi- 
thelium (fig. 19). Posteriorly the walls of the cesophagus thicken considerably. The 
two distinct gizzards are in viii and ix, as usually connected by a very thin wall of 
the same general nature as csophagus (fig. 7). The muscles of the gizzards are 
columnar but not bipinnately arranged, the ribbons running parallel with the short 
diameter of the body (figs. 7 and 21). 

The tubular intestine extends through somites x to xiv, being of irregular out- 
line. Sacculated intestine commences in xv, and is furnished with a typhlosole in 
somites xviii to xxiii, or thereabout. 

Epithelial cells of the alimentary canal surround glands of various forms. In 
the epithelial lining of the gizzard we find club-like glands (fig. 21 g/.) consisting 
each of one (seldom of more) large cell with round nucleus, a narrow duct reaching 
between the epithelial cells, and ending witha large chamber, in very much the same 
way as the pharyngeal and septal glands. 

In the narrow thin walled part between the two gizzards I find a few clusters 
of glands (?) similar to those I have described in Argilophilus. At the bottom of the 
cluster we find a glandular cell, upon which are butting the peculiar lunate cells, 
which again surround a lumen, which is much wider than in the corresponding organ 
in Argilophilus (fig. 22 g/. and c.) The lining of the sacculated intestine and the 
typhlosole are composed of three distinct kinds of cells, two of which are glandular 
(figs. 24 and 25.) The common epithelial cells offer nothing of particular interest. 
The glandular cells are of two distinct kinds (fig. 25). One kind is the one most 
common in oligocheeta (fig. 25 g/.) and its enclosed granules are much smaller. The 
other kind is probably identical with the T-shaped cells described by Benham in 
Eminiodrilus, though the T-shaped form is not quite so prominent. The granulation 
is coarse and highly refractive and the distal part of the cell stains intensely, especially 
with methyl green. The other or first mentioned glands remain at the same time un- 
affected by this stain. These dark staining cells are much less numerous than the 
other kind, and are scattered about in a rather regular way. Fig. 24 4 represents 
one of the lobes of the typhlosole, showing the absence of glands at the apex as well 
as the general distribution of the dark staining cells. 

Caleiferous diverticula. I found in the two specimens dissected and sectioned 
only two pairs of calciferous glands in somites xv and xvi, but I am unable to say if 
this will be found constant, as all other species of Benhamia possess three pairs 
of calciferous pockets, which Beddard claims are characteristic of this genus. 
Michaelsen’s observation that the two diverticula are of different nature is confirmed 
here, as in the posterior pair no lime crystals were found, either in this or in the other 
species described below. Also the histological structure of the posterior and anterior 


PACIFIC COAST OLIGOCH ETA. 1s3i 


diverticula is distinct. The nuclei of the anterior pair are round while those of the 
posterior pair are more oval, and the cells of the latter pair are not separated by dis- 
tinct walls. They contain also numerous vacuoles or bladder-like bodies, some of 
which deatch themselves from the main body of the organ. 

Figures 27 and 28 represent the anterior pair, 29 and 30 the posterior pair of 
calciferous diverticula seen respectively under high and low power. The posterior 
pairs only were found covered by chloragogen cells. These differences in structure I 
also found in the calciferous diverticula of the following species: Benhamia palmi- 
cola and rugosa, and they probably hold good with all the species of Benhamia. 

Typhlosole exists in somites xvii or xviii to xxii, and is greatly developed as 
regards length, width and depth (figs. 7 and 14). It is widest and deepest in somite 
xix, tapering or diminishing both anteriorly and posteriorly. In the following species 
the typhlosole is much smaller. 

Spermathece. 'The two pairs as usual in vii—yiii and viii-ix. The form of the 
spermatheca and its diverticula may be best understood from the figures 34 and 35. 
The pores come rather close together in a general groove below the ventral ganglion. 
There is a muscular and a glandular part, and the spermatozoa were fonnd as usual 
principally in the diverticula. 

Spermducts and rosettes. The only peculiarity of the spermduets is that they 
are thickened in somites xviii, xvii and xvi. In xv the duct narrows, and continues 
forward, with about half the thickness it possesses in the above posterior somites. The 
thickening is due to an increase of a circular muscular layer (figs. 7, 11, 12, 13, 36, 
37). The rosettes are in x, xi, as usual. 5 

Prostates. The usual two pairs are in xvii and xix. The lower part is long, 
slender and muscular, while the upper thicker part is glandular. This glandular part 
consists of only one layer of cells, covered by a thin epithelium. The glandular cells 
are of different length, as shown in fig. 32. The muscular part consists mainly of a 
thick layer of circular muscles (fig. 36), lined by a very thin layer of interior epi- 
thelial cells (fig. 38). 

Penial sete. A sac with two penial setze open jointly with each prostate. At 
least one of the sete is sculptured as shown in the figures 31 and 33, with a number 
of bristles, 12 or more. The smaller seta was broken in all the specimens, and I 
could not ascertain if it was sculptured or not. 

Nephridia. Benhamia nana belongs to a group in which the nephridia are 
arranged in three rows on either side of the median line. There is not a diffuse 
nephridic condition, but each nephridium is well developed, and upon the same 
principle as the mega-nephridia of Acanthodrilus. Nephridia a. 6. are ventral and 
in front of sete 3 and 4, while the other nephridium c., is lateral and partly dorsal. 
The posterior nephridial ducts, from xvii to xviii, are superposed each on a large oval 
sac of coelomic cells, while those in front of the male-pore are not furnished with any 
celomic mantle. Of the anterior nephridia those in the genital somites are some- 
what smaller than those in the vicinity of the pharynx, the increase in size forward 


being gradual. Blood capillaries are found in great numbers on all the nephridia, but 
Memorrs, Vou. II, 5. December 14, 1895. 


132 CALIFORNIA ACADEMY OF SCIENCES. 


especially so on the posterior ones. The general structure of the nephridial canals 
and their windings resembles those of fully developed mega-nephridia (figs. 39, 40, 
41), but the smallness of these organs prevented me from definitely ascertaining the 
relative size and structure of all the canals. The canal leading to the outlet duct is 
strongly ciliated. Fig. 41, which represents the inner part of one of the anterior 
nephridia is held somewhat diagrammatic, as the respective thicknesses of the canals 
could not be properly delineated. I believe the most anterior nephridia are found in 
iii, in which somite they are very large, crowding each other. 


Benhamia palmicola n. subsp. 
Figs. 45-55. 


Halitat. Miraflores, in the Cape Region of Baja California, some 40 miles 
north of San José del Cabo; also around Tepie, territory of Tepic, on the Pacific 
Coast of Mexico. The altitude of Miraflores is about 1,000 feet, while that of the 
City of Tepic is about 4,000. Probably the species does not descend to the hot low- 
lands of the coast belt, at least no specimens have been found either at San José del 
Cabo or at San Blas. 

Color. The specimens are pink flesh, much less opaque than 6. nana, no pig- 
ment cells. 

General Remarks. For the present I arrange this as a subspecies under 
B. Bolavi, from which it differs in several points of considerable importance. These 
are the different form of spermathece, the situation of the first dorsal pore, the differ- 
ent sculpture of the largest penial seta, the location of the gizzard, the number of 
sperm-sacs, etc. 

EXTERIOR CHARACTERS. 

Size 50 to 60 mm. 

Somites 110. Seas 

Clitellum incomplete in all somites xili—xx,. 

Dorsal pores, most anterior one between iv—y. 

Spermathecal pores in line with sete 1 and 2, between vii/vili and viii/ix. 

Oviducal pore. One central median ovidueal pore in xiy. 

FPenial sete, two in each sac. The longer seta has apex curved and furnished 
with four blunt, bristle-like crenulations. The smaller seta has the point slightly 
forked, one prong being longer, and a thin membrane extended between the two forks. 

Common sete. About equal distance between the dorsal and ventral couples 
as between the ventral couples. 


INTERNAL CHARACTERS. 


Buccal cavity with a dorsal pocket of sense organs and glandular cells. 
Pharynx not situated far forward. 

Gizzards both apparently in viii. 

Calciferous diverticula, 5 pairs in xy, xvi and xyill. 

Septal glands, small in ix and x. 


PACIFIC COAST OLIGOCHETA. 133 


Typhlosole small. 

Sperm-sacs in xi and xii. 

Spermducts not thickened in the somites near the pores. 

Prostates are larger than in 6. nana, but mostly confined to one somite each. 

Spermathece in vii-vili and yili-ix. The two divisions are of equal size, and 
globular; small tubercular diverticle pointing forwards. 

Nephridia in three distinct rows on either side of the median line. Nephri- 
dium a nearest the ventral ganglion consists of two unequal parts, the most ventral 
of which is the smallest. These two parts together are much larger than either of 
the other two nephridia. The dorsal nephridia are far apart. 


DETAILED DESCRIPTION. 


Body-wall and sense organs. We find the same zone of sense organs in the 
body-walis as in B. nana. In B. palmicola this zone is found in all the somites, 
while in B. nana I found none in somite i. 

The sense-organ zone with pellucid cubic cells is in this species larger than in 
B. nana, and stands out most prominently on the upper side of the buccal cavity. 

Septal glands. As has already been stated small septal glands are found in 
somites ix and x or posterior to the gizzard. These glandular masses are quite small, 
are situated in the anterior part of the somite, close to the intestine, but hardly pro- 
jecting above the latter. I cannot find that they are in anyway connected with the 
pharyngeal system of glands. 

Intestine, etc. The portion of the pharynx is not far forward. The region of 
its epithelial columnar cells is thick and the discharge chambers of the pharyngeal 
glands are tubular, instead of globular as in B. nana. The pharyngeal glandular 
mass is less lobed than in B. nana. 

The region between somite iand pharynx is much longer in B. palmicola than 
in nana. The specimen sectioned longitudinally showed the two gizzards as both sit- 
uated in viii, and there was no septum separating them. If this character is constant 
is undecided as I did not wish to sacrifice another specimen. The tubular intestine 
is long and cylindrical and furnished with the regular three pairs of calciferous 
diverticula in somites xy, xvi, xvii. No lime crystals in the posterior pair. The 
sacculated intestine commences in xvill or xix. The typhlosole is smaller than in 
B. nana, but owing to sand I could not definitely ascertain its location. 

Spermathece. ‘The four spermathece are of the same form and size. The 
contraction at the center is deep, dividing the organ in two equal, globular sacs, each 
sac being confined to one somite; that is, the apical part is situated entirely in the 
somite posterior to the pore. The basal part carries a short narrow cylindrical and 
tubular diverticulum pointing forwards. The equality in size and the globular form 
of the two parts of the spermatheca appears very constant and characteristic of the 
species. 

Sperm-sacs are sac-like, not racemose. They are found in xi and xii, while 
in B. nana they are placed in x and xi. 


134 CALIFORNIA ACADEMY OF SCIENCES. 


Spermducts are considerably thickened, but the lower part is not any thicker 
than the upper part, as in B. nana. 

Gonads are affixed high up on the septum and not at the junction with septum 
and body-wall. 

Prostates. These bodies are larger than in B. nana, but nevertheless generally 
confined to one somite each. The penial sete are, as usual, of unequal size. The 
larger seta is furnished with four notches, the smaller is spoon-like and forked at the 
apex. 

Nephridia. These organs are arranged in three distinct rows on either side of 
the median line, but the nephridia in each somite are of unequal size. The nephri- 
dium nearest the ventral ganglion or nephridium @ consists of two, more or less, 
separate parts, evidently a tendency to diffusion or an imperfect centralization. The 
most ventral part is the smallest and the most distal the largest. The ducts run con- 
tinuously between these parts, but the ecelomic cells are grouped in such a way that 
the bridges between the two parts are quite narrow. The nephrostome of nephridium 
a was always plain and readily seen, but I never succeeded in finding the nephro- 
stomes of bande. Still, these nephridia appear perfectly formed on the meganephrie 
principle, and I could never see any connection by canals between a and d, and } and 
c, though sometimes the ccelomic cell masses extended more or less continuously over 
and between the respective nephridia a, 6 and ce. 

Nephridia @ open in front of setee 1 and 2, while in &. nana they open in front 
of 3and 4. The most anterior nephridium possessing a ccelomic covering I found in 
xxl. Through the courtesy of Dr. Michaelsen, I have received specimens of Bb. 
Bolavi for comparison. The nephridium of B. palmicola resemble that of B. Bolavi, 
but is much larger, and the respective nephridia cover each other slightly, while in 
the two specimens of 6. Bolavi which I dissected the respective nephridia were sep- 
arated by considerable distance; the latter nephridia are also smaller. I had at first 
intended to assign these species of Benhamia possessing several nephridia of a perfect 
form under a separate subgenus, when my attention was called] to the fact, by Dr. 
Michaelsen, that 6. Stwh/manni sometimes possessed a similar arrangement of nephri- 
dia as those in B. Bolavi, ete. As the nephridia of B. Stuh/manni are generally 
plectonephric or diffuse, it became at once evident ‘that this distinction could not be 
used as a generic character of value, and that it really is impossible to draw any dis- 
tinct line between a plectonephric and a micronephric condition. It is, however, en- 
tirely incorrect to characterize these nephridia as a mass of tubules, etc., as wherever 
they are separated one from the other, as in Bolavi, palmicola, nana, rugosa and 
probably great many other species, each micronephridium is perfect in itself, and built 
on the same general principle as the meganephridia of the other terricole. I would 
therefore propose to make a distinction between plectonephidia, or really diffuse 
nephridia, and micronephridia, or nephridia of small size, but perfect, or built on the 
meganephric plan. Such a distinction may be useful in descriptions, even if they are 
not morphologically distinct. 


PACIFIC COAST OLIGOCHATA. 135 


Benhamia papillata n. sp. 
Figs. 43 a, B, c, D, E; 52. 

Habitat. epic, Territory of Tepic, 4000 feet; Mexico. 

Tam unable, on account of want of time, to add any detailed description to the 
short definition of this species given above. When I described Benhamia palmicola 
I possessed only a single specimen of 6. papil/ata,and I supposed that the differences 
in the structure of some of the organs, as well as the presence of the prostate papille, 
were due exclusively to individual variation. But while this paper was being printed 
I became possessed of about twenty specimens from the same locality, all of which 
resemble each other in all the points referred to in the description, and I therefore 
do not hesitate now to assign to them a specific name, especially as I find that this 
species is much more distinct than [ at first suspected. While it in many respects 
resembles 5. Golavi, it differs in other points which must be considered of specific im- 
portance. 5. papillata differs thus from 2. Bolavi in possessing four exterior tubercles, 
one for each prostate pore. The smaller penial seta in B. Bolavi is flat and somewhat 
forked, while in B. papillata the smallar seta is furnished with a sigmoid tip of ex- 
ceeding thinnesss. The clitellum in Bb. Bolavi and Bb. palmicola is incomplete, but 
in B. papillata it is complete even on the ventral side, showing several rows of glan- 
dular cells, but the width of the layer of clitellar cells is much narrower in the central 
part. Clitellum is ventrally complete only in somites xiv, xv and xvi. 

The following are the other points of interest as regards the character of this 
species: It is larger than Bb. palmicola. The four papillae, each one of which 
carries a prostate pore, are very distinct and prominent, and they stand close together 
in the sunken genital pit of the clitellum. The median central papilla, on which open 
the ovipores, is elevated and oval. The two ovipores are situated in the center of the 
papilla, but entirely separate. They are in the very center of the somite and ina 
line drawn between sete 1 and 1. In B. palmicola the ovipores join, and the common 
pore is situated somewhat in front of a line drawn between sete 1 and 1. Longitu- 
dinal section of a specimen shows that the septum separating somites xiii/xiv is very 
much cupped, and the oviducal funnel is situated exactly above the central papilla in 
xiv and dips down straight to the ovipore. The anterior septe are hardly thicker 
than the posterior ones, but they are all very much cupped. Sacculated intestine 
commences in xix. There is a strong superior typhlosole in xx to xiv. Of the three 
pairs of calciferous diverticula the anterior one in xv is very much, or about four 
times, smaller than each of the posterior ones. The glandular part of the prostate is 
folded and quite thick, but confined to one somite. 

Penial sete. 'The penial sete are the most distinct character of this species, 
besides the tubercles of the prostate pores. In general shape the two sete are much 
more slender than those of 5. palmicola, and several times narrower at the apex. They 
are so thin that it required an oil-im. 1/12 to show their structure sufficiently to enable 
me to sketch it. The larger seta is slightly curved, gradually tapering from the root 
to the apex. The apex is much less curved than that of B. Bolavi or B. palmicola, 
and furnished with seven or eight shallow but still distinct notches. But the smailer 


136 CALIFORNIA ACADEMY OF SCIENCES. 


seta differs entirely from that of the B. Bolavi group. Instead of being more or less 
distinctly forked and wider at the apex, as in those species, it is drawn out to an ex- 
ceedingly thin hair-like point, which is sigmoid (fig. 43 ¢). In shape the seta is almost 
straight, with the two ends slightly curving. 

The papille of the genital region are most characteristic. Each papilla forms 
the exterior apex of a prostate. With a low power lens these papille appear round, 
but under a higher power they appear rosette-like, and consist of two or three divisions, 
one on top of the other. The ventral part of segments xvii, xviii and xix, carrying 
the genital pores, is much contracted, and the papille appear as ina bunch. The 
distance between the male or spermaducal pore and the prostate papilla is just about 
the same as the base of the papilla. 

In a section the structure of the oviducal median papilla appears to greatly 
resemble that of the tubercula pubertatis of Sparganophilus. I[ find there the same 
relatively large number of tall, narrow cells, radiating from the ovipore, but I cannot 
discern any sense cells. 

The spermathece differ in shape from those of B. palmicola. The apical sac 
is not globular, but pointed and narrow, as represented in fig. 52 G and H. 

The nephridia also differ some from those of 6. palmicola. In this species the 
inner nephridium, or @, consists of two parts, each one with a coelomic mantle separate 
from the other, but in 6. papil/ata nephridium a consists of three distinct parts, each 
one with a ceelomic mantle. ‘This, of course, refers only to the posterior nephridia; 
the anterior ones are not covered by ccelomic mantle. 

Sperm-saes in x and xi, not racemose. In other respects, as far as I can judge 
by a hasty examination, this form resembles 6. palmicola. 


Benhamia rugosa np. sp. 
Figs. 56-63. 

Habitat. Native habitat unknown. The eight specimens in my possession 
were found in the orchid house in the Golden Gate Park of San Francisco, California, 
under pots. Two were adult, two imperfectly developed, the others immature. 
July, 1895. 

Color reddish flesh. 

EXTERIOR CHARACTERS. 

Size, 830 mm. by 15 m. 

Segments, 118. Cephalic lobe very long, pointed. 

Clitellum incomplete. 

Dorsal pore, most anterior y—vi. 

Oviducal pore, one single in the median line in xiv. 

Prostate pores, the anterior ones elevated on tubercles. 

Penial sete. Largest seta with five notches, smallest seta forked with prongs 
of equal size. 

Common sete. All ventral in four couples; the inner sete are present in xviii. 


PACIFIC COAST OLIGOCHATA. 137 


INTERIOR CHARACTERS. 


Gizzards, two in vii and viii. 

Calciferous diverticula. Three pairs in xy, xvi and xvii. 

Sacculated intestine commences in xviii. 

Typhlosole present. 

Prostates slender, confined each to one somite. 

Spermathece. The apical part a trifle smaller than the basal. Both parts 
mnch flattened. A very small diverticulum pointing forwards. 

Nephridia in four rows on either side of the median line. The posterior ne- 
phridia with ccelomic glandular mantle. Nephridia in each somite of about equal 
size. Nephridia d and d about equidistant as a and a. The most ventral part of 
nephridium a not covered by the mantle. The clitellar nephridia larger than those 
anterior to clitellum. 

DETAILED DESCRIPTION. 


Size. The specimens had been slowly killed and extended before hardening. 
All the specimens were curved backwards, thus with the dorsal line on the inner 
margin of the crescent, causing the male pores to be situated at the greatest bend on 
the outer margin of the crescent. Most terricolee curve the opposite way. 

Shape of segments. ‘The first ten somites are very nearly of the same diameter 
in the direction from head to tail. Somites xi and xii are slightly narrower. The 
clitellum which comprises somites xili to xx shows plainly the intersegmental grooves. 
The posterior somites are of a somewhat shorter diameter than xi and xii. All the 
anterior somites including the most posterior ones of the clitellum are sculptured by 
longitudinal furrows running in the diameter from head to tail. This corrugation is 
seen all around the body, and gives the anterior part of the worm a very marked 
appearance. The posterior somites show a fainter corrugation. All somites are 
3-ringed, except those of the clitellum. 

Prostomium is long and pointed and slightly curved upwards (figs. 57 and 58) 
and is a trifle longer than somite 11. Inall the specimens but one, somite i, was nearly 
entirely retracted in the buccal cavity, and when viewed from the exterior only a 
short portion of its dorsal part could be seen (fig. 58) and the prostomium appeared 
as if projecting from somite li. One specimen had somite i extended as figured 
in fig. 59. Dr. Michaelsen has remarked a somewhat similar retraction of somite i in 
B. kafuruensis. Spermathecal pores as usual, vii—viii and viii—ix. 

Clitellum is narrower than the surrounding somites. It is strongly corrugated, 
especially at the anterior and posterior margins. The prostate pores are, as usual, in 
xvii and xix. The exterior part of the genital region is very characteristic. In 
young specimens the two prostate pores are connected by a deep furrow in the 
center of which is seen the slight depression for the male pore. In the two fully 
adult specimens, however, the two anterior prostate pores were each situated on a 
very large globular papilla, surrounded by a deep fossa; while no such papilla and 
fossa characterized the posterior pair of prostate pores. In fact, these posterior pores 


138 CALIFORNIA ACADEMY OF SCIENCES. 


were not visible from the exterior and could only be. ascertained by dissection and 
transparent light. Viewed from the exterior the clitellum appears incomplete. 

Penial sete. The longer seta is hooked at the apex and furnished with five 
notches on the inner side, very much as in B. Bolavi. The shorter seta is less curved 
and about ? as long as the larger sete. It is not hooked and the free apex is slightly 
forked, each prong being of the same length. There is no flare or wing between the 
two prongs (figs. 60, 61 and 62). 

Oviducts. Both open together in a central pore in the median line in xiv, 
very close to the groove between xiii-xiy. In the largest of the two adult specimens 
the ovipore was situated abnormally between xv and xvi. : 

Intestine and glands. A strong supra-pharyngeal gland as usual. Gizzard in 
vii and viii, separated by a thin wall, as in other species. Tubular intestine extends 
from gizzard to xvii, and is in xy, xviand xvii furaished with the three pairs of 
calciferous diverticula, of which the two anterior ones are of different construction 
histologically just as in other species. The sacculated intestine commences in xviii. 
There is a typhlosole. 

Spermathece. As usual, in vii and viii. Those in vii were slightly smaller, 
and entirely confined to their somite, while those in viii projected their distal part 
through the septum into ix. The distal part is only a little smaller than the other 
part. The spermatheca is much flattened. Seen from below it appears as in fig. 52 m, 
while seen from the side it looks as in fig. 52 n. The lower part is furnished with 
a very small diverticulum, pointing forwards. The duct leading to, the pore is very 
short, almost not set off from the lower sac. 

Oviducts are slender, tubular, with a small and narrow funnel. 

Nephridia. Those posterior to xix are furnished with a coelomic cell mantle 
of oval form, even and regular, and of the same size in the various nephridia. Pos- 
terior to xix we find four rows of nephridia on each side of the median line, the rows 
being reguiar and parallel. The most ventral nephridium or a is situated between 
the setze couples 1 and 2 and 3 and 4. The other nephridia 4, ¢ and d are dorsal and 
lateral. The two dorsal nephridia d and d are about as equidistant as nephridia a and 
a, and much closer together than the dorsal nephridia of ‘B. nana, palmicola and 
Bolavi, the three other species which I have examined. The more ventral part com- 
prising the outlet duct, ete., of nephridium a is longer than the corresponding parts 
of the other nephridia, and not covered by the ecelomic mantle. The clitellar nephri- 
dia are larger and their duets thicker than the other anterior nephridia. The 
nephridia anterior to clitellum are very small, except those in somites iv and v, where, 
as usual, the nephridial ducts are long. Each nephridium appears perfect, and built 
on the meganephric plan (fig. 63). 

Benhamia rugosa is readily recognized and characterized by the large papille 
on which open the two anterior prostate pores, by the four rows of nephridia, by the 
forked smaller penial sete, by the corrugation of the anterior somites, inclusive of 
clitellum, by the pointed prostomium, by the flattened spermathece. 

Since this was written Rosa has described a new American Benhamia from 


PACIFIC COAST OLIGOCHZETA. 139 


Paraguay, viz.: B. octonephra, which appears to resemble my B. rugosa in several 
points, especially in having four rows of nephridia on either side of the yentral median 
line, or eight parallel rows in all. But it also differs in several others, sufficiently to 
be arranged under separate species. . octonephra has about 90 somites, B. rugosa 
about 118. B. rugosa possess penial setee, which, as far as I can judge from Rosa’s 
description are different from those of L. octonephra. 

Unfortunately, the characteristics of Benhamia species must be founded on 
most minute characters, which cannot always be expressed in words, as long as so com- 
paratively few species are known. Too few of these characters have been figured, 
and many have been entirely overlooked. The penial setee, which are of the utmost 
importance as species characters, should be in all instances figured. They vary much 
less than do the spermathecze, the shape of which is only approximately constant. 


ACANTHODRILUS Perrier. 


Out of thirty-five species of this genus, which have been described to date and 
recognized by Beddard, none have been found as far north as Central America or 
Mexico. The two new additions to the genus which I am able to describe below will 
therefore prove of considerable interest on account of their habitat so far north, while 
one of the species shows a combination of characters rarely met with in this genus. 


DEFINITIONS OF SPECIES. 


Acanthodrilus tamajusi n. sp. 


Derinition. Length 15 cm., width 1-cm. in thickest part in front of clitellum; 
number of somites about 218. Clitellum xviii to vx. First dorsal pore posterior to clitel- 
lum. Penial setw ornamented, one seta only in each sac. Common sete paired.  Pros- 
tate pores each on a round papilla. Gizzard inv. Caleiferous diverticula 3 pairs in 
ou, out and iw. Saculated intestine commences in xiv. Typhlosole present. Sperma- 
theca large sac-like, with a few small warty diverticula. Sperm-sacs not racemose, in ix, 
x, vt, aii. Prostates four, much folded, opening with penial sete. Nephridia large, 
meganephridia, not alternate. Color reddish, partly pigmented. Habitat, Guatemala, 
lowlands. 

Acanthodrilus Vasliti n. sp. 

Derrition. Length 6 cm. by 2mm. in the region of somite viii; number of 
somites 92. Prostomium distinct, divides somite 1 about 4. Olitellum unknown. — First 
dorsal pore vii/viii; first large dorsal pore ia/x. Penial sete absent; common. sete 
paired, 1 and 2 not present in xvvi and xix. Prostate pores not on papilla; each pore a 
duplex one. Ovipores in front of sete 1 and 2. Spermathecal pores in front of 
sete 1 and 2, and between somites vii—viii, viti-ix. Spermiducal pores in xviii in line 
with sete 1and 2. Septa v to wii slightly thickened. Gizzard in v.  Sacculated intes- 
tine in xv. No calciferous diverticula. Peritoneum covered with a single layer of very 
large glandular cells. Very large suprapharyngeal gland, and a very small subpharyn- 


geal gland. Spermatheca long tubular in viii and ix, no diverticula. Prostates 8 in 
Memorrs, Vou. II, 5. December 16, 1895. 


140 CALIFORNIA ACADEMY OF SCIENCES. 


number, 2 and 2 opening close together in the prostate pore. Typhlosole. Nephridia large, 
meganephridia, not alternate, opening in front of sete 3 and 4. Color milky white, no 
pigment. Habitat, Tepic, Mexico, 4000 feet altitude. 


Acanthodrilus tamajusi n. sp. 
Figs. 87-96. 


Habitat. Of this species I possess one adult and half a dozen young speci- 
mens, principally from Tamaju on the rio Polochic in Guatemala. The largest in- 
dividual measures about 150 mm., but even this one did not have the clitellum per- 
fectly developed, but was otherwise seemingly mature. One of the smaller specimens 
measuring only 70 mm. in length possessed a more perfect clitellum than the larger 
specimen. ‘These specimens were found by myself several years ago in June and 
July on top of the ground after rain. It is probably a common species in tropical 
Guatemala. On account of imperfect preservation of the specimens, some histological 
details are wanting. 

Color. Deep violet-brown and iridescent when alive. Skin pigmented strongly 
on the tail, slightly on anterior part, but only on the upper side in the two mature 
specimens. A large immature specimen possessed no pigment, and was of a grayish 
color, but there may be some doubt as to its belonging to this species. 

Size. Largest specimen 15 em. by about 1 em. in front of clitellum. The 
specimen had not been extended and killed before being placed in alcohol and must 
therefore have been considerably longer. 

Number of somites 218 about, those in front of clitellum much larger than 
those posterior to it. 

First dorsal pore in the pigmented specimens only posterior to clitellum. In 
the unpigmented between vii and xiii. 

Prostomium doubtful (injured), but appears to incompletely or not at all divide 
somite i. 

Clitellum complete, xiii to }xx. 

Penial sete. A sac with one penial seta at each prostate. This seta has 
numerous short rows of teeth, decreasing in number towards the apex. ‘The seta is 
quite short, blunt at one end and broad, tapering gradually toward the apex which is 
slightly recurved. The whole seta is irregularly sickle-shaped. The teeth are on 
the concave side. One seta possessed four notches with short teeth also on the con- 
vex side posterior to the other, or about where the thickest part begins (figs. 93 4, B, c). 

Common sete strictly paired on ventral side of body. The distances between 
the couples about the same. The ventral couples of sete are present near to the 
spermiducal or male pores. They are the ordinary form but slightly larger than the 
common sete. 

Oviducal pores in xiv situated somewhat nearer the median line than to the 
inner couples of sete, and the distance between the pores is a trifle more than the 
distance between setee 1 and 2 in the same somite. 

Prostate pores situated each on a small papilla. 


PACIFIC COAST OLIGOCHATA. 141 


Gizzard is in v. 

Calciferous diverticula, three pairs, one each in yii, vili and ix, all of the same 
structure. 

Sacculated intestine commences in xix and furnished with a typhlosole. 

Spermathecw in viii and ix with a few warty diverticula, one of which is 
larger and furnished with two swellings. Spermathecal pores in front of sete 1 and 
2. The spermathece are large, filling the space between the septa on the ventral side 
of the body. No spermathecal copulary sete. 

Sperm-sacs sac-like, not racemose, occupying the whole space left in ix—xii. 

Prostates much folded, opening with penial sete; each prostate confined to 
one somite each, the free distal end pointing backwards, and situated close behind the 
male pore. 

Nephridia, strictly meganephridia, not alternating. 


DETAILED DESCRIPTION. 


Each prostate pore is situated on a round globular papilla, between which and 
the male pore runs a copulatory fossa. In xyiand xx there is in each a copulatory 
ridge, running parallel with the intersegmental grooves and somewhat longer than the 
space between sete 2 and 2 (fig. 91). There is a shorter ridge in xviii extending 
between the male pores. 

The prostomium and somites i and ii are transversely suleated, most so somites 
iand ii. Also somite ili shows this sulcation in a smaller degree. The anterior nine 
somites are more or less distinctly segmented in three parts, that is, they show a central 
ridge on which are situated the sets. The following posterior somites show a seg- 
mentation in five parts. In the clitellium the segmentation is clear, as far as regards 
the intersegmental grooves, but the segmentation of the somites is not very distinct. 

The supra-pharyngeal glands form a very large body, superposing the pharynx. 
It is rounded posteriorly, and consists of four larger and five smaller lobes, this when 
viewed in a longitudinal section just outside the median line (fig. 96). 

- The calciferous diverticula show all the same structure. 

The spermathece are large, globular masses, as seen in figs. 94 and 95, with 
warty diverticula, one of which is much larger. The spermathecz overlap each other 
and are bunched in a solid mass between the septa on the ventral side of the body. 

Testes are in two pairs, quite or very small when viewed in sections—in x and 
x1, in front of the ciliated rosettes, on the posterior face of the anterior septa. 

The ciliated rosettes or sperm-funnels are in xi and xii, and are very large, 
crimped and furnished with a wide muscular duct. The funnels in xii lie much 
higher up than the one in xi, much closer to the intestine. The posterior part of the 
funnel is very thick, and so is the duct. The ducts join and open as usual in xviii. 
The rosettes lie free, but are connected by connective tissue with the anterior septa, 
They thus point backwards. 

The body-wall is very thick, and the longitudinal muscles are not bipinnately 
arranged. The septa between v, vi and so on until xi, xii, are much thickened, and 
thicker than the other, though the one between xi, xii is not as thick as the anterior 


142 CALIFORNIA ACADEMY OF SCIENCES. 


ones. The gizzard is connected by several powerful muscular strands with the body- 
wall in x1. 

There is a very large mass of pepto-nephridial tubes situated immediately above 
the posterior part of the pharyngeal gland. Longitudinal sections show the anterior 
nephridial canals to be greatly folded and very narrow. There are very large masses 
of free ccelomie cells situated in the somites containing the sperm-sacs and the funnels 
of the spermducts. 

Acanthodrilus Vasliti n. sp. 


Figs. 148-154. 


General Remarks. This species is one of those abnormal forms which occur 
in almost every large genus, and whose organization and characteristics are not readily 
accounted for. It is also the most northern of any Acanthodrilus found so far, though 
undoubtedly true Acanthodrili will be found much further north. Acanthodrilus 
Vasliti differs from any other Acanthodrilus in possessing eight prostates or spermi- 
ducal glands, arranged in four pairs, two and two prostates opening together in each 
one of the four prostate pores in somites xviii and xx. Also in one other respect does 
this species show an interesting characteristic. The peritoneum lining the septa and 
body-wall is covered with enormous glandular cells, very much resembling those 
forming the nephridial mantle in many species. The duplication of the prostates is 
also found in Kerria MeDonaldi, while abnormal development of peritoneal cells re- 
mind us of certain coelomic organs found in Perichsta and some species of Acantho- 
drilus (Beddard, page 29, Monograph of Oligothzeta). Four pairs of prostates have 
been described by Ude in Geodrilus singularis, but details are wanting. 

Of the specimens in my possession one was sectioned longitudinally, two were 
dissected and afterwards sectioned vertically. None of the specimens were fully 
adult; the various generative organs were developed, but there was no trace of 
clitellum. 

Habitat. Tepic, Territory of Tepic, Mexico, at 4000 feet altitude, in the 
moist ground immediately under decaying logs, in the shade of a stone-fence, about 
one mile north of the city. October, 1894, Eisen and Vaslit, col. 

Color. Milky white, like an Enchytreus, without trace of pigment. 

Size. Length 6 em. by 2 mm. in the region of somite viii. Slightly tapering 
towards the tail, the end of which is thickened. 

Number of somites 92 in the largest specimen, all of about the same size, ex- 
cept the last few caudal ones, which diminish in width towards the most posterior 
somite. All are smooth, the anterior ones with a faint trace of trisegmentation. 

Dorsal pores. The most anterior one that is distinct is seen between ix—x. 
Between yii/viii and viii/ix there is respectively a much smaller but still distinct pore. 
The most anterior pore is thus between yvii/viil. 

Prostomium is distinet, dividing somite i about $. The anterior somites are 
more distinctly set than the others. A long narrow groove begins on the ventral 
median line between somites xvi and xyiii, and extends backwards about 20 to 26 - 


PACIFIC COAST OLIGOCHETA. 143 


somites, then changes into a narrow keel. This in the largest specimen, all having 
been slowly killed and straightened out. 

Clitellum unknown, all specimens being immature. 

Genital pores. Two pairs of spermathecal pores between ix—viii and vili—vii, in 
front of the respective septa, the spermathecz thus being in ix and viii. The pores 
are in line with sete 1 and 2, but are not prominent. Oviducal pores are in front of 
sete 1 and 2 between xii/xiv. 

Prostate pores in two pairs in the center of somites xvii and xix, in line with 
setee 1 and 2. But each pore is really a duplex of two joint pores, which are only 
separated at the very epidermis by a thin wall. Thus the two prostates of each 
couple run parallel through the body-wall, each one opening separately, but the pores 
being so closely joined that they appear almost as one. No setze of any kind near these 
pores (figs. 151 and 152). 

Spermiducal pores are seen in line with sets 1 and 2, close posterior to the 
setee, only visible by strong transmitted light. The genital region in my immature 
specimens did not show any particular structure with papille, ete. The median fossa 
already referred to causes these pores to be situated on a slight ridge, which was not 
marked off laterally. Close to each prostate pore on their ventral side is a small 
tubercula pubertatis, only visible in sections. 

Penal sete absent. 

Common sete. Strictly paired ventral and lateral; 3 and 4 being situated 
slightly ventral, below the horizontal line. Sete: 1 and 2 are missing in somites xvii 
and xix. In xviii the ventral setze 1 and 2 are present, but do not differ from the 
other setze, which are all plain and sigmoid. 

Nephropores in front of sete 3 and 4. 


INTERNAL CHARACTERS. 


Body-wall offers no prominent characteristics, except that the peritoneal layer 
is enormously developed in all somites posterior to xiii. . Peritoneum is strongly yas- 
cular, but in addition to the usual small peritoneal cells we meet with a thick layer 
consisting of a single row of tall peritoneal cells of varying but fairly even height. 

Peritoneal cells are in places as high as the other layers of the body-wall com- 
bined, while in other places they are shorter. Similar peritoneal cells cover also the 
septa, principally those posterior to xiii. The center of the septa bear as a rule the 
tallest cells (figs. 149 and 153). These cells show a round nucleus and a granulation 
which resembles that of chloragogen cells, but which does not stain deeply as does 
the one of the latter cells. 

The hearts in x, xi and xii, as well as the septal glands in vii and viii, are also 
surrounded by similar cells. “The dorsal blood vessel again is covered by regular 
chloragogen cells. 

Septa. Those separating respectively somites v to xii are slightly thickened 
and very strongly cupped, but not covered by any peritoneal cells. The septa sep- 
arating xii to xiv are less cupped, not thickened, but all covered by a few peritoneal 


144 CALIFORNIA ACADEMY OF SCIENCES. 


cells. The septa posterior to somite xiy again are not much cupped, but each one is 
lined on each side by a single layer of very thick tall peritoneal cells (fig. 153). 

Alimentary canal.. Pharynx is furnished with a large upper chamber and is 
apparently only developed superiorly, although it possesses a small subpharyngeal 
gland close to the ventral nerve cord. 

(Esophagus rises diagonally upwards and joins the single gizzard situated in 
v. The tubular-intestine extends to xiv. The sacculated-intestine commences in xy. 
There is a typhlosole in the dorsal wall of the intestine in xvi to xix. The typhlo- 
sole presents a network of fibres resembling in a general way the structure of a 
sponge. This typhlosole does, however, not descend into-.the canal, but partakes 
more of the nature of a wide continuous blood-sinus. 

Salivary glands. The suprapharyngeal glands form a mass with five distinct 
lobes, of which the posterior one, as usual, is the largest and the anterior one the 
smallest (fig. 149). There is also a subpharyngeal gland very low but rather long 
(fig. 149). A thin but wide septal gland is found in vi, posterior to the gizzard, 
while smaller septal glands, which are principally developed ventrally are found in 
vii and viii. 

Spermathece consist of two pairs of long and narrow organs in viii and ix 
opening in the intersegmental grooves between vi and viii and viii and ix. In my 
specimens they were probably rather undeveloped and did not show any trace of 
diverticula either externally or in the wall. 

Testes are in x and xi and ovaries in xiii. The oviduct opens between the 
sete and septum. Spermiducal funnels or rosettes are very small, thick and compact, 
and situated in x and xi. The spermducts run separately backwards between the 
longitudinal layer and peritoneum and open jointly on the center of xviii as usual. 

Prostates are in four pairs as has been already stated, two and two opening in 
each pore in line with sete 1 and 2, these sete, however, not being present in these 
somites (fig. 150). The prostates showed no glandular part, the whole being muscu- 
lar (fig. 154). They were very thin, tubular, the two prostates in each twin couple 
running entirely parallel and close together along the septa, as far as the line of sete 
3and 4. A large part of this distance the muscular part is surrounded by regular 
peritoneal cells, not by the large glandular ones. Each prostate remains separate 
from the other and even their external pores, though situated close to each other, are 
not strictly joined, though they are surrounded by a common thicker lip (figs. 150 
and 151). I have already referred to the duplication of prostates in one species of 
Kerria, otherwise it has not been found with certainty in Acanthodrilid. Some of the 
earliest described Acanthodrilides were, however, supposed to haye a prostate and 
spermducts open jointly, and it does not seem unreasonable to suppose that in some 
case at least a duplicate prostate existed, and one was mistaken‘for a spermduct. This 
would be quite easily done in specimens poorly preserved, especially if the prostates 
should be entirely muscular as in Acanthodrilus Vasliti, in which species they are 
also very narrow and thin and not really wider than the spermducts of many forms. 
Ude, in describing Geodrilus singularis from Danville, Illinois, mentions as one of the 


PACIFIC COAST OLIGOCHATA. 145 


generic characteristics four pairs of prostate glands, but does not describe nor figure 
these in detail. The only way I can understand the presence of four pairs of pros- 
tates opening into four pores, is that two and two prostates open together. But on 
the following page (70) we are told that there are two pairs of prostates in segments 
18 to 22, which open in segments 17and19. The figure shows us four prostate pores. 
If the first statement is not a misprint, we would in Geodrilus singularis have an 
analogy similar to what we find in Acanthodrilus Vashti. But Beddard states in his 
large monograph on Oligochieta that Geodrilus singularis is probably identical with 
Diplocardia communis. This could, of course, not be the case if the prostates were 
duplicated in the former. 

Tubercula pubertatis. Although no large and prominent elevation in the 
genital region is found, I have, however, satisfied myself that a tubercula pubertatis 
is really present even in my undeveloped specimens. Adjoining the prostate pore is 
a small tubercle, consisting of some tall supporting cells surrounding a bundle of sense 
cells, the latter characterized by the usual long, oval nuclei. These nuclei are 
situated much deeper than they are in the sense cells of the epidermis or in the buccal 
cavity, and they are also narrower than those. Otherwise this organ shows a very 
great resemblance to those in Benhamia, except for the absence of the glandular 
refractive cells. At the base of the sense cells in the tubercle I find numerous smaller 
round nuclei, the relationship of which my sections do not fully explain. Numerous 
nerve fibrils are seen to connect with the sense cells. Although no clear glandular 
cells are seen around the sense-organ in the tubercle, it may be possible that some 
may develop later at the same time as the clitellar cells, none of my specimens _pos- 
sessing any. 

Vascular system. The last heart is in xi. The dorsal vessel is single; no 
subneural vessel. A large blood sinus in xvi to xix inclusive. 

Nephridia are strictly paired, opening in front of setee 8 and 4. ‘Those posterior 
to somite xiv are surrounded by large peritoneal cells, while those anterior to xv con- 
sisted of the usual narrow ducts, free of any coelomic peritoneal cell mantle. Numer- 
ous blood vessels cover the nephridii. The most anterior nephridium is found in iii. 
No pepto-nephridium. The anterior nephridia gradually diminish in size towards the 
anterior part of the body. 


ALEODRILUS pn. gen. 
Figs. 66 to 86. 


Derrition. Acanthodrilide. Paired meganephridia, not alternating. Pros- 
tate pores, two pairs; one pair in wx and one par in xxii. Svermiducal pair of pores 
in wat. No calciferous glands; no penial sete. Two pairs of spermathece in viii and 
ix. Clitellum witi—vx. Two gizzards. 

General remarks. I possess only one single specimen of this interesting oli- 
gocheta, collected at Ensenada de Todos Santos in the northern part of Baja Califor- 
nia. My time for searching was very limited and the season unfavorable, and this 
will account for the want of specimens. The single specimen was found in the dry 


146 CALIFORNIA ACADEMY OF SCIENCES. 


bed of the river south of town at a water-hole in the otherwise dry creek bottom 
sand. It occurred here with Deltania and Limnodrilus. The sand was merely moist 
on account of overlying rubbish and sacks. The species is undoubtedly a native one 
and the only Acanthodrilid found on the coast so far north in the open ground, and 
and on this account even of geographical interest. ‘The intestine of the worm was 
gorged with the coarse white sand of the river bed. The anterior part of the worm 
was cut lengthwise, one-half dissected and the other half sectioned crosswise. The 
want of specimens made a full investigation impossible, though I believe none of the 
important points remains in doubt. I have no reason to believe the species is scarce, 
though probably it issharing the fate with all native worms, that of being displaced by 
European importations. The species is dedicated to Professor W.S. Keyes, my com- 
panion in many travels in tropical Mexico. 

Affinities. It is interesting to note that Aleodrilus shows considerable affinity 
to the only other North American genus of this family, viz.: Diplocardia. It re- 
sembled this genus by having two gizzards, no calciferous glands, meganephridia, no 
sacs with penial sete. It resembles Benhamia in having two gizzards, ete., but it 
differs from these two, as well as from all other Acanthodrilide, by the far backward 
position of the prostate and spermidueal pores, these being in xx, xxi and xxii re- 
spectively, while all other genera of this family have these respective pores in xvii, 
xvili and xix. The genital male pores are thus in Aleodrilus pushed three somites 
further back. Considering these and some minor characters I believe I am justified 
in placing this worm in a new genus. 


Aleodrilus Keyesi n. sp. 


Derinition. Length 7 cm., by 5 mm. wide; number of somites 80. First dor- 
sal pore viii-ir. Clitellum complete in anterior, incomplete in the posterior somites, 
Lvititex. No penial sacs and sete. Common setw paired, those of the inner couple 
closer than those of the outer couple. Spermathecal pores between vii/viii and viti/ix. 
Gizzards inv and vi. No caleiferous glands. Nephridia not covered by a celomic man- 
tle. Nephropores outside of sete 4. Hearts in x, wi, xii. Sperm-sacs racemose iD) Hibs 
vi, wii. Testes ina, wi. Color pale flesh, no pigment. Habitat, Northern Baja Cal- 
ifornia, at Ensenada de Todos Santos. 


EXTERNAL CHARACTERS. 


Color is very pale, mottled and marbled, showing clearly the intestines and 
blood vessels. When collected this worm resembled in delicacy of color and trans- 
parency Deltania elegans and I supposed it to be this species. Spermathecal pores are 
separate in front of sete 1 and 2 between vii/viii and viii/ix. Setw are ventral and 
lateral, 8 in each somite approached in couples. ‘The sete of the inner couple is 
closer than those of the outer couple. The distance between the couples is about twice 
as large as the distance between sete 1 and 2, and one and one-half us wide as the dis- 
tance between sete 3 and 4. No penial sete in special sacs. All sete are sigmoid 
without sculpture. The anterior five somites are two-ringed, that is with a single 
groove in the equatorial region, while all the following somites are four-ringed, or 


PACIFIC COAST OLIGOCH ETA. 147 


with two parallel grooves in the equatorial region. The posterior somites are very 
wide and four-ringed, but the segmentation is irregular. 
Ovipore a little interior to seta 1; nephropores outside of seta 4. 


INTERIOR CHARACTERS. 


Body-wall. There is a special zone of sense organs in the anterior somites as in 
Benhamia, but they are more scattered than in this genus. The longitudinal muscles 
run irregularly, with no trace of bipinnate arrangement. The strands are rather 
marrow. The longitudinal layer is very narrow, especially in the clitellum. The 
transverse layer consists of only 3 to 4 strands. The hypodermis offers nothing of 
special interest. 

The longitudinal layer in the clitellum is greatly diminished in thickness, most 
so in the lateral and dorsal region of the clitellum. Anteriorly in somites ii and iii, 
the longitudinal strands leave the body-wall and spread out fan-shaped to the inner 
wall of the prostomium, forming retractor muscles for the upper and lower lips. 

Arciform muscles. In somites xx, xxi and xxii we find on each side of the 
ventral nerve-chord several oblique or aciform muscles, running from the region of 
the copulatory grooves to the region above the lateral sete, thus serving to depress 
and relax the two grooves. These muscles are confined to a single row, and do not 
show a complex arrangement, as is so frequently shown in oligocheta. 

Septa. The septa between somites vii and xiv are much thicker than the others, 
especially thickened are those separating somites from vii to x. Those between x and 
xii are less thick than the anterior ones. The thickest septa, vii/viii, vili/ix, ix/x, are 
much thicker than the body-wall. The most ‘anterior thick septum is the one which 
posteriorly bounds the gizzard (fig. 74). 

The septum next anterior to this, the one which separates the two gizzards, 
presents the peculiarity of not being attached to the body-wall, between vi and 
vii, but it extends forward parallel with the intestine and passes in front of the brain 
on the upper side, while the ventral side is attached to the cesophagus below the 
pharynx. It forms thus a sac, as in various species of Benhamia. Anterior to this 
septum I find no trace of others. 

Alimentary canal. The pharynx is well developed and superposed by a very 
large glandular mass, which consists of about six layers of lobes, attached to muscular 
strands, as usual. The most posterior mass is the thickest. The discharge pockets of 
these glands into the pharynx are much thicker than any I have seen in other species, 
but are otherwise not of any characteristic construction. 

Septal glands are situated far back in somites vii to xi. They are of the same 
nature as those which discharge in the pharynx, but I have good reasons to believe that 
the glands in this species discharge in the tubular intestine. I have been able to 
follow the discharge duct as far as to the muscular layers of the intestine, which would 
hardly have been the case if the ducts had continued forwards into the pharynx, as 
do those of the forward septal glands in many genera. A peculiarity of these glands 


is that they are especially developed on the ventral side of the intestine (fig. 75) and 
Memoirs, Vou, II, 5. January 6, 1896. 


148 CALIFORNIA ACADEMY OF SCIENCES. 


are distinctly paired. On the median line below the intestine the ends of each 
glandular mass meet and join into one duct, the one I have just referred to. The free 
ends of the glandular masses are attached by mesenteric tissue to the ventral or sub- 
intestinal blood trunk. 

A much smaller tubular gland of the same nature runs between the dorsal 
vessel and the intestine; discharge duct unobserved. A yet smaller gland is seen 
above the subventral vessel in the same somite as the former; its discharge duct could 
not be followed. 

These glands stain exactly as the common supra-pharyngeal glands and septal 
glands, but they show no similarity as regards reagents with the chloragogic cells of 
the intestine and blood vessels, in corresponding places in the somites posterior to xi. 
The septal glands appear to be of about equal size, a close examination being im- 
possible, from want of sufficient material. 

Other glands are found in the epithelial walls of the intestine, arranged in 
clusters, like the cloves ina garlic. They are scattered about at short intervals among 
the epithelial cells, and appear of the same nature as those I have described in Ar- 
gilophilus, but they are not as numerous as in that species. They do not stain freely, 
but stand out bright and pellucid among the darker staining cells. 

Typhilosole not present. 

Gizzards are connected by a very thin wall of the same nature as the cesophagus. 
As far as I can make out the gizzard must be in v and vi, at least the posterior gizzard 
is bound by the septum separating vi and vil. The circular muscular layer is about 
30 strands wide, and is at the widest place about four times thicker than the epithelial 
layer and cuticle together. The whole width of the gizzard wall is little more than 
twice that of the body-wall in that somite. The longitudinal muscular layer of the 
gizzard is only one single strand thick, and the thickness of this strand is less than 
any one strand of the circular layer of the gizzard. The epithelial layer is com- 
paratively thick, about one-fifth of the whole gizzard. It contains the same peculiar 
glands as I figured in Benhamia. 

Spermathece. The absence of diverticula is interesting in as much as most 
species of related genera possess them. There are, however, some warty elevations. 
The muscular or basal duct is very long, slender and tubular, several times longer 
than the upper ovoid sae (fig. 74). The muscular part offers no peculiarities of 
structure. The spermathecz occupy each only one somite. They stand upright fol- 
lowing close to the anterior surface of the septum. 

Testes are greatly lobed and are situated high up on the septum, just as are the 
ovaries. 

Spermducts and ciliated rosettes. The spermducts are separated, but enclosed 
in a common muscular sheath until somite xxi is reached. Between xx and xvi the 
two ducts fuse into one lumen, which opens out into the center of somite xxi. The 
double lumen runs along the circular muscular layer forward until somite xili/xi, 
when the respective ducts rise upwards following the septum to the ovary and testes. 
From here on forward each duct is thicker and muscular, and instead of following the 


PACIFIC COAST OLIGOCHATA. 149 


body-wall passes straight through the ccelomic cavity to the anterior septa, which they 
pierce immediately at the base of the ovaries and testes (fig. 78). A double lumen 
is seen only after it passes posteriorly to the ovary. The rosettes are not large, 
but thick with a wide base; the latter is furnished, where it passes through the sep- 
tum, with several small sac-like glands, each with a distinct lumen, which I followed 
through the muscular layer of the spermduct and which I suppose empties into the 
neck of the rosette (fig. 79). The rosettes are not enclosed in the sperm-saes. 

Sperm-sacs. ‘The three pairs of sperm-sacs are racemose, but not exceedingly 
so. There are four or five large lobes of globular shape seen in every section sus- 
pended from the septum and on the under side of the intestine, though somewhat 
projecting above it. The two anterior sperm-sacs in x and xi are smaller than the 
posterior one in xii, the one in x being the smallest of the three pairs. Those in 
x and xi are suspended from the anterior septum separating x/xi and xi/xti. Of the 
position of the posterior sperm-sacs I am uncertain, but it appears suspended from 
the posterior septum, the one separating xii/xili, as far as I can judge from a eross- 
section. 

Spermatogonia. In the two anterior sperm-sacs the spermatogonia offered 
nothing peculiar as regards the development of the spermatozoa. There is in each 
sperm-sac a large central zone of peculiar cells, staining differently. They re situ- 
ated very close together, and possessed in my preparation rather shrunken nuclei 
(figs. 63 and 75). This zone exists in all the various sperm-sacs in the anterior as 
well as in the posterior ones. The development of the spermatozoa in the two an- 
terior pairs appeared entirely normal, the spermatogonia possessing the same form as 
in other species, the large nuclei standing out freely like beads above the wall. But 
in the posterior sperm-sacs the spermatogonia, one and all, looked quite differently. 
They were here of many varying sizes, some small, some enormously large, and the 
nuclei, instead of standing out from the wall of the spermatogonium, were always 
bunched in the center, or strung across it as a central band (75). In other spermat- 
ogonia the nuclei were arranged as ina ring along the cell walls, but without pushing 
out. When the nuclei were in the center there appeared always a row of large and 
small vacuoles along the cell-wall. I find two sizes of nuclei, the smaller being al- 
ways less in number, and about four times smaller than the large nuclei. When 
counter-stained with orange, the larger nuclei give quickly up part of their heema- 
toxylon, the smaller ones giving it up slowly and not at all. The cytoplasma of the 
spermatogonia presented a very strong polarity as regards its position, it being always 
massed towards the cell-wall nearest the central germinative area (fig. 63). A some- 
what similar-process of developing spermatogonia have been described by Vernon in 
the sperm cells of Bombyx. The central germinative cell in Bombyx appears to 
correspond with the central area or cell agglomeration in Aleodrilus, but I have seen 
no such budding out of the germ cells as figured by Vernon, but this may depend 
on the insufficient material at my command. 

A large number of similar spermatogonia were found in very large numbers 
in nearly all the anterior somites, either loose in the coelomic cavity, or in a 


150 CALIFORNIA ACADEMY OF SCIENCES. 


peculiar tissue which fills much of the ccelomic cavity. This tissue consists of large, 
more or less connected cells, very much dike the ccelomic glandular cells surrounding 
so many nephridia. These cell masses are more or less diffuse, and appear not dis- 
tinetly connected with any of the interior organs of the body. Among these cells 
are found scattered about large masses of spermatogonia, all in about the same stage 
of development. There are also numerous small free ccelomic cells, such as found in 
all earthworms. 

Nephridia. There is one pair of mega-nephridia in each somite. The 
nephridium is very large, projecting above the lateral line of the body. The nephro- 
pores—at least some of them—are seen outside of or more dorsal than setee 4. The 
nephrostome again is as usually seen very near sete 1. There are no glandular 
ceelomie mantles on any of the nephridia. The anterior fold is much wider then the 
posterior one. The windings are very deep and twisted, and-the spur generally sigmoid. 
The outlet duct, which is tapering towards the pore, is much darker than the other 
ducts. Its prolongation into the anterior fold forms there the central wider canal, 
which is also darker than the other two. The canal in the bridge continues forwards 
and upwards through the anterior fold, and is there the posterior one of the three 
canals. 

Vascular system. The ventral and dorsal longitudinal vessels are both single. 
There are connecting vessels in somites vii to xii... In x, xi and xii, these vessels are 
very large, and take the form of so-called hearts. Each heart consists of four or five 
links, increasing in thickness upwards. Between each two links there isa thick 
circular valve. Similar valves are seen also in the dorsal vessel at the junction with the 
hearts. At the base of some of the valves are seen two rows of very large cells, the 
nuclei of which are about 3 to 4 times larger than the nuclei in the valve cells (fig. 
83). All the valves in the hearts point downwards or ventrally. There are no 
glandular cells in the vessels, such, for instance, as are found in Pontodrilus, ete. 

Posterior to the hearts we find long, tubular connecting lateral vessels, between 
the dorsal and ventral trunks, in the anterior part of each somite. These vessels, one 
pair in each somite (fig. 86), are of even thickness throughout, but with two short 
knob-like diverticula, one above the other, about equal distances from each other and 
from the longitudinal vessels. Both the dorsal vessel, as well as the laterals, are 
thickly covered with ceelomic chloragogen cells of a yellowish opaque color. These 
cells do not cover the ventral longitudinal vessel. These lateral vessels contain no 
valves. 

Each valve consists of several circular rings, each containing a number of 
muscular strands enclosed within a common membrane. In longitudinal sections of 
the vessel most of the nuclei lie parallel with each other, but the outside ones run as 
the periphery in a circle. The smaller valves consist each of only one such lobe (fig. 
73a), while the large ones are composed of several (fig. 730). 


PACIFIC COAST OLIGOCHETA. 151 


SPARGANOPHILUS Benham. 


Derinition. Aquatic oligocheta. Hight sete in four couples. Prostomium 
not marked off from peristomium, but furnished with a superior pit. Few dorsal pores. 
Clitellum very large, from eight to twelve somites. No penial sete. Spermathece in 
vii, vii and ix, from two to eight in each somite. Sperm-saes in xi and xii, racemose. 
Specmduct always subepidermal. Male pore xviii/ix, or anterior part of wix.  Pros- 
tates generally present in three or four pairs situated several somites posterior to the 
spermiducal pores. No gizzards, wo calciferous diverticula, no typhlosole. Four pairs 
of hearts. Two pairs of lateral, longitudinal integumental vessels extending forward 
from somite xiv, not connecting anteriorly with the gut-wall and median trunks. 
Nephridia meganephric, commence in somite wit or xiii. 

Principal species characters are derived from the position of the setee; number 
and shape of spermatheci in each somite; lobulation of the sperm-sacs; course of the 
spermducts; shape and position of tubercula pubertatis, whether dorsal or ventral, to 
the male pore or spermduct; extent of clitellum; presence of a subpharyngeal integ- 
umental gland in iii; length of worm and number of somites, ete. 


GENERAL REMARKS. 

The first species belonging to this genus was described by Benham from the 
river Thames in England. Later, Frank Smith described another species from North 
America and it became doubtful, as first suggested by Benham, if the genus was 
originally an American or European one. The very restricted locality or habitat of 
the European species would indicate its probable importation from some other country, 
and when the American species was found it became almost certain that we had to 
search for the original home of this genus on the American continent. Already 
when Benham’s paper was published [ had in my possession specimens of this genus 
from Guatemala, California and the Central North American States, and there 
remained no doubt in my mind as to the native habitat of the genus. As it now stands 
we have seven species sufficiently well defined to be recognized and one more of which 
no detailed description can be had, but of which we know enough to be able to 
recognize it should it again be observed. Of these eight species then seven are 
American and one European, and everything points to the probability that the latter 
one is a lately imported species to England most likely from this country, as Ben- 
ham originally supposed. 

The species described here below are not of equal value as species as might 
be naturally inferred. Four of the species are well defined, which principally 
is due to better preserved and abundant material for study. These species are: Spar- 
ganophilus tamesis, Hiseni, Benhamiand Smithi. . Two are less well-known, due entirely 
to want of sufficient well preserved material. ‘These species are Sparganophilus car- 
neus and guatemalensis, both of which may prove only varities of Sp. Benhami. The 
remaining species Sparganophilus sonome may prove a yariety of Sparganophilus 
Smithi, but just as such all the more interesting. 

A point of unusual interest in this genus is the presence of prostates, or, as 


152 CALIFORNIA ACADEMY OF SCIENCES. 


Beddard calls them, spermiducal glands. These glands are, in Sparganophilus, situ- 
ated much further from the male or spermiduéal pores than in any other genus, 
apparently quite independent of the pores. They are also subject to considerable 
variation, in some species being three, in others four; in one species none. Of the 
same nature I consider the forward parietal pair of glands in somite ili of Spargano- 
philus Hiseni, and it seems not unlikely that originally this genus possessed many 
more pairs of spermiducal glands, perhaps one in every somite. This location of the 
spermiducal glands favors greatly Beddard’s view that these glands were originally 
independent of the spermduct, and according to this view these glands are in 
Sparganophilus the most primitive of any. There is also much difference as to the 
development of these glands in the various species. Thus in Sp. Benhami the gland 
consists of two distinct parts and is both glandular and muscular, while in Sp. Smithi 
the muscular part is degenerate or undeveloped, the whole gland being very dimin- 
utive. Another point showing the primitive arrangement of the spermiducal glands 
is the absence of any copulatory sete, the common set in their vicinity being 
unmodified. 

There is no doubt that a large number of species of this genus will soon be 
found on the American continent, especially as specimens appear numerous and widely 
distributed. In Guatemala I found them everywhere in springs and lakes, and a 
hasty examination of live specimens satisfied me that there were among them about 
three species, characterized by the position of the hearts, whether in vili—xi, ix—xi, or 
x-xiiil. Unfortunately my Guatemala collection was destroyed and only few speci- 
mens in poor condition remain. 

In this connection I will also call attention to another species of Sparganophilus 
not described below, but found by me several years ago in the small lake known as 
Laguna Puerea, situated near the ocean at San Francisco, California, or in the same 
lake as I now find Sparganophilus Smithi. This species which [I have since been 
unable to re-collect, although I have made repeated excursions to the place and dug at 
the identical spot, offered much of interest and was undoubtedly a different form 
from any now described below. It was much shorter and thicker, about 5 cm. long 
by 4 mm. thick, or about half as long and twice as thick as our smallest species now 
known. The hearts were, according to a fieldnote, situated in vii, vill, 1x, x. The 
longitudinal lateral vessels began in xiv, but most interesting of all, the species 
possessed five distinct and large eye spots on the first somite and prostomium, an 
occurrence not recorded in any other of the higher earthworms. 

The specimens were very rare—I could only find two or three in an hour; they 
occurred in the clay soil near the shore at shallow depth. 

In connection with this it is most interesting to note that at that time I found 
no trace of Sparganophilus Smithi, which now occurs in the same lake in countless 
numbers, a worm many times longer and much thinner. It is impossible that I could 
be mistaken in regard to my former observations. My explanation is this: Formerly 
the Laguna Puerca was much lower, the bottom soil near the surface was clayey, while 
now the lake is several feet higher, the bottom soil near the surface is sandy. A 


9 
3 


PACIFIC COAST OLIGOCHETA. 1 


oO 


number of vegetable gardens have of late years been established at the headwaters 
along the creek furnishing the lake, and it is probable that Sparganophilus Smithi 
has been introduced with watercress, etc., from some other locality within the last few, 
say ten years, while the eye-bearing species may yet be living in the bottom soil. 


KEY TO THE SPECIES OF SPARGANOPHILUS. 


I. Sets# 3 and 4 ventral; no prostates, spermducts dorsal to tubercula pubertatis. 
SPARGANOPHILUS TAMESIS Benham. 
II. Sets 3 and 4 dorsal, prostates present, situated several somites posterior to the male pores. 
A. <A pair of yentrally situated parietal glands in somite iii. Spermducts and spermiducal pores dorsal to 
tubercula pubertatis. SPARGANOPHILUS EISENT Smith. 
B. No ventral parietal gland in any somite. 
x. Spermiducal or male pore ventrally situated to the tubercula pubertatis. 
a. 8 spermathecm in each of somites vil, viii, ix. 2 ditto in vi. SPARGANOPHILUS SMITHI 0. sp. 
b. 4 spermathec# in each of somites vii, vili, ix. 2 ditto in vi. SPARGANOPHILUS SONOM& n. subsp. 
xx. Spermiducal pore dorsally situated to tubercula pubertatis. 


ec. Spermathecw trowel-shaped, outline crenulated. SPARGANOPHILUS BENHAMI 0. sp. 

d. Spermathece trowel-like, but outline smooth. SPARGANOPHILUS GUATEMALENSIS 1. subsp. 
xxx. Spermiducal pore unknown, but probably dorsal to tubercula pubertatis 

e. Spermatheci club-like, apical end globular, smooth. SPARGANOPHILUS CARNEUS 0. sp. 


Sparganophilus tamesis Benham. 


Derinition. Length 7to 10 em. Clitellum 1-2 av—1-2 wav. Tubercula puber- 
latis xvii to xxir ventral to spermiducal pore. Sete 3 and 4 ventral. Spermiducal pore 
between voit and via dorsal to tubercula pubertatis. Sperm-sacs in wi and wii minutely 
lobulate. Spermathece one pair in each of somites vit, viii, ix, ventral. No prostates. 
Anterior nephridium in ait. Septal glands inv and vi. No subpharyngeal integu- 
mental gland. Blood capilaries on nephridia are numerous. Continuous blood sinus in 
the sacculated intestine. Hearts in viii, ix, x, xi, not filling the cwlom. Color pinkish 
with violet iridescence. Habitat, England, Thames. Description is compiled after 
Benham’s paper only, as I have not seen any specimens. 


Sparganophilus Eiseni Smith. 


Derinition. Length 18 to 20 cm.; width 26 cm. Clitellum dorsally 1-4 xv- 
wav, ventrally 1-2 wiv—vavi. Tubercula pubertatis 1-2 vvii-1-2 wait, ventral spermathecal 
pores. Sete 3 and 4 dorsal. Spermiducal pore in anterior part of xix, dorsal to tubercula 
pubertatis. Sperm-sacs minutely lobulate. Spermathece, one pair each in vir, viii and 
ix, all dorsal. Prostates 4 pairs in «xxiii—avi; muscular duct present. Anterior 
nephridia commence in wit. Septal glands in iv, v, about the same size, while the one in vi 
is very small. A subpharyngeal parietal pair of glands present, opening in front of 
sete 1 and 2 in somite iii. Blood capillaries on nephridia are many. Continuous blood 
sinus in sacculated intestine present. Hearts in viii, ix, x, vi small. Color violet flesh. 
Habitat, Illinois, North America. 

Prof. Frank Smith has kindly sent me for comparison a full set of sections of 
this species, also several mature specimens in alcohol and formalin. As, however, 
Prof. Smith writes me that he intends to further work out the details of the anatomy 
and histology of this species, I haye not considered myself at liberty to more than 


154 CALIFORNIA ACADEMY OF SCIENCES. 


touch upon a few essential points of importance in further characterizing this species 
from its other American allies. The species is nearest related to Sp. Benhami and its 
subspecies. The main points of difference and similarity may be found in the com- 
parative table of the species of Sparganophilus. Sp. Benkami is a much more slender, 
form than Sp. Hiseni, especially as regards the body posterior to clitellum. The ven- 
tral side of the genital regions in the two species differ. Thus in Sp. Benhami the 
ventral depressed area between the tubercula pubertatis and the oviducal pores is 
circular or oval, while in Sp. Hiseni it is contracted in the middle, the shape of a sand 
time-glass. The presence of a small parietal gland in somite ii in Sp. Hiseni 
distinguishes this species from all others known so far, This gland is paired, extends 
considerably into the ccelomie cavity, is racemose and opens to the exterior in front 
sete 1 and 2, and projects its free apex back through septum into iv. Another 
character is the small size of the septal glands in vi, the corresponding pair in 
Sp. Benhami being larger. 
Sparganophilus Smithi n. sp. 

Dermition. Length 20 cm. by 3 1-2 mm. wide; number of somites 185. Dor- 
sal pores, the most anterior one between i/ii, the most posterior one between xu/xii. Ch- 
tellum dorsally 1-2 xvi-veviii, ventrally via—wav. Tubercula pubertatis xiz—xaxvit, dorsal 
to spermiducal pores. Sete 3 and 4 dorsal. Spermiducal pore between xviti/xix, ventral 
of tubercula pubertatis. Sperm-sacs in xi and wii. Spermathece 4 pairs in each of 
somites vii, viii, ix, and 1 pair in somite vi, slender, dorsal and lateral. Prostates 3 pairs 
in wati, wviii/aviv. No muscular duct. Anterior nephridium xvi. Septal glands in 
iv, v, vi and vii about equal insize. No subpharyngeal parietal gland. Blood capillaries 
on nephridia numerous. Blood sinus in sacculated intestine not continuous. Hearts in 
viii, ix, 2, vi, small. Color violet flesh, somewhat iridescent. Habitat, San Francisco, 
California. 

Sparganophilus sonome n. subsp. 

Derinition. Length 20 cm. by 3 1-2 mm. wide; number of somites about 200. 
Dorsal pores. Clitellum dorsally xvii-waviii, ventrally xvi-waix. Tubercula pubertatis 
viv—xaviii, dorsal to spermiducal pore. Sete 3 and 4 dorsal. Spermiducal pores 
between aviii/wix, ventral of tubercula pubertatis. Sperm-sacs in ai and xu.  Sperma- 
thece, two pairs in each of vii, viti, ia; one pair in vi, not slender. Prostates 3 pairs 
in weiti, xxiv, wav, no muscular duct. Anterior nephridium in «vi. Septal glands in 
iv, 0, vi, vii, about equal in size. No subpharyngeal parietal gland. Blood capillaries 
on nephridia numerous. Blood sinus in sacculated intestine not continuous. Hearts 
in viii, iz, xv, vi, small. Color violet flesh, less iridescent than Sp. Smithi. Habitat, 
Sebastopol, Sonoma County, California. 


Sparganophilus Benhami n. sp. 


Derrition. Length 17 em. by 2 1-2 mm. in the posterior part of the clitellum; 
namber of somites about 250. No dorsal pores. Clitellum, dorsally xv—xxiw; ventrally 
avii-axvi. Tubercula pubertatis xviti-xaii, ventral to spermiducal pores, Sete 3 and 4 


 Spermathece ... 


i. 


Sperm-sacs . 
a4. 


Beene peters da | 


: | Anterior nephridia 


commence CA ope 


es Septal glands in 


- 


Parietal celomic 
<o gland in iti... . 


‘he ‘ RG 


Seer sinus in the 
sacculated intestine... 


7 


 Hearts........ see 


Table B. 


COMPARATIVE 


CHART OF 


SPECIES OF SPAR 


GONOPHILUS. 


To face page 154, 


) (01 


Length and width... +++» 


Sete 3 and 4 
Clitellum....seseeeeceees 


Tubercula pubertatis. .... 


Spermiducal pore......-- 


Spermathece 


Sperm-sac8.......+++ +++: 


Prostates 


Anterior nephridia 
commence in.... 


Septal glands in 


Parietal celomic 
gland in iii... . 


Blood sinus in the 
sacculated intestine... . 


Capillaries on the ne- 
phridia and in the 
body-wall . 


tamesis Benham. 


England, Thames. 


7 to 10 cm. 
Ventral. 
dxy-}xxv. 


xvii-xxii, ventral to 
spermiducal pore. 


Between xviii/xix, dor- 
sal to tubercula puber- 
tatis; so are the sperm- 
duets. 

Slender; one pairin each 
of somites vii, viii, ix; 
ventral in line with 
3 and 4. 


In xi and xii, minutely 
lobulate. 


Not present. 


xiii. 


iv, v, vi; the posterior 
are the largest. 


Absent. 


Continuous. 


viii, ix, x, xi; not filling 
the ccelomic cavity. 


Very numerous. 


+ 


Hiseni Smith. 
Illinois River, U.S. A. 


18 to 20 cm. by 2 mm. 
Dorsal. 


Dorsally {xv-xxv; 
ventrally 4xiv—xxvi. 


4xvii-}xxii, ventral to 
the spermiducal pore 
and to the spermducts. 


On anterior part of xix, 
dorsal to the tubercula 
pubertatis; so are the 
spermducts. 


One pair in each of so- 
mites vii, viii, ix, dor- 
sal in line with 3 and 
4; slender. 


In xi and xii, minutely 
lobulate. 


Four pairs in xxiii-xxvi. 


xiii. 


iv, v, vi. Those in iv 
and vy about equal size; 
the one in yl is very 
diminutive. 


One pair of glands open 
in front of ventral sets 
in ili; distal end ex- 
tending into iy. 

Continuous. 

In viii, ix, x, xi not fill- 
ing the ecelom. 


Very numerous. 


Benhami n. sp. 


Tepic, Mexico; 4,000. 


17 em. by 24 mm. 
Dorsal. 


Dorsally xy-4xxv; 
ventrally xvii-xxvi. 


xvili-xxii, ventral to 
spermiducal pore and 
to spermduct. 


On anterior part of xix, 
dorsal to tubercula pu- 
bertatis. 


Robust, trowel-like, with 
wavy outline; one pair 
in each of somites vii, 
viii, ix; dorsal. 


xi, xii, less minutely 
lobulate. 


Four pairs, one each in 
xxili-xxvi; along mus- 
cular basal duct. 


xii. The first two pairs 
smaller; the first large 
pair in xiv. 


iv, v, vi. About equal 
in size. 

None. 

Continuous. 


viii, ix, x, xi, not filling 
the celom. 


Very few. 


| 


| guatemalensis n. subsp. 


| Guatemala. 


10 cm. by 2 mm. 
Dorsal. 


XVi-XXVi. 


Parallel ridges, xviii-xxii}.... 


Robust, trowel-shaped, 
with smooth outline, 
in vii, viii, ix. 


Four pairs in xxiy, xxv, 
XXvi, xxvii. 


None. 


viii, ix, x, xi, not filling 
the celom. 


carneus X. subsp. 


Mississippi River, Iowa. 


9 em, by 2mm. 
Dorsal. 


Dorsally xv-xxiv; 
ventrally ? 


Dorsal, one pairin each 
of somites vii, viii, ix. 
Apical end globular, 
not trowel-like. 


xi and xii, lobulate, but 
not minutely so, as in 
Sp. Eiseni. 


Present; four pairs in 


Xxlil, xxiv and prob- 
ably in xxv and xxvi. 


iv, v, vi; the one in yi 
the smallest. 


None. 


viii, ix, x, xi, completely 
filling the celom. 


Numerous and large; on 
the body-wall very nu- 
merous in the six an- 


terior somites. 


Smithi n. sp. 


Laguna Puerca, San 
Francisco, California, 


20 cm. by 3} mm.(or less) 
Dorsal. 


Dorsally 4xvi- xxviii; 
ventrally xix-xxy. 


xix-xxvii, dorsal 
spermiducal pore. 


to 


Between xviii/xix, ven- 
tral to tubereula pu- 
bertatis. 


Slender; four pairs (8) 
in each of somites vii, 
vili, ix, dorsal and lat- 
eral; one pair only in 
vi dorsal. 


xi and xii, more lobulate 
than in Benhami, but 
less so than in Biseni. 

Three pairs, one each in 
somites xii, xiii, xiv; 
no muscular duct. 

xvi. 


In iv, v, vi, vii; about 


equal in size. 


None. 


Not continuous. 


viii, ix, x, xi, quite small, | 


not filling the celom. 


sonome n. subsp. 


Sonoma Valley, California 


25 to 30 em. by 3} mm. 
Dorsal. 


Dorsally xvii-xxviii; 
ventrally xvi-xxix. 

xix~xvili, dorsal to 
spermiducal pore and 
to spermduct. 

Between xviii/xix, ven- 


tral to tubereula puber- 
tatis. 


One pair in vi; two pairs 
(4) in each of somites 
Vli, vili, ix; not slender. 


xi and xii, more lobulate 
than Benhami, but less 
than Eiseni. 

Three pairs in xxii, xiii, 
XIv. 


Xvi. 


iv, V, Vi, vii, about equal 
in size. 


| None. 


Not continuous. 


Viii, ix, x, xi, quite small, 
not filling the ccelom. 


Numerous, giving the 
nephridia a pink color. 


Very numerous. 
| 


PACIFIC COAST OLIGOCH#ETA. 155 
are dorsal. Spermiducal pore on anterior part of xia. Sperm-sacs in xi and wii, not 
minutely lobulate. Spermathece one pair in each of vii, viii and ix are dorsal. Sperm- 
iducal glands or prostates 4 paws in waiti-vaxvi, with a long muscular duct. Most 
anterior nephridium in wii. Septal glands in iv, v, vi of about equal size. A pair of 
subpharyngeal parietal glands not present. Blood capillaries on nephridia few. A 
continuous blood sinus in sacculated intestine. Blood glands many. Hearts in viii, ix, 
x, ai. Color violet, strongly iridescent. Habitat, Tepic, Mexico, 4000 feet. 


Sparganophilus guatemalensis n. subsp. 


Derinition. Length 10 em. by 1 1-2 mm. wide; number of somites 260.  Dor- 
sal pores none. Clitellum xvi-vxn. Tubercula pubertatis xviii-avxii, parallel ridges. 
Sete 3 and 4 dorsal. Spermiducal pores unknown. Prostates four pairs in wxiv, wav, 
wxvi, veou. Spermathece one pair each in vit, viii, ix; wide free end, wide, flat, outline 
smooth. Hearts very strong. Color deep bluish violet ividescence. Habitat, Guatemala 
City. 

Sparganophilus carneus n. subsp. 

Derrinition. Length 9 em. by 2 mm. wide; number of somites 160. Clitellum 
dorsally 1-2xy—vxv. Tubercula pubertatis unknown, Sete 3 and 4 dorsal. Spermidu- 
cal pores unknown. Sperm-sacs in xvi and «xii, lobulate, but less minutely than in Sp. 
Hiseni. Spermathece one pair each in vii, viii, ix dorsal; free end globular. Prostates 
unknown. Septal glands in iv, v, vi, the one in vi the smallest. No subpharyngeal 
integumental glands. Blood capillaries enormously large and many in the anterior six 
somites. Hearts very large in viii, ix, v, wi. Color reddish flesh, much darker when in 
alcohol than Sp. Benhami. Habitat, Mississippi River, near Clayton, Iowa. 

After these preliminary definitions I will now describe in detail the various 
species of Sparganophilus examined by me. 


- 


Sparganophilus Smithi n. sp. 
Figs 120-122, 124, 129-39. 


Habitat. This species is very abundant in a small lake or pond known as 
Laguna Puerca and situated between Lake Merced and Golden Gate Park at San 
Francisco. Adult form in June to October, more frequent in the latter month. It 
occurs at or near, above and below the water’s edge, the earth being thrown up in 
abundance and in heaps. I have already referred to nonpresence of this species 
some ten years ago in the above locality. 


EXTERIOR CHARACTERS. 


Color. A brownish flesh with violet reflex, but much less so than Sp. Benhama 
or Sp. guatemalensis. In alcohol this species becomes much paler than either of the 
two just referred to, almost pure white, looses its irridescence entirely. In formalin 
the color is pale, the irridescence very faint, strongest on the clitellum, and hardly 
perceptible on any other part of the body. 


Length varies some, but generally reaches in fully grown and largest speci- 
Memorrs, Vou. II, 5. January 6, 1896. 


156 CALIFORNIA ACADEMY OF SCIENCES. 


mens 20 cm., while the width is about 35 mm. posterior to clitellum. This and the 
following species are the longest forms so far known of the genus. The worms are 
not remarkably lively when out of the ground. 

Sew. The couple 3 and 4 are distinctly dorsal, and it may be of interest to 
note that in all species found in America the dorsal portion of these sete appear 
characteristic, while in Sp. tamesis the setee 3 and 4 are ventral or sublateral. I can 
find no character in the form of the sete in the various species. They are all slightly 
sigmoid, slightly hooked and devoid of ornamentation. There are no penial sete and 
no modified sete in any part of the genital region. The sete of Sp. tamesis as figured 
by Benham are more hooked than those of my new forms. : 

Prostomium and pygidium. I cannot find any good characters in the form of 
these part in the respective species. The small pit in the prostomium of Sp. tamesis, 
described by Benham, is found in all the species. The pygidium varies greatly. 
While some specimens have the anal orifice elevated and dorsal others have it central. 
The shape and size of the last somite varies also to such an extent that no species 
character can be derived from it. I will then in the following not refer to these part 
in any of the species described below. 

Clitellum. In all the species of this genus the clitellum is very large, oecupy- 
ing from 8 to 15 somites. In Sp. Smithi it is located as follows: dorsally, $xvi-$xxviil; 
ventrally, xix—xxy. Continuous all around the body. In all preserved specimens 
the body is bent towards the ventral side just at the clitellum, and special pains must 
be taken if a straight specimen is desired. 

Tubercula pubertatis consists of a very elevated, continuous ridge, which is 
broken or depressed at the intersegmental grooves. It extends through somites 
xix—xxvil and is situated dorsally to the spermiducal duct and its pore. The ridge is 
further generally concave on the dorsal and convex on the ventral side. The ventral 
area between the two tubercula pubertatis ridges is thus much wider than in any of 
the other three new species described here. Anteriorly the tubercula pubertatis 
ridge is continued forward in a kind of semicircle which ends at the groove between 
somites ix/x. But these anterior ridges are much lower than the tubercula pubertatis 
proper, but nevertheless very sharply defined and distinct. 

Spermiducal pore is situated between xix/xx, laterally to the sete 1 and 2 and 
about three times as far from 2 as 2is from 1. But the pore is ventral to the tuber- 
cula pubertatis ridge. This is an important characteristic and only shared with Sp. 
sonore. In Sp. tamesis, Kiseni and Benhami, the spermiducal pores are dorsal to the 
tubercula pubertatis ridge. 

Oviducal pores are plain in front of setee 1 and 2 on somite xiy. 

Prostate pores externally not visible, but situated in front of sete 1 and 2. 

Somites. All the anterior somites are three-ringed, except i, ii, 11, which are 
smooth. 

Dorsal pores. The most anterior pore is situated between i/1i, and the most 
posterior one between xii/xiil. 


PACIFIC COAST OLIGOCHETA. 15 


~I 


INTERIOR CHARACTERS. 

Septa. Thickest septum is found between somites xi/xii. Anterior septa are 
gradually diminishing in thickness forward, and similarly the septa posterior to xi/xii 
are diminishing in thickness backwards. The most posterior thickened septum, the 
one between xiv/xy, is hardly thicker than the one next posterior. The central 
part of the thickened septa is very much thicker than the part near the body-wall. 
(fig. 127¢.m.) The septa correspond more to the intersegmental grooves than do those 
in Sp. Benhami. When the body is viewed in cross-section it will be seen that the 

‘septa are principally attached to the body-wall at four points nearly half way between 
the sete (fig. 131). 

Suprapharyngeal and septal glands. There are dorsal septal glands in iv, v, 
vi, vii, about equal in size, and ventral septal glands in y, vi, vil, also of about equal 
size. The latter are much larger than in Sparganophilus Benhami, and reach above 
the center of the body (fig. 120). In addition to these glands I find an accessory 
septal gland attached to the central anterior surface of septum ix/x, just above the 
intestine and extending downwards. ‘This gland is closely attached at every point to 
the septum, and projects only slightly into the ecelomic cavity. In height this gland is 
not much thicker than the central part of the septum (fig. 152). Its free surface is very 
smooth and even. I could follow ducts with precipitated secretions running along the 
septum downwards towards the intestine, but its connection with the latter, if any, I 
could not ascertain. Beddard suggests that as the nephridia only commence in the 
thirteenth segments the septal glands described by Benham in Sp. tamesis are homo- 
logous with the nephridia. In all species of Sparganophilus investigated by me I find 
no such mucous glands as in Pontoscolex, only regular salivary glands, which open 
into the pharynx, and of the same structure as the supra-pharyngeal glands generally. 
These salivary glands in Sp. tamesis occur exactly in the same somites as in all other 
Sparganophilus species, viz.: iv, v and vi. 

Body-wall offers no other characteristics, except that it is throughout of almost 
the same thickness, and not thinner along the dorsal parts of some of the central 
somites, as in Sp. Benhami. 

Clitellum is continuous all around the body, but it is much thicker dorsally 
than ventrally. This thicker dorsal portion commences with the tubercula pubertatis, 
gradually increasing in thickness dorsally towards the median line, where it is four 
times thicker than at the ventral median line between the tubercula pubertatis. This 
refers only to the clitellar cells in the central clitellar somites. The long clitellar 
cells are confined to the latero-dorsal part, while the ventral part contains only 
the short clitellar cells, the point where the former cease being the tubercula 
pubertatis. This is the case in all the species examined by me. 

Tubercula pubertatis. It has already been stated that they form a continuous 
ridge, only broken in the anterior two somites xvii and xviii. Their structure differs 
considerably from that of Sparganophilus Benhami. We find the tubercle cells less 
differentiated (fig. 138), and more resembling the short clitellar cells, of which they 
are only a variety. Another characteristic is the presence of a large blood vessel in 


158 CALIFORNIA ACADEMY OF SCIENCES. 


the center of the tubercle. The most interesting feature, however, consists in the 
presence of three or four arciform muscular bands which connect the opposite sides 
of the tubercular projection, and which, of course, serve to further push out the tip 
of the tubercle, or to relax it, as the occasion may demand. These bands are entirely 
confined to the clitellar cell layer, and penetrate to the very cuticle, to which at least 
some of the muscles are attached, while others seem attached to the cells themselves. 
Besides these epidermal arciform muscles, there are numerous coelomic arciform 
muscles, which also penetrate the glandular layers and serve to connect the two sides 
of the clitellum on either side of the tubercula pubertatis (fig. 158). While the 
arciform muscles of the epidermal layer have been described by Cerfontaine in Lum- 
bricus, they are by no means known from many species. Their presence varies con- 
siderably in different species of Sparganophilus, but they are especially numerous 
and strong in this one. 

(Hsophagus and intestine. As regards the general shape of the alimentary canal 
I can see no marked difference in the various species so far examined. Again as to 
structure I find two points worthy of mention. The chloragogen cells which. are 
everywhere covering the intestine are much larger and more numerous in this species 
than in Sp. Benhami, covering as they do both tubular and sacculated intestine. 
Another point is the absence of a continuous blood-sinus. A continuous blood-sinus 
in the sacculated intestine has been described by Benham in Sp. tamesis, by Smith in 
Sp. Hiseni and is also found in Sp. Benhami, as will be recorded further on. In this 
species, Sp. Smithi, the sinus is not quite continuous, the respective blood lacunes are 
quite close and in places run together, but they are nowhere continuous in the same 
way as in the species referred to above. The sacculated intestine commences in xiii. 

Spermathece (fig. 130). The most characteristic feature of Sparganophilus 
Smithi is the occurrence of numerous spermathece in at least three somites, while in 
one somite there is found only one pair. Sevenadult specimens from Laguna Puerca 
were opened and agreed in the following arrangement and number of spermathece: 

Somite vi: 2 spermathec, in front of or slightly dorsal to setee 3 and 4. 

Somites vii, vili, ix: 8 spermatheee in each, in front of, slightly dorsal and 
lateral of setee 5 and 4. 

The location of these spermathec:e is not strictly constant asin some specimens, 
as well as insome somites of the same specimen, the spermathece were shoved a little 
dorsally or ventrally of setee 38 and 4. One specimen possessed three spermathece 
on one side and four on the other side in the same somite, but all the other specimens 
possessed eight spermathecee in each of the somites, except in vi where inyariably 
only one pair was found. 

These spermathece are very large (fig. 125), tall and slender, and viewed 
in cross-sections of their body they are seen to extend from one end of the ccelom to 
the opposite side, touching both body-walls. They are generally directed forward 
and crowding each other; they occupy nearly all of the available room in the, somite. 
I believe there is no other case known in oligocheta where so many and so large 
spermathece are known to occupy the same somite. The size of the respective sperm- 


PACIFIC COAST OLIGOCHATA, 159 


athec varied some, but the general shape appeared quite constant within certain 
limits. The figures appended represent the spermathece taken from one specimen. 
There is a narrower muscular basal tube and a more swollen non-muscular chamber 
as usual, the structure of which offers no characteristics. 

Sperm-sacs. Of the two pairs of sperm-sacs the anterior one in somite xi is 
much the narrowest in the direction of the median longitudinal diameter of the body. 
It is principally ventral in location, but extends upwards and joins the one from the 
opposite site. It is closely surrounded by a sperm reservoir without enclosing mem- 
brane. The sperm-sacs in xii are much broader and longer, but are generally only 
dorsal and do not enclose the intestine in any of the specimens sectioned. This rela- 
tive size of the two pairs of sperm-sacs appeared constant. Somites ix and x are 
transformed into two very extended sperm reservoirs. 

Ovaries in the various species are long and flat, not lobed, and in both longitu- 
dinal and transverse section offer to view a single undivided surface. 

Oviducts are very broad and furnished each with an ovisae consisting of an in- 
vagination of the septum, which in all my specimens were filled with large gregarinz, 
rarely containing any ova. 

Ciliated rosettes are similar to those of Sp..Benhamiin size. The spermducts run 
ventrally to as the tubercula pubertatis, about four times as far laterally and dorsally 
from seta 2, as 2 is distant from seta 1. The spermiducal pore is similarly situated 
ventrally to the tubercula pubertatis, This characteristic is in all the species only 
shared by Sparganophilus sonome. Allother species are characterized by having the 
spermducts run dorsally as to the tubercula pubertatis. The spermiducal pore is simi- 
larly in Sp. Smithi situated ventrally to the tubercula pubertatis, and is found in the 
intersegmental groove between somites xix/xx. 

Prostates or spermiducal glands (Beddard). There are three pairs of spermi- 
ducal glands, one pair each in somites xxii, xxili, xxiv. One specimen possessed only 
one gland in xxiv, and there may possibly be found some variation in number when 
more specimens have been investigated. In all cases these glands were much smaller 
than in Sparganophilus Benhami, but otherwise almost similar in structure, except for 
the entire absence of the basal muscular duct. The glandular part was never large 
enough to be folded on itself lengthwise, but its tube was much twisted in the direction 
of its long diameter, often to such extent that cross-sections always showed the tube 
as three or four circular openings, surrounded by a wall two cells thick. The muscu- 
lar basal part is absent, and the lower tube, where it enters the muscular layers, being 
very short, is surrounded by a thin layer of connective tissue. The whole organ in 
mature specimens projects only slightly above the body-wall, and the glandular part 
is in either diameter not any thicker than the body-wall. 

Nephridia are in this species covered and perforated by numerous blood 
capillaries, to a much greater number than in Sparganophilus Benhami, but similar to 
what is described in Sp. tamesis and Sp. Hisent. Otherwise the nephridia of the re- 
spective species appear to be of the same general size and structure. 

Vascular system. The hearts begin in xi, and extend forwards to vill. In xii 


160 CALIFORNIA ACADEMY OF SCIENCES. 


and xiii there are no connecting vessels. In xiv to xvii we find the dorsal vessel en- 
larged and folded on itself in a zigzag manner. Both hearts and dorsal vessels are 
thickly covered with chloragogen cells. I have already pointed out the absence of 
a continuous blood sinus in any of the various parts of the intestine. All the various 
organs of the body are thickly covered with blood capillaries, especially so the neph- 
ridia, spermathece, the clitellum, in which strong capillaries separate every two or 
three rows of the large clitellar cells. In the center of the tubercula pubertatis is 
always seen a dense mass of larger and smaller capillaries. 

A characteristic of the vascular system is also the great scarcity of blood 
glands, these being very numerous in Sp. Benhami. 


Sparganophilus sonome n. subsp. 
Figs. 123, 126. 


Habitat. Creeks and springs around Sebastopol, Sonoma County, California. 
Adult specimens very numerous in April. 

General Remarks. 1 consider Sparganophilus sonomew as probably a sub- 
species under Sp. Smithi, which it resembles in most points, two of which are of 
the greatest importance, viz.: The position of the spermduct and spermiducal pore, 
and the duplication of the spermathec: in several somites. But as long as I found 
the number of spermathece constant, and some other minor points of difference, I 
thought it best to describe this form more carefully, leaving the question of species or 
subspecies as a matter of choice, and for future consideration and study. 


EXTERNAL CHARACTERS. 


Tubercula pubertatis are the only external organs which differ from Sp. Smithi. 
While in Sp. Smithithey form a continuous ridge on either side, they are in Sp. sonoma 
broken up in numerous tubercles, generally two or three on each somite (fig. 123). 
I believe there is another characteristic worthy of mentioning. The single puberty 
groove is in the center of the tubercle, while in Sp. Smithi there are two grooves, one 
at the base on either side of the tubercle. But I freely confess that I have not sec- 
tioned up a sufficient number of specimens in order to know if these points are really 
constant, or if not rather they are subject to considerable variation. My supply of 
mature specimens of Sp. sonome was very limited, but in all I found the tubercula 
pubertatis broken into a succession of little knobs, but all together forming a ridge 
in outline like that of Sp. Smithi. 


INTERIOR CHARACTERS, 


Spermathece. While in all the specimens of Sp. Smithi I found eight sperma- 
thece in each of somites vii, viii, ix, the subspecies possess them as follows: Somite 
vi: one pair with the pore in front of seta 4. Somites vii, vill, ix: two pairs (4) in 
each, with pores in front of sete 4, and between 3 and 4. Each pair consists of two 
separate spermathece, one of which opens in the intersegmental groove in front of 
seta 4, while the other opens similarly in front of a line drawn 4 the distance between 
sete 3 and 4, the 3 being towards 4 and the 3 towards 3. Both pairs are, therefore, 


PACIFIC COAST OLIGOCHXETA,. 161 


dorsal. The direction of the spermathece is in the diameter of head to tail. They 
lie closely pressed to the body-wall. As regards size, the most anterior pair is the 
smallest, and the most posterior the largest. 


Sparganophilus Benhami np. sp. 
Figs. 97-119. 

Habitat.—In the mud of springs, October, November, 1894. Tepic City, Ter- 
ritory of Tepic, Mexico, 4000 feet altitude. Of some fifty specimens collected only 
three were well developed as regards clitellum and tubercula pubertatis. 

EXTERIOR CHARACTERS. 

Color, reddish-yiolet, very strongly iridescent. The violet tint is very pro- 
nounced, much more so than in any other species that I have examined except Sp. 
guatemalensis, where the bluish-violet tint is very deep and intense. 

Body. This isa long and slender species tapering towards the tail end. The 
cephalic lobe is superiorly not distinct from somite i, but there is a small pit on the 
boundary between the two. Somites i, i, iii are less prominent. Somites iv to ix are 
strongly convex, but x to xv are much less so. The clitellar intersegmental grooves 
are only distinct between the ventral sete. 

Dorsal pores. The most anterior dorsal pore is seen between i/ii; the most 
posterior one between v/vi. There is a central pit in the dorsal side of somites 1, i, iil. 

Clitellum, as in other species; the extent of the clitellum is different on the 
dorsal and ventral sides. This I think can best be expressed thus: dorsally, $xiv— 
xxiv; ventrally, xvii—xxvi. The elitellum is continuous, but much less developed 
ventrally between the tubercula pubertatis than dorsally. This is also characteristic 
of the other species I have examined. Viewing the clitellum exteriorly, it appears 
as discontinued between the tubercula pubertatis ridges. 

Tubercula pubertatis begin in xviii and end in xxii. Each one consists of a 
straight ridge on the top of which runs a groove parallel to the ridge. Posterior to 
the tubercula pubertatis proper the ventral part of the clitellum is concave deepening 
posteriorly and ending in a deeper pit in the posterior part of xxvi. Anteriorly the 
tubercula pubertatis ridges continue forwards almost straight and outside of the ventral 
sete 1 and 2 to the center of xii, where they end in the region of the ventral sete. These 
two almost parallel ridges are very thin and sharp, several times narrower than the 
tubercula pubertatis proper. They only appear in fully adult specimens, and are very 
characteristic. I believe that the underside of the clitellum will furnish good species 
characteristics if carefully noted. But in order to bring out this region properly the 
specimens must be slowly killed and then as rapidly as possible be passed through the 
alcohols into that of 967%. Formalin specimens do not show the region as well as 
alcoholic specimens. 

Male pore or spermiducal pore is situated on the anterior + of xx, just lateral 
to the tubercula pubertatis. The pore is not prominent, but may be readily seen. 

Prostate pores are not prominent, but in adult specimens they may be seen in 
front of sete 1 and 2 in somites xxili, xxiv, xxy, Xvi. 


162 CALIFORNIA ACADEMY OF SCIENCES. 


Ovipores are separate between somites xili/xiy, or slightly on the anterior part 
of xiv. 
Nephropores prominent in front of setze 1 and 2 in all somites posterior to xii. 


INTERIOR CHARACTERS. 


Body-wall. In somites x to xiy the body-wall is much thinner on the dorsal 
side of the body than on the ventral side. As Benham does not mention this peculi- 
arity in Sp. tamesis, and as I have not observed it in Sp. Smithi, I take it for granted 
that this character is peculiar to this species. The clitellum is complete, but much 
thinner on the ventral side between the sete, as shown by figure 119, which repre- 
sents the ventral body-wall in somite xix. It will be seen that the longitudinal muscular 
layer is several times thicker than the transverse layer. The strands are separated 
by irregular layers of granulated tissue of various thicknesses. The circular layer 
again consists of only one row of fascicles, each fascicle being tubular, and inclosing 
numerous irregularly distributed muscular fibers, strongly striate. When viewed in 
cross-section it becomes apparent that the strands of the longitudinal muscles are of 
varying thickness, the thicker ones being situated nearest the ccelomic cavity, from 
there gradually decrease in size towards the circular muscles. 

Clitellar cells. The clitellar cells are especially interesting on account of their 
relationship to the cells of the tubercula pubertatis, which will be described later on. 
Figs. 107, 108 represent a longitudinal section of the body-wall in somites xxiii and xxiv. 
The muscular layers are much thinner, the clitellar cells are only developed in the 
vicinity of the prostates. Below the clitellar cells are seen long tubular cells, some- 
what like those of the tubercula pubertatis, and between them are seen, now and then 
(fig. 107 d.ch.), the deeply staining discharge chambers of the clitellar cells. The uni- 
cellular glands, which are here very irregular, are entirely confined to the zone 
surrounding the intersegmental grooves. In fig. 108 is seen a more magnified part 
of the region close to the prostate. Only one unicellular gland, though of unusual 
size, is seen. It isa stray one, as theyare rare in this particular zone. The structure 
of the clitellum of Sparganophilus has been well described by Benham in his paper 
on Sparganophilus tamesis, and I can only add a few points of interest. For studying 
the clitellar and tubercula pubertatis cells I have used principally the iron-lack 
Heidenhain process, but by adding a light tint of Ehrlich-Biondi to the ammonio- 
ferric-alum solution the most perfect differentiation may be had of the various clitellar 
elements. The clitellam in Sparganophilus contains the following various varieties 
of cells: 

1. Regular epidermal cells, secreting the cuticle, furnished with oval nuclei. 
Even on the fully developed clitellum of Sparganophilus Benhami they occur all 
around, both on dorsal and ventral sides. Their inner ends are drawn out into one or 
more threads, some of which extend to the innermost margin of the epidermis 
(fig. 103, 1). 

2. Unicellular goblet glands, entirely confined within the outer layer of the 
foregoing cells. They are very irregular as to size. These cells are the only ele- 


PACIFIC COAST OLIGOCH ETA. 165 


ments in the clitellum which stain deeply with hematoxylin. These cells offer 
nothing characteristic of the species. They are the last to be affected by tropceolum 
and orange, their mucin remaining intensely blue. Their nuclei are compressed and 
placed in the posterior part of the cell in the triangular mass and extremity, staining 
black in the iron-lack (fig. 103, 2). 

3. Short club cells. These are of very uniform size, their free ends forming a 
marked line all around the clitellum interior to the goblet cells. The free end of 
this cell is club-shaped and generally regular in outline. The heights of the indivi- 
dual cells are quite even, and, as these cells stain darker than the following kind, 
they form a marked contrast to all others. The granulation is coarse and regular, but 
does not stain readily with hematoxylin. It takes, however, deeply the red aniline 
stains. On the dorsal side of the clitellum these cells reach inward about 4 of the 
whole width of the epithelium, while on the ventral side, they constitute the only 
so-called clitellar cells. The nuclei of these cells are oval and always prominent 
(fig. 103, 3). 

4. The pre-eminently clitellar cells consist of several, 3 or 4, different lengths. 
This class of cells are in many genera, such as Lumbricus, ete., massed in columns, 
often very regular, and separated by septa. In Sparganophilus these cells are not 
arranged in columns, but distributed almost regularly all through the epidermis, of 
which they occupy about % of the whole mass. At certain intervals, however, may 
be seen a. thin triangular septa of connective tissue with nuclei, projecting down- 
wards or outwards from the muscular layers, dividing this part of the epidermis in 
numerous irregular pyramidal masses, which latter are homologous with the columns 
in Lumbricus, ete. The nuclei of these clitellar cells are shrivelled up and situated far 
back in the triangular apex where the cellular plasma is especially agglomerated. 
There is no such granular secretion in this class as in the former, and they stain much 
less intensely than No. 3. These cells occur only dorsally to the tubercula pubertatis, 
and are entirely absent from the ventral side between the tubercula pubertatis. 

5. Here and there we find intermediate varieties of oval and club-shaped 
cells longer and wider than the epidermal supporting cells and furnished with large 
round nuclei. 

6. Connective tissue cells with large round nuclei, especially prominent 
between the circular muscular layer and the tall clitellar cells. At intervals this 
tissue forms triangular projections outwards, separating bunches of cells four or five 
wide and of different sizes. This tissue is generally accompanied by numerous 
capillaries. 

Tubercula pubertatis. These organs are very prominent, and situated on a 
nearly continuous ridge occupying somites xviii-xxii. Benham has described this 
structure in detail in his species Sp. tamesis, and called attention to the fact that it is 
composed of long narrow cells, which are modified clitellar cells. In Sparganophilus 
Benhami these cells are of two kinds, as far as regards the form. 

a Some which are very narrow, longer than the others, and narrowest at the 


free end. All of this class are grouped in such a way that they radiate outwards 
Memoirs, Vou. II, 5. January 7, 1896. 


164 CALIFORNIA ACADEMY OF SCIENCES. 


towards and around a small concave pit or groove in the outer edge of the epidermis 
(fig. 105). 

}. Interior to these taller cells are shorter and thicker cells, indicated asd in the 
above figure. 

Exterior to these, the regular and characteristic tubercula pubertatis cells 
proper are seen numerous unicellular glands, the same as the goblet cells of the 
epidermis. 

In addition to these glandular cells, I find at certain intervals a few sense cells 
opening out into the narrow pit or groove just referred to. They are everywhere very 
few in number, tall, narrow, with now and then the tip projecting through the cuticle 
into the shallow pit (fig. 105s. ¢./.) There is not a continuous row of these cells, but 
here and there are found bunches of half a dozen cells, opening close together. They 
do not seem strictly parallel, but bulge and diverge in such a way that in sections, 
which show the common cells parallel, rarely more than one single sense cell is in 
view to any great part of its length. This, together with the small number of these 
cells, is undoubtedly the reason why they are frequently overlooked. 

In connection with these sense organ cells, I have re-examined the ventral 
papille found in Argilophilus marmoratus, and I find that these structures are really 
nothing but tubercula pubertatis nature, or at least sense organs furnished with sense 
cells, a description of which will be deferred to a future paper. As a general con- 
clusion, I may state that the tubercula pubertatis are really sense organs, furnished 
with sense cells of the same nature as those found in other parts of the epidermis, and 
described by me in Benhamia, and by Vejdoysky in Rhynchelmis, by Cerfontaine 
and Langdon in Lumbricus, by Hesse in Lumbricus and Allolobophora, ete. 

Septa. The septa are not all of the same thickness, neither is each individual 
septum of the same thickness throughout. The first distinct septum is found between 
iv/v. It is of regulation thickness, or as thick as septum x/xi and those following 
posteriorly. The anterior septa increase in thickness forwards and backwards in such 
a way that septa vi/vii, vii/viii are of about equal thickness, while those in front and 
behind these are thinner in proportion as they are more distant. The anterior five 
septa are much thicker in their central area, and thin out towards the periphery and 
body-wall, but even then, at the thinnest part, they are about four times thicker than 
the posterior septa. The six anterior septa do not strictly correspond with the seg- 
ments, but are attached to the body-wall about one-fifth the distance forward from the 
posterior intersegmental groove. 

Suprapharyngeal and septal glands. The glandular mass superposed on the 
pharynx and opening into it, is prominent, but situated far back, and with its lobes 
pointing forward, or in the same direction as the septal glands. The opposite is gen- 
erally the case, and has been so in all other species examined by me. 

Longitudinal sections show that the regular septal glands are present in three 
somites. The most anterior pair is in iv, immediately behind the suprapharyngeal 
glands, and not separated from the latter by any septum. ‘The other two pairs are in 
v and vi. In longitudinal sections the suprapharyngeal glands are seen to be com- 


PACIFIC COAST OLIGOCH ETA, 165 


posed of two distinct lobes of about equal size, one situated closely posterior to the 
other. This is also the structure of the septal glands in iy and y, but the one in vi is 
solid, consisting of only one lobe, when viewed in longitudinal section. The main 
lobes are situated on the same longitudinal muscular band, with their discharge ducts 
running forwards into the pharyngeal cavity. In each of the three somites there are 
two distinct glands on either side of the median line. One larger supraintestinal 
consisting of three parts extending laterally, and one smaller subintestinal. These 
latter ones also open into the pharynx, but into its ventral side, the pharynx 
thus being partially developed even on the ventral side. The discharge tubes and 
chambers are very large, and the latter occupy more than one-half the width of the 
pharyngeal wall. They stand very close together, and are all of about the same height 
and form (fig. 112). The dorsal wall of the pharynx is much thicker and denser 
than the ventral one, and the discharge chambers stand closer and are of more uni- 
form size, more tubular. The discharge chambers on the ventral side are thicker, 
more pear-shaped and much fewer in number. Fig. 112 represents the lining of 
the dorsal wall of the pharynx; fig. 115 the ventral wall of the same, and fig. 114 
the wall next posterior to the main dorsal section. 

(Hsophagus and intestine. ‘The description given by Benham of the histology 
of the intestine of Sp. tamesis may in the main points be applied to this species. 
The cwsophagus is cylindrical with parallel sides and slightly nipped by the septa. It 
is sparsely covered by chloragogen cells, which latter are more numerous on the 
intestine. he cilix of the inner epithelium are much longer in the cesophagus, 
and so are the epithelial cells themselves. Seen in longitudinal section (fig. 110) 
we find that the transverse muscles surrounding the cesophagus are more numerous and 
present in several rows, while in the intestine they are few and far between, and 
arranged only in one row. There is thus a distant approach to a gizzard in the 
cesophagus. The longitudinal muscular layer is reduced everywhere in the intestine 
to a single strand, but in the cesophogus it is double, sending out strands to the septa 
and to the mesenteric sac. 

Blood sinus. Both Benham and Smith have shown the existence of a conti- 
nuous blood sinus in the intestine of the species described by them. Inthe cesophagus 
of our present species we find only a vascular network and confluent smaller lacunes, 
but in the intestine proper we meet with a continuous blood sinus all around the 
epithelial cells. The radiated appearance of the blood in the sinus, as well as in the 
other vessels, must as Benham suggested be due to the erystalization of hzematine. In 
Sp. Benhami and Sp. Smithi these crystals are so many and so heavy that they invari- 
ably destroyed the edge of the microtome knife and made sectioning most difficult. 
But the erystalization presents some peculiarities, and in places appears as if there 
was a mass of radiating fibers always from the side nearest the center of the body 
radiating towards the periphery, but never the contrary. In Sp. benhami these crystals 
are much more slender than in Sp. Smithi. In the latter species their nature is. not 
to be doubted. They are also found in the hearts, but more rarely in the main longi- 
tudinal vessels. 


166 CALIFORNIA ACADEMY OF SCIENCES. 


In the intestine we find between the inner ends of the epithelial cells a layer 
of connective tissue, strengthening as it were that side of the perienteric sinus (fig. 
111). Both w@sophagus and intestine are enclosed by a thin mesenteric sac, which 
is nipped by the septa (fig. 111, mes.) and pressed close to the chloragogen cells. 

Spermathece. These are much broader than those of Sp. Smithi or Sp. tamesis, 
but resemble more those of Sp. Hiseni and Sp. guatemalensis. Their position in 
somites vii, viil, ix, is in the anterior part of the somite, opening in the intersegmental 
eroove and pointing backwards. Their pores are in line with the dorsal couple of 
sete, or 3and 4. The muscular part is tubular, smaller, and consists of two layers, 
one inner of epithelial cells, one outer thicker, of circular muscles. The inner epi- 
thelial layer, which is a direct continuation of the epidermis of the body-wall, consists 
of very tall, narrow, columnar cells, while the thick muscular coating is a direct con- 
tinuation of the circular muscular layer of the body-wall. The free end of the 
spermatheca is wavy and warty in outline, and consists of much shorter epithelial 
cells, simply covered by the peritoneum. The spermatheca is very broad and very 
flat (fig. 118). 

Sperm-sacs. There are two pairs of lobulate sperm-sacs in xiand xii projecting 
from the anterior septum. They are situated principally dorsally, and resemble those 
of Sp. tamesis, but are much less lobulate than those, and very much less lobulate 
than the sperm-sacs of Sp. Hiseni, judging from sections of the species sent me by 
Prof. Frank Smith for comparison. 

Large masses of free spermatogonia and spermatozoa are seen in front of the 
ciliated rosettes. 

Ciliated rosettes (fig. 109) are large and very regularly folded. Each rosette 
sends out a long tubular lip into the sperm-sacs, in way which I have figured in fig. 
119ce. Ina cross-sectioned specimen I found this lip far back in the posterior part 
of xii. The lower convolute lip of the funnel is almost absolutely regular, and similar 
in each of the four rosettes. The spermducts appear to resemble those of Sp. tamesis 
and Sp. Hiseni. The spermidueal pore is situated just outside of the tubercula puber- 
tatis in the anterior part of xx, just as in Sp. Hiseni. 

Prostates. Smith is the first to describe the prostates in Sparganophilus. In 
Sp. tamesis they appear not to be present, as Benham does not mention them. In Sp. 
Benhami there are four pairs opening in somites xxili, xxiy, xxv and xxvi, in front 
of setee 1 and 2. The prostates are constructed on the same principle as the prostates 
in Acanthodrilide and Cryptodrilide, and consist of two parts; one basal and 
muscular, one apical and glandular. The glandular part is tubular, straight or folded, 
of considerable length, but confined to one somite. The glandular part contains an 
inner epithelium, and surrounding it club-like glandular cells of varying lengths, 
giving to the surface of the prostate a wavy and irregular appearance. The prostates 
in this species differ from those of Sp. Smithi by having a muscular duct or basal 
part. This latter does not exist in Sp. Smithi, the glandular part in the latter species 
being immediately attached to the body-wall. 

Nephridia. There is undoubtedly some difference in the location of the most 


PACIFIC COAST OLIGOCHETA. 167 


anterior nephridia in the various species. In Sp. Benhami the most anterior pair is 
in xii, but they are smaller than those in xiii. In xv we find the first very large 
nephridium covered with a thick coelomic cell mantle. This mantle covers the ducts; 
sometimes we also find a mass of similar cells attached to the nephridium as a rounded, 
nearly separate mass, only connected with the nephridium by means of a very narrow 
part. Owing to the great opaqueness of the nephridia, I have not been able to make 
out the run of all the canals, and can illustrate only a little more than the outline of 
the organ. It appears, however, that some of the canals are doubled. The outlet 
duet is very heavy, wide, and its walls are thick. It runs far up into the windings. 
There is a bridge, and I can distinguish all the various principal parts described by 
me in the nephridium of Pontodrilus. There are comparatively few blood vessels on 
the nephridia, and in this respect the species differs from Sp. Hiseni and Sp. Smithi, 
in which the nephridia are thickly covered and penetrated by capillaries, causing 
them to be of a deep pink color. As regards size, the nephridia are as high as the 
diameter of the celom. The nephropores are in front of setee 1 and 2, and very 
wide. 

Vascular system. Benham’s description of the vascular system of Spargano- 
philus is so complete that I can add but little; the various species seem to agree to a 
very great degree. Sp. Benhami is distinguished by a scarcity of capillaries on the 
various organs of the body, such as clitellum, nephridia, ete., while in Sp. Simithi 
and Sp. Hiseni capillaries are so abundant that they, for instance, almost obscure the 
nephridial surface. ‘The erystalization of the blood has already been described. It 
is found in all the vessels of the body, but especially in the mesenteric blood sinus. 
I have found swimming free in the blood two distinct cell elements, some of which 
are very large-—possibly leucocytes, while others are extremely minute, more round 
and dense—possibly erythrocytes. But as their description requires more time and 
study, I will defer it to a separate paper. 

The walls of the blood vessels present a banded appearance, caused by par- 
allel bands of thicker tissue, furnished with large circular nuclei, arranged in rather 
regular rows. ; 

A characteristic feature of the capillaries of this species are the numerous 
blood glands, similar to those I have described in Pontodrilus and Argilophilus. They 
are especially numerous in the nephridia and in the septal glands. They also contain 
a large number of nuclei. 


Sparganophilus guatemalensis n. subsp. 


Habitat. Guatemala. While in this Central American State several years 
ago I found a great number of specimens pertaining, as I believe, to at least three 
distinct species of Sparganophilus. These, as well as my other oligochzetological col- 
lections made there, were mostly destroyed by accident, few specimens being saved. 
These are now not in good condition for description, and this must account for the 
imperfect data I am able to furnish for all species now described from Guatemala. I 
found Sparganophili in that country in the most varied localities—City of Guatemala 
at Los Baios, Los Arcos, Laguna Amatitlan, Coban, Panzos Ysabal, Duenas, ete. 
The present form is from Los Bafos and from Coban. 


168 CALIFORNIA ACADEMY OF SCIENCES. 


GENERAL REMARKS. 


I consider this a subspecies of Sparganophilus Benhami, at least until a closer 
investigation of better material may reveal other characters, if any there are. Spar- 
ganophilus guatemalensis is one of the smaller species about 10 em. long by 2 mm. 
wide; the specimens were considerably stretched. Color is deep flesh, with an intense 
deep, dark violet lustre, much darker than in any other species and well preserved in — 
alcohol, in which the specimens appear blackish violet. 

Setw are lateral and dorsal. 

Chitellum extends from xvi-xxvi, but cannot be well defined on the ventral 
side. The tubercula pubertatis are in the shape of two parallel ridges extending from 
Xvili to xxii or specimens from Patal and Coban, which I take to belong to the same 
species as those from Los Bajios in the City of Guatemala. 


INTERNAL CHARACTERS. 


Of these I could only distinguish a few. The spermathece are in three pairs, 
one each in vii, viii, ix, opening in the anterior intersegmental groove. ‘The apical 
end in the fully developed spermatheca is very wide, flattened like a mason’s trowel. 
Several of the spermathece possessed a slit or opening in the apical end which com- 
municated with a longer or shorter sac, continuous with the exterior lining of the rest 
of the spermatheca. It is possible that this is only a result of maceration, as I have 
seen nothing like it in other species of Sparganophilus, and it is not probable that we 
here have an analogy with the spermathece of Enchytreeus or Sutroa, where this 
organ communicates with the intestine. 

Prostates are found in four pairs and situated in somites xxiy—xxvil or one 
somite further back than in Sp. Benhami. This in specimens from Sapote. If this 
character holds good it is one of considerable importance. 

Hearts were in yili, ix, x, xi strongly developed, but not filling the ccelom. 
Some of the Guatemala Sparganophili possessed only three pair of hearts in somites 
ix, X, xi, with the ventral vessel branching in xyv/xvi. Those were specimens from 
Los Arcos. Others again from Amatitlan possessed the hearts in x, xi, xii, xiii, with 
the ventral vessel divided in xv/xvi, while those from Los Baiios, Guatemala City, had 
the ventral vessel branched in xiv/xv. I believe those from Amatitlan and Los 
Arcos belong to different species, but, as all the specimens are lost, I can only call 
attention to the differences and to the importance of further investigations. 


Sparganophilus carneus n. subsp. 


Habitat. Mississippi river near Clayton, Iowa, in soil at the water’s edge, 
under pieces of boards and lumber. Numerous specimens at end of August, 1890, 
only few of which were adult. 

General remarks. It is probable that this is only a northern form of Sparga- 
nophilus Benhami, from which species it differs principally in the form of the sperm- 
athece, and by a much lighter color. Thespecimens at my command were not in proper 
condition and the shape and position of the tubercula pubertatis could not be made out. 


PACIFIC COAST OLIGOCHATA, 169 


My object is merely to call attention to this form which after all may be more distinct 
as more of its structure is known. In regards to the form of the spermathece it 
differs much more from Sp. Benhami than does the other subspecies Sp. guutemalensis. 
It is also much smaller. 

CHARACTERISTICS. 


Color is much lighter, more flesh and less violet than that of Sp. Benhami and 
Sp. guatemalensis, and the irridescence more faint. The species is smaller than 
Sp. Benhami, being about 9 em., about one-half the size of that species. 

Clitellum in the most developed specimen occupied xy—xxiv ventrally, dorsally 
it could not be properly defined. Prostates present with certainty in xxiii, xxiv, but 
owing to the somites being torn in xxv, xxvi I could not with certainty ascertain their 
presence there. 

Sete are similar to those in Sp. Benhami. The anterior xxii somites have 
much larger sete than the other, except the most posterior ten. The sets increase 
in size from somite 1i to somite xy, then diminish backwards to xxiii, in which somite 
they are smaller yet, this size continuing to the tail, where at about ten somites from 
the anus the sete again begin to increase. There is no parietal gland in somite 
lil as in Sp. Kiseni, and this is the chief reason why I class this species under Sp. 
Benhami. The spermathece are in three pairs and open in front of 3 and 4 dorsally. 
Their shape resembles much more those of Sp. Hiseni than they do those of Sp. 
Benhami. The apical end is globular and very much larger than the basal part. 
The spermathece are not trowel-like. I give some outline drawings of the sperm- 
athecz taken from one and the same individual. 

Sperm-sacs are very large in xi, xii, the one in xxii projecting into xxiii filling 
the somite also. The sperm-sacs are coarsely lobulate, much less so than in Sp. Hiseni. 

The capillaries resulting trom the lateral integumental vessels are very large 
in the anterior six somites. The hearts also are very large and seem to fill all available 
space on the coelomic cavity in vill, ix, x, xi. 


Deltania Troyeri Eisen, n. var. crassa. 
Figs. 142, 143. 


Derrinition. Size 45 mm. by 2 1-2 mm. Septal glands in v and vi, about twice 
as thick as the septum. Spermathecw each with two diverticles, which have the apical 
ends globular and inflated. Prostates thick, with the glandular part about 1 1-2 times 
longer than the thin muscular duct. Habitat, Baja California. 

In other respects like the type, judging from two dissected specimens. 

Habitat. Of this variety I possess only three specimens, from Ensenada de 
Todos Santos, in the northern part of Baja California, on the Pacific Coast side. They 
were taken in the same locality as Aleodrilus Keyesi, in the pure sand in the creek 
bottom, in a spot which had been kept moist by the accumulation of some sacks and 
rubbish. The specimens became strongly contracted, on account of the necessity of 
immediately placing them in alcohol. 


170 CALIFORNIA ACADEMY OF SCIENCES. 


EXTERNAL CHARACTERS. 

The variety resembles the main form in most respects, but is much larger, 
especially thicker. The contracted specimens were as long as the longest of the main 
form from San Francisco, when fully extended, and twice as thick, and must, when 
alive, have measured 45 mm. by 24 mm. Other external characters agreed exactly 
with those of Deltania Troyeri type from San Francisco, California. The deltoid 
arrangement of the ventral sete in the genital region was identical, which satisties me 
that we in this arrangement have most important species characters between the 
species of this genus. 

INTERNAL CHARACTERS. 

The variety differs from the type in two particular points. 

The septal glands are present only in two somites, v and vi, and are exceedingly 
thin, or about one-fourth as thick as those of D. Troyeri type. This refers to those 
superior to the intestine. Those below the intestine I could not discern, as no sections 
were made. The glands were lamelle-like, and only about twice as thick as the 
septum. 

The spermathece were thicker than those of the type, with more inflated diver- 
ticula. 

The prostate was thick, and its glandular part about 13 times longer than the 
very thin muscular duct. Two or three fully developed penial sete with each 
prostate. 

Deltania Troyeri Kisen, n. var. lagune. 
Figs. 144-147. 


Derinition. Size 32 mm. by 1 1-2 mm. No copulatory papiila. Setw a and 
b newt posterior to clitellum, reach their final distance in the fifth somite posterior to clitel- 
lum. Spermathece in ix twice as large as those of the type, contracted at center, apical 

/ Y i L » Of 
nart swollen. Prostates more slender than in the type. Large ovisacs. Habitat, Baja 
l i; , 
California, Sierra Laguna. 

L ? g 
DETAILED DESCRIPTION. 


Habitat. The exact locality in the Cape Region of Baja California and Sierra 
Laguna is a camping-place called La Joya. There, where the trail crosses the creek 
the water had backed up behind some fallen logs causing a miniature mudflat covered 
with leaves and sediment. The specimens were found in the rotten wood and under 
the decayed leaves in the mud. Altitude about 5500 feet. 


EXTERIOR CHARACTERS. 


The average specimen is larger than the type from San Francisco, the largest 
specimen of the latter being smaller than the average mature specimens of the 
variety Jagunw. The two round copulatory palpille figured by me from D. Troyert 
are not seen in the variety. The sacs of penial setee in several dissected specimens 
possessed each two penial sete fully developed, one having three, but of which one 


PACIFIC COAST OLIGOCHMTA. 171 


was undeveloped. One of the sete is a trifle more curved than the other. The 
point of each penial seta shows a slight and indistinct ornamentation (fig. 144). 
Since my first description of D. Troyeri I have found some more mature specimens at 
the old locality in San Francisco. One of these specimens possessed two penial setze 
in each sac, both slightly sigmoid, with trace of ornamentation at the free apex. In 
the variety /agune the ornamentation is more distinct. 

Common sete resemble the type, but setee a and 4 next posterior to clitellum 
reach their proper and final distance from each other only in the fifth somite posterior 
to clitellum, while in D. Troyeri type they reach the final distance in the third somite 
posterior to clitellum. 

INTERNAL CHARACTERS. 


Spermathec are much larger in the variety /agune, or about twice the size of 
those in D. Troyeri, with the main body distinctly contracted at the center, and with 
the apical part much enlarged. The diverticula are distinctly tri-digitate and reach 
up to the narrow contraction of the main sac. 

Prostates or spermiducal glands are much more slender and longer than in 
D. Troyeri type, and generally but not always confined to one somite. 

Ovisacs are two, projecting from the septum xiii/xiv into xiv, occupying the 
largest part of the ccelomic cavity in the latter somite. The structure of the ovisac 
is characterized by numerous trabecula forming an extensive system of rouad pockets 
of various sizes. Where there are no pockets there are solid zones of round or slightly 
oval nuclei (fig. 147) imbedded in a dense mass of tissue without distinct cell-walls. 


PHQE:NICODRILUS Eisen. 


The finding of a new species of this genus enables me to more properly define 
it. The genus was originally based on the absence of a prostate at the male-pore, 
there being, however, a short muscular atrium, in which opened the spermduct. The 
type was Phanicodrilus taste, a species from the Cape Region of Baja California. 
The new species described below under the name of Phenicodrilus tepicensis possesses 
amuch more developed atrium, in shape and structure resembling a spermatheca. 
There is a total absence of glandular cells, such as are found in prostates, the whole 
atrium being muscular. There is besides another difference between Phcenicodrilus 
and Oecnerodrilus, though of much smaller importance. In Phcenicodrilus we find a 
large number of muscular fascicles divided in three paired groups in somite xvii, radiat- 
ing from the male pore. In Ocnerodrilus these arciform muscles are few in number. 
In Pheenicodrilus these muscles are so thick and numerous that the atrium itself can 
only be seen in sections, not by suface view of the inner side of the body-wall. 

Through the possession of a muscular atrium this genus comes rather close to 
Nannodrilus Beddard. This genus, of which so far only one species is known from 
Africa, possesses a muscular bursa copulatrix, in which opens separately the anterior 
prostate and the anterior spermduct. The great similarity between the” atrium in 


Pheenicodrilus and the bursa copulatrix of Nannodrilus is very striking, especially as 
Memoirs, Vou. II, 5. January 28, 1896. 


172 CALIFORNIA ACADEMY OF SCIENCES. 


the spermducts in Pheenicodrilus also open intothe atrium. But the question that still 
remains to be settled is this. Is the atrium in Phcenicodrilus a modified bursa copu- 
latrix, or is it simply the remains of deteriorated prostate. The absence of a penis 
in Phoenicodrilus makes the latter alternative seem most probable. While in Oene- 
rodrilus the reduction of the prostate consists in the loss of one layer of glandular 
cells in the apical part, this reduction has proceeded further in Phcenicodrilus, the 
whole glandular apical part here being wanting. 

The principal differences between the genera Ocnerodrilus and Pheenicodrilus 
are then as follows: 

Ocnerodrilus. A prostate of varying size is always present at the male pore. 
This prostate consists of two parts, one glandular, lined by a single layer of cells, and 
one muscular basal part. 

Phenicodrilus. No prostate, only a short or rudimentary muscular atrium, in 
the lower part of which opens the spermduct. 


Pheenicodrilus taste Eisen. 


Derinition. Atrium rudimentary, muscular and bulbous, only about twice as 
thick as the spermducts. No ovisac. Spermathece covered with warty excrescences. 
Arciform muscles in xvii comparatively few. 

This species, which I first described from the west coast of the Cape Region 
of Baja California, has since been found by Messrs. Eisen and Vaslit in two widely 
separated localities. It was found in great numbers at Miraflores, on the east side of 
the Sierra of the Cape Region, some 30 miles north of San José del Cabo, together 
with Ocnerodrilus Beddardi, Sept., 1894. It was, however, quite unexpected to find 
this species also common in garden soil at San Blas, the seaport in the territory of 
Tepic, several hundred miles south of the Cape Region peninsula. I can find no 
differences in the anatomy, nor in the exterior of the specimens from these respective 
localities. The small copulatory papilla in somite xiv, lateral to the oviduct, is present 
in all. So are the numerous muscles in the copulatory region around the male pore. 


Pheenicodrilus tepicensis n. sp. 
Figs. 155-160. 


Derrinition. Atrium about twice as long as the body wall, but hidden by arciform 
muscles in somite xvii. A median ovisac in axiv/xiti. Spermathece entirely smooth, 
without warty excrescences. Arciform muscles in xvii very numerous. 

Halitat. Numerous specimens in and around the city of Tepic, territory of 
Tepic, Mexico, 4000. In moist soil under logs, in gardens, ete. November, 1894, 
Found together with Acanthodrilus Vasliti. Only few specimens were adult in 
November. 

EXTERIOR CHARACTERS. 


Size4 cm. by 15 mm. Number of somites 75. 
Chitellum incomplete, comprising somites xili—xix. 


Oo 


PACIFIC COAST OLIGOCHAETA. il 


Sete strictly paired, sigmoid plain. The pair 1 and 2 is wanting in xvi. 

Spermathecal pores in ix in front of sete 1 and 2. 

Oviducal pores in xiy in front of sete 1 and 2. 

Spermaducal pores in the center of xvii, where otherwise would be the sete 
land 2. A small copulatory papilla around each male pore, but it is not connected 
with a zone. No dorsal pores. 


INTERNAL CHARACTERS. 


Muscles. Numerous arciform muscles in the copulatory region, but especially 
in Xvi, connecting the copulatory papille with the body-wall in line with sets 3 and 4. 
These arciform muscles are arranged in three distinct groups. The central group, 
which branches out fan-shaped from the inner surface of the copulatory papille, is by 
far the largest, consisting of about 10 to 12 distinct fascicles. Anterior to this is 
another group of 6 to 7 fascicles, and in the posterior part of the somite we find a 
swollen group of about 5 fascicles. To what extent these vary as to numbers, ete., is 
uncertain, but my observations in Phwnicodrilus taste are that they are quite constant, 
and might be used as valuable characters in determining the species, as with a change 
in the muscular strands is also connected one in the exterior copulatory zone. Between 
xvii and xiii, as well as between xvii and xix we find several pair of arciform muscles 
in each somite. 

Septal glands are well developed. The suprapharyngeal glands are very iong 
and extend far backwards. The septal glands in the following somites diminish 
posteriorly. 

Spermathece. One pair in ix open between viii/ix in front of sete 1 and 2. 
Form sac-like, very thin walls, pellucid, outline smooth, without any warty diverticula. 
No distinction between a muscular and glandular part. The size is large, about as 
long as the somite is wide. 

Testes, ovaries and ciliated rosettes not characteristic. 

Ovisac. There is a median ovisac in xiii/xiv (fig. 157), consisting of a 
pouching backwards of part of the septum. This is the only species in the three 
genera, Ocnerodrilus, Gordiodrilus and Pheenicodrilus, which possesses an ovisae. 

Spermducts are of the same width throughout, without any muscular enlarge- 
ment near the male pore. The two ducts run together, but their lumens are separate 
until they reach the muscular atrium, where, just before they enter it, the two lumens 
fuse into one (figs. 158, 159). 

Muscular atrium. As has already been stated the usual prostates in Oenerodrilus 
are replaced by a pair of muscular pouches, entirely covered up by the numerous 
arciform muscles. There are no glandular cells, and these atria can best be compared 
to the basal muscular parts of the Ocnerodrilid prostate only they are very much 
thicker, and consist entirely of muscular cells with the lumen in the body-wall lined 
by columar epithelium. In the main pouch this epithelium consists of short cells with 
round nuclei, around which extends a thin musenlar layer, which becomes wider at 
the base. This atrium is very short and cannot readily be seen when the opened 


174 CALIFORNIA ACADEMY OF SCIENCES. 


body cavity is viewed from the interior surface. It is attached to the at this point 
very much thickened body-wall (fig. 158) by numerous muscles and connective 
tissue. The various species of Ocnerodrilus, especially those with short prostates, 
require reinvestigation in order to ascertain the structure of the prostates. Should 
intermediate forms be found it might become proper to merge the two genera into one. 

Nephridia. Those anterior to clitellum are not covered with a ccelomic cell 
mantle, while those posterior to the clitellum are surrounded by a very thick one. . 
The structure of the nephridia appears to be similar to those of Phwnicodrilus taste, 
at least in a general way. 


LITERATURE REFERRED TO. 


Benuam, W.B. A new English Genus of Aquatic Oligochwta (Sparganophilus), belonging to the family of Rhino- 
drilide. Quarterly Journal Micros. Science, vol. xxxiy. 

Bennam, W.B. Report on an Earthworm collected for the Nat. Hist. Dept. of British Museum by Emin Pasha in 
East Africa. Journal Roy. Microse. Society, February, 1891. 

Bepparp, Fr. E. Two New Genera, Comprising Three New Species of Earthworms from Western Tropical Africa. 
Proceedings Zoological Society, 1894, pages 379, etc. 

Bepparp, Fr. E. A Monograph of the Order of Oligochwta. Oxford, 1895. 

Crerrontarne, P. Recherches sur le Systeme Cutané, etc., du Lombric Terrestre. Archives de Biologie, x, 1890. 

Eisen, G. California Eudrilidw. Memoirs Cal. Academy of Sciences, vol. ii, No. 3. 

Ersen, G. Pacific Coast Oligochwta I. Memoirs Cal. Acad. of Sciences, vol. ii, No. 4. 

Eisen, G. Anatomical Studies on New Species of Ocnerodrilus. Proceedings Cal. Academy of Sciences, ser. 2, 
vol. iil. 

Garman, H. On the Anatomy and History of a New Earthworm (Diplocardia communis). Illinois State Labora- 
tory of Nat. History, ili, 1888. 

Hessg, Ricuarp. Uber the Septaldriisen der Oligochwten. Zoological Anzeiger, No. 456, 1894. 

Hessz, Ricnarp. Zur Vergleichende Anatomie der Oligochwten, Zeitschrift f. Wissensch. Zoologie, 1894, pages 
406. 

Horst, R. Earthworms from the Malay Archipelago. Zoolog. Ergebn. e. Reise in Niederland. Ost-Indien. Dr. 
Max Weber, B. ii, Leiden, 1892. ; 

Lanepon, F. E. Sense Organs of Lumbricus. Journal of Morphology, vol. xi. 

Micnarrsen, W. Oligocheten des Nat. Hist. Museum, Hamburg, iii. 

MicuarEtsen, W. Oligochewten des Nat. Hist. Museum, Hamburg, No. iv. 

MicHakEtsenN, W. Regenwiirmer der Thierwelt Ost-Afrikas, Bd. iv, Berlin, 1895. 

Rosa, Dr. D. Annailen d. K. K. Nat? Hist. Hofmuseums. Die Exotischen Terricolen, Wien, 1891. 

Rosa, Dr. D. Contributo allo Studio dei Terricoli Neotropicali. Accademia Reale delle Scienze di Torino, 1895. 

Swirn, Frank. A Preliminary Account of Two New Oligochata from Illinois. Bulletin Illinois State Laboratory 
of Nat. History, vol. iv, 1895. 

Upr, H. Beitrage zur Kentniss auslandischer Regenwirmer. Zeitschr.f. Wissens. Zoologie, 57, 1894, page 69, etc. 

Vespovsky, Fr. Organogenie der Gordiodrilen, Zeitschr. f. Wissensch. Zoologie, Bd. lvii, p. 642. 

Verspovsky, Fr. System u. Morphologie der Oligochawten, Prag., 1884. 

Verson, E. yon. Zur Spermatogenesis, Zool. Anzeiger, xii, 1889, p. 100. 


PACIFIC COAST OLIGOCHETA. 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buceal cayity. 

ce. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. ceelomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cel. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c, t, connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d.v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. 8. gr. intersegmental groove. 
l. c. large cells. 

1. m. longitudinal muscles. 
m. muscles. 

m. atv. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n. c, nerve cells. 

ne. nerve ganglion. 


nec, neck of nephridium. 
neph. nephridium. 

neph. p. nephropore. 

@s. esophagus. 

ov. ovary. 

ovd. oviduct. 

org. Sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p.f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 
pr. prostate. 

p. spd. posterior spermducet. 
pr. sto. prostomium. 

ps. penial sets. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 
sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 


s. org. b. sense organ cells in equatoreal zone. 


8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

s. ps. Sac with penial seta. 

sp. 8. Sperm-sacs. 

8. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u. c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. ‘windings ” of nephridial ducts. 


175 


176 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLVI. 


BENHAMIA PALMICOLA. 
0. Longitudinal section of the genital somites. 


BENHAMIA NANA. 
1. A specimen; natural size. 


2. The ventral side of the anterior somites, showing the exterior pores, etc. 


The yentral side of the most anterior somites, showing the everted pharynx and the very narrow somite i, etc. 


Pee 


Side view of the same somites. 


co 


The anterior twenty somites laid open, exposing the various organs, ete. The alimentary canal is removed 
and the whole figure is held somewhat schematic. aftr. thickened part of the spermducts. 

Part of a posterior somite viewed from the inner side, showing the relative position of sets and nephropores. 
1, 2, 3, 4 refer to the sets; a, b, c to the nephridia. 


(>) 


MeM.CAL.Acan.[l FLare XLVI 


1H wo 


non wu 
‘ia —— a 
5 now wow 
ae 


non 


win ohn 


BENHAMIA PAPILLATA Fic 0. ~ BENHAMIA NANA !'6'5. 170 6. 


LITH, BRUTTON & REY, &.R 


=I 


PACIFIC COAST OLIGOCHETA. 17 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrvium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 
c.c. columnar cells. 

c. ep. celomie epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. s. gr. intersegmental groove. 
l. c. large cells. 

l. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n. c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium., 

neph. nephridium. 

neph. p. nephropore. 

es, cesophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p. c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
pha. pharynx. 

p- neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium. 

ps. penial seta. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

s. org. 6. sense organ cells in equatoreal zone. 
8. ory. pr. sense organ cells in prostominm. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. sac with penial seta. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

irb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u. c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. “ windings ” of nephridial ducts. 


178 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLVII. 
BENHAMIA NANA. 


7. A somewhat diagrammatic figure representing the anterior part of a specimen sectioned lengthwise, composed 
from several sections, showing the location and size, etc., of various organs. wz. beginning of the muscular 
part of the spermduct. 


8. Cross-section through the most anterior part of the body, through prostomium and buccal division, showing 
the pharyngeal glands in four distinct divisions. The section is anterior to the cephalic ganglion. 


9. Section posterior to the former, through somite ii. 
10. Section through somite vii. 


11. Section through somite xiv; the calciferous gland with the intestine belongs to somite xiv; the ovary to xiii. 
The section passes just in front of the oviduct. 2. the limit of the clitellum. 


12. Section through somite xvi. The calciferous diverticula belong to somite xv. ~ 
13. Section through somite xvii and the anterior prostate pore. zx. ending of clitellar cells. 
14. Section through somite xx, showing the typhlosole. 


15. Longitudinal section through the most anterior part of the body, showing the projected pharyngeal division 
with the gland ducts. pha. pharyngeal division. The dark line indicates the row of discharge chambers of 
the ducts from the glands. A more magnified view is represented in fig. 18. 


16. A couple of strands of the unicellular salivary septal glands with their muscles. 

17. Some of the septal unicellular glands more highly magnified. 

18. Section through pharynx, showing the discharge ducts and pockets from the septal and pharyngeal glands. 
19. Lining of the esophagus below the salivary giands, showing its exceeding thinness. 


20. Longitudinal section through the body-wall, showing the sense organ region in the equatorial line of the 
somite. c.e. central lumen or organ proper, in which are seen the sense cells, together with nuclei of 
supporting cells. These supporting cells are surrounded by peculiar lunate or sac-like transparent cells, also 
found in the sense organ of the pharynx. 7. c. nerve cells and their nuclei. 


21. Longitudinal section through gizzard, showing glands with their discharge tubes and small terminal pockets. 


22. Section through the alimentary canal between the two gizzards, showing glands. c. cluster of lunate cells 
surrounding a lumen. 


23. Longitudinal section of the tubular intestine from somite xvi. u.c. unicellular glands. 6/. s. blood sinus. 


24.. One of the lobes of the typhlosole, showing distribution of glands, some of which are dark staining, others 
not. 


24b. Section through the intestine adjoining the typhlosole. 
25. Two glandular cells from the former section, more magnified. The two glands belong to different types. 


26. A pair of calciferous diverticula from xiv. The right-hand gland is much smaller, which is an exception, as 
generally they are of equal size. 


27. Detail of a section of lamella from one of the anterior calciferous diverticula. 
28. Two cells more highly magnified. 

29. Section through one of the posterior calciferous diverticula. 

30. Part of the same more highly magnified. Zeiss Hom. Im. Eyp., 2. 

31. One of the prostates with sac with penial sete. 

32. Cross-section through glandular part of prostate. 

33. Part of a penial sete near the apex; the latter is broken. 


LITR. BUTTON & REY RE 


Mem. CAL.Acan.U. PLATE XLVI 


AAs 


as 
= 
™ 
Z 
[a 
S 
= 
: 
a 


RUUD) 


f 
at oy 
_ 
- a 
é i > 
- = 
; > i 
” 
. 
* 
+ 
- 
. 
\ 
’ 


PACIFIC COAST OLIGOCH@HTA. 179 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior caleic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buceal cavity. 

ce. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. celomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chioragogen cells. 

cil. ciliated epithelium. 

el. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v, dorsal vessel. 

¢. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. 8. gr. intersegmental groove. 
l. c. large cells. 

1. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. Inesenteric tissue 

m. pr. muscular prostate. 
n. nephridia. 

n. c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

ws. esophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p. J. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium, 

ps. penial sete. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gb. Suprapharyngeal glands. 

s. org b. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. 
spe. Sperm-cells, 

spd. spermduct. 

spgn. spermatogonia, 

spr. spur of nephridium. 

8. ps. Sac with penial seta. 

sp. 8. Sperm-sacs. 

8. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellar glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. “windings” of nephridial ducts. 


150 


36. 


37. 
38. 


39. 


40. 
41. 
42. 


43A. 
43B. 
43C. 
43D. 
43E. 


44A. 
44B. 
44C. 
45. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLVIII. : 


BENHAMIA NANA. 
Section through one of the prostate pores, showing prostate, sac with penial sete and the thickened spermduct. 
x. at this point the clitellar cells cease. c. ¢. connective tissue between the two muscular layers. ; 
Cross-section of the thickened part of the spermduct, showing the two lumens separate. 


Section through the muscular part of the prostate, showing the epithelium and some of the muscles nearest 
the lumen. 


One of the posterior nephridia; the shaded sac-like part represents the celomic mantle. It is generally covered 
with capillaries to such an extent that the structure of the organ is obscured. br. bridge. mec. neck of nar- 
row duct. sp7v.spur. o./f. anterior fold. p.f. posterior fold. 


One of the anterior nephridia, showing absence of ccelomic glands, also the arrangements of blood vessels. 
Part of the nephridium more magnified. No capillaries are represented. 
Part of celomic mantle of one of the posterior nephridia. se. secreted matter staining very deep. mn. nuclei of 

the glandular cells. 

BENHAMIA PAPILLATA. 

A specimen, natural size, from Tepic. 

The ventral zone of the clitellum. 

The same more highly magnified, showing the papillw and prostate pores. 

The two penial set in one sack. Zeiss D. Eyep., 2. 


The tips of the two penial sete. Zeiss 12, Eyep., 2, 1. 


BENHAMIA PALMICOLA. 
The ventral side of the anterior somites. 
The ventral region of the clitellum. 
A specimen, natural size. 


Penial set. A. apex of the shorter sete. B. apex of the longer setw. C. two sets, whole. 


Mew. Cab-Acan II. PLATE XLVIEL 


= 
= 
= 
= 
= 
a 
= 
> 
= 
= 
= 
= 
x 
= 
= 
2 
= 
= 
= 
= 
= 
= 


Gusmv Een TzL. LTH, BRITTEN & REY SF 


BENBAMIA NANA Figs. 367042. HENHAMIA PAPILLATA Fic. 43. HENHAMIA PALMICOLA He's. 44 & 45. 


, “288 


PACIFIC 


COAST OLIGOCHEHTA. 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 

buc. buccal cavity. 

c. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. celomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 

c. r. ciliated rosettes or sperm-funnels. 
c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 

d. pr. dorsal pore. 4 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. s. gr. intersegmental groove. 
1. c. large cells. 

1. m. longitudinal muscles. 

m. muscles. 

m. atr. muscular atrium. 

mes. mesenteric tissue. 

m. pr. muscular prostate. 

n. nephridia. 

n. c. nerye cells. 

ne. nerye ganglion. 


nec. neck of nephridium. 
neph. nephridium. 

neph. p. nephropore. 

ws. esophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p. c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 
pr. prostate. 

p. spd. posterior spermduct. 
pr. sto. prostomium. 

ps. penial sete. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 
sep. gl. septal gland. 

sep. septum, 

s. gl. suprapharyngeal glands. 


s. org. b. sense organ cells in equatoreal zone. 


8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells, 

spd. spermduet. 

spgn. spermatogonia. 

spr. spur of nephridium. 

s. ps. sac with penial seta. 

sp. 8 sperm-sacs, 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m, transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. ‘windings ” of nephridial ducts, 


181 


182 


46. 


47. 


52. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE XLIX. 


BENHAMIA PALMICOLA. 

A somewhat diagrammatic figure, composed from several longitudinal sections. s. org. pr. sense organ zone 
of the prostomium. ss. org. b. the sense organ zone of the body wall, on the equatorial line of the somite. 

Some of the sense organ cells of the prostomium more highly magnified. Zeiss. 12; Hom. Im. Eyrp.2. The 
sense cells are seen in the center between the pellucid sac-like cells. Between the sense cells are seen nuclei 
of supporting cells. 

Part of a posterior somite laid open, showing the three nephridia on one side. 

A part of a nephridial cell mantle, showing various kinds of cells, some of which appear to contain caleuli. 

Cephalic ganglion, ete. 

The spermathecal zone seen from the inner side of the body. The large globular apical part of each sperma- 
theca is situated in the somite posterior to the pore. 


Outlines of the spermathecs of various extra African species of Benhamia, some of which are copied from 
various sources. 


52A-B-C-D. Spermathecw of Benhamia Bolavi, from specimens from Hamburg, kindly furnished by Dr. W. 


52E. 
52F. 
52G-— 
521. 


52K- 


Michaelsen. 
Spermatheca of Benhamia malayana after Dr. Horst. 
Spermatheca of Benhamia floresiana after Dr. Horst. 
H. Two spermathecw from the same side of a specimen of Benhamia papillata from Tepic, Mexico. 
Spermatheca of Benhamia octonephra after Rosa. 


L. Spermathece of Benhamia palmicola from Miraflores, Baja California, 


52M-N. A spermatheca of Benhamia rugosa seen from two different sides. 


Mem.CAL.ACAn Il. PLATE XLIX 


Benham Bolavt 


ov. 
Penhamia Folavt 


PBenhamia Bolavt 


Benham Malayana 


Benhama papillata 


Benhamia Florestana. 


Lenhamea nana. 


é ©) (Gam 


LITH, BRITTON & REY, SE 


_ HENHAMIA PALMICOLA Fig's 46 Td 51 


Gusmav Eisen Det 


wy eh ra 


- 


PACIFIC COAST OLIGOCH ETA. 183 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcice gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buceal cavity. 

c. clusters of lunate cells. 

ec. c. columnar cells. 

c. ep. celomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cel. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d.v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. 8. gr. intersegmental groove. 
l. c. large cells. 

1. m. longitudinal muscles. 
m. muscles, 

m. atv. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n.c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

@s. esophagus. 

ov. Ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermducet. 

pr. sto. prostomium. 

ps. penial sete. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

8. org. b. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. sac with penial seta. 

sp. 8. Sperm-sacs. 

8. ph. sgl. sabpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

¢. m. transverse muscles. 

irb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u. c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

sgl. ventral gland. 

w. “ windings ” of nephridial ducts. 


= 


184 CALIFORNIA ACADEMY OF SCIENCES. 


53. 
54. 


PATE 
BENHAMIA PALMICOLA. 


Part of the intestine and hearts dissected out. 
Prostate and sac with penial set. 
BENHAMIA RUGOSA. 
The ventral side of clitellum, showing the copulatory papilla on which are situated the anterior prostate pores 
in xvii. The clitellum is much crenulated. 
Anterior somites seen from above, showing prostomium, somite i, ii and iii, the first of which is much swollen. 
The same seen from the side. In these two figures somite i is retracted in the buccal cavity. 
Another specimen; the anterior somites, but i, is fully extended. 
Larger penial sets, free end. 
Smaller penial setzx. 


A partly diagrammatic representation of the interior surface of ‘a posterior somite, showing the nephridia. 
1, 2, 3, 4, small, refer to setw; 1, 2, 3, 4, large, refer to nephridia. 


ALEODRILUS KEYESI. 
A specimen, natural size. 


The anterior somites seen from the ventral side, showing the generative pores and the position of set. 
Prostomium and somite i. 
The sexual fosse and papillx. 


A somewhat diagrammatic view of the anterior somites, composed from several longitudinal sections. 


“& dpr- “+ 2. 65... ft. 4 
ay : a = 43 2¢ aw) 
NF = 9 es ‘ | wh 
Guemyv Ersen Teh. LITH. BRITTON & REY SF: 


HENHAMIA PALMICOLA Pig's 53.%54 HENHAMIA RIGOSA Figs. Sh TO 69. Atgonrius Kevesi Mies. 66 TO 70. 


PACIFIC COAST OLIGOCHETA,. 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 
c.c. columnar cells. 

c. ep. ecelomie epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

cl. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. bypodermis. 

i. s. gr. intersegmental groove. 
l. c. large cells. 

l. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n. c. nerve cells, 

ne. nerve ganglion. 


nec. neck of nephridium. 
neph. nephridium, 

neph. p. nephropore. 

es, cesophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p. ¢. gl. posterior calciferous diverticulum. 
p.f. posterior fold, nephridium. 
pha. pharynx. 

p. neph. posterior nephridia. 
pr. prostate. 

p. spd. posterior spermduct. 
pr. sto. prostomium. 

ps. penial seta. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 
sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 


s. org. b. sense organ cells in equatoreal zone. 


8. ory. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

s. ps. sac with penial seta. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u. c. unicellular glands. 

v. g. ventral ganglion, 

v. v. ventral vessel. 

v. sgl. yentral gland. 

w. ‘ windings ” of nephridial ducts. 


185 


186 CALIFORNIA ACADEMY OF SCIENCES. 


71. 


77. 


78. 


79. 


80. 


81. 


PLATE LI. 
ALEODRILUS KEYESI. , 


Cross-section through somite ix, showing the septal glands and the thickened septa. ss. gl. septal gland, the 
prolongation of which probably connects with the intestine and the median blood vessel. zx. at this point the 
ventral vessel and the septal gland have been bent and cut twice. 


Cross-section through somite xii, showing the posterior sperm-sac. tra. trabecula. v. v. ventral vessel in two 
successive somites. spc. sperm-ce]l germinative area of cells with narrow nuclei. 


Section through apical part of spermatheca. 


Section through intestine and septal gland in somite x, anterior to fig. 72, showing discharge duct of septal 
gland. The ventral vessel and heart have been folded and cut three different times; similarly part of the septal 
gland has been cut twice. d. discharge duct of septal gland; it probably reaches in between the epithelia, 
cells of the intestine. 


Section through dorsal median vessel, where it connects with two lateral vessels; the section goes only through 
one of the points of connection. The lateral vessels are covered with chloragogen cell. 

Section through the suspended posterior part of the spermduct in the anterior part of somite xii. The duct is 
seen suspended between two mesenteric strands, one connecting with the heart, the other with the septum 
close to the body wall. h.v. heart valve. gl. gland emptying in spermduct. 


Section through spermduct in somite xi, the cut going through testis, which is immediately superposed on the 
duct or rather the posterior part of the ciliated rosette. 

Section through spermduct going through the ovary in xiii. a. spd. spermduct from anterior rosette, already 
attached to the body wall. p. spd. spermduct from posterior rosette, yet suspended and passing through the 
ovary. ov. ovary with full-grown ova, with germ-cells and with a germinative area where cells are in mitosis. 
The nuclei in the germ-cells are oval, while those in the grown ova are round. Zeiss D. Eyp. 2. 


Section through spermduct through one of the posterior somites where the two ducts have joined in the body- 
wall. clit. clitellum in xiy. 


bbw 


Guspy Ensen [ut 


%. 


; 
, 


PACIFIC COAST OLIGOCHETA. 


REFERENCE TO LETTERS ON PLATES. 


a.c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. celomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chioragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

cel. clitellum. 

c. m. circular muscles. 

com, commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
giz. gizzard. 

h. heart. 

hy. hypodermis. 

i. s. gr. intersegmental groove. 
1. c. large cells. 

1. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium, 
mes. mesenteric tissue 

m. pr. muscular prostate. 
n. nephridia. 

mn. c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

@s. esophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium. 

ps. penial sete. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

s. org b. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. * 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. Sac with penial seta. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

irb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellar glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. ‘windings ” of nephridial ducts. 


187 


84. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE LII. 


ALEODRILUS KEYEST. 
Section through the epithelium of tubular intestine, showing a cluster of glands between the epithelial cells. 
b1. blood lacune. 
Section through one of the valves of the heart. J/. c. large cells at the base of the valves. 
Some spermatogonia from the sperm-sacs in xii. 
Cells from the central germinative and in the sperm-sac. 


One of the nephridia with an anterior lateral vessel, connecting dorsally with a branch from the ventral vessel. 
d.v. diverticula of the lateral vessel. Jat. v. lateral vessel. 


ACANTHODRILUS TAMAJUSI. 
The largest of my specimens, natural size. This specimen is strongly contracted, and was when alive con- 
siderably longer. 
The anterior somites, ventral view. 
The anterior somites, dorsal view. 
The anterior somites, side view. 


The ventral part of the genital region in the clitellum, showing genital papillw, the three transverse genital 
ridges, penial sets, common sete, etc. The clitellum is not well developed, but in somite xx may be seen its 
posterior termination. The other one of my specimens, which possessed a more developed clitellum, had 
these papille less pronounced, though of the same form and arrangement. 


Cross-section of the body-wall, showing the common sete. 
The free end of the penial seta. 
The spermathece, dorsal view. 


One of the spermathece dissected out. 


al 


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ACANTHODRILUS TAMAJUSI 


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ATEODRILUS Keyes! Fiss. 6 


Guetay Excen Dez. 


PACIFIC COAST OLIGOCH/TA. 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior caleice gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

b]. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 

ce. c, columnar cells. 

c. ep. celomic epithelium. 

ce. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

el. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. 8. gr. intersegmental groove. 
1. c. large cells. 

l. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n. c. nerve cells. 

ne. nerye ganglion. 


nec. neck of nephridium. 
neph. nephridium. 

neph. p. nephropore. 

@s. esophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p. c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 
pr. prostate. 

p. spd. posterior spermduct. 
pr. sto. prostomium. 

ps. penial sets. 

prt. peritoneum. 

se. secreted matter. 

s. nt. sacculated intestine. 
sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 


s. org. b, sense organ cells in equatoreal zone. 


8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. sac with penial seta. 

sp. 8. sperm-sacs. 

8. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel, 

v. sgl. ventral gland. 

w. “windings ” of nephridial ducts. 


189 


190 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE LIII. 
ACANTHODRILUS. TAMAJUSI. 


96. A somewhat diagrammatic view composed from several longitudinal sections of the body. r. m. retractor mus- 
cles of gizzard. c.c. masses of free ecelomie cells. 


SPARGANOPHILUS BENHAMI. 

97. Adult specimen, natural size. - 

98. The anterior somites, ventral view, including clitellum, tubercula pubertatis, etc., magnified four times. 

99. Longitudinal section of the anterior somites passing through the spermathecal and prostate pores, composed 
from several sections. Somites xix-xxii are not figured. sep. gl. septal glands which open in the dorsal part 
of the pharynx. J. sep. gl. lower septal glands which open in the ventral part of the pharynx. The sperma- 
thecw have been cut through diagonally and the figure does not give any idea of their real shape. 

101. Section of the body-wall in ventral part of somite xii. p. peritoneum. 2. ep. nuclei of supporting cells. 

102. Longitudinal section of the clitellum. 7. Supporting cells. 2. Unicellular glands. 3. Short clitellar cells, 


staining deeply, and furnished with perfect nuclei. 4. Largest clitellar cells, staining less deeply and with 
imperfect nuclei. Zeiss D. 


103. A part of the same section, more magnified. Iron, hematoxylin, Haidenhein, Ehrlich 3 acid. Numerals indi- 
cate the same as in fig. 102. d. n. degenerate nuclei of the 4 cells. x. sec. cell plasma around the nuclei 
pushed far back in the apical part of the cell. mn. sh. c. nuclei of the short clitellar cells surrounded by cell 
plasma and mucin. ec. ¢. connective tissue. 61. c. blood capillaries which are very scarce in the clitellum of 
this species. 


104. Section of the body wali in somite xiv. 

105. Cross-section through clitellum and tubercula pubertatis. 7. Supporting cells. 2. Unicellular glands. 3. 
Shorter clitellar cells staining darkly. 4 The longer clitellar cells. 5. tpc. tubercula pubertatis 
cells, largest kind. A few of these are seen to contain free but shrunken nuclei. 6. tpc. tubercula puber- 
tatis cells of the narrow kind, with cell plasma in the apical end, but also with shrunken nuclei. ¢. p. tuber- 


cula pubertatis groove. n. f. nerye fibers. cl. c. 2. large unicellular glands, a large form of the usual uni- 
cellular epidermal glands. Above these is seen a wide lumen, probably a blood vessel. 


106. Longitudinal section of the body-wall, through spermatheca. 

107. Longitudinal section of body-wall through prostate and pore in somite xxiv. ¢.c. tubular supporting cells. 
g.c. glandular cells confined to the zone around the intersegmental grooves. 

108. The same as 107, more magnified. 

109. The inner whorl of the ciliated rosette. 

110. Section through the esophagus. 7. c. interstitial cells. gl. interstitial glands. 

111. Section through intestine in somite xii. fbr. blood crystals and fibrine. w. bl. w. wall of blood sinus. 


112. Section through upper pharyngeal wall, showing epithelial cells, and between them the discharge tubes of the 
glands. Zeiss 12, hom. im. 


113. Section through the same part, showing a three-forked duct. 


114. Section through the intestinal wall, next posterior to the real pharynx, showing larger discharge tubes and 
chambers. 


115. Section through ventral wall of pharynx, with ducts and storage chambers of glands, and between them shorter 
epithelial cells. m. and d. muscles along which run the ducts. n. d. nuclei of ducts. 

116. Surface view of one of the larger blood vessels. 

117. Outline of one of the nephridia. 

18. Outline of a spermatheca. 


CPARGANOPHILUS 


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PACIFIC COAST OLIGOCHEHTA. 191 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buceal cavity. 

c. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. ceelomic epithelium. 

c. gl. ealciferous glands. 

ch. discharge chambers of glands. 
chlo. chioragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t, connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d.v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

7, 8. gr. intersegmental groove. 
l. c. large cells. 

l. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n.c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

es. esophagus. 

ov. Ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p.f. posterior fold, nephridium. 
phew. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium. 

ps. penial setz. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

s. org. b. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. Sac with penial sete. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. ‘windings ” of nephridial ducts. 


192 


119. 


a. 
bd. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE LIV. 


SPARGANOPHILUS BENHAMI. 


Five cross-sections through various parts of the body. 


Through posterior part of pharynx in somite v. 


Through spermathecs and somites yili and ix. 
Through somite x. ¢.c. r. tube of ciliated rosette. 


Through oviduct, somite xiv with sperm-sacs. 
Through clitellum in somite xix, showing tubercula pubertatis. 


clitellar cells. 


cl. c. 1 long clitellar cells. 


spd. spermduct running below the muscular layers. 


s. gl. dorsal septal glands. /. s. gl. lower septal glands. 


cl. c. 2 short 


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PACIFIC COAST OLIGOCHETA. 193 


REFERENCE TO LETTERS ON PLATES. 


c. gl. anterior calcic gland. 
. f. anterior fold of nephridium. 
nephr. anterior nephridia. 
. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 
c.c. columnar cells. 

c. ep. celomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chloragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

cl. clitellum. 

c. m, circular muscles. 

com. commissures. 


ae ea 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d. v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. s. gr. intersegmental groove. 
1. c. large cells. 
l. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 

n. nephridia. 

n. c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

es. cesophagus. 

ov. Ovary. 

ovd. oviduct. 

org. sense organ. 

p. c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium. 

ps. penial set. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

s. org. b. sense organ cells in equatoreal zone. 
8. ory. pr. sense organ cells in prostomium. 
spe. Sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

s. ps. sac with penial seta. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u. c. unicellular glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. “windings ” of nephridial ducts. 


194 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE LV. 
SPARGANOPHILUS SMITHI. 


120. A large specimen, natural size, ventral view. 


121A. Anterior part of the body, side view, showing tubercula pubertatis ridges, and their anterior extension to 
the center of somite xi. The dorsal and ventral termini of clitellum marked by x. 


121B. The anterior somites, ventral view. The spermiducal pore is seen about half way between sete 1 and 2 aid 
the tubercula pubertatis in xix/xi. 
122. Two of the clitellar somites, side view, showing tubercula pubertatis ridge. v. ventral. d. dorsal side. 


SPARGANOPHILUS SONOM 2. 

123. The same as 122, but from Sparganophilus sonome, showing the tubercula pubertatis ridge to consist of a suc- 
cession of separate tubercles, two or three on eyery somite, with the grooye very nearly in the center of each 
papilla. 

SPARGANOPHILUS SMITHI. 


124. Longitudinal section of the anterior somites, composed from seyeral sections. v. sg/. ventral septal glands in 
vy, vi, vii. s.v.spermtanks. s. s. sperm-sacs. f. four large folds of the dorsal vessel in somites xiv—xviii. 
sp. gl. prostates. 

125. A series of outline drawings of spermathecw from one specimen. 


SPARGANOPHILUS SONOM 2. 
126. Outlines of spermathece from a single specimen. 
SPARGANOPHILUS SMITHI. 
127. Cross-section of septum xi/xii. 


128. Cross-section of the body; the Roman numeral indicates the somite. 


129. Cross-section of the body; the Roman numerals indicate the somites. 


8. 


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PACIFIC COAST OLIGOCHETA. 195 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcice gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct 

atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 

buc. buceal cavity. 

c. clusters of lunate cells. 

c. ¢, columnar cells. 

c. ep. celomic epithelium. 

c. gl. caleiferous glands. 

ch. discharge chambers of glands. 
chlo. chioragogen cells. 

cil, ciliated epithelium. 

cl. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com, commissures. 

c. 7. ciliated rosettes or sperm-funnels. 
c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 

d. pr. dorsal pore. 

d, v. dorsal vessel. 

¢. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 

gl. pr. glandular part of prostate. 
gizz. gizzard. 

h. heart. 

hy. hypodermis. 

i. 8. gr. intersegmental groove. 
l. c. large cells. 

1. m. longitudinal muscles. 

m. muscles. 

m. atr. rouscular atrium. 

mes. mesenteric tissue 

m. pr. muscular prostate. 

n. nephridia. 

n. c. nerve cells, 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

@s. cesophagus. 

ov. ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p.f. posterior fold, nephridium. 
phx. pharynx. 

p. neph. posterior nephridia. 

pr. prostate. 

p. spd. posterior spermduct. 

pr. sto. prostomium, 

ps. penial seta. 

prt. peritoneum. 

se. secreted matter, 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

8s. org 6. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. Sac with penial seta. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellar glands. 

v. g. ventral ganglion. 

v. v. ventral vessel. 

v. sgl. ventral gland. 

w. ‘‘windings” of nephridial ducts. 


196 CALIFORNIA ACADEMY OF SCIENCES. 


PLATE LVI. 
SPARGANOPHILUS SMITHI. 


130-136. Cross-sections through various somites. The details are indicated diagrammatically. The Roman numer- 
als indicate the number of the somite or somites through which the section passes. s. m. septal muscles. 
a. s. gl. accessory septal gland of septum between somites ix and x. spz. ¢. spermatozoa in spermtanks. spz. s. 
spermatozoa and spermatogonia in sperm-sacs. os. ovyisac. a. m. t. arciform muscles of tuberecula puber- 
tatis, connecting the opposite surfaces of the tubercula pubertatis. a.m. c. arciform muscles passing from 
opposite sides of clitellum through the ecelomic cavity. 

137. Section through sacculated intestine. /. c. blood capillary reticulum. 


138. Tubercula pubertatis, Zeiss Hom. Im. ,';. af. m. arciform muscles passing through ccelom. af. ¢. arciform 
muscles confined to the epidermal cells of the tubercula. 


139. Section through the body-wall through one of the prostates. 7. m. retractor muscles passing to the cuticle 
and the outermost epidermal cells. ct. connective tissue. 


SPARGANOPHILUS CARNEUS. 


140. Two spermathece from the same specimens. The free or apical end is seen to be almost globular. 


SPARGANOPHILUS GUATEMALENSIS. 


141, a and b. Two spermatheecw. The apical end is flat and trawl-like, bearing a sac-like pouch which connects 
with the interior of the spermatheca, through a small slit. 


DELTANIA TROYERI var. CRASSA. 
142. A prostate gland with penial sete. 
143. A spermatheca. 

DELTANIA TROYERI yar. LAGUNA. 
144. One of the penial sete with the tip more highly magnified. 


145. Two spermathecs, both from the same specimen. A and B are seen from the broad side, C from the narrow 
side. 


146. Two prostates with penial setw. A is seen from the flat side, B is seen from the broad side. 
147. Section of one of the ovisacs. Zeiss 4, Hom. Im. 


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PACIFIC COAST OLIGOCHETA. 197 


REFERENCE TO LETTERS ON PLATES. 


a. c. gl. anterior calcic gland. 
a. f. anterior fold of nephridium. 
a. nephr. anterior nephridia. 
a. spd. anterior spermduct. 
atr. atrium. 

bd. body-wall. 

bl. blood vessel or capillaries. 
bl. s. blood sinus. 

br. bridge in nephridium. 
buc. buccal cavity. 

c. clusters of lunate cells. 

c. c. columnar cells. 

c. ep. ccelomic epithelium. 

c. gl. calciferous glands. 

ch. discharge chambers of glands. 
chlo. chioragogen cells. 

cil. ciliated epithelium. 

cl. c. clitellar cells. 

el. clitellum. 

c. m. circular muscles. 

com. commissures. 


c. r. ciliated rosettes or sperm-funnels. 


c. t. connective tissue. 

cu. cuticle. 

d. gl. dark staining glands. 
d. pr. dorsal pore. 

d.v. dorsal vessel. 

e. epithelial cells. 

ept. epithelium. 

gl. glands. 

gl. c. unicellar glands. 


gl. pr. glandular part of prostate. 


giz. gizzard. 

h. heart. 

hy. hypodermis. 

i. s. gr. intersegmental groove. 
l. c. large cells. 

1. m. longitudinal muscles. 
m. muscles. 

m. atr. muscular atrium. 
mes. mesenteric tissue. 

m. pr. muscular prostate. 
n. nephridia. 

n.c. nerve cells. 

ne. nerve ganglion. 


nec. neck of nephridium. 

neph. nephridium. 

neph. p. nephropore. 

@s. esophagus. 

ov. Ovary. 

ovd. oviduct. 

org. sense organ. 

p.c. gl. posterior calciferous diverticulum. 
p. f. posterior fold, nephridium. 
phx. pharynx. 

p- neph. posterior nephridia. 

pr. prostate. 

p- spd. posterior spermduct. 

pr. sto. prostomium. 

ps. penial seta. 

prt. peritoneum. 

se. secreted matter. 

s. int. sacculated intestine. 

sep. gl. septal gland. 

sep. septum. 

s. gl. suprapharyngeal glands. 

s. org. b. sense organ cells in equatoreal zone. 
8. org. pr. sense organ cells in prostomium. 
spe. sperm-cells. 

spd. spermduct. 

spgn. spermatogonia. 

spr. spur of nephridium. 

8. ps. sac with penial set. 

sp. 8. Sperm-sacs. 

s. ph. sgl. subpharyngeal glands. 
spth. spermatheca. 

spth. p. spermathecal pore. 

t. testes. 

t. m. transverse muscles. 

trb. trabecula. 

th. spd. thicker part of spermduct. 
tu. tubular intestine. 

tub. p. tubercula pubertatis. 

ty. typhlosole. 

u.c. unicellular glands. 

v. g. ventral ganglion. 

v. v, ventral vessel. 

v. sgl. ventral gland. 

w. ‘windings ” of nephridial ducts. 


198 


148. 
149. 


150. 


155. 
156. 


157. 
158. 


159. 


160. 


CALIFORNIA ACADEMY OF SCIENCES. 


PLATE. DVI. 


ACANTHODRILUS VASLITI. 
The largest specimen, natural size, but not yet adult; the clitellum not developed. 


Anterior part of a young specimen; clitellum not developed. The nephridia and some other detail not figured. 
s. pl. sgl. subpharyngeal salivary gland. prc. ss. peritoneal cells forming solid masses, probably surrounding 
rudimentary sperm-sacs. prc. large peritoneal cells lining the septa as well as the peritoneum. a. p. pr. 
anterior pair of prostates opening close together. p. p. pr. posterior pair of prostates. 

Part of the ventral part of the body in somites xvii, xviii, xix laid open, showing prostates and thick septa 
with peritoneal cells. m. copulatory muscles. «a. pr. anterior prostate. p. pr. posterior prostate both of 
the same pair. 

Cross-section of the body-wall in somite xix, showing the prostate pore, and the two prostates opening close 
together. 


Section through the same pore, three or four sections further back, showing the tubercula pubertatis, sense 
organ, etc. deb. debris. sup. c. supporting cells. 


Part of a septum, showing the very large peritoneal cells, some of which are broader, others longer and nar- 
rower. : 


Cross-section of the two prostates. p. pr. posterior prostate pore. a. pr.anterior prostate pore. prt. perito- 
neal cells and connective tissue connecting the prostates with the body wall. 


PHCNICODRILUS TEPICENSIS. 
Spermathece, side and front view of one and the same. 


Cross-section of the body in somite xvii through the male pore, showing the copulatory region with muscular 
atrium and spermduct. The longitudinal muscular layer is greatly increased at the male pore. The muscu- 
lar atrium is seen in cross-section, the cut passing through the whole length of that organ. This is also 
seen in figures 158, 159, the latter being taken from a section posterior to the former. 

The oviduct, ovisac and part of the ovaries, interior view of the body. 

Section through muscular atrium in cross-section of the body. The spermducts have not yet united, nor have 
they entered the atrium. The section passes through the whole length of the atrium. The arciform mus- 
cles must have been torn, as they did not show in the section. 

The same parts as seen in fig. 158, but four sections further back, the section passes through the outer layers 
of the atrium. The two spermducts have united and are entering the atrium in line with the longitudinal 
muscular layer. 


Arciform muscles around one of the male pores, viewed from the interior surface of the body. Only one side 
is shown. 


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