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Contributions from the Zoological Laboratory

DESCRIPTIONS

OF

Two New Leeches from Porto Rico

BY

J. PERCY MOORE

Instructor in Zoology

Extracted from U. S. Fish Commission Bulletin for 1900, Volume 2. Pages 211 to 222

Plates 12 and 13

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Contributions from the Zoological Laboratory

DESCRIPTIONS

OF

Two New Leeches from Porto Rico

BY

J. PERCY MOORE

Instructor in Zoology

Extracted from U. S. Fish Commission Bulletin for 1900, Volume 2. Pages 211 to 22a

Plates 12 and 13

19OI

U. S. COMMISSION OF FISH AND FISHERIES,

GEORGE M.. BOWERS, Commissioner.

DESCRIPTIONS.

OF

TWO NEW LEECHES FROM PORTO RICO.

BY

J . P K R C Y A I O O K K

Extracted from U. S. Fisli Commission Bulletin for 1900, Volumo 2. Pages 211 to 222.
Plates 12 and 13.

WASHINGTON:

GOVERNMENT PRINTING OFFICE.

1 9 01.

DESCRIPTIONS OF TWO NEW LEECHES FROM PORTO RICO.

BY

J. F'ERCY IVrOORE,

histrudor in Zoology^ University of Pennsylvania,

268332

211

DESCRIPTIONS OF TWO NEW LEECHES FROM PORTO RICO.

By J. PERCY MOORK,
Instructor in Zoology, University of Pennsylvania.

The material under consideration was collected by the expedition sent with the
steamer Fish Ilawh to Porto Rico in the winter of 1898-99, under the auspices of
the United States Fish Commission.

The leech fauna of the West India Islands is very imperfectly known and no
fresh- water forms have hitherto been described from the island of Porto Rico. Con-
sequently the two species composing this collection both prove to be novelties. One
certainly has and probably both have a much wider distribution.

HIRUDINARIA Whitman (= PCECILOBDELLA Blanchard ('93) subgenus).

mrudinaria was established by Whitman ('86, p. 373) for the Hirudo javanica of Wahlberg, the
generic characters assigned being the large size of the posterior sucker, together with the long interval
(7 J rings) separating the male and female genital orifices. The great increase in our knowledge of the
species of leeches, which we owe so largely to the labors of Blanchard during the last ten years, has
rendered such characters, when taken alone, unavailable for generic distinctions. Accordingly Blanchard
( '97 ) has discarded Ilirudinaria and has referred the type species to his subgenus Pcecilohdella of the genus
Limnatis. Pcecilohdella appears to stand for a very natural assemblage of forms typified by Hirudo
granulosa Sav. and especially characterized by the very striking and constant color pattern, which is
similar in all of the species included by its author. The described species have hitherto been known
only from the tropical and subtropical East Indian islands and Indo-China. Blanchard has indeed
mentioned, but without characterizing, a species from the island of Martinique.^

The leech described below under the name of Mrudinaria hlanchardi maybe the Martinique species.
It has the typical color pattern of Pcecilohdella and resembles Ilirudinaria javanica very closely in
almost all important features of external organization, but the sex pores are separated by but five rings.
I have dissected IL hlanchardi and find, among other peculiarities of the reproductive organs (pi. 12,
fig. 7), that the vagina and the common oviduct open separately into the female bursa (pi. 12, fig. 9).
Professor Whitman has very generously placed at my disposal for dissection one of his specimens of
II. javanica, in which the female organs, though less mature, present the same pecuUarity.

The female organs of Limnatis nilotica (the type species) have been figured by Moquin-Tandon ('46)
and Leuckart ('94). In this species the common oviduct opens into the vagina, the mouth of which,
therefore, becomes the only internal opening into the bursa, as in Hirudo, etc. On the other hand, L.
nilotica resembles //. javanica in the numerous small uniserial denticles and the papilla? which are found
on the sides of the jaws. Limnatis, as understood by Blanchard, is naturally divided by the character of
the female reproductive organs into two genera, the one typified by L, nilotica, the other by H. javanica.
The latter is the Po'.cilohdella group and will probably be found to include all of the species which have
been referred to that subgenus; but for this genus the name Hirudinaria Whitman has priority.

A revised diagnosis of Ilirudinaria based upon an examination of the type species and IL hlanchardi
is as follows: Resembling Limnatis in external annulation, lips, and jaws; the common oviduct and
vagina have separate openings into a small bursa; the segmental sensill«i are large and usually elongated

1 From this island also three nominal species, referred to Hirudo anrl Hscmopis Moq.-Tan., were long ago described
by Moquin-Tandon and Blainville. The habits of at least one of these suggest a Limnatis or Ilirudinana, but there is
nothing in the dosc^-iptions of any of the three sufficient to identify it with the Porto Riean species.

213

214 BIJLLETIlSr OF THE UKTTED STATES FISH COMMISSION,

in shape; the denticles are small and very numerous; some of the pharyngeal glands open on conspicuous
papillse on the sides of the jaws. To this the color pattern as described by Blanchard for Pcecilobdella
will probably need to be added.

Hirudinaria blanchardi, sp. no v.

Diagnosis. — Genital pores separated by five annuli, the male being situated at xi65/6, the female
at XI166/6. Annulation as in H. javanica (Wahlberg) Whitman, except that xxiv65 and 66 and xxv?>l
and b2 are developed (that is, the annuli numbered 95 and 96 in Whitman's figure ('86, pi. xx, fig. 56)
are in the present species each represented by two partially separated rings).

Named for Prof. Raphael Blanchard, of Paris.

External characters. — All of the examples are very much contracted. The type specimen, which
is of medium size, measures as follows: Length, 39 mm.; greatest width (xviii or xix), 11 mm.; depth
at same point, 5.6 mm.; diameter of mouth at base of upper lip, 2.5 mm.; diameter of sucker, 6 mm.
The species reaches a much larger size, probably equaling our Macrohdella decora. The largest example
is 64 mm. long and its greatest width is 18.5 mm., but most of those in this collection are much smaller,
being from 15 mm. to 25 mm. long. "

Owing chiefly to their contracted state the larger examples are short and thick, slightly depressed
behind but becoming more and more terete toward the mouth. The greatest width occurs at the
beginning of the posterior third, in which region the dorsal surface is much more convei: than the
ventral, as indicated in fig. 4, plate 12, by the greater distance m dm than m vm. A strong annular
contraction occurs in all the smaller examples at about somites vm and ix, resulting in the appearance
of a short roimded head united by a neck to the body, which is of a flattened, narrow, elliptical form.
The lips and margins of the mouth are so much contracted that the folds which lie anterior to the jaws,
and in some examples even the jaws themselves, are plainly visible. This would indicate that under
normal conditions the mouth is of large size, as in II. javanica. No distinct clitellum is indicated.

There are 104 annuli, beginning with the prostomium and ending with the postanal annulus at
the base of the acetabulum (pi. 12, flgs. 2 and 3). Owing to contraction they become very short and
crowded at the posterior end, but much less so anteriorly. The annuli are marked off into quadrate
areas, each of which contains one or more pointed papillae and very numerous smaller sense organs
(pi. 12, fig. 4). The larger papillae become very prominent posteriorly and at the margins of the
body. Some further details are added below in connection with a description of the annulation and
metameric seasillse.

The broad upper lip consists of the preocular region, here counted as one somite and one annulus,
together with the first three eye-bearing annuli and the fifth annulus (pi. 12, figs. 1 and 2). Like the
remainder of the body, this region is roughened by numerous non-segmental papillae. A deep longi-
tudinal median sulcus (pi. 12, fig. 1) divides the lower surface of the lip, as in Limnatis, into equal
halves. Posteriorly this sulcus widens into a triangular depressed area from which the median jaw
rises. On each half of the lip are about three less deep sulci about equidistant and extending parallel
to the median one. Between these the lip is nearly smooth, but is studded with numerous minute sense
organs, which are especially plentiful near the anterior margin. The mouth is bounded posteriorly
by the coalesced sixth and seventh annuli (pi. 12, fig. 1).

A full somite separates the sex pores, the male being situated at xi65/6, the female between the
corresponding annuli of xii. Both of these orifices are inconspicuous and not indicated by any eleva-
tions, glandular areas, or pits. The male pore is somewhat the larger, but in no case does the penis
protrude. No clitellum is visible even on the largest specimens. Nephridiopores are found in the
position and number usual in the family (pi. 12, fig. 4, np.). They are placed exactly on the inner
margin of the ventral black band.

The posterior sucker (pi. 12, fig. 3) , like the posterior region generally, is much contracted, but still
large. Its anterior margin reaches as far forward as somite xxiii, so that when relaxed it would probably
have the size and proportions of the corresponding part in II javanica. Dorsally it is marked by six
or eight irregular transverse wrinkles, and is somewhat roughened and bears papillae, as in the body
annuli. Numerous elongated sensillae are also found here, but they vary much in number, size, and
position in different individuals. Ventrally, the sucker in its contracted state is more or less funnel-
form, with a deep central depression and sloping sides which are marked by numerous faint radiating
furrows. The anus is of small size and is situated between somite xxvii and the base of the sucker.

DESCKIPTIONS OF TWO NEW LEECHES FROM PORTO RICO. 215

Annulation. — In the following description of the annulation, the annul us bearing the metameric
sensillge is regarded as the middle one of the complete somite (see Castle, 1900, and Moore, 1900).
The description further serves to show how this view works out when applied to one of the five-ringed
Ilirudinidse. This leech possesses fifteen typical complete somites, being Nos. ix to xxiii, inclusive
(pi. 12, figs. 1, 2, and 3). The five rings constituting each are of approximately equal length, but,
as indicated by the symbols on figures 2 and 3, they have very different morphological values, the
middle annulus being equal to the first plus the second, and to the fourth plus the fifth. (See Moore,
'98 and 1900.)

The structural details of the exterior of the complete somite xv are shown in figure 4, plate 12,
the portion included between the lines dm and m representing the dorsal surface and between vm and m
the ventral surface. Owing probably in part to the accidents of contraction, the annuli are of irregular
length; they overlap somewhat at the posterior margin, and are in places marked by slight wrinkles,
usually running transversely. More definite and constant longitudinal furrows, which tend to alter-
nate in position on adjacent annuli, divide the surface into quadrate areas. According to their size,
these areas bear one, two, or more of the conical sense organs each elevated on a papilla and surrounded
by a grouj) of small goblet-shaped organs and sometimes by smaller conical organs. When not too
much displaced by contraction, these papillae form a range along the middle of each annulus. They
vary much in size in different places and individuals, but are especially rough and prominent at the
margins and posterior end of the body.

The middle annulus, in addition to these non-metameric organs, bears seven pairs of very
conspicuous metameric sensillge which have the large size, the shape, and inclined position so well
represented in Whitman's ('86) figure of H. javanica. Typically four pairs of these are dorsal (pi. 12,
fig. 4), the dorso-median {md), the dorso-lateral (dl), the dorso-marginal (dma), and the supra-
marginal (sm); three pairs are ventral, the submarginal (sbm), the ventro-marginal (vma), and the
ventro-lateral {vl). They are of elliptical outline with a rather prominent axial ridge, along which a
narrow white line (of transparent cells) runs. The dorso-median pair are broadly elliptical, situated
c^lose together with only one quadrate area intervening and inclined to the median plane at an anterior
angle of about 30°. The dorso-lateral pair have the same form, but are of slightly larger size, being
the largest of the segmental sensillse. They are separated from the dorso-median by three or four
quadrate areas, and lie almost exactly midway between the marginal line and the latter. They incline
to the median plane at an angle of approximately 45°. The dorso-marginal are long and narrow and
their inclination to the median plane approaches closely, or even reaches, 90°; they are commonly
separated from the last described by three quadrate areas. Much smaller are the supramarginal
organs, which vary much in shape, size, and position.

The ventral sensillse are all of approximately equal size and similar shape; the ventro-lateral have
an inclination of about 35° and the others of about 90° to the median plane. The ventro-lateral and
ventro-marginal lie closer together and nearer to the median line than the corresponding organs above.
Of course, the angles of inclination vary and in any case can not be measured very accurately, so that the
angular inclinations given are only approximate. The sensillse situated at the margins of the body are
peculiarly inconstant. Frequently one is represented by two or three smaller ones, or may become minute
or altogether suppressed, or two may apparently unite into one, which occupies an intermediate position.
Toward the anterior end the first incomplete somite met with is No. viii. Annuli a2, 55, and 66
are precisely as in the following somites. On the dorsal side 51 and 52 are still distinct, but much
shorter, and the furrow which separates them has become faint. On the ventral side the furrow
extends only a short distance mesiad from the margins, but the two rows of sense organs are apparent
almost to the middle line, at which point the ring becomes entirely undivided, representing al. On
somite vii the corresponding annulus exhibits a slight trace only of the furrow 51/52 on the middle of
the dorsal surface, but two rows of sense organs persist to the margins or even onto the ventral surface.
Annuli 55 and 56 present exactly the same condition as do 51 and 52 of the succeeding somite (viii).
VI is a typically triannulate somite above, but on the middle part of the ventral surface the furrow
between al and a2 has disappeared. The distinction between these two annuli is preserved across
this space, however, by the persistence of two series of sense organs. On the dorsal side the presence
of double series of sense organs on al and aS suggests the growth potentiality of these rings also. On
this somite the sensillge of the dorso-lateral pair become modified as the last pair of eyes, which are
the smallest of the series and have their axes directed outward and backward. Somite v is biannulate
dorsally, witli a faint jmrtial furrow incoivi])letely dividing the anterior annulus into two. Anterior to

216 BULLETIN OF THK UNITED STATES FISH COMMISSlOlSr,

this furrow is a complete transverse series of non-segmental organs, and posterior to it another and
the metameric organs, including a pair of eyes. Ventrally this somite bounds the mouth posteriorly
and is represented by a single annulus. Somite iv is also biannulate. The two annuU are of nearly
equal width, or the first slightly the wider. That the latter represents al and a2 seems evident from
the presence of traces of a furrow passing anterior to the eyes and lateral sensillse, and by the presence
of an additional series of organs anterior to this furrow. The dorso-median sensillse have moved
forward slightly anterior to the traces of this imperfect furrow. Somite iii is very imperfectly
biannulate, the faint cross furrow not continuing to the margins. The non-segmental sense organs,
though reduced in number, form two complete transverse rows. This somite is further interesting
because it bears two pairs of eyes, the outer belonging to the dorso-lateral series and being the largest
and most anterior of^this series. They are crowded toward the posterior margin of the anterior incom-
plete ring. The second pair of eyes represent the dorso-median sensillf5e and have moved forward to
become included in a common pigment mass with the eyes on somite it. Whitman ('92) has described
a similar condition in Clepsine \piacohdella) . The eyes of this sixth pair are of small size and were
detected only upon the examination of sections. Somites i and ii are only imperfectly separated by a
partial and slight furrow, which in many cases appears to be wanting altogether. The first pair of
eyes is situated on the posterior part of the annulus representing ii, but the other metameric sensillse
are very small and diflficult to distinguish from the scattered organs.

Considering the anterior end as a whole it will be seen that the transition from the uniannulate
or even coalesced somites of the prostomium and lip to the complete quinque-annulate somite is a very
gradual one. The appearance of a new ring is first heralded by the gradual separation of an additional
row of non-metameric sense organs, then by the appearance in the dorso-median region of a faint fur-
row, which travels toward the margins, becomes gradually deeper, and creeps around to the ventral
side toward the median line, where it finally becomes complete. These processes are always more
advanced^ in the posterior than the anterior portion of the somite, and in every stage of development,
from the beginning biannulate somite iii to the nearly complete somite vttt, the portion of the somite
posterior to a2 is more fully developed than the anterior portion.

At the posterior end (pi. 12, fig. 3) the series is run through more rapidly, and owing to the
crowding of the annuli the arrangement of the non-metameric organs could not be satisfactorily worked
out. Somite xxiii is complete. On xxiv the two posterior annuli (55 and 66) are short and the
dividing furrow faint or sometimes obliterated in the middle part. Somite xxv is quadri-annulate,
55 and 56 being represented by aS; 51 and 52 are also shorter and not fully developed on the ventral
sides. These two somites, therefore, are more differentiated anteriorly than posteriorly; that is, like
the anterior somites they are developed from the end toward the center of the body. There is a
sudden change from the four rings of xxv to the two of xxvi, and there is here no definite clue to the
whereabouts of the rings (51 and 52) which, as such, are wanting in somite xxvi and in xxvii, which
is precisely similar. These two somites also differ from those which immediately precede them in
their stronger posterior development. They may, therefore, be compared with the anterior bian-
nulate somites iii, iv, and v, in which, the easy transitions show conclusively that the annuli 51 and
52 belong potentially to that portion of the anterior and usually larger annulus which lies anterior to
the metameric sensillge. The almost universal position of the latter on the posterior part of the annulus
gives added force to this view. The resemblance of such biannulate somites to the complete somites
of MicrohdeMa has already been jwinted out (Moore, 1900) , and it need only be added that the persist-
ence of this type of somite at both ends of the leech's body, under mechanical conditions which appear
to stimulate neighboring somites to growth in opposite directions, is significant of a probably phyletic
meaning.

Three and sometimes even all four pairs of the dorsal sensilh^e may be traced serially on the pos-
terior sucker; but they are very irregularly developed, sometimes multiplied by division, sometimes
reduced in number by concrescence or suppression and always variable in position, so that they have
little value in the determination of metamerism (pi. 12, fig. 3).

Color. — The smaller examples only present a distinct color pattern, which becomes more obscure
and diffuse with increase in size. Some specimens have been preserved in formalin, and the colors
are described from these; but they probably have undergone some alteration, so that only the pattern
is significant. Of a specimen measuring 22 mm. in length the ground is a clear reddish clay color,

1 It is assumed for descriptive purposes that the process of development is a progressive one by increasing- complexity
from before backward.

\,

DESCBIPTIOJSrS OF TWO KEW LEECHES FROM PORTO RIOO. 217

becoming a dull orange on the ventral surface, where it forms a continuous uniform area occupying
a,bout two-thirds of the width of the body and entirely uninterrupted by any markings. At the sides
this area is delimited by a pair of sharply contrasting broad bands of dull black, in the borders of
which are scattered some irregular spots of deeper black. External to the black bands, and marking
the exact margins of the body, are two narrower bands of the ground color, but somewhat paler than
the ventral surface. The dorsal color pattern produces a very beautiful effect. Briefly stated, it consists
of five dark longitudinal bands, separated by four of the ground color, which also appears more or less
within the dark bands. Of the five latter the unpaired one is the widest and is divided along the
median line into halves by a regularly broken line of vivid black, which is flanked at intermetameric
intervals by pairs of black spots. A pair of narrower supramarginal bands bear upon their outer
flanks regularly arranged outstanding black spots, between which and the yellow marginal stripes the
ground color becomes of a nearly pure olive. The intermediate bands, or second pair, are still narrower
and are situated about midway between the median and supramarginal bands.

Inasmuch as the color pattern of this species throws much light on the character of the segmenta-
tion of the body a further description, especially of its metameric features, is now given. When fur-
ther analyzed each of the dark bands is seen to be composite, being constituted of longitudinal elements
or narrower lines, which are the result of the greater or less admixture of black pigment with the
ground color. In general the dark pigment is more dense along the margins of the bands, resulting in
the formation of narrow black borders to sooty or clouded olive stripes. The bands are further made
up of serial units, the metameric and intermetameric distribution of which is expressed in each by an
evident tendency to widen in the middle and to shrink or even become suppressed at the ends of
somites. Esch band consists, therefore, of a series of metameric enlargements, alternating with con-
strictions, which are here termed intermetameric because they extend over the contiguous portions of
two adjacent somites (as somites are determined in this paper) . In complete and typical somites the
metameric elements belong to the three middle annuli (62, a2, and 65), while the intermetameric
are confined to the first and fifth (61 and 66) ; but in the entire series the last annulus of one somite
is united with the first of the succeeding somite in respect to color effectiveness (pi. 12, fig. 4).

The metameric elements (fig. 4) are found: (1) In both borders of the middle and intermediate
bands and the ental border of the lateral, all of which become much more sharply defined or of a
deeper black in the three middle annuli and more diffuse and obscure or entirely suppressed on the
terminal annuli. (2) In the median black line, which, as above noted, is not continuous, but formed
of a series of short black dashes, each of which is a bold distinct stroke extending over the three
middle annuli and interrupted by light areas on the first annulus and the fifth. Usually there is no
blending of successive dashes, but sometimes the intervening areas become more or less suffused with
black pigment, which is more likely to occur on the first than on the fifth annulus. (3) In the
black spots which lie on the outer borders of the lateral bands. These are of a deep black color and
occur constantly on the second (62) and fourth (65) annuli, respectively anterior and posterior to
the dorso-marginal sensilloe, around the internal side of which they are connected by a delicate arch
of black pigment. To these positive elements may be added some negative elements: the light spots of
more or less pure ground color which are included within the widened portions of all the bands, viz.,
a pair flanking each segment of the median black line of the unpaired band, a series of similar spots in
each of the intermediate bands, and less distinct ones in corresponding positions in the lateral bands.

The intermetameric elements (pi. 12, fig. 4) of the lateral and intermediate bands are the rather
negative features of the contracted regions on the first and fifth annuli. Here the black borders lose
their intensity, and the black pigment becomes diffused and distributed almost uniformly across the
whole width of the bands, thus separating from one another the light serial spots above mentioned.
In the corresponding parts of the median band the black pigment becomes largely concentrated into a
pair of intense spots, extending over the fifth (66) and first (61) annuli, and including between them
a more or less clear light spot, the repetition of which causes the series of breaks in the median black
line above mentioned.

On the incomplete somites at the anterior end all of these markings may be distinguished, and
as they retain their exact relative positions they afford an important clue to the homology of the
developed annuli. All five of the bands become more distinctly constricted intersegmentally, the
intermediate pair finally breaking into two series of elliptical spots and the lateral similarly into
series of crescents, the horns of which embrace the dorso-marginal sensilhe. On somite vin, in which
the annuli 61 and 62 are incompletely developed, the median black dash extends over a2, 65, and tlie

218 BULLETIN OF THE XJKITED STATES FISH COMMISSION.

posterior half (representing 52) of the first anniilus, while the paired spots are confined to the anterior
half (hi) of this annulus. The black pigment of the intermediate band is entirely, and that of the
lateral band nearly, absent from the fifth ( h 6) annulus, while the anterior portion of the first or
double annulus (61 and 62) is also free from black pigment. On the triannulate somites vii and vi
this arrangement is still more striking, and the exact composition of the first and third annuli is
indicated by the extent of the metameric and intermetameric pigmentation. The median black dash,
the intermediate ellipses, and the lateral crescents on each occupy the middle annulus and about the
adjacent one-half of the third (a3) and two-thirds of the first (al) annuli, while the paired inter-
metameric spots are confined to exactly the remaining fractions of annuli al and aS. These parts
have exactly the values which were assigned on other grounds to 61 and 66 in the composite annuli.
Even on the biannulate somite v the pattern remains; the median dash becomes a mere spot, which
occupies about the posterior three-fourths of the first (al and a2) and the anterior half of the second
(a3) annulus. Anterior to this the median line becomes continuous over somites iv and in to a point
between the first pair of eyes, where it meets the light color of the lip margin. The remains of the
intermediate and lateral bands, and esp»ecially the intermetameric spots of the median band, serve to
define the potential annuli even on somite iv, the last named extending onto the posterior part of iii.
It will be seen that the study of the color pattern of this species confirms to the last detail the values
which were assigned to the annuli as a result of the study of sense organs, integumental furrows, etc.
Further, it afiords support to the neuromeric standard for limiting the somite.

At the posterior end of the body a similar relation between the color markings and the somite
constituents appears to be maintained, but the condition of the material prevents its being satisfactorily
worked out. . The posterior sucker is marked above by an irregular ring of black inclosing a light
spot and flanked by a few black rays; the ventral surface is ash-colored, with a few small scattered
black spots, mostly near the margin.

Small specimens are still brighter in color. The ground is purer and clearer, sometimes paler,
sometimes redder, and the black markings relatively deeper and more vivid. As the size increases
there appears to be a progressive tendency for the black pigment to diffuse over the entire dorsal
surface, changing the ground color to a dull olive and entirely obscuring the markings, though the
more important black spots may always be found. The largest specimen is of a nearly uniform brown
olive, with faint darker cloudings, spots, and broken lines, among which may always be distinguished
the median series of dashes, the paired intermetameric spots, and the lateral spots. They are very
obscure, but sufficient to show the persistent impress of the original pattern.

Alim£ntary Canal. — The mouth cavity is bounded posteriorly by three broad triangular folds, a
single median ventral and a pair of dorso-lateral, together forming a diaphragm, through the limbs of
the trifid aperture of which the anterior edges of the three jaws are usually visible. Slight grooves
run along the bases of the dorsal folds and meet in a triangular expansion of the median labial l : -'^ns.

Except that the median dorsal jaw is somewhat lower, the three jaws (pi. 12, fig. 5) are of simiiax
shape — long, low, and compressed, with the edge only slightly convex. They are strongly angulated
anteriorly, where they pass abruptly into the supporting plate, but recede more gently behind. The
denticles (pi. 12, figs. 5 and 6) are very small, triangular plates set transversely on the jaws, and
number from 160 to 180 on each jaw in the two examples which were examined. They are largest
anteriorly, where they measure 0.03 mm. in height, and diminish gradually to one-half that height
at the posterior end. Along the sloping sides of each jaw are considerable numbers of button-shaped
papillae supported on narrow stalks. They vary in size, the largest being about 0.07 mm. in diameter,
the smallest about one-fourth that size. Most of them are arranged somewhat in three irregular rows
(fig. 5) of about ten each, between which some smaller ones are scattered. They appear not to be
sense organs, but serve as places of exit for some of the ducts of the pharyngeal glands, most of which
traverse the interior of the jaw and open on its ridge between the teeth. Possibly it is due to the
great number and crowding of the denticles that additional outlets are required for the glands.

The muscular oesophagus is marked by six longitudinal folds, the three which correspond to the
jaws being larger than those which alternate with them. It reaches to somite ix. There are ten
pairs of gastric cseca developed, as in Ilirudo, The last originates in xix, is large, sacculated, and
extends caudad. None of the smaller Cc^eca are much sacculated. That this species is a true blood-
sucker is shown by the presence of blood in the caeca of all specimens examined.

Reproductive Organs. — A general view of the reproductive organs is shown in pi. 12, fig. 7. The
male organs do not differ in any important respect from IRruxfo, though tlie sperm sa(^s {de) are larger.

r>ESCRIPTI03NrS OF TWO NEW LEECHES FROM PORTO RiOO. 219

There are eleven pairs of testes {t 1-t 11) resting on the ventral longitudinal muscles on each side of the
nerve cord and immediately behind their corresponding ganglia. The sperm ducts (vd) lie on the
floor of the body just external to the testes, and, owing in part to the contraction of the body, are
greatly convoluted. At the posterior end they extend beyond the last pair of testes as short, blind
tubes, but a sufficient number has not been dissected to show that this is a constant feature. At the
anterior end the ducts become much smaller, less glandular, and less convoluted. In somite xi they
bend back on themselves and immediately become wide, irregular, closely convoluted tubes, forming
rather compact masses, the epididymis {ep). At their posterior end these again pass into the small
ends of the conspicuous fusiform, muscular-walled sacs, the ducti ejaculatorii, or sper^ sacs (de).
Finally the two sperm sacs unite in a common penis sheath (figs. 7 and 8 ps.), which is a thick,
muscular, pyriform sac bent sharply forward on itself and attached to the ventral body wall at the
male orifice. There is a long coiled-up penis, but in no cases was it found to be protruded. Prostate
glands are scarcely evident, only a few solitary unicellular glands opening into the bulb of the penis
sheath.

The pecuharities of the female organs (pi. 12, figs. 7 and 9) have already been mentioned in the
generic description. The vagina (vg) is cylindroid in younger specimens, but becomes much enlarged
at the blind end, with a relatively narrow neck (fig. 9) in fully mature examples, in which also it may
become doubled on itself. The two ''ovaries" (ov) lie about equidistant from the middle line and
their ducts empty into a common oviduct with a small albumen gland {gla). The common duct is
short and folded into a compact coil which is bound to the anterior wall of the vagina, with which it
opens, but by a separate orifice, into a small and probably eversible female bursa {h).

The female organs of JJ. javanica are shown on pi. 13, figs. 18, 19, and 20. The specimen,
although much larger than that from which fig. 7 was drawn, is much less mature. The structure
of the organs is in every respect similar. The vagina is marked by longitudinal wrinkles and it pro-
jects upward almost vertically from the ventral body wall, carrying the nerve cord with it.

Several specimens (about fifteen) of various sizes were taken from a bottle labeled '' Cagua 1-10-99."

DIPLOBBELLA, gen. nov.

The leech for which this genus is proposed belongs to the monostichodont Eirudmidse and differs
in several respects from all members of this group whose internal anatomy has been described. The
sperm ducts of the two sides remain entirely distinct almost to the external orifice, with which they
are connected by a very short canal. There is no definite and distinct penis sac, and if the common
canal is eversible at all it can form only a very short penis. There is a remarkable development of
gastric c^ca, most of the complete somites as far back as xviii having two pairs in place of the one
usually present. These are^ always very obvious and the two pairs are of equal length in some of the
postgenital somites. There is frequently a tendency for most of the annuli to become faintly
subdivided. (See pi. 18.) This disposition toward the doubling of organs has suggested the name.

The nearest ally of Diplobdella known to me appears to be Philobdella VerrilL This genus has the
sperm ducts similarly separated almost to the external orifice, the prostate glands very large, the jaws
high and few toothed, and the color pattern similar. But it differs in the complicated copulatory
glands and pits, in the much simpler lobing of the stomach, the form of the ovaries and vagina, and,
in the known species, in the location of the sex pores and some details of annulation. Macrohdella
Verrill possesses the two pairs of gastric c^eca per somite, but the male reproductive organs are quite
dissimilar.

None of the numerous descriptions of Hirudmidm from South and Central America, etc. , appear to
fit the type species, which is therefore described as

Diplobdella antellarum, sp. nov.

Diagnosis,— The sex pores are separated by 3J rings, the male being situated at ^n, the female at
the middle of xii 65. There are about 35 denticles on each jaw. Annulus xxvi al is developed at
the margins. Annulus iv aS is frequently imperfectly differentiated. The colors are plain ventrally,
longitudinally striped above.

External Characters.—When fully extended this leech is slender and the sides of the body nearly
parallel for a long distance, as in a nephelid; when strongly contracted, which is the state of most of
the specimetis, the body is nearly elliptical w4th bluntly rounded ends. Whether contracted or

220 BULLETIN OF THE UNITED STATES FISH COMMISSION,

extended, the body is always flattened, quite unlike the thick rounded form assumed by Hsemopis, etc.,
when contracted or resting. The largest extended specimens are over 3 inches long. The selected
type specimen has the following dimensions: Length, 55 mm.; greatest width (about xviii), 8.5 mm.;
greatest depth, 3.8 mm. (about); diameter of sucker,'5.4 mm.; width of upper lip at base, 2.3 mm.

There are 105 distinct annuli, counting one which appears posterior to the anus. In many
specimens most of them are marked by a transverse depressed line, making them faintly double. The
annuli are smooth, in no case exhibiting the prominent papillae so characteristic of the species last
described, although in some cases examples of the two species had been killed and preserved together.

In marked contrast to Hirudinaria Uanchardi the mouth is relatively small. The upper lip is
slightly wrinkled on the margin, but nearly smooth below, and undivided by a median sulcus except
at the base. Dorsally it is seen to consist of the first four somites, the coalesced annuli of the fifth
bounding the mouth posteriorly.

The five pairs of conspicuous eyes (pi. 13, fig. 10) are disposed as usual in the family. The dif-
ferent directions toward which their pigment cups face is especially evident. The first pair look
directly forward, the second forward and slightly outward, the third outward, the fourth outward and
backward, and the fifth backward and slightly outward.

The male pore is situated on a small, more or less prominent cone rising from the contiguous
parts of somites xi and xii, between which the orifice lies. In no case does a penis protrude, and
there are no copulatory glands. The female pore opens through the middle of the fourth annulus
(65) of somite xii. (PI. 13, fig. 12.) None of the specimens show a clitellum.

The anus has the usual position behind somite xxvii, but xxviii appears as a distinct ring bound-
ing it behind. The sucker is relatively small, thin, very flat, rather narrowly attached, and much
more broadly free posteriorly than anteriorly (pi. 13, fig. 11). The usual 17 pairs of nephridiopores
appear as short oblique slits on the posterior margins of annulus 52 of every somite from viii to xxiv,
inclusive (figs. 10 and 11). The first 16 pairs are just ectad of the ventro-marginal sensilbe; the last
pair is peculiar in being distinctly more mesiad.

Annulation. — The details of annulation are sufficiently shown in the diagrams (pi. 13, figs. 10, 11).
Attention is directed to the following points: Somites ii and in are very short, with no trace of more than
one annulus each. The second annulus (a3) of iv is sometimes very imperfectly separated. On the ven-
tral surface v is uniannulate and vi biannulate. viial is very large, but always entirely undivided;
XXVI al is separated at the margins at least, the first annulus {a\ + <^2) of xxvii is very broad, and
its sensilke are far posterior, and, lastly, xxviii appears as an annulus distinct from the sucker.

The metameric sensillse (pi. 13, figs. 10 and 11) are small and round as in Hsemopis and
Hirudo medicinalis, but are generally easily found. The dorso-lateral are much larger than any of the
others. Their arrangement is shown in the figures, the three ventral pairs on somite xiv (fig. 10).
Sensillse are very numerous on the dorsal surface of the sucker, all of the eight dorsal series being well
represented. As usual, they are variable in number and position and show evidences of multiplication.

Color. — The pigmentation is described from formalin specimens, which have probably faded
considerably. The colors are very quiet and the pattern simple, but both tint and arrangement
vary greatly. The ventral surface is of a uniform ash, which sometimes has a distinctly reddish tint,
in which case there is likely to be a slight submarginal band clouded with black. A narrow yellowish
band varying in intensity extends along each margin. Dorsally, the ground color may be just like
the ventral or it may be darker or tinged with yellow. In a typical example there are three pairs of
dark longitudinal bands, which together occupy about two-thirds of the width of the dorsum and
unite at the ends of the body. They are of an olive color bordered by narrow margins of brown. The
innermost pair are the broadest and are separated by a narrow but distinct line of yellowish. The
intermediate pair is slightly and the lateral pair much narrower, and the latter is frequently broken
and interrupted by intrusions of the ground color. Between the dark bands the intervening strips of
ground color are very narrow, and broken here and there by bridges of the dark pigment, which cross
more frequently between the lateral and intermediate bands. Such connections are most likely to
occur on the fourth annuli of somites, but the pigmentation of this species appears to have no constant
metameric characters. All of the dark bands coalesce in a single spot of pigment which surrounds
the anus and extends onto the sucker in the form of a crescent.

The extreme of color on the one hand is found in a type in which the pigmented bands become
of an inky black, the median yellow line is almost obliterated, and an additional dull black supra-

DESCRIPTIONS OF TWO NEW LEECHES FKOM PORTO RICO. 221

marginal line apx)ears, and on the other in an example in which the dorsal markings have become
very o})scure and pale, so that the entire animal is of a nearly uniform reddish ash color. Intermediate
examples are found.

Alimentary Canal. — The jaws are much like those of Hirudo, conspicuous, high, short, and com-
pressed, bearing a relatively small number (about 35) of denticles arranged in one series (pi. 13, fig. 13).
They are much larger on the anterior part of the jaw, becoming smaller and relatively broader posteri-
orly, where they are succeeded by several imperfectly developed ones. All of the functional denticles
bear recurved calcareous tips. There are no lateral papillae, but all of the gland ducts empty on the
jaw ridge between the teeth (fig. 13).

A very remarkable feature of the alimentary canal is the extensive development of gastric ca^ca.
Each complete somite from viii to xviii contains two pairs instead of the one pair only, which is
usual. In the more anterior of the somites named these cseca are of relatively small size and are
irregularly developed, but in the more posterior the caeca of the two pairs are of equal and large size,
and considerably lobed. The last caeca arising in xix, as usual in the Hirudinidx, are very large and
extend posteriorly by the sides of the intestine into somite xxiv (pi. 13, fig. 17).

Re'productive Organs. — Ten pairs of testes (pi. 13, fig. 14, tl-tlO) occur in as many somites (xi to xx).
The last are very small and terminate the sperm ducts. The latter (vd) have a narrow muscular (?)
section in somites xi and xii. The epididymes (ep) are large curved masses which open into sperm sacs
(de). The sperm sacs themselves (figs. 14, 15, and 16, de) are comparatively small, and present the
remarkable feature of being closely bound together by muscular and especially by connective tissue
sheaths. Moreover, they are concealed from above by a thick layer of prostate glands (pgl) which com-
pletely cover them dorsally, laterally, and posteriorly, and which open by numerous ducts into the
sperm sacs themselves. In a dissection the epididymes appear to terminate in a somewhat massive
median body which, by sectioning, is found to include the sperm ducts,-as just described. The common
median portion of the sperm ducts, corresponding to the penis sheath of most liirudinidsa^ is a very
short atrium with yqyj thick muscular walls {mh) and a rather wide lumen lined by a somew^hat
folded epithelium (m). There is no indication of a long filiform penis, such as occurs in most jawed
leeches, and while the lining of the atrium is probably eversible, it could form only a very short, rather
wide penis. The structure of the whole region would appear to adapt it better to the production and
placing of spermatophores, as in Glomphonia, rather than for copulation as practiced by Ilirudo.

The female organs are like those of Hirudo, etc. The vagina {vg) and slightly folded common
oviduct {ode) lie to the right of the nerve-cord, ventral to which the left oviduct crosses to join the
latter.

Like IL blanchardi, this species is a blood feeder, as is evidenced by the presence of blood in the
cgeca, etc. A large number of specimens were collected at San Juan on January 12, 1899, and at Cagua
on January 10, 1899. It also occurs on the American continent, as I have received from Professor
Harold Heath, of Leland Stanford University, examples collected by R. 0. McGregor in Jaxiuary,
1896, at Panama, Colombia. The drawings of the internal anatomy were made from the latter.

LIST OF PAPERS CITED,

Blanchard, R., 1893. Revision de tlirudinees du Musee de Turin. Bollettino dei Musei di Zoologia
ed Anatomia comparftta della R. Universita di Torino, viii (1893), 1-32.

Blanchakd, R., 1897. Hirudinees du Musee de Leyde. Notes from Leyden Museum, xix (1897),
73-113.

Bayek, E., 1898. Hypodermis und neue Hautsinnesorganeder Rhynchobdelliden. Zeitschrift f. wis-
senschaftliche Zoologie. lxiv (1898), 648-696.

Castle, W. E., 1900. The metamerism of the Hirudinea. Proceedings American Academv of Arts
and Sciences, xxxv (1900), 285-303.

Leuckart, R., 1894. Die Parasiten des Menschen. 2d ed. Leipzig, 1894. Bd. v. Leif., 5.

Moore, J. Percy, 1898. The leeches of the IJ. S. National Museum. Proceedings United States
National Museum, xxi (1898), 543-563.

Moore, J. Percy, 1900. A description of Microbdella biannulata, with especial regard to the consti-
tution of the leech somite. Proceedings Academy Natural Sciences Phila., 1900, 50-73.

Moquin-Tandon, 1846. Monographic de la famille de Hirudinees. 2d ed. Paris, 1846.

Whitman, C. O., 1886. The leeches of Japan. Quarterly Jour. Micro. Science, xxvi (1886), 317-416.

Whitman, C. 0., 1892. The metamerism of Clepsine. Festschrift zum 70ten Geburtstage R. Leuckart^
(1892), 385-395.

222 BULLETIN OF THE UNITED STATES FISH COMMISSION.

DESCRIPTION OF FIGURES.

Plate 12.

Figs. 1 to 9 represent Ilirudinaria blanchardi.

Figs. 1, 2. Ventral and dorsal views, respectively, of a typical large specimen, X 5. The somites
are indicated by Koman numerals on the left; the numbers on the right are of the annuli,
counted from the prostomium, while the symbols included within parentheses show their
theoretical value. The eyes and metameric sensillse are shown in heavy black, the ^'goblet-
shaped sense organs" and the "conical sense organs" as small dots. Except that all of
the goblet-shaped organs are not represented, the position and number of the sense organs
is indicated as accurately as possible. An attempt has also been made to have the boldness
of the lines correspond with the depth of the interannular furrows.

Fig. 3. A semidiagrammatic view of the dorsum of the posterior end of the same specimen, X 5. The
metameric sense organs only are represented. The dotted line across annulus xxiv a3,
indicates its partial subdivision into two. The somites and annuli are indicated as in fig. 2.

Fig. 4. Slightly more than the right half of somites xv and xvi of the largest specimen, X 3. The
whole surface, dorsal and ventral, is shown as though flattened out, and every topographical
feature of xv has been drawn with the greatest care. On xvi and on the middle portion of
XV the color pattern of a small specimen has been plotted to show its relation to the sen-
sillse, etc. The intensity of the black pigment is indicated by the closeness of the stippling.
dm, dorsi-meson; w, extreme lateral margin; vm (the exact margin of the figure), the ventri-
meson; 7ip,_nephridiopore; md, median-dorsal; dl, dorso-lateral; dma, dorso-marginal; sm,
supra-marginal; sbm, submarginal; vma, ventro-marginal; and vl, ventro-lateral sensillse.

Fig. 5. A median dorsal jaw, X 33. Outline as seen from the right side. The teeth and papillge are
shown.

Fig. 6. Transverse section of a jaw shown in outline, X 33. The cut is near the anterior end, and
shows one of the larger denticles and the central mass of gland ducts which open partly on
the dentigerous margin, partly on the lateral faces of the jaw. d, denticle; p, papillse; gd,
axial mass of ducts.

Fig. 7. Partial representation of the reproductive organs from the dorsal aspect, X 6.5. On the left
side the first testis only is shown, and the ductus ejaculatorius is drawn forward to show
the epididymis, tl-tll, the eleven testes; vd, the vas deferens; de, ductus ejaculatorius
(sperm sac); ep, epididymis; ps, penis sheath (atrium); ov, ovary; ode, common oviduct;
vg, vagina.

Fig. 8. Outline of terminal portion of male duct, seen from the left side, X 6.5. ^, male pore; other
letters as in fig. 7.

Fig. 9. Outline of the female organs of the largest specimen, somewhat dissected, X 6. 5. lov and rov,
left and right ovaries; g/a, albumen gland; ode, common oviduct; vg, vagina; b, female
bursa; 9 ? female pore.

Plate 13.

Figs. 10 to 17 represent Biplobdella antillarum.

Figs. 10, 11. Semidiagrammatic figures of the anterior and posterior ends of the body viewed from

the dorsum. X (about) 5. The somites and annuli with their values are indicated as in figs.

1, 2, and 3, but the metameric sensillse are indicated by circles. On somite xiv the ventral

sensillse are shown. Sensillse lettered as in fig. 4.
Fig. 12. The sex pores and neighboring structures. X (about) 5.
Fig. 13. A left lower jaw shown in outline and as a transparent object. X 56. The denticles, the

course of the gland ducts from the central mass, and the arrangement of some of the muscles

are shown.
Fig. 14. Reproductive organs from above. X 6.5. The left testes are omitted, and the lettering is as

in fig. 7, except pgl, prostate glands.
Figs. 15, 16. Two transverse sections through the atrium. X 26. The section shown in fig. 15 is

fiNQ sections anterior of 16. $ , male pore opening from bursa; m, median common part of

duct opening into male bursa; m5, muscular wall of atrium; de, ductus ejaculatorius; ep,

epididymis; pgl, prostate glands; Im, longitudinal muscle layer; em, circular muscle layer;

71, nerve cord.
Fig. 17. Outline of alimentary canal. X (about) 5. The Roman numerals indicate somites, and show

the extent of the several regions.
Figs. 18, 19, and 20 are, respectively, nearly anterior, ventral, and right lateral views of the female

reproductive organs of Hiriidinaria javaniea, X 10. Lettering as in fig. 9.

Bull. U. S. F. C. 1900, Vol. 2.— J. P. Moore. (To face page 222.)

Plate 12.

Bull. U. S. F. C. 1900, Vol. 2.— J. P. Moore. (To face page 222.)

Plate 13.

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