SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME
98,NUMBER
17POLYCLAD WORMS C0LLEOrep
,B6W " 39
THE PRESIDENTIAL CRUIS^'"* mm
OF 1938
BY
LIBBIE H.HYMAN
American Museum of Natural History
NewYorkCity
(Publication 3542)
CITY
OF WASHINGTON
PUBLISHED
BY THE SMITHSONIAN
INSTITUTIONJUNE
17, 1939SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME
98,NUMBER
17POLYCLAD WORMS COLLECTED ON
THE PRESIDENTIAL CRUISE
OF 1938
BY LIBBIE H.
HYMAN
American Museum of Natural History
NewYorkCity
(Publication 3542)
CITY
OF WASHINGTON
PUBLISHED
BY THE SMITHSONIAN
INSTITUTIONJUNE
17, 1939Z§t£orfcQ0afttmore (prees BALTIMORE, MS., D.S.A.
POLYCLAD WORMS COLLECTED ON THE PRESIDENTIAL CRUISE OF
1938By LIBBIE
H.HYMAN
American
Museum
of Natural History,New
York CityThe
material consisted of four vials of polyclacls collected at vari- ous points alongthe shores of the Galapagos Islandsand Old
Provi- denceand
Clipperton Islandsby
Dr.Waldo
L. Schmitt during the Presidential Cruise of 1938.Each
vial contained a different species represented in three vialsby
a single specimen, in the fourth vialby two
specimens. All of theworms were
badlycrumpled and
foldedand
itwas
necessaryto straightenthem
out forcibly beforeany work
could be
done on
them. This has resulted insome
distortion of partsand
hasproduced
foldsand
wrinkles in thebody
surface, but the material is adequate for the determination of the necessary points of anatomy. All of the specimenswere
stained aswhole mounts
with dilute acidifiedborax
carmineand, aftertheywere
studiedand
drawn, the sexual regionwas removed
in the case of three speciesand
cut into sagittal serial sections.The
studyof the material hasshown
thatit comprises
two
species ofN
otoplana, one of Euplana,and
one ofProsthiostomum, all of
which
appear to be undescribed forms.JfOTOPLANA INSULARIS,
n. sp.Figs. 1-3
Description.
— The
singlespecimen,somewhat
contractedand
folded(fig. 1), is 9
mm.
longand
3mm. wide
at the widest part, obviouslymuch
longer in life, of elongated oblong shape, taperingsomewhat
posteriorly. Colornot determinable.
Pharynx
small with afew
folds,embraced by
the uteriwhich come
to a point anteriorly behind the brain.Eyes
(fig. 2) in paired elongated bands, tentacular clusters thus not separatedfrom
cerebral groups; tentacular groups evident asrounded group
in eachband
at about the level of the middle of the brain.Eyes
unusual in that cerebral eyes are of thesame
size as tentacular eyes. Posterior half of specimenmade
into serial sections.Copulatory apparatus (fig. 3) typical of the genus. Seminal vesicle oval, with thick muscular wall of fibers parallel to the surface con- tour. Ejaculatory duct curves
backward and
after a short courseSmithsonian Miscellaneous Collections, Vol. 98, No. 17
2
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL.98
enters the oval prostate vesicle,which
is smaller than the seminal vesicle. This has a moderately thick muscular wall of fibers parallel to the surface contour, penetratedby
the ducts of the extracapsular prostate glands. Ejaculatory duct projects into prostate vesicle for about three-fourths the length of the latter.No
penis papilla.A
relatively short stylet springs directly
from
the prostate vesicle ; it lies in a moderately long tubularmale antrum which
curves grace- fully ventrallyand
posteriorlytothe malegenital pore.Male antrum
with a muscular wall of inner longitudinaland
outer circular fibers.Female
pore shortly behind male pore; canal leading infrom
surface probably artificial causedby
straightening of the specimen.Vagina
ofmoderate
width with coating of circularmuscle fibers; curves pos- teriorlyand
widenswhere
itreceives theuteri;beyond
this,wide
stalk of Lang's vesicle curvesdown,
then posteriorly,and
opens into the very long large Lang's vesicle.Remarks. — Noto
plana insularis does not closely resembleany
of theknown
speciesfrom
adjacent islands of the Caribbeanand West
Indies, but
seems more
nearly related toN.
stylifcrafrom Juan
Fer- nandez,from
which, however, it differs in several details,which
will be apparenton
comparison ofmy
figures with those of Bock.Differential diagnosis.
—
Tentacularand
cerebral eyes in one band,cerebral eyes of
same
size as tentacular eyes, seminal vesicle larger than prostate vesicle,no
penis papilla or penis sheath, penis styletshort,
male antrum
tubular, vaginawidened
at entrance of uteri.Locality.
—
Shores,Old
Providence Island. Collected by Dr.Waldo
L. Schmitt,
August
6, 1938.Holotype.
—
Anterior half aswhole mount
; posterior half as sagit- tal serialsections,U.S.N.M.
no. 20423.NOTOPLANA CARIBBEANA,
n. sp.Figs. 4-8
Description.
— One
large specimen, 33mm.
long, 13mm. wide
atwidest part, of elongated
wedge
form, widest anteriorly, tapering gradually to obtuse posteriorend
(fig. 4). Color not determinable.Eyes
in paired bands inwhich
the tentacular clusters are included (fig. 5).Some
indications of tentacles in center of tentacular eye clusters; tentacular eyes obviously larger than cerebral eyes; cerebral eyes small,forming
elongatedgroup
anterior to tentaculargroup
andfew
scattered eyes behind tentacular group.Pharynx
long, very nar- row, withmany
small folds,embraced by
uteriwhich come
to a point in front ofpharynx
(fig. 4). Sexual regionremoved and
cut intoNO. 1/
POLYCLAD WORMS HYMAN
3sagittal serial sections.
Male and
female pores considerably sepa- rated,forming
large oval openings. Crescentic seminal vesicle im- mediately behind posteriorend
ofpharynx
has very thin wall of muscle fibers paralleling its external contour. Seminal vesicle opens directly into large sphericalchambered
prostatic vesicle without the intervention of a duct; ejaculatory duct projects only a short dis- tance into prostatic vesicle (fig. 6). Small penis papilla at beginning ofmale
canal leadingaway from
prostate vesicle. This canal, similar topenis pocket of other Notoplanas, is remarkably longand
slender;
runs
backward
above excessively muscularmass
surrounding the maleantrum and
opens intoposterodorsal angleof thisantrum.Male antrum
largewith longanterior extensionand
linedby
atallepithelium filledwith granules of glandular nature;male antrum
surroundedby
thick powerful muscle fibers running mostly lengthwise filling allspace
between
antrum, prostate vesicle,and
penis pocket. Posterior wall ofantrum
also muscular.Antrum
opens belowby
large oval male genital pore.Large
oval female pore considerably behindmale
pore (relative positionsshown
in fig. 4).Pore
opens intowide
funnellike femaleantrum
(fig. 7) with very folded walls; this appearancemay
be due tothe straightening of the specimen. Tall liningepithelium of femaleantrum
has glandular appearance.Vagina
proceeds anteriorlyfrom
female antrum,makes
sharp posterior bend,and some
distance be-yond
thebend
receives the uteri.Beyond
uteri long slender stalk of Lang'svesiclerunsbackward and downward and
opens into elongated oval Lang's vesicle.Vagina and
stalk linedby
ciliated epithelium, outside ofwhich
is afibrous coatwhich
does not appear tobemuscu-
lar.
Numerous cement
glandsopen
into vaginaand
beginning of vesicle stalk. Lang's vesicle linedby
very tall epithelium, the distal two-thirds ofwhich
is filled with eosinophilous spherules (fig. 8).These
suggest an eosinophilous secretion, but it ismore
probablethat they are digesting material. Lang's vesicle probably serves to digest excess sex cells as well as for a seminal receptacle.Remarks. —
This species undeniably bears a very great resemblance insize,shape,and
thedetailsof thecopulatory apparatustoNoto
plana binoculata (Verrill) 1901 (syn.Noto
plana bahamensisBock
191 3)of
which Bock
has given agood
description, supplementedby some remarks
ina recent paperofmine (Hyman,
1939).The
similarityofN.
caribbeanatoN.
binoculatais not surprising, in view of the prox- imity ofOld
Providence Island to theBahamas, and
there still re-mains
a little doubt inmy mind
that Iam
justified inmaking
a dis- tinct species ofN.
caribbeana.The
following differencesmay
be4 SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 98 noticedbetween
thetwo
species. InA
T. caribbeana the cerebral eyes aremuch
smaller than the tentacular eyesand on
thesame
level,whereas
inN.
binoculatathey are of nearly thesame
size as the ten- tacular eyesand
so deeply buried that theywere
overlooked alto- getherby
Verrill(whence
thename
binoculata).The
seminalvesicle of JV. caribbeana ismuch
largerand
thinner walled than that of JV. binoculata, itsmale
canal is longerand
the maleantrum
is of dif- ferent shapeand
lacks thetwo
hillocksfound
in JV. binoculata towhich
Ihavecalled attention.The
female apparatusofthetwo
speciesis very similar, but in JV. caribbeana the vagina
and
stalk of Lang's vesicleappeartobelongerthaninJV. binoculata(althoughthesediffer- encesmay depend
simplyon
the sizeof thespecimen)and
the cement glands extend farther along the female apparatus than in JV. binocu- lata.These
differencesseem
to justify the separation of theforms
as distinct species.Bock
has placed JV. binoculata in hisgroup
B, typifiedby
JV. atomata, but admits that it is not closely related to themembers
ofthisgroup. Isuggest thatJV. binoculataand
JV.caribbeana be considered to constitute a separategroup
of the large genusN
otoplana, characterizedby
the long male canaland
the large extremely muscular male antrum.Differential diagnosis.
— N.
caribbeanadiffersfrom
otherNotoplanas exceptA
T. binoculatainthelong slendermalecanal, largemaleantrum
with an anterior extension,and
excessive muscularity of the anterior wallof themale
antrum,whose
musclefibers fill allthe spacebetweenantrum and
seminal vesicle. Differencesfrom
JV. binoculata are listedabove.Locality.
—
Shores ofOld
Providence Island. Collectedby
Dr.Waldo
L. Schmitt,August
i, 1938.Holotype.
— Whole mount
with sexual regionremoved;
set of sagittal sectionsof sexual region,U.S.N.M.
no. 20424.EUPLANA CLIPPERTONI,
n. sp.Figs. 9-12
Description.
— Two
specimens, larger 17mm.
long, 7mm.
wideacross widest part, not quite fully
mature
sexually, elongated, widest at about the level of the brain, tapering posteriorly,much
foldedand
contracted, evidentlymuch
longerand more
slender in life (fig. 9).Smaller specimen very young, 6.5
mm.
in length, obovate, withno
trace of sex organs. Color not determinable.
With
evident tentacular groups of eyes, consisting of 3 large eyesand
4-6 smaller ones(fig. 10). Cerebral eyes loosely arranged, scattered, radiating in a
NO. 1/
POLYCLAD WORMS HYMAN
5linear
arrangement
along the principal nerve trunks (fig. 10).Of
the cerebral eyes, there aretwo
large ones close to the cerebralmass
of granules characteristic ofmany
Acotylea,and some
large ones in arow
along the bases of themain
anterolateral nerve trunks (fig. 10).The
smaller eyes are peripheral to the large ones, radiatingalong the nervetrunks. Smallspecimen (fig.n)
hassame number and
arrange-ment
of large eyes as the larger specimen, but the smaller eyes have not yet appeared.Pharynx
relatively small with small lateral folds.Sexual region of larger
worm removed and
sectioned. Conclusionfrom
study ofwhole
animalthat sexual maturityhad
not quite been attained confirmedby
study of sections, but essential parts appear to be present. Sexual apparatus close behind pharynx, farremoved from
posteriorend
of body. Sexual poressome
distance apart(fig.9).
Male
copulatory apparatus (fig. 12) typical of thegenus,i.e.,themale canalis not definitely divisible into seminal vesicle
and
pros- taticvesicle,and may
be considered toconsist entirely of the seminal vesicle.Vasa
deferentia enter separately the lateral angles of the seminal vesicle; seminal vesicle then turns dorsally, then bends pos- teriorlyand
extends as slender tube to penis papilla. Seminal vesicle hasnarrow lumen and
thick muscular wallcomposed
chiefly of circu- lar fibers.Xo
prostatic glands seen, probably because of immaturity of the specimen. Penis papilla small, rounded,armed
with a short stylet in process of being secreted. Penis papilla lies in short penis pocketwhich forms
a penis sheathwhere
it joins themale
antrum.Lattermoderatelylarge cavity, of
moderate
length.Female
apparatus(fig. 12) has large genital pore, long muscular vagina bent into an S-shape,receiving
cement
glands along itscourse. Followingentrance of uteri, female canal continues as a small Lang's vesicle,which may
not yet be fully formed.
Remarks. —
Inan
articleon
Atlantic coast polyclads(Hyman,
1939 b), I haveshown
that"Prosthiostomum"
gracile Girard 18501is not a
Prosthiostomum
at all, but fits into DiscoplanaBock
1913.However,
since Girard in 1893 himselfremoved
this speciesfrom Prosthiostomum and
created for it anew
genus Euplana, Discoplanamust become
asynonym
of Euplana.The
copulatory apparatus of1In a recent publication Pearse (1938) has grossly misidentified Euplana gracilis (Girard) consideringitanewspecies,typeofanewgenus,Conjuguterus.
Conjuguteriis becomes a synonym of Euplana and C. parvus a synonym of Euplana gracilis. Pearse's account of the male system of this species is also erroneous. These corrections are based on an examination of the type speci-
men of C. parii<s and a number of other specimens labelled by Pearse with this name.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL.98 Euplana (=Discoplana)
isvery similarto thatof thegenus Stylocho- planaand
in factit isdifficulttodistinguishbetween
thesetwo
genera.The
sole difference is that inEuplana
there isno
distinct prostatic vesicle,whereas
one exists in Stylochoplana. E. clippertoni certainly resembles Stylochoplana in general external appearance, but hasno
definiteprostaticvesicle
and
henceseems
to fit better intoEuplana. It differsfrom
otherknown
species ofEuplana
{•=Discoplana) in the presenceofapenisstylet.Euplana
gracilis (Girard) isthemost
simpli- fiedmember
of the genus, havingno
penis papilla, stylet, or penis sheath.Differential diagnosis.
—
Loosely arranged radiating cerebral eyes, small penispapilla,penis pocket, penissheathand
short styletpresent, longmuscular vagina with S-bend, small Lang's vesicle, genital pores well separated.Locality.
—
Clipperton Island,under
rocksto south of landingplace.Collected
by
Dr.Waldo
L. Schmitt,July 21, 1938.Holotypc.
— Whole mount
with sex regionremoved
; set of sections of sex region,U.S.N.M.
no. 20425.Young
specimenon same
slide with holotype.PROSTHIOSTOMUM PARVICELIS,
n. sp.Figs. 13-15
Description.
— One
specimen,6 mm.
long, of usual slender shape typical of the genus, fully mature, bent in sexual region, part of an- teriormargin
missing (fig. 14). Color not determinable.Eyes
as usual in the genus, comprising paired cerebral groups,and band
of eyes along anteriormargin
(fig. 13). Cerebral eyes consist ofan
irregular curvedrow
of 7 eyeson
one side,8 on
the other, with a single isolatedeye lateral totheanteriorend of the rows,on
eachside.Marginal
row
of eyes unfortunately imperfect because of missing anteriormargin
but obviouslyfew
innumber compared
tomost
other species of the genus, of about 2rows
in central part of band, thin- ning outaround
thesidesto afew
widely spaced eyesin a single row.Pharynx
typicalof thegenus (fig. 14). Becauseofbend
inbody mak-
ing
good
sagittal sections impossible, sexual apparatuswas
not sec- tioned. Becauseof eversion of themale
apparatus,most
of thisappa- ratus can be seen in thewhole
mount.Male
apparatus completely everted as in copulation (result of fixation?) (fig. 15). Inside the large cylindricaleverted structure can be seenthe oval seminalvesicle withthick muscularwalland
Indian-club shaped lumen, the 2 spheri- cal thick-walled accessory vesicles (ofunknown
function),and
theNO. 17
POLYCLAD WORMS HYMAN
Jsinuous ejaculatory duct.
The
ducts of the accessory vesicleswere
not seen.At
the distal end of the evert is the penis sheath; throughits
lumen
extends the curved stylet typical of the genus, protrudingfrom
thepenisend
forabout one-third of its length.The sperm form
aclub-shaped sharply definedmass
in thelumen
of the seminal vesicleand from
thismass
a stream ofsperm
can be followed along the ejaculatory ductand
styletand
out of thetip of the stylet.The mode
of entrance of the vasa deferentia into the seminal vesicle is
shown
inthefigure.
Not much
can bemade
out of the female apparatus in thewhole mount
; genital pore appears to lead into acement pouch which
re- ceivesthe encirclingcement
glands (fig. 15).The
eversion of themale
apparatus in this specimen is of great interest. It is undoubtedlynormal
since thesame
eversionwas
seen in aBermuda Prosthiostomum,
P. cy clops (Verrill) where, however, the specimen,through long preservation,was
so darkand opaque
that nothing could bediscernedintheprotrudedapparatus. This Galapagos specimen indicates that inProsthiostomum
the entiremale
apparatus everts in copulation. Thismeans
that themale antrum must
turn inside out, so that its liningbecomes
the surface epithelium of the evert,and
thewhole
male copulatory complex, including the seminal vesicleand
the 2 accessory vesicles,becomes
pulled into the evert. It is further seenfrom
figure 15 that the so-calledpenis sheath actually serves as penis papilla while the penis papilla remains insideand
ap- pearsto function merely to anchor the stylet.One
begins towonder
if these structures have not been
misnamed and
if the penis sheath should not be regarded as the penis papilla,and
the projection at present considered the penis papilla should not receivesome
othername. Of
course,many
polycladsdo
nothave apenis sheath.Whether
it occursonly inthose species witha stylet deserves investigation.
Remarks. — The
species of thegenus Prosthiostomum
resemble each other so closely in external charactersand
details of the copulatorycomplex
that separation into species is oftendifficult.As
pointed out byBock
(1913) onemust depend
chieflyon
the colorationand
the eye arrangement.Some
differences also appear to exist as regards the lengthand
shapeof themale
antrum, shape of theseminal vesicle,and
points ofentrance of thevasa deferentia into the seminal vesicle.On
thewhole
it is not advisable to omit the study of serial sections in determining the species of this genus. In the Galapagos specimen, thedamage
tothe marginalset ofeyes, the lack of dataon
the colora- tion in life,and
the eversion of the male apparatus, preventingcom-
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL.98
parisons with other specieswhere
the male apparatus inknown
only in the resting condition, have increased the difficulties of decidingwhether
one is here dealing with anew
species or not.The
great resemblance, even almost the identity, of the eyearrangement
of the present species with that of thecommon
Mediterranean Prosthiosto-mum,
P. siphunculus (Delle Chiaje) 1828, fully describedby
Lang, 1884, will not have escaped the notice of students of polyclads.On
theotherhand,itseemshighlyimprobable thatthe
same
specieswould
befound
intwo
such widelyseparated localities asthe Mediterraneanand
the Galapagos Islands. It further appears that P. siphunculus isvery
much
larger than P. parvicelis.According
to Lang, 1884, the formermay
reach alength of30 mm. and
the majorityof the sexual individuals rangefrom
12-18mm. The
specimen of P. parvicelis isalready fully
mature
atalengthwhich
could nothave beenmore
than 8mm.
in the extended condition.The
principal difference, however,between
thetwo
species, concernstheshape of the seminal vesicleand
the points of entrance of the vasa deferentia. Lang, 1884, plate 30, figure 20,shows
the seminalvesicleof P. siphunculusas curved, with a simple fusiformlumen and
muscular wall ofuniform
width; the vasaenterat itsproximalend. In P.parvicelisthe wall of the seminal vesicle is very thick proximallyand
thins out distally, thelumen
isof peculiar shape (fig. 15),
and
the vasa deferentia enter the sidesand
then turn proximally to join the beginning of the lumen.The
shapeof thecement pouch
appearsalso to differ in thetwo
forms.On
these
grounds
I withsome
hesitation consider the Galapagos species distinctfrom
the Mediterranean one.Differential diagnosis.
— Eye arrangement
as in P. siphunculus;smaller than this species; seminal vesicle with thicker wall
and
smalllumen
proximally, thin walland
largerlumen
distally, receiving the vasa deferentia into its sides.Locality.
—
Sulivan Bay,James
Island, Galapagos Islands, shore collecting. Collectedby
Dr.Waldo
L. Schmitt, July 24, 1938.Holotype.
— One whole
mount,U.S.N.M.
no. 20426.REFERENCES
Bock, Sixten
1913. Studien iiber Polycladen. Zool. Bidrag, vol. 2, pp. 1-343.
1923. Polycladen aus Juan Fernandez. The Natural History of Juan Fernandez and Easter Island, vol. 3, pp. 341-372.
Girard,Ch.
1893. Recherches sur les Planaires et les Nemertiens de l'Amerique du Nord. Ann. Sci. Nat.,vol. 15, No.7, pp. 145-310.
NO. 17
POLYCLAD WORMS HYMAN
9Hyman,
L. H.1939a. Acoel and Polyclad Turbellaria from Bermuda and the Sargassum.
In press.
1939b.
Some
Polyclads from theNew
England coast, especially from theWoods
Hole region. Biol. Bull., vol. 76, pp. 127-152.Lang, A.
1884. Die Polycladen (Seeplanarien) des Golfes von Neapel und der angrenzenden Meeresabschnitte. Fauna und Flora des Golfes von Neapel, Monog. 11.
Pearse, A. S.
1938. Polyclads of the east coast of North America. Proc. U. S. Nat.
Mus., vol. 86, pp. 67-98.
Verrill, A. E.
1901. Addition to the fauna of the Bermudas from the Yale Expedition of 1901, with notes of other species. Trans. Connecticut Acad.
Sci., vol. 11, pp. 15-62.
EXPLANATION OF NUMBERING OF TEXT-FIGURES
1, cerebral eyes; 2, tentacular eyes; 3, cerebral granule mass; 4, uterus;
5, pharynx; 6, seminal vesicle; 7, prostate vesicle; 8, stylet; 9, male genital pore; 10, female genital pore; 11, Lang's vesicle; 12, stalk of Lang's vesicle;
13, vagina; 14, entrance of uteri into vagina; 15, female antrum; 16, male antrum; 17, ejaculatoryduct; 18, extracapsular prostate glands; 19, vasa defer- entia; 20, cement glands; 21, penis papilla; 22, muscular wall of male antrum;
23, penis pocket; 24, penis sheath; 25, cement pouch; 26, accessory vesicles
;
27, everted male organ; 28, sucker; 29, marginal glands; 30, sperm mass in seminal vesicle; 31, openingofprostate glands into penis pocket.
Whole mounts and figures of eyes drawn with the aid of the camera lucida;
others free-hand.
IO
SMIT„SON IAN MISCELLANE0
"SCOLLECTIONS
•f i
VOL. 98
sexual reg10n of the
holotype
"
PW m ™
l°ris,NO. 17
POLYCLAD WORMS HYMAN
II"..'
-0: Pi
Figs. 4-6-
12
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL.98
l\ /,
:#
)\ i: I..U
u
/ '
-
31
U
J ,',-.r•>'
'M
:'f\\11
Figs. 7-11.
sef'offectTon^aT fit''?
f
«™
V^n*",
°f Notoplana caribbeana, from same vesicle ofNotoplan!\akLin U^
from.th^
.ining epithelium of Lang's clippertoni,hootyp™ Eyes oft"Zf
aS^'-^ f
herU'eS; 9'^
/a"«young specimen oF
Eupfam^l
ippertonicUppertom
> ho
^^
"> Eyes of aNO. 17
POLYCLAD WORMS HYMAN
13Figs. 12-15.
12, Semidiagrammatic sagittal view of the copulatory complex of Euplana clippcrtoni,basedon sagittal serial sections of the sexual regionof theholotype
;
13, Eyes ofProsthwstomum parvicclis, holotype; 14, Prosthwstomum parvicehs, holotype; 15, Everted male copulatory organ of Prosthwstomum parvtcelts containingthe maleapparatus.