THE MIERICAN SPECIES OF SPHYRADimi WITH AN IN-QUIRY AS TO THEIR GENERIC RELATIONSHIPS.By G. Dallas Hanna,Of the United States Bureau of Fisheries, Washington, D: C.Land snails belonging to the genus Sphyradium are found in thoseparts of Europe, Asia, and North America generally designatedas Holarctic. One species extends into the Neotropical realm, andtwo forms which, by the shell, apparently belong to the genus, arefound in the Hawaiian Islands.As the shell is distinctly Pupillid in character the group was orig-inally included in the family Pupilhdas (or Pupidse), but when thejaw and radula became such factors in the classification of the landsnails, the genus was removed from this family and of late yearsit has been placed along with Punctum in a subfamily of the Endo-dontidse, one of the divisions of the Aulacopoda.Some live animals of Sphyradium edentulum ( Vertigo simplex Gould,of Binney and others), were recently received from Caribou, Maine,collected by Mr. O. O. Nylander. A study of this material in con-siderable detail has revealed the fact that the form, by lacking pedalgrooves, does not belong to the Aulacopoda and all the other anatom-ical characters other than those of the jaw and radula show thatit is closely allied to the genus Vertigo of the family Pupilhdse.The genus was removed from the PupilUdse by Sterki ^ upon thestructure of the radula. The jaw is by Sterki stated to be composedof numerous separate plates loosely joined together in a arcuatedseries. This is the condition of the jaw in Punctum and for thisreason the two genera have been placed together in the Endodontidse.According to Lehman,^ however, the jaw is constructed of a singlepiece and the radula is but slightly different from what is found inVertigo, 12?1?12, teeth in a transverse row. Different investi-gators seem to have obtained different results in their w^ork on these
1 sterki, Nautilus, vol. 10, 1896, p. 75.2 Lehman, Lebenden Schnecken und Muscheln der umgegend Stettins und in Pommem, 1873, p. 142,pi. 14, fig. 49. Proceedings U. S. National Museum, Vol. 41?No. 1865 371
372 PROCEEDINGS OF THE NATIONAL MUSEUM. vol.41.organs.^ When the rest of the anatomy is considered, however, itis seen that the genus can not belong to other than the PupilUdseand in this family it is closely related to Vertigo.Genus SPHYRADIUM Charpentier.Shell brown, translucent, cylindrical or subconical. Spire elevatedand apex obtusely pointed. Axis minutely perforate. Aperturebasal, semi-circular and without teeth. Peristome thin and acute,its plane radial from the axis, not reflected or thickened witliin withcallus. No indentation in the upper palatal wall.Animal without a lower pair of tentacles and foot without pedalgrooves.Kidney lying parallel to the rectum and with the urethra leadingdirect from the anterior end to the mantle margin. The granularmatter of the kidney is not arranged in a series of longitudinal fila-ments as in most of the other Pupillidas.Penis with the vas deferens attached at the apex; flagella absent.Hermaphroditic gland composed of a single mass of granules. Nodemarcation between the oviduct and vagina.The genus is closely related to Vertigo, from which it differs mhaving no teeth in the aperture of the shell, and the peristome thinand without a callus deposit. Also the surface of the foot is cov-ered with a network of incised lines which are not found in anyof the species of Vertigo examined by the writer. It is allied to thisgenus by the animal lacking a low^er pair of tentacles and by thevas deferens being attached to the apex of the penis, not do^^^l onits side as in Bifidaria or Pwpoides.In America there are three species of Sphyradium . One of these hasthus far been found only as a fossil and another is almost extinct.Key to the American species.
a. Shell more than 5 mm. in height hasta.a '. Shell lass than 3.50 mm. in height.h. Shell long and slender; whorls seven to eight alticolum.b ^ Shell shorter and more conical; whorls five to six edentulum.SPHYRADIUM HASTA, new species.Shell more than 5 mm. in height, long and cylindrical. Lightbrown in color and glossy. Spire greatly elevated but obtuselypointed on the apex. Whorls 8^ to 9, rather flattened on the faceand the last subangulated around the periphery. The last six
? Note by William H. Dall.?Mr. Olaf O. Nylander of Caribou, Maine, having kindly furnished somefresh material, with the help of Miss Mary Breen the jaw and radula of S. edentulum from that localitywere Isolated, and prove the correctness of Doctor Sterlci's observations. The jaw is narrow antero-posterially, arcuate, and composed of about 16 rhomboidal overlapping plates. A high magnification isrequired to bring out these features. The cusps of the teeth on the mature part of the radula wereblackish instead of horn color, which is very unusual in the Pulmonata.
NO. 1865. AMERICAN SPECIES OF SPffYRADIUM?HANyA. 373whorls are of about equal diameter; the first three increase rapidly.Lines of growth faint and oblique; apex smooth and white. Aperturesomewhat angulated at the base of thecolumella. Peristome thin and acute, form-ing a regular curve without an inden-tation in the upper palatal region such asis present in most of the Vertigos. Theaperture is very slightly thickened withcallus on the inside of the peristome inthe basal region. Peristome not reflectedand with no callus crest back of the aper-ture. Teeth and lamellsB entirely absentfrom the aperture. Umbilicus with a verysmall perforation.Length, 5.81 mm.; diameter, 2.03 mm.This species differs from all others ofSjyhyradium by its much greater size andthe relatively smaller aperture.Type.?Cat. No. 214302, U.S.N.M., fromthe Pleistocene of Long Island, PhillipsCounty, Kansas. The specimens of thisspecies were collected by Mr. Edward C.Johnston and the writer in the autumnof 1910. They occur in deposits ofsandy, green marl of undoubted Pleisto-cene age. With them were large num-bers of other Pupillidse, as well as other land snails. The entirefauna of the beds is foreign to the region at the present time, butis allied to the present day Canadian fauna.From this it is supposed that the animalslived just before or during the Glacial epoch.SPHYRADIUM ALTICOLUM IngersoU.Pupilla alticola Ingersoll, U. S. Geol. Surv. of theTerr., Bull. 1, No. 2, ser. 2, p. 128 (1874); Eighth Ann.Rep. Hayden Surv., 1874, p. 391.Pupa alticola (Ingersoll) Binney, Man. Amer. LandShells, 1885, p. 174, fig. 166.Shell between 2.50 mm. and 3.50 mm. inheight, long and cylindrical. Light brown incolor with the apex much lighter, sometimeswhite. Spire elevated and with the apex ob-tusely pomted. Wliorls 7 to 8, well rounded onthe face and with the sutures well impressed.The last whorl in adult shells is of greater diameter than the onepreceding it, while the next three above it are of about equal size;
Fig. 1.?Sphyradium hasta Hanna,TYPE, cat. no. 214302, U.S.N.M.Draavn by g. d. hanna.
Fig. 2.?Sphyradium alti-coLUM Ingersoll.Drawn by g. d. hanna.
374 PROCEEDINGS OF THE NATIONAL MUSEUM. vol.41.this expansion of the last whorl makes the shell slightly iinsymmet-rical. Peristome thin and acute, without callue, thickenings orindentations in the palatal wall. Lines of growth weak and oblique
;
apex smooth. Aperture larger in proportion to the shell than ineither S. hasta or 8. edentulum; entirely without teeth or lamellae.The umbilicus is very slightly perforate.Height. Diameter.3. 33 1. 742.81 1. 11 ^2. 70 1. 14This species is found at various places as a fossil in the Pleistoceneand Loess deposits of the Mississippi Valley.^ It is also found inseveral States of the Rocky Mountain region at the present time;i. e., in Utah, Colorado, and Wyoming.Its abundance in the Pleistocene beds of Long Island, PhillipsCounty, Kansas, shows that at one time this was a very commonshell. Many specimens were found there in 1910 by ^Mr. Johnstonand the writer. They were associated with S. Tiasta and other Pupil-lidae which are at the present time apparently extinct.This species can not be confused with any other United States landmollusk. The much greater height of the shell and the greater num-ber of whorls shows it at once to be distinct from aS'. edentulum.SPHYRADIUM EDENTULUM Drapamaud.Pupa edentula Draparnaud, Hist. Moll., 1805, jj. 52, pi. 3, figs. 28, 29.Pupa simplex Gould, Bost. Journ. Nat. Hist., vol. 3, 1840, p. 403, pi. 3, fig. 21.Vertigo simplex (Gould) Binney, Man. Amer. Land Shells, 1885, p. 191, fig 195.Shell, smaller than either of the two preceding species, not morethan 2 mm. in height; translucent brown in color, the apex notlighter than the body of the shell in live specimens. Outline conicalrather than cylindrical; apex obtuse. Wliorls four to five, wellrounded on the face and wnth the sutures well impressed. Lines ofgrowth very faint and oblique. Shell not spoiled in symmetry by thelast whorl increasing in size unproportionately . Aperture well roundedand proportionately not so large as in S. alticolum; entirely withoutteeth or lamellae. Peristome thin and acute, without callous thick-enings or indentations. Umbilicus minutely perforate.Height, 1.60 mm.; diameter, O.SO mm.The shell does not vary appreciably in size. It is a resident in thatpart of North America designated by the United States BiologicalSurvey ^ as Transition and Canadian. It is also a resident of thenorthern parts of Europe and Asia.
1 Pilsbry, Nautilus, vol. 11, 1898, p. 142.2 U. S. Biological Survej'. Fourth Provisional Zone Map of North America, by C. Hart Merriam, VernonBaUey, E. W. Nelson, and E. A. Preble, 1910.
NO. 1S65. AMERICAN SPECIES OF 8PHYRADIVM?HANNA. 375
Fig. 3.?Animal ofSPm-KADIUM EDEX-TULUM DrAPAE-NAUD WITH THESHELL REMOVED,SHOWING THE KID-NET. Dra^'n byCD. 11ANNA.
Animal (fig. 3) , with a rather short oval foot. Color dark about thehead region, hghter posteriorly and on the sole of the foot. Lowertentacles absent. Foot without pedal grooves or traces of such struc-tures. Surface of body covered with a network of incised lines, themeshes of which are very large. Tliis network is absent in at leastsome of the Vertigos, but is present in all of the Bifi-daria and Pupoides which the writer has had the op-portunity to examine.The breatliuig pore is situated at the upper right-hand angle of the mantle and with the anus imme-diately to the right. There are no black markingson the mantle or lung wall.Kidney, a long, slender, granular walled pouch at-tached to the outer wall of the pallial cavity and withthe posterior end lying against the anterior end of thealbumen gland. The glandular matter of the kidneyis not arranged in longitudinal filaments or folds as inmost of the other Pupillids, but is evenly distributed over the entireinner surface. There is in the anterior end an indication of transversebars which are more noticeable on the upper part of the urethra. Theurethra leads directly from the anterior end of the kidney to theexcretory pore, which is situated as in the other Pupdlidse. It opensinto the anterior end of the pallial cavity, immediately posterior tothe breathing pore. The size of the urethra and rectum is about thesame and the two are separated by a considerabledistance.The salivary glands are united in one, buthave a pair of ducts leading to the buccal mass,and passing beneath the cerebral nerve commis-sure and tlirough the nerve collar.The genitalia (fig. 4) are distinctly Pupillid incharacter. Hermaphroditic gland, composed ofa single mass of grape-like granules as in Vertigo.In the Endodontidse, where Sphyradium has here-tofore been placed, this gland is composed ofmany individual bunches of elongated tubules,the bunches arranged in a diminishing series towardthe apex. The gland of S. edentulum is embeddedin the anterior end of the liver, which is lightHermaphroditic duct slightly swollen in its lowerhalf, but not convoluted in any part. It discharges into the oviductat the junction of the latter with the albumen gland.The albumen gland is very large and massive, white and transparent,and on the posterior end there are a few scattering specks of blackpigment. These are on the outer surface of the gland only. Theoviduct is large at the posterior end and gradually diminishes in size
Fig. 4.?Genitalia ofsphyradium edentu-lum Draparnaud.Spec. no. 142, of thewriter's collectionOF alcoholics. Muchmagnified. Drawn byg. d. hanna.brown in color.
376 PROCEEDINGS OF THE NATIONAL MUSEUM. vol.41.
anteriorly, passing imperceptibly into the vagina. It contains nofolds or pouches, nor is there upon it such a shell gland (prostate) asis found in the Endodontidse. The vagina is a wide tube, free of con-volutions, and it opens to the exterior at the base of the right eye-stalk.Spermatheca, small and globular with a slender duct almost aslong as the oviduct and vagina. It discharges into the vagina nearthe lower end of the latter. There is no point of demarcationbetween vagina and oviduct, but for purposes of study and comparisonin the Orthurethra it is convenient to call that part of the duct belowwhere the vas deferens leads away, vagina, and that which is above,oviduct.The penis is very small in comparison to the size of the other organs.TMs, however, is characteristic of the Pupillidae. It is not attachedto the vagma as usual in this family, and therefore there is no atrium;instead it opens to the exterior, on the vaginal lip. The penis is veryslender at the point of exit, but enlarges gradually toward the upperend, where it is half the diameter of the vagina. The vas deferensleads from the center of the female organs, the junction of the vaginaand oviduct, to the apex of the penis. The retractor muscle isattached to the vas deferens a little way down from tne apex of thepenis and also to the floor of the pallial cavity. Above the attachmentof the vas deferens there is no continuation of the penis as in Bifidariaand no long flagella-like appendage as in Pupoides. In S. edentulumthe condition is nearer what is found in some of the Vertigos as V.ovata.The genitalia are most closely related to the Vertigos of any groupof the Pupillidae, thus bearing out the resemblance of the shells.The j aw, radula, and genital organs of the European ^S'. edentulumhave been described and figured by R. Lehman.^ His figures are notvery good, but they show that the genitalia of the animal he had werevery different from those of the form now under consideration. Ashe describes and figures these organs there is one long flagellum on thepenis and two short thread-like ones. The larger one has a bulb-likeenlargement on the outer end. He also says the oviduct is pouchedand folded, and his figure shows a distinct demarcation between theoviduct and vagina. If the species he had was S. edentulum and if hisdissections are correct, then the American form is specifically if notalso generically distinct from the European, and the name, ^S". simplexGould, will apply to our snail.However, until modern methods of dissection are used on the Euro-pean form, it is perhaps best to continue to refer the American snailto S. edentulum.
I Lehman, Lebenden Schnecken, etc., p. 142, pi. 14, fig. 49.