Urn
BBAtf
MMMM_MM
KELEY
RARY
ERSfTY OF
IFOKNIA
FRONTISPIECE
SECOND SERIES: PULMONATA.
MAN UAL
OP
ONCHOLOGY:
STRUCTURAL AND SYSTEMATIC.
WITH ILLUSTRATIONS OF THE SPECIES.
BY GEORGE W, TRYON, JR.
CONTINUATION BY
HENRY A. PILSBRY,
CONSERVATOR OF THE CJNCHOLOGICAL SECTION AND PROFESSOR OF MALACOLOGY
IN THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA.
Vol. IX.
(HELICID^E, VOL. 7.)
GUIDE TO THE STUDY OF HELICES.
PHILADELPHIA :
3?iablislied by Concliologioal Section
ACADEMY OF NATURAL SCIENCES,
OF PHILADELPHIA.
1894.
tJNIVERSITT
*. Of
EARTH
SCIENdt
LIBRARY
PREFACE
The group of Pulmonate genera familiarly known as Helices,
forms an important factor in the land mollusk fauna of every coun-
try, in point of numbers exceeding any other group of snails. This
numerical and faunal pre-eminence has caused the authors of the
MANUAL to devote eight volumes to Helicoid genera, the earlier
three (Vol. II to IV) being prepared by Mr. Tryon, the later vol-
umes by the writer.
During the progress of the work it became obvious that the
established system of grouping required revision, not alone in the
details of many minor divisions, but in those broader principles
underlying our conceptions of the entire classification and genealogy
of the group. The object of this volume is to formulate in compact
form the new classification of Helices, and incidentally to indicate
some general principles upon which a new grouping of all land pul-
monates must be based.
In the systematic portion of the work (pp. 1-344) I have
attempted to show the main characters of the genera, both in hard
and soft anatomy, giving illustrations as copious as the limits of the
work would permit ; for while fully pursuaded that, as Darwin has
said, naturalists " never read each other's works," I am sure that
they look at the pictures illustrating them. In the Introduction the
larger groups are defined (p. xxxii) and their probable genealogy
suggested (p. xxxi). Finally, the geographic distribution of Helices
is discussed with reference to the genesis and migrations of the
principal groups, and the origin of modern faunas (p. xxxviii).
Few will dispute the general proposition that until the systematic
classification of a group is placed upon a secure basis, all discussion
of the larger questions of geographic and geologic distribution is
futile. A sound systematic zoology is at once the key and the test
of zoogeographic speculations ; and without this check, zoologist
and geologist are alike at the mercy of mere opinion and specula-
tion, too often based upon false notions of affinity, or upon a decep-
IV PREFACE.
tive external likeness which may mask fundamental differences.
These considerations justify, I believe, the stress placed upon mere
system in this volume. The treatment of minor groups may be ob-
jected to as unduly minute ; and it is true that most groups seem
over-divided. As my predecessors are responsible for most of this,
I have been satisfied to reflect their labors faithfully. Those groups
having important structural characters I have considered generic;
grouping under these as subgenera and sections the various smaller
assemblages, which specialists find useful, but which are usually of
little systematic value, and not much utility to the general malacol-
ogist. These remarks imply no disrespect to the founders of this
multitude of groups. Their labors were necessary in pointing out
the differential features of Helices. They sought differences, for the
establishment of new groups ; the modern systematist seeks more
profound likenesses, in order to establish lines of descent. The
splitting of faunas into minute groups has taught us the compara-
tive value of characters, paving the way for more philosophical
study of the genealogy of faunas. The torch of analysis lights the
path for synthesis.
It will, of course, be obvious that a general idea of the contents of
the principal divisions of Helicidse as here distinguished, must be
obtained before the geographic hypotheses can be rightly under-
stood.
Acknowledgements and Thanks. That a large number of Helicoid
groups are made known anatomically in this work is primarily due
to the kindness and generosity of many conchologists who have sup-
plied living or alcoholic material for dissection ; and while it would
be impossible to name here all those who have thus assisted me with
specimens, notes on distribution, synonymy, etc., I must express
my obligations for material for investigation to W. G. Binney, John
Brazier, Alfred Caruana Gatto, Dr. J. C. Cox, Wm. H. Ball, Henry
Hemphill, J. B. Henderson, C. W. Johnson, O. von Mollendorff,
Morris Schick, Dr. Benj. Sharp, Dr. H. Simroth, Frederick Stearns,
Henry Suter and Rev. R. Boog Watson. A series of mounted rad-
ulse which I owe to Rev. Prof. H. M. Gwatkin, has enabled me to
illustrate the teeth of many interesting genera, among them Oxy-
chona, Macrocyclis, Albersia, Planispira, Entodina, Acavus and
others. My friend, Charles Hedley, of Sydney, has contributed not
a little to views both systematic and theoretical expressed herein,
but my main debt to him is for help more subtle than this.
PREFACE. V
To Mr. John Ponsonby, of London, thanks are due for numerous
rare or new species of Helices, many of which have been figured in
the Manual, and more especially for the correction of errors in
synonymy, localities, etc., occurring in previous volumes of this
work. Mr. G. K. Gude has rendered me a similar service ; and
from a very large number of conchologists both in America and
abroad, I have received information upon particular species and
genera, for all of which I would here express my gratitude.
Summary. In this volume the author has essayed to indicate the
primary groups of the Helicidae, arranging the genera accord-
ing to a few main types of internal structure, in place of the chaotic
or arbitrary sequence of groups hitherto prevailing. The multitude
of groups recognized are shown to be reducible to about fifty genera
distinguished by structural features of importance, which are de-
scribed and illustrated, lists of the living species of each genus being
given. An outline of the distribution of the main groups is offered,
with hypotheses of the probable migrations and phylogeny of these
groups. Incidentally, the comparative value of the genitalia, shell,
jaw and radula in classification, and the laws of their modification
are worked out in some detail. Finally, the nomenclature of Helices
has been thoroughly revised, and, it is hoped, placed upon a sound
basis.
It rests with the critical and discriminating conchological public
to decide whether the author of this volume shall undertake a com-
panion work on the genera of Zonitidse and Agnatha.
H. A. P.
MANUAL OF CONCHOLOGY, IX.
GUIDE TO THE STUDY OF HELICES.
INTRODUCTION.
I. NOTES ON THE GENERAL MORPHOLOGY OF HELICES.
SHELL.
In Helices the shell is always a well developed spiral, capable ol
containing the entire animal when retracted, It is generally wider
than high, and coiled loosely so that the central column is hollow or
umbilicate, but in some forms it is much higher than wide, and the
umbilicus is closed in the adult by an expansion of the lip, or the
whorls are coiled in close contact, forming a solid columella,
The general contour of the shell is excessively variable in all gen-
era containing many species ; and as the number of main types of
form is limited, parallel groups or species occur in the various gen-
era as shown in the following table :
Genera. -Shell globose, Shell depressed, Shell keeled.
Helix
Pomatia,
vermiculata,
gualtierana.
Helicigona
Arianta,
11 Campylcea"
lapicida.
Epiphragmop
hora calif or niensis,
mormonum,
circumcarinata.
Eulota
Acusta,
Euhadra,
Pleetotropis.
Helicostyla
Calocochlea,
Corasia,
Axina.
Polygyra
" Mesodon"
tridentata,
obstricta.
Tliei sites
Xantlwmelon,
Badistes,
Glyptorhagada.
Camcena
Phcenicobius.
xanthoderma,
saturnia.
Obba
papilla.
planulala,
marginata.
Pleurodonte
nuxdenticulata,
Isomeria,
Caracolus.
The list is capable of indefinite extension ; and even those minor
groups called " sections " often show the same series of changes in
form, thus:
(vii)
Vlll MORPHOLOGY OF SHELL.
Sections. Shell globose, Shell depressed,
" Dentellari't " nuxdenticulata, deniiem, lychnuchus.
Thelidomus emarginata, petltiana, lima.
Pleurodonte bronni, anomala, ' pvracutissima.
Stenotrema stenotrema, monodon, spinosa.
Axina montfortiana, magister, siquijorensis,
That characters of contour are valueless for distinguishing gen-
era in Helices is now conceded by students of the living groups,
but palaeontologists still use them ; and for this reason the above
tables are given.
The sculpture of Helices, like the contour, affords valuable spe-
cific characters, being subject to a wide range of mutation. Shells
may be either smooth, obliquely striate, ribbed, decussated, granu-
lated, malleated or hairy ; and frequently several varieties of sculp-
ture characterize different species of one genus, thus :
Genus. granulate, spirally striate,
Helicigona lapicida, arbustorum,
Polygyra palliata, albolabris,
Epiphragmophora tudiculata, inter cisa,
Pleurodonte lima, petitiana,
ribbed, hairy, smooth.
gobanzi, setosa, cingulata.
obstricta, hirsuta, jejuna,
eircumcarinata, remondi, mormonum.
scabrosa, auridens, marginella.
Sometimes upon a smooth or granulate surface there are papillae
or hairs arranged in regular obliquely decussating series, or in
quincunx. This occurs in some species of Chloritis, Helicigona,
Thysanophora, Lysinoe, Hygromia, etc. Some genera exhibit a
wide range of variation in texture and color, but in most cases this
is correllated with the habits of the species. Tree living snails are,
as a rule, bright colored and tend to become elevated or conical,
while ground snails are duller or brown, and usually depressed*
Some genera, like Helieostyla in the Philippines and Cepolis in the
West Indies, contain both arboreal and terrestrial forms, and con-
sequently appear, on superficial observation, to be composed of very
incongruous elements.
The embryonic shell (the portion formed within Ihe egg), is found
to vary greatly in size, and its extent compared to that of the adult
MORPHOLOGY OF SHELL. IX
shell is a character of considerable value in classification. In
Helicophanta, Acavus&nd their allies it is very large, sometimes one-
third the diameter of the adult shell, and its junction with the post-
embryonic growth is distinctly marked. In Polygyra it is very
small and indistinct. In Camcena and allied groups it is of medium
size. Some genera have the embryonic shell sculptured, as Ano-
glypta, Chloritis, certain species of Helicigona and Pleurodonte, but
it is usually smooth and polished.
The apertuie is usually crescentic, half-round or round, but in
keeled species becomes angular, and in those having teeth it is often
ear-shaped. The outer lip is expanded, reflexed or thickened within
in nearly all the genera, but in some (Sagda, Glyptostoma, etc.) it is
simple and sharp as in Zonitidce. Tooth-like processes are fre-
quently developed upon the lip and parietal wall, and sometimes
these become excessively complex. Usually there are two teeth
upon the lip and one upon the body wall ; totally diverse genera
having independently evolved this arrangement. In a few groups
there are internal plates or septa, far within the mouth.
The banding of Helices, although variable as a specific character,
often shows considerable constancy in a genus or subgenus. Thus,
in Helix the five-banded plan of coloring is usual. In Helicigona
one- or three-banded ; Epiphragmophora is one-banded. The band
just above the periphery is the most constant, and may be found in
most genera of Belogona. The Epiphallogona have their own
band-arrangement, noticed on p. 103. Snails inhabiting dry situa-
tions or arid regions, deposit more lime in the shell than those liv-
ing in moister places, and there is a strong tendency to split the
bands into many narrow lines, as in Euparypha, Helicella, Rhagada,
Micrarionta.
A convenient formula was invented by Georg von Martens many
years ago, for the designation of band variations in Helices, espe-
cially the five-banded forms. The bands are numbered 1, 2, 3, 4,
5, beginning above. The absence of any band is indicated by a
cypher ; the coalescence of bands by parenthesis ; and the splitting
of a band by repetition of its number. Thus, the specimen shown
in fig. 5, of plate 44, is Helix nemoralis, 12345. Fig. 4 is H. nemo-
ralis 00000. Fig. 12 is H. desertorum 123(45). PI. 43, fig. 44, is
H. saulcyi 1(23) 40. A specimen with the bands united to conceal
all the ground color would be (12345) ; and one with the third
iband split would stand 123345.
Vlll MORPHOLOGY OF SHELL.
Sections. Shell globose, Shell depressed,
" Dentellarix " nuxdenticulata, dentiens, lychnuchus.
Thelidomus emarginata, petitiana, lima.
Pleurodonte bronni, anomala, peracutissima.
Stenotrema stenotrema, monodon, spinosa.
Axina montfortiana, magister, siquijorensis,
That characters of contour are valueless for distinguishing gen-
era in Helices is now conceded by students of the living groups,
but palaeontologists still use them ; and for this reason the above
tables are given.
The sculpture of Helices, like the contour, affords valuable spe-
cific characters, being subject to a wide range of mutation. Shells
may be either smooth, obliquely striate, ribbed, decussated, granu-
lated, malleated or hairy; and frequently several varieties of sculp-
ture characterize different species of one genus, thus :
Genus. granulate, spirally striate,
Helicigona lapicida, arbustorum,
Poly gym palliata, albolabris,
Epiphragmophora tudiculata, inter cisa,
Pleurodonte lima, petitiana,
ribbed, hairy, smooth.
gobanzi, setosa, cingulata.
obstricta, hirsuta, jejuna,
circumcarinata, remondi, mormonum.
scabrosa, auridens, marginella.
Sometimes upon a smooth or granulate surface there are papillae
or hairs arranged in regular obliquely decussating series, or in
quincunx. This occurs in some species of Chloritis, Helidgona,
Thysanophora, Lysinoe, Hygromia, etc. Some genera exhibit a
wide range of variation in texture and color, but in most cases this
is correllated with the habits of the species. Tree living snails are,
as a rule, bright colored and tend to become elevated or conical,
while ground snails are duller or brown, and usually depressed*
Some genera, like Helicostyla in the Philippines and Cepolis in the
West Indies, contain both arboreal and terrestrial forms, and con-
sequently appear, on superficial observation, to be composed of very
incongruous elements.
The embryonic shell (the portion formed within Ihe egg), is found
to vary greatly in size, and its extent compared to that of the adult
MORPHOLOGY OF SHELL. IX
shell is a character of considerable value in classification. In
Helicophanta, Acavus and their allies it is very large, sometimes one-
third the diameter of the adult shell, and its junction with the post-
embryonic growth is distinctly marked. In Polygyra it is very
small and indistinct. In Camcena and allied groups it is of medium
size. Some genera have the embryonic shell sculptured, as Ano-
glypta, Chloritis, certain species of Helicigona and Pleurodonte, but
it is usually smooth and polished.
The aperture is usually crescentic, half-round or round, but in
keeled species becomes angular, and in those having teeth it is often
ear-shaped. The outer lip is expanded, reflexed or thickened within
in nearly all the genera, but in some (Sag da t Glyptostoma, etc.) it is
simple and sharp as in Zonitidce. Tooth-like processes are fre-
quently developed upon the lip and parietal wall, and sometimes
these become excessively complex. Usually there are two teeth
upon the lip and one upon the body wall ; totally diverse genera
having independently evolved this arrangement. In a few groups
there are internal plates or septa, far within the mouth.
The banding of Helices, althongh variable as a specific character,
often shows considerable constancy in a genus or subgenus. Thus,
in Helix the five-banded plan of coloring is usual. In Helicigona
one- or three-banded ; Epiphragmophora is one-banded. The band
just above the periphery is the most constant, and may be found in
most genera of Belogona. The Epiphallogona have their own
band-arrangement, noticed on p. 103. Snails inhabiting dry situa-
tions or arid regions, deposit more lime in the shell than those liv-
ing in moister places, and there is a strong tendency to split the
bands into many narrow lines, as in Euparypha, Helicella, Rhagada,
Micrarionta.
A convenient formula was invented by Georg von Martens many
years ago, for the designation of baud variations in Helices, espe-
cially the five-banded forms. The bands are numbered 1, 2, 3, 4,
5, beginning above. The absence of any band is indicated by a
cypher ; the coalescence of bands by parenthesis ; and the splitting
of a band by repetition of its number. Thus, the specimen shown
in fig. 5, of plate 44, is Helix nemoralis, 12345. Fig. 4 is H. vemo-
ralis 00000. Fig. 12 is H. desertorum 123(45). PL 43, fig. 44, is
H. saulcyi 1(23) 40. A specimen with the bands united to conceal
rail the ground color would be (12345) ; and one with the third
*band split would stand 123345.
X EXTERNAL ANATOMY.
EXTERNAL FEATURES OF ANIMAL.
The general form of the animal in Helicidce is similar to that of
Zonitidce, etc. The shell is carried on the middle or somewhat
behind the middle, its axis being held oblique or vertical to the
plane of the sole. The head has the usual eye-peduncles and ten-
tacles, and more or less distinct labial lobes (see frontispiece, fig. 7).
The mantle rarely projects beyond the lip-edge of the shell, and is
generally provided with right and left body lobes (frontispiece, fig.
7, r.l. right lobe, II. left lobe). Sometimes the latter emits one or
two small tongue-like processes on the left side (pi. 33, fig. 7). The
back, from mantle to head, generally shows one or several dorsal
grooves. The sides are granulated in various patterns, and often a
groove extends from the lips obliquely upward to mantle on each
side, the facial grooves (see pi. 33, figs. 7, 8 ; frontispiece, fig. 7).
The tail in some genera has a median longitudinal groove (espe-
cially in Epiphallogona) or sometimes a serrate keel (Lysinoe, Oxy-
chona). Usually, however, it is rounded above and shows no special
features, being granulated like the sides, but more finely. In the
Endodontidce and Zonitidce a deep longitudinal furrow runs parallel
to the foot-edge on each side a short distance above it. These are
the parapodial or pedal grooves (see pi. 14, fig. 46). They are
absent in Helicidce. In Zonitidce and Endodontidce these furrows are
often associated with a mucus-secreting pore at the tail. The sole or
creeping disc is divided longitudinally into three bands or areas in
some genera, but in most Helices such division is absent, or indica-
ted by coloring only.
DIGESTIVE TRACT.
The jaw is well developed and usually strong and orange-colored
in Helices. The types of jaw occurring in Helicidce, Endodontidce f
and Zonitidce are
Polyplacognath (or unsoldered type of jaw, see pi. 1, figs. 4, 5, 6 r .
9) consisting of numerous separate plates, overlapping at their edges y
and united by a common membrane only (Punctum').
Stegognath (or plaited, pi. 15, fig. 6, 7) composed of similar or
narrower vertical plates soldered together, but with free, overlapping
outer edges (Flammulina, Sagdd).
Goniognath (or converging-plaited, pi. 42, fig. 36) same as stego-
gnathous type, but outer imbricating edges of each plate converg-
MORPHOLOGY OF JAW. XI
ing toward the middle below, the median plate or plates triangular,,
not reaching the cutting margin (Plectopylis).
Aulacognath (or striated, pi. 15, figs. 1, 2) primary elements or
plates completely soldered together, vertically striated (Pyramidulay
Oxygnath (or smooth, pi. 21, fig. 8) completely soldered, smooth^
(Leucochroa).
Odontogrtath (or ribbed, pi. 21, fig. 11) completely soldered, hav-
ing convex vertical ribs, projecting at one or both edges {Helix}.
The most primitive type of jaw occurring in recent terrestrial
Pulmonata is found in the Polyplacognatha, Punctum and Laoma..
By the partial union of the loose plates of this sort of jaw, the
Stegognathous type is formed. The goniognath form as seen in
Liguus, Orthalicus, etc., is a mere variant of this low stegognath
type, and can hardly be considered a primary type. In the Aulaco-
gnatha the plates have become completely soldered, although their
edges still show as strise ; and finally in the Oxygnatha these striae
disappear, leaving a completely smooth jaw. In the Odontognatha,
vertical ribs are developed upon its anterior face. The data supplied
by anatomy and embryology indicate the above as the general phy-
logenetic sequence of the various types of jaw ; but the Oxygnatha
consist of two sections of different genesis. In some forms (such as
the typical Sagdas) the jaw has apparently been evolved directly
from the stegognathous type ; and this is probably true likewise of
the HelicopTianta group. In others (such as some species of Pleu-
rodonte, and Helicostyla,t\\e genera Obba, Cepolis, Leucochroa, Allo-
gnathus, etc.) a smooth jaw has resulted from the degeneration of
the ribs on an odontognathoustype. The ribbed orodontognathous
type has in some cases been formed upon a plaited jaw. In other
cases it may have been formed upon a smooth jaw, but evidence is
lacking to establish this. In certain cases (such as Hygromia) the
degeneration of a ribbed jaw has resulted in one approaching the
plaited type. It must also be understood that the distinction
between the goniognathous, stegognathous, aulacognathous and
oxygnathous types is in some cases not well defined, and often it is
not possible to distinguish between & primarily or secondarily oxy-
gnathous or smooth jaw, although it is practically demonstrated that
the Oxygnatha are diphyletic.
It therefore appears that at the time the main phyla of monotre-
mate, jaw bearing land snails diverged, they were provided partly
with a jaw of unsoldered plates, partly with one of the incompletely
Xll MORPHOLOGY OF JAW.
united type (stegognathous or plaited). In the Helicoids the major-
ity of forms acquired the firmer and completely united smooth or
ribbed type, although some still retain the primitive, incompletely
united forms, as seen in Punetum, Flammulina, Thysanophora, etc.
In the Zonitidce the oxygnathous type has been very generally
acquired, although a few forms retain a modified plaited jaw. In
JBulimulidce (which includes the " Orthalicidae ") the plaited type of
jaw has been retained with various modifications, and the same is
found in Cylindrellidte. The Papidce have a completely united,
striated jaw. The Achatinidce have a striated or ribbed jaw. It
appears that the various families, starting with an incompletely
united jaw, have been very unlike in the degree of development
attained ; some preserving the ancestral form until to-day, but in
most a stronger, solid jaw has been acquired through various well
understood successive stages, occasionally parallel in several phyla.
These considerations show that the various classifications of land
mollusks by jaw characters are artificial ; the various " types " of
jaw on which it is founded representing merely successive stages of
progress from an incoherent or incompletely united, to a solid jaw,
and these stages have been independently reached or passed through
by several totally diverse branches of the pulmonate trunk. The
ihistory of the various jaw types is shown in the following diagram-
ribbed smooth
striated
plaited goniognath
Jaw of distinct plates
The two lower stages were probably passed through by the majority
<of families in common ; the others were reached by various groups
independently and by their own special routes. In most families of
land snails, two or more of these types are represented among the
various genera.
THE RADULA in Helicidce is of the strap-like form usual in Pul-
monata, the individual teeth having squarish basal plates. In even
the lowest types now existing, the multicuspid form of tooth of the
primitive Pulmonates has given way to the tricuspid type (see pi.
MORPHOLOGY OF TEETH. Xlll
15, figs. 3, 4), although in some forms more cusps remain on the
outermost teeth. The individuality of these three cusps is remark-
ably fixed ; for however completely the typical tricuspid form may
be changed, it is always possible to identify the three primitive ele-
ments, or such of them as are retained.
In the study of Helicid raduhe, and especially those departing
widely from the typical structure, it is essential to recognize at the -
outset
The law of mesometamorphosis : All modifications in the teeth
proceed from the median line of the radula outwards toward the edges,
the o Liter marginal teeth being the last to be modified.
A study of the marginal teeth, therefore, gives a clue in many
cases to the ancestral condition of a much modified radula ; although
in certain groups the change has been so long established and has
proceeded so far that even the outermost teeth no longer retain
their primitive form. In such cases recourse must be had to the
radube of young individuals or embryos still uhhatched, which
sometimes retain an ancestral type of teeth (see Sterki, Proc. Acad. ..
Nat. Sci., Phila., 1893, p. 388).
The evident reason why the order of tooth-changes stated above
should obtain, is that the median portion of the radula is the part
most used on account of its position and the convex boss-like shape
of the subradular cushion.
The most frequent departure from the tricuspid type of tooth is
seen in the lateral teeth of most Helices, in which the inner cusp
(eutocone) is lost, cr more commonly its cusp is united with that of
the middle cusp (mesocone) as a lateral extension of the latter. In
many groups both inner and outer cusps of rhachidian and lateral
teeth are suppressed in this manner (see pi. 34, fig. 9), but all three
cusps reappear on the marginal teeth, which are less modified.
Usually the outer marginals have the ectocone, or outer cusp, split
or bifid, a reminiscence of the early multicuspid teeth which were
part of the heritage of the Pulmonates from their Tectibranch
ancestors.
Radulw with teeth tricuspid in whole or part. In many Endodonti-
dee and minute forms of other groups, the teeth are all tricuspid (see
plates 8, 9). This form of teeth is usually correllated with small
size and strictly terrestrial habits.
RadulcK with all teeth unicuspid. In a few genera the loss of side
cusps has extended to even the outermost teeth of the radula (see-
XIV MORPHOLOGY OF TEETH.
pi. 51, figs. 1, 2, and pi. 48, all figs.). This modification is especially
characteristic of one of the primary divisions of Helices, but occurs
also on a few isolated genera, such as Allognathus, of other phyla.
Radulw of arboreal snails. Data presented in the systematic por-
tion of this volume establish the fact that arboreal snails always
assume teeth with broad, gouge-like cusps, in place of the slender,
pointed cusps of ground snails, and regardless of the form of teeth
prevailing in the family stocks whence they were derived. Cases
in point are Polymita, Amphidromus, Orthalicus, Papuina, Cochlo-
ttyla, Oxychona, etc., etc. Some apparent exceptions are due to the
very recent assumption of arboreal habits by certain forms ; the
change of teeth lagging behind the change of station, as in the
arboreal forms of the genus Cepolis.
This modification goes hand in hand with the change in shell feat-
ures ; arboreal forms always becoming light or bright colored, often
having a color-scheme in vivid hues of green, yellow, orange or
pink; while the most nearly allied terrestrial species or genera have
the shell of dusky or inconspicuous shades of brown.
In some tree snails the middle cusp only is modified into a broad
gouge, the side cusps remaining as rudimentary basal spurs, which
become larger on the outer edges of the radula, in accordance with
the general law formulated above. An instance is Oxychona. pi.
51, figs. 9, 10, (o being the rhachidian tooth). Again, the three
cusps are retained and enlarged on all the teeth, as in Polymita, pi.
51, figs. 5, 6, 7. (Fig. 7, outermost marginals; compare pi. 57, fig.
48, a marginal of Cepolis, the genus most nearly allied). The same
has occurred in Papuina, pi. 37, figs. 1, 10.
As a general rule, groups of greater value than genera cannot be
based upon these special modifications of the tricuspid type of teeth.
And on account of the fact that similar modes of life produce simi-
lar tooth-forms in widely different groups, these peculiarities can
have comparatively little weight in fixing the place in the general
system or the family affinities of any genus.
The salivary glands, stomach, liver and intestine have not been
observed to offer differences of taxonomic value in the Helices,
although I have observed variations in certain genera. An extended
series of observations of these organs is necessary.
KEPRODUCTIVE SYSTEM.
General considerations: Helicidse, like all pulmonates, are her-
maphrodites, the male and female genitalia uniting below in a com-
MORPHOLOGY OF GENITALIA. XV
nion cloaca, the atrium or vestibule. It is now held that the herma-
phrodite condition is secondary in mollusks, the male organs being
superimposed or grafted upon the female individual (see Pelseneer
Quart. Jouru. Mic. Sci. 1894, p. 19). The proofs for this view com-
ing from many sides, all indicate that in the primitive mollusks the
sexes were separate.
Embryological data indicate that the entire generative system
except atrium, penis sack and their special appendages, are of mesp-
dermal origin. Simroth is probably right in holding that the
atrium and evertible penis (but not epiphallus) are ectodermal evag-
inations. The case of Limax primitiviis which he cites to prove that
the penis has been " pulled out" from the atrium, is, however a case
of degeneration in all probability. It is very probable that the
penis in land mollusks is strictly homologous with that of Tecti-
branchs, and its union with the female organs at the atrium has
been brought about by the gradual moving forward of the female
orifice, originally posterior in position.
It seems likely that the dart apparatus is primarily an outgrowth
from tlie atrium, although in some cases it has moved upward on
the vagina. It is not homologous with the dart sack of Philomycus,
nor with that of certain Zonitidce. The gland or sack upon the
penis, called the appendix, is probably a very ancient character, and
is homologous with that sometimes developed upon the atrium (see
Helicella), but not with the blind sack found high on the vagina in
such forms as Panda, etc., which seems to be an independent growth
from the vagina, probably serving as a temporary receptacle for
spermatophores (packets of spermatozoa), analogous to the diverti-
culum of the spermatheca duct. Although both male elements
(spermatozoa) and female (ova) are produced in the same acini of
the hermaphrodite gland, the former ripen first, and passing down
are enclosed in a leathery or chitinous case, the spermatophore
(" capreolus ") secreted by flagellum or epiphallus. In forms
lacking these the spermatophore is absent. In the female system
these spermatophores are stored in the spermatheca and its ap-
pendages, pending the ripening of eggs and their passage down-
ward. The dart apparatus is only a stimulating organ, the dart
being thrust from one individual into another during copulation.
Von Ihering considers the papilla in the penis also a sensory organ.
The function of the penis-gland is unknown. During copulation the
penis is everted in most Helices, but in some there are reasons for
XVI MORPHOLOGY OF GEX1TALIA.
believing that the atrium only is thrust outward. Further investiga-
tions of snails during breeding are needed.
Description of organs : The external opening of the genitalia lies-
a short distance behind and below the right (or in sinistral species
the left) eye-peduncle. This opens into a short chamber the atrium-
(Frontispiece, ar.), from which the penis (p.) branches toward the
digestive tract, and the vagina (vag.) toward the outer side. The
penis (p.) is a tube with muscular walls, usually corrugated within,
and sometimes having longitudinal fleshy pillars (pilasters, pi. 21
fig. 14, 1 5) adherent along one side to the wall of the cavity. At its
distal end the vas deferens (v. d.) enters, its opening being sometimes
at the base or summit of a papilla (the penis papilla, pi. 28, fig. 2)^
The penis retractor muscle (r.) is inserted on the penis or its append-
ages, and attached distally to the floor of the lung. The vagina(vag.y
branches above into the spermatheca duct (sp. d.) which terminates
in the spermatheca (sp.~) ; the other branch (uterus', ut.) becoming en-
larged and sacculated. At the apex of the uterus the albumen gland
(a. gl.) supplying the albumen of the eggs, is attached ; from near
its base the ovisperm duct springs, and terminates in the hermaphrod-
ite gland (h. gl.).
Besides the above essential organs, the genitalia of many snails
are complicated by the presence of various accessory organs. On
the male side the penis may bear a gland or sack of unknown func-
tion, called the appendix (see pi. 21 fig. 1, 2, 3). This structure may
be near its apex, at its base, or even on the atrium. In some groups-
the vas deferens does not enter the penis directly, but becomes
modified into a larger tube the epiphallus (epi.~) which is continued
beyond the apex of penis and frequently bears a long blind duct, the
flagellum (fl.).
The female side in some groups is provided with a muscular sac
upon the vagina (or atrium), the dart sack (d. s.), containing a
needle or dagger-like calcareous dart (see frontispiece, fig. 5, sec-
tion of dart sack, showing dart). Associated with this apparatus are
found one or several glands, various in form, the mucus glands (m.
gl.~). In certain forms there is a .curved hollow appendage high
upon the vagina, which probably serves as a receptacle for sperm-
atophores, and has been called the appendicula (see pi. 17, fig. 1)..
The duct of the spermatheca in some Helices bears a long blind
tube, the diverticulum (see pi. 63, fig. 8).
HISTORICAL SKETCH. XV11
The musculature of the genitalia is often a character of some value.
The penis retractor may be inserted either on the penis itself, or on
the epiphallus ; and in a few cases it is split, having a double or triple
insertion. Distally it is attached normally to the lung floor, but in
a few cases to the vagina, or to the main columellar retractojr .of_
foot and buccal mass. In a few groups the penis retractor is absent.
The vagina in some cases is attached to the adjacent body wall by a
broad band-like muscle. The dart sack has no retractor, but in
certain genera its apex is connected with the vagina. The retractor
of the right eye-peduncle in most genera passes between the penis
and vagina; but in a few it passes to the left of the penis. These
myologic features are of considerable importance in classification;
and the variation in the distal insertion of 'the penis retractor in some
forms, as well as the abnormal position of the eye-retractor in others,
are difficult to explain.
II. HISTORICAL SKETCH OF THE CLASSIFICATION ,OF HELICES.
Five epochs may conveniently be recognized in the taxonomic
history of land mollusks. I, Linnsean epoch; II, Lamarckian
epoch; III, Ferussacian, IV, Beckian, V, Albers-Martensian ; each
of these being initiated by the appearance of some work largely re-
modelling the system of classification.
I, 1758-1799. The LINN^AN EPOCH was characterized by the
wide limits^and heterogenous contents of its genera, although
in a broad sense most of them have proved to be natural groups.
Linnseus himself and his successors in Germany, France and Eng-
land until the time of Lamarck, are the exponents of this period.
II, 1799-1819. LAMARCKIAN EPOCH. The genus Helix of
Linnaeus was much restricted about the beginning of the present
century by the segregation of its most diverse elements by LAMARCK
and DRAPARNAUD ; the Limnophila, Clausilia, Pupa, Succinea,
Achatina, etc. being removed to form distinct genera. Within the
group of forms retained in Helix, but few divisions were made, and
such genera as were instituted during this epoch were mainly based
on one or a few peculiar species, no attempt being made to classify
the entire series. Fischer de Waldheim (about 1808), Montfort
(1810), Schumacher (1817) are the principal contributors to this
literature.
IH, 1819-1837. FERUSSACIAN EPOCH. The Tableaux Systema-
tique de la Famille des Limacons presented the first consistent attempt
XV111 HISTORICAL SKETCH.
to classify the Helices into subgeneric groups. After dividing the
shell-bearing terrestrial inoperculate pulmonates into six genera,
Helixarion, Helicolimax, Helix, Polyphemus, Vertigo and Partula,
Ferussac proposes the following system for Helix :
t Redundantes.
Volutatse, Helicoides, subgenus Helicophanta [ Daudebardia,
Aerope, Helicophanta].
Evolutse, Cochloides, subgenus .Cochlohydra [ Succinea].
ft Inclusse.
Volutatae, Helicoides, subgenus Helicogena [=all globose Helices],
subgenus Helicodonta [=all toothed Helices],
subgenus Helicigona [=all keeled toothless
Helices],
subgenus Helicella [depressed, mostly simple
lipped Helices and Zonitidce].
subgenus Helicostyla [^elevated Helices, not
keeled].
Evolutse, Cochloides, subgenus Cochlostyla [=Bulimoid forms, im-
perforate, with entire mouth],
subgenus Cochlitoma [ Liguus, Achatina].
subgenus Cochlicopa [ Glandina, Stenogyra,
Ferussacia].
subgenus Cochlicella, [=Cochlicella, Rumina,
etc.],
subgenus Cochlogena [=Limicolaria, Bulimus,
Achatinella, etc.].
subgenus Cochlodonta [=Pupa, Strophia, Gib-
bus, etc.],
subgenus Cochlodina ^Cylindrella, Clausilia,
Buliminus].
Each of these subgenera is divided into several groups designated
by terms expressive of their peculiarities, thus :
{LomastomsB, f Aplostomse.
Aplostomse, TT T * / I Lamellatse.
Hygromaa^s. ^ *%H Canaliculato.
Jleliomanes, [ Marginatse.
These divisions of the subgenera were not intended in the sense of
sub-subgenera and should not be used in such sense. Many of them
HISTORICAL SKETCH. XIX
were repeated in several subgenera, and it is only by accident that
any of them are acceptable ill form. Such names as Hyalina (Hyal-
inae, Fer.), Heliomanes, etc., cannot date from the Tableaux. The
subgeneric divisions of Ferussac's system are based almost wholly
upon contour, one of the least stable characters of Helices. -Xhe
system is, therefore, wholly artificial. Other writers of this epoch
are Kisso (1826), who by restricting the heterogeneous subgenera of
Ferussac, fixed their types ; Leach, whose subgeneric names are
quoted in the synonymy of Turton's work (1831) ; Fitzinger (1833),
who proposed generic names for many European groups ; andChar-
pentier (1837) who publishes certain names proposed by Agassiz.
The latter three authors did not work on Ferussacian lines, but may
rather be regarded as foreshadowing the next epoch.
IV, 1837-1860. BECKIAN EPOCH. A great advance in Helicol-
ogy marked the year 1837. The period of artificial classification
waned ; and with the works of HELD and of BECK a new period
dawned. Held's work applied only to the European Helices ; but
Beck included all known species in his classification. Discarding
the arbitrary contour-grouping, Beck formed his subgenera upon the
elusive and less striking, but far more stable features of shell struct-
ure and texture, form of lip and columella, etc. A large proportion
of the groups proposed in the Index Molluscorum are still retained
in essentially their original limits. Although founded upon shell
characters only, Beck's classification is a vast advance upon previous
work; and indicates a mind of rare subtlety and discrimination.
During the decade following Beck's publication, several notable
works upon Helices appeared. Swainson (1840) attempted to apply
the "quinary system," proposing at the same time some new genera.
Hartmann (1840-1844) also made additions to the list of names
and PPEIPFER, whose name was to be henceforth so intimately asso.
ciated with all departments of Pulmonate species-work, published
the Symbolse ad Historian! Heliceorum (1841-'42), and in 1848 the
first volume of the famous Monographia Heliceorum. PfeifFer's main
strength was in the discrimination and concise, explicit, description
of species, and in the careful sifting of synonymy ; and in these lines
his work has been of incalculable benefit to science. As a systemat-
ist his views were not especially original.
J. E. Gray issued in 1847, a list of genera with their types; and
this publication fixes definitely the type species of a number of old
genera of Helices, such as Obba, Cochlostyla, etc.
XX HISTORICAL SKETCH.
The publication of Albers' Die Heliceen, in 1 850, marked a dis-
tinct advance in the discrimination of natural groups throughout
the land snails ; but the general principles followed do not differ
radically from those of Beck. In 1855 Pfeiffer published a some-
what amplified arrangement, with some new subgeneric names ; and
in the same year the brothers Adams reached the Helices in their
Genera of Recent Mollusca. The classification adopted in this work
differs widely from previous arrangements; but as its original
features are nearly all either retrogressive or founded upon fallacious
characters, the generic and subgeneric scheme need not be quoted
here. Reeve's monograph of Helix in the Conchologia Iconica
(1851-1854) supplied the first illustrations of a multitude of species,
chiefly those of Pfeiffer. Dr. Binney's Terrestrial Mollusks of the
United States (1851-1857) gave a magnificent series of plates of
American forms, among the best portraits of snails ever published ;
and the work of Dr. Joseph Leidy therein, was the first anatomical
investigation to be made on American Mollusks.
In France, Moquin-Tandon was preparing a faunal work of the
same thorough character, which was issued in 1855, with sumptuous
colored plates and well-drawn anatomical details of the snails of
France.
Simultaneous with the last, Adolph Schmidt published his Ges-
chlechtsapparat der Stylommatophoren in taxonomischer Hinsicht
(Berlin, 1855), a classic work, ranking with that of Semper in the
grasp of principles, and laying a broad foundation for the compara-
tive study of snail genitalia. Schmidt establishes upon anatomical
data the groups Pentatcenia (=Helix s. str.), Fruticicola, Xerophila^
Campylcea, shows the true relationships of the carthusiana and
nummus groups and of H.pisana and personata, separates H. obvoluia
from the personata group, etc. Many of these notable improvements
in classification have since been completely lost sight of by recent
European conchologists, and are only of late fully appreciated.
The work of Schmidt belongs to the Beckian period only chrono-
logically. In insight and genius it is altogether modern.
V, 1860- . ALBERS-MARTENSIAN EPOCH. While several
works of the decade preceding 1860 were far in advance of the stand-
point of Beck, yet their scope was not sufficiently wide to create any
general change in the views of Helix classification held in various
countries. The appearance of the second edition of Albers' Die
HISTORICAL SKETCH. XXI
Heliceen, edited by von Martens, marked a period closed, and a new
<epoch begun.
As the classification given in this work has been the basis of nearly
all subsequent systematic arrangements, it is here quoted in full
The brackets indicate that groups so united are supposed to be-
closely allied. For purposes of comparison I have given in Roman
type the names of the super-generic groups of this volume, under
which each of the Albers Martensian subgenera falls, these groups
being as follows :
Endodontidce : Haplogona, Polyplacognatha.
Helicidce: Protogona, Teleophallogona, Epiphallogona, Belogona
(with two divisions, Bel. Euadenia and Bel. Siphonade-
nia), Macroogona.
Vitrinea.
Oenus SAGDA Beck (Teleophallogona).
Hyalosagda, Proserpinula, Odontosagda.
Genus LEUCOCHROA Beck (Belogona).
Helicacea,
' Acanthinula, Belogona.
Vallonia, Belogona.
Petasia, Belogona.
Frutiricola, Belogona.
Dorcasia,Protogon a & Belogona.
Epiphallogona.
Genus HELIX L.
( Amphidoxa, Haplogona.
j Mierophysa, Teleophallogona.
j Aerope, Rhytididse.
^ Pella, Haplog. & Zonitidse, etc.
f Patula, Haplogona.
j Charopa, Haplogona.
j Stephanoda, Haplogona. C Xerophila, Belog. Siphonadenia.
[Rhytida, Rhytididse. -j Turricula, Belog. Siph.
Jatiulus, Zonitidse. (_ Cochlicella, Belog. Siph.
Endodouta, Haplogona. ( Ochthephila, Belogona.
Sesara, Zonitidse. < Actinella, Belogona.
Pelia, Zonitidse. (_ Tectula, Belogona.
Gonostoma, Belog. si phonadenia. | Plectotropis, Belog. Euadenia.
Ophiogyrd, Protogona? \ Aegista, Belog. Euad.
< Polygyra, Protogona. ( Aglaia, Belog. Euad.
\Stenotrema, Protogona. j Campy Icea, Belog. Siph.
{Triodopsis, Prolog. & Belog. j Eurycampta Belog. Euad.
Mesodon, Protogona. \^Arwnta^ Belog. Siph. & Euad.
Laoma, Polyplacognatha. Eurystoma, Epiphallogona.
XX11
HISTORICAL SKETCH.
Euparypha, Belog. Siph.
C lachea, Belog. Siph.
x Macularia Belog. Siph.
( Iberus, Belog. Siph.
Coryda, Belog. Euadenia.
( Hemicycla, Belog. Siph.
j Plebecula, Belog., Si ph.?
(_ Leptaxis, Belog. Siph.
Pomatia, Belog. Siph.
( Thelidomus, Epiphallogona.
Cystieopsis, Belog. & Teleoph.
Plagioptycha, Belog. Euad.
Polymita, Belog. Euad.
Liockila, Epiphallog. & Belog.
^Eurycratera, Epiphallogona.
Polydontes, Epiphallog.
f Helicophanta, Macroogona.
Panda, Macroogona.
Stylodon, Macroogona, Belog.
Erepta, Zonitidse.
Dentellaria, Epiphallogona.
Cepolis, Belog. Euad.
^Pleurodontn, Epiphallogona.
Anostoma, Pupidse.
f Labyrinthus, Epiphallog.
j Isomeria, Epiphallogona.
| Caracal us, Epiphallogona.
(_ Phania, Macroogona ?
f Thersites, Epiphallogona.
I Merope, Epiphallogona.
f Obba, Epiphallogona.
-< Trachia, Epiphallogona.
(^Planispira, Epiphallogona.
P/iasis, Haplogona ?
Cfiloritis, Epiphallogona.
Pedinogyra Macroogona.
Ampelita, Macroogona.
Solar apsis, ?
( Camena Epiphallog., & Belog.
< Hadra, Epiphallog., & Belog.
(_Papuina, Epiphallog.
f Leptoloma, Belog. Euadenia.
j Geotrochus Epiphallog. & Belog.
(_ Cymotropis, Epiphallogona.
f Chlorata, Belog. Euadenia.
Cbrcuto, Belog. Euadenia.
Axinn, Belog. Euadenia.
Cullicochlias, Belog. Euadenia.
Genus COCHLOSTYLA Fer. Belogona Euadenia.
The general plan of this arrangement is to establish a series lead-
ing from Zonitoid to Bull moid shells; and the characters mainly
depended upon in the formation of groups are texture, form of lip,
and general contour of shell. In the appreciation of that indefin-
able something, which counts for so much in classifying Helices, the
authors of Die Heliceen are far beyond all previous work ; and it is-
this quality this accurate feeling for subtle affinities for which no
good reason can be given in words that has rendered this work the
basis of classification for three and a half decades, a long period in
so changeable a science as malacology.
It would be obviously unfair to criticise this great work by stand-
ards of the new anatomical classification, for excepting the Haplo-
gona, Protogona and Belogona, the Helices were practically unknown
anatomically in 1860. Compared with the new system, it is note-
worthy that the Haplogona are mostly grouped together near the
Zonitidse, where they unquestionably belong; and many other felic-
ities of grouping will be obvious to one looking over the list, besides-
the genius shown in forming natural subgenera, already referred to.
For the rest, the Epiphallogona, Belogona, Teleophallogona, Proto~
HISTORICAL SKETCH. XX111
gona and Maerooyona are indiscriminately grouped; but with the
exception of the last named, which has good conchological peculiari-
ties, one would expect this ; for there are no diagnostic characters
of these super-generic groups to be found in the shells alone.
The work of PfeifFer, although begun in the last period, extended
through the greater portion of this one. Final results of this great
series of monographs are given in the Nomenclator Heliceorum
Viventium, edited by Clessin (1878). The system of classification
differs but little from that of Albers-Martens.
The successive papers and volumes of Binney and Bland upon the
land shells of America, although based on Die Heliceen, have made
notable improvements in the treatment of cis-Atlantic groups,
largely the result of Binney's work upon the jaws and radulae of
United States and West Indian species. The work of Tryon upon
Helices has been based upon conchological studies only, and is
essentially a modified form of the Albers-Martensian. Fischer like-
wise gave no weight to anatomical characters in his treatment of
Helices.
The systematic work of Morch, although begun in 1859 (Mai. Bl.
vi, 109), belongs to this epoch rather than the last. Fully recogniz-
ing the unreliability of groupings based upon shell-contour, he pro-
poses to use the jaw as a basis for dividing land snails into primary
groups. The arrangement given is as follows, the genera of Helic-
idse being italicised:
1. OXYUNATHA. Jaw with a projecting tooth, Liniax, Vitrina,
Succinea, Helicella, Zomtes,Leucochroa, Ryssota, Obba, Caracolla,
Otala, Pleurodonta.
2. AULACOGNATHA. Jaw striated, with crenulated margin. E'ury-
omphala, Bradybcena, Sogda, Cochlicella, Rum in a, Pupa, Glaus-
ilia.
3. ODONTOGNATHA. Jaw with separated cords which form teeth at
its margin, Arion, Ariolimax, Nanina, Teba, Pomatia,Helicogena,
Helicogona (Campy Isea), Achatina, Limicolaria, Bulimus.
4. GONIOGNATHA. Othalicus, Pseudostrombus ( Liguus).
5. AGNATHA. Oleacina, Testacella.
In 1835 (Journ. de Conchyl.) this idea is further elaborated and
the Elasmognitha added.* As I have shown on a previous page
(xi), the jaw is as unreliable as the shell ; and the family groups
XXIV HISTORICAL SKETCH.
based upon it are almost always artificial. Still, the attempt to use
internal features was in itself a move in the right direction.
The above classification paved the way for the great work of Dr.
Carl Semper, Reisen im Archipel der Philippine!!, Landmollusken.
In this, the most extensive work yet published upon the soft anato-
my of land mollusks, a great number of genera in all families of
snails are made known anatomically, the following scheme of classi-
fication being adopted.
Family ZONITID^E : tail with gland ; marg. teeth aculeate, etc.
Family HELICID^E : no caudal mucus-gland.
Vitrininse: Sole divided, margined; jaw smooth; marginal teeth
thorn-like. Limax, Vitrina, Parmacella, Vitrinoconus,
Vitrinoidea, Hyalina.
Helicinse: Sole undivided ; jaw various; marginal teeth short,
several-cusped.
Oxygnatha :
Teeth unicuspid : Acavus, Gorilla, Caryodes, Panda, Cara-
colus, Labyrinthus.
Teeth broad, several-cusped.
Tentacles 2 . Janella.
Tentacles 4 ; jaw with accessory plate : Succinea.
Tentacles 4 ; jaw with no accessory plate: Oopelta,
Trochomorpha, Planispira, Obbina, Strophia,
Sagda.
Aulacognatha : Philomycus, Cionella, Tornatellina, Stenogyra,
Endodonta, Buliminus, Pupa.
Odontognatha :
No accessory organs on genitalia: Achatina, Amphidro-
mus, Bulimus, Otostomus, Partula, Hadra,
Pleurodonta, Polygyra, Trachia.
Genitalia with accessory organs : Cochlostyla, Chlorsen,
Eulota, Xerophila, and other genera [this group
of Semper's is the foundation of v. Ihering's
"Helicidse" and Pilsbry's "Belogona"].
Goniognatha : Orthalicus.
Agnatha : Rhytida, etc., etc.
Family ONCHIDID^.
Family VAGINULID^E.
Although founded upon the arrangement of Morch, this classifica-
tion exhibits a distinct advance, not only in the recognition of the
HISTORICAL SKETCH. XXV
-subordinate value of the jaw structure (which Semper considered of
much less moment them would be thought from the above table),
but in the partial recognition of the value of features of the
genitalia, teeth, mantle, foot-grooves, etc., here for the first time made
much use of in classification. The great number of genera investig-
ated anatomically, and the admirable way in which the work was
done, have made. Semper's work a classic in malacological literature.
The principal defects of the classification are the exaggerated im-
portance given to the mucus tail gland, and the structure of the
jaw. Moreover, shell characters were practically ignored an ex-
treme view, not borne out by broader investigations.
During the .Albers-Martensian epoch, much good detail work
upon the anatomy of Helices has been done by investigators using
Die Heliceen and Semper's Reisen as their main reference books.
Among these may be mentioned the work of W. G. Binney, Wieg-
mann, Pfeffer, Schuberth, Brancsik, Lehmann, Fischer, Tapparone-
Canefri, Hutton, Hedley, Suter, Hesse, Pollonera, Braun, Morse and
others referred to in the text of this volume. Moreover, the advance
in knowledge of the shell has been unparalleled, many acute and
talented conchologists giving their energies to the elucidation of the
Helix faunas of every quarter of the world, and bringing to scientific
knowledge a vast number of interesting species, as well as adding
enormously to the data for zoogeography.
During the years 1889-1892 the writer published anatomical data
upon various Helices bearing upon a new classification of the entire
group, these memoranda being practically the basis of the present
volume.
The Morphologic und Systematik des Genitalapparates von Helix,
by Dr. H. von Ihering, appearing in 1892, has exercised a wide in-
fluence upon views of Helix classification, and placed the main
European genera upon a firm basis. In this powerful essay, v. Iher-
ing adopts the second division of Sempers' Odontognatha as a group
of family rank, the Helicidce, with the following genera: Xerophila,
Fruticicola, Helix [ Pentatsenia Schm.], Campylcea, Gonostoma,
Dorcasia [=Eulota], Cochlostyla. He also treats of Neohelix
(new name for Poiygyra Say), but does not attempt to show its
affinities ; and the exotic Helices of which the relationships were
unknown to him are placed under the new genus Parahelix.
The great merit of this work lies in its advanced views re-
garding the value of the various modifications of the genitalia in
; UNIVERSITY
\ ** OF
XXVI CLASSIFICATION OF HELICES.
systematic malacology, the role played by degeneration, and in form-
ally adopting and suitably characterizing the main European genera
as originally outlined by Schmidt. In the preliminary classification-
proposed by the writer (Proc. Acad. Nat. Sci. Phila. 1892, p. 392)
these European groups were placed as subgenera of Helix, but a
fuller study of the subject has resulted in the adoption of the genera
defined by von Ihering.
III. NEW CLASSIFICATION OF HELICES.
It will be seen by reference to the preceding pages that the classi-
fication of Helices has been based hitherto mainly upon the modifi-
tions of a single organ, such as the shell or the jaw ; and that even
the best of these classifications have yet given no clue to the rela-
tions the various groups of different life-areas bear toward one an-
other, nor have they even remotely suggested any phylogenetic lines.
In the present volume the attempt has been made to found a system
of grouping based upon several organs, and one expressive of the
facts of phylogeny and zoogeography.
Single-organ classifications are even more than usually dangerous
in Pulmonates for we find that they have, like their ancestors the
Tectibranchs, an extremely plastic shell which shows many cases
of parallel or " converging" development, and frequently becomes-
reduced to a functionless remnant, in members of widely different
families, and their mouth parts are subject to great changes in
nearly allied groups. The Prosobranchs show no such wide range
of mutability in either shell or radula.
It is generally held by biologists that a classification which takes
cognizence of several totally diverse, uncorrellated organs, is more
reliable than one based upon a single organ ; for the reason that
while some one organ or system of directly correllated organs, may
independently assume similar forms in members of different stocks
or phyla, when they are subjected to similar conditions of life, the
probabilities are remote that several organs not directly correllated
will be simultaneously so modified. Again, the ancestral form of a
certain organ may be retained in several groups widely diverse in
other respects; and moreover, the taxonomic value of a given struc-
ture varies widely in different families or genera.
Another consideration of weight in selecting characters for a phy-
logenetic classification, is the fact that peripheral organs, or those-
directly acted upon by external forces, are most readily remoulded.
CLASSIFICATION OF HELICES. XXV11
or modified by these influences, while internal organs are much less
directly acted upon, and lag behind in the process of transforma-
tion. For this reason, specific characters as well as those of sections
or subgenera are mainly drawn from the shell, while generic features
are usually found in the dentition, jaw and genitalia. As a rule,
these internal organs in any genus, show a far smaller range of
variation than the shells. In this connection it may be noted that
the appendages or organs of the foot (such as operculum, mucus
glands, pedal grooves, etc.) show much less variation in any natural
genus or family than the mantle organs (shell, mantle lobes, etc.).
The generally acknowledged facts recited above, with the conclu-
sions reached regarding the taxonomic value of the shell (page vii),
the jaw (p. xi) and the teeth of the radula (p. xiii), have as their
logical outcome, caused us to form a classification of the land snails
based upon all the main features of the animal economy, special de-
pendence being placed now upon one, now upon another system of
organs. Former arrangements of the genera based upon one or two
organs, must sooner or later be abandoned. Neither jaw, teeth or
shell, taken singly, prove to be sufficiently stable, nor is v. Ihering's
primary division of Pulmonata into Micronoten (small-mantled, such
as Helix, Limax, Papa), and Meganoten (large-mantled, as Vagin-
ulus, Philomycus), any more naturaL
In the opinion of the writer, a natural classification of Pulmon-
ates should be based upon :
Organs of protection (shell, mantle, integument of body).
Organs of locomotion (foot with pedal-grooves, tail gland, etc.).
Organs of reproduction (genitalia, comparative size of eggs, etc.).
Organs of nutrition (jaw and teeth, intestinal tract, kidney).
Nervous system (including sense-organs such as tentacles, etc.).
Muscle system.
In applying this scheme to the Helices, I have not attempted to
use characters of the nervous system, partly because neither the re-
quisite time or space is at my command, partly because other organs
promised results of more immediate utility. The other organic
systems named I have tried to study impartially. Although the
foundation of this system throughout rests upon comparative anato-
my, I have been influenced in some cases, where anatomical data are
wanting- or insufficient, by the facts of geographical distribution ;
but this class of facts I have purposely held subordinate to anatom-
CLASSIFICATION OF HELICES.
ical affinities, even when as in the case of Pleurodonte (p. 86), I
could not at the time of writing, see the slightest connection between
the clearly expressed organic characters, and the apparently anom-
alous distribution.
CLASSIFICATION OF SNAILS WITH JAW AND A HELICOID OR
ZONITOID SHELL.
Key to families.
[It will readily be understood that for purposes of a key, only the
most obvious peculiarities are chosen ; too much space would be re-
quired were the diagnostic characters of all organs to be given].
I. Foot-edges with no trace of pedal grooves ; no tail gland ; sole
undivided. Side teeth unicuspid, thorn-shaped, with narrow
basal-plates. Shell with simple lip and without opaque mark-
ings, SELENITID^E.
II. Foot margin defined by a pedal groove. Shell sharp-lipped.
a. Marginal teeth with narrow, elongated basal-plates, and
either unicuspid and thorn-shaped by suppression of side
cusps, or bicuspid by elevation of outer on middle cusp.
Tail gland often present, and sole frequently tripartite,
ZONITIDjE.
b. Marginal teeth with wide, short and squarish basal-plates,
with one or several cusps, the outer cusp never elevated on
middle cusp. Shell with opaque, brown coloring or flam-
mules, usually rib-striate, the lip thin, unexpanded and
sharp, ENDODONTID^E.
III. Foot-edges without pedal grooves; no tail gland. Marginal
teeth with wide, short, squarish basal -plates and one or several
cusps, the outer cusp never elevated on middle cusp. Shell
usually with expanded or reflexed lip, HELICLD JE.
This series of families is allied on one side to groups which have
undergone degeneration of the jaw, such on Rhytididce, and on the
other to the families to be monographed in succeeding volumes of
the MANUAL. The Selenitidce and Zonitidce will not further be con-
sidered in this volume, with the exception of a single genus of the
'latter (TROCHOMORPHA, page 1), which Tryon and Fischer errone-
ously intercalated among the Patuloid snails.
CLASSIFICATION OF HELICES.
Synopsis of genera of Endodontidce.
XXIX
Polyplacognatha j ^ anctum > P- 6.
\ Laoma, p. 8.
f Flammulina, p. 10.
I Phasis, p. 36.
Endodontida3<{ Haplogona { Amphidoxa, p. 39.
j Endodonta,p. 20.
(^ Pyramidula, p. 42.
| Pararhytida, p. 52.
Synopsis of genera of Helicidce.
Protogona <(
f Penis retractor trifid, Praticolella, p. 67-
Jaw ribbed | ( lip well reflexed, Pofy-
N. American j p retr gi , ! ^ra, p. 68
| lip unexpanded, Po/t/-
t gyrella, p. 78.
Anat. unknown, f whorls rounded, S. American, Poly-
S. Amer. Papua. -< gyratia, p. 81.
Shell many-( whorls keeled, Papuan, Coxia, p. 83.
whorled.
Jaw smooth, S. African, Dorcasia, p. 172.
f No blind sack
I on vagina \
Macro-
ogona
A blind sack-}
on vag. or |
sp. duct.
Keeled, emb. whorls decussate, columella short,
with convex lobe or tooth, Stylodonta, p. 149.
f Eggs or young Not keeled, columella concave, lip narrow, Heli-
very large cophanta, p. 151.
Not keeled, lip very wide, colored, Acavus, p.
153.
Keeled; lip and columella wide, colored, Pyrochi-
Eggs of mod- lus, p. 154.
rate size Columella narrow, lip not bright, Atnpelita, p.
166.
Quoit-shaped, yellow, finely striate, Macrocyclis,
p. 165.
! Quoii-shaped, dark and solid, Pedinogyra.. p. 158.
Helicoid "j Subtrochiform, rough above, smooth below, Ano-
1 glypta, p. 159.
f Apex spirally lirate, suture crenulate, Caryodes,
Shell Bulimoid I p. 161.
j Apex beaded or smoothish, suture even, Panda.
( p 163.
Shell
f Thysanophora, p. 54.
Teleophallogona -< Sagda, p. 58.
{Zaphysema, p. 65.
XXX11 CLASSIFICATION OF HELICES.
aaa. No mucus gland at tail.
b. Holarctic forms Pyramidula, p. 42^
bb. E. Indian, Australian and Oceanic forms En-
dodonta, p. 20.
3. Jaw solid and smooth ; penis with flagellum (?) and appendix ;
shell moderately large, solid and strong. Pararhytida p. 52.
The genera of this group rest upon much slighter characters than
those of Helicidw. Flammulina, Phasis and Amphidoxa are sepa-
rated mainly upon their geographic distribution, and many prove
to constitute but one genus. Endodonta and Pyramidula are re-
tained distinct for the same reason. The genitalia of very few of
the southern hemisphere forms are known, and the jaws and teeth
are not especially characteristic.
Key to genera of Helicidce.
The author has purposely abstained from assigning subfamily
rank to the natural tribes of Helices defined below. If they be
considered subfamilies, they may bear the names 1 Polygyrinse ; 2
Acavinas ; 3 Sagdinse ; 4 Camseninse ; 5 Helicinse.
I. Genitalia simple; vas deferens inserted directly on the well-
developed, long penis, which has no epiphallus or flagellum ;
no dart sack or mucus glands; no diverticulum on spermatheca
duct ; eggs small and numerous. Jaw solid, ribbed or smooth;
marginal teeth with more than one cusp. Shell with lip thick-
ened within, expanded or reflexed, the embryonic whorls not
distinctly differentiated PROTOGONA.
II. Genitalia simple, the vas deferens inserted directly on penis or
enlarged into an epiphallus ; no flagellum. No dart sack or
mucus glands, but sometimes having a blind sack or appendic-
ula high on vagina; eggs or young at birth very large, hard-
shelled. Jaw solid, smooth or vertically striate ; teeth all uni-
cuspid ; embryonic shell large, generally distinctly differentia-
ted from later growth by diverse sculpture or a terminal wrin-
kle. Shell large and solid MACROOGONA.
III. Genital system having an epiphallus and flagellum developed
on penis, and a complicated, flagellum-like appendix, or penis
gland; female side without dart sack or other accessory appen-
dages; eggs calcareous-shelled, or young born alive. Jaw
smooth or plaited ; rhachidian teeth tricuspid, laterals bicuspid.
CLASSIFICATION OF HELICES. XXX111
Tail with a longitudinal groove above. Shell somewhat Zoni-
toid, unicolored, with sharp simple lip, neither thickened or
reflexed TELEOPHALLOGONA;
IV. Genital system having an epiphallus and flagellum on penis
(but these structures obsolete in some Pleurodontes and Plain- "
spiras) ; appendix or penis gland small if present ; female side
with no accessory appendages. Eggs small or moderate in size.
Jaw smooth or ribbed ; radula with two or more cusps on part
of the side teeth. Shell usually solid, the lip expanded or
reflexed EPIPHALLOGONA.
V. Genital system having epiphallus and flagellum (rarely want-
ing) on penis ; a dart sack and mucus glands (rarely wanting)
on vagina, and frequently a diverticulum on the long sperma-
theca duct. Eggs of small size. Jaw ribbed, smooth or plaited ;
teeth with several cusps on marginals (except in Allognathus).
Shell solid or thin, often conspicuously banded BELOGONA.
Some few exceptions to the above scheme are due to degenerative
groups of the higher tribes, which simulate lower tribes, and are
only to be correctly placed by attention to the totality of their
characters. Of this sort are Cristigibba, which by degeneration of
penis and its appendages is like the Haplogona ; and Oiliella, Meta-
frutiticola and Cochlicella, unquestionably Belogona, by the loss of
their dart apparatus resemble Epiphallogona.
Tribe I, PROTOGONA.
a. Jaw ribbed. North American forms.
b. Penis retractor with trifid insertion ; a large ac,
cessory sack on penis; shell globose, unkeeled, white-
with translucent or brown bands, lip narrowly re-
flexed Praticolella, p. 67.
bb. Penis retractor not split ; no large sack on penis ;
shell yellowish or brown.
c. Lip well reflexed, often toothed, but no inter-
nal teeth ; striate above ; spermatheca duct
short Poly gym, p. 68.
cc. Lip not in the least reflexed, but thickened
within ; texture glassy ; spermatheca duct
long Polygyrella, p. 78.
aa. Jaw and soft anatomy unknown ; shell discoidal, with many
narrow whorls,
iii
XXXIV CLASSIFICATION OF HELICES.
b. Whorls rounded at periphery; South American
Polygyratia, p. 81.
bb. Whorls carinated at periphery; Papuan region
Coxia, p. 83.
aaa. Jaw solid, smooth ; penis sack continued slightly beyond
insertion of vas deferens ; duct of spermatheca long; shell
globular or depressed-globose; S. African, Dorcasia, p. 172.
Tribe II, MACROOGONA.
a. No blind sack or appendicula on vagina or spermatheca duct.
Eggs or young at birth very large, about one-third the
diameter of adult shell. ; shell with more less reflexed lip, the
embryonic whorls distinctly demarked from after growth.
b. Shell keeled, at least when young, imperforate,
finely wrinkled, the embryonic 3$ whorls spirally
grooved or decussate ; columella short, vertical, its
inner edge with a convex lobe or acute fold. Vivi-
parous, Seychelles Is. Stylodonta, p. 149.
bb. Shell capacious, not keeled ; embryonic whorls
over one-third diameter of adult, post-embryonic
growth 1 whorls or less. Aperture large, lip nar-
row, dilated at columellar insertion ; columella
concave, toothless. Madagascar
Helicophanta,p. 151.
bbb. Shell imperforate, globose-depressed or trochoidal,
not carinated, solid, bright colored; embryonic
shell about i diameter of adult. Lip broadly
reflexed, vividly colored Acavus, p. 153.
aa. No blind sack on vagina or spermatheca duct ; junction of
nuclear shell with after growth not distinct ; lip expanded or
reflexed.
b. Shell acutely keeled, at least when young; lip
usually bright colored, the columella widened into
a flat plate. Moluccas Pyrochilus, p. 154.
bb. Shell umbilicate (except in Poacilostylus), the lip
not bright colored, not widened at columella.
Madagascar Ampelita, p. 155.
aaa. Vagina or spermatheca duct bearing a blind sack. Lip of
shell narrow or simple. Eggs and embryonic shell smaller.
b. Shell Helicoid, umbilicate, wider than high.
CLASSIFICATION OF HELICES. XXXV
c. Shell solid, dark colored, quoit-shaped with
wide umbilicus, flattened spire and subhori-
zontal, oblong mouth, the lip blunt, subex-
panded, rounded ; vagina with appendicula ;
jaw smooth. Australian, Pedinogyra, p. 158.
cc. Shell subtrochiform, conoidal and tuberculate-
lirate above, polished and one-banded below
the peripheral keel ; outer lip with a deflexed
angle ; spermatheca duct with a sack ; jaw
striate. Tasmanian Anoglypta, p. 159.
bb. Shell bulimoid, higher than wide ; outer lip neither
expanded or reflexed.
c. Upper whorls spirally lirate, with crenulated
suture. Tasmanian Caryodes, p. 161.
cc. Upper whorls finely beaded or smoothish,
suture even. Australian Panda, p. 163.
aaaa. Soft anatomy unknown ; teeth all unicuspid ; shell quoit-
shaped, with wide umbilicus and low spire, uniform yellowish,
densely striate ; lip narrowly expanded throughout. S. Amer-
ica Macrocyclis, p. 165.
Tribe III, TELEOPHALLOGONA.
a. Shell smooth, depressed or trochoidal, light yellowish, having
the texture of Hyalina, composed of 6-9 narrow, closely
coiled whorls. Aperture small, narrowly lunate, often with
internal laminae, the lip thin, sharp and simple Sagda, p. 58.
aa. Shell globose, imperforate, of 5-6 convex whorls, the last
large, inflated, brown ; aperture large, rounded-lunate, tooth-
less, the lip thin and sharp, closely appressed at the white-
calloused columella Zaphysema, p. 65.
uaa. Shell conic or depressed, thin, not opaque, pale brownish or
corneous, umbilicate ; surface rather dull, often bristly or with
delicate riblets; whorls 4-6, separated by deep sutures.
Aperture round-lunate or oblong, toothless ; lip thin, often a
trifle expanded Thysanophora, p. 54.
Tribe IV, EPIPHALLOGONA.
a. Epiphallus developed, flagellum usually present, but short.
b. Penis retractor inserted at apex of penis ; Ameri-
can Pleurodonte, p. 84.
XXXVI CLASSIFICATION OF HELICES.
bb. Penis retractor inserted on epiphallus ; Old World*
c. Penis with a feather-shaped appendix ; jaw
smooth Ob ba, p. 107.
cc. No such appendix.
d. Apex or whole shell with points in
quincunx ; jaw ribbed Chloritis, p. 117.
dd. Genitalia unknown ; jaw ribbed ; shell
imperforate, with columella wide above
Albersia, p. 124.
ddd. Not so sculptured.
6. Solid, capacious, rough sculp-
tured ; embryonic shell rather
large Camcena, p. 101.
ee. Rather solid, depressed, depress-
ed-globose or keeled Thersites,
p. 125 ; Planispira, p. 110.
eee. Trochoidal, thinner and mostly
light colored Papuina, p. 136;
Ganesella, p. 168.
aa. Epiphallus or flagellum more or less obsolete.
b. Epiphallus more or less obsolete, flagellum present
Polydontes, etc., p. 87-
bb. Epiphallus and flagellum obsolete Cristigibba,
p. 112.
Tribe V, BELOGONA.
a. Mucus glands sacculated, club-shaped, bulbous or flattened,
glandular, inserted on dart sack or at its base, never on vagina
above dart sack (except in Lysinoe, p. 191, in which there are
3 club-shaped glands on vag.) Belogona Euadenia, p. 175.
aa. Mucus glands tubular or finger-like (except in Elona, p. 307),
and always inserted on vagina, never on dart sack or accessory
sacks ' Belogona Siphonadenia, p. 235.
KEY TO GENERA BY SHELL CHARACTERS.
A key to groups of Helices based upon shell features only, cannot
be made without numerous double entries, and even then to be
exhaustive it would be extremely complicated, probably too com-
plex to be of use to beginners in the science, for whom alone it
CLASSIFICATION OF HELICES.
XXXV11
be intended. The following table simply shows the genera
arranged according to some of the more obvious shell characters.
I. Shell with lip thin and sharp, as in Zonites, not expanded,
reflexed or with a rib-like thickening within.
1, American : a, no internal teeth or laminae : Pyramidula 42,
Punctum 6, Amphidoxa 39, Hyalosagda 61,
Thysanophora 54, Zaphysema 65, Glyptos-
toma 192, Polymita 184.
6, with internal teeth or laminae : Helicodiscus
51, Sagda 58.
2, Old World : a, no internal laminae or teeth : Pyramidula,
Punctum 6, Phrixgnathus 9, Flammulina 10,
Phasis 36, Charopa 22, Pararhytida 52,
Anoglypta 159, Caryodes 161, Panda 163,
Pupisoma 52, Acanthinula, Chalepotaxis 167.
b, with internal laminae or teeth : Atlantica 50,
Laoma 8, Endodonta 20.
II. Shell with lip blunt, hardly or not at all expanded, usually
thickened within.
1, American : Polygyrella 78, Polymita 184.
2, Old World : Pedinogyra 158, Leucochroa 232, Helicella 245,
Geomitra 238, Hygromia 269, Acanthinula
280.
III. Shell with lip expanded, not flatly reflexed.
1, American: Praticolella 67, Polygyratia 81, Macrocyclis
165, Thysanophora 54, Pleurodonte 84, Cepolis
177, Lysinoe 191, Epiphragmophora 193, Oxy-
chona 189, Vallonia 282, SoJaropsis 166.
2, Old World: Coxia 83, Dorcasia 172, Stylodonta 149, Heli-
cophanta 151, Ampelita 155, Camaena 101,
Obba 107, Chloritis 117, Albersia 124, Thersi-
tes 125, Planispira 110, Papuina 136,Ganesella
168, and most genera of Belogona.
IV. Shell with the lip decidedly reflexed, often toothed.
1, American : Polygyra 68, Vallonia 282, Pleurodonte 84,
Lysinoe 191.
2, Old World : Acavus 153, Pyrochilus 154, Camama 101, Obba
107, Chloritis 117, Thersites 125, Planispira
110, Papuina 136, Helicostyla 216, Chloraea
XXXV111 GEOGRAPHIC DISTRIBUTION.
214, Eulota 200, Vallonia 282, Helicodonta
284, Helicigona 296, Helix 311, Plectopylis-
143, Gorilla 147.
IV. DISTRIBUTION OF HELICES IN TIME AND SPACE.
The bare facts of distribution of the several genera and species-
are sufficiently stated in the systematic portion of this work ; it
remains to draw the more obvious conclusions which they indicate.
As to means of distribution, there is much reason to believe that
upon continental areas, land snails, like mammals, Lave been mainly
dependant upon their own powers of locomotion, although rivers
with their flood-carried debris have doubtless been effective. Such
island faunas as are not traceable to former land connections, are
probably due to drift wood and " floating islands " swept from rivers ;
for although in rare cases the agency of birds or cyclones may have
been efficient, still the evidence of such means of transport of land
snails is extremely slight, and the facts now known do not warrant
or call for any extensive invocation of means so extraordinary and
exceptional, and so completely hypothetical. It will readily be
understood that the case with freshwater snails is quite a different
subject.
The key to the wide distribution of many genera or super-generic
groups of terrestrials, is the known fact of their vast antiquity,
which has enabled them to take advantage of the various land com-
binations of several geological periods, and also of the rarely occur-
ring means of transport mentioned above.
The fact must constantly be borne in mind that the evolution of
Pulmonates has been excessively slow ; and although the terrestrial
forms have changed more rapidly than the freshwater mollusks,
they cannot be compared with mammals or birds in this respect..
Many genera of Helices dominant to-day, are known to have existed
in the early Miocene, and apparently as distinct then as now. In
the Eocene, forms less close to the recent occur, but in many cases
they cannot be generically different. In the mammalia we find the
roots not only of families, but of orders in Eocene strata, while
even the genera of Helices have scarcely changed since that time.
The super-generic groups must, therefore, strike deep into Mesozoic
time. As the means of transport of land snails are very limited
and slow, they lag far behind such freely mobile creatures as mam-
mals and birds ; and, therefore, we do not find, nor can we expect to-
GEOGRAPHIC DISTRIBUTION. XXXIX
find that the life areas defined by mollusks and those based on the
vertebrates named, correspond in all respects ; although the much
greater time limit in the case of mollusks to some extent offsets their
slower movements. The same factor of greater antiquity introduces
another disturbing quantity into the equation ; for land mollusks _
have been able to take advantage of early continental and insular
connections which no longer existed when the modern orders of pla-
cental mammals came upon the stage.
In the following pages, the distribution of the Helices will be dis-
cussed in order of groups. It will readily be understood that the
hypotheses offered, whether borrowed or original, are simply sugges-
tions, subject to such changes as the study of other groups or of
palaeontology demands, or to complete rejection. They are based,
however, on a careful consideration of the facts now known, with
regard to land snails generally; and are, I trust, fair inferences
from these facts.
Endodontidce. As will be seen in the systematic part of this work,
this family is intermediate between Zonitidce and Helicidce in its
characters, and it is decidedly less specialized than either. While it
may not be in the direct line of descent of these two families, it is
certainly nearer than either of the others to the common ancestor of
the three, as is shown by its unspecialized jaw, teeth, genitalia and
shell. Palaeontology has yet given but little to the history of the
group, but that little is significant; the Carboniferous of Nova Sco-
tia has afforded a small Helicoid described as Zonites prisons Cpr.,
which in form and ribbed-striate sculpture can only be compared to
such Endodontidce, as Pyramidula or Charopa. In my opinion this
species is to be regarded as the oldest form of Helicoid yet known,
and as a probable member of the genus Pyramidula.
Agreeing with this view of the antiquity of the group is the fact
that the Endodontidce have a wider geographic range now than
either Helicidce or Zonitidce, inhabiting the entire Holarctic realm,
the southern extremities of S. America and Africa, Australo-Zea-
landic land, and almost all oceanic islands of the entire globe.
Upon the continents they are very scarce or absent in the tropics,
probably from the competition of numerous newer groups ; and it
is mainly in island faunas, where they do not compete with true
Helicidce, that snails of this family abound. The presence of very
similar forms in southern South America and Tasmania and New
Zealand, has been accounted for by the hypothesis of a former more
xl GEOGRAPHIC DISTRIBUTION.
extensive Austral continent or " Antarctica," which may have been
supplied with these snails as well as with certain marsupials, fishes,
etc., from Australia, and subsequently became united at Cape Horn,
transferring the fauna. The connection could hardly have been in
a reverse order, or why should not Edentates and Hystricomorph
Rodents have invaded Australia ? The principal papers bearing on
such continental connections in relation to mollusks are those of
Hutton, von Ihering and Hedley. It is obvious that the Endodon-
tidcB and ffelicidce alone are insufficient to base much speculation
upon regarding former extensions of Austral land. A similar ques-
tion occurs with regard to the fauna of South Africa, which in the
presence of Endodontidce, JKhytididce, Cceliaxis, etc., shows affinity
to that of New Caledonia, Australia and Tasmania. The flora,
according to Hooker, also has affinities with the West Australian.
Helicidce-Protogona. This group, as the name implies, is believed
to be cearer the ancestral stock of the family than the other groups,
mainly, because of the simplicity of the genitalia, which are as in
Endodontidce, the less modified Zonitidce, the Rhytididce, etc. The
palseontological history of the group is very scant, a few species
entirely modern in aspect being found in Miocene strata of Florida.
Some forms of equal or greater age are reported from the western
United States, but none of them are really known to belong to this
group. The references to Triodopsis and Mesodon by writers on the
European Tertiary are groundless, the supposed Triodopsis belong-
ing to Isognomostoma, the Mesodons to Mesodontopsis, a group near
Tac heocampylcea.
Of the living forms, Polygyra, Polygyrella and Praticolella are
exclusively North American, the first named having a few species
in the West Indies, and a few which have penetrated from the head
valleys of the Missouri to those of the Columbia, and thus reached
the northwest coast, the others being East American. There can-
not be much doubt that the ancestors of this group of genera have
occupied East American soil ever since it had a fauna of ffelicidce,
and with the Pyramidulas, to the exclusion of other groups of Hel-
ices. In South America the genus Polygyratia occurs ; and while it
is likely that its affinities and past history are similar to the preced-
ing North American forms, no safe conclusions can be drawn until
the anatomy is known. The species from New Guinea and New
Ireland, grouped under Coxia, are also beyond the limit of profitable
speculation.
GEOGRAPHIC DISTRIBUTION. xl
The South African genus Dorcasia, although so widely separated
geographically, seems to be a member of this group of genera. It
is probably a remnant of a large number of Protogona which may
have had a wide range in the Eastern Hemisphere in Mesozoic
times.
Macroogona. This group comprises all the large Helices, in fact
all the Helicidce of Madagascar and the Seychelles, with genera in
Ceylon and Moluccas, and another group of genera in eastern
Australia and Tasmania (see page 148). No profitable speculations
can now be based upon this peculiar range, which probably dates
from Mesozoic time. The largest known Helices belong to this
group, as well as some very handsome forms, such as Acavus hcema-
stomus and Pyrochilus lampas, described in the last century. As a
temporary expedient, we have placed the N. Chilian group Maerocy-
clis here, but it may prove to belong elsewhere, possibly to Proto-
gona, when the genitalia come to be examined.
Teleophallogona. As stated on p. 56, this group, consisting of
three genera only, is essentially West Indian. Zaphysema is re-
stricted to Jamaica ; Sagda is nearly as local, although a few species
from Hayti and Cuba are referred here ; while Thysanophora is
universally diffused throughout the West Indies, and occurs on the
mainland from Trinidad to Florida.
Epiphallogona. The range of this group of genera includes Aus-
tralia (but not Tasmania), the Solomon Islands (but neither New
Caledonia or New Zealand), New Guinea north throughout the East
Indies, and the mainland of Asia from India to Japan. In Amer-
ica it covers the West Indies and northern South America. The
majority of genera and species are insular.
Arising from an Oriental Protogonous stock now extinct, prob-
ably a remnant of the same which had much earlier given birth to
the Macroogona, this tribe seems to have radiated in all directions.
There is no evidence showing that it ever extended further west than
at present; but in the north it evidently passed over a Bering
bridge, and travelled southward in America, becoming established
in the West Indies, probably in Secondary times. In this invasion
of American soil, the ancestors of the West Indian and Mexican
genera of Cyclostomatidce and Cyclophoridce probably shared, the
nearest allies of these groups being Oriental forms.
Whether the American Clausilias accompanied this early exodus,
or a later one, remains uncertain ; and the same is true of the
xlii GKOGHAPHIC DISTRIBUTION.
Gl'indinidce and StrcptaxidcB, which, indeed, may have originated in
America. On the south and south-east, the Oriental area of Epi-
phallogona overlaps somewhat that of the much older Austral fauna
of Endodontidce, Rhytididce etc., which lies mainly south and east
of the range of the other group. Similarly, the Epiphallogona extend
southward far beyond the range of the Belogona. The succession
of these faunas from south to north in this Asio-Australian belt of
islands, is extremely significant, and clearly indicates the compara-
tive ages of the groups in that region. The chronological order of ap-
pearance of Endodontidse, Macroogona, Epiphallogona and Belo-
gona, as determined by theoretical grounds from their comparative
anatomy, coincides with the evidence given by their geographic dis-
tribution.
Belogona. By comparing the organs of such an Epiphallogonous
form as Chloritis (pi. 28, figs. 1-4) with some Asiatic or American
Belogona, such as Monadenia, pi. 59, figs. 81, 86, or Mastigeulota^.
pi. 66, fig. 26, it will be noticed at once that the structure of the
male genitalia is identical in the two groups; each having a short
penis continued in an epiphallus which bears the retractor and ends-
in a flagellum. The female side is alike in the two groups in hav-
ing the spermatheca duct long and branchless, the other organs
being identical except that in the Belogonous groups the dart ap-
paratus is added. The jaw, teeth and shell show no features diagnos-
tic of the groups Epiphallogona and Belogona. It is, therefore,
highly probable that the latter group originated from the former,
merely adding the dart apparatus to the characters already pos-
sessed by the parent stock. There is no especial reason for believ-
ing that this transformation took place in any other area than that
now occupied by the most nearly allied modern forms of each of
these groups, viz. southeastern Asia or the adjacent island groups.
The evidence derived from comparative anatomy tends to show that
the dart apparatus of the Helices was evolved de novo in this group,
and while analogous to that of the Zonitidce, it is not homologous.
As in Zonitidce, the glands associated with the dart sack were origin-
ally proliferations from that sack ; and this structure is still retained
in the Oriental and American genera constituting the BELOGONA
EUADENIA. In the European group of genera the glands have
moved from the dart sack to the vagina, and are generally found
inserted above, never below, the insertion of the dart sack. This is
a purely secondary change, and together with the modification of
GEOGRAPHIC DISTRIBUTION. xliii
the glands into the tubular or finger-like form, is characteristic of
the BELOGONA SIPHONADENIA.
The Belogona Euadenia in the Old World extend from Japan
and India southward throughout the East Indies, with a few Cor-
asia-like forms in New Guinea and the Solomons. That they are
chronologically a later element than the EpipJiallogona is shown by
the fact that they are represented in the southern and southeastern
portion of this range by only one genus (ffelicostyla*), and even this
is much restricted, being absent in Australia, the Louisiades and
New Hebrides, etc., where Epiphallogona are well represented. On
the north, the mainland of Asia offered easy passage to Japan ; and
during a period of mild climate in high latitudes, and of elevation
of the Bering Sea region, the Euadenia penetrated westward to
America and south east to California, Mexico and South America,
crossing to the West Indies by way perhaps of a Yucatan-Cuba ridge
of elevation.
The date of this exodus of Asiatic life we are unable now to fix ; but
it could hardly have been later than the beginning of the Eocene, and
there are good reasons for believing it earlier. At the same time, while
it may have been coincident with the ingress of Epiphallogona into
America, it was probably later ; for no Belogona reached the Caribean
chain (where a well differentiated group of the other tribe is uni-
versally represented), and its distribution eastward in South Amer-
ica is less great. In North America the barrier to eastward dis-
tribution has apparently been due to extensive inland seas in the
Rocky Mountain tract, and upon their disappearance to arid clima-
tic conditions. At all events, we now have in America several
sharply defined generic types : Cepolis, the peculiarities of which
have been evolved on Antillean soil, and which gave rise to a side
line of arboreal snails, Polymita, the early origin of which is shown
by its retention of three cusps on all teeth ; a feature now lost .in
the other genus, some divisions of which have also assumed arboreal
life, with its consequent remodelling of the radula. On the main-
land the Mexican genus Lysinoe offers characters clearly telling of
ancient divergence; and this is supported by the discovery of a
species apparently allied to L. ghiesbreghtiiin the Puerco Group or
New Mexico, this Eocene horizon being below the Wasatch Group,
immediately above the Laramie (H. naeimientensis White, Bull. TL
S. Geol. Surv. no. 34, 1886, pi. 5, f. 3-7). Associated with this
Lysinoe in the Puerco are Holospira and numerous fresh-water forms.
xliv GEOGRAPHIC DISTRIBUTION.
Several Eocene species from Utah and Wyoming are probably refer-
able to Epiphragmophora ; and perhaps the Miocene Helix leidyi
Hall & Meek belongs here also ; though the condition of preserva-
tion of these fossils of the fresh -water strata of the West, is quite in-
sufficient for positive generic identification, which must await the
finding of more perfect material.
Returning to the Palsearctic region, we observe that a few species
of Eulota have penetrated into Central Asia, and one, E. frvticwn,
as far as eastern Europe. This form is evidently a late-comer,
being absent from the loess fauna, and belonging to a section of
Eulota characterized by the degeneration and loss of the flagellum.
Its late advent in Europe may be correllated with the presence in
China of a few European types such as Helicodonta and Metodontia.
The Belogona Siphonadenia are par excellence the Helices of Eu-
rope. Judging purely by the present distribution of the group, its
diagnostic peculiarities seem to have been assumed in the European
or adjacent tracts, whither the ancestral stock of Belogona Euadenia
had emigrated from the Orient. Probable companions of these
Belogona were the terrestrial operculates (some of which have been
erroneously referred to West Indian genera), and perhaps the Agn-
atha, although the origin of these is problematic. In this European
extension of the Palsearctic fauna the Siphonadenious phylum has
split into numerous genera, and apparently has crowded out any
earlier Helices of simpler structure, if such ever existed in that
quarter of the world. The old families Endodontidce and Zonitidce
retained their place owing probably to the notably different stations
occupied by them. Very early branches of the European Belogona
were Leucochroa, a probable remnant of the original stock which
did not share the changes resulting in modern Siphonadenia; and
Vallonia, a genus well differentiated in the early Eocene of Europe,
now more widely dispersed than any other genus of Helicidce, and
possibly antedating the European immigration. Further notes upon
the Belogona Siphonadenia will be found on pages 235-237. The
only Siphonadenia which have strayed far from the area now
occupied by the majority of the genera, are certain Chinese forms
referred to Helicodonta and Hygromia (q. v.), which from their close
resemblance to European types are probably recent colonies moving
eastward through Siberia. Thus, Metodontia seems closely allied to
Dibothrion, a group of middle Europe and Siberia ; and H. bicon-
cava of China is nearly allied to the European Miocene H. involuta
GEOGRAPHIC DISTRIBUTION. xlv
as the Chinese H. binodata is to certain living and tertiary European
species.
* * *
Summary by Continents. The Americas are poor in autochthonous
types of Helices (and land snails generally), the genera Polygyratia,
Solaropsis and Macrocyclis being the only South American forms of
great antiquity, the genera Epiphragmophora, Pleurodonte and
probably Oxychona having been derived from the north in compara-
tively recent times, and the Amphidoxa forms are in all probability
stragglers from the Australian tract.
The West Indies claim one group of genera, Sagda, Thysanophora
and Zaphysema of evidently great age and unknown ultimate affini-
ties, but the other elements, Pleurodonte, Cepolis and Polymita are
Mesozoic or early Eocene immigrants from the mainland, .and
primarily from Asia.
North America possessess in Polygyra, Polygyrella and Praticol-
ella a primitive fauna, to which has been added from Asia, the be-
logonous forms Vallonia, and the stock now differentiated into Epi-
phragmophora, Lysinoe, Glyptostoma and the West Indian genera
mentioned ; this addition can scarcely have been later than Creta-
ceous or base of the Eocene.
Africa is in the north practically a part of Europe ; but at the
Cape a Helix-fauna of as primitive a type as that of eastern North
America is found, consisting of the genus Phasis of Endodontidce and
Dorcasia, a type allied to Polygyra, and probably a remnant of the
early wider distribution of the Protogona. S. Africa has real
affinities with Australia, but whether these are due to the preserva-
tion of antique types in both tracts, or to some actual connection, re-
mains to be solved. Madagascar is much more allied to Ceylon and
Australia than to S. Africa.
Europe and western Asia. The western portion of Asia together
with Europe and North Africa, is peopled by a peculiar, highly
organized type of Helices practically confined to these regions, but
evidently derived ultimately from extreme south-east Asia or the
East Indies, by a Cretaceous (?) migration.
Eastern Asia, from Japan and China southward to Australia,consti-
tutes another great division in Helix distribution, and the middle of
this area has been in all probability the birth-place of the groups
Epiphallogona, Belogona and Macroogona. These three divisions
still occupy the region, various genera of the first, Camcena, Chloritis,
GEOGRAPHIC DISTRIBUTION.
Ther sites, Obba, Planispira, Papuina, Ganesella, being character-
istic of all portions of the tract. The Belogona have a smaller range
southward, but in the genera Helicoslyla, Eulota and their allies, ex-
tending over the central and northern portions of the region. The
several genera of Macroogona, such as Helicophanta and Ampelita
in Madagascar, Acavus in Ceylon, Panda, Pedinogyra, Anoglypta,
etc. in Australia and Tasmania, have a much broken, discontinuous
range, indicating a high antiquity and much extinction ; but the
origin of the group from Protogonous ancestors, within the general
region now covered by the several genera, is probable.
In conclusion : We find that the distribution of Helices in space
and time is not hap-hazard or erratic, as has been supposed from the
earlier classifications, and from the erroneous generic and subgenenc
references contained in works on the fossil forms, but that it is
orderly and comprehensible. We find that, whenever the data are
sufficient for judgment, the genera and species of any given life-
area exhibit such affinities to each other and to those of adjacent
areas, that no orographic changes or continental extensions other
than those recognized by geologists as either demonstrated or prob-
able, are necessary to account for the various snail faunas of to-day.
We find that not only is it unnecessary to throw land bridges across
the depths of Atlantic and Pacific to account for the distribution of
Helices, but that such hypotheses are contrary to many facts indicat-
ing that such groups of snails as are common to America and Europe,
have radiated from an Oriental center westward to Europe and east-
ward via the Bering Sea route to America, while in the far south
a hypothetical extension of the Antarctic continent fulfils the con-
ditions asked by the zoogeographer. Another fact worthy of remem-
brance is that in each faunal region, one or a few types of Helices
have been modified to fill the several stations available, and that the
most highly modified forms are generally found to be nearest akin
to the normal Helices of the same region, not to similarly modified
Helices of other regions. Thus, the groups Phengus, Papuina,
Oxychona and Leptoloma are strikingly similar, yet they are not
related to each other, but to less abnormal snails occupying their
several areas. The same is true of Caracolus and T her sites ;
Camcena, Euhadra and Hadra, Stylodonta and Columplica, Isognomo
-stoma and Triodopttis, and scores of other groups.
METHOD OF PREPARATION. xlvii
V. PREPARATION OF LAND SNAILS FOR ANATOMICAL.
STUDY.
Land snails intended for anatomical examination should be placed
-when collected in a vessel of water from which air is excluded^-
Usually twenty-four hours is a sufficient time to drown the animal,
when they may transferred to 50% alcohol and after a day to 60 and
then 80%. It is often impossible on account of lack of facilities to
observe this rule ; and in such cases the animal may be thrown into
about 60% alcohol when drowned. If time or facilities cannot be
had for drowning the snails in water, they should be killed by the
usual method, by scalding with boiling water, and then placed in
spirit not stronger than 60 % . The one process to be avoided is plung-
ing the living animal into spirit ; as this causes so much contraction
that subsequent work is very difficult. Of course even a badly con-
tracted specimen is vastly better than none ; and no malacologist
should neglect to preserve some sort of specimen of a species not
known anatomically, in view of the present condition of malacology,
and the advantage to be gained for science by the expenditure of
the small amount of time involved in preserving the soft parts.
The dissection of land snails is very easy, a shallow vessel with a
floor of blackened wax, some small scissors, a scalpel and pins being
all the material required. After removing the shell and observing
external features, an incision may be made extending from the top
of the head backward, laying open the visceral mass. The genitalia
will then be seen on the left (the head being toward the observer),
the digestive tract in the middle. Each of these systems may be
readily removed and pinned out separately for examination. Jaw
and radula may be mounted in glycerine jelly in the usual manner.
NOTE ON NOMENCLATURE.
The numerous changes from previous usage in generic and sub-
generic names of Helices, which have been introduced in this volume,
are mainly due to a rigid adherence to the rule of priority. The
older generic and subgeneric names were nearly all proposed for
miscellaneous and artificial assemblages of species ; and in these
cases we are compelled to accept these names in the sense in which
subsequent authors understood them and restricted them. For ex-
ample : Ferussac's Helicigona comprised all keeled and edentulous
Helices ; but as Risso retains under that name only the H. lapicida
and H. cornea, we must accept this restriction ; and as cornea was
NOMENCLATURE.
not included by Ferussac in his group, while lapicida was, we are
obliged to consider the latter species the type of Relicogona Fer.
Some authors demand that a generic name to be accepted, must be
not only appropriate in meaning, but also be correctly limited by its
describer; but such a course would only result in utter confusion.
Thus, if correct limitation be insisted upon, we might have given
new names to about half the genera as recognized herein, for fully
that many are composed of materials never before brought into the
present associations and groupings. Instead of such a course, we
have invariably tried to select for each group, the oldest name ap-
plied to any of its members.
Regarding specific nomenclature, we believe that the dictum,
" once a synonym, always a synonym," is the only satisfactory
course. Thus, Helix edwardsi Cox was changed to H. nigrilabris
because there was a prior Helix edwardsi of Bland ; and this change
holds, even though the shells of Cox and of Bland are now known
to belong to different genera. On the other hand, Polygyra hemp-
hilli W. G. B. is not held to be preoccupied by the earlier Helix
hemphilli Newc., because Binney described his species as a Trio-
dopsis, not a Helix; and as hemphilli W. G. B. is a Polygyra, and
hemphilli, Newc. a Pyramidula, there has never been a duplication
of the binomial term " Helix hemphilli"
(UHIVERSITY
TROCHOMORPHA.
Genus TROCHOMORPHA Albers, 1850.
Trochomorpha ALBERS, Die Heliceen, p. 116. MARTENS, Die
Hel. (edit. 2), p. 60, type trochiformis Fer.; Ostasiat. Zool., Land-
schn. p. 245. Discus ALBERS, /. c., p. 117. MARTENS 1. c., p. 61-
type metcalfei Pfr. Not Discus Fitz., q. v. Nigritella MARTENS~
Die Hel. (edit. 2), p. 63, type nigritella Pfr. ; Ostas. Landschn., p.
246. Videna H. & A. ADAMS, Gen. Rec. Moll, ii, p. 115. MAR-
TENS, Ostas. Landschn., p. 247. Sivella BLANFORD, Ann. and Mag.
Nat. Hist. (3) xi, p. 86 (1863), type castra Bens. Geotrochusv.
HASSELT, Algemeene Konst- en Letterbode voor bet Jaar 1823, p.
233 (= Trochomorpha sp. aud'Sitala sp.)
Shell varying from high trochiform to depressed lens-shaped, umbil-
icate or at least perforate ; solid and opaque, or thin and subtranslu-
cent ; carinated, at least in the young. Having 5-6 whorls. Sur-
face rather smooth. Embryonal whorl not marked off from the
after growth. Aperture basal, the upper lip terminating at the keel
or periphery ; peristome simple and sharp, or thickened and blunt,
the basal margin arcuate ; columellar margin arcuate, short, not
dilated or reflexed; ends of lip distant. Type T. trochiformis Fer.,
pi. 7, figs. 8, 9. (See also pi. 7, figs. 1-3, T. quadrasi Hid. ; pi. 7,
figs. 4-6, T. merzianoides Grt. ; pi. 7, fig. 7, T. meleagris Pfr.)
Animal : Foot long and rather narrow ; sole flat, with no trace of
longitudinal division ; parapodial groove distinct, bounding a wide
vertically grooved foot margin, and having a shallower groove
above it. Tail depressed above, rounded behind, without a mucus
gland. Back with several indistinct longitudinal rows of granules;
sides irregularly granular. Shell lappets none ; but mantle having
a wide body-lappet on the right and a small one on the left. Lung
orifice to the left of the superior angle of aperture, (pi. 8, fig. 12, 21
assimilis Grt. ; fig. 13, T. beckiana Pfr. ; pi. 9, figs. 32, 33, T. timor-
ensis Mts.).
Genitalia simple, the penis moderately long, somewhat twisted, the
retractor muscle and vas deferens entering at the apex. Sperma-
theca on a short duct. (PI. 8, fig. 9, T. assimilis; fig. 14, T. beck-
iana; fig. 17, T.troilus; fig. 19, T. subtrochiformis ; figs. 15, 16, T.
metcalfei; pi. 7, figs. 14, 15, T. planorbis~). Orifice of genitalia near
the pedal groove, below and slightly back of the right eye-peduncle.
Right eye peduncle retracted between branches of genitalia. Kidney
long and narrow.
Xlviii NOMENCLATURE.
not included by Ferussac in his group, while lapicida was, we are
obliged to consider the latter species the type of Helicogona Fer.
Some authors demand that a generic name to be accepted, must be
not only appropriate in meaning, but also be correctly limited by its
describer; but such a course would only result in utter confusion.
Thus, if correct limitation be insisted upon, we might have given
new names to about half the genera as recognized herein, for fully
that many are composed of materials never before brought into the
present associations and groupings. Instead of such a course, we
have invariably tried to select for each group, the oldest name ap-
plied to any of its members.
Regarding specific nomenclature, we believe that the dictum,
" once a synonym, always a synonym," is the only satisfactory
course. Thus, Helix edwardsi Cox was changed to H. nigrilabris
because there was a prior Helix edwardsi of Bland ; and this change
holds, even though the shells of Cox and of Bland are now known
to belong to different genera. On the other hand, Polygyra hemp-
hilli W. G. B. is not held to be preoccupied by the earlier Helix
hemphilli Newc., because Binney described his species as a Trio-
dopsis, not a Helix; and as hemphilli W. G. B. is a Polygyra, and
hemphilli, Newc. a Pyramidula, there has never been a duplication
of the binomial term " Helix hemphilli"
THf
{ UHIVEBSITY
TROCHOMORPHA.
Genus TROCHOMORPHA Albers, 1850.
Trochomorpha ALBERS, Die Heliceen, p. 116. MARTENS, Die
Hel. (edit. 2), p. 60, type trochiformis Fer.; Ostasiat. Zoo]., Land-
schn. p. 245. Discus ALBERS, /. c., p. 117. MARTENS I. c., p. 61-
type metcalfei Pfr. Not Discus Fitz., q. v. Nigritella MARTENS,
Die Hel. (edit. 2), p. 63, type nigritella Pfr. ; Ostas. Landschn., p.
246. Videna H. & A. ADAMS, Gen. Rec. Moll, ii, p. 115. MAR-
TENS, Ostas. Landschn., p. 247. Sivella BLANFORD, Ann. and Mag.
Nat. Hist. (3) xi, p. 86 (1863), type eastra Bens. Geotrochus v.
HASSELT, Algemeene Konst- en Letterbode voor het Jaar 1823, p.
233 (Trochomorpha sp. audSitala sp.)
Shell varying from high trochiform to depressed lens-shaped, umbil-
icate or at least perforate ; solid and opaque, or thin and subtranslu-
cent ; carinated, at least in the young. Having 5-6 whorls. Sur-
face rather smooth. Embryonal whorl not marked off from the
after growth. Aperture basal, the upper lip terminating at the keel
or periphery ; peristome simple and sharp, or thickened and blunt,
the basal margin arcuate ; columellar margin arcuate, short, not
dilated or reflexed ; ends of lip distant. Type T. trochiformis Fer.,
pi. 7, figs. 8, 9. (See also pi. 7, figs. 1-3, T. quadrasi Hid. ; pi. 7,
figs. 4-6, T. merzianoides Grt. ; pi. 7, fig. 7, T. meleagris Pfr.)
Animal : Foot long and rather narrow ; sole flat, with no trace of
longitudinal division ; parapodial groove distinct, bounding a wide
vertically grooved foot margin, and having a shallower groove
above it. Tail depressed above, rounded behind, without a mucus
gland. Back with several indistinct longitudinal rows of granules;
sides irregularly granular. Shell lappets none; but mantle having
a wide body-lappet on the right and a small one on the left. Lung
orifice to the left of the superior angle of aperture, (pi. 8, fig. 12, T.
asvimilis Grt.; fig. 13, T. beckiana Pfr. ; pi. 9, figs. 32, 33, T. timor-
ensis Mts.).
Genitalia simple, the penis moderately long, somewhat twisted, the
retractor muscle and vas deferens entering at the apex. Sperma-
theca on a short duct. (PI. 8, fig. 9, T. assimilis ; fig. 14, T. beck-
iana; fig. 17, T. troilus ; fig. 19, T. subtrochiformis ; figs. 15, 16, T.
metcalfei; pi. 7, figs. 14, 15, T. planorbis). Orifice of genitalia near
the pedal groove, below and slightly back of the right eye-peduncle.
Right eye peduncle retracted between branches of genitalia. Kidney
long and narrow.
2 TROCHOMORPHA.
In T. castra and T. timorensis (pi. 9, fig. 31) the duct of the sper-
matheca is very long. In all other features of genitalia, jaw and
teeth, they resemble the typical Trochomorphas. The length of this
duct may warrant the retention of the section Sivella Blanf.
Jaw arcuate, smooth, with a small median projection, or none.
(PL 8, fig. 10, T. assimilis; pi. 7, fig. 13, T. planorbis ; pi. 9, fig. 30,
T. timorensis.)
Radula : Central and lateral teeth having the strong mesocones pro-
jecting well over the posterior borders of their basal-plates, and lacking
ecto- and entocones. Outer lateral teeth at first siuuated outside, the
sinuation increasing to a denticle on the transition teeth, and
ascending on the mesocone to form the long bifid cusps of the marginal
teeth, which become very oblique (pi. 8, fig. 11, T. assimilis, central,
lateral and transition teeth, with several adjacent marginals and an
outer marginal drawn). See also fig. 18, T. subtrochiformis, show-
ing central and 1st, 12th, 13th and 24th teeth.
Of the names quoted in the reference paragraph above, none ante-
dates Trochomorpha except v. Hasselt's Geotrochus, dating from
1823 ; but as the species included by the Dutch author were not
described nor figured, and in fact remained unrecognized until v.
Martens identified them by the aid of v. Hasselt's unpublished
drawings, his names cannot have precedence for either genus or
species.
The prominent features of this genus are its simply conical or
lens-shaped, smooth shell, with toothless aperture and non-expanded
lip ; the undivided sole of the foot, bordered above by parapodial
grooves, without caudal mucus gland; the simple genitalia; smooth
jaw ; and unicuspid central and lateral, and bifid, Nanina-like mar-
ginal teeth.
Our knowledge of the anatomy of this genus hitherto has been
due to Semper's investigations. Gould has given figures of the liv-
ing animal of tentoriolum, troilus and conijormis, and Quoy and
Oaimard figure that of solarium. All of these figures agree with my
own observations and figures of T. assimilis Grt., from which the
above account is mainly drawn. Wiegmann has recently dissected
a specimen of T. planorbis Less. (Webers' Zool. Ergebnisse einer
Reise in Niederliindisch Ost-Indien, iii, p. 152, 1893). This species
shows the lower portion of the vas deferens to be dilated beyond the
apex of the penis, where the retractor muscle is inserted (pi. 7, figs.
14, 15, showing penis, etc. from both sides). The vagina is much
TROCBOMORPHA. 6
swollen between the lower end of the uterus and the opening of the
spermatheca duct, and at the upper end of this swollen portion there
is inside a whitish gland formed of one-celled club-shaped follicles (pi.
7, fig. 14a). This internal vaginal gland has not been noticed in
other species. Stoliczka lias published the anatomy of T. castra and
T. timorensis (Journ. As. Soc. Beng. xlii), finding these species to
have the structure of typical Trochomorpha except for the very long
duct of the spermatheca.
The genus Trochomorpha inhabits a vast area, and is excessively
prolific in specific and varietal forms. Its range extends from India,
central China and the Liu Kiu Is. on the north, southward to New
Guinea, the Louisiades and New Hebrides, and east to the Society
Islands. It is not known to occur in Australia, New Caledonia, or
any island having the Australo-Zealandic fauna, such as Norfolk
and Lord Howe. The species are in many cases founded upon
slight differences, and may become subject to some reduction as our
knowledge of their variation increases. For the present, it is
necessary to use great care in their description ; the width of
umbilicus compared with that of the base should always be stated.
The only genus with which species of Trochomorpha are likely to be
confused is the East Asian group Plectotropis ; this however differs
in the dilated columellar lip of the shell, etc.
Subdivisions.
Trochomorpha may be divided in to three sections: (1) TROCHO-
MORPHA s. str. (of which Nigritella is a synonym), containing the
solid, opaque, trochiform species, mainly Polynesian, (2) VIDENA
Ads. for depressed, acutely keeled, thin shelled forms, with wide
umbilicus, and (3) SIVELLA Blanf. for species having the shell like
Videna, but with a very long duct to the spermatheca.
The species of Videna occupy the entire area inhabited by the
genus, but are especially characteristic of the Philippines and east
Indies generally. Sivella is an Indo-Chinese group.
Systematic position.
The family relationships of Trochomorpha have been variously
estimated ; v. Martens (Albers, edit. 2) placing it under Nanina as
a subgenus, while Pfeiffer (Nomencl. Hel. Viv.) considers it a genus
between Leucochroa and Patula. Semper also places it among the
true Helices. The facts at present known incline me to view
TROCHOMORPHA.
Trochomorpha as a somewhat aberrant genus of Zonitidce ; and as
such it can properly claim no place in this volume. It is a signifi-
cant fact, that, so far as I know, all Zonitidce which possess a bifid
cusp upon the marginal teeth, jorm it by the elevation of the ectocone
upon the mesocone, while in those Helicidce having a long bifid inner
cusp on the marginals it is formed by the union of the entocone with
the mesocone. One of the earliest modifications of the Zonitid stock
was the loss of entocones from the marginal teeth ; but in the
Ilelicoids they persist in most genera.
Species of India, China, Farther India and adjacent islands.
T. benigna Pfr. iii, 84.
T. borealis Mlldff. viii, 119, 133.
T. cantoriana Bens, iii, 83.
T. caryx Bens, iii, 75.
T. castra Bens, iii, 84.
v. galerus Bens, iii, 75.
T. fritzei Bttg. viii, 194.
T. haenseli Sch. & Bttg. viii, 119.
T. paviei Mori, iii, 82.
Species of Andaman and Nicobar Is.
T. billeana Morch, iii, 84. T. sanis Bens., iii, 84.
T. iopharynx Morch. T. subnigritella Bedd. viii, 127.
T. kjellerupi Morch. iii, 74. T. sulcipes Morch. iii, 84.
Philippine Island species.
T. percompressa Bens, iii, 84.
T. saigouensis Crse. iii, 84.
T. sapeca Heude.
T. shermani Pfr. iii, 84.
T. subtricolor Mab. viii, 134.
T. timorensis Mts. iii, 83.
thieroti Morg. viii, 133.
T. tonkinorum Mab. viii, 120.
T. acutimargo Pfr. iii, 85.
T. albocincta Pfr. iii, 86.
T. bagoensis Hid. viii, 134.
T. beckiana Pfr. iii, 86.
v. kierulfii Morch. iii, 86.
T. bintuanensis Hid. viii, 134.
T. boettgeri Mlldff. viii, 134.
T. conomphala Pfr. iii, 84.
T. costellifera Moll, viii, 125,
T. crossei Hid viii, 134.
T. curvilabrum Rve. iii, 86.
T. gouldi Pfr. iii, 77.
T. granulosa viii, 125.
T. luteobrunnea Moll, viii, 120.
splendens Hid. non Semp.
T. metcalfei Pfr. viii, 121.
solaroides Rv. iii, 85.
T. neglecta Pils. viii, 124.
T. quadrasi Hid. viii, 122.
stenogyra Mlldff.
T. radiila Pfr. iii, 85.
T. repanda Moll, viii, 123.
T. rufa Mlldff. viii, 133.
T. sibuyanica Hid. viii.
T. splendens Semp. viii, 123.
T. splendidula Moll, viii, 123.
TROCHOMORPHA.
T. infanda Semp. viii, 120. T. stenozona Mlldff. viii, 133.
T. loocensis Hid. viii, 120. T. strigilis Pfr. iii, 85.
Species of Java, Celebes and the Moluccas.
T. bicolor Marts, iii, 82. T. planorbis Less.
T. concolor Bttg. viii, 126. v. lardea Mts. iii, 83.
T. ? costulata Marts. zollingeri Mouss. not Pfr.
T. gorontalensis Mts. iii, 83. T. sculpticarina Marts, iii, 80.
T. hartmanni Pfr. iii, 83. T. staudingeri Anc. viii, 134.
T. planorbis Less, iii, 82. T. strubelli Bttg. viii, 126.
syncecia Mlldff. viii, 133. zonatus v. Hasselt.
v. appropinquata Marts, iii, 82. T. ternatana Guill. iii, 76.
v. lessonii Marts, iii, 82. v. batchianensis Pfr. iii, 76.
v. javanica Marts, iii, 82. T. tricolor Marts, iii, 83.
v. nummus Issel. iii, 82. T. zollingeri Pfr. iii, 82.
Species of New Guinea and dependencies.
T. exclusa Fer. iii, 85. T. nigrans Sm. viii, 128.
T. infrastriata Sin. iii, 80. v. cornea Hedl. viii, 296.
T. lomonti Braz. iii, 82. T. papua Legs, iii, 89.
T. morio Canefri, viii, 128. . T. solarium Q. & G. iii, 80.
Species of the Solomon and New Hebrides groups.
T. apia Jacq. iii, 88. T. meleagris Pfr. iii, 81.
T. belmorei Cox. iii, 76. v. sebacea Pfr.
T. catinus Pfr. iii, 74. cerealis Cox.
T. convexa Hartm. viii, 131. thorpeiana Braz.
T. crouanii Guill. iii, 90. T. membranicosta Pfr. iii, 76.
T. crustulum Cox. iii, 90. T. merziana Pfr. iii, 89.
T. deiopeia Ang. iii, 89. T. partunga Ang. iii, 81.
T. eudora Ang. iii, 88. T. rhoda Ang. iii, 88.
T. exaltata Pfr. iii, 76. T. rubens Hartm. viii, 129.
T. fatigata Cox, iii, 76. T. sanctseannse Sm. iii, 89.
T. gassiesi Pfr. iii, 8'9. T. scytodes Pfr. iii, 77.
T. godeti Sowb. viii, 129. T. semiconvexa Pfr. iii, 88.
T. henschei Pfr. viii, 130. T. serena Cox. iii, 77.
T. juanita Aug. iii, 77. T. xiphias Pfr. iii, 89.
T. matura Pfr. iii, 88. T. zenobia Pfr. viii, 131.
PUNCTUM.
Polynesian species, Pelew
T. abrochroa Crse. iii, 90.
v. pseudoplanorbis Mouss. iii, 91.
T. accurata Mouss. iii, 80.
T. alta Pse. iii, 73.
T. approximata Guill. iii, 90.
T. assimilis Garr. iii, 92.
T. concentrica Guill. iii, 81.
T. contigua Pse. iii, 78.
congrua Pse. not Pfr.
T. corallina Mouss. iii, 93.
T. cressida Gld. iii, 91.
vahine H. & J.
T. electra Semp. iii, 86.
T. entomostoma H. & J. iii, 79.
T. eurydice Gld. iii, 90.
T. fessonia Aug. iii, 79.
T. fuscata Pse.
T. goniomphala Pfr. iii, 78.
T. kantavuensis Garr. viii, 127.
T. kiisteri Pfr. iii, 80.
T. latimarginata Sm. iii, 92.
T. ludersi Pfr. iii, 92.
T. luteocornea Pfr. iii, 90.
T. marmorosa H. & J. iii, 90.
T. merzianoides Garr. viii, 132.
T. navagatorum Pfr. iii, 90.
to Marquesas groups.
T. nigritella Pfr. iii, 78.
v. oppressa Pse. iii, 78.
T. oleacina Semp. iii, 77.
T. pagodula Semp. iii, 77.
T. pallens Pse. iii, 91.
T. planoconus Mouss. viii, 132.
T. prostrata Pse. iii, 93.
T. rectangula Pfr. iii, 73.
hapa H. & J.
T. samoa H. & J. iii, 81.
T. sansitus Cox. iii, 81.
T. subtrochiformis Mouss. iii,79.
v. albostriata Mouss.
T. swainsoni Pfr. iii, 91.
v. lenta Pse.
v. scuta Pse.
T. taviuniensis Garr. viii, 133.
T' tentoriolum Gld. iii, 79.
T. therm's Garr. viii, 134.
T. transarata Mouss. iii, 79.
T. trochiformis Fer. iii, 79.
circumdata Miihl.
T. troilus Gld. iii, 92.
T. tuber Mouss. iii, 81.
T. tumulus Gld. iii, 91.
Species of unknown habitats.
T. conferta Pfr. iii, 81. T. securiformis Dh. iii, 78.
T. hidalgoana Crse. iii, 93. T. valenciennesii Guill. iii, 93.
T. pagodula Pfr. iii, 73. guilloui Pfr.
T. planissima Pfr. iii, 93. T. virgulata Sowb. iii, 77.
T. rudiuscula Pfr. iii, 93.
Genus PUNCTUM Morse, 1864.
Punctum MORSE, Obs. on the Terrest. Pulm. Maine, Journ. Portl.
Soc. Nat. Hist. 1864, p. 27. Type P. minutissimum Lea. See also
BINNEY, Second Suppl. Terr. Moll, v, Bull. Mus. Comp. Zool. xiii,
no. 2, t. 3, f. 4, 6. SCHAKO, Mai. Blatter xx, p. 178, f. A-D.
PUNCTUM. 7
JICKELI, Fauna der Land- und Siisswasser Moll. N.-O. Afrika's, in
Verb. K. Leopoldinisch-Carolinisch Deutschen Akad. der Naturfor-
scher, xxxvii, p. 54, t. 1, f. 4.
Shell minute, thin, subdiscoidal but with convex spire, openly_
umbilicated ; unicolored ; whorls about 4, convex, the apical 1
smooth, rather distinctly demarked from the following whorls, which
have oblique striae or irregular riblets and excessively fine spiral
stripe ; the last whorl cylindrical, not descending in front. Aperture
lunate, rounded; lip simple, thin. Type P. pygmceum var. minutis-
simum, pi. 1, figs. 11, 12, 13.
Jaw arcuate or horse-shoe shaped, composed of numerous (13-19)
separate rhomboidal plates, more or less overlapping, the outer
imbricating over the inner plates ; the median two or three plates
slightly separated, not overlapping.
The individual plates are composed of vertical chitinous fibers
forming a fringe at the edges (fig. 6, 7, P. pygmceum) ; the plates are
bound together by a thin transparent membrane. The number of
plates varies somewhat, P. pygmceum (fig. 6) having 19 (/Sc/m&o) ;
P. pygmceum var. minutissimum having 16 (Morse); P. conspectum
(fig. 9) having 14 to 16, P. cryophilum (fig. 5) having 13 plates.
Radula rather long and narrow ; teeth rather separated, not in
the least overlapping. Central tooth tricuspid, the mesocone longest,
but not as long at the narrow basal-plate, side cusps small. Lateral
teeth having wider rhombic basal-plates and bicuspid, the mesocone
having a longer cusp. Marginal teeth not differentiated in any way
from the laterals, but becoming lower with shorter cusps (pi. 1, fig.
8, conspectum.)
The number of transverse rows of teeth is 75 in P. conspectum. the
formula 17-1-17 (Pilsbry) ; in P. pygmceum there are 114 rows of
19-1-19 teeth (Schako) ; in P. pygmceum var. minutissimum, Morse
counted 54 rows of 13-1-13 teeth ; in P. cryophila there are 75 rows
of 16-1-17 teeth, according to Jickeli. Each transverse row bends
forward in the middle, as shown in the line above fig. 8, represent-
ing the curve of a half row.
Distribution : North America, Europe, northern Asia and north-
eastern Africa.
This genus differs from the other Patuloid Helices in having the
jaw composed of broad rhombic plates which are not in the least
soldered together, and in the peculiar form of the bicuspid lateral
teeth. It is evidently a type of vast antiquity, and probably has
8 LAOMA.
actual affinity to the Neozealandic genus Laoma ; both may perhaps
be regarded as remnants of a Palaeozoic fauna.
The minute species of Discus-\ike shells must all be re-examined
with especial reference to the characters of the jaw before a complete
list of the species of Punctum can be made. It is not unlikely that
micropleuros Paget, elachia, debeauxiana, poupillieri, aucapitaineana
and massoti Bgt., etc., will be found to belong here. For the pre-
sent it seems the wisest course to group in Punctum only such species
as are known to have the characteristic anatomical features of that
genus, leaving unexamined minute Patuloid forms in Patulastra.
The species of Punctum live upon rotten or decaying logs in forests.
P. pygmaBum Drap. iii, 29. P. conspectum Bid. ii, 203.
schwerzenbachiana Calc. P. cryophilum Mts. iii, 32.
v. minutissimum Lea.
Genus LAOMA Gray, 1849.
Laoma GRAY, Proc. Zool. Soc. Lond. 1849, p. 167 ; type Bulimus f
(Laoma) leimonias. Phrixgnathus HUTTON, Trans. N. Z. Inst. xv,
p. 136, 1882; types H. fatua=P. celia Hutt., and P. marginatus
Hutt. See also'HuTTON, Tr. N. Z. Inst. xvi, p. 168. SUTER, Tr.
N. Z. Inst. xxiii p. 92 and xxiv, p. 297.
Shell more or less trochiform, thin, perforate or umbilicate, the
periphery keeled, at least in the young ; horn-colored, striped radially
with tawny. Aperture rhombic, provided with entering lamellae, or
without them; lip thin, simple. Type L. leimonias Gray, pi. 1,
fig. 1.
Animal heliciform, the mantle subcentral, its edge slightly
reflected over the peristome ; no locomotive disc nor mucus pore.
Jaw arcuate, composed of 20 to 24 rhombic or oblong plates
which are hairy-papillose and fringed at the upper and lower mar-
gins (pi. 1, fig. 4, L. glabriuscula Pfr.)
Radula having the central tooth rather narrow, unicuspid (or tri-
cuspid), the mesocone much shorter than the basal-plate. Lateral
teeth wider, rectangular, with two cusps which are either subequal
or the inner one larger. Marginal teeth low, wide, with two short
cusps, becoming obsolete on the outermost teeth (pi. 1, fig. 3, L.
glabriuscula', pi. 1, figf. 2, L. acanthinulopxis.')
The number of teeth in a transverse row is 35.1.35 in L. marginata,
21.1.21 in acanthinulopsis, 26.1.26 in marina; the last named
species has 110 straight transverse rows.
LAOMA.
9
Distribution : All of the species known to belong to this genus
inhabit New Zealand and Tasmania. Two sections may be distin-
guished :
Section Laoma Gray, 8. sir. Aperture provided with an entering
lamella upon the columella only, or with lamellae upon columella,
parietal wall and outer and basal lips (pi. 1, fig. 1, L. leimonias).
The writer has examined specimens of all of the species ; L. pcecilo-
sticta froms a transition to Phrixgnathus.
Section Phrixgnathus Button. Shell and animal the same as in
Laoma, except that the aperture has no teeth or folds within. (Type
L. celia Hutt., pi. 1, fig. 10). This name cannot be used in a generic
sense on account of the priority of Gray's Laoma. The mere
absence of aperture-teeth is, of course, not sufficient for generic dis-
tinction.
The fibrous jaw, composed of rhombic plates bound together by
a tbin membrane only, and the peculiar bicuspid side teeth, agree
exactly with the genus Punetum ; and upon these grounds the two
genera were associated by the writer, forming the group Polyplaco-
gnatha (Proc. Acad. Nat. Sci. Phila. 1892, p. 403.)
(Section Laoma Gray.)
L. leimonias Gray, iii, 68.
L. poecilosticta Pfr. iii, 68.
pcecilocostata Pfr. olim.
L. marina Hutt. viii, 57.
nerissa Hutt.
L. pirongiaensis Sut.
Section Phrixgnathus Hutt.
L. marine Gray, iii, 37.
umbraculum Pfr.
L. conella Pfr. viii, 58.
L. ariel Hutt. viii, 59.
L. marginata Hutt. viii, 60.
L. celia Hutt. viii, 60.
L. regularis Pfr. iii, 37.
L. erigone Gray, iii, 37.
heldiana Pfr.
L. microreticulata Sut. viii, 63.
L. pumila Hutt. viii, 63.
L. raricostata Sut. viii, 100.
L. allochroida Sut. viii, 63.
v. sericata Sut. viii, 64.
v. lateumbilicata Sut. viii, 64.
L. campbellica Filh.
L. phrynia Hutt. viii, 61.
L. fatua Pfr.
L. glabriuscula Pfr. iii, 37.
L. sciadium Pfr.
L. titania Hutt. viii, 62.
L. haasti Hutt. viii, 62.
L. acanthinulopsis Sut. viii, 61
L. transitans Sut. viii, 59.
UNIVERSITY
. y
10 FLAMMULINA.
( Tasmanian Species.}
L. csesa Cox. iii, 261. L. henryana Pett.
coesus Cox. L. pictilis Tate.
v. occulta Cox. iii, 264.
Genus FLAMMULINA Martens, 1873.
Flammulina MTS., Crit. List Moll. N. Z., p. 12.
Gerontia HUTTON, Trans. N. Z. Inst. xv, p. 135. PILSBRY,
Nautilus vi, p. 55 ; Manual viii, p. 64. Family Phenacohelicidce
SUTPJR, Trans. N. Z. Inst. xxiv, p. 270, 1892.
Shell thin, varying from discoidal to subtrochiform, umbilicated or
perforated, the perforation sometimes closed ; generally somewhat
translucent ; surface striate or ribbed, often decorated with reddish
flamrnules. Embryonal 1-H whorls smoother, often spirally
striated. Aperture rounded lunar, lacking folds, teeth or internal
callus ; the lip thin and simple, somewhat dilated at the columella.
Animal with a narrow foot bearing a mucous gland at the tail,
sometimes surmounted by a papilla. Genitalia unknown ; mantle
subcentral, its margin even, and slightly reflexed over the peristome
of the shell.
Jaw delicate, composed of thin vertical laminse firmly soldered
together but showing more or less of the overlapping edge of each
plate.
The radula exhibits a considerable amount of variation in the
different species, but the extremes are connected by all intermediate
forms. That of F. (Thalassoheliji) ziczac, drawn by the author from
an Auckland specimen, on pi. 3, fig. 28, may be taken as an example-
The central tooth has a moderate or long mesocone, ectocones being
entirely lacking in some forms, present and well developed in others.
The laterals are not crowded, and generally have a long mesocone
and short ectocone, but often the entocones also are developed, mak-
ing the tooth tricuspid. The marginals are formed by the shorten-
ing of the basal-plate and increase in size and obliquity of the cusps,
the mesocone in most forms remaining distinctly larger, sometimes
becoming bifid, probably by fusion with the entocone. The ectocone
persists on the marginal teeth, either as a simple cusp, as in F. ziczac,
or becoming split into several distinct points, as in the Allodiscus
species, and in the latter the tooth becomes very wide. In one sub-
genus, Phacussa, the ectocone is lost on the marginals, but they retain
the characteristic rhomboidal basal-plate ; and Thalassohelix exhib-
its a form of marginals connecting Phacussa with the more normal
FLAMMULINA. 11
Flammulinas. In no case does the ectocone unite with and ascend the
mesocone on the marginals as is the case in all genera of Zonitidce
which retain ectocones upon the marginal teeth.
Distribution, New Zealand, New Caledonia, Lord Howe Island*
Australia and Tasmania.
A group of rather small shells differing from Zonitidce in the forms
of the marginal teeth of the radula and the plaited jaw, and from
Endodonta and its subdivisions in the possession of a well developed
caudal mucous gland. The numerous species have been distributed
into many groups which are considered genera by some authors, but
which intergrade so closely in all essential characters that I am com-
pelled to class them as sections or at most subgenera. Their differ-
ential characters seem no more generic than those distinguishing
Tachea, Macularla and Pomatia among European Helices, or Meso-
don, Triodopsis and Stenotrema among American. Genera should,
it is believed, be founded upon really tangible structural differences,
either in shell, animal or both ; and such differences these groups do
not seem to possess. They are however of value as subgeneric divis-
ions. The investigation of the genitalia may lead to more satisfactory
results, but I expect to find but little differentiation within the
genus.
Our knowledge of these forms and their anatomy is mainly due to
Professor F. W. HUTTON and Mr. H. SUTER, who have investigated
the dentition of a majority of the New Zealand species. See Hutton,
in Trans. N. Z. Inst. xiv, xv, and xvi, and Suter, in Tr. N. Z. Inst.
xxiii and xxiv. Messrs Hedley and Suter have revised the nomen-
clature in Proc. Linn. Soc. N. S. Wales (2) vii.
The characteristics of Flammulina may warrant the surmise that
they have been modified to occupy in New Zealand the place filled in
the economy of nature by Zonitidce in other regions. If this be true,
the anomalous dentition of Phacussa must be regarded as a recent
adaptive modification.
For the generic title of this group the writer, in 1892, selected
Hutton's name GERONTIA, this being the oldest designation pro-
posed for species then known to him to belong to the genus. There are,
however, three prior names, Flammulina Martens, which being the
earliest is now adopted, and Monomphalus and Rhytidopsis of Ancey,
which are also believed to apply to members of this genus. The
presence of a caudal mucous-secreting gland, however, has not been
ascertained in the species of these New Caledonian groups, so that
their relationship to Flammulina is uncertain.
12 FLAMMULINA.
The following sectional or subgeneric divisions may be distin-
guished :
Phacussa Hutt., p. 12. Phenacohelix Sut., p. 16.
Thalassohelix Pils., p. 12. Flammulina Mart., p. 17.
Gerontia Hutt., p. 14. Suteria Pils., p. 17.
Allodiscus Pils., p. 14. Hedleyoconcha Pils., p. 18.
Pyrrha Hutt., p. 15. Monomphalus Anc., p. 19.
Therasia Hutt., p. 15. Rhytidopsis Anc., p. 20.
Subgenus PHACUSSA Hutton, 1883.
Phacussa HUTTON, Trans. N. Z. Inst. xv, p. 138 (proposed for
Zonites (?) helmsi and/tt^nunota.)
Shell depressed, umbilicated ; the spire convex, periphery
rounded. Whorls striate or with fine ribs, the apical li whorls
smooth. Aperture rounded-crescentic, peristome simple. Type F.
helmsi Hutt. (pi. 3, figs. 10, 11, F. hypopolia Pfr.)
Animal elongated ; *foot very narrow and long, compressed, not
tapering, truncated posteriorly and with a caudal gland ; mantle
slightly reflected ; eye peduncles long and thick, tentacles moderate
(Hutton, Ph. helmsii.)
Jaw arcuate, slightly tapering toward the ends, with numerous
flat imbricating vertical plaits (20-25 in helmsi, about 45 in hypo-
polia) which denticulate the margins (pi. 2, figs. 2, 3, F. hypopolia.)
Dentition : About 110 nearly straight transverse rows of 15-11-
1-11-15 to 17-13-1-13-17 teeth. Central tooth with a wide meso-
cone and minute ectocones. Lateral teeth tricuspid, the entocone
minute, obsolete on the outer ones. Marginal teeth having the basal-
plate subquadrate, mesocone very long and oblique, lacking side
cusps (pi. 2, fig. 1, F. hypopolia.)
Distribution, New Zealand. The shell in this group is very similar
to that of Phenacohelix, but the marginal teeth lack side cusps, the
mesocone being strongly developed, giving it a Zonitoid aspect.
F. helmsi Hutt. F. hypopolia Pfr. ii, 181.
v. maculata Hutt. F. fulminata Hutt.
Subgenus THALASSOHELIX Pilsbry, 1892.
Thalassohelix PILS., The Nautilus, Sept. 5, 1892, p. 56, type H.
zelandice, (Gray) Hutton. Thalassia of HUTTON and other New
Zealand authors. Not Thalassia ALBERS, Die Hel. 1860, p. 59-
Not Thalassia Chevrolat, 1834, a genus of Coleoptera.
Shell umbilicated, thin, depressed or trochiform, the periphery
acutely keeled (zelandice), bluntly angled (obnubila), or rounded
FLAMMULINA. 13
(ziczac). Apical whorls most minutely spirally striated or smooth.
Aperture rather large, lip thin, simple, subreflexed at columella.
Type F. zelandice, pi. 3, fig. 29.
Animal with narrow foot bearing a caudal mucous gland with a
papilla above it ; mantle slightly reflected over the peristome.
Jaw arcuate, with flat plaits.
Dentition : central tooth with a short mesocone, the ectocones
obsolete ; laterals with a short ectocone, which disappears on the
marginals, leaving a long, oblique mesocone only (pi. 3, fig. 27, F.
zelandice.)
Distribution New Zealand and Tasmania. The shells included by
Messrs Hedley and Suter in this division are rather dissimilar in
form. The dentition resembles Phacussa in the prominence of the
mesocones and obsolescence of ectocones on the marginal teeth, and
this peculiarity also serves to distinguish Thalasso helix from Therasia,
the shell of which is of similar form. Certain Tasmanian forms have
recently been referred by Suter to this group, a relationship
previously suspected by the writer.
New Zealand species.
F. ziczac Old. ii, 210. sigma Pfr. MS.
portia Gray, ii, 213. F. propinqua Hutt. viii, 72.
kappa Pfr. F. zelandise Gray, ii, 214.
collyrula Rve. neozelanica Hutt.
F. igniflua Rve. i, 129. v. antipoda H. & J. ii, 214.
lambda Pfr. F. aucklandica (Le Guill.) Hutt.
v. obnubila Rve. i, 120. auklandica Guill.
Australian and Tasmanian species.
[Compiled by Charles Hedley.]
F. fordei Brazier. F. hamiltoni Cox.
allporii Legrand, iii, 263. ccepta Cox. iii, 263.
austriniis Cox. iii, 264. dvcani Cox. iii, 46.
fernshawensis Petterd. floodi Brazier, iii, 46.
helice Cox, iii, 261. irvince Cox. iii, 46.
macoyi Petterd. kingi Brazier, iii, 46.
medianus Cox, iii, 264. langleyana Brazier.
petterdi Cox. milligani Brazier.
positura Cox, iii, 262. pascoei Brazier, iii, 46.
tabescens Cox. plexus Cox. iii, 262.
tranquilla Cox. savesi Petterd. iii, 46.
14 FLAMMULINA.
F. georgiana Quoy & Gaimard. F. hamiltoni Cox.
F. trajectura Cox. iii, 264. scrupulus Cox. iii, 46.
F. wynyardensis Petterd. spoliata Cox. iii, 46.
stephensi Cox. iii, 46, 262.
Subgenus GERONTIA Hutton.
Gerontia HUTTON, Trans. N. Z. Inst. xv, p. 135. Gerentia
(typog. err.) in N. Z. Journ. of Sci. i, p. 476.
Shell depressed and openly umbilicated, having the contour of
Patula ; thin, rather fragile, the surface delicately sculptured with
fine cuticular riblets. Apical whorl minutely granular, or showing a
few weak spirals, having a minute perforation at the tip. Type F.
pantherina Hutton, pi. 3, figs. 1-3.
Animal heliciform, mantle rather posterior, included ; tail acute
with a mucous pore but no papilla.
Jaw vertically striated (pi. 2, fig. 5, F. pantherina.)
Dentition : central teeth tricuspid, cusps with moderate cutting
points. Laterals similar, but the ectocones larger than the ento-
cones ; transition teeth bicuspid by fusion of entocone with meso-
cone. Marginals with a broad bifid cusp (pi. 2, fig. 4, F. panther-
ma.)
The shell is like a thin Selenites with delicate close riblets. It is
more broadly umbilicated than in the other subgenera of this genus.
The two species are from New Zealand.
F. pantherina Hutt. viii, 65. F. Cordelia Hutt. viii, 66.
Subgenus ALLODISCUS Pilsbry, 1892.
Psyra HUTTON, Trans. N. Z. Inst. xvi, p. 201, 1884. First species
P. dimorpha. Not Psyra Stal, 1876. Allodiscus PILSBRY, Nautilus
vi, p. 56, Sept. 5, 1892 ; Man. Conch. (2), viii, p. 66.
Shell thin, orbicular and depressed, with low or fiat spire, rounded
periphery, and narrow or subimperforate umbilicus ; surface radially
rib-striated, not hairy, the embryonic H whorls spirally striated (pi.
3, fig. 12, F. tullia). Aperture crescentic scarcely oblique ; per-
istome thin, shortly reflexed at the columella; parietal wall nude.
Type H. dimorpha Pfr. (See pi. 3, figs. 4, 5, 6, F. planulata Hutt.)
Jaw slightly arcuate, not tapering toward the ends, flatly ribbed
or plaited (pi. 2, fig. 11, F. tullia ; pi. 2. fig. 13, F. godeti.)
Dentition : central tooth with tricuspid reflection, mesocone long.
Lateral teeth bicuspid, the entocone being suppressed, or tricuspid.
FLAMMULINA. 15
Marginal teeth broad, with 3 to 5 cutting points (pi. 2, fig. 14, F.
godeti). In F. tullia the side cusps of the centrals are minute ;
inner marginals tricuspid, outer bicuspid (pi. 2, fig. 12, F. tullia;
pi. 2, fig. 14, F. godeti). In F. dimorpha the side cusps of the
central tooth are minute ; the marginals have a long bifid inner cusp
(mesocone, or fused mesocone and entocone), and by splitting, two
ectocones.
F. dimorpha Pfr. ii, 211. F. adriana Hutt. viii, 67.
F. venulata Pfr. ii, 211. F. miranda Hutt. viii, 68.
F. cassaudra Hutt. viii, 66. F. godeti Sut. viii, 68.
F. tullia Gray, ii, 211. F. wairoaensis Sut.
F. planulata Hutt. viii, 67. F. urquharti Sut.
Subgenus PYRRHA Button, 1884.
Pyrrha HUTTON, Trans. N. Z. Inst. xvi, p. 200.
Shell depressed-globose, thin, translucent, striated and minutely
reticulated, imperforate; the periphery rounded, spire convex.
Peristorne simple, reflexed over the minute perforation. Type P.
cressida Hutt., pi. 3, figs. 17, 18, 19.
Animal heliciform, mantle subcentral, reflected over the peristome
with an even margin ; tail truncate, with a large papilla and mucous
gland.
Jaw arcuate, flatly ribbed (pi. 2, fig. 9, .P. cressida.)
Dentition : central tooth with the mesocone only developed.
Laterals bicuspid, the entocones suppressed. Marginal teeth with
several cusps (pi. 2, fig. 10, F. cressida.)
The single species inhabits New Zealand.
G. cressida Hutton, viii, 72.
Subgenus THERASIA Hutton, 1883.
Therasia HUTT., N. Z. Journ. of Sci. i, p. 477 (proposed for tamora,
Valeria and thaisa.)
Shell depressed, perforate or umbilicate, thin, with conoidal
spire; the periphery angular or subangular ; aperture round-lunar ;
lip thin, slightly reflexed at the columella. Surface striated.
Embryonic whorls spirally striated. Type T. thaisa Hutton, pi. 3,
figs. 14, 15, 16.
Resembles Allodiscus in the dentition, and the spirally striated
apex of the shell, but differs in the form and sculpture of the latter,
which is much as in section Thalassohelix.
16 FLAMMULINA.
Animal elongated ; mantle subcentral, included ; foot long and
narrow, reaching beyond the shell, rounded behind, slightly truncated,
and with a mucous gland situated under a caudal papilla (Hutton,
F. thaisa.)
Jaw membranaceous, arcuate (F. thaisa) or horse-shoe shaped
(F. decidua, pi. 2. fig. 19), with broad imbricating plates.
Dentition : central teeth narrow, with small reflection, the meso-
cone long; ectocones hardly visible. Lateral teeth with larger
reflection, the inner ones without side cusps, the outer tricuspid.
Marginal teeth in thaisa (pi. 2, fig. 21) multicuspid ; in decidua (fig.
20) first bicuspid, then tricuspid, rounded at the anterior margin ;
the outer 2 or 3 marginals are bicuspid.
Distribution : New Zealand.
F. celinde Gray, ii, 211. F. thaisa Hutt. viii, 70.
F. Valeria Hutt. viii, 69. F. decidua Pfr. viii, 71.
F. ophelia Pfr. ii, 211. F. traversi Smith, ii, 214.
F. tamora Hutt. viii, 70.
Subgenus PHENACOHELIX Suter, 1892.
Phenacohelix SUTER, Trans. N. Z. Tnst. xxiv, p. 270. Fruticicola
HUTTON, olim, not of Held.
Shell depressed, umbilicated, subdiscoidal, the spire slightly convex
orconoidal, periphery broadly rounded. Whorls finely ribbed, the
apical one smooth except for microscopic spiral strise. Aperture
lunate, the lip simple. Type F. pilula Rve., pi. 3, fig. 13.
Animal elongated, the foot narrow, projecting behind the shell ;
mantle subcentral, rather anterior, included ; eye peduncles long,
rather clavate ; tentacles moderate. Hutton, from whose paper the
above description of the animal of F. granum is quoted, does not
mention a caudal mucous pore, but it is doubtless present.
Jaw arcuate, with about 35 flat ribs which indent the concave but
not the convex margin (pi. 2, fig. 6, F. pilula.)
Dentition : central tooth with small side cusps (F. granum) or
none (F. pilula) ; laterals similar, [lacking entocones. Marginal
teeth multicuspid, the inner cusp larger (pi. 2, fig. 7, F. pilula.}
Distribution, New Zealand. The shell is very like that of Pha-
cussa but the marginal teeth differ widely.
F. pilula Rve. ii, 212. F. granum Pfr. ii, 212.
iota Pfr. F. chordata Pfr.
FLAMMULINA. 17
Subgenus SUTERIA Pilsbry, 1892.
Suteria PILS., The Nautilus, Sept. 5, 1892, p. 56, type H. ide
Gray. Ckaropa HUTTON, olim, Dot Albers. Patulopsis SUTER,
Trans. N. Z.lnst. xxiv, p. 270, 1891, type H. ida Gray ; not Patu-
opsis Strebel, 1879, a Mexican group of Zonitidce.
Shell thin and rather opaque, openly umbilicated ; discoidal, the
spire flat; periphery broadly rounded. Surface having low spirals,
and radial, undulating cuticular lamellce bearing hairs', 1* apical
whorls smooth. Lip thin, simple. Type H. ide Gray, pi. 3, figs.
24-26.
Animal rather short and narrow ; mantle subcentral, rather ante-
rior, slightly reflexed over the peristome of the shell ; foot narrow,
extending behind the shell, the tail truncated and furnished with a
mucous gland ; no locomotive disc. Eye peduncles very long,
cylindrical, approximate at their bases; tentacles long. (Hutton
for H. ide.')
Jaw with 30 flat plaits, each transversely striated.
Dentition : centrals tricuspid, the mesocone long, ectocones short
and constricted on the outer sides. Lateral teeth similar, but the
entocone smaller than the ectocone. Inner marginals with one bifid
cusp, the outer with several subequal cusps (pi. 2, fig. 8, (F. ide.}
The principal feature of the umbilicated discoidal shell is its
hairy, undulating ribs. The dentition is characterized by the pres-
ence of entocones on the lateral teeth ; but Gerontia pantherina,
Allodiscus planulata and other forms have this same feature. The
single species is from New Zealand.
F. ide Gray, ii, 210.
ida auct.
Subgenus FLAMMULINA v. Martens, 1873.
Flammulina MART., Critical List of N. Z. Moll., p. 12. HEDLEY
& SUTER, Proc. Linn. Soc. N. S. Wales, (2) vii, p. 643, 1892.
Amphidoxa of N. Z. authors, not of Alb. Calymna HUTTON, Tr. N.
Z. Inst. 1884, p. 199, not of Hiibner, 1816.
Shell narrowly umbilicated or imperforate, globose or depressed,
thin, fragile, subpellucid, composed of few rapidly widening whorls,
which are either smooth and glossy or striated. Aperture large,
rounded-lunar ; lip thin, simple, slightly expanded at the columellar
insertion. Type F. zebra Le Guill., pi. 3, fig. 23.
18 FLAMMULINA.
Animal carrying the shell subcentrally, mantle edge slightly
reflected over the peristome of the shell, with an even margin ; tail
depressed, rounded, with a mucous gland (Hutt.')
Jaw delicate, more or less arcuate, with numerous vertical plaits,
which generally crenulate the lower margin (pi. 2, fig. 14, F. corneo-
fulva. PI. 2, fig. 18, F. chiron.)
Dentition : Rhachidian teeth with the mesocone well developed,
ectocones small (absent in corneofulva). Lateral teeth similar to the
central. Marginal teeth tricuspid (sometimes 4-cuspid), in some
species the cusps coalescing on the outer teeth (pi. 2, fig. 16, F.
corneofulva. PI. 2, fig. 17, F. chiron.)
Distribution : New Zealand and Lord Howe Island.
The contour of the shell is between Hyalina and Vitrina, and in
texture it is nearly as fragile as the latter. Both striped and uni-
colored species occur. In typical Flammulina the surface is smooth
and polished. In the section Calymna Hutton (Tr. N. Z. Inst.
xvi, p. 199, 1884), the surface is finely striated. PL 3, figs. 20-22
represent G. costulata Hutt, the type of Calymna.
F. compressivoluta Rv. i, 128. F. jacquenetta Hutt. viii, 76.
omega Pfr. F. perdita Hutt. viii, 76.
F. cornea Hutt. viii, 75. F. crebriflammis Pfr. i, 130,
F. zebra Le Guill. viii, 76. F. corneofulva Pfr. viii, 76.
phlogophora Pfr. F. novarse Pfr.
flammigera Pfr. F. chiron Gray, viii, 77.
multilimbata H. & J. F. masters! Braz. viii, 294.
F. costulata Hutt. viii, 73. F. feredayi Sut. viii, 74.
F. lavinia Hutt. viii, 74. v. glacialis Sut.
F. olivacea Sut. viii, 75.
Subgenus HEDLEYOCONCHA Pilsbry, 1893.
Shell perforated, trochiform, keeled, thin, with oblique riblets and
minute spiral lines. Aperture angu late-lunate, peristome simple,
thin, slightly expanded at the columella. Type H. delta Pfr.
Animal 13 mill, in length, 'color white almost translucent ; pos-
terior part of body sharply keeled, terminating in a mucous gland; a
shallow furrow starts from the end of the tail and runs forward on each
Me to the lips, the surface below this furrow being smooth, above it
FLAMMULINA. 19
finely tuberculate. Tentacles moderately long, cylindrical. Habits
very active ; emitting, when crawling, abundance of transparent
mucous. PL 9, fig. 27.
Jaw low, arcuate, the ends rounded, recurved ; with a blunt
median projection below ; crossed by numerous fine folds (pi. 9~
fig. 28.) _
Dentition : all teeth having basal-plates of the usual quadrate
form. Centrals tricuspid, the mesocone projecting beyond the lower
margin of the basal-plate, side cusps not quite reaching half the
length of the plate. Inner laterals similar but slightly oblique ; on
he outer laterals the entocone increases and the ectocone diminishes.
Marginals with the basal-plate low and wide, bearing the large, sub-
equal ento and mesocones, and a bifid or trifid ectocone ; the extreme
marginals having an irregularly serrated edge (pi. 9, fig. 29.)
The trochoidal shell resembles that of the keeled Thalassohelix
species, but the low, wide and multicuspid marginal teeth offer a
contrast to those of that group.
Our knowledge of the anatomy of this group is due to Charles
Hedley's researches (Proc. Roy. Soc. Queensl. v, p. 152, and vi, p.
250) ; the figures were drawn from specimens collected on Little
Nerang Creek, Queensland, where it was found abundantly, on the
trunks of trees.
F. delta Pfr.ii, 215.
conoidea Cox.
fenestrata Cox.
Subgenus MONOMPHALUS Ancey, 1882.
Monomphalus . . . . Le Naturaliste 1882, p. 86 (M. bavayi and
heckelianus) ; ANCEY, Bull. Soc. Mai. Fr. v, p. 370. TRYON,
Manual i r p. 114.
Shell thin, discoidal, the spire slightly concave, umbilicus reduced
to a mere chink ; periphery broadly rounded. Sculptured with fine
riblets, the embryonal whorl showing very fine spiral striae. Aper-
ture vertical, lunate, lip thin, dilated over the perforation. Type
F. rossiteriana Crosse, pi. 3, figs. 7, 8, 9.
Soft parts unknown. Distribution, New Caledonia.
This group is very similar in shell characters to Allodiscus, and
the two may require to be united. They are here retained separate
because the anatomy of the New Caledonian forms is unknown, and
may prove sufficiently different.
20 . ENDODONTA.
G. rossiteriana Crosse. i, 114. G. gentilsiana Crosse. i, 115.
heekeliana Crse. G. cerealis Crosse. i, 114.
G. bavayi Crosse. i, 114. G. lifuana Montr, i, 115.
Subgenus RHYTIDOPSIS Ancey, 1882.
Ehytidopsis Le Naturaliste 1882, p. 85; ANCEY, Bull.
Soc. Mai. Fr. V, p. 371, 1888.
Shell globose-depressed, narrowly urabilicated, thin but rather
strong. Whorls about 5, slowly increasing, the last rounded at the
periphery. Aperture subvertical, oblong-lunate ; lip sharp, gently
sinuous below, dilated at the columella. Type H. chelonites Crosse,
pi. 6, figs. 69, 70.
Jaw widely arcuate grooved by 18 indistinct rather wide lamellae
which denticulate its cutting edge.
Lingual ribbon 1 mill, long, J mill, wide ; teeth according to the
formula 12-8-1-8-12. The central teeth are as large as the lateral,
tricuspid, the side cusps small, median cusp elongated. Lateral teeth
having a rudimentary entocone, a large mesocone, and a sloping
bicuspid ectocone. Marginal teeth forming an angle with those of
the middle field, they are spaced, gradually increasing, and bear
three cusps, the entocone and mesocone being united toward their
bases, the ectocone smaller.
Our knowlege of the dentition of H. chelonites rests upon a note
by Saint-Simon, in Bull. Soc. d'Hist. Nat. de Toulouse 1880, pp. 171,
174. The jaw and teeth agree well with those of other sections of
the genus Flammulina, but whether a caudal mucous gland is present
or not remains to be ascertained. Ancey's name appeared anony-
mously in 1882. It is not easy to decide what effect this should
have on nomenclature.
F. chelonites Cr. i, 117. F. corymbus Cr. i, 117.
v. major Anc. F. (?) minutula Cr.
F. prevostiana Cr. i, 123.
Genus ENDODONTA Albers, 1850.
Endodonta ALBERS, Die Hel., p. 89. MARTENS, Die Hel. (edit.
2), p. 90. Not Endodonta Lansb., Notes Ley den Mus. viii, p. 108
(Coleoptera), 1886.
ENDODONTA. 21
Shell small, varying from discoidal to trochiform, generally
umbilicated ; the surface striate or ribbed. Aperture varying from
multidentate to toothless; peristome simple. Type, E. lamellosa
Fer.
Animal having distinct grooves above the margins of the foot,
but no caudal mucous gland. Eye peduncles club-shaped. Genital
system simple, lacking all accessory appendages. Jaw delicate,
vertically sparsely striated. Radula having the basal-plates of
central and lateral teeth large and square ; central teeth tricuspid ;
lateral teeth tricuspid or lacking the endocones; marginal teeth
having a low, wide basal-plate, bearing 3 or 4 cusps, the endocone
and mesocone generally united at base, ectocone simple or bifid.
Distribution, Australasia and Polynesia.
This genus differs from Flammulina (and its subgenera) in lacking
caudal mucous pore, and in the striated rather than plaited jaw ;
from Pyramidula in the clavate eye-peduncles.
No one, I believe, who examines large series of the species from
various regions, will claim that the groups included as sections under
Endodonta, can be admitted as genera. They have no anatomical
differential characters whatever, as far as is now known, and the
shell features integrade by easy stages throughout.
The distinction between Charopa and Endodonta is of little value,
on account of the degeneration of the teeth in some forms of the
latter, producing species which technically fall under the former
group. In this genus, therefore, as in many others (such as Gas-
trodonta, Polygyra, Lucerna, Sagda, etc.), the presence or absence of
teeth or lamellae within the aperture is of scarcely more than specific
value, or at most, serves to define groups of no more than sectional
rank.
The principal authorities upon the shells now assembled here are
Pease, Garrett, Cox, Brazier, Hutton, Semper, Suter and Hedley.
Mousson, Gassies, Crosse and Pfeiffer have also contributed to the
literature. Notwithstanding the great amount of work which has
been done, a vast field for future labor remains. The anatomy is
but little known except in the New Zealand species, and very few
acceptable figures of the shells have been published. In this genus,
figures should be drawn of sufficient size to show clearly all features
of the shell, and this cannot be done with figures much smaller than
those illustrating the present work.
22 ENDODONTA.
KEY TO SUBGENERA.
a. Aperture provided with teeth, folds or spiral lirse.
b. Parietal callus elevated at edge into a transverse lamella ;
base glossy,
Brazier ia.
bb. Teeth or lamellae internal, spirally entering or tubercular,
c. Form elevated conical ; columella calloused, Diglyptus.
cc. Form not high conic, diam. greater than alt. ; no heavy
columellar nodule.
d. Umbilicus pouch-shaped, wide within, constricted at
opening, Libera.
dd. Umbilicus not pouch-shaped, open or imperforate,
Endodonta.
aa. Aperture lacking teeth, internal folds or lirse.
b. Spire elevated, the alt. nearly equalling or exceeding the
diam.
c. Shell cylindrical, pupiform, Phenacharopa.
cc. Shell convex-conoidal or thimble shaped, hairy,
Aeschrodomus.
ccc. Shell pyramidal-conic, spirally sculptured and pitted, not
hairy, Paratrochus.
bb. Spire depressed, convex, flat or concave ; diam. much
exceeding the alt., Charopa.
Subgenus DIGLYPTUS Pilsbry, 1893.
Diaglyptus PILS., Manual viii, p. 86, not Diaglypta Foerst., Verh.
Ver. Rheinl. xxv, p. 1 76 (Insecta.)
? Pitys BECK, Index Molluscorum, p. 9, 1837 (name only), type P.
oparana B. (undescribed). MCERCH, Catal. Yoldi, p. 6, 1852 (no
description ; H. bilamellata Pfr. mentioned.)
Not Pitys PEASE, P. Z. S. 1871, p. 450.
Shell elevated-trochiform, rather narrowly but openly umbilicated,
the two apical whorls spirally striated the remaining whorls strongly
obliquely ribbed. Aperture armed with a strong entering parietal
lamella, and two close columellar plicse terminating in a large callous
nodule on the columellar lip ; peristome expanded below. Type
Helix bilamellata Pfr.,=pagodiformis, pi. 5, fig. 54.
Anatomy unknown. The single species inhabits the island of
Opara, one of the Austral group. It was doubtless derived from the
ENDODONTA. 23
Endodonta stock, but the elevated contour and the aperture arma-
ture render it quite distinct in aspect.
E. pagodiformis Smith, viii, 86.
bilamellata Pfr. not Sowb.
? oparana Beck (undesc.)
Subgenus LIBERA Garrett, 1881.
Libera GRT., Journ. Acad. Nat. Sci. Phila. (2), viii, p. 390; ix,
p. 33. Not " Cephalopoda Libera " DEHAAN, Monographic Ammo-
niteorum et Goniatiteorum, p. 18 (1825), which was not proposed as
a generic name, and is in no sense such.
Shell depressed, widely umbilicated in the young, the umbilicus
strongly constricted in adults to form a pouch- like cavity, in which the
eggs are carried. Whorls 7-9, closely-coiled, the last generally
angular. Aperture subrhombic, provided with folds within ; lip
thin, sharp ; the columellar margin dilated, emarginate. Type E.
subcavernula, pi. 5, figs. 45, 46, 47.
Animal small, ovoviviparous ; eye peduncles long and slender,
tentacles small ; foot short, narrow, pointed behind.
Genitalia entirely simple, lacking all accessory organs (L. bursa-
tella, teste Semp. Phil. Reise, p. 135.)
Jaw of L. bursatella distinctly striated, narrow, as if composed of
fully 20. narrow lamellae; entirely similar to that of P. rotundata
Mull.
Radula consisting of 15-1-15 (recedens) to 10-7-1-7-10 (tumulo-
ides) teeth. Centrals tricuspid. Laterals lacking the entocones or
having it excessively small. Marginal teeth having a long bifid
inner cusp (entocone plus mesocone) and a small ectocone (pi. 9,
fig. 34, E. recedens Grt. ; pi. 9, fig. 26, E. tumuloides Grt.)
The prominent feature of this radula is the lack of entocones on
the lateral teeth. The jaw corresponds exactly with that of the
typical Charopas. Semper has examined the animal of bursatella ;
Binney the teeth of tumuloides, and I have examined the radula
of recedens.
This group is distinguished from Endodonta and other toothed
Patuloids by the constriction of the umbilicus. The young (pi. 5,
fig. 48, E.fratercula Pse.) contained in the umbilical pouch consist
of about H rounded, ventricose whorls, which are regularly and
finely rib-striate, showing no trace of spiral stride. The figure shows
24 ENDODONTA.
the shell seen from above. The half grown shells are widely umbil-
icated, and resemble the normal Endodontas in form and teeth.
This group also has descended from the Endodonta stock, being
differentiated only by the constriction of the mouth of the umbilicus.
Garrett writes as follows . " Remarkable for their singular habit
of ovipositing into the cavernous umbilicus. The eggs usually from
four to six, or the same number of young shells, may frequently be
seen closely packed in the cavity. The peculiar constriction of the
umbilicus does not occur until the last two whorls are completed,
previous to which it is very open or cup-shaped. Certain species
more completely secure the safety of their eggs by the formation of
a very thin shelly plate, which projects from the columellar and
parietal region and nearly closes the umbilical opening. It is sub-
sequently either broken away or absorbed by the animal to facilitate
the escape of the young shells. All the species are gregarious, liv-
ing under loose stones, rotten wood, and less frequently buried in
decaying leaves. They range from the low lands near the sea-shore
to upwards of two thousand feet above sea-level. So far as known,
the genus, which comprises about a dozen species, is peculiar to
the Society and Cook's Islands. In the former group they are con-
fined to Tahiti and Moorea."
E. cavernula H. & J. iii, 69. E. coarctata Pfr. iii, 71.
E. sculptilis Pse. iii, 70. turricula H. & J.
fratercula Pse. streptaxon Rv.
E. subcavernula Tryon, iii, 70. E. bursatella Old. iii, 71.
cavernula Garr. not H. & J. E. retunsa Pse. iii, 71.
E. tumuloides Garr. iii, 70. E. heynemanniPfr. iii, 72.
E. jacquinoti Pfr. iii, 71. E. gregaria Garr. iii, 72.
excavata H. & J. E. recedens Garr. iii, 72.
Subgenus ENDODONTA Albers.
Shell more or less depressed, varying from rounded to acutely
keeled at the periphery, umbilicus generally open, rarely minute or
closed, and never contracted at its opening. Aperture armed within
with teeth or entering plates (rarely absent by degeneration).
This group comprises a great number of species, and is especially
characteristic of the Polynesian fauna, although a few forms are
found as far to the west as New Zealand, New Caledonia and the
Philippine Islands. The species are unequally related, as is usually
the case in large groups; and several minor divisions (Thaumato-
ENDODONTA. 25
don, Piychodon, Helenoconchd] have received names. These divis-
ions or "sections" are of doubtful value, as they are practically
undiagnosable ; but still they are natural groups of species, and as
such have their uses.
Sections of Endodonta.
a. St. Helena forms Helenoconcha.
aa. Australo-Polynesian forms
b. Shell acutely keeled ; teeth generally large, rarely wanting
Endodonta s. s.
bb. Shell rounded at periphery
c. parietal wall with one or many lirse, outer lip toothless
Nesophila.
cc. outer lip toothed or lirate; parietal wall generally
toothed Ptyehodon, Thaumatodon.
Section Endodonta s. str.
Shell openly or widely umbilicated, depressed, carinated, opaque;
aperture obstructed by internal lamellae, of which there are one or
two on the parietal wall and several on the basal wall (but in E.
fabrefacta lamellae are absent). Type E. lamellosa Fer., pi. 4, figs.
40, 41. (see also E. obolus Old., pi. 4, fig. 39 ; and E. fabre/acta Pse.,
pi. 5, figs. 52, 53).
Radula having 12-6-1-6-12 teeth. Centrals square, tricuspid ;
laterals of the same size as the centrals, bicuspid, the entocone
being absent. Marginal teeth having a long bifid inner cusp and a
short bifid ectocone (PI. 9, fig. 22, E. huaheinensis Pfr.).
Sandwich Island species.
E. apiculata Anc. viii, 95. E. lamellosa Fer. iii, 67.
E. binaria Pfr. iii, 61. E. laminata Pse.
E. fricki Pfr. iii, 67. E. rugata Pse. iii, 67.
Society Island species.
E. cretacea Grt. iii, 66. E. obolus Old. iii, 61.
E. fabrefacta Pse. iii, 45. acetabulum Pse.
conicava Mouss., Schm. celsa Pse.
v. picea Grt. barffi. Grt.
E. ficta Pse. iii, 62. intermixta Mouss.
E. garrettii Anc. viii, 95.
26
E. huaheinensis Pfr. iii, 61.
aranea Behn.
ENDODONTA.
E. tanese Grt. iii, 62.
janece Schra. & Pfr.
boraborensis Pse. ms.
Pelew Island species.
E. constricta Semp. iii, 67. E. kororensis Bedd. viii, 84.
E. fuscozonata Bedd. viii, 83. E. lacerata Semp. iii, 67.
E. irregularis Semp. iii, 67.
Section Thaumatodon Pilsbry, 1893.
Pitys PEASE, P. Z. S., 1871, p. 450 (in part). GARRETT, Journ.
Acad. Nat. Sci. Phila. viii, p. 388 (1881). Not Pitys Beck, Index
Molluscorum p. 9 (1837).
Shell discoidal, the spire low, convex ; umbilicus open or closed ;
periphery generally broadly rounded ; surface rib-striate, unicol-
ored or flammulate. Aperture having internal teeth or folds upon
the outer wall, and the parietal wall, sometimes lacking upon the
latter. (PI. 4, figs. 35, 36, 37, 38, E. multilamellata Grt. See also
pi. 4, figs. 33, 34, E. derbesiana Or.).
In E. multilamellata Grt. the lamellae within the outer lip exhibit
a peculiar structure; at frequent intervals they bear long curved
hook-like processes, directed toward the aperture as shown in pi. 4,
fig. 38. This structure is well adapted to prevent the entrance of
insect enemies of the snail. No similar formation has been
described in other land snails, except in the genus Strobilops ; but
in that group the processes are upon the parietal lamellae only,
while in Thaumatodon the palatal lamellae alone are armed.
Polynesian species.
E. acuticosta Mouss. iii, 60.
E. analogica Pse. iii, 63.
E. anceyana Grt. viii, 96.
E. baldwini Anc.
v. albina Anc.
E. boraborensis Grt. iii, 66.
E. consimilis Pse. iii, 60.
societatus Mouss., Schm.
E. consobrina Grt. iii, 66.
E. contorta Fer. iii, 63.
intercarinata Migh.
E. decemplicata Mouss. iii, 63.
E. marquesana Grt. viii, 96.
E. maupiensis Grt. iii, 65.
maupitiensis Pfr.
E. multilamellata Grt. iii, 63.
E. octolamellata Grt. viii, 95.
E. opanica Ant. iii, 67.
oparica auct. iii, 67.
E. parvidens Pse. iii, 64.
incerta Mouss., Pfr.
E. paucicostata Pse. iii, 60.
E. punctiperforata Grt. iii, 66.
E. radiella Pfr. iii, 38.
ENDODONTA. 27
E. decussatula Pse. iii, 60. pardalina Dh.
E. dsedalea Old. iii, 64. undulata Fer.
E. degagei Grt. iii, 65. E. raratongensis Pse. iii, 64.
E. distans Pse. iii, 60. E. rotellina Pse. iii, 60.
E. elisse Anc. viii, 95. E. rubiginosa Gld. iii, 59.
E. filocostata Pse. iii, 60. E. rurutuensis Grt. iii. 61.
E. graffei Mouss. iii, 65. E. sexlamellata Pfr. iii, 63.
E. harnyana Anc., viii, 95. E. stellula Gld. iii, 61.
E. hystricelloides Mouss. iii, 65. E. subdsedalea Mouss. iii, 64.
E. hystrix Migh. iii, 59. E. subtilis Grt. iii, 66.
setigera Gld. ms. E. tiara Migh. iii, 38.
E. imperforata Pse. iii, 68. E. unilamellata Grt. iii, 60.
aitutakiana Mouss., Schmeltz. E. verecunda Pse. iii, 63.
E. jugosa Migh. iii, 59. E. woapoensis Grt. viii, 95.
E. lamellicosta Grt. E. zebrina Grt. iii, 64.
Species of New Zealand, New Caledonia, Tasmania and Philippine
Islands.
E. berlieri Cr. iii, 59. E. philippinensis Semp. viii, 82.
E. cryptobidens Sut. viii, 85. E. timandra Hutt. viii, 84.
E. derbesiana Cr. iii, 63. E. varicosa Pfr. iii, 23.
E. dispar Braz. iii, 59. E. vincentina Cr. iii, 59.
E. Jessica Hutt. viii, 85.
Section Nesophila Pilsbry, 1893.
Shell discoidal with open umbilicus, rounded periphery and
depressed spire ; surface generally ribbed, unicolored or flamrnu-
late. Aperture round-lunar, the parietal wall sculptured with one or
many spiral entering lirce; outer wall toothless. Type H. tiara
Migh., pi. 6, fig. 66.
The species are Polynesian in distribution. See list under Thau-
matodon, in which they are included.
Section Ptychodon Ancey, 1889.
Ptychodon ANC., Bull. Soc. Mai. Fr. v. p. 372. HEDLEY & SUTER,
Proc. Linn. Soc. N. S. Wales (2) vii, p. 652. Maoriana SUTER,
Trans. N. Z. Inst. xxiii, p. 96. PILSBRY, Manual, viii, p. 87. Stro-
bila HUTTON, olim, not of Morse. Huttonella SUTER, Tr. N. Z.
Inst. 1890, not of Pfr., 1855.
28 ENDODONTA.
Shell urabilicated, discoidal, with low-convex spire, rounded
periphery and rib-striated surface. Aperture crescentic, subverti-
cal ; outer lip thin, simple, armed a short distance within with
numerous low folds ; columellar lip bearing one or two larger enter-
ing lamellae, and parietal wall bearing one or two stout entering
plates, sometimes emarginate, or several smaller folds. Type E.
leioda Button. (PI. 4, figs. 30, 31, 32, E. aorangi Sut.).
Jaw membranaceous, slightly arcuate, with distant vertical stride
(pi. 8, fig. 6, E. microundulata).
Radula consisting of 90-100 slightly sinuous transverse rows of
teeth, the formula varying from 6-4-1-4-6 (wairarapa) to 10-7-1-
7-10 {aorangi}. Central tooth tricuspid. Lateral teeth similar,
tricuspid. Marginal teeth tricuspid or quadricuspid, the cusps
showing a tendency to coalesce on the outer ones (PI. 8, fig. 5, E.
microundulata).
This group is closely allied to the Polynesian section Thaumato-
don. The species live under bark and rotten wood, in the bush.
Our knowledge of them is due to Professor F. W. Button and Mr.
H. Suter.
E. leioda Hutt. viii, 87. E. hectori Sut. viii, 89.
E. pseudoleioda Sut. viii, 88. magdalence Anc.
E. wairarapa Sut. viii, 88. E. aorangi Sut. viii, 90.
E. microuudulata Sut. viii, 89. E. hunuaensis Sut.
Section Helenoconcha Pilsbry, 1892.
Manual of Conch. (2), viii, p. 91.
Shell discoidal, umbilicated ; aperture armed within with small
folds. Type H. polyodon Sowb., pi. 4, figs. 42, 43. Distribution,
St. Helena.
Soft parts unknown. This group is distinguished from Thauma-
todon mainly on account of its different distribution. Its generic
relationships cannot be affirmed with certainty until the soft parts
are examined. It is not improbable that the species of Patula
described from St. Helena are toothless forms of this group.
E. polyodon Sowb. viii, 93. E. pseustes Sm. viii, 92.
alexandri Fbs. E. biplicata Sowb. viii, 92.
helenensis Pfr. E. vernoni Sm. viii, 91.
E. minutissima Sm. viii, 94. E. bilamellata Sowb. viii, 91.
E. leptalea Sm. viii, 95. v. unilamellata Sm. viii, 91.
E. cutteri Pfr. viii, 93.
ENDODONTA. 29
Subgenus BRAZIERIA Ancey, 1887.
Brazieria ANC., Conch. Exch. ii, p. 22, August, 1887. Not Bra-
zieria Petterd, Proc. Roy. Soc. Tasrn. for 1888, p. 76 (Amnicolidce).
Shell depressed, narrowly but openly umbilicated, ribbed aboye,_
smooth and shining beneath. Whorls 4J-5, the earlier H reticu-
lated (fig. 51), the last strongly keeled. Aperture securiform, lack-
ing internal lamellce. Peristome thickened within, obtuse, the pari-
etal callus elevated into an erect tongue-like transverse process. Type
H. velata Hombr. & Jacq., pi. 5, figs. 49, 50, 51.
Soft parts unknown. The specimens before me were collected by
Mr. John Brazier at Lugunar, one of the Caroline Islands. He
found it also at Hagolu, Carolines, whence Hombron and Jacqui-
uot procured it. We cannot regard the generic relationships of
this snail as established until the soft parts are investigated ; it may
prove to belong to Zonitidce. The elevated parietal tooth is formed
on the plan of that of Polygyra cereolus, etc.
E. velata H. & J. Hi, 61.
Subgenus PHENACHAROPA Pilsbry, 1893.
Tesseraria BTTG., in v. Martens' Conchol. Mittheil., i, p. 69 (1881).
HEDLEY & SUTER, Proc. Linn. Soc. N. S. Wales (2), vii, p.
660, 1892. Not Tesseraria Hseckel, Das System der Medusen, in
Denkschr. Med.-Naturwissensch. Gesellsch. zu Jena i, p. 633 (1879
or 1880). Pupa sp., PFR., et al.
Shell pupiform, cylindrical, the altitude nearly double the diame-
ter; apical end obtusely rounded; base slightly wider, convex, nar-
rowly perforated. Surface ribbed and maculated as in Charopa s.
str. Aperture subvertical, higher than wide, toothless ; peristome
simple, thin, the columellar margin dilated. Type Pupa novosee-
landica Pfr., pi. 6, fig. 60.
Jaw arcuate, ends blunt with distant vertical strise ; upper mar-
gin slightly denticulated ; a blunt median projection on the cutting
edge (pi. 8, fig. 2).
Radula consisting of about 90 straight transverse rows of 11-5-
1-5-11 teeth. Central tooth tricuspid. Lateral teeth larger, simi-
lar to the centrals, but slightly asymmetrical and with longer ineso-
cones. Marginals broad, the 6th to 12th tricuspid, the mesocone
largest; 13th to 15th with four cusps, the ectocone being split, mes-
ocone still longest ; last marginal with one broad low cutting point
(pi. 8, fig. 1).
30 ENDODONTA.
To Mr. H. Suter, is due our knowledge of the dentition and jaw
of this peculiar shell, as well as the determination of its systematic
position. Anatomically it presents no divergence from the typical
Charopas, but the elevated pupiform shell resembles Pupa far more
than Charopa.
E. novoseelandica Pfr. ix, pi. 6, f. 60. New Zealand.
Pupa neozelanica auct.
Subgenus ^ESCHRODOMUS Pilsbry, 1892.
JEschrodomus PILS., Nautilus, vi, p. 55, footnote (Sept. 5, 1892),
-Them HUTTON, Trans. N. Z. Inst., xvi, p. 193. Not Thera
Stephens, 1831.
Shell elevated, dome-shaped, the altitude about equal to the diame-
ter. Whorls rather narrow, the apical !$ forming a light colored
spirally striated distinct embryonal shell ; the lower whorls
sculptured with oblique lamellar riblets which bear hairs where they
cross the angular periphery ; base flattened ; umbilicus small but
open. Aperture toothless, the peristome thin, simple. Type E.
stipulata Rv., pi. 6, figs. 67, 68.
Animal (of E. stipulated) like that of Charopa coma ; mantle sub-
central, slightly reflected over the peristome ; eye peduncles long
and cylindrical ; tail short, pointed, and with no mucous gland.
Jaw thin and delicate, but little arched, broadly and faintly verti-
cally striated, sometimes showing a line of reinforcement parallel to
the cutting edge (pi. 8, fig. 4, E. barbatula}.
Radula consisting of about 100 almost straight transverse rows of
teeth, the formula varying from 9-3-1-3-9, 10-4-1-4-10 or 10-6-
1-6-10 (in stipulata) to 15-1-15 (in barbatula). Central teeth tri-
cuspid, the mesocone attaining the anterior margin of the basal
plate or shorter. Laterals similar but with longer mesocones.
Inner marginals tricuspid, the outer quadricuspid by splitting of the
ectocone; the outermost having one broad low cusp (pi. 8, fig. 3,
E. barbatula Rv.
This group differs from typical Charopa in its elevated, thimble-
like contour, and the peripheral fringe of hairs borne by the riblets.
Both of the species are from New Zealand.
E. stipulata Rve. iii, 94. E. barbatula Rve. iii, 95.
alpha Pfr. beta Pfr.
ENDODONTA. 31
Subgenus PARATROCHUS Pilsbry, 1893.
Paratrochus PILS., Manual viii, p. 295.
Shell high-conic, having numerous (8i) whorls; narrowly umbil-
icated and well sculptured. Aperture nearly round, the peristome
continued in a thin callus across the parietal wall. Type H. dalber-
tisi Braz., pi. 6, figs. 55, 56.
Soft parts unknown. The single species is from Yule Island,
British New Guinea.
E. dalbertisi Brazier, viii, 295.
Subgenus CHAROPA Albers, 1860.
Shell depressed, umbilicated, discoidal or subdiscoidal. Aperture
toothless ; lip thin and simple.
This subgenus differs from Nesophila in entirely lacking spirally
entering line upon the parietal wall. It is likely that some of the
species included herein have descended from toothed forms ; although
the toothless Charopas are doubtless nearer than the toothed types
to the ancestral form of the genus Endodonta.
Sections of Charopa.
a. Shell ribbed or rib-striate, Patuloid ; whorls rounded,
Cliaropa, s. s.
act. Shell often hairy or shaggy ; whorls keeled,
Acanthoptyx, Tropidotropis, Pterotropis.
Section Charopa Albers, s. sir.
Charopa ALB., Die Hel. (edit. 2), p. 87, type H. coma Gray.
ANCEY, Bull. Soc. Mai. Fr. v, p. 364. PILSBRY, Manual viii, p. 96.
HEDLEY, Proc Linn. Soc. N. S. Wales (2) vii, p. 157. Simplic-
aria Mouss. MS., Suter, Tr. N. Z. Inst. xxiii, p. 90.
Shell depressed, subdiscoidal, the spire varying from convex to
concave ; openly umbilicated ; whorls rather cylindrical, the last
rounded or subangular (never keeled) at the periphery. Surface
sculptured with oblique or sigmoid rib-strise ; unicolored or painted
with radiating reddish flames. Aperture lunate, oblique, the lip
thin and simple, more or less sinuous ; parietal wall covered by a
varnish of callus, the riblets being removed by absorption. Type
E. coma Gray, pi. 6, figs. 57, 58, 59 (pi. 6, figs. 63, 64, 65, E. tapir-
32 ENJDODONTA.
Animal small, the mantle rather posterior, tail not produced
behind the shell. Eye peduncles large, club-shaped, approximated
at their bases; tentacles short. Foot margined by a .parapodial
groove.
Jaw delicate, thin, more or less arcuate, sculptured with fine
spaced subvertical striae (pi. 9, fig. 24, E. coma; pi. 9, fig. 21, E.
sylvia= buctinella.*)
Radula having the teeth in somewhat sinuous transverse rows.
Central tooth tricuspid, the mesocone reaching about to the anterior
border of the basal-plate, side cusps small. Laterals similar but
somewhat asymmetrical, the entocone becoming larger outwardly
until it becomes joined at the base with the mesocone. The marginals
are very low and wide, by shortening of the basal-plates ; tricuspid,
the ento- and mesocone often joined at base ; ectocone smaller, simple
or split into two. Cusps variously degenerate on the outermost mar-
ginals (pi. 9, fig. 23, E. coma ; pi. 9, fig. 20, E. sylvia=buccinella.)
In some species, such as E. dispersa Gassies, the entocone remains
minute upon all of the lateral teeth ; and in some the ectocone splits
on the marginals ; but otherwise the dentition of the species does
not differ from that of E. coma.
The shell is like Goniodiscas in being umbilicated, depressed, rib-
striate ; whorls tubular, aperture round-lunar or crescentic. It
differs from Goniodiscus in the tendency of the upper lip to recede
toward its insertion, forming a Pleurotomoid sinus or notch between
outer lip and body-whorl ; and in the more or less depressed (some-
times concave) inner whorls.
The species are very numerous, and they occupy a vast territory ;
but New Zealand, New Caledonia, and Australia with Tasmania are
their especial home.
New Zealand species.
E. anguiculus Rv. iii, 23. E. huttoni Sut. viii, 104.
theta Pfr. E. infecta Rv. iii, 23.
E. bianca Hutt. viii, 97. zeta Pfr.
v. montana Sut. v. irregularis Sut. viii, 98.
E. biconcava Pfr. i, 130 ; viii, 104. v. alpestris Sut. viii, 99.
E. brouni Sut. viii, 102. E. lucetta Hutt. iii, 22.
E. buccinella Rv. iii, 23. stokesi Sm. iii, 262.
gamma Pfr. E. moussoni Sut. viii, 105.
sylvia Hutt. viii, 98. E. mutabilis Sut. viii, 101.
ENDODONTA.
33
E. caputspinulse Rv. iii, 102.
epsilon Pfr.
E. colensoi Sut. viii, 99.
E. coma Gray, iii, 22.
v. beta Pfr.
'v. globosa Sut. viii, 96.
E. corniculum Rv. iii, 24.
eta Pfr.
v. maculata Sut. viii, 96.
E. egesta Gray, iii, 23.
E. eremita Sut. viii, 103.
E. pseudocoma Sut.
E. raricostata Sut. viii, 100.
E. segregata Sut.
E. serpentinula Sut. viii, 103.
E. sterkiana Sut. viii, 101.
v. major Sut.
v. reeftonensis Sut.
E. subantialba Sut. viii, 104.
E. tapirina Hutt. iii, 23 ; viii, 97.
E. tau Pfr.
E. variecostata Sut. viii, 100.
New Caledonian species.
E. alveolus Gass.
E. bazini Cr. i, 121.
E. calliope Cr. iii, 36.
E. confinis Gass. iii, 35.
E. costulifera Pfr. i, 120.
v. major Cr.
E. decreta Gass. iii, 26.
E. dispersa Gass. iii, 45.
E. inculta Gass. iii, 26.
E. kanakina Gass. i, 122.
E. koutoumensis Gass.
E. lamberti Cr. iii, 26.
E. melaleucarum Gass. iii, 26.
E. melitse Gass. iii, 45.
E. rnorosula Gass.
E. noumeensis Cr.
E. ostiolum Cr.
E. pinicola Pfr. i, 121.
E. rhizophorarum Gass. iii, 36.
E. rusticula Gass. iii, 26.
E. saburra Gass.
E. subcoacta Gass. iii, 26.
E. subtersa Gass. iii, 35.
E. taslei Cr. iii, 36.
E. vetula Gass. iii, 36.
Species of Lord Howe and Norfolk Is.
E. depsta Cox, iii, 46.
E. exagitans Cox, iii, 46.
E. unwini Braz. viii, 106.
E. wilkinsoni Braz. viii, 105.
E. whiteleggei Braz. viii, 106.
v. balli Braz. viii, 107.
v. ledgbirdi Braz. viii, 107.
Species of Australia and Tasmania.
[The following synonymic list was furnished by my valued corre-
spondent and friend, CHARLES HEDLEY, of Sydney, N. S. W.]
E. agnewi Cox, iii, 263. E. microscopica Cox.
petterdi Brazier. microcosmos Cox.
var. peroni Brazier. E. millestriata Smith, i, 130.
UNIVEBBITT
OF
34
ENDODONTA.
E. albanensis Cox, ii, 209 ; viii,
[pi. 37, f. 43-46.
eastbournensis Beddome &
[Petterd.
petterdiana Taylor,
var. stanleyensis Petterd.
var. albida Taylor.
E. antialba Beddome viii, 107.
E. barrenensis Petterd.
E. belli Cox, iii, 25.
E. biretracta Mousson, ii, 208.
E. bischoffensis Beddome, viii,
[109.
E. brazieri Cox, iii, 24.
E. cochlidium Cox, iii, 25.
E. corticicola Cox.
E. cupera Cox, iii, 24.
napera Brazier.
E. curacose Brazier.
ramsgatensis Cox, iii, 265.
E. cygnea Benson, ii, 213.
E. diemenensis Cox, iii, 24.
thomsoni Cox.
daveyensis Cox, iii, 265.
atkinsoni Cox, iii, 266.
camillce Cox.
wellingtonensis Cox.
midsoni Brazier.
E. funerea Cox, ii, 209.
E. furneauxensis Petterd.
E. gadensis Beddome viii, 109.
E. halli Cox.
E. hookeriana Johnston.
E. iuloidea Forbes ii, 209.
omicron Pfeiffer.
ammonitoides Reeve.
legrandi Cox, ii, 209.
ricei Brazier.
onslowi Brazier.
E. kershawi Petterd.
E. mimosa Petterd.
E. mucoides Tenison-Woods. iii,44.
E. murphyi Cox, iii, 46.
E. murrayana Pfeiffer.
E. nautiloides Cox.
inusta Cox, ii, 209.
E. neglecta Brazier.
luckmani Brazier,
var. siliens Cox.
var. jungermamse Petterd.
var. trucanini Petterd.
E. officieri Cox, iii, 266.
E. otwayensis Petterd.
var. alpina Johnston.
E. paradoxa Cox.
morti Cox, iii, 34.
hobarti Cox, iii, 34.
arenicola Tate, iii, 52.
E. pexa Cox, iii, 25.
E. retepora Cox, iii, 34.
E. reteporoides Tate, viii, 110.
E. roblini Petterd.
E. rotella Brazier.
E. saturni Cox, iii, 24.
E. sericatula Pfeiffer, ii, 208.
melbournensis Cox, iii, 35.
E. spaldingi Brazier,
var. carinata Brazier.
E. parvissima Cox.
E. spiceri Petterd.
E. spectra Cox, iii, 266.
architectonica Brazier.
gunni Brazier.
assimilis Brazier.
E. similis Cox.
stellata Brazier, iii, 34.
derelicta Cox.
E. stroudensis Cox, iii, 25.
E. subdepressa Brazier.
dandenongensis Petterd.
ENDODONTA. 35
E. limula Cox. E. sublesta Benson.
E. lottah Petterd. E. subrugosa Brazier.
E. raacdonaldi Cox. E. tamarensis Petterd.
kingstonensis Cox, iii, 266. E. tasmanise Cox, iii, 34.
gouldi Cox. E. vigens Cox, iii, 263.
juliformis Cox, iii, 263. ammonitoides Brazier.
E. marchianse Cox. bassi Brazier.
fuscoradiata Cox, iii, 265. E. vinitincta Cox, i, 115.
E. mathinse Petterd.
E. lizardensis Pfr. iii, 86.
New Guinea, Am and Tenimber Is. species.
E. brunnescens Mlldff. viii, 82. E. texta Hedley. viii, 294.
E. demani Tap.-Can. iii, 26.
Polynesian species.
E. adposita Mouss. iii, 41. vicaria Mouss.
E. canalis Grt. iii, 39. v. vicinalis Mouss. iii, 39.
E. complementaria Mouss. iii, 40. E. monstrosa Anc. viii, 82.
E. decorticata Grt. iii, 40. irregularis Mouss. not Semp.
v. otarese Grt. E. oualanensis Pse. iii, 41.
E. filiola Fer. iii, 38. E. planospira Grt. iii, 41.
E. glissoni Anc. viii, 82. E. princei Liard. iii, 27.
E. harveyensis Grt. iii, 40. E. proxima Grt. iii, 39.
E. helva Cox, iii, 262. E. radicalis Mouss.
E. ignava Pfr. iii, 36. E. rotula Hombr. iii, 67.
E. inermis Mouss. iii, 41. E. rudis Grt. iii, 39.
E. lamellicostata Grt. iii, 39. ? sublaminata Mss, Schra.
E. modicella Fer. iii, 38. E. tenuicostata Grt. iii, 39.
v. atiensis Pse. E. youngi Grt. iii, 40.
Section Acanthoptyx Ancey, 1888.
Acanthoptyx ANCEY, Bull. Soc. Mai. France, v, p. 370.
Shell discoidal, thin, openly umbilicated ; whorls few (3-4) and
rapidly increasing, sculptured with fine close lamellar striae and
unevenly spaced elevated ribs, rising into lamellae as they cross the
36 PHASIS.
subangular periphery. Aperture large, oblique, toothless ; peristome
fragile. Type H. acanthinula Crosse, pi. 6, figs. 71, 72, 73.
Jaw and soft parts not examined.
Dentition : centrals as wide as long, tricuspid, the mesocone attain-
ing the anterior border of the basal-plate, side cusps small. Laterals
similar. Marginal teeth low, wide, with the entocone and mesocone
long, united at base, the ectocone split into three minute cusps (pi.
9, fig. 25, E. acanthinula. ,)
E. acanthinula Crosse. iii, 124. New Caledonia.
Section Tropidotropis Ancey.
Tropidotropis ANC., Bull. Soc. Mai. Fr. v, p. 370.
Shell broadly umbilicated, discoidal, the spire nearly flat; whorls
flat above, the last acutely carinated ; epidermis^laciuiate-lamellose.
Aperture securiform, toothless, the peristome simple, acute. Type
H. trichocoma Crosse, pi. 6, figs. 61, 62.
E. trichocoma Cr. iii, 45. New Caledonia.
Section Pterodiscus Pilsbry, 1893.
Tropidoptera ANC., Bull. Soc. Mai. Fr. vi, p. 191. Not Tropido-
pterus Blanch. 1845 {Coleoptera.}
Shell umbilicated, depressed, thin or fragile, horny brown.
Whorls finely, densely striated, the last acutely keeled at the periph-
ery, carinated around the umbilicus. Aperture oblique, toothless ;
lip thin and simple. Type H. alata Pfr., pi. 4, fig. 44.
Shells of this section have the appearance of the New Caledonian
groups Acanthoptyx and Tropidotropis. The species are from the
Sandwich Is.
E. alata Pfr. E. depressiformis Pse.
E. prostrata Pse. E. digonophora Anc.
Genus PHASIS Albers, 1850.
Phasis ALB., Die Hel., p. 92. Type and only species H. menke-
ana Pfr. PILSBRY, Manual viii, p. 135.
Shell resembling Xerophila; depressed, umbilicated, solid, white
and opaque, generally with brown bands or dots, the apex dark; last
whorl not descending ; aperture rounded-lunate, but little oblique ;
PHASIS. 37
lip thin, simple, its eolumellar margin dilated. Type H. menkeana
Pfr., pi. 10, figs. 1, 2, 3.
Distribution, South Africa.
Under this genus as subgenera may be ranged two groups : Tra-
chyeystis and Sculptaria, both belonging to the S. African fauna. The
anatomy of typical Phasis is unknown. That of Trachycystis is
described below. The diagnosis given above applies to the restricted
subgenus Phasis only, to which the following species belong :
P. capensis Pfr. iii, 103. P. namaquana Mts. viii, 297.
irrorata Zieg. P. paludicola Bens, iii, 104.
littoricola Bens. P. sturmiana Pfr. vi, 317.
P. menkeana Pfr. iii, 108. P. uitenhagensis Kr. iii, 104.
Subgenus TRACHYCYSTIS Pilsbry, 1892.
Trachycystis PILS., Man. of Conch, viii, p. 136. Pella Alb. (in
part), Die Hel. (2), p. 84, 1860. Not Pella Steph. 1832.
Shell small, thin, generally somewhat translucent, horny or earthy
brown in color, usually sculptured with oblique riblets or rib-striae,
the apical whorl spirally striated (fig. 7) ; aperture lunate ; lip
simple, thin, dilated at the eolumellar insertion. Type P. biseulpta
Bens., pi. 10, figs. 5, 6, 7 ; see also P. browningi Bens. pi. 10,
figs. 8, 9.
Animal (of P. rariplicata) having a rather long slender foot, the
sole apparently undivided ; foot-margins wide, not crenulated nor
more coarsely granulated than the rest of the surface, defined by a
pair of shallow grooves; tail lacking a mucous pore.
Jaw thin, having numerous flat plaits.
Radula having the transverse rows of teeth crowded, so that the
cusps of one row project over the bases of the next. Central teeth
tricuspid, the mesocone longer than the basal plate, slender, side cusps
small. Lateral teeth altogether similar, but slightly asymmetrical,
the entocones increasing in length outwardly. Transition from
lateral to marginal teeth very gradual, the latter tricuspid, the
ento- and mesocones subequal, long, oblique and united at their
bases, the ectocone smaller, simple (in P. biseulpta'} or bifid (P. rari-
plicata'). PI. 15, figs. 3, 4, P. biseulpta.
All of the teeth are tricuspid ; the|central and inner lateral teeth
are so similar that it is difficult to distinguish which is the rhachi-
dian row, and the mesocones are long and slender. The inner mar-
ginal teeth are remarkable for their long ento- and mesocones.
38 PHASIS.
Binney has figured the teeth of P. rariplicata, but judging by the
radula before me he makes the median teeth too short for their
length. He correctly figures the ectocone of the outer marginals
as bifid. The radula of P. bisculpta has not before been examined.
These shells are shaped like Phasis from which they differ in the
thin texture and sculpture. Some species resemble the New Zea-
land group Allodiscus and others are like Thysanophora or Patula.
All of them belong to the South African fauna, with the exception
of a few species from Mauritius and adjacent islands, which present
the same shell characters, but have hitherto been grouped in Patula
or Charopa. The affinities of the genus are with the Charopoid
Endodontas as far as present knowledge enables us to judge; and
they are separated from that type mainly by their distribution and
certain features of the teeth described above. The differences in
the radula are, however, of but little importance.
P. actinotricha Melv. & Pons. P. microscopica Kr. iii, 106.
[viii, 143. P. minythodes Melv. & Pons.
P. amea Kr. iii, 105. [viii, 144.
P. aprica Kr. iii, 107. P. newtoni Nev. iii, 27.
P. arachne Morel. P. perplicata Bens, iii, 106.
P. aulacophora Anc. viii, 138. P. petrobia Bens, iii, 107.
P. bathycoele Melv. & Pons. viii, P. planti Pfr. viii, 142.
139. platti Pfr. olim.
P. bisculpta Bens, iii, 105. v. africse Bra. viii, 142.
P. browningii Bens, viii, 136. P. prionacis Bens, viii, 137.
P. burnupi Melv. & Pons. viii, P. rariplicata Bens, iii, 107.
[140. P. rhysodes Melv. & Pons. viii,
P. caldwelli (Barcl.) Bens, iii, 27. [141.
paulus Mor. P. rivularis Kr. iii, 107.
P. conisalea Melv. & Pons. viii, P. rodriguezensis Cr.
[145. P. sabuletorum Bens, iii, 107.
P. crawfordi Melv. & Pons. viii, P. somersetensis Melv. & Pons.
[146. [viii, 295.
P. epetrima Melv. & Pons. viii, P. strobilodes Melv. & Pons.
[146. [viii, 147.
P. erateina Melv. & Pons. viii, P. tabulae Chap, viii, 139.
[137. P. trichosteiroma Melv. & Pons.
P. farquhari Melv. & Pons. viii, [viii, 143.
[147. P. tuguriolum Melv. & Pons.
P. glanvilliana Anc. viii, 147. [viii, 145.
SCULPTARIA, AMPHIDOXA. 39
P. hottentota Melv. & Pons. P. turmalis Morel, viii, 144.
[viii, 141. P. viridescens Melv. & Pons.
P. inops Morel, viii, 144. [viii, 78.
P. liricostata Melv. & Pons. P. vorticialis Bens, iii, 107.
[viii, 140. P. vorticella H. Ad. iii, 35.
P. loveni Kr. iii, 106. P. zanguebarica Craven.
P. lygsea Melv. & Pons. viii,
[138.
Subgenus ? SCULPTARIA Pfr., 1856.
Sculptaria PFR., Malak. Blatter ii, p. 135, type If. sciilpturata
Gray.
Shell small, discoidal, carinated, widely umbilicated ; last whorl
becoming free at the aperture ; aperture very oblique, rounded,
with continuous slightly expanded peristome, and having several
teeth on the outer lip and an entering parietal lamina. Type H.
SGidpturata Gray, pi. 10, fig. 4.
Anatomy unknown. A group of problematic relationships, rep-
resented by a few species in southwestern Africa (Damaraland).
S. damarensis H. Ad. iii, 138. S. chapmanni Anc. viii, 152.
S. sculpturata Gray, iii, 138. S. retisculpta Mts. viii, 152.
v. collaris Pfr. iii, 138.
Genus AMPHIDOXA Albers, 1850.
Amphidoxa ALB., Die Hel. p. 110 (for H. marmorella and heli-
cophantoides') ; Edit. Martens, p. 82.
Shell thin, depressed-globose or discoidal, perforated or umbili-
cated; aperture lunar-rounded or ovate; peristome simple, thin.
Type H. marmorella Pfr., pi. 7, figs. 10, 11, 12.
Distribution: southwestern shore of South America and adjacent
islands, Juan Fernandez, Chiloe, etc., Cape Horn region and Ker-
guelen Is.
These shells resemble some forms of the genera Flammulina and
Endodonta\ the typical Amphidoxas recalling Flammulina or
Calymna, the Stephanodas being like Allodiscus, Suteria or Charopa.
The anatomy of the South American forms is unknown, but that of
A. hookeri of Kerguelen Island shows an affinity to Charopa in the
possession of parapodial grooves. Two sections compose this group.
40 AMPHIDOXA.
Section AMPHIDOXA Alb.
Shell small, perforate, depressed-globose, thin and pellucid, costu-
late-striate, Whorls 8-82, rapidly enlarging. Aperture ample.
Anatomy unknown. Distribution, Juan Fernandez.
A. marmorella Pfr. iii, 46. A. helicophantoides Pfr. iii, 46.
Section STEPHANODA Albers, 1860.
Stephanoda ALB., Die Hel. (2) p. 88. Type H. dissimilis Orb.
Stepsanoda PFR., Nomencl., p. 93.
Shell umbilicated, thin, costulate, sometimes spirally striated ; in
shape like Discus or Charopa. Whorls 5-7, the last cylindrical, not
descending. Aperture rounded lunar ; lip thin, simple. Type H.
dissimilis Orb., pi. 7, figs. 19, 20, 21. See also pi. 7, figs. 16, 17, 18,
A. hookeri Reeve.)
Anatomy of the typical forms unknown ; of A. hookeri as follows,
the living animal according to Eaton's observations (Philos. Trans.,
1879, p. 183), the internal anatomy according to Schako and Pfeffer
(Monatsber. K.-P. Akad. Wissensch. Berlin, 1877, p. 269.)
Animal (in spirit) with a narrow foot, rather narrower posteriorly
than in front. The sole of a pale livid olive, sides dark slate color.
Mantle above the head pale livid, dotted with dark slate spots.
During life the animal (viewed through a lens), is black, reticulated
with gray; tentacles either black above and dark gray beneath
longitudinally, or dark gray throughout. Foot bordered above by
a ribbon-like stripe which is composed of long oblong tessellations
whose interstices are gray, which is separated by a thin pale irreg-
ular line from the more finely reticulated upper portion of the sides
and back. The interspaces of the reticulation of these last are
slightly raised and black, and cause the surface to be somewhat
granulated. Some of the lines of growth of the shell are occasion-
ally straw color (Eaton). Sole tripartite, divided into areas by two
longitudinal and many transverse grooves, the outer areas darkly
pigmented. No appendages upon the mantle margin.
Genitalia simple, without accessory organs ; vas deferens inserted
at the apex of penis, passing gradually into it ; spermatheca terminat-
ing in a short straight or bent appendage, and situated upon a rather
long duct (pi. 1, fig. 16, A. hookeri.}
Jaw measuring *7 x '68 mill., rather narrow, low-arcuate, sculpt-
ured with fine, somewhat wavy transverse striae and numerous nar-
AMPHIDOXA. 41
row vertical grooves, which hardly crenulate the cutting edge. In
young examples it seems as if composed of narrow plates held
together by the underlying membrane (Schako). PI. 1, fig. 15, A.
hooker i.
Kadula measuring 2'41 x '68 mill., consisting of 205 closely placed
transverse rows, each with 35, 51, 57 or 65 teeth. Formula 25-11-
1-11-25. Rhachidian tooth with a broa'd, blunt, rounded meso-
cone, no side cusps. Laterals similar, the cusp often extending
beyond the thin basal-plate. Marginals tricuspid, the side cusps small
but distinct, obsolete on the outer marginals (pi. 1, fig. 14, A.
hookeri, showing teeth R, 1, 12, 17, 22, 25.)
The principal peculiarity of the radula is that the central and
lateral teeth possess mesocones only, in this respect differing from
the genera Endodonta and Phasis ; but as the dentition of but one
species is known, too much stress should not be laid upon this feature.
The close alliance of the toothless Endodontas (Charopa), the S.
African group Trachycyatis, the northern genus Pyramidula, and the
S. American Amphidoxa-Stephanoda series, is evident.
A. arctispira Pfr. iii, 47. A. lirata Couth, iii, 42.
A. binneyana Pfr. iii, 48. A. magellanica Sm. iii, 42.
A. bryophila Ph. iii, 42. A. musicola Ph. iii, 43.
A. ceroides Pfr. iii, 47. A. ordinaria Sm.
A. cliilensis Miihlf. iii, 42. A. pazii Ph. iii, 43.
A. coiquecana Ph. iii, 43. minviellei Ph.
A. coppingeri Sm. iii, 42. A. pleurophora Moric. iii, 53.
A. corticaria Ph. iii, 43. A. pusio King, iii, 47.
A. costellata Orb. iii, 41. A. quadrata Fer. iii, 47.
A. dissimilis Orb. iii, 48. Idngi Pfr.
histrio Miihlf. A. rigophila Mab. & Roch. viii,
plagiata Beck. . [81.
A. epidermia Ant. iii, 42. A. selkirki Sm. iii, 47.
A. exigua Ph. iii, 43. A. spirillus Gld.
A. germaini Ph. iii, 43. A. stelzneriana Ph. iii. 43.
A. gratioleti Hupe, iii, 48. A. strobeliana Ph. iii, 43.
A. holmbergi Dor. iii, 43. A. tenuistriata Ph. iii, 48.
A. hookeri Rve. iii, 48. A. tessellata Miihlf. iii, 47.
A. hypophloea Ph. -iii, 43. contortula Fer.
A. jungermanniarum Ph. iii, 43. A. zebrinaPh. iii, 48.
A. leptotera Mab. & Roch. viii, 81 .
42 PYRAMIDULA.
Genus PYRAMIDULA Fitzinger, 1833.
Pyramidula FITZ., System atisches Verzeichniss der im Erzher-
zogthume Oesterreich vorkommenden Weichthiere, als Prodrom
einer Fauna derselben, p. 95 (for H. rupestris Drap.)
-f Gonyodiscas and Discus FITZ., 1833 ; Patula HELD., 1837 >
Delomphalus Ag., 1837; Eyryomphala Beck, 1837 ; etc., etc.
=Patula of most modern authors.
Shell openly umbilicated, varying in contour from flattened and
disk-like to conoidal. Generally opaque, often rib-striate. Uni-
colored, spirally banded or flammulate. Whorls subcylindrical or
keeled, the apex generally smooth. Aperture rounded-lunate ; lip
simple and thin. Type P. rupestris Drap.
Animal having the sole undivided; lateral margin of the foot with
a distinct border bounded by a groove, the grooves meeting above the
tail. No caudal mucous pore. Eye-peduncles long and slender
(pi. 14, fig. 40, 46, P. alternata.')
Genital system lacking all accessory organs ; vas deferens and
retractor muscle inserted near or at the apex of the penis ; duct of
the spermatheca very long; hermaphrodite duct very long, but
shortened by its extreme convolution (pi. 11.)
Jaw arcuate, its component laminae generally compactly soldered,
and indicated only by fine stride which diverge slightly from the
middle.
Radula (1) having only the mesocones developed upon central
and inner lateral teeth, or (2) having the centrals tricuspid, laterals
bicuspid lacking the entocones, marginal teeth similar but with short
basal-plates ; this being the usual form. In some species the mar-
ginal teeth are multicuspid by the splitting of their ectocones.
The dentition, as usual, shows considerable variation, even in
species otherwise closely related. As a general rule, the lateral teeth
completely lack entocones, differing in this respect from Trachycystis
and the Endodonta- Charopa series ; but in the section flelicodiscus,
entocones are well developed. The dentition is quite unlike Tra-
chycystis in the forms of the marginal teeth.
The genus Pyramidula consists of dull-colored ground-living snails,
species of which occur over the whole northern temperate land area.
Its nearest relatives are Charopa, Trachycystis and Stephanoda, genera
occupying the southern temperate regions of Australasia, Africa and
South America respectively. All may be regarded as the remnants
of an early fauna, now replaced in the tropics, and to a large extent
PYRAMIDULA. 43
in temperate regions also, by higher groups of Helices. The latter
differ widely from these Pafculoid genera in lacking parapodial
grooves, in the solid, ribbed jaw, complex genital system, and other
features to be described later.
In treating of the subgenus Patula it will be shown that that name
is not available as a designation for the present genus as a whole.
Pyramidula is the earliest name, and should be accepted. It may be
objected that no diagnosis of Pyramidula was published by Fitzinger,
but the same may be said of Beck's genera. Let those who repudi-
ate Beck's names cast the first stone at Fitzinger !
Pyramidula is divisible into eight subordinate groups, which may
be tabulated thus :
a. Shell lacking internal teeth or folds,
b. Spire conical ; size very small, shell thin,. Pyramidula s. s.
bb. Spire depressed,
c. Shell rather large and solid, Patuia.
cc. Shell small or minute,
d. Surface spirally lamellate, Lyrodiscus.
dd. Body-whorl with 20-25 spaced oblique laminae,
Planogyra.
ddd. Surface striate or rib-striate, Gonyodiscus, Patulastra.
aa. Body-whorl having one or several pairs of internal teeth,
b. Internal teeth tubercular ; surface spirally sculptured,
Helicodiscus.
bb. Internal teeth lamellar ; surface obliquely sculptured,
Atlantica.
Besides these, another group, Pupisoma, has been referred provi-
sionally to this genus.
Subgenus PYRAMIDULA Fitz.
Shell small, moderately or widely umbilicated, lacking internal
folds or teeth.
The following sections may be grouped under this subgeneric
head : Pyramidula s. sir., Patulastra, Planogyra, Gonyodiscus and
Lyrodiscus.
Section Pyramidula Fitz., s. sir.
Pyramidula FITZ., Syst. Verz., p. 95.
Shell minute, openly umbilicated, with pyramidal spire and obtuse
44 PYRAMIDULA.
smooth apex. Whorls tubular, obliquely striated. Aperture round
or nearly so ; lip simple. Type H. rupestris Dr., pi. 10, figs. 15, 16.
Jaw arcuate, finely striated vertically.
Radula having the central teeth unicuspid, the side cusps being
represented by a slight sinuation. Laterals bicuspid. Marginals
with low wide basal-plate, the inner bearing two cusps, the outer
becoming multicuspid by splitting of the cusps, (pi. 11, fig. 25, P.
rupestris Dr.)
Distribution, Europe and Central Asia.
This section differs from Gonyodiscus and Patulastra in having the
spire conically elevated, and from the former in lacking rib-strise.
P. rupestris Dr. iii, 51. f. dalmatina Cl.
umbilicata Mont. f. pinii Ad. iii, 51.
aliena Zieg. P. chorismenostoma Bl. & West.
spirula Villa. P. hierosolymitana Bgt. iii, 52.
myrmecidis Scac. P. humilis Hutt. iii, 22.
f. rupicola Stab. P. orphan a Hde.
f. saxatilis Hm. P. euomphalus Blf. iii, 32.
f. subglobosa Bgt. P. abbadiana Bgt. iii, 52.
f. conoidea Bgt. P. brucei Jick. iii, 52.
f. meridionalis Iss. P. amblygona Reinh. iii, 52.
f. jsenensis Cl. iii, 51. P. lepta West, viii, 81.
Section Patulastra Pfeiffer, 1878.
Patulastra PFR., Nomencl. Hel. Viv., p. 87.
Shell having the form of Patula, but minute, with fewer whorls,
the surface unicolored, with or without riblets.
This section may be retained to include the minute forms similar
in general characters of the shell to Punctum, but with the anatom-
ical features of the genus Pyramidula. The limits of the group are
uncertain, as part of the species might be placed in the sections
Gonyodiscus or Pyramidula, and others are likely to prove Punc-
tums. Of course the melange included here by Pfeifier and by
Tryon must be assorted into many diverse groups.
P. abyssinica Jick. iii, 32. P. debeauxiana Bgt. iii, 28.
rivularis Mts. P. carotae Bgt. Serv. iii, 31.
P. aranea Parr, iii, 31. P. elachia Bgt. iii, 28.
P. aucapitainiana Bgt. iii, 29. P. galla3ciana Silv.
P. balatonica Serv. iii, 31. J P. henriquesi Silv.
P. bussacona Silv. P. lederi Bttg, iii, 31.
PYRAMIDULA. 45
P. luseana Paiv. iii, 31. P. pusilla Lwe. iii, 31.
P. massoti Bgt. iii, 29. hypocrita Dohrn.
P. micropleuros Pag. iii, 28. servilis Sh.
P. microstigmsea Silv. P. servaini Bgt. iii, 31.
P. uemesiana Bgt. iii, 31. P. simoniana Bgt iii, 3J.
P. pornse Serv. iii, 31. P. sororcula Ben. iii, 29.
P. poupillieri Bgt. iii, 29. P. tenuicostata Sh. iii, 28.
Section Planogyra Morse, 1864.
Planogyra MORSE, Obs. Terrest. Pulm. Maine, p. 24, type P.
asteriscus Mse.
Shell minute, discoidal, openly umbilicated, the spire flat. Whorls
bearing thin, sharp, spaced lamince, parallel to growth -striae. Aper-
ture rounded-lunar, lip simple. Type P. asteriscus Morse, pi. 10,
figs. 10, 11.
Jaw slightly arcuate, bluntly rounded at the ends, irregularly
vertically wrinkled, the concave margin having a slight median
projection.
Radula consists of 77 transverse rows containing about 13.1.13
teeth. Centrals tricuspid. Laterals lacking the entocone. Mar-
ginal teeth multicuspid, the mesocone largest, bifid (pi. 1J, fig. 21,
P. asteriscus Morse).
The radula differs from that of Pyramidula s. sir. only in the
development of side cusps on the central tooth, and the shorter
mesocone of the same. But one species is known ; it is widely dis-
tributed in Canada and northern New England, living in very wet
places.
Morse represents the eye-peduncles of this species as short, thick,
and club-shaped (pi. 10, fig. 10) ; his observation should be checked
by an examination of the living animal, as that form of eye stalk
is widely different from the other Pyramidula species.
Section Gonyodiscus Fitzinger, 1833.
Gonyodiscus FITZ. Syst. Verz. p. 98, proposed for G. perspectivus
Fitz. IT. solaria Mke. Discus FITZ., Syst. Verz., p. 99 ; proposed
for H. rotundata, ruderata, pygmcea, cristallina (not Discus Less.
1837, nor of Hald. 1840, nor of Alb. 1850, nor of Campb. 1879).
Patula HELD, in part. Delomphalus AGASSIZ, in CHARP., Catal. des
Moll. Terrest. et Fluv. de la Suisse, p. 12, in Nouv. Mem. de laSoc.
Helvetique des Sci. Nat. i, Neuchatel, 1837 ; proposed for H.
rotundata, ruderata pygmcea. Eyryomphala (in part) BECK, Index,
46 PYRAMJDULA.
p. 8. Patularia CLESSIN, Die Molluskenfauna Oesterreich-Ungarns
und der Schweiz, p. 104 (proposed for P. rotundata, hauffeni, ruderata,
solar ia,pygmcea). Spelceodiscus BRUSINA. Mittheil, naturwissensch.
Ver. Steierruark, 1885, p. 37, type, H. hauffeni. Allerya BOUR-
GUIGNAT, Atti Ac. Palermo, 1876 (^embryonic shells of H. rotund-
ata, etc.).
Shell rather small, depressed, with low but convex spire and open
umbilicus. Apical H whorls smooth, the rest obliquely rib-striate,
rather tubular, rounded or keeled at the periphery, unicolored or
flamed with reddish. Aperture wide-lunate, the lip simple. Type
P. solaria Mke., pi. 10, fig. 14. See also pi. 10, figs. 12, 13, P.
rotundata Mull.
Animal (of P. perspectives Say) long and narrow, the foot white,
head and back dusky blue. Sole equal in length to the diam-
eter of the shell, undivided (having a central longitudinal sulcus
when entering the shell or in alcohol); margins of foot having a
wide border, bounded by a distinct groove, the grooves meeting
above the tail. Upper surface coarsely granulated. Eye pedun-
cles long and slender, from one-third to one-half as long as the foot
(pi. 14, fig. 45).
Genital system lacking all accessory organs. The penis is short,
having the retractor and the vas deferens inserted at its apex.
Spermatheca small, situated upon a very long simple duct, which
enters the vagina very low. At the base of the albumen gland
there is a rather large talon. The albumen gland is small and
adherent to the lower part of the hermaphrodite duct ; the latter
being large and very much convoluted (pi. 11, fig. 22, P. persjiec-
tiva).
The genital system of P. rotundata as figured by Lehmann is sim-
ilar. Leidy's figure of that of perspectiva is incorrect in showing
an appendicula.
Jaw arcuate, with a slight median projection, finely striated,
the striae subvertical, diverging below toward the outer basal
angles of the jaw (pi. 11, fig. 19, P. perspectiva). The jaw of
rotundata, according to Lehmann and Moquin-Tandon, has fewer,
more spaced striae than I have found in P. perspectiva. That of P.
balmei (pi. 15, fig. 2) is very distinctly and closely striated, and dif-
fers from the jaw of perspectiva is being incompletely soldered, the
edges of the component vertical plates being slightly free, as in some
charopoid snails.
PYRAMIDULA. 47
Radula bearing crowded teeth (i n P. perspectives, arranged accord-
ing to the formula 12.8.1.8.12). Centrals having a long mesocone
and small side cusps. Laterals having no entocone, the mesocone
oblique, ectocone small. Marginals similar, but with short, broad
basal plates (pi. 11, fig. 26, P.perspectiva).
In P. balmei the marginal teeth are like those of Planogyra
aster iscus.
This section is distinguished from Pyramidula s. str. by its low
spire, discoidal form, and the rib-striation, which is often obsolete
below the periphery, but generally persists on the upper surface
and within the umbilicus. The typical species of Gonyodiscus are
carinated at the periphery, and those with rounded whorls have
been separated under the name Discus, but such a separation does
violence to the facts in the case, for all intermediate stages of contour
between the most acutely carinated aud the rounded types occur. As
well might one separate Papuina brumeriensis from diomedes as a
distinct section, or Pyramidula (Patula) cumber landiana fromalter-
nata. Such classification may be left for those who point the small
end of the telescope at nature.
Eurasian species.
P. abietana Bgt. iii, 21. P. omalisma Bgt.
P. aperta Mlldff. viii, 80. P. pallens Gred. viii, 82.
P. assarinensis Calc. iii, 51. P. pauper Gld. iii, 20.
P. balmei P. & M. iii, 30. P. putrescens Lwe. iii, 31.
flavescens Parr. P. retexta Sh. iii, 44.
flavida Zieg. P. rotundata Mull, iii, 19.
striolata Ph. brocchiana Calc., Ben.
P. bianconii Dh. iii, 32 cupaniana Calc., Ben.
P. carpetana Hid. radiata DaC.
P. concinna Lwe. iii, 21. v. pyramidalis Jeffr.
P. costulata Mlldff. iii, 266. v. globosa Friedl.
P. engonata Shuttl. iii, 43. v. turtoni Flem. iii, 19.
v. pallidior Mouss. P. ruderata Stud, iii, 20.
P. erdeli Roth, iii, 30. umbilicus Mark.
P. flocculus Mor. perspectives Fer.
P. frivaldskyana Rm. iii, 21. v. angulosa Mouss. iii, 26.
convexa Fer. v. opulens West, iii, 20.
P. gortschana Mouss. iii, 20. P. solaria Mke. iii, 43.
P. hauffeni Schm. iii, 30. perspectiva Miihl.
P. luseana Paiv. iii, 31. megerlei Jan.
48 PYRAMIDULA.
P. sudensis Pfr. iii, 30. P. zapateri Hid.
P. textilis Sh. iii 31.
American species.
P. perspectiva Say. iii, 20. P. striatella Anth. iii, 20.
patula Dh. v. catskillensis Pils.
P. bryanti Harp, iii, 43. v. cronkhitei Newc. iii, 21.
P. horni Gabb. iii, 21.
Section Lyrodiscus Pilsbry, 1893.
Lyra MOUSSON, Rev. Fauue Malac. Canar., p. 26. Not Lyra
Cumberl., 1816.
Shell depressed, with large open umbilicus and low-convex spire,
in form being like Patula; surface sculptured with slight growth-lines
and numerous elevated cuticular spiral threads. Type H. circumsessa
Shuttlew. Anatomy unknown. Distribution, Canary Islands.
P. circumsessa Sh. P. torrefacta Lwe.
Subgenus PATULA Held, 1837.
Patula HELD, Isis, 1837, p. 918 (proposed for alternata, rotundata,
solaria, perspectiva, ruderata, pygmcea, rupestris). Eyryomphala
BECK, Index Moll. p. 8 (proposed for solitaria, alternata, perspectiva,
ruderata, solaria, rudis, rotundata, rupestris, pygmcea, pusilla, lineata
and some undescribed Amphidoxa or Stephanoda species). Euryom-
phala HERM. et al.Anguispira MORSE, Obs. Terr. Pulm. Maine,
p. 11, type H. alternata Say.
Shell ratherjlarge and solid, with convex spire and open umbilicus ;
whorls rounded or carinated at the periphery. Surface striate,
ribbed-striate or spirally ribbed, obliquely flamed, unicolored or
spirally banded ; lip thin, simple. Type P. alternata Say, pi. 14,
figs. 34, 35, 36.
Animal having a large foot, its length greater than the diameter
of the shell, the tail rounded ; sole without any traces of longitudinal
divisions ; the foot-margins having a wide border above, bounded by
a distinct groove, the grooves meeting over the tail (fig. 40). Eye-
peduncles long and slender, tentacles minute. Mantle edge thick
(pi. 14, figs. 40, 46, P. alternata).
PYRAMIDULA. 49
Genital system simple, lacking accessory organs. Penis receiving
the vas deferens and the retractor muscle at its summit. Spermatheca
bulbous, its duct very long. Ovi-sperm duct very much convoluted,
the ovo-testis consisting of small groups of large club-shaped follicles.
Eye-peduncle retracted between the branches of the genitalia (pl.-
11, fig. 20, P. alternate Say. PL 11, fig. 27, P. strigosa Gld.)
Jaw strong and opaque, arcuate, with a slight or obvious median
projection; surface rather faintly subvertically striated (pi. 11, fig.
18, P. alternata. PI. 11, fig. 17, P. strigosa).
Radula: Central teeth having the mesocone long, side cusps
small. Laterals having a large mesocone and a well developed
ectocone ; no entocone. Marginals similar, but with the basal plate
short, as usual (pi. 11, fig. 23, P. alternata}. This type of dentition
is common to P. alternata, solitaria and idahoensis. In P. cumber-
landiana the side cusps are obsolete on central and inner lateral
teeth.
In P. strigosa and haydeniihe central and lateral teeth lack ecto-
cones. The outer marginal teeth have an ectocone developed, and
sometimes it is split into two minute cusps (pi. 11, fig. 28, P,
strigosa).
The Patulas of eastern America are oviparous, the eggs small.
round, not hard-shelled. P. strigosa and its allies are viviparous,
four to six young occupying the uterus at the same time, the most
mature having a shell of 2! whorls, 3 to 4 mill, diameter, the earlier
2 whorls with fine oblique and spiral stride, marked off by a distinct
line from the latter third of a whorl, which is spirally lirate and
more or less hirsute. The viviparous mode of reproduction has
probably been assumed on account of the aridity of the Rocky
Mountain region. The rains are in this area uncertain, and for
snails mainly unseasonable ; and probably insufficient to insure the
development of eggs committed to the earth in the usual way.
Snails of this section are distributed over the whole of the United
States except the California!! slope. Individuals of the species are
numerous, P. alternata in the East and strigosa in the West being
among the commonest of land snails. They live by preference in
rocky places, the talus of a limestone cliff being a favorite station.
The species are polymorphic to a degree inconceivable to those
who have not actually seen large series of the shells. P. alternata
fergusoni and P. cumber landiana seem to be the extremes of one
series of forms, and P. idahoensis and haydeni of another.
XJNIVEBSIT1
50 PYRAMIDULA.
The name Patula, as well as Eyryomphala, was intended to include
all of the forms referred now to the genus Pyramidula ; and most
recent authors have adopted Patula as the generic name. Such a
course is inadmissible on account of the earlier names Pyramidula,
Gonyodiscus and Discus of Fitzinger ; and there is, moreover, another
difficulty, for Patula, Delomphalus and Eyryomphala were all
proposed in the same year (1837), and it is now impossible to decide
which should be given priority. In von Martens' edition of Albers,
the type of Patula is said to be If. rotundata ; but as that species
was already the type of a prior group (Discus'), we cannot accept
^such a selection. We are, therefore, obliged to consider Held's first
.species, If. alternata, the type.
Species.
P. alternata Say, iii, 57. P. strigosa Gld. (PL 14, f. 37-39.)
scabra Lam. /. depressa Ckll.
strongy lodes Pfr. f.fragilis Hemph. viii, 117.
infecta Parr. /. carnea Heraph. viii, 117.
v. fergusoni Bid. iii, 57. /. rugosa Hemph. viii, 117.
v. mordax Shutt. iii, 57. /. albida Hemph. viii, 117.
P. cumberlandiana Lea, iii, 58. /. buttoni Hemph. viii, 117.
P. solitaria Say, iii, 58. /. globulosa Ckll. viii, 118.
kochi Pfr. v. jugalis Hemph. viii, 117.
subrudis Pfr. v. subcarinata Hemph. viii, 118.
P. idahoensis Newc. iii, 55. bicolor Hemph. viii, 118.
v. newcombi Hemph. viii, 115. lactea Hemph. viii, 118.
/. wasatchensis Hemph. viii, picta Hemph. viii, 118.
[116. v. cooperi W. G. B. viii, 118.
v. binneyi Hemph. viii, 116. P. haydeni Gabb. iii, 57.
/. multicostata Hemph. viii, 116. /. hemphilli Newc. viii. 119.
/. castanea Hemph. viii, 116. /. gabbiana Hemph. viii, 119.
/. albofasciata Hemph. viii, 116. j. bruneri Anc. viii, 119.
/. gouldi Hemph. viii, 116. oquirrhensis Hemph.
P. strigosa Gld. viii, 117. hybrida Hemph.
parma Hemph.
Subgenus ATLANTICA Ancey, 1887.
Atlantica ANC., Conch. Exch. i, p. 54, April, 1887, type H. semi-
plicata Pfr.
Shell small, discoidal, with wide shallow umbilicus and low-convex
spire ; whorls narrow, obliquely ribbed above, polished below, the
PYRAMIDULA. 51
last obstructed far within by several pairs of elevated lamellce upon
the basal-outer wall (fig. 32). Lip thin, simple. Type H. semi-
plicata Pfr. pi. 14, fig. 32, 33.
Anatomy unknown. Distribution, Madeira. This group is prob-
ably a modification of Goniodiscus.
P. semiplicata Pfr. iii, 44. P. calathoides Paiv. iii, 44.
gueriniana Lwe.
Subgenus HELICODISCUS Morse, 1864.
Helicodiscus MSE., Obs. Terrest. Pulm. Maine, p. 25, type H.
lineata Say.
Shell small, disk or coin-shaped, with flat spire and broad, shallow
umbilicus. Whorls numerous, convex and closely coiled, spirally
striated or lirate, the last whorl having one or several pairs of tuber-
cular teeth within, situated upon the basal-outer wall. Aperture
lunate, lip thin, simple. Type P. lineata Say, pi. 14, figs. 29, 30, 31.
The shell lies perfectly flat upon the posterior end of the foot, the
eye-peduncles standing nearly vertically ; posterior end of the long
and narrow foot conspicuously furrowed above, very short behind
the mantle (pi. 14, figs. 47, 48, P lineata).
Jaw arcuate, striate, the strise diverging somewhat from the median
line; median projection inconspicuous (pi. 15, fig. 1, P. lineata).
Morse's figure represents the jaw as less arcuate and pointed at
the ends. -The jaw figured on my plate, however, seems to be per-
fect, although the ends are blunt.
Radula having about 77 rows of 12-M2 or IS'MS teeth. The
central tooth is decidedly narrower than the laterals, its mesocone
very short, side cusps minute. Laterals with large square basal-
plates, the mesocone as long as the basal-plate, eutocone and ectocone
equally developed, strong, with short cutting points. Marginals low,
wide, the ectocone bifid or trifid (pi. 11, fig. 24, P. lineata).
These minute snails live upon decaying wood. The most con-
spicuous features of the dentition are the tricuspid lateral teeth,
recalling those of Stephanoda or Charopa, and unlike the teeth of
Pyramidula generally, in which the entocones are as a rule absent.
The splitting of the ectocones of the marginal teeth is correllated
with the small size of the creature, snails of many groups assuming
the Pupa-like form of marginal teeth when the size of the animal
becomes minute.
52 PYRAMIDULA-PARARHYTIDA.
P. lineata Say, ii, 200. P. fimbriatus Weth. ii, 200.
v. salmonensis Hem ph.
salmonaceus Hemph., W. G. B.
v. sonorensis Coop.
Subgenus ? PUPISOMA Stoliczka, 1873.
Pupisoma STOL., Journ. Asiat. Soc. Beng. xlii, p. 32. PFR.-CLESS,
Nomencl. Hel. Viv., p. 352. v. MOLLENDORFF, Bericht Senck.
naturforsch. Ges., 1890, p. 223.
Shell minute, thin, brown, perforated ; varying from Pupiform,
almost cylindrical, to globose-conoidal ; apex obtuse ; whorls
rounded, with delicate, irregular, cuticular riblets. Aperture
oblique, truncate-oval or rounded, the lip thin, simple, or a little
expanded, broadly dilated at the columella, nearly closing the
umbilical perforation ; the columellar edge sometimes slightly project-
ing, but hardly dentate. Type Pupa lignicola Stol., pi. 14, figs. 41,
42. See also P. philippinicum Mlldff., pi. 14, figs. 43, 44.
Animal having very short eye peduncles and barely a trace of
tentacles. (Stol.}. Jaw, radula and genitalia unknown.
Distribution, India, Borneo, Philippines.
A group of uncertain position. Stoliczka referred it to Pupidce;
v. Mollendorff to the Fruticicola series, near Acanthinula and Zoo-
genites. For the present I prefer to consider it a modification of
Pyramidula, comparable to the American group Ptychopatula ; but
I am not sure that it is not a group of Pupidce.
P. lignicola Stol. P. pulvisculum Iss. iii, 191.
P. orcella Stol. P. philippinicum Mlldff.
P. orcula Bens, ii, 177. P. miccyla Bens, ii, 176.
Genus PARARHYTIDA Ancey, 1882.
Pararhytida ANC., Le Naturaliste 1882, p. 85 ; Bull. Soc. Mai.
Fr. v, p. 360. Platystoma ANC., 1882, Not Platystoma of Klein or
Homes, nor Platyostoma Conr. Saissetia (Bayle) ANC., Bull. Soc.
Mai. Fr. v, p. 368, 1888.
Shell perforate or umbilicate, solid and strong, depressed, acutely
keeled (but periphery rounded in section Saissetia). Baso-columellar
lip thickened by a callus within, and dilated at the insertion. Type
H. dictyodes Pfr.
PARARHYTIDA. 53
Under this generic head may be comprised two groups, as
follows :
Section Pararhylida s. sir.
Shell thick lens-shaped, in form like Trochomorpha. AVhorls about
6, slowly increasing, acutely keeled, basal lip somewhat sinuous. Type
H. dictyodes Pfr., pi. 7, figs. 25, 26, 27.
External anatomy unknown. Jaw arcuate, quite strong, without
median projection, and absolutely smooth (pi. 9, fig. 35, P.
dictyodes').
Radula composed of 22-14-1-14-22 teeth in nearly horizontal
series. Central tooth tricuspid, the mesocone attaining the anterior
border of the basal-plate, side cusps small. Lateral teeth tricuspid,
slightly asymmetrical. Marginal teeth also tricuspid, the entocone
and mesocone united at their bases (pi. 9, fig. 36, P. dictyodes}.
Genitalia : Penis stout, extending into a long flagellum (?), the
vas deferens inserted high upon it ; the stout lower portion bearing
a globose appendix, at the base of which the retractor is inserted.
Vagina is short, muscular and swollen. Spermatheca very large
and long, its duct short ; (in the figure is shown a spermatophore
within it). Albumen gland small ; hermaphrodite duct long, not
convoluted (pi. 9, fig. 37, P. dictyodes).
The notable generic features of the anatomy are that all of the
teeth are tricuspid (as in many Endodontas) ; the jaw is smooth, not
vertically striated ; the penis bears a flagellum and apparently an
appendix. The most important shell characters are the solidity, and
the callous thickening of the baso-columellar lip.
I have considered Pararhytida a genus separate from Endodonta,
mainly on account of the smooth jaw. In Endodonta, Pyramidula,
etc., the jaw is always laminate or striate. In Pararhytida its com-
ponent laminae seem to be completely fused. The characters of the
foot must be examined before we can intelligently discuss the system-
atic position of Pararhytida. Our knowledge of its anatomy is due
to Fischer (Journ. de Conchyl., 1875).
P. dictyodes Pfr. iii, 95. P. mouensis Cr. iii, 95.
v. dictyonina Euth. viii, 134.
Section Saissetia (Bayle) Anc., 1889.
Shell solid, depressed-globose or subdiscoidal, the spire slightly
convex ; umbilicus rather narrow. Whorls rapidly increasing, the
54 PARARHYTIDA-THYSANOPHORA.
last one ivide, rounded at the periphery. Lip generally somewhat
retracted at the upper insertion, thickened on the baso-columellar
margins, dilated at the basal insertion. Surface smooth or rib-striate
above. Type H. saisseti Montr., pi. 7, figs. 22, 23, 24.
The soft anatomy is unknown. Binney has figured the jaw and
teeth of P. astur. The jaw is low, wide, slightly arcuate, ends hardly
attenuated, blunt; anterior surface without ribs; having a wide,
blunt median projection ; a line of reinforcement runs parallel to
the cutting edge ; upper margin with a strong muscular attachment
(pi. 8, fig. 7). The radula has 21-9-1-9-21 teeth. Centrals tri-
cuspid ; laterals lacking the entocone, at least on the inner teeth ;
marginals tricuspid, the entocone and mesocone united.
It will be seen that this differs from typical Pararhytida in the
median projection of the jaw and the loss of entocones on the lateral
teeth (pi. 8, fig. 8).
As no type was designated by Ancey, I have considered H. sais-
seti Montr, as such, for I suppose this was Bayle's intention.
Species.
P. baladensis Souv. i, 116. P. occlusa Gass. i, 122.
P. oriunda Gass. i, 121. P. astur Souv. i, 117.
P. bruniana Gass. i, 119. P. saisseti Montr, i, 117.
P. perroquiniana Cr. P. goulardiana Cr. i, 122.
P. turneri Pfr. i, 119.
Genus THYSANOPHORA Strebel & Pfeffer, 1880.
Thysanophora S. & P., Beitr. Mex. Land- und Susswasser-
Conchylien, iv, p. 30 (proposed for impura, paleosa, conspurcatetla).
PILSBRY, Proc. Acad. Nat. Sci. Phila. 1889, p. 192.
Microphysa MARTENS in Albers, Die Hel., p. 82 ; type Helix
boothiana Pfr. Not Microphysa Westw., 1834 (Hemiptera), nor of
Guen. 1841 (Lepidoptera).
Acanthinula STREBEL & PFEFFER, /. c., p. 31, and of v. MAR-
TENS, Biol. Centr. Amer., p. 130. Not Acanthinula Beck. Ptycho-
patula PILSBRY, Proc. Acad. Nat. Sci. Phila. Sept. 17, 1889, p. 191 ;
Nautilus iii, p. 62 (proposed for cceca, dioscoricola, punctum, plagio-
ptycha, etc.).
Euclasta v. MARTENS, Jahrb. D. M. Ges. 1877, p. 347 (for H.
musicola Sh.). CROSSE, Journ. de Conchy]. 1892, p. 14. Not
THYSANOPHORA. 55
Euclasta Lederer, Verh. Zool.-bot. Vereinsin Wein, v, p. 252, 1855,
and Weiner En torn. Monatschr. vii, p. 423, 1863 (Microlepido-
ptera).
Shell varying from flat and discoidal to depressed-globose and to
conical or pyramidal ; thin; pale brown, yellow or corneous, some-
what translucent or at least not opaque; narrowly umbilicated ; sur-
face rather dull, smooth or with slender riblets (generally cuticular),
or densely, minutely bristly. Embryonic whorl not distinctly
demarked from the after-growth, smooth or granular. Whorls 4-
6 , convex, separated by deep sutures, the last whorl rounded or car-
inated. Aperture lunate or oblong; lip thin, simple or a trifle
expanded, the columellar margin more or less dilated. Type T.
conspurcatella Morel., pi. 16, fig. 3. (See also pi. 16, fig. 4, T.
caca. PL 16, figs. 5, 6, 7, T. hypolepta. PI. 16, figs. 8, 9, 10, T. stig-
matica. PI. 16, figs. 1, 2, T. turbiniformis).
Foot (of T. peraffinis) narrow, the sole not tripartite ; upper sur-
face granulated, the tail having a median sulcus above (pi. 15, fig.
8), sides granulated, with oblique grooves but no distinctly differ-
entiated foot-margin (fig. 9). Tail without mucus pore.
Genital system unknown, but oviduct (of T.peraffinis) containing
several hard and brittle-shelled white eggs. T. vortex has been
observed by Morse to be viviparous. In this genus, therefore, as in
Sagda, both viviparous and oviparous species occur.
Jaw thin and delicate, flexible, strongly arcuate, composed of many
flat, narrow lamellae, the free edges of which appear as vertical sir ice ;
lower margin of jaw denticulated by the lamella (pi. 15, fig. 7, T.
peraffinis. PI. 15, fig. 6, T. turbiniformis).
Dentition: Rhachidian tooth with square basal-plate and three
stout cusps, the mesocone projecting beyond the basal-plate. Lateral
teeth bicuspid, the entocone completely absent. Marginal teeth
various in form ; having either (1) a long oblique mesocone, and a
small simple or bifid ectocone (T.peraffinis pi. 15, fig. 10, and also
T. incrustata, T. ingersolli) ; or (2) the mesocone is bifid by union
with the entocone (T.turbiniformispl. 15, fig. 5, and also T.granum,
T. vortex, T. pubescens). In T. granum, incrustata and vortex the
ectocone is trifid ; in the others it is either simple or bifid.
The jaws and teeth of turbiniformis and pubescens, and the teeth
of T. cceca have been figured by W. G. Binney, Ann. N. Y. Acad.
Sci. iii, pp. 105, 106, 113 ; those of T. incrustata, T. ingersolli and T.
56 THYSANOPHORA.
vortex in Terr. Moll, v, p. 170-173, and Man. Amer. L. Sh., p. 356.
The jaws and teeth of T. perdepressa and T. peraffinis have been
examined by myself. All of these species have essentially the same
type of jaw. The teeth vary only in the denticulation of the mar-
ginals, as noted above. The jaw is distinctly stegognathous in type,
being more like that of Flammulina than that of Pyramidula.
The absence of a parapodial groove widely sunders this genus
from Pyramidula, Cfiaropa, Phasis and Amphidoxa. The first of
these groups differs also in the structure of the jaw. Thysanophora
agrees with Hyalosagda in characters of the jaw, dentition, foot and
the calcareous-shelled eggs.
The shell of Thysanophora somewhat resembles that of Pyrami-
dula ; but it is less opaque, never flame-painted nor strongly rib-
striate. The columella moreover is generally dilated as in Trachy-
cystis.
The species inhabit the Greater Antilles, with a few in Bermuda,
Florida and the Gulf States, and extending to the Middle American
mainland from Vera Cruz and Yucatan south to Trinidad. The
forms from the periphery of this area are small or minute, but in
the large West Indian islands species of considerable size occur.
These snails live upon the ground, under leaves or stones.
More than any other group of Antillean Helices, the Sagda-Thy-
sanophora-Zaphysema group impresses us as being an original West
Indian element. The other main genera of the Antilles, Pleurodonte
and Hemitrochus, with the allies of each, show far-reaching affinities
with Old world Helices ; and Polygyra has been derived from the
North American fauna ; but not only is the Sagda-Thysanophora-
Zaphysema group characteristic of the Antillean region now, but no
Helices known to approach them in morphology of genitalia and
shell have been found in any other part of the world. Thus, as far
as present knowledge enables us to judge, of the three main stocks
into which the West Indian Helix fauna is sharply divided, the
Thysanophora, etc., phylum is that which has longest occupied the
region, and probably developed its peculiar features therein, arising
from some very early, un differentiated Helix stock of the Poly-
placognathous type. The other two great groups are much later
(although still ancient) elements, which reached the Antillean tract
after their essential anatomical features had become well estab-
lished.
THYSANOPHORA. 57
The forms of this genus are so little known anatomically that any
attempt at sectional division would now be premature. By purely
conchological standards, three sections are indicated : (a) Thysano-
pJwra restricted, including small forms having cuticular riblets
more oblique than, and crossing, the growth-lines. This may include-
Ptychopatula (type c.ceea, pi. 16, fig. 4) which differs in being globosely
elevated with only a minute umbilicus. Acanthinula of Strebeland
v. Martens (in Biol. Centr. Amer.) is a synonym. (2) forms of the
type of vortex, with smoother surface, the spire varying from flat to
pyramidal ; mostly Antillean. . T. ptychodes, T. turbiniformis (pi.
16, figs. 1, 2), etc., belong here also. (3) Larger forms, with the
spire mostly depressed, sometimes concave, the surface minutely
roughened or bristly, such as T. stigmatica (pi. 16, figs. 8, 9, 10), T
suavis, T. velutina, etc., from the Greater Antilles, and T. sigmoides
from Guatemala.
Species of the mainland, Trinidad to Texas and Florida, etc.
T. conspurcatella Morel, iii. 50. T. venezuelensis Jouss. viii, 112.
T. impura Pfr. iii, 50. T. rojasi Jouss. viii, 112.
T. incrustata Poey. ii, 204. T. vortex Pir. iii, 93.
T. ingersolli Bid. iii, 101. T. turbinella Morel, iii, 51.
T. paleosa Streb. iii, 50. T. csecoides Gupp. iii, 55.
T. granum Streb. iii, 55. T. guatemalensis C. & F. ii, 174.
T. ierensis Gupp. iii. 55. T. coloba Pils.
T. plagioptycha Sh. ii, 174. T. punctum Morel, iii, 53.
T. dioscoricola C. B. Ad. ii, 174. T. intonsa Pils. viii, 111.
T. cseca Guppy. iii, 55. T. sigmoides Morel, iii, 101.
T. bactricola Guppy. iii, 55. vitrinoides Tristr.
T. hornii Gabb. iii, 21.
[Of the above species, T. granum and ierensis are probably mere
varieties of plagioptycha, and T. cceca and punctum varieties of dios-
coricola. Specimens of all the above, except turbinella, guatemalensis,
venezuelensis, rojasi and punctum are in the collection of the
Academy.]
Species of the West Indies and Bermuda,.
T. alveus C. B. Ad. iii, 98. T. musicola Shutt. iii, 97.
T. angustispira C. B. Ad. iii, 97. v. major Crosse.
T. anthoniana C. B. Ad. iii, 96. T. peraffinis C. B. Ad. iii, 98.
58
THYSANOPHORA-SAGDA.
T. arecibensis Pfr. iii, 58.
T. boothiana Pfr. iii, 97.
v. vitrina C. B. Ad. iii, 97.
T. brevior C. B. Ad. iii, 99.
depressa Ad.
T. cyclostomoides Pfr. iii, 100.
T. debilis Pfr. iii, 101.
fragilis Pfr.
T. desiderata Pfr. iii, 96.
T. diminuta C. B. Ad. iii, 99.
T. dioscoricola C. B. Ad. ii, 174.
T. elatior Weinl. & Mts. iii, 97.
T. euclasta Shutt. iii, 97.
wuifti Pfr.
T. fuscula C. B. Ad. iii, 98.
T. gracilis Poey.
T. hilum Weinl. & Mts.
T. hypolepta Shutt. viii, 111.
T. iramunda C. B. Ad. iii, 99.
T. inaguensis Weinl. iii, 41.
T. inconspicua C. B. Ad. iii, 99.
T. incrustata Poey, ii, 201.
inerassata Rve.
saxicola Gld.
T. jeannereti Pfr. iii, 53.
T. krugiana Mart.
T. leucoraphe Pfr. iv, 77.
T. montetaurica Pfr. iii, 97.
T. perdepressa C. B. Ad. iii, 100.
T. plagioptycha Shutt. ii, 174.
T. portoricensis Pfr. iii, 96.
T. prominula Pfr.
T. pruinosa Pfr. iii, 186.
T. ptychodes Pfr. iii, 100.
T. pubescens Pfr. iii, 184.
T. raripila Morel, iii, 101.
T. rufula Pfr. iii, 99.
T. sincera C. B. Ad. iii, 99.
T. spreta C. B. Ad. iii, 98.
v. errans Ad. iii, 98.
T. stigmatica Pfr. iii, 100.
T. suavis Gundl. iii, 100.
T. subaquila Shutt. iii, 98.
T. tichostoma Pfr. iii, 100.
lamellina Newc.
T. translucens Gundl. iii, 96.
T. turbiniformis Pfr. iii, 96.
subpyramidaJis C. B. Ad.
macnabiana Chitty.
pyramidatoides d'Orb.
T. velutina Lam. iii, 100.
T. virescens Pfr. iii, 96.
T. vortex Pfr. iii, 98.
selenina Gld.
otellina Rose.
v. bracteola Fer.
Genus SAGDA Beck, 1837.
Sagda BECK, Index Molluscorum p. 9 (for alveolata B. and
aiistralis Ch. B.). A LEERS-MARTENS, Die Hel. p. 76. SHUTTLE-
WORTH, Bern. Mittheil. 1853, p. 85. See also BINNEY, Ann. N. Y.
Acad. Sci. iii, p. 88. SEMPER, Reisen im Archip. Phil., Landmoll.,
p. 128, and PILSBRY, Proc. Acad. Nat. Sci. Phila. 1892, p. 213.
Epistilia SWAINS. Malacol., p. 165, type E. conica Swains., I. c., f.
18a \_S.jayanaT\. Epistyla SWAINS., /. c., p. 331, type E. conica
Sw. [ $. cookiana /].
-\-Hyalosagda ALB. and Odontosagda MARTENS, Die Hel., p.
77, 78.
SAGDA. 59
Shell having the texture of Zonites or Hyalinia, imperforate or
umbilicate, varying in form from depressed and subdiscoidal to
globose-conic or trochoidal ; whorls 6-9, narrow and slowly increas-
ing, the last not deflexed in front. Aperture nearly vertical, lunate,
either with or without internal laminse; lip thin, sharp and simple,
slightly dilated or reflexed at the axis ; columella short, having a
callous fold, or thin and simple. Type S. cookiana Gmel., pi. 16,
figs. 11, 12, 13. (See also pi. 16, figs. 14-20).
Animal viviparous, or oviparous with hard-shelled eggs.
Foot extremely long and narrow (the sole in S. similis measuring
length 20, width 3 mill.), strongly granulated above, the tail having
an impressed median longitudinal line or groove, acute behind ; sides
of foot without longitudinal grooves, but marked by a zigzag line
(pi. 35, fig. 7) ; sole not divided longitudinally (pi. 35, fig. 8, S.
similis).
Genital system much elongated, the vestibule short. Penis long,
the vas deferens and aflaaellum inserted at its apex (pi. 35, fig. 2),
and an elongated appendix inserted at the lower third (appendix of
S. similis seen convoluted in the normal manner in pi. 35, fig. 2,
partially straightened out in pi. 35, fig. 3). Vagina long and nar-
row ; uterus larger, containing eggs or young shells; dud of sper-
matheca very long, expanded near the base (pi. 35, fig. 4, S. similis).
See also pi. 21, fig. 9, penis of S. cookiana, p. penis, a. appendix, r.
p. retractor muscle,^, flagellum. Fig. 10 shows the appendix parti-
ally uncoiled.
Jaw thin, arcuate, smooth, with a slight median projection or none
in S. foremaniana, haldemaniana, jay ana and cookiana (pi. 21, fig. 8).
In S. similis (pi. 35, fig. 6) it is thin, arched, and of the stegognath-
ous type, being composed of 27 narrow flat vertical plates, soldered
together, the outer imbricating edges of which are distinctly visible.
Radula having the transverse rows of teeth nearly straight.
Central teeth having a square basal-plate and three cusps, the meso-
cone longer than the basal-plate. Lateral teeth bicuspid, the meso-
cone long. Marginal teeth also bicuspid. (PI. 21, fig. 7, S. cook-
iana ; pi. 35, fig. 5, S. similis; pi. 35, fig. 1, S. haldemaniana').
The jaw of foremaniana has been described by Semper, that of
haldemaniana smdjayana by Binney, that of cookiana by myself;
all agree in being smooth (oxygnathous) as described above. The
jaw of S. similis, examined by the writer, is of the plaited type. The
teeth of for email iana are described by Semper, those of connectens,
60 SAGDA.
haldemaniana andjayana by Binney. All agree with the descrip-
tion given above, and with those of S. similis and S. cookiana
examined by the writer.
The prominent features of the shell in this genus, are its Zonites-
like texture, the subvertical aperture, and sharp simple lip. The
genital system is peculiar for its appendix and flagelium on the
penis, and the long spermatheca duct. The teeth of the species
investigated agree in the long mesocones, constantly present ecto-
cones, and bicuspid marginals. The jaw in the typical forms is
smooth, by the complete union of its component laminae; in the
section Hyalosagda, which is nearer the ancestral stock, the jaw shows
vertical imbricating plates, as in Tkysanophora, Flammulina, etc.
The genus Sagda is by no means so isolated in the family of
Helices as has been supposed. Its relationship with Thysanophora
and especially with Zaphysema, is moderately intimate.
The analogy of the shell of Sagda with that of the Zonitid genus
Gastrodonla is remarkable. Both contain forms with spiral internal
laminae, and depressed forms without laminae ; the section Hyalo-
sagda being quite comparable to the section Zonitoides. Gastrvdonta
too, has elevated forms (G. ligera, etc.) recalling Sagda in contour.
Sagda is confined to the island of Jamaica, with the exception of
the subgenus Odontosagda inhabiting Haiti and Cuba. The
species and forms are numerous, extremely variable, and correspond-
ingly difficult to determine. This difficulty is enhanced by the fact
that some of the best specific characters can be seen only by break-
ing an opening in the last whorl a half volution behind the aperture ;
the vicinity of the suture being the best place for the incision. By
this means only may the form and length of the lamellse be observed,
as is the case with Plectopylis and some other groups. The lamellae
are present in young specimens, but are progressively absorbed as
the animal grows, so that in adults they do not extend inward much
beyond the last whorl. The basal lamina is sometimes totally
absent in species normally possessing it, just as in Gastrodonta ; but
as in that genus, it is a comparatively rare condition in most species.
Subdivisions.
Section Sagda (restricted). Shell imperforate, the axis solid ;
aperture having a spiral lamina within the last whorl and generally
a fold on the columella. Type S. cookiana, pi. 16, figs. 11-13. (See
also pi. 16, figs. 16, 17, S. connectens, and pi. 16, figs. 14, 15, S. alligans.)
SAGDA. 61
Animal viviparous, the young at birth being depressed-globose,
flattened above, thin, translucent, perforated ; composed of 2
whorls; measuring nearly one-fifth the diameter of the adult. We
have observed young shells in specimens of cookiana, foremaniana
and ambigua.
Section Hyalosagda Martens. Shell perforate or umbilicate,
glassy, thin and depressed. Aperture with no internal lamellaB or
teeth. Type S. similis, pi. 16, figs. 18, 19, 20.
Animal oviparous, the eggs short-oval, with a hard, white, smooth
calcareous shell ; the length of its longest axis contained 5 to 6 times
in the diameter of the shell. We have found eggs in S. haldemani-
ana and S. similis.
Subgenus ODONTOSAGDA Martens. Small, thin and umbilicated ;
internal laminae interrupted ; columella thin.
The extreme difficulty of the genus, and the inadequacy of the
accounts of it in the works of PfeifFer, Reeve and Tryon, induce me
to offer the following key to the species. Shuttle worth has published
an excellent revision of the group. All known Jamaica species are
represented in the collection of the Academy.
Key to the species of Sag da.
I. Aperture provided with internal teeth or laminae.
a. Basal lamina either interrupted, or less than i whorl in
length ; base very convex ; form subglobose or globose-conic.
6. Base deeply impressed at columella ; basal lamina
interrupted.
c. Globose-conic ; solid, strong, yellow ; col-
umellar fold weak or obsolete; basal lamina
interrupted forming several teeth ; whorls 9 ;
alt. 21, diam. 22 mill. cookiana.
bb. Base not deeply impressed ; basal lamina continuous ;
columella with a nodule-like fold.
c. Globose-pyramidal ; base globosely convex,
not impressed at columella ; solid, strong,
yellow ; columellar fold a strong nodule,
not entering; basal lamina very strong, i
62 SAGDA.
whorl long ; whorls 7 ; alt. 16-18, diam. 17
mill. foremaniana.
cc. Globose ; thin, subtranslucent, corneous ;
base globosely convex, hardly impressed ;
columellar fold strong and heavy, spirally
entering; basal lamina strong, short, one-
fourth whorl long ; whorls 6; alt. 12, diam.
12-13 mill. pila.
GGC. Globose-subconic ; thin but rather solid,
translucent ; base very convex, only slightly
impressed ; columellar fold a stout nodular
callus, somewhat entering; basal lamina
strong, about i whorl long ; an additional
small fold developed between basal and col-
umella folds; whorls 6-7 ; alt. 16, diam. 16
mill.; alt. 11, diam. 13 mill, triptycha.
aa. Basal lamina 3 to whorl long, continuous ; base not not-
ably convex.
b. Large, solid and elevated.
G. Trochoidal ; base not excavated in the
middle; no columellar fold; basal lamina
deep-seated, about * whorl long ; whorls 8 ;
alt. 24-26, diam. 27-30 mill. alligans.
cc. Elevated trochoidal ; base deeply excavated
in the middle ; columellar fold and basal
lamina strong within, spirally entering, but
sometimes neither is visible from the aper-
ture ; whorls 9 ; alt. 25, diam. 27-28 mill. ;
alt. 18, diam. 24 mill. jayana.
bb. Small, thin, depressed.
c. Depressed-subglobose, thin, subtranslucent
greenish-yellow, the surface minutely spic-
ulose ; base slightly excavated ; columella
calloused ; lamina peripheral in position,
extending nearly to lip-edge ; whorls 6 ; alt.
9, diam. 13 mill. lamellifera.
cc. Subdiscoidal, thin, translucent, polished ;
base excavated ; columella calloused ; basal
lamina not deep-seated, i $ whorl long;
whorls 6J ; alt. 6J, diam. 11 mill.
osculans.
SAGDA. 63
aaa. Basal lamina one whorl long or more.
b. Lamina peripheral in position, lamellifera.
bb. Lamina basal in position.
c. Much depressed-trochoida}, solid, somewhat
translucent, smooth ; base but little excav-
ated; columellar fold small or obsolete;
basal lamina more than a whorl long;
whorls 7-1-9 ; alt. 16, diam. 23-25 mill.
connectens.
cc. Trochoidal, solid, strong, costulate-siriate
above the periphery ; no columellar fold;
basal lamina a full whorl long; whorls 7;
alt. 17, diam. 20 mill.; alt. 13, diam. 16
mill. epistylioides.
ccc. Globose-trochoidal, solid, strong ; base some-
what excavated ; columellar fold strong and
lamellar within ; basal lamina strong, about
one whorl long; whorls 8; alt. 18, diam.
19 mill. Smaller and paler than jay ana,
with less excavated base, but probably a
variety ofjayana. alveare.
cccc. Elevated trochoidal, solid, strong and
opaque; base deeply excavated ; columellar
fold and basal lamina strong within, but
often not visible from the aperture ; whorls
9 ; alt. 25, diam. 27-28 mill. ; alt. 18, diam.
24 mill. jayana.
ccccc. Globose-trochoidal, solid but thin, covered
with a yellow cuticle bearing minute spicules
or pitted; base depressed, excavated; col-
umella with a strong lamellar fold ; basal
lamina long; whorls 8; alt. 12, diam. 14J
mill. spiculosa.
cccccc. Elevated, pyramidal, solid, subtranslucent ;
upper whorls finely costulate-striate; base
narrowly and but little excavated ; columella
having a blunt callous fold, spirally enter-
ing ', ^ basal lamina strong, about one whorl
long; whorls 8-9 ; alt. 17, diam. 16 mill.,
torrefacta.
64
SAGDA.
II. Aperture lacking internal teeth or laminae.
a. Umbilicus moderate, its breadth contained 9-15 times in
diameter of shell.
b. Diam. 13-16, alt. 7-9 mill. similis.
bb. Diam. 9-10, alt. 5-5 J mill. hollandi.
bbb. Diam. 3-4, alt. 1-H mill. brevis.
aa. Umbilicus reduced to a perforation partly closed by the
reflexed columelJa, or wholly closed and imperforate.
b. Imperforate; base depressed; alt. lli-12, diam.
5 i-7 mill. osculans v. delaminata.
bb. Perforated ; base convex, well impressed in the
middle.
c. Diam. 11-12 mill. haldemaniana.
cc. Diam. 9-10 mill. ambigua.
Species of Sagda.
[See pi. 16, figs. 11-13, S. cookiana \ pi. 16, figs. 14, 15 f 8. alligans;
pi. 16, figs. 16, 17, S. connectens-, pi. 16, figs. 18-20, S. similis.]
S. cookiana Gmel., iii, 6.
austral is Chem., auct.
conica Swains.
epistylium Dillw., Sowb.
foremaniana Rve.
S. foremaniana C. B. Ad., iii, 7.
S. pila C. B. Ad., iii, 8.
S. triptycha Shuttl., iii, 7.
S. alligans C. B. Ad., iii, 6.
epistylium Pfr. & Rve., not
/ alveolata Beck. [Mull.
S. connectens C. B. Ad., iii, 6.
S. osculans C. B. Ad., iii, 8.
v. delaminata Ad.
S. ambigua C. B. Ad., iii, 9.
S. lamellifera C.B. Ad., iii, 8.
S. epistylioides Fer., iii, 6.
S. jayana C. B. Ad., iii, 6.
f alveolata Beck (undesc.).
cookiana Pfr.
alligans Rve.
sayana Alb.-Martens.
f conica Swains.
S. alveare Pfr., iii, 7.
S. spiculosa Shuttl., iii, 7.
S. torrefacta C. B. Ad., iii, 7.
(Section Hyalosagda*).
S. similis C. B. Ad., iii, 9.
S. haldemaniana Ad., iii, 8.
arboreoides Ad.
S. hollandi C. B. Ad., iii, 9.
S. ? brevis C. B. Ad., iii, 9.
SAGDA-ZAPHYSEMA. 65
Subgenus ODONTOPAGDA Martens, 1860.
Odontosagda MARTENS, in Albers, Die Hel., p. 78.
Shell small, depressed, thin, whitish, smooth, umbilicated, with 5-6
convex narrow whorls ; base convex. Aperture subvertical, lunate, -
the lip thin and simple ; interior having upon the basal wall several
spiral laminw interrupted into teeth, or with such a spiral lamina and
a series of transverse blades ; eolumella thin, not calloused nor toothed.
Type S.polyodon (see pi. 20, figs. 35, 36, S. hillei GundL).
Anatomy unknown. Distribution, Haiti and eastern Cuba. This
group differs from the toothed Sagdas of Jamaica in the perforated
or umbilicate shell and the interrupted laminae.
S. polyodon Weinl. & Mart, ii, S. blandi Weinl. iii, 8.
[201. S. hillei Gundl. ii, 199.
Species erroneously referred by authors to Sagda : H. epistyliulum
C. B. Ad. is a Guppya. H. circumfirmata and discrepans belong to
the Zonitidce, genus Pcecilozonites.
Genus ZAPHYSEMA Pilsbry, 1894.
Cysticopsis, in part, of authors.
Shell globose, thin, unicolored brownish, smooth except for slight
growth-wrinkles ; imperforate, the axis solid ; composed of 5 to (5
convex whorls the embryonic shell consisting of two whorls, its junc-
tion with the after-growth marked by an indistinct oblique line ; the
last whorl much wider, large and inflated, hardly deflexed in front.
Aperture large, round-lunate, moderately oblique, and toothless ; the
lip thin, sharp and simple, dilated and closely appressed at the
white-calloused eolumella. Type Helix tenerrima C. B. Ad., pi. 16,
fig. 21.
Foot black, rather short, granulated and obliquely grooved above,
as in Thysanophora and Sagda, the tail obtuse, having a median lon-
gitudinal groove above ; anterior half of the foot traversed on each
side by an obliquely descending groove arising about the middle of
the mantle insertion. Sole indistinctly tripartite.
The figures of the foot of Thysanophora peraffinis (pi. 15, figs. 8,
9, upper and lateral views) well represent that of Z. tenerrima also.
Genital system having a short vertibule. Penis long, the vas def-
erens inserted near the apex, where a long flagellum and a curved
appendage are inserted ; at the lower third of the penis arises an
, UNIVERSITY
r^.-" /
66 ZAPHYSEMA.
appendix, which seems to be glandular, and terminates in two long
flagellum-like organs ; the retractor-muscle arises from a median
dilation of the penis. Vagina short, narrow ; uterus enormously
distended with young shells. Spermatheca globular, situated on a
very long duct, which is apparently branched (pi. 35, fig. 12, Z. tener-
rima).
Jaw wide, arcuate, with a slight median projection ; composed of
narrow vertical flat plates soldered together, their outer imbricating
edges appearing as delicate spaced vertical strise ; above projects a
narrow conical process, springing from the middle of its surface (pi.
35, fig. 10, Z. tumida).
Radula composed of short teeth with square basal-plates. Centrals
having the mesocone about as long as the basal-plate, and very
broad, side cusps small but well developed. Lateral teeth similar,
but lacking entocones. Marginal teeth low and wide, the mesocone
large, sometimes bifid at the apex ; ectocone simple or bifid (pi. 35,
fig. 11, Z. tenerrima', pi. 35, fig. 9, Z. tumida').
Distribution, Jamaica.
The shell in this genus is globose, with large body-whorl, spire
convex or low-conoidal, lip sharp and thin. The jaw is like that of
Thysanophora and Sagda in structure, being of the stegognathous
type. The dentition closely resembles that of the two genera named,
but in Sagda the mesocones are longer. The foot in the three genera
Thysanophora, Sagda and Zaphysema is practically the same in
structure. The genital system is similar in general features to that
of Sagda. The modes of reproduction are identical in the three
groups.
Binney has examined the jaw and teeth of Z. tumida; the writer
has figured the teeth and genitalia of Z. tenerrima. The other species
are still unknown anatomically.
The group Cysticopsis, in which these forms have hitherto been
placed, differs widely from them in anatomical features. It must be
included in the genus Hemitrochus as a sectional division.
In the single individual of Z. tenerrima examined, the thin-
walled uterus contained 27 young shells, and an egg, which was
globular, with thin brittle white shell. The young shells are
depressed-globular, translucent, often iridescent, and measure alt.
1'5, diam. 2 mill. ; whorls two. It would seem that in Thysanophora,
Sagda and Zaphysema eggs are normally formed, having the shell
.hard and calcareous. In some species of each group the eggs
PRATICOLELLA. 67
develop and hatch within the uterus, the young snails consume their
egg-shells, using the lime for shell-building; they attain a growth of
about two whorls or more before birth. In other species this pro-
longation of the antenatal period has not been established, and hard-
shelled eggs are brought forth.
Species of Zaphysema.
Z. macmurrayi C. B. Ad., v, 7. Z. tumida Pfr., v, 8.
Z. buddiaua C. B. Ad., v, 7. tmiicata C. B. Ad.
Z. muuda C. B. Ad., v, 9. Z. tenerrima C. B. Ad., v, 8.
Z. columellata C. B. Ad. v, 9.
Genus PRATICOLELLA v. Martens, 1892.
Praticola STREBEL & Pfeffer, Beitr. Mex. Land- und Su'ssw.
Conch, iv, p. 38, 1880, type P. ocampi. Not Praticola Swains.,
1837.Praticolella v. MART., Biol. Centr. Arner., Moll., p. 138. Dor-
casia BINNEY, Terr. Moll, v, p. 346, not of Gray. See for anatomy,
W. G. BINNEY, I. c. ; SEMPER, Phil. Archip. p. 246 ; STREBEL &
PFEFFER, I. c.
Shell of the ordinary Helix shape; narrowly umbilicated, globose,
shining, opaque white or yellowish with translucent corneous and
brownish spiral bands, the most constant band supraperipheral in
position. Aperture lunate-rounded, slightly oblique, lip narrowly
reflexed, dilated at the columellar insertion, sometimes thickened
within. Type P. ampla Pfr. (see pi. 20, figs. 26, 27, P. griseola ; pi.
20, fig. 28, P. berlandieriana ; pi. 20, figs. 29, 30, 31, P. flavescens).
Mantle having both right and left body-lappets; sole indistinctly
tripartite, the central area not sharply separated from the sides, but
darker colored (in spirit).
Genitalia (pi. 21, figs. 1-4, P. ocampi=ampla) Female organs as
in Polygyra, without dart sack, mucus glands or other accessory organs ;
spermatheca oval, its duct simple and very short. Penis large, the
vas deferens inserted at its apex ; retractor trifid, one branch inserted
at apex and one at middle of penis, with a small branch to vas defer-
ens (fig. 2). Cavity of penis containing a tongue shaped papilla (pi.
21, fig. 3), inserted near apex of cavity ; a fleshy 'ridge arising at the
insertion of the vas deferens runs nearly to the base of penis. At
the lower third of the penis is inserted a large, club-shaped appendix,
68 PRATICOLELLA-POLYGYRA.
opening into the penis by a narrow aperture, and containing two
strong longitudinal fleshy ridges (pi. 21, fig. 3). Talon coronated
(pi. 21, fig. 4).
Jaw arcuate without median projection, sculptured with numerous
(12-14) broad, crowded ribs, denticulating both margins, (pi. 21,
fig. 5, P. ampla).
Radula having the central teeth tricuspid, mesocones with a long
reflection, the cutting points projecting beyond the basal-plates, ecto-
cones shortly reflected with long cutting points. Laterals similar
but lacking entocones. Marginal teeth low, wide, the mesocone and
ectocone both bifid (pi. 21, fig. 6, P. griseola).
Distribution : eastern Mexico and Texas. The species live in
open fields and chaparral.
The most important anatomical features of this group are the
simplicity of the female generative system, which is like Polygyra in
its' short spermatheca duct, lobed talon and other characters; the
male system being also like Polygyra except that the retractor has a
triple insertion, and the penis has a large appendix. Jaw as in
Polygyra, section Stenotrema ; teeth of radula as in Polygyra.
External features also like Polygyra. Our knowledge of the anat-
omy is due to the investigations of Leidy,Binney, Semper and Pfeffer.
Von Martens is in error in attributing a dart sack to this group, and in
placing it as a subgenus under Helix s. str. ; it is intimately allied to
Polygyra, the large appendix and split penis retractor being the only
anatomical features separating Praticolella from Polygyra, the texture
of the shell offering another differential feature.
P. griseola Pfr., iv, 76.
P. ampla Pfr. cicercula Fer., Dh.
ocampi Streb. iv, 76. pisum Beck.
P. flavescens (Wiegm.) Pfr., iv, albocincta Binn.
[75. albozonata Biun.
P. berlandieriana Moric. iv, 76. albolineata Old.
pachyloma Mke. splendidula Ant.
Genus POLYGYRA Say, 1818.
Polygyra SAY, Journ. Acad. Nat. Sci. Phila, i, p. 278 (proposed
for auriculata, avara and seplemvolva). PILSBRY, Proc. Acad. N. S.
Phila. 1889, p. 193 ; 1892, p. 400.
Plus Dcedalocheila BECK, Index, p. 21 (for auriculata, avara and
implicata). Triodopsis RAFINESQUE, Journ. de Phys., etc., Ixxxviii,
POLYGYRA. 69
p. 425, 1819 ; Enum. and Acct. etc., p. 3, 1831 (type Tr. lunula,= H.
tridentata Say). -\-Menomphis RAF., /. c. Xolotrem,a RAF., I. c.,
(proposed for X. lunula, X. triodopsis and X. clausa, all undescribed
and unidentified). Odotropis, Chimotrema and Toxotrema RAF.,
Jotirn. de Phys., t. c., p. 425 (y=Stenotrema). Stenotrema RAF., /. c.~
(type S. convexa=H. stenotrema Fer.). Aplodon RAF., I. c. (type A.
nodosum ; undescribed and unidentified). Stenostoma RAF., Enum.
and Account, 1831 (type S. convexa Raf.). Mesodon RAF., /. c.
(type M. maculatum Raf., unidentified). Trophodon and Odomph-
ium RAF., 1. c. unidentified. Ulostoma ALBERS, Die Hel. 1850, p.
95 (=Polygyra s. str., Stenotrema, Triodopsis, etc. Not Ulostoma
TRYON!). Patera ALBERS, I. c., p. 96 (=Mesodon auct.). Cyclo-
doma SWAINS, (part), Malacol., p. 193. Tridopsis BECK, Index
Moll., p. 22 ; GRAY, P. Z. S. 1847, p. 173, type If. plicata.
Helicodonta (in part) FER., Prodrom., p. 33. Anchutoma (in part)
H. & A. AD., Gen. Rec. Moll, ii, p. 205, 1858. Angystoma (in part)
KLEIN, Tent. Meth. Ostr., p. 10, 1753 (pre-Linnsean).
Neohelix v. IHERING, Zeitschr. f. Wissensch. Zool. liv, p. 482,
1892 (=Polygyra Pils.).
Con/. W. G. BINNEY, Terr. Moll, v, and STREBEL & PfeflFer,
Mex. Land- u. Siisswasser-Moll. (anatomy).
Shell helicoid, varying from globose or depressed-globose to lens-
shaped or planorboid, the periphery carinated or rounded ; umbilicus
either open or closed. Surface striated or hirsute ; corneous, yellow
or brown, generally unicolored, but sometimes with many bands, the
most constant being supra-peripheral, the others when present being
wholly indefinite in number and position. Lip well reflexed; aper-
ture typically obstructed by three teeth, one parietal, two upon the
lip ; but any or all teeth often wanting.
Animal externally as in Helix, the mantle subcentral, foot rather
long and narrow, not distinctly tripartite below, and without longitu-
dinal grooves above the lateral margins, although a sort of foot-
margin is produced by the tessellated granulation of the edge. Sur-
face rather coarsely irregularly granulated, the granulation finer
posteriorly ; back with a pair of indistinct grooves extending from
mantle to facial area; sides of foot, and sides and top of tail without
any distinct oblique or longitudinal lines, irregularly granulated ; tail
rounded above, obtuse behind. Mantel edge reflexed to correspond
with the lip of the shell, its edge even ; shell lappets none ; body-
70 POLYGYRA.
|appets small, the right one long, giving off a short ascending branch
behind the lung-pore ; left lappet very small, short.
Genitalia completely lacking accessory organs ; retractor and vas
deferens inserted at the apex of the penis. Spermatheca oval or
oblong, situated upon a short simple duct (pi. 30, fig. 6, P. troost-
iana; pi. 30, fig. 12, P. inflecta; pi. 30, fig. 20, Rclausa-, pi. 31,
fig. 27, P. spinosa ; pi. 21, figs. 12-16, P. albolabris^ The penis
is divided internally into two parts: (1) a lower, .invertible portion,
the inner surface of which shows few or many longitudinal folds,
which are smooth and may be either weak or strong and acute ; and
(2) an upper portion the cavity of which has finely corrugated walls
and is partially filled by one or two fleshy pillars adherent along
the sides.
Jaw arcuate, solid and strong, sculptured with 7 to 20 strong con-
vex ribs; cutting edge without median projection, but denticulated
by the ribs (pi. 30, fig. 19, P. sayi Binn. ; pi. 30, fig. 21, P. kiawaen-
sis Simp.; pi. 21, fig. 11, P. albolabris Say).
Distribution : North America (exclusive of some parts of the south-
western U. S.) ; Cuba, Bahamas and Bermuda.
The white-lipped Helices of North America form a very distinct
and homogeneous genus, well distinguished by characters of the shell
and still more by those of the soft parts. The group, in practically
its present limits, was first defined in 1889, by the writer; subse-
quently the European forms supposed by former authors to be allied
to Triodopsis were shown to differ generically (Journ. de Conchyl.
1891, p. 22). Dr. H. v. Ihering has more recently discussed the
genus, under the new name, Neohelix (Zeitschr. f. wissenschaftl.
Zool. 1892, p. 482). This name must be considered superfluous, on
account of the priority of no less than twenty other more or less
available generic or subgeneric names proposed by various authors.
No snails referable to Polygyra have been found in any part of the
Old World, or in South America, either living or fossil. It is there-
fore highly probable that the genus arose and developed its peculiar-
ities upon eastern North American soil. The West Indian species
are to be regarded as stragglers from the continental fauna, just as
Hemitrochus, Liguus and Thysanophora in Florida are emigrants
from the Antillean fauna. A former connection between southern
Florida and the Great Antilles is demonstrated by the Pliocene
fauna of the former; but the connection was probably not direct,
POLYGYRA. 71
but by way of the Bahama bank, which had previously been
connected with Cuba and Haiti.
The question of the relationships of Polygyra is beset with diffi-
culties. I had formerly grouped the genus with Pyramidula, etc,
but the characters of the foot peremptorily forbid such association.
Dr. v. Ihering suggests the possibility that it may be either a modi-
fied branch of Arionta in which thegenitalia have become simple by
degeneration, or a further development of Patula. The latter hypoth-
esis is untenable. The former has as yet no facts to support it.
No fossils now known throw light upon the problem. From what
we know of the living forms of Polygyra, it is likely that their
common ancestor possessed a shell with tridentate aperture, reflected
lip, and a color-band above the periphery. It is not unlikely that
the group represents an early stage of the true Helix phylum, which
did not share the evolution of the accessory organs of the genitalia
now characteristic of the Pentatcenia, Campylcea, Cochloetyla, etc.
Polygyra divides into three sections, typically very distinct in
in appearance, but closely connected by more or less intermediate
species. The anatomy is practically the same throughout.
Section Polygyra Say, (restricted).
Shell depressed ; umbilicated, or having a curved groove caused
by the tangential deviation of the last whorl. Aperture somewhat
kidney-shaped or ear-shaped, the lip continued in an elevated v-shaped
callus across the parietal wall : outer lip having two teeth or none.
Type P. septemvolva Say, pi. 30, figs. 1, 2, 3. (See also pi. 30, fig.
4, P. auriculata Say).
Central teeth tricuspid, the side cusps well developed ; laterals
bicuspid ; marginal teeth generally having the mesocone bifid at tip,
at least on the extreme margin of the radula, ectocone simple (pi.
30, fig. 5, P. septemvolva ; pi. 30, fig. 7, postelliana*). Genital system
as described above (pi. 30, fig. 6, P. troostiana).
This section comprises some very aberrant species, but the
extremes are so closely connected by intermediate forms that no use-
ful subdivisions can be maintained. The synonymy of the restricted
section Polygyra comprises the names Dcedalocheila, Ulostoma and
Cyclodoma.
The species inhabit the Southern States, a few ranging as far north
as South Carolina, Kentucky and Missouri, extending southward
throughout Mexico. In the West Indies species are found in the
72 POLYGYRA.
Bermudas, Bahamas and Cuba. Most species, such as cereolus, aurifor-
mis, mooreana, etc. are gregarious, and occur in great numbers. All
are ground snails, living at the roots of grass, or under bits of wood
or leaves ; and while some forms such as auriformis are found only
in the immediate proximity of water, others occur in very dry situa-
tions, the arid mesquite chaparral of southern Texas being inhabited
by texasiana and mooreana.
Species without teeth on the outer lip.
Bland has published an excellent essay upon these forms in
Annals N. Y. Lyceum vii, 132, 1860, but his material was not
extensive enough to show the intermediate forms now known. The
forms included under P. cereolus are absolutely connected by a series
of transitions, in which the supposed specific characters found in the
striation or ribbing, the degree of carination, number of whorls, form
of umbilicus and presence or absence of an internal lamina, blend
by imperceptible degrees.
The typical cereolus is found on the Florida keys and adjacent
mainland ; it passes into the smaller form carpenteriana, which con-
tinues up the coast, mainly westward ; occurring also at Matanzas,
Cuba! In central and eastern Florida septemvolva occurs, its small
race volvoxis spreading north to St. Simon's I., Georgia, and to the
west (under the name febigeri) it occurs at New Orleans, La., and
Galveston, Texas. Var microdonta, which is typically quite distinct
in its fine striation, occurs abundantly in Bermuda, and also on New
Providence (at Nassau), Bahamas. At the latter locality transition
forms occur; and it must also be noted that some specimens of
volvoxis from Florida (Tampa) and carpenteriana (Key Biscayne)
show striation equally fine. Species of this group inhabit the neigh-
borhood of the sea, and generally occur in great numbers. Besides
the species enumerated below there is another Polygyra with tooth-
less outer lip, P. anilis ; but its relationships are with an entirely
different group of forms.
(Key to species and varieties).
a. Parietal tooth minute, not connected with columellar lip by a
raised callus ; no internal lamina. paludosa.
aa. Parietal tooth connected with a raised parietal callus.
b. Internal lamina present; upper surface strongly ribbed.
c. Size large ; whorls 7-10. cereolus.
POLYGYRA.
73
cc. Size small ; whorls 6, the last contracted in its first half,
its last half notably swollen. carp enter iana.
bb. No internal lamina.
c. Upper surface coarsely ribbed.
d. Size large, whorls 7 or more. septemvolva.
dd. Size smaller, whorls 5-7, volvoxis.
cc. Upper surface very finely striated. microdonta.
P. cereolus Miihlf.,iii, 128.
laminifera W. G. B.
/. carpenteriana Bid.
microdonta W. G. B., olim.
/. septemvolva Say.
planorbula Lam.
polygyrata " Binn." Pfr.
/. volvoxis Pfr.
febigeri Bid.
/. floridana Hemph.
Var. microdonta Desh.
delitescens Shutt., undesc.
cheilodon Say, Bid.
f plana Dkr.
P. paludosa Pfr. iii, 129.
lingulata Fer., Dh.
ramonis d'Orb.
ramondi d'Orb., Atlas.
insularum Beck, undesc.
; bardenflehtii B., Villa.
Species with teeth on the outer lip.
With the exception of P. johannis of Cuba, the species of this sec-
tion are all continental. The auriculata series inhabits the southern
tier of Gulf States, from Florida to Texas; the dorfeuilliana series
is confined to the more or less mountainous region south of the Ohio
River, from Tennessee to Oklahoma; the texasiana acutedentata
series is from Mexico, extending into Texas along the northern
-continuation of the Sonoran fauna and flora.
P. auriculata Say, iii, 137.
v. microforis Dall, iii, 138.
P. uvulifera Shutt, iii, 137.
florulifera Rve.
P. auriformis Bid., iii, 137.
/ sayii Wood, DeKay.
P. hazardi Bid., iii, 131.
plicataS&y.
f finitima Dh.
P. dorfeuilliana Lea, iii, 133.
v. sampsoni Weth., viii, 152.
P. postelliana Bid., iii, 137.
P. espiloca (Rav.) Bid., iii, 136.
P. avara Say, iii, 136.
P. pustula Fer., iii, 131.
P) pustuloides Bid., iii, 132.
P. leporina Gld., iii, 131.
# #
P. fastigans Say, iii, 131.
fatigiata Say.
fastigiata DeK.
P.jacksoniBld.,iii, 134.
v. deltoidea Simp., viii, 152.
P. troostiana Lea, iii, 131.
<
Of THg
74 POLYGYRA.
P. implicata (Beck) Mart, iii, P. ventrosula Pfr., iii, 136.
[133. v. hindsii Pfr., iii, 136.
P. oppilata Morel, iii, 133. 'P. texasiana Moric., iii, 135.
P. dysoni Shuttl., iii, 132. tamaulipasensis Lea,
dorfeuilliana Pfr. not Lea. tridonia Beck.
P. chiapensis Pfr., iii, 138. P. triodontoides Bid., iii, 135.
P. mooreana W. G. B., iii, 135. P. behri Gabb., iii, 134.
v. tholus W. G. B., iii, J35. P. ariadnie Pfr., iii, 132.
P. yucatanea Morel., iii, 146. couchiana Lea.
v. helictomphala Pfr., iii, 130. P. acutedentata W. G. B., iii,
P. plagioglossa Pfr., iii, 133. loisa W. G. B., iii, 134. [134.
P. dissecta v. Mart., viii, 151. q-uinquedentata F. & C.
P. couloni Shuttl., iii, 134. v. unguifera Mouss., iii, 132.
P. bicruris Pfr., iii, 136. P. anilis Gabb., iii, 130.
P. richardsoni v. Mart., viii, 151. P. bippocrepis Pfr., iii, 134.
* # *
P. johannis Poey, iii, 130. notata Poey.
Section Triodopsis Rafinesque.
Triodopsis plus Mesodon of authors.
Shell varying from depressed to globose-conoidal, timbilicate or
imperforate ; surface generally striated ; whorls 5-6, the last wider,
more or less deflexed in front. Aperture lunate, typically obstructed
by three teeth, two on the lip, one on the parietal wall ; but any or
all of the teeth often absent. Type P. tridentata Say, pi. 30, fig. 8
(see also pi. 30, figs. 9, 10, P. appressa; pi. 30, figs. 13, 14, P. albo-
labris var. maritima; pi. 30, figs. 17, 18, P. sayi).
Jaw sculptured with numerous moderately spaced ribs (pi. 30, fig.
19, P. sayi; pi. 30, fig. 21, P. kiawaensis-, pi. 21, fig. 11, P.
albolabris).
Radula having (1) ectocones with cutting-points developed on
central lateral and marginal teeth, as in P. tridentata, pi. 30, fig. 11,
and P. albolabris, pi. 30, fig. 16, or (2) no side cusps or cutting-
points whatever on any of the teeth, as in P. clausa, pi. 30, fig. 15.
Genital system as described for the genus (pi. 30, fig. 12, P.
inflecta; pi. 30, fig. 20, P. clausa ; pi. 21, figs. 12-16, P. albolabris').
In P. albolabris Say (pi. 21, figs. 12-1 6) the lower third of the penis-
POLYGYRA. 75
(the portion everted during copulation) is smooth inside (fig. 15) ;
it extends upward in a sort of sheath over the base of the upper
portion (figs. 12, 15). This sheath is what Leidy and Biuney call
the " prepuce." The upper portion has fleshy walls which are
densely corrugated or subgranulated within, and the cavity is almost
filled by a thick longitudinal corrugated column,adnate throughout
its length to one side (fig. 15, penis slit open longitudinally ; fig. 13,
14, transverse sections of penis with fleshy column). At the apex of
the cavity there is a perforated papilla (pi. 21, fig. 13, transverse
section), free at its lower end. The retractor muscle is inserted on
the vas deferens a short distance above the apex of penis ; its distal
end being attached to the floor of the lung cavity. The lower
part of the spermatheca duct (pi. 21, fig. 15) is swollen, with fleshy
walls which inside are strongly corrugated lengthwise (pi. 21, fig.
16, transverse section).
Distribution : Eastern North America from Canada to Florida,
west to central Texas and Dakota; in the northwest occurring in
Idaho, and on the Pacific slope from Sitka to Santa Cruz, California.
Most of the species live around decaying logs or under and upon
decaying leaves in forests. Some, like multilineata occur in great
numbers on the low, weedy, willow covered flood-plains of rivers;
others, like profunda, prefer shady, leaf-carpeted and rocky hill-
sides. P. dentifera and P. palliata are found under the loosened
bark of hemlock boles, sharing these retreats with Philomycus.
Most species come from their hiding-places in the warm days of
early spring, and during rainy weather in summer. They may
then be found crawling upon the dead leaves, or ascending nettles,
etc., the leaves of which they eat. In sunny days after rain, they
are found adhering to the lower surfaces of nettle leaves. They
never ascend trees.
The species enumerated below have been divided by authors into
two sections, Triodopsis and Mesodon ; but such division seems to be
artificial. Some species of Triodopsis. &YZ known to have varieties
lacking lip-teeth, and these would technically fall into Mesodon.
In other cases, such as the group of Ida.no and Washington species,
all the transitions from tridentate to toothless apertures occur. The
group of P. appressa is also a transition group. Tryon has resusi-
tated the section-names Xolotrema and Ulostoma. The first of these
is a Rafinesquian name totally unidentifiable ; the second was pro-
posed by Albers for species of Polygyra s. s. and Triodopsis s. s., and
76
POLYQYRA.
did not include either of the forms Tryon uses the name for ! Aplo-
don, Raf., has also been used in this connection ; it is positively uni-
dentifiable.
Species.
v. obsoleta Pils.
HP. hopetonensis Shutt. iii, 144.
ephabus Say, ms.
P. vaunostrandi Bid., iii, 145.
P. vultuosa Gld., iii, 144.
143. v. henriettse Maz., iii, 144.
V^copei Weth., iii, 144.
v. cragini Call, iii, 144.
P. edentata Samp., viii, 154.
edentula W. G. B.
. tridentata Say, iii, 143.
lunula Raf.
v. juxtidens Pils.
^v. edentilabris Pils.
P. fraudulenta Pils.
fallax auct., not Say, iii,
-'P. fallax Say.
^introferens Bid. iii, 145.
"P. rugeli Shuttlw.,iii, 147.
i/P. inflecta Say, iii, 146.
P. mullani Bid., iii, 145. P. columbiana Lea, iii, 154. t-
t^v. hemphilli W. G. B., iii, 146. v. labiosa Gld.
binominata Tryon, iii, 146. v. armigera Anc., viii, 155.
olneyce Pils. P. roperi Pils., viii, 154.
v. blandi Hemph. ^P. loricata Gld., iii, 145.
v. harfordiana .W. G. B., iii, lecontii Lea.
commutanda Anc. [146. P. levettei Bid., iii, 143.
salmonensis Tryon, iii, 146. thomsoniana Anc.
v. oregonensis Hemph. orobcena Anc.
P. devia Gld., iii, 154.
baskervillei Pfr.
P. profunda Say, iii, 155.
richardi Fer.
/ bulbina Dh.
P. sayii Binn., iii, 155.
diodonta Say, not Miihlf.
v. chilhoweensis Lewis, iii,
[155.
*
P. albolabris Say, iii, 150.
rufa DeK.
v. maritima Pils.
P. kiowaensis Simp., viii, 155.
v. arkansaensis Pils., viii, 156.
P. townsendiana Lea, iv, 72.
pedestris and ruida, Gld.
v. ptychophora A. D. Br., iii,
[154.
f. castanea Hemph.
* *
v. traversensis Leach.
t^^T. major Binn., iii, 150.
POLYGYRA.
77
P. andrewsi W. G. B., iii, 150.
i P. exoleta Binn., iii. 151.
^zaleta Binn., olim.
A-P; multilineata Say, iii, 150.
A-F. appressa Say, iii, 148.
linguifera Lam.
v. peri^rapta Pils.
P. sargentiana J. & P., viii, 153.
sargenti J. & P., not Bid.
P. subpalliata Pils.
#
t^P. elevata Say, iii, 148.
tennesseensis Lea.
knoxvilliana Fer.
P. divesta Gld., iii, 152.
dejecta and abjecta Gld.
"P. wetherbyi Bid., iii, 152.
^P. roemeri Pfr., iii, 152.
P. dentifera Binn., iii, 152.
* *
i-P; obstricta Say, iii, 148.
helicoides Lea.
v. carolinensis Lea, viii, 153.
,^FC palliata Say, iii, 147.
denotata Fer.
notata Dh.
P. clarki Lea, iii, 149.
**F~. pennsylvanica Green, iii, 151,
/ P. thyroides Say, iii, 152.
thyroidus Say.
L&-. bucculenta Gld., iii, 153.
/P. clausa Say, iii, 153.
ingallsiana Shutt.
jugallsiana Alb.
*-?. wheatleyi Bid., iii, 151.
^P. christyi Bid., iii, 151.
/-P. mitchelliana Lea, iii, 151.
P. downieana Bid., iii, 153.
P. lawi Lewis, iii, 153.
P. mobiliana Lea.
/<P. jejuna Say, iii, 153.
Section Stenotrema Rafinesque.
Shell small, compact, imperforate or umbilicate ; subglobose,
globose-depressed or lens-shaped the periphery varying from rounded
to acutely keeled ; surface dull, smooth, generally hairy. Whorls 5-
6, closely revolving, the last suddenly deflexed in front. Aperture
basal, narrow, obstructed by an oblique blade-like parietal tooth
parallel to the reflexed basal Up, the latter often notched in the
middle. Last whorl generally having in its last fourth a short
transverse partition on the axis. Type P. stenotrema Fer. (see pi.
31, figs. 22, 23, 24, P. monodon var. alicice').
Animal externally as in Triodopsis.
Genital system having the penis notably longer than the recepta-
culum seminis and its duct, the latter being quite short (pi. 31, fig.
27, P. spinosa).
78 POLYGYRA-POLYGYRELLA.
Jaw having the ribs wide and rather more crowded than is usual
in the other sections of the genus (pi. 31, fig. 25. P. monodon).
Radula having ectocones developed on all the teeth ; basal plates
short and square, slightly shorter than the mesocones of central and
lateral teeth ; marginals with the basal plates short, wide, mesocone
bluntly bifid at tip, ectocone simple or bluntly bifid (pi. 31, fig. 26,
P. hirsuta).
Distribution: Entire Gulf and Atlantic drainages, north to
Canada and south to southern Texas (San Antonio) ; Oregon'. The
species live under and around decaying logs and bits of wood.
This is a well-marked section, distinguished by the compact nar-
row-mouthed shell as well as the crowded, wide ribs of the jaw. The
hairs of the shell collect a coat of earth, which renders the snails
difficult to see, the dusky shade of the animal also assimilating their
color to the surrounding earth or rotten wood. P. monodon ranges
over nearly all of eastern North America ; P. hirsuta has almost as
wide a distribution ; but the other species are rarer and more local ;
P. maxillata, bar big era, edvardsi, edgariana, labrosa and spinosa
being confined to certain localities in the Cumberland system of
mountains. P. germana is ( found in Oregon, but it may prove
related to the mullani group of Triodopsis, rather than to Stenotrema.
Species of Stenotrema.
HP. spinosa Lea, iii, 141. P. hirsuta Say, iii, 140.
P. labrosa Bid., iii, 141. f porcina Say.
P. edgariana Lea, iii, 141. v. altispira Pils.
P. edvardsi Bid., iii, 141. P. maxillata Old., iii, 141.
P. barbigera Redf., iii, 142. ^ P. monodon Rack., iii, 142.
-P. Stenotrema Fer., iii, 140. v. fraterna Say, iii, 142.
hirsuta var. a. Fer. v. alicise Pils., viii, 152.
convexa Raf. v. cincta Lewis, viii, 152.
v. subglobosa Pils., viii, 152. P. leai Ward, iii, 142.
P. germana Old., iii, 143.
Genus POLYGYRELLA Binney, 1863.
=Polygyrella Binn. & J$ld.-t-Ammonitella Cooper.
Shell discoidal, openly umbilicated, the spire slightly convex, flat,
or concave; texture glassy, somewhat translucent. Aperture sub-
POLYGYRELLA. 79
triangular or crescentic, the lip not in the least expanded, blunt,
thickened within the edge by a white rim, simple or two-toothed ;
parietal wall smooth or with an erect tooth. Type P. polygyrella,
pi. 31, figs. 28, 29, 30.
External anatomy unknown. Genital system (in P. polygyrella,
the only species yet investigated) without accessory organs, like that
ofPolygyra (pi. 31, fig. 31).
Jaw low and wide, with no median projection, having numerous
strong vertical ribs, denticulating its margins (pi. 31, fig. 32).
Central teeth tricuspid, laterals bicuspid, marginal teeth bicuspid,
the ectocone simple or bluntly bifid (pi. 31, fig. 41).
Distribution : California, Washington, Idaho and Montana.
The anatomy of the species of this genus is, as far as it is known,
the same as in Polygyra except that the jaw is wider with more ribs.
The shell differs from Polygyra in its absolutely unexpended blunt
lip and its glassy texture. It may be distinguished from the Palse-
arctic genus Gonostoma by the characters of the shell just mentioned
(Gonostoma having an opaque shell with expanded or reflexed lip),
and by the simplicity of the generative system. The relationship of
Polygyrella to Polygyratia cannot be decided until the anatomy of
the South American genus is made known.
[Note. Mr. Binney's classified Synopsis of North American land
shells, in which the name Polygyrella first appeared, is referred to
as " a mere proof" by Professor Joseph Henry, Secretary of the
Smithsonian Institution, who adds that it " should not be quoted as
authority or referred to as a published work." This suggestion
cannot be followed. The Synopsis is not in any ordinary sense a
proof-sheet. A. large edition of it was printed and widely circulated,
as an official publication of the Smithsonian Institution.]
Subgenus POLYGYRELLA Binney, 1863.
Polygyrella BINNEY, Smithsonian Miscellaneous Collections, no.
000, p. 5, Dec. 9, 1863 (no description; type H. polygyrella).
Polygyrella Bland, BINNEY & BLD., in Land and Fresh-Water
Shells of North America, p. 112, type H. polygyrella Bid. & Coop.
W. G. BINNEY, Terr. Moll, v, p. 289 (jaw and dentition), and
Second Supplement to the same, p. 36 (genitalia). PILSBRY, Proc.
80 POLYGYRELLA.
Acad. Nat. Sci. Phila. 1890, p. BQO.Adelodonta ANCEY, Le
Naturaliste, Dec., 1880, p. 334, type H. polygyrella.
Shell disk-shaped, the spire flat or nearly so, periphery rounded,
even in the young ; umbilicus wide within, showing all the whorls ;
texture somewhat glassy and subtranslucent, polished beneath ; color
yellow, greenish or light brown ; whorls 6-8, narrow, slowly widen-
ing, the last slightly descending in front. Aperture subtnangular,
oblique ; peristome blunt, not expanded, thickened within, with or
without lip-teeth or internal teeth ; parietal wall bearing an erect
triangular tooth. Type P. polygyrella, pi. 31, figs. 28, 29, 30, (see
also pi. 31, figs. 33, 34, 35, P. harfordiana, enlarged).
Jaw very wide, arcuate, without median projection below ; surface
with numerous (17-36) broad, slightly separated ribs, denticulating
either margin (pi. 31, fig. 32, P. polygyrella).
Radula long and narrow, with teeth according to the formula 22.
5.1.5.22. Teeth as in Polygyra, the centrals tricuspid the mesocone
longer than the basal-plate ; laterals bicuspid, marginals bicuspid,
the ectocone bifid on the outer teeth (pi. 31, fig. 41, P. polygyrella*).
Genital system like that of Polygyra, but having the duct of the
spermatheca rather longer (pi. 31, fig. 31, P. polygyrella).
This group agrees with Polygyra in essential features of dentition,
jaw and genitalia; it differs from that group in the glassy texture
of the shell and its totally unreflexed lip. The texture of the shell
is like Systrophia, but that South American type has the lip-edge
slightly expanded.
P. polygyrella Bid. & Coop., iii, 129. Coeur d'Alene Mts., Idaho.
P. harfordiana Coop, iii, 130. Fresno Co., California.
Subgenus AMMONITELLA Cooper, 1869.
Ammonitella J. G. COOP., Amer. Journ. of Conch, iv, p. 209.
(Issued February 4, 1869). Gonostoma W. G. BINNEY, Terr. Moll,
v, p. 261.
Shell Ammonite shaped, with sunken, concave spire, and open
umbilicus showing all the whorls; periphery broadly rounded;
texture glassy, subtranslucent ; whorls about 6, very narrow and very
closely revolving, the last whorl embracing the greater part of the pre-
ceding, deflexed in front, its suture somewhat tangentially produced.
POLYGYRELLA-POLYGYRATIA. 81
Aperture narrowly crescentic; lip blunt, thickened within except
toward the upper termination, not in the least expanded, toothless;
parietal wall toothless. Type P. yatesi Coop., pi. 20, figs. 32, 33,
34.
External characters and genitalia of animal unknown.
Jaw low, wide, slightly arcuate, without median projection below ;
surface with a strong transverse line of reinforcement, and about 12
wide crowded ribs, denticulating either margin (pi. 36, fig. 12, P.
yatesi).
Radula long and narrow; teeth after the formula 18.6.1.6.18.
Teeth like those of Polygyrella, but ectocone of marginals simple
(pi. 36, fig. 11, P. yatesi).
This group has been united with the European genus Gonostoma,
but erroneously. It is readily distinguished from that type by the
non-expanded lip and glassy texture of the shell. The dentition also
differs widely. The genital organs of Gonostoma present character-
istic features, but as the soft anatomy of Ammonitella is unknown,
no comparison can now be made. The American species will be
found to have the genitalia simple, as in Polygyrella, if my estimate
of its affinities proves to be correct.
P. yatesi Coop., iii, 115. Calaveras Co., California.
yatesiana W. G. B., olim.
Genus POLYGYRATIA Gray, 1847.
Polygyratia GRAY, Proc. Zool. Soc. Lond., 1847, p. 173, type H.
polygyrata. Ophiogyra ALBERS, Die Hel. 1850, p. 91, type H.
polygyrata Born. Systrophia PFR., Malak. Blatter, ii, 1855, p. 136,
for H. helicyc hides, systrophia, heligmoidea. Entodina ANCEY, Con-
chologists' Exch., i, p. 64, May, 1887, type H. reyrei.
Shell disk-shaped, flat or nearly so above, concave beneath, com-
posed of 5-10 narrow, closely coiled whorls, equally visible above and
below, the last descending in front. Aperture oblique, rounded or
subtriangular, the lip generally narrowly expanded, sometimes
toothed ; parietal callus inconspicuous or raised into a tooth-like pro-
cess. Type H. polygyrata Born, pi. 20, figs. 37, 38.
Animal unknown. The species are said to live in forests. The
typical subgenus is confined to the central portions of South Amer-
ica. The affinities of the genus are problematical. It may per-
haps prove to be allied to Polygyrella.
6
82 POLYGYRATIA.
A number of forms agreeing with Polygyratia in general characters
of the shell are found in Papua and New Ireland. Whether they
have actual affinity to the South American types can be decided
only by an examination of the soft parts. The excessively peculiar
shell argues great antiquity for the group ; and the somewhat similar
distribution of Marsupials and Struthious birds suggests the theory
of an ancient migration in the case of Polygyratia. Such a theory,
however, rests on no known facts of palaeontology or anatomy.
Subdivisions.
Subgenus I. POLYGYRATIA Gray.
Shell having the whorls rounded at the periphery, the spire flat
or concave. South A merican. Three sections, showing slight differ-
ences have been named :
Section Polygyratia. Last whorl provided with an internal barrier
of short spiral lamellae ; parietal callus thin, appressed.
Section Systrophia. Last whorl without internal laminae ; parietal
callus thin, appressed.
Section Entodina. Last whorl without internal laminae ; edge of
parietal callus raised, connecting the ends of the lip, and forming a
sort of parietal tooth.
Subgenus II. COXIA Ancey.
Shell having the whorls flat above, acutely keeled at the shoulder ;
spire subconcave, flat, or conical. Papuan region.
Subgenus I. POLYGYRATIA Gray.
Section Polygyratia Gray (restricted).
Shell solid, typically with opaque dark coloring ; lip expanded,
its margin toothless; parietal callus thin, appressed, body-whorl
having an internal barrier of short spiral lamellae, on both outer and
parietal walls. Type P. polygyrata, pi. 20, figs. 37, 38.
The internal lamellae are like those of Gorilla.
P. polygyrata Born, iii, 124. P. quinquelirata Sm., viii, 150.
charybdis Morch. P. pollodonta d'Orb., iii, 126.
POLYGYRATIA. 83
Section SystropUa Pfr., 1855.
Shell yellowish or corneous, thin, the lip slightly expanded, often
having one or two teeth ; parietal callus slight, not elevated nor
toothed ; no internal lamellae. Type P. helicycloides d'Orb. (see pi.
20, figs. 41, 42, 43, P. stenogyra).
Distribution : Brazil, Bolivia, Equador, Peru.
P. calculina Pfr., iii, 125. P. pseudoplanorbis Lub., iii, 126.
calculus Pfr. not Lowe. P. stenogyra Pfr., iii, 124.
P. decagyra Phil., iii, 125. P. stenostrepta Pfr.
P. gyrella Morel., iii, 126. P. systropha Alb., iii, 127.
P. helicycloides d'Orb., viii, 150. P. tortilis Morel., iii, 125.
P. ortoni Crosse, iii, 127. P. wallisiana Mouss., iii, 126.
P. polycycla Morel., iii, 125.
Section Entodina Ancey.
Shell planorboid, many whorled ; lip narrowly expanded, tooth-
less or toothed, its ends connected across the parietal wall by an
elevated, toothed callus. Type P. reyrei Souv., pi. 20, figs. 39, 40.
Distribution same as Systrophia.
The parietal callus is shaped somewhat like that of Polygyra
cereolus.
P. cheilostropha d'Orb., iii, 128. P. janeirensis Pfr., viii, 150.
P. entodonta Pfr., iii, 126. P. platygyra Alb.
P. heligmoidea d'Orb., iii, 125. P. reyrei Souv., iii, 127.
Subgenus? II. COXIA Ancey, 1887.
Coxia ANC., Conchologists' Exchange, i, p. 75, June, 1887. Type
Helix macgregori Cox. Calostropha ANC., Conch. Exch. ii, p. 38,
Sept., 1887. Type Helix raffrayi T.-C.
Shell many (about 10) whorled, the volutions acutely carinated at
periphery or shoulder, equally visible above and below. Spire either
flat, slightly concave, or conoidal. Aperture oblique, subquad-
rangular, the lip expanded and slightly thickened, its ends connected
by a parietal callus. Type P. macgregori, pi. 20, figs. 44, 45, 46.
Soft parts of animal unknown.
The shells in this group differ from those of the South American
many-whorled Helices in the flat upper surface of each whorl, and
its acute peripheral keel.
84 POLYGYRATIA-PLEURODONTE.
Helix multispirata Hombr. & Jacq. (Manual, iii, 127) and H.
microphis Crosse, have been referred to Polygyratia by authors. The
first is probably a Charopa. The other has been made the type of
a group Microphyura by Ancey (Bull. Soc. Mai. Fr. v, 375). It
belongs to the genus Diplomphalus in Khytididse (Manual i, p. 114).
Species.
P. macgregori Cox, iii, 127. New Ireland.
P. raffrayi Tap.-Can., iii, 128. Western New Guinea.
# # *
The series of genera next to be considered comprises a majority
of the large, solid-shelled Helices of the tropics and southern hemi-
sphere, both in the Old World and America. All discussion of this
and other primary divisions of the Helices is reserved for tie intro-
ductory portion of this volume, but certain brief notes may be of
use in this place. These genera share certain peculiarities of the
generative system : the female branch is without dart sack or other acces-
sory organs ; the male side has the penis continued beyond its papilla-
bearing apex in a narrow tube called the "epiphallus" which terminates
in a flagellum and vas deferens. In most forms the epiphallus is as
long or longer than the penis itself ; but in some (such as Thelido-
mus') it is so shortened as to be inconspicuous, or even absent. In
Cristigibba this process of shortening has resulted in the com-
plete degeneration of both epiphallus and flagellum. In these and
similar cases we must not mistake the structure resulting from de-
generation for a primitive condition. Such an error would be like
holding Ancylus to be a primitive gastropod on account of its (at
present) non-spiral shell, or like grouping the limbless lizards, An-
guis or Amphisbcena with the snakes.
In the American forms the penis retractor is inserted upon the
penis ; in the Asiatic and Australian it is usually upon the epiphal-
lus.
The jaw is generally stoutly ribbed, but often by degeneration of
the ribs, smooth ; and this modification is certainly in some cases
not a generic or even subgeneric character.
Genus PLEURODONTE Fischer de Waldheim, 1808.
=Pleurodonte -j- Lucerna-{- Deniellaria-}- Caraeolus -\-Isomeria-\-
Labyrinthus -\- Polydontes -\- Thelidomus -j- Liochila + Eurycrat era,
etc., of authors.
PLEURODONTE. 85
Shell imperforate or umbilicate, rather large and solid, varying
from globose-depressed to lens-shaped, the periphery rounded or
keeled ; surface striate or granular. Whorls 4 to 6. Aperture with or
without teeth, the lip more or less expanded or reflexed. Eggs
rather large, oval, hard-shelled, the newly hatched young having ar
shell of 2 to 2} whorls. Type P. sinuata Mull. (See pi. 22, figs. 1
to 10 ; pi. 25, all figs except fig. 9.)
Animal having the sole undivided ; lateral edges of foot with no
traces of foot margin; tail rounded, convex above; sides of foot
with granules arranged in oblique rows or irregular ; back with some
indistinct longitudinal lines or none; mantle-edge generally having
small body lappets.
Jaw solid, arcuate, with blunt ends, and either smooth with a
slight median projection, weakly ribbed, or with strong rounded
ribs on its median moity (plates 21, 24, 26).
Teeth of radula in nearly straight transverse rows; central and
lateral teeth unicuspid, the lateral expansions of the cutting point
occupying the place of ectocones, or having side cutting points
developed. Marginal teeth either unicuspid or having a bifid meso-
cone and a simple or bifid ectocone (plates 21, 24, 26).
Genitalia: Penis large, muscular, having the retractor and
epiphallus inserted at its apex; interior with many longitudinal folds
and usually a papilla ; sometimes provided with a short appendix.
Epiphallus varying from long to very short, ending in a short
ftagellum. Female system lacking all accessory organs.
Distribution, West Indies and northern South America. All of
the species are ground snails.
The essential features of this genus are anatomical : (1) the in-
sertion of the retractor on the penis itself; (2) the continuation of
the penis in an epiphallus, into which the vas deferens enters, and
which terminates in a flagellum ; (3) the entire simplicity of the fe-
male system as in Pyramidula or Polygyra ; (4) the rather large,
hard shelled eggs; (5) the tendency of the teeth to develop meso-
cones at the expense of ectocones.
The jaw varies from the ribbed (odontognathous) to the smooth
(oxygnathous) type. The shell exhibits a wide range of variation in
the several sectional groups; but notwithstanding the eonsiderable
variations of both shell and soft parts, the genus is a well character-
ized one, the forms being unquestionably of common ancestry, al-
86 PLEURODONTE.
lied by a strong bond of affinity, and well distinguished from all
other recent genera.
The genus stands isolated in the New World fauna, its relation-
ships being decidedly with the groups of China, the East Indies,
Papua and Australia. Its advent in Middle America is one of the
most interesting problems in Helix distribution. The solution of
this mystery is not lightened by the known distribution of Glandina,
Clausilia, etc., in both the Old and the New Worlds, for no shells
in the least allied to this genus of large Helices have been found in
European tertiary strata.
A thorough study of the nomenclature of this group reveals the
necessity of several unwelcome but apparently inevitable changes.
The well known generic name Caracolus, must be replaced by Pleu-
rodonte, which was proposed and defined in a perfectly proper and
regular manner by Fischer de Waldheim. It is impossible to use
the anonymous, undefined name Lucerna, of Humphrey's sale cat-
alogue Museum Calonnianum, unless we disregard the universally
recognized canons of nomenclature.
Subdivisions.
Pleurodonte may be divided primarily into two subgenera, each
of which is split into several minor groups or sections. The latter are
practically impossible to recognizably define in words, although not
difficult to learn by sight. It will readily be understood, therefore,
that no great importance attaches to these lesser groups. They are
the natural result of late geological changes in the West Indiesr
which broke the parent stock into island colonies. The whole
series tells clearly of a former period of greater elevation of the
Antillean region, and closer connection with the middle American
mainland. The fact that all of the main modifications are found
upon the greater Antilles would lead us to believe that here the group
first became established ; that the Caribbees were peopled from the
northwest, and the mainland of South America also from the north ;
and that the sections grouped below under subgenus Polydontes are
a later modification of the stock, which took place subsequent to the
migration to the southward. The full understanding of the distri-
bution of these Helices, awaits the explanation by geologists of the
main orographic changes of the West Indies during tertiary time
an inquiry beset with difficulty, and as yet but little understood.
PLEURODONTE. 87
Subgenus PLEURODONTE Fischer.
Shell generally solid, dark, depressed and opaque, the aperture
generally toothed. Genitalia characterized by the long epiphallus,
and lack of appendix on the penis.
^Section 1, Pleurodonte (sensu stricto). Shell granulate, at-least
above ; imperforate ; aperture with compressed teeth on the basal lip
only, or if not toothed the shell is not acutely keeled. Jamaica.
^Section 2, Caprinus Montf. Shell solid, imperforate, with thick-
ened peristome, sometimes armed with nodular teeth. Lesser An-
tilles.
Section 3, Gonostomopsis Pils. SheH thin, hirsute, umbilicate ;
aperture trilobate-lunar, outer and basal lips each with a tooth.
Section 4, Caracolus Montf. Shell large, solid, carinated ; aper-
ture lacking teeth. Cuba, Haiti, Porto Kico.
Section 5, Isomeria Alb. Shell depressed, large, dark, solid, not
acutely keeled ; aperture generally with small teeth. Ecuador,
Columbia.
* Section 6, Labyrinthus Beck. Shell carinated, depressed, with an
entering parietal lamella and two processes on the basal lip.
Subgenus POLYDONTES Montf.
Shell depressed or globose, often light colored or variegated with
many bands, the aperture generally toothless. Genitalia having
the epiphallus very short or obsolete, and often with a swollen ap-
pendix near the base of the penis.
Section 7, Thelidomus Swains. Shell globose-depressed, baso-col-
umellar lip of the peristome wide and plate-like, sometimes toothed ;
aperture otherwise lacking teeth.
* Section 8, Polydontes Montf. Shell large, depressed, carinated >*
aperture toothless or with nodular teeth on the peristome ; lip thick.
11 Section 9, Parthena Alb. Shell capacious, unicolored or multi-
lineate. Aperture large, toothless; lip expanded.
Section 10, Luquillia Crosse. Shell similar, but dark colored,
with conoidal spire.
Section 11, Eurycratera Beck. Shell large, globose, with few
whorls. Aperture very large, toothless.
Section 1, Pleurodonte Fischer de Waldheim.
Pleuredonte F. de W., Tab. Synopt. Zoogn. p. 129 (Moscow,
1808) ; proposed for H. sinuata Gm., lyclinuchus Gm., lucerna Gm.,
88 PLEURODONTE.
incequalis Fiseh., lapicida L., isognomostomos Gin. Pleurodonta
BECK, Index Molluscorum p. 33, and of subsequent authors. Den-
tellaria SCHUMACHER, Essai d'un Nouv. Syst. des Hab. des Vers
Test., p. 69, 230, proposed for D. globularis Sebum, (undescribed
and unfigured) and D. sinuata Mull. (1817). Lucerna "Humph."
SWAINS., Malacology, p. 329 (in part). Man. of Conch, v. p. 97.
Not Lucerna Humphrey, Museum Callonianum p. 61, 1797.
Shell imperforate or umbilicated, solid, opaque, varying from sub-
globose to lens-shaped ; surface densely granulated, at least above.
Whorls 4-6, the last deflexed in front. Aperture wider than
high ; peristome broadly expanded, toothless or having from one to
five teeth upon the basal lip; parietal wall calloused but without
teeth. Type P. sinuata Mull. (See pi. 25, figs. 6, 7, P. sloaneana
var. vendryesi; pi. 25, fig. 8. P. acuta var nobilis.)
Animal having the sole undivided, foot edge with no trace of
border ; tail rounded behind ; back with a few indistinct grooves from
mantle to head, the sides irregularly granulated.
Genital organs as in Caprinus. Penis stout, cylindrical, the re-
tractor muscle and epiphallus inserted at its apex ; epiphallus long,
flagellum short. Spermatheca oval, situated on a long duct (pi.
24, fig. 5, P. invalida ; pi. 24, fig. 6, P. acuta).
Jaw arcuate, solid, having unequal, strong, rounded ribs denticu-
lating both margins, the ends blunt and free from ribs (pi. 24, fig.
4, P. acuta'}.
Dentition as in Caprinus. Central and lateral teeth having the
mesocones large and long, expanded laterally. Marginal teeth
having an oblique cusp, formed by the united ento-, meso- and ecto-
cones, which are indicated by slight notches (pi. 24, fig. 7, P. acuta').
Distribution, Jamaica.
Pleurodonte is allied to Caprinus in characters of dentition and
genitalia, the anatomical features of the two groups being practi-
cally alike. The shell differs from that of Caprinus somewhat in
the arrangement of the teeth, which in Pleurodonte are restricted
to the basal lip ; but chiefly by the general fades something diffi-
cult to define, but readily recognized in the shells themselves. The
group is developed with a wonderful exuberance and variety of spe-
cific and subspecific forms, perhaps unparalleled in any tract of like
extent in the world. The anatomy has been investigated by Sem-
per (Reisen), Binney (Ann. N. Y. Acad.), and myself.
PLEURODONTE.
89
This group has hitherto been called Pleurodonta or Lucerna; but
Fischer's Latin form of the word, as well as his French version, was
" Plenrodonte." His name was accompanied by a sufficient diagnosis.
He included several species of the Jamaica group, and also H. lych-
nuchus, lapicida and isognomostomus (=personatoC) ; but as these
three have been made the types of subsequent groups, we obtain by
elimination a residue of several congruous species, of which the first
one of his list has been selected as the type. Dentellaria Schu-
macher was proposed for two species, the first one of which was unde-
scribed and unfigured, but compared with an old illustration prob-
ably representing a small form of H. acuta ; the second being H.
sinuata Mull. Lucerna, proposed anonymously by Humphrey in
the sale catalogue of M. de Callonne's collection, was not defined,
and contains none of the Jamaica group, so far as one may judge
by the fantastic list of species given under impromptu names of the
auctioneer's manufacture. He seems to have included Labyrinthus,
Anostoma and Phania among other shells ; but the work is not
worthy of quotation in scientific literature, and its introduction
therein by the Adams brothers has caused nothing but confusion.
Species.
P. carmelita Fer., v, 99.
mora Gray.
redfieldiana C. B. Ad.
P. bainbridgei Pfr., v, 99.
lamarckii v. unidentata Fer.
v. pretiosa C. B. Ad., v, 100.
v. splengleriana Pfr., v, 100.
P. subacuta Pfr., v, 100.
P. acuta Lam., v, 100.
v. acuta Lam., v, 100.
acutissima Lam.
heteroclites Lam.
v. lamarckii Fer., v, 102.
v. sublucerna Pils., v, 102.
v. patina C. B. Ad., v, 102.
f. goniasmos V A. D. Br., v, 102
f. nannodonta A. D. Br., v, 103
v. oxytenes A. D. Brown, v, 103
v. ingens 0. B. Ad., v, 103.
v. nobilis C. B. Ad., v, 103.
P. abnormis Pfr., v, 104.
P. chemnitziana Pfr., v, 104.
fluduata C. B. Ad.
P. lucerna Mull., v, 105.
v. Julia Fer., v, 105.
v. fuscolabris C. B. Ad., v, 106.
P. rhynchaana A. D. Br., v, 106.
P. peracutissima C. B. Ad., v, 106.
straminea Alb.
martiniana Pfr.
P. cara C. B. Ad., v, 107.
amabilis C. B. Ad.
v. media Ad., v, 107.
P. soror Fer., v, 107.
quadridentata Mke.
P. schroeteriana Pfr., v, 108.
v. chittyana C. B. Ad., v, 108.
90 PLEURODONTE.
P. tridentina Fer., v, 109. P. valida C. B. Ad., v, 113.
sivainsoniana C. B. Ad., v, 109. P. picturata C. B. Ad., v, 113.
v. browneana Pfr., v, 109. sinuata Deless., Chenu, etc.
v. subsloaneana Pils., v, 110. P. pallescens Shuttl., v, 114.
P. okeniana Pfr., v, 110. P. sinuata Mull., v, 114.
fortis C. B. Ad., Rv. v. propenuda Ad., v, 115.
P. atavus Shuttl., v, 110. P. sinuosa Fer., v, 115.
P. sloaneana Shuttl., v, 111. consanguinea C. B. Ad.
bronni v. ft Pfr. v. simson Pfr., v, 116.
schrceteriana Rv. P. invalida C. B. Ad., v, 117.
v. vendryesi Ckll., viii, 263. v. candescens C. B. Ad., v, 117.
P. bronni Pfr., v, 112. P. anomala Pfr., v, 117.
P. strangulata C. B. Ad., v, 112.
Section Caprinus Montfort, 1810.
Caprinus MONTF., Conch. Syst., ii, p. 142, type Caprinus recognitus
Montf. (=H. lychnuchus Miill.). Lucidula SWAINS., Treatise on
Malacol., p. 329, type barbadensis (=isabella Fer.). Lucernella-
SWAINS., t. c., p. 330, type hippocastaneum (=nuxdenticulata). Den-
tellaria BECK, Index Molluscorum p. 34, (1837), and of subsequent
authors. Not Dentellaria Schumacher, Essai, p. 230 !
Shell imperforate, solid, opaque, globose or depressed, the spire
convex or conoidal, periphery rounded or keeled. Surface generally
granulated. Aperture transverse, wider than high, oblique, iheper-
istome thick, expanded, the basal lip widened and generally toothed ;
parietal wall covered with a callus, sometimes toothed. Type
see also (P. lychnuchus P. Isabella Fer., pi. 25, fig. 11 ; pi. 25, fig.
10, P. nuxdentieulata^.
Animal (of P. orbiculata) having the sole undivided ; edges of
foot with no trace of a foot-margin. Entire upper surface rather
evenly granulated, the granules arranged in rather indistinct lon-
gitudinal rows on the back, elsewhere irregularly placed. Mantle
margin without shell-lappets, the right body-lappet well developed,
the left minute, subobsolete.
Jaw arcuate, solid, and either having few strong ribs (pachygastra,
orbiculata, Isabella, dentiens, nucleola, badia, nuxdentieulata), or
without ribs, but having a median projection (formosa, josephince).
P. orbiculata, perplexa and lychnuchus have weak ribs or traces of
ribs, thus connecting the two extremes of jaw structure (pi. 24, fig.
2, P. Josephines ; fig. 3, nuxdentieulata ; fig. 9, orbiculata; fig. 11,
dentiens').
PLEURODONTE. 91
Dentition characterized by the absence of side cusps on central
and lateral teeth, a lateral continuation of the reflexed cutting edge
of the raesocones representing the absent side cutting points. Mar-
ginal teeth having a large, bifid mesocone and a small simple or bifid
ectocone (pi. 24, fig. 8, P. orbiculata} pi. 24, fig. 12. P. dentiens^.
Genitalia without accessory organs on the female side, the duct of
the spermatheca long. Penis having the retractor muscle inserted
at its apex, and continued above in a long epiphallus terminating in
a flagellum (pi. 24, fig. 10, P. orbiculata').
Distribution, Lesser Antilles.
In this group the shell is solid and opaque, as in Caracolus s. str.,
but the basal lip is widened and more or less distinctly toothed. It
is closely allied to the Pleurodonte series, of Jamaica ; and while it
is easy to distinguish the two groups on sight, it is extremely difficult
to point out the differences in words. Anatomically Caprinus and
Pleurodonte are similar.
It is much to be regretted that the well-known name for this sec-
tion had to be rejected ; but it is better to correct the mistakes of
early systematists than to perpetuate them.
P. nuxdenticulata Chemn., v, 82. v. guadeloupensis Pils., v, 87.
punctata Born not Mull. P. lychuuchus Mull., v, 87.
hippocastaneum Lam. recognitits Montf.
P. nucleola Rang, v782. P. josephinse Fer., v, 88.
erassidens Pfr. scabrella Mke.
P. parilis Fer., v, 83. v. nevisensis Pils., v, 89.
pseudoparilis Grat. P. perplexa Fer., v, 89.
P. obesa Beck, v, 83. granifera Gray.
P. dentiens,Fer., v, 84. P. formosa Fer., v, 90.
v. isabellina Pils., v, 85. lenocinia Fer.
P. Isabella Fer., v, 85. P. pachygastra Gray, v, 90.
barbadensis Lm. fuscoviridis Grat.
guildingi Pfr. dolata Fer.
P. orbiculata Fer., v, 86. P. nigrescens Wood, v, 91.
P. badia Fer., v, 86. fuliginea Fer.
Section Gonostomopsis Pilsbry, 1889.
Gonostomopsis PILS., Man. Conch, v, p. 92. Chrysodon ANC.,
Conchol. Exch. i, p. 54, 1887, not Chrysodon Oken, 1815.
92 PLEURODONTE.
Shell narrowly umbilicated, rather thin, opaque, hirsute, the spire
depressed, body- whorl depressed, rounded at periphery. Aperture
as high as wide, trilobate-lunar] peristome narrowly expanded, the
outer and basal margins each with one tooth. Type P. auridens
Rang, pi. 25, figs. 12, 13.
Anatomy unknown. The single species inhabits Martinique. It
resembles in form H. obvoluta Mull, of Europe.
Section Caracolus Montf.
Caracolus MONTF., Conch. Syst. ii, p. 138. PILSBRY, Man. of
Conch, v, p. 113. Caracolla SCHUM., Essai, p. 192, 1817. Serpent-
ulus (KLEIN, Tent. Meth. Ostr., p. 8, 1753 ; in part) H. & A. AD.,
Gen. Rec. Moll, ii, p. 201. Lampadion BOLT, in part. Discodoma
SWAINS., Malacol., p. 329, 1840.
Shell depressed, carinated, imperforate or umbilicate ; thick, solid
and opaque ; spire conical, apex obtuse. Whorls 5-6, the last but
little or not deflexed in front. Aperture oblique, wider than high ;
peristome not toothed, its basal margin expanded or narrowly
reflexed, terminations remote. Type P. caracolla L., pi. 25, fig. 1.
Jaw arcuate, stout, and either smooth with a low median projec-
tion (P. caracolla, P. marginella, P. semiaperta), or furnished with
stout ribs (P. bornii). See pi. 26, fig. 3, P. marginella', pi. 26, fig.
6, P. marginella var. rostrata.
Radula having the central and lateral teeth furnished with a single
broad obtuse cusj? (coalescent meso- and ectocone). Marginal teeth
having an oblique cusp, which is simple as in the lateral teeth, or
split into mesocone and ectocone (pi. 26, fig. 8, P. caracolla; fig. 1,
P. maginella ; fig. 2, P. marginella var. semiaperta}.
Genital organs having the vagina more or less swollen, sperma-
theca oval, on a rather long (P. caracolla) or a short duct (P. mar-
ginella). Penis long, the retractor inserted at its summit; continued
in a long epiphallus which terminates in a short flagellum (pi. 26, fig
7, P. caracolla, penis everted ; pi. 26, figs. 4, 5, P. marginella var.
rostrata viewed from both sides, the extremely short flagellum seen
in fig. 4).
Distribution : Eastern Cuba ( P. marginella and varieties), Hayti
(P. caracolla, excellens, insititia, sarcocheila, angistoma, bizonalis and
semiaperta), and Porto Rico and Vieque (P. bornii).
PLEURODONTE. 93
The strong, opaque, carinated shell, with toothless aperture, uni-
colored or with few, broad bands, is characteristic, as is also the very
long epiphallus and short flagellum, and the blunt, broad cusps of
the teeth. The jaw is either smooth or ribbed, as in Caprinm.
A fuller knowledge of the genitalia is necessary for the final _settle-
ment of specific limits; meantime the following arrangement is
offered.
P. caracolla Linn., v, 120. sagemon Beck.
tornata Born. arangiana Poey.
albilabris Lam. marginatoides d'Orb.
ocnlatus Montf. f fasciata Blv.
P. excellens Pfr., v, 120. f indiscreta Beck.
P. insititia Shutt., v, 121. v. gutierrezi Poey, v, 125.
P. sarcocheila Morch, v, 121. v. schwartziana Pfr., v, 125.
P. angistoma Fer. v. mina Pfr., v, 12,5.
angystoma Dh. , marginata Orb.
anchistoma v. Mart. jactata Gundl.
P. bornii Pfr., v, 127. v. rostrata Pfr., v, 126.
marginella Pfr not Gmel. pazensis Poey.
P. bizonalis Desh., v, 127. cupulata Pfr.
v. gaskoini Pfr. v, 127. v. marginelloides d'Orb., v, 126.
P. marginella Gmel., v, 124. transitoria Pfr.
marginata Born. v. semiaperta v. Mart., v, 125.
Section Isomeria Albers, 1850.
Isomeria ALB., Die Hel., p. 126, type H. oreas Koch. v. MART.,
Die Hel., p. 155. PILSBRY, Manual of Conch, v, p. 135.
Shell depressed, solid, opaque, chestnut or chocolate colored, rounded
or obtusely carinated at the periphery, im perforate or umbilicated.
Spire depressed, convex, with 6 or fewer whorls, the last deflexed or
straight in front. Aperture wider than high, very oblique ; peristome
expanded or reflexed, toothless or with small teeth, of which one is
situated near the termination of the periphery; ends of peristome
remote, joined by a parietal callus, the parietal wall often having
an oblique tooth. Type P. oreas Koch. (pi. 25, figs. 2, 3, P. faunus
var. ritchieana).
Animal unknown.
A group of large and beautiful dark colored helices confined to
the valleys of the higher Andes of Ecuador and Columbia, where
^4 PLEURODONTE.
they replace Labyrinthus of the lower regions of northern South
America. The shells differ from Labyrinthus in the more or less
transversely dilated contour, the swollen base of the latter portion
of the body-whorl, and the less developed aperture-teeth. In a few
species (cenigma, vexans) the teeth are strongly developed ; but these
are to be regarded as an independent line of evolution from typical
Isomeria, rather than as an intermediate or ancestral form between
Isomeria and Labyrinthus.
Species.
P. oreas Koch, v, 136.
procera Pfr.
P. faunus Phil., v, 1 37.
v. ritchieana Pils., v, 138.
P. subelliptica Mouss., v, 139.
P. continua Pfr., v, 137.
P. aloagana Jouss., v, 139.
P. peritropis Pils., v, 140.
P. fordiana Pils., v, 141.
P. calomorpha Jonas, v, 142.
P. sequatoriana Hid., v, 142.
P. scalena v. Mart., v. 143.
P. meobambensis Pfr., v, 144.
P. atrata Pfr., v, 144.
P. mauritii Jouss., v, 145.
atrata Rv. not Pfr.
P. cymatodes Pfr., v, 146.
P. parietidentata Mill., v, 147.
P. kohlbergi Mill., v, 148.
P. martinii Bern., v, 149.
morula Hid.
P. granulatissima Mill., v, 148.
P. gealei E. A. Sm., v, 149.
P. stoltzmanni Lub., v, 150.
P. sequatoria Pfr., v, 150.
P. equestrata Moric., v, 151.
P. triodonta d'Orb., v, 152.
P. juno Pfr., v, 152.
P. neogranadensis Pfr., v, 153.
P. hartwegi Pfr., v, 153.
loxensis Mill.
P. basidens Mouss., v, 154.
P. bituberculata Pfr., v, 154.
bourcieri Rv. not Pfr.
v. tridentula Mill., v, 155.
v. latideutata Mill., v, 156.
P. bourcieri Pfr., v, 156.
bituberculata Rv. not Pfr.
P. subcastanea Pfr., v, 157.
globosa Brod. not Sowb.
P. senigma Dohrn, v, 158.
P. vexans Dohrn, v, 158.
Section Labyrinthus Beck, 1837.
Labyrinthus BECK, Index Moll., p. 33, type L. otis=labyrinthus
Dh. PILSBRY, Manual of Conch. (2), v, p. 159. MARTENS, Biol.
Centr. Amer., Land Moll., p. 175. Lyrostoma SWAINS., Malacol.
p. 329, type L. labyrintha. Lyriostoma SWAINS., I. c., footnote
<1840).
Shell umbilicate, depressed, carinated, microscopically granulated,
not transversely dilated. Whorls less than 6, the last descending in
PLEURODONTE. 95
front, constricted behind the peristome. Aperture transverse, sub-
horizontal, obstructed by three primary folds or teeth, one long
parietal fold, one tooth on the outer, another on the inner portion of
the basal lip ; peristome expanded or reflexed in every part, con-
tinuous across the parietal wall. Type P. labyrinthus, pi. 25, figs. 4,
5. (See also pi. 22, figs. 7, 8, P. sieversi).
Soft parts unknown. Jaw slightly striated (Morch, Journ. de
Conch. 1865, p. 381), with a slight median projection. Teeth all
uni-cuspid (Semper, Reisen, p. 105) as in the Cuban Caracolus.
(PI. 26, fig. 9, P. plicata Born, after Semper).
This group is characteristic of northern South America, extend-
ing from the Amazon River and its western tributaries in eastern
Peru, northward in Central America to Costa Rica. It inhabits less
elevated regions than the allied group Isomeria. Its complicated
aperture-armature has doubtless been evolved for protection against
predaceous insects (c/. Man. of Conch, v, p. 159; Biol. Centr. Amer.
Moll., p. 175 ; Pop. Sc. Monthly, 1892, p. 191).
Labyrinthus agrees with the restricted section Caracolus in teeth
and jaw, as well as in the general features of the shell. It stands in
about the same relation to Caracolus that Triodopsis and the auri-
culate Polygyras hold toward the toothless Mesodons. There seems
no sufficient reason for considering Labyrinthus a distinct genus, as
von Martens has done.
Species.
P. labyrinthus (Chem.) Dli. v, P. leucodon Pfr., v, 167.
subplanata Petit. [161. P. sieversi v. Mart., viii, 263. .
v. erecta Mouss., v, 162. P. quadridentata Brod., v, 168.
v. sipunculata Forbes, v, 162. P. tamsiana Dkr., v, 169.
annulifera Pfr. P. tarapotonensis Moric., v, 170.
P. plicata Born, v, 163. P. bifurcata Desh., v, 170.
hydiana Lea. P. furcillata Hupe, v, 171.
hydeanus v. Mart. P. raimondii Phil., v, 172.
P. uncigera Petit, v, 164. P. yatesi Pfr., v, 173.
conoidea Anc., viii, 264. P. ellipsostoma Pfr., v, 173.
anopla Anc., viii, 264. P. leprieurii Petit, v, 174.
v. creveauxiana Anc., viii, 264. auriculina Petit.
P. triplicata v. Mart., v, 165. P. dunkeri Pfr., v, 174.
cesopus Ang. P. isodon Pfr., v, 175.
96 PLEURODONTE.
P. manueli Higg., v, 166. P. bogotensis Pfr., v, 176.
manoeliPfr. P. otostoma Pfr., v, 176.
manseli Pfr.-Cless. stostoma Rv.
Section Thelidomus Swain son, 1840.
Thelidomus SWAINS., Malacology, p. 191, 192,330, type IT. incerta
Fer. Not Thelidomus Swains., t. c., p. 228, 353,=larva-cases of Heli-
copsyche, (Neuroptera). Otala BECK and others, not of Schumacher.
Pachystoma ALBERS, Die Hel., 1850, p. 125. Not Pachystoma
Guilding, Zool. Journ. 1828, p. 536. .? Thelidonta SWAINS, t. c.,
p. 194.
Shell imperforate, globose-depressed, with few whorls, the last
deflexed in front, swollen beneath, carinated or rounded at the
periphery. Surface granulated, costulate-striate or decussated.
Aperture very oblique; peristome expanded, thickened within, the
lower margin straightened, with a plate-like callus inside. Type P.
incerta Fer. (See pi. 22, fig. 5, P. lima ; pi. 22, fig. 4, P. trinitaria').
Jaw arcuate, having 7-15 strong ribs, sometimes not denticulating
the lower margin (pi. 23, fig. 23, P. auricoma var. havanensis^.
Radula either with or without ectocones on central and inner
lateral teeth. Marginal teeth obtusely and obscurely bicuspid. PI.
23, fig. 22, P. auricoma var. havanensis.
Animal having the sole undivided ; lateral edges without trace of
pedal grooves or margins. Sides of foot granulated, granules
arranged in vertical series in the middle, obliquely descending series
in front and behind ; back irregularly granular, without longitud-
inal grooves.
Genital system having the penis stout, with a flagellum at apex ;
vas deferens and retractor muscle also inserted at apex, the latter
slender, and attached distally to the integument of the vestibule ; a
small appendix sometimes present; no internal papilla, the opening
of the vas deferens being a simple orifice at the base of the flagellum
(pi. 23, fig. 19, showing opened penis and vestibule). Spermatheca
oval or oblong, enveloped in the folds of the uterus, its duct short,
bearing at the base a broad muscle connecting with the integument
of the body-wall near the genital orifice (pi. 23, fig. 21, v. m.) ; ovo-
testis composed of one compact tuft of long creca, (pi. 23, figs. 19
21, P. auricoma var. havanensis ; pi. 23, fig. 24, P. lima ; pi. 23, fig.
25, P. aspera).
PLEUROBONTE. 97
The principal peculiarity of the shell of Thelidomus is the plate-
like baso-columellar lip, somewhat like that of Acavus or Macularia.
The anatomy exhibits considerable variation in some details, such
as the presence (pi. 23, fig. 24, lima) or absence (pi. 23, figs. 20-21,
auricoma v. havanensis*) of an appendix. The spermatheca duct is
much shorter than in Parthend. Many more species must be invest-
igated before a satisfactory account can be given of the peculiarities
of the genitalia of Thelidomus and related groups. See Poey,
Memorias ; W. G. Biimey, Proc. Acad. Nat. Sci. Phila. 1875, and
Ann. N. Y. Acad. ; Semper, Reisen, pi. 15. The eggs are oval,
white and calcareous-shelled ; the embryonic shell is densely gran-
ula^kd in the typical forms, shining and radially grooved in the
Cuban group which I have called Zachrysia. The latter are said to
form a calcareous epiphragm.
(Thelidomus s. sir., species of Jamaica, Porto Rico and Lesser
Antilles}.
P. incerta Fer., v, 57. punctifera Lm.
notabilis Fer. asperula Beck.
curvidens Pfr. v. castrensis Pfr., v, 59.
striolata Guild. P. aspera Fer., v, 59.
alutaceaZgl. granosa Wood.
vehitinoides Anton. P. cognata Fer., v, 59.
ravnii Beck. P. discolor Fer., v, 60.
P. lima Fer., v, 58. P. ? sanctselucise Smith, v, 198.
(Zachrysia ; species of Cuba and Bahamas).
P. petitiana Orb., v, 60. P. emarginata Gundl., v, 64.
P. guanensis Poey, v, 61. P. bayamensis Pfr., v, 64.
P. scabrosa Poey, v, 61. P. guantanamensis Poey, v, 65.
P. auricoma Fer., v, 62. v. proboscidea Pfr., v, 66.
microstoma Lm. P. rangelina Pfr., v, 66.
v. noscibilis Fer., v, 63. P. trinitaria Gundl., v, 67.
v. havanensis Pils. P. baracoensis (Gut.) Poey, v,.
" zeta Pfr." v, 63. lamellicosta Pfr. [67.
v. provisoria Pfr., v, 63.
Section Polydontes Montfort.
Polydontes MONTF., Conch. Syst. ii, p. 154, type P. imperator.
Shell large, depressed, imperforate or narrowly umbilicated, solid
and heavy ; the surface microscopically decussated. Whorls 4^-5,,
98 PLEURODONTE.
the last often carinated, slightly deflexed in front. Aperture
oblique, the peristome thick, expanded, simple or bearing obtuse
teeth, and having an obtuse fold near the eolumellar insertion. Type
P. imperator, pi. 22, fig. 9.
Anatomy unknown. Eggs large, oblong, with a hard calcareous
shell, that of P. imperator (pi. 22, fig. 10) measuring 8i by 12 mill.
Young having when hatched a granulated, umbilicated shell of
about 2 whorls, measuring about one-fifth the diameter of the adult
shell.
The shell in this section is generally marked with many spiral
lines of brown, the widest and most conspicious being immediately
below the periphery. P. apollo is sometimes unicolored. It will be
seen that in coloration, the relationship of Polydontes to Parthena
(P. dominicensis, etc.) is extremely close. All three species of
Polydontes are known to voluntarily amputate their tails when
captured (Journ. de Conchyl. 1860, p. 226). They live under dead
leaves. Distribution, eastern Cuba.
P. imperator Montf., v, 79. P. sobrina Fer., v. 80.
magica Fer. crassilabris Pfr.
P. apollo Pfr., v, 79.
Section Parthena Alberp.
Parthena ALB., Die Hel., p. 112 (first species If. angulata).
Shell imperforate, globose or depressed, the periphery rounded or
carinated; spire short, whorls 4-4, the earlier 1| forming a gran-
ulated or radially grooved embryonic shell, the last notably inflated
and capacious, unicolored or begirt with many brown lines ; surface
granulated. Aperture large ; peristome expanded ; columella
arcuate. Type P. angulata, pi. 22, fig. 2. (See also pi. 22, fig. 3,
P. dominicensis).
Animal (of P. dilatata') having the sole undivided, with no
indication of lateral borders or pedal grooves. Upper surface and
sides coarsely granulated, the granules arranged in descending rows
on the sides, finer and irregular on the back ; tail rounded above ;
back from mantle to face irregularly granulated, lacking longitud-
inal grooves. Mantle-edge lacking shell-lappets ; body-lappets well
developed, the right one short, the left extending the entire length
of the outer lip of the shell (pi. 23, fig. 15, showing posterior angle
of aperture, respiratory opening and lappets).
PLEURODONTE. 99
Jaw strong, arcuate, sculptured with high, rather narrow ribs
crenulating the upper margin only, or both margins (pi. 23, fig. 16,
P. dilatat(t). In P. angulata Binney found 7 ribs on the jaw ; in
vrispata 10 ribs. In P. dilatata we find 9-11 ribs, which crenulate
the upper but not the lower margin, the latter having a -slight
median projection.
Dentition : Central and lateral teeth having stout, longmesocones
projecting beyond the basal-plates ; ectocones represented by a lateral
bulging of the reflection of the cusp, or by distinctly developed cut-
ting-points. Marginal teeth having the mesocones stout, oblique,
blunt or sub-bifid, the ectocone simple, minute (pi. 23, fig. 18, P.
angulata ; pi. 23, fig. 17, P. dilatata').
Genitalia : Female system presenting no accessory organs ; the
spermatheca short, globular, situated upon a long duct. Penis stout,
cylindrical, having a large globular appendix near its base. At its
apex is situated a short, curved, obtuse flagellum, near the base of
which is inserted the vas deferens, and a short teat-like organ which
is solid and fleshy, not perforated or hollow. No retractor muscle
seen. When opened lengthwise the walls of the cavity of the
appendix and of the penis are seen to be longitudinally folded (fig.
14), the folds disappearing in the upper part of the penis-cavity.
The upper part of the cavity is occupied by a large, free, pestle-
shaped penis-papilla, perforated at the end, the perforation leading
to the cavity of the vas deferens and flagellum the latter being
corrugated inside (pi. 23, fig. 13, 14, P. dilatata).
Distribution: Hayti.
P. angulata Fer., v, 69. P. dissita Dh., v, 71.
inflata Dh. P. undulata Fer., v, 72.
acutangula Beck. lineolata Lam.
P. obliterata Fer., v, 69. v. crispata Fer., v, 72.
P. angustata Fer., v, 70. P. dilatata Pfr., v, 73.
P. dominicensis Pfr., v, 70.
extensa Pfr. not Mull.
Section Luquillia Crosse, 1892.
Luquillia CROSSE, Journ. de Conchyl. 1892, p. 19, typeiT. luquil-
lensis. Leiostoma SWAINS. Malacol., p. 328 (preoc.), 1840.
Shell imperforate, solid, subglobose with rather conoidal spire, of
about 5i whorls, the earliest 2i forming the large, coarsely gran-
100 PLEURODONTE.
ulated embryonal shell, the following whorls microscopically decus-
sated ; the last whorl rounded at periphery. Color yellowish-brown
with dark oblique streaks and sometimes a subperipheral girdle.
Aperture wider than high, the thick lip expanded ; columella short,
arcuate, with an obscure callus fold. Type P. luquillensis Shutt.
(See pi. 22, fig. 1, P. gigantea).
Soft anatomy and jaw unknown. Radula (of P. luquillensis) as
in Parthena angulata, q. v.
Distribution, Haiti and Porto Rico.
P. gigantea Scop., v, 73. P. audebardi Pfr., v, 74.
cornumilitare auct. not L. P. luquillensis Shutt., v, 74.
Section Eurycratera (Beck) Gray.
Eurycratera BECK, Index Moll., p. 45, in part. GRAY, P. Z. S.
1847, p. 171, type H. jamaicensis. Lejocheila or Leiocheila ALBERS,
Die Hel., p. 109, lS50.Liochila v. MART., Die Hel., p. 146, 1860,
type H. jamaicensis.
Shell imperforate, globose, solid, the surface finely wrinkled,
embryonal whorls H, large, shining, not granulated. Whorls 4,
the last very large, rounded, having few color bands. Aperture very
large, oval, the outer lip expanded, columella long, arcuate ; parietal
and columellar callus spreading upon the base. Type P. jamaicensis
Gmel., pi. 22, fig. 6.
Jaw thick, arcuate, attenuated toward the ends, the anterior sur-
face sculptured with 14 decided but unequal ribs, irregularly dis-
posed, and denticulating either margin. Lingual membrane with
41.1.41 teeth. Side cusps and cutting points wanting on central and
inner lateral teeth, but represented by an expansion of the reflexed
sides of the mesocones. The single species is confined to Jamaica.
Beck selected no type for Eurycratera, and his list of speciesincludes
forms belonging to many diverse groups. Gray, in 1847 selected
H. j amaicemis as type of the group, and I do not see how we can
avoid following his selection ; especially in view of the fact that v.
Martens, in 1860, selects as type H. dominicensis, a species not
included by Beck in his list, and therefore certainly not the type of
his group.
CAM^NA. 101
Genus CAMJ5NA (Alb.) Pils. & v. Moll.
Camcena ALBERS, Die Heliceen p. 85, 1850, in part. Camena
Alb., v. MARTENS, Die Hel. 2d. edit., p. 165, type cicatricosa Mull,
(restricted to sinistral species of Cam sen a and Euhadra). Camcena
Alb., PILSBRY, Man. of Conch, vi, p. 197, and viii, 265. v. MOLL-
ENDORFF, Nachrichtsbl. d. D. M. Gesellsch. 1891, p. 195. PILSBRY,
/. c., 1892, p. 71 ; Proc. Aoad. Nat. Sci. Phila., 1892, p. 398 (anat-
omy). Eucochlias THEOB. in NevilPs* Handlist Moll. Ind. Mus. pt.
1, p. 81, 1878 (type, ochthoplax; contains also bougainvillei, illustris
sulcocincta smdpyrostoma).
-\-Pseudobba v. Moell. and Phoenicobius Morch.
Shell rather large, varying from depressed-globose or conoidal to
elevated and short pupiform ; dextral or sinistral, solid, yellow or
brown usually encircled by chestnut bands or lines. Surface closely
malleated or wrinkled all over ; whorls about 5-5 z, the upper ones
flattened, the last subglobose or carinated ; peristome expanded or
reflexed, its ends not converging, columellar margin dilated over or
partly over the rather narrow umbilicus. The columella is rounded.
The nuclear shell is rather large, (about one-fifth the diameter of the
shell), consisting of 2 to 2J whorls, its junction with the after-growth
marked by a (generally) distinct line. The young shells are acutely
carinated. Type cicatricosa Mull., pi. 1 9, fig. 8.
Animal having the solemn/ indistinctly tripartite ; lateral edges of
foot with no trace of a foot-margin, sides of foot granulated in irregular
pattern, the tail rather long, rounded above, with an indistinct slightly
impressed longitudinal median line ; anteriorly there are a few indis-
tinct longitudinal grooves from mantle to head. Mantle-margin
with a small triangular right body-lappet, and a longer left one.
Jaw arcuate, strong, typically with numerous strong, separated
ribs (pi. 18, fig. 5).
Dentition : Central and lateral teeth having the mesocones only
developed, the cusps large, cutting-points small (pi. 18, fig. 6). Mar-
ginal teeth with a long, oblique, bifid mesocone united at base with
the ectocone, which becomes bifid on the outer teeth (pi. 18, fig. 7).
Genitalia: Vestibule short; penis stout, continued above in an
epiphallus, in which the retractor and then the vas defer ens is inserted,
and terminating in flagellum ; penis corrugated within, and having
a large papilla at its apex (pi. 18, figs. 2, 4). Vagina stout, bound
102 CAMCENA.
to the body-wall by a band of muscles ; duct of the spermatheca
long (pi. 18, figs. 1 and 3).
Distribution of the typical forms, southern China (provinces
Kwang-Tung and Kui-chu) to Burmah and southward.
The most important features of this genus are found in the genital
system, which is of the type called by the writer epiphallophorous.
The penis is continued upward in a flagellum-like extension (epiph-
allus), in which the vas deferens enters, and which enters the penis
itself through a penis-papilla (pi. 18, fig. 2). Thus far, the structure
is exactly like Caracolus of the West Indies; but Camcena differs
from Caracolus in having the retractor muscle inserted upon the
epiphallus instead at the apex of the true penis. The penis-retractor
is attached to the floor of the lung-sack. The female system lacks
all accessory organs, and the duct of the spermatheca is longer than
in Pleurodonte. The vagina has strong muscular walls, and is bound
to the adjacent right body-wall by a band of muscles (shown in pi.
18, fig. 1) ; this structure occurs also in the West Indian Thelidomus
(see p. 96). The teeth are of the Caracolus type, being character-
ized by the total absence of side cusps on centrals and inner laterals.
The specimens dissected were received from Dr. v. Mollendorff, with-
out the shell. I take them to be C. xanthoderma.
The shell is rather large, capacious, solid, and generally roughly
sculptured. The Japanese sinistral helices (H. qucesita, etc.) have
been associated with Camcena, but they belong to a totally different
phylum of Helix. The columellar lip is rounded in Camcena, not
expanded in a flat plate as in Phania or Acavus.
Perhaps no group of Helices has been less understood by systema-
tists than this. Albers included several very diverse types in his
original list of species. Martens restricted the group to large,
capacious sinistral helices of true Camcena and the very different
group Euhadra. Pfeiffer united the whole Oriental and Australian
series of Euhadra, Camcena and Hadra under the one name
Camcena. The present writer, in 1890, defined the natural
groups of Oriental Helices, and indicated the conchological char-
acters upon which they rest, restricting Camcena to forms having a
large nuclear shell. This work was criticised by v. Moellendorff
(Nachrbl. D. M. G. 1891, p. 195), and several improvements in
classification were suggested. These were in large part adopted in
a later paper by the writer (Nachrbl. 1892, p. 71).
CAM^NA. 103
Subdivisions.
Subgenus CAM^NA (restricted). Shell capacious, narrowly um-
bilicated, depressed-globose, often carinated. Surface malleated or
wrinkled. Last whorl not descending in front. Distribution south-
ern China and Farther India.
Section Phcenicobius Morch. Shell differing from Camsena in
the generally more elevated, conoidal or pupiform shape, and in
having (typically) four dark spiral bands; the surface varying from
smooth to rib-striate, sometimes slightly malleated beneath. Dis-
tribution, Philippine Is.
Subgenus PSEUDOBBA v. Moell. Shell rudely sculptured, with
wrinkles or furrows oblique to the growth-lines ; solid ; umbilicus
large; peristome thickened, the terminations joined by a cord of
callus across the parietal wall. Distribution, Northern Celebes and
Sangir Is.
Subgenus CAM^NELLA Pils. Shell smoothish, depressed sub-
globose, banded and maculated with brown on a white ground ; sur-
face smoothish; whorls about 5J, the last deflexed in front; embry-
onal shell less than one-sixth the diameter of the adult. Columella
with an obtuse tooth. Distribution, Island of Hainan.
Subgenus NEOCEPOLIS Pils. Shell smoothish, globose-conical,
with 6-7 closely revolving whorls, the last deflexed in front. Aperture
having an internal fold within, marked by a pit outside ; the col-
umellar lip obtusely toothed. Distribution, Tonquin.
Subgenus CAM^NA Alb.
The shell is more wrinkled or malleated than in Phcenicobius ;
the last whorl does not descend in front. Of the four principal color
bands of this phylum of Helices, band ii (supraperipheral), or band
iii (subperipheral) is retained, bands i and iv being absent. Some
species show many fine spiral lines of color in addition. The anat-
omy is described above. The jaw is ribbed. The subgenus is Indo-
Chinese in distribution. Many more species will probably be dis-
covered. Type C. cicatricosa, pi. 19, fig. 8.
Species.
C. cicatricosa Mull., vi, 198. v. inflata Mlldff., vi, 199.
senegalensis Fer. v. ducalis Anc., vi, 199.
chinensis Voigt.
104
CAMCENA.
C. longsonensis Mori., viii, 265.
C. jaculata Mab., vi, 120.
C. seraphinica Heude, vi, 199.
C. hahni Mab., vi, 200.
broti d'Ham. & Dautz.
C. subgibbera Mlldff., vi, 200.
C. leonhardti Mlldff., vi, 201.
C. vulpis Gredl., vi, 116.
C. pachychila E. A. Sm., viii, 265.
C. gabriellse Dautz. & d'Ham.,
bathmophora Mab. [vi, 205.
v. subhainanensis Pils., vi, 205.
C. hainanensis H. Ad., vi, 204.
C. xanthoderma Mlldff., vi, 206.
v. polyzona Mlldff., vi, 207.
C. illustris Pfr., vi, 201.
C. ochthoplax Bens., vi, 202.
C. saturnia Gld., vi, 203.
Section PHCENICOBIUS Morch, 1852.
Phoenicobius MORCH, Cat. Yoldi, p. 32, type H. arata. MLLDFF.
'Nachrbl. D. M. Ges. 1891, p. 202. PILSBRY, Man. of Conch, viii,
p. 266.
The shell is like Camcena in the large embryonal portion, consist-
ing of about 2? whorls. It differs from Camcena in being generally
more elevated, sometimes pupiform ; and most species have all of
the four bands (i subsutural, ii supraperipheral, iii subperipheral
and iv umbilical) developed. Type C. arata Sowb., pi. 19, fig. 13 ;
(See also pi. 19, fig. 12, C. monochroa').
The anatomy is unknown. The species are said to inhabit the
Philippine islands Tablas, Mindoro, Luban, Busuanga and Palawan ;
but the localities of some of the pupiform species are not certain.
This group has generally been considered a section of Cochlostyla.
Dr. v. Mollendorff first pointed out the true affinities of the forms.
C. arata Sowb., viii, 267.
v. lutea Pils., viii, 267.
C. brachyodon Sowb., viii, 267.
v. naujanica Hid., viii, 268.
C. adusta Sowb., viii, 268.
C. oblonga Sowb., viii, 269.
lubanicus Pfr.
C. oomorpha Sowb., viii, 269.
C. bintuanensis Hid., vi, 237.
C. campanula Pfr., vi, 236.
C. auacardium Dohrn, vi, 238.
C. ceres Pfr., vi, 239.
C. trailli Pfr., vi, 207.
C. monochroa Sowb., vi, 208.
palawanica Pfr.
saulice Pfr.
lagunce Hid.
dorice Dohrn.
C. palumba Souv., vi, 209.
C. egregia Dh., vi, 210.
C. avus Pfr., vi, 210.
CAMCENA. 105
Subgenus PSEUDOBBA MollendorfF, 1891.
Pseudobba v. MOELL., Nachrbl. D. M. Ges. 1891, p. 202, type H.
mamilla. Obba (typical part) MARTENS, not Gray.
The shell in this group is heavy, solid, rudely sculptured, with a
rather large umbilicus. The subgenus is evidently more nearly
allied to Phcenicobius than to the continental Cam&enas. Type C.
mamilla, pi. 19, fig. 9.
The living animal, as figured by Quoy and Gaimard (Voy. de
1'Astrol. pi. 7), agrees with that of Camcena in external features.
Jaw of C. quoyi horse-shoe shaped, the ends attenuated; cutting edge
with a distinct median projection (pi. 15, fig. 11). Viewed in profile,
the anterior surface is concave (pi. 15, fig. 12). The color is dark
chestnut. Anterior surface smooth ; showing by transmitted light
fine wavy lines parallel with the margins. (Schako, from a half-
grown specimen; Mai. Bl. xx, p. 169).
Central and lateral teeth with the mesocones only developed, as
in Camcena. Marginal teeth with a large, oblique, bifid rnesocone
and an ectocone ; also closely resembling the teeth of Camcena (pi.
15, fig. 13, central, lateral and marginal teeth, and fig. 14 a lateral
of C. quoyi, seen in profile).
The species of this subgenus inhabit northern Celebes and the
Sangir Is. It is a satellite group of the Philippine Island Camsenas,
which has spread southward like a few Obbas and Cochlostylas.
The dentition is the same as in Camcena, but the jaw (o/aw imma-
ture specimen of quoyi) lacks ribs ; so it seems that in this genus, as
in the West Indian Caracolus, the presence or absence of ribs is not a
generic character. From Quoy's remark that the jaw of mamilla
Is not different from that of French Helices, we presume that it is
ribbed in that species.
Species.
C. mamilla Fer., vi, 212. C. linmeana Pfr., vi, 214.
C. quoyi Desh., vi, 213. C. tirmaniana Anc., viii, 269.
undulata Q. & G., not Fer.
Subgenus CAM^ENELLA Pilsbry, 1893.
Camcenella PILS., Proc. Acad. Nat. Sci. Phila. 1892, p. 398, type
Helix platyodon Pfr. (Feb. 14, 1893).
Shell depressed, subglobose, solid, imperforate, banded and macu-
lated, with about 5J whorls, the last descending in front. Surface
106 CAMJENA.
minutely granular. Peristome well reflexed, the baso-columellar
margin toothed. Embryonal shell smooth, two-whorled, between
one-sixth and one-seventh the diameter of the adult, its junction
with the after growth indicated by a widening of the whorl. Type
C. platyodon Pfr., pi. 19, fig. 10.
Animal resembling Camsena. Sole very indistinctly tripartite, the
edges of foot lacking all appearance of a foot-margin ; sides irregu-
larly granulated ; tail rounded above, with a median, impressed lon-
gitudinal line, which does not extend quite to the tail.
Jaw strong, dark orange colored, having eight strong ribs (pi. 39^
fig. 3).
Central and lateral teeth of the radula (pi. 39, fig. 1) having a
single large cusp, which extends about to the posterior edge of the
basal plate. Marginal teeth (pi. 39, fig. 2) becoming quadri-cuspid,
by the splitting of both mesocone and ectocone.
Genital system (pi. 39, fig. 4) having a very short vestibule.
There is no dart-sack or other accessory gland upon the female side ;
spermatheca duct very long, without diverticulum. Penis stout,
exhibiting, when cut open (fig. 5), a very large penis-papilla ; from
the apex of the penis arises the slenderer epiphallus, which receives
the penis retractor at the middle, the vas deferens at the fourth of
its length ; beyond the insertion of the vas deferens it is continued
in a short flagellum.
In anatomy, Camcenella platyodon resembles Camasna in all
respects save that the penis-papilla is larger (a trifling difference)
and the cusps of the marginal teeth are much shorter. The shell
has a smaller nucleus than in Camcena more as in Obba, but not
so indistinctly defined ; and the maculated white and brown colora-
tion and deflexed last whorl are also as in Obba. It differs from
Obba in lacking an appendix on the penis, and in the ribbed jaw.
C. platyodon Pfr., vi, 239. Island of Hainan.
tournoueri Crosse.
Subgenus NEOCEPOLIS Pilsbry, 1891.
Neocepolis PILS., Man. of Conch, vi, p. 235, type H. merarcha.
Shell globose, solid, narrowly umbilicated, with elevated spire and
narrow, slowly widening whorls, the last deflexed in front. Aperture
truncate-rounded, the entire lip reflexed, its ends joined by a callus.
Columella dilated, thickened and obtusely toothed within. Typically
CAM^ENA-OBBA. 107
having a strong fold within the outer lip, marked outside by a deep
pit, as in Cepolis. Type C. merarcha, pi. 39, figs. 9, 10.
Anatomy unknown. The two species are from Tonquin. The
relations of the group are problematical, but judging from shell
characters, it belongs near either Obba or Camcenella.
C. merarcha Mab., vi, 235. C. morleti Dautz. & d'Ham., vi, 240.
mercatorina Mab., vi, 121.
Genus OBBA Beck, 1837.
Obba BECK, Index Moll. p. 30 (proposed for H. planulata, papilla,
mamilla) GRAY, P. Z. S. 1847, p. 172 (H. planulata selected as
the type). See also PILSBRY, Manual of Conch, vi, p. 211, and
viii, p. 270. Gallina HARTM., Erd u. Susswasser Gast. Schweiz,
1840, p. 197 (H. rota Sowb.). Philina (in part) ALBERS, Die Hel.
1850, p. 119 (preoc.). Pusiodon SWAINS., in part. Obbina SEM-
PER, Reisen im Archip. Phil., Landmoll., ii, p. 123 (type H. planu-
lata}, 1873.
Shell varying from trochoidal to depressed lens-shaped ; carinated,
at least in the young; umbilicated ; the surface striated or wrinkled
in the direction of growth-lines. Nucleus composed of about two
polished whorls, not distinctly marked off from the after-growth.
Last whorl very deeply deflexed in front, aperture sub horizontal ; ends
of the expanded peristome approaching, and connected by a cord or
raised callus ; basal lip reflexed and often bearing a tooth. Whit-
ish, buff or light brown, banded or speckled with brown. Type 0.
planulata, pi. 19, figs. 14, 15, 16. (See also pi. 19, fig. 17, 0. basi-
dentata).
Animal having a very broad flat foot, the tail short and quite flat,
sole undivided; tentacles short (pi. 39, fig. 7, 0. planulata).
Jaw smooth, without median projection, or with it small and blunt.
Dentition : Central and inner lateral teeth having wide mesocones,
no side cusps. Marginals developing a bifid mesocone and an ecto-
cone, the outer marginals having both mesocone and ectocone split
showing four short cusps, as in Camcenella platyodon. (PI. 39, fig
8, central and marginal teeth of 0. planulata; pi. 39, fig. 6, an
inner marginal of 0. basidentata}.
Genitalia lacking accessory appendages on the female side, the duct
of the spermatheca short. Penis short, continued in alongepiphallus
bearing the retractor muscle, vas deferens and a flagellum. The
108 OBBA.
cavity of the penis is strongly, irregularly plicate or papillose. It
is encircled by a feather-like glandular appendix (pi. 39. fig. 12),
the follicles of each side of which unite into two separate ducts
sunken into the cavity of the penis (pi. 39, fig. 13, section of the
gland). See pi. 39, fig. 11-13, 0. planulata.
Distribution, Philippine Islands, with a few forms in northern
Celebes, Halmaheira, and Ceram.
This group, like Cochlostyla, seems to have originated in the Phil-
ippine Island area. A few stragglers are found to the southward,
as is also the case with Camoena and some other typically Philippine
groups.
Obba differs from Camcena chiefly in the less capacious shell, with
subhorizontal aperture and continuous peristome ; in the possession
of a glandular appendix on the penis, the short spermatheca duct,
and the smooth jaw. It differs from Planispira in the raised parie-
tal callus and the keel of the shell, which is characteristic of all the
species when young, and most of them when adult. The teeth are
like those of Camcenella and Planispira. The anatomy of 0. plan-
ulata, listeri and basidentata has been examined by Semper (Reisen,
Landmoll., ii, p. 120) ; the jaw of rota by Morch.
In Vol. VI of this work the writer stated that Helix mamilla was
the type of Obba, following v. Martens. But in 1847 Gray selected
H. planulata for the type. The genus must, therefore, be restricted
to species allied to planulata.
(Group of O. planulata.')
O. papilla Mull., vi, 216. O. planulata Lam., vi, 220.
v. heroica Pfr., vi, 217. aurieulata Swains.
O. listeri Gray, vi, 218. papilionacea Val.
v. costata Semp., vi, 219. eollapsus Perry.
O. gallinula Pfr., vi, 219. v. sarcochroa Pils., vi, t. 68, f.
v. morongensis Mlldff. viii, p. [85.
[270. O. calcar v. Mart., vi, 221.
( Group of 0. moricandi.)
O. moricandi Sowb., vi, 222. O. scrobiculata Pfr., vi, 224.
O. basidentata Pfr., vi, 223. O. rota Sowb., vi, 225.
O. livesayi Pfr., vi, 223. O. platyzona Mlldff.
OBBA. 109
(Group of 0. marginata.')
O. bigonia Fer., vi, 226. O. kobeltiana Pfr., vi, 228.
samarensis Pfr., olim. O. parmula Brod., vi, 229.
bizonia H. & A. Ad. discus Dh., vi, 230.
O. marginata Mull., vi, 227. f. obscura Mlldff., vi, 230.
grayana Pfr. f. elevata Mlldff., vi, 230.
grayi Hombr. & Jacq. f. trochoidea Mlldff., vi, 230.
scabrosa Fer. O. bustoi Hid., vi, 230.
v. griseola Mlldff., vi, 228. O. saranganica Hid., vi, 230.
v. sororcula v. Mart., vi, 228. O. kochiana Mlldff, vi, 231.
devincta Tap. Can. O. bulacanensis Hid., vi, 226.
(Group of 0. horizontalis.*)
O. horizontals Pfr., vi, 232. O. lasallii Eyd., vi, 233.
O. reeveana Pfr., vi, 233. meretrix Sowb.
O. columbaria Sowb., vi, 234.
Subgenus? OREOBBA Pilsbry, 1 894.
Janira ALBERS, Die Heliceen, 1850, p. 124, only species H. codo-
nodes. Not Janira Leach, 1813 (Crustacea), of Oken, 1815 (acale-
pha) or of Schumacher, 1817 (Pecteri).
Shell globose-conoidal, bullet- shaped, composed of about 5 whorls
which are carinated in immature shells ; the embryonal portion not
differentiated ; last whorl defiexed in front. Surface shining, micro-
scopically spirally striated. Aperture truncate-rounded ; entire lip
well reflexed, at the columella expanded partly over the narrow
umbilicus, and armed with a callous tooth on the inner edge. Type
H. eodonodes Pfr., pi. 19, fig. 11.
Animal unknown. 0. eodonodes inhabits the Nicobar Islands.
It resembles the Philippine Island group Phcenicobius in contour, but
not in texture or minute sculpture, nor in the apical whorls. The
sculpture is like that of the Obba horizontalis group. Of the two
species I have seen only eodonodes. A knowledge of the anatomy
is necessary to the exact location of this group in the system. It
cannot, in my opinion, be united to Phcenicobius.
O. eodonodes Pfr., vi, 236. O. camel us Pfr., vi, 237.
110 PLANISPIRA.
Genus PLANISPIRA Beck, 1837.
= Planispira-\- Cristigibba-\-Angasella-\- Trachia-\- Eurystoma.
Planispira BECK, Index Moll., subg. 25, p. 29. MARTENS in
Albers' Die Heliceen, p. 160, type H. zonaria L. SEMPER, Reisen,
ini Arch. Phil., Landmoll., p. 120. TAPPARONE-CANEFRI, Ann.
Mus. Civ. Genov. xix, p. 162, 181, 1883. PILSBRY, Manual, vi, p.
"274. Pusiodon (in part) SWAINS., Malacol., p. 330 (for H. zonaria
and auriculata). Philina (in part) ALBERS, Die Hel., p. 119.
Shell depressed, generally umbilicated, having four to five rapidly
widening whorls, the first not granulated nor marked by hair points,
the last deeply deflexed in front. Aperture very oblique or subhori-
zontal; outer and upper lips expanded, basal lip reflexed, often
toothed. Type P. zonaria L., pi. 12, figs. 4, 5, 6.
Animal having the sole undivided (pi. 12, fig. 2, P. zonaria).
Genital system lacking all accessory organs on the female side,
the large spermatheca situated on a very long duct. Penis thick
and long, the retractor muscle apparently inserted at its apex ;
epiphallus ending in a short flagellum (pi. 12, fig. 1, P. zonaria).
Jaw smooth, arcuate (pi. 12, fig. 7 P. zonaria.)
Middle tooth and inner laterals with a single obtuse cusp ; outer
laterals and marginal teeth with the ectodont developed (pi. 12, fig.
3, P. zonaria).
Distribution, southern Celebes, Moluccas, New Guinea ; aberrant
groups extending over the Indo-Malayan and part of the Australian
regions.
This genus differs from Chloritis in the white or pale colored,
banded shell, very oblique aperture, and lack of quincuncial sculpt-
ure on the apex, and in the smooth jaw. It differs from Papuina in
the depressed earlier whorls of the shell and the ribless jaw. It
agrees with Obbain the jaw, but differs in lacking an appendix upon
the penis, and in the typically thinner, smoother shell, depressed at
the apex.
The anatomy is imperfectly known from the work of Semper and
Tapparone-Canefri. Investigation should be directed to the penis
in order to ascertain whether a papilla is present (denied by Semper),
the true limits of penis proper and epiphallus, and the point of inser-
tion of the penis retractor, v. Martens describes the jaw of P. loxo-
tropis as weakly ribbed. It is probable that 'the complete absence
PLANISPIRA. Ill
-of ribs will prove to be a character not without exceptions in this, as
in some allied genera.
The genus is divided into four well-defined subgenera :
* Shell white or light colored, generally banded, smooth, the
earlier whorls flat or concave.
Subgenus PLANISPIRA (restricted). Shell with no crest-like ridge
behind the lip ; aperture decidedly wider than high, the basal lip
usually somewhat thickened or toothed. Penis having a flagellum.
Subgenus CRISTIGIBBA Tap.-Can. Shell with a crest or swollen
ridge behind the lip; aperture about as high as wide, the basal
lip narrow, not thickened or toothed. Penis very short, the retrac-
tor and vas deferens inserted at its apex ; no flagellum.
* Shell often roughly sculptured, the earlier whorls not
especially depressed.
Subgenus ANGASELLA A. Ad. Shell depressed, the whorls tub-
ular, costulate or granulated ; aperture rounded or angular, the lip
well expanded, reflexed below. Australian species.
Subgenus TRACHIA Alb. Shell varying from discoidal to de-
pressed globose, generally banded on an opaque whitish ground.
Last whorl deeply deflexed in front. Lip expanded, reflexed below.
Anatomy as in Planispira except that the jaw is ribbed. Indo-Ma-
layan species.
Subgenus PLANISPIRA Beck.
Anatomy described above.
( Group of P. zonaria).
P. zebra Pfr., vi, 275. v. instricta Mart. [280.
guttata LeGuill. edentata Mart.
v. embrechtiana Mouss., vi, P. aurita Mart., vi, 281.
P. iaddse Pils., vi, 276. [275. P. biconvexa Mart., vi, 281.
P. halmaherica Strub., viii, 284. P. scheepraakeri Pfr., vi, 282.
P. chariessa Pils., vi, 279. P. zonaria Linn., vi, 277.
P. quadrifasciata LeGuill., vi, /. lineolata Mart.
112 PLANISP1RA.
P. zonaria Linn. v. fasciata Mart.
/. fulminata Mart. /. collis Mouss.
/. obliquata Mart. /. nitidiuscula Bttg., viii, 284.
/. inaculosa Mart. v. fasciolata Less.
/. coluber Beck. v. martini Schepm. Leyd. Mus.
lunulata Mart. [xv.
( Group of P. endoptycha).
P. endoptycha Mart., vi, 282. P. porcellana Grat., vi, 283.
f compta H. Ad.
(Group of P. zonalis).
P. zonalis Fer., vi, 284. P. loxotropis Pfr., vi, 285.
leucostoma A. & R. /. bernsteinii Mart.
P. atrofusca Pfr., vi, 285. /. laticlavia Mart.
P. latizona Pfr., vi, 285. /. angusticlavia Mart.
P. atacta Pfr., vi, 287. /. pluricincta Mart.
v. lorquini Pfr., vi, 286.
(Group of P. Icurri).
P. kurri Pfr., vi, 287. P. surrecta Bttg.
P. tietzeana Rolle, Nachrbl. '93, P. flavidula Mart, vi, 288.
[p. 33. flaveola Mts. not Kryn.
(Group of P. exceptiuncula}.
P. exceptiuncula Fer., vi, 289. P. thetis Pfr., vi, 290.
/. phryne Pfr. (see Nachrbl. 1892, p. 43).
/. aspasia H. Ad.
Subgenus CRISTIGIJBBA Tapparone-Canefri.
Cristigibba T.-C., Ann. Mus. Civ. Genov. xix, 1883, p. 161.
With the general appearance of Planispira, these shells differ in
having a crest or swollen ridge behind the lip, or a strong swelling
on the base immediately behind the constriction preceding the lip.
The spire is flat, a little concave in the middle. Type P. corniculum.
(See pi. 12, fig. 13-15, C.macgregori Hedley.).
The group is characteristic of New Guinea, but a few species range
as far north as Ceram, Batjan, and even Borneo and Sumatra.
PLANISPIRA. 113
Jaw arcuate, smooth (pi. 12, fig. 9, C. dominula). In C. mac-
gregori the lower margin shows traces of denticulation, and the
median portion is transversely wrinkled (pi. 12, fig. 11).
Central and inner lateral teeth with a single cusp, shorter than
the basal-plates. Marginals having a long, oblique, bifid mesoeone
and a small ectocone.
Genital system like that of Planispira on the female side. Penis
extremely short, stout, the retractor and vas deferens inserted at its
apex (pi. 12, fig. 8, P. plagiochila ; pi. 12, fig. 12, P. dominula}.
In this group the epiphallus and flagellum have evidently been
lost by degeneration. The anatomy of several species is known
through the researches of Tapparone-Canefri and Charles Hedley.
The following list of species will probably suffer considerable
reduction when sufficient material for comparative study is brought
together.
(Group of P. corniculwn).
P. corniculum H. & J., vi, 291. P. deaniana Ford, vi, 292.
/ kiesneri LeGuill. P. dominula Tap.-Can., vi, 293*
. P. purpurostoma LeGuill., vi, P. macgregori Hedl., viii, 285.
[177.
(Group of P. tortilabia).
P. tortilabia Less., vi, 294. P. rhodomphala T.-C., vi, 297.
torticollis (LeGuill.), T.-C. P. semirasa Mouss., vi, 295.
gibbosula H. & J. moluccensis Pfr.
P. plagiocheila T.-C., vi, 295. P. leptocheila T.-C., vi, 296.
( Group of P. margaritis).
P. margaritis Pfr., vi, 297. P. expansa Pfr., vi, 298.
v. zonulella Mouss. anozona Mart.
P. mersispira Mart., vi, 298. P. quadrivolvis Mart., vi, 299.
Subgenus ANGASELLA A. Ad.
. Angasella A. AD., P. Z. S. 1863, p. 521, only species, cyrtopleura.
Pleuroxia ANCEY, Conch ologists' Exchange, ii, p. 38 (Sept., 1887),
same type. Not Angasiella Crosse, 1864 (Nudibranchiata).
Shell depressed, umbilicated, plicate-striate ; whorls 4 to 5, the
last wide, deflexed in front. Aperture oblique, oval-truncate, the
114 PLANISPIRA.
peristome expanded, reflexed below, not toothed, margins approach-
ing and joined by a parietal callus. Type P. cyrtoplcura, pi. 19, figs.
20, 21, 22.
Distribution, South Australia. Anatomy unknown.
This group contains snails allied to the P. tuckeri group of the
Islands off the north coast of Australia, but modified by the condi-
tions of life in an arid region. Still it is doubtful whether the separa-
tion of the two groups serves any useful purpose.
Species.
P. cyrtopleura Pfr., iv, 65. P. eyrei Ad. & Ang., iv, 66.
P. phillipsiana Ang., iv, 66. P. subsecta Tate, iv, 66.
Section Trachiopsis Pilsbry.
Trachiopsis PILS., Manual of Conch, viii, p. 284.
Shell small, depressed, umbilicated, the whorls rather cylindrical,
covered with a brown cuticle, the last deflexed in front and more
or less constricted behind the lip. Aperture round or angular,
oblique, the lip thin, well expanded or reflexed, sometimes toothed.
Type P. tuckeri Pfr., pi. 19, fig. 18, 19.
Anatomy unknown. These small Planispira-like shells have
hitherto been classed in Trachia, an Indian group. They inhabit
the northern coast of Australia and adjacent islands. It is doubtful
whether this group should be separated from Angasella. It differs
mainly in the lighter, thinner texture of the shell, and the tendency
to form a tooth upon the basal lip.
P. tuckeri Pfr. iv, 65. P. delessertiana LeGuill, iv, 66.
strangulata H. & J. taranaki Gray.
P. cyclostomata LeGuill., iv, 65. torresiana H. & J.
P. dentoni Ford, viii, 285. P. endeavorensis Braz., P. Z. S.
P. dryanderensis Cox, P. Z. S., [1871, 640.
[1872, p. 19. P. baudinensis Smith, viii, 286.
P. collingii Smith, viii, 287.
Subgenus TRACHIA Albers, 1860.
Trachia ALB., Die Hel., edit. 2, p. 160. STOLICZKA, Journ.
Asiat. Soc. Beng. xl, (2), p. 223 (anatomy). Eurystoma ALB., Die
Hel. 1850, p. 126; edit. 2, 1860, p. 1 29, typeH.vittata. Cf. SEMPER,
PLANISPIRA. 115
Beisen im Archip. Phil., Landmoll., p. 163, anatomy of H. vittata.
Not Eurystoma Raf. 1818, nor Eurystomus Vieill., 1816. Philidora
<ie MORGAN, Bull. Soc. Zool. France, 1885, p. 384 (proposed for P.
wrayi and hardouini).
Shell varying from discoidal to depressed-globose, umbilicate, Hie~
surface rather roughly sculptured, hairy when young, microscopically
granulated, sometimes ribbed when adult ; the apex typically show-
ing no distinct sculpture. Last whorl strongly deflexed in front.
Aperture very oblique, small, the lip well expanded, reflexed below,
the terminations approaching and sometimes connected by a raised
callus. Type P. asperella, pi. 19, fig. 25. (See also pi. 19, fig. 24,
P. vittata; and pi. 19, fig. 23, P. vittata var. spinolce).
Animal (of P. delibrata} having the left body-lappet of the mantle
represented by a simple thickening; right lappet reaching anteriorly
over the back and rapidly becoming narrower below. In P. vittata
the sole is indistinctly tripartite.
Jaw arcuate, the entire anterior surface ribbed, the seven median
ribs stronger (pi. 32, fig. 44, P. delibrata. PI. 34, fig, 5, P. trovhalia).
In P. vittata there are five very high ribs, strongly denticulating
the margin.
Radula (of P. delibrata) very long, with 124 transverse rows of 22
(to 18) 20.1.20.18 (to 22) teeth. Central and inner lateral teeth with
a large mesocone and obsolete side cutting-points ; outer laterals and
marginal teeth with the ectocone developed. In P. vittata the
formula is about the same ; central and inner 14 laterals unicuspid ;
outer laterals with an ectocone. At the 25th tooth the mesocone
becomes bifid, and outwardly the bifid mesocone becomes shorter,
the outermost marginals having three subequal cusps. See also pi.
34, fig 4, P. asperella, and pi. 34, fig. 6, P. trochalia.
Genitalia having the female side free from all accessory organs,
the duct of the spermatheca very long. Penis terminating in an
^piphallus near the root of which the retractor is inserted ; epiph-
allus long, terminating in a short flagellum and the vas deferens (pi.
32, fig. 45 P. delibrata'). The genitalia of P. vittata are similar ;
penis with a spirally coiled flagellum. In P. penangensis (pi. 42,
fig. 6) the penis bears an epiphallus ending in a short flagellum, and
has an accessory sack, perhaps an " appendix."
Distribution, India, Burmah, Ceylon, Mergui Archipelago and
Sumatra.
116 PLANISPIRA.
These shells are characterized by the deeply descending whorl at
the aperture, and the strongly converging ends of the lip. The
anatomy is in essential agreement with either Chloritis or Plani-
spira, although the strong ribbing of the jaw is most like the former
group. On the other hand, the general form of the shell, the deep
descent of the last whorl to the very oblique aperture, and the
system of coloring, agree more nearly with Planispira. The sculp-
ture of the shell varies considerably in the different species. The
more typical, such as falladosa, nilagirica, proxima, as well as vittata
exhibit an apparently smooth apex ; but propinqua, tanqueryi and
a few others, show an excessively fine quincuucial puncticulation of
the apical whorls, such as occurs in Chloritis, in combination with
the characteristic shell contour of Trachia. Until we know by the
examination of numerous species, how and to what extent the char-
acters of jaw and genitalia are correllated with the above-men-
tioned shell structures, no consistent zoologist will be justified in
drawing rigid lines of demarcation between the Chloritis and Plan-
spiras of southeastern Asia. It is better to frankly recognize the
fact that in this area the two groups are represented by some forms
which, so far as shell characters show, are undifferentiated or separ-
ated by feeble characters only.
(Group of P. falladosa').
P. albicostis Pfr., iv, 65. P. helferi Bens., iv, 63.
P. asperella Pfr., iv, 62. P. nilagirica Pfr., iv, 65.
granifera Bens. P. penangensis Stol., iv, 63.
P. atkinsoni Theob., iv, 56 P. proxima Fer., iv, 63.
P. contracta Hutt., iv, 65. P. ruginosa Fer., iv, 63.
P. delibrata Bens., iv, 64. v. crassicostata Bens., iv, 64.
procumbens Gld. P. sordida Pfr., iv, 65.
v. fasciata G.-A., iv, 64. P. vittata Mull., iv, 120.
v. khasiensis Nev., iv, 64. zonula Wood.
P. fallaciosa Fer., iv, 64. v. spinolse Villa, iv, 120.
P. footei Stol., iv, 64.
(Group of P. c/abata).
P. trochalia Bens., vii, 88. P. pilisparsa Mart, viii, 192.
bigsbyi Tryon. P. smithii Bock, iv, 57.
P. gabata Gld., iv, 57. P. wrayi Morg., vii, 86.
v. merguiensis Phil. P. hardouini Morg., vii, 86.
CHLORITIS. 117
Genus CHLOKITIS Beck, 1837.
Chloritis BECK, Index Moll. subg. 24, p. 29. GRAY, P. Z. S.
1847, p. 172, type H. ungulina. v. MART., in Alb., Die Hel. 1860,
p. 161, type H. ungulina L. Erigone ALB., Die Hel. 1850, p. 92
(for discordialis Fer.). Semicornu " Klein," H. & A. ADAMS, Gen.
Rec. Moll, ii, p. 202, 1855. Cf. PILSBRY, Man. of Conch, vi, p.
242; viii, p. 270; and v. MOELLENDORFF, P. Z. S. 1891, p. 335,
336.-\-Sulcobasis Tap.-Can., Austrochloritis Pils., Trichochloritis Pils.
and Plecteulota Mlldff.
Shell varying from discoidal and biconcave to depressed subglo-
bose with convex spire ; the apical whorl flattened or sunken, and
showing under a lens regularly arranged granules or hair-points,
which often persist over the whole shell. Aperture lunate, the lip
reflexed. Type C. ungulina L., pi. 29, figs. 1, 2, 3.
Animal (of C. porteri) with undivided sole, the edges of the foot
lacking a foot border ; sides irregularly granulated ; tail rounded,
above with an impressed longitudinal median line ; back from mantle
to head having a few longitudinal grooves. Mantle edge bearing a
small right body-lappet.
Jaw strong and ribbed.
Radula having the middle cusp only developed on central and
inner lateral teeth, the cutting points about as long as the basal
plates ; side cusps completely absent, but represented by small cut-
ting points. Lateral teeth with a long, oblique, bifid mesocone and
a small ectocone.
Genital system characterized by the lack of dart sack or other
accessory organs on the female side, the spermatheca duct rather
long and closely bound to the uterus. Penis without appendix, its
cavity containing at the apex an iinperforate fleshy papilla (pi. 28,
fig. 2), situated beside the opening of the epiphalltis ; epiphallus
(pi. 28, figs. 1, 2, C. porteri) long, the penis retractor inserted upon
it ; terminating in a flagellum and vas deferens.
Distribution, Northern Australia and Solomon Is., north to south-
ern China. No fossil forms are known. All of the species live
upon the ground, as far as known.
The genus Chloritis was originally proposed for flat, biconcave
Helices ; but modern systematists have widened the group to con-
tain allied forms having the spire convex. Early in 1891 the
writer discussed the group, fixing upon the previously unnoticed
character of a quincuncially granulated apex as the true generic
118 CHLORITIS.
criterion, and considerably widening the limits of the genus. At
about the same time Dr. v. MollendorfF redefined Chloritis, and con-
cluded that the sculpture of " impressed points placed in quite
regular quincunx," and the presence of a " keel or angle round the
umbilicus" were diagnostic generic characters. In this connection it
should be noticed that the hairs or hair-points are totally lack-
ing upon the outer whorls of many undoubted Chloritis, and that
the umbilical angle completely fails in C. circumdata, maforensis, per-
cussa, etc. It therefore appears that the most we can say of the
sculpture is : apical whorls and mually the whole shell sculptured
with hair-points arranged in quincunx. It is probable that when
hairs or hair-points are present on the last whorl, they are always
disposed in regular oblique sweeps or quincunx, but this cannot be
said to be demonstrated. Some species show a granulation between
the hair points. The European Oligocene and Miocene species
which have been referred to Chloritis such as H. lepidotricha A.
Br., have no relationship to the Oriental Chloritis ; the H. lepido-
tricha is a Campylcea. In this connection it must be emphatically
stated that while the character of surface-sculpture discussed above
distinguishes Chloritis from other groups inhabiting the same quar-
ter of the globe, it is not in itself a feature of much importance,.
nor in itself diagnostic of this genus alone. In Europe the hairy
forms of the Campylcea planospira group (as well as some other
Campylaeas, such as setosa Ziegl.) show absolutely the same surface
sculpture, from the apex out. On the other hand, the Australian
group Hadra is extremely close to Chloritis in anatomy, but lacks
the quincuncial sculpture. We may, therefore, regard the quin-
cuncially arranged hairs as a secondary character, which has arisen
independently in several widely different groups of Helices. The
function of the hairs is evidently to gather dirt, thus disguising the
snail from its bird enemies.
Chloritis has the essential internal organization of Camcena, Cam-
cenella, etc. It differs from these groups and from Obba, mainly in
the non-differentiation of the embryonal whorls, and the smaller
size of the shell at the time its independent life begins. The spe-
cies referred by Semper to Chloritis belong to an entirely different
group. His anatomical characterization of the genus therefore
falls.
Chloritis may be divided into several sectional groups Chloritis,
Sulcobasis, Austrochloritis, Trichoehloritis probably natural, but
.
CHLORITIS. 119
blending at their confines into one another. The typical forms of
the first two represent the more divergent and presumably modern
lines of differentiation.
Section Chloritis (restricted).
Shell with the spire sunken, flat or somewhat convex with flat
earlier whorls. Type C. ungulina, pi. 29, figs. 1, 2, 3.
But two species of the typical group of Chloritis have been inves-
tigated anatomically, C. dinodeomorpha Tap.-Can., Ann. Mus. Civ.
Genov. xix, 1883, p. 168, and C. leei Cox, Hedley, Proc. Linn. Soc.
N. S. W. (2), vi, p. 687. They agree essentially with Austrochlo-
ritis, q. v.
Jaw arcuate, having about 8 strong ribs separated by narrow
intervals (pi. 32, fig. 43, C. leei). Central and inner lateral teeth
unicuspid ; marginal teeth having a long bifid mesocone and an
ectocone. Genitalia lacking appendages on the female side, the
duct of the spermatheca long. Penis long, the retractor apparently
inserted at its apex ; epiphallus very long, dilated where it receives
the vas deferens, and ending in a flagellum (pi. 28, fig. 10, C. dino-
deomorpha, after Tap.-Can. ; pi. 32, fig. 42, C. leei, after Hedley).
Distribution, New Guinea and Moluccas (typical forms) ; Solo-
mons, New Ireland, Louisiades and Celebes (divergent forms).
( Group of ungulina.)
C. ungulina Linn., vi, 243. C. biomphala Pfr., vi, 244.
v. minor Fer. C. martensi Pfr., vi, 244.
C. unguiculina v. Mart., vi, 244. C. cheratomorpha Tap.-Can., vi,
[245.
( Group of circumdata.)
C. circumdata Fer., vi, 246. C. maforensis Tap.-Can., vi, 247.
mollweta Pfr., vi, 246. ' v. micromphalus Pils., vi, 247.
C. lansbergiana Dohrn, vi, 247.
(Group of unguicula.)
C. unguiculastra v. Mart., vi, C. ceramensis Pfr., vi, 249.
v. buruensis Mart. [248. C. unguicula Fer., vi, 249.
v. amboinensis Mart. yoldii Morch.
v. pilosa Mart. C. gruneri Pfr., vi, 250.
C. flexuosa Pfr., vi, 249. C. exacta Pfr., vi, 250.
120 CHLORITIS.
(Group of eustoma.)
C. erinacea Pfr., vi, 251. C. ursina Pfr., vi, 253.
C. leei Cox, vi, 251. C. dinodeomorpha Tap.-Can., vi,
v. sudestensis Hedley. [254.
C. subcorpulenta Sm., vi, 251. C. delphax Dohrn, viii, 271.
C. discordialis Fer., vi, 252. C. silenus Angas, vi, 254.
C. eustoma Pfr., vi, 252. C. gaimardi Dh., vi, 255.
C. dentrecasteauxi Sra., vi, 253. adustus Hinds.
C. mendanse Cox, vi, 255.
(Group of tuba Celebes species.)
C. bulbulus Mouss., vi, 258. C. tuba Alb., vi, 258.
bulbus Mouss. C. zodiacus Fer., vi, 259.
Section Sulcobasis Tap.-Can.
Sulcobasis T.-C., Annali del Museo Civico di Storia Naturale di
Geneva, xix, 1883, p. 161.
Shell large, solid, globose-depressed or depressed ; spire convex,
the inner whorls (and apex when not worn) showing minute hair -scars
arranged in oblique series; body-whorl more or less distinctly spirally
sulcate beneath. Lip well reflexed. Type C. sulcosa Pfr., pi. 29,
figs. 9, 10.
Distribution, Aru Is., New Guinea, New Ireland, Solomon Is.
Anatomy unknown. Tapparone-Canefri has given a crude figure
of the central and inner lateral teeth of C. beatrids, showing them
them to lack side cusps, as usual in the genus. Doubts have been
expressed as to the relationship of this group of large solid Helices
to Chloritis (Jahrb. D. M. G. 1892, p. 94) ; bufc those who see the
shells themselves, will agree with Tapparone-Canefri that the group
is simply a section of Chloritis.
(Typical group).
C. sulcosa Pfr., vi, 260. C. lepidophora Dohrn. viii, 273.
C. rubra Alb., vi, 260. C. rehsei v. Mart., vi, 261.
C. concisa Fer., vi, 262. gerrardi Sm.
C. beatricis Tap.-Can., vi, 260. genardi Braz.
C. rohdei Dohrn, viii, 273. v. obtecta Reinh., vi, 262.
CHLORITIS. 121
(Aberrant group).
. bougainvillei Pfr., vi, 128. C. quercina Pfr., vi, 257.
angasiana Newc. v. hombroni Pfr., vi, 258.
. majuscula Pfr., vi, 255. janellii Hombr. & Jacq.
. isis Pfr., vi, 256.
Section Austrochloritis Pilsbry.
Austrochloritis PILS., Man. of Conch, vi, p. 262. ? Plecteulota v.
MOELL., Jahrb. D. M. Ges. 1892, p, 92, type Eulota goniostoma
Mlldff.
Shell rather small, depressed, but with convex spire and obtuse
apex, umbilicated, unicolored ; surface hairy or marked with regular
series of hair-scars to the apex. Aperture round-lunar, the lip
expanded, thin, ends of peristome converging; sutures well-im-
pressed. Type C. porteri Cox, pi. 29, figs. 4, 5.
Animal (see under Chloritis).
Jaw arcuate, with numerous ribs (pi. 28, fig. 3, C. porteri).
Dentition : Central and inner lateral teeth with the mesocones
only developed, slight lateral cutting-points upon it representing the
absent ectocones. Marginals having a long, oblique mesocone and
a small ectocone (pi. 28, fig. 4, C. porteri).
Genitalia (of C. porteri) lacking all accessory appendages on the
female side; spermatheca lying beside the albumen gland, its duct
therefore very long, bound closely to the oviduct thorougnout its
length. Penis club shaped, the walls of its cavity corrugated, with
a large, fleshy papilla at the apex, beside the opening of the epiphal-
lus (pi. 28, fig. 2). Epiphallus long, the retractor inserted at its
middle; ending in a rather long flagellum. Penis retractor
attached to the floor of the lung cavity ; right eye-peduncle retrac-
tor passing between primary branches of genitalia (pi. 28, fig. I C.
porteri Cox. Fig. 2 penis of same opened, epiphallus and
flagellum).
Distribution, Queensland, New Guinea and adjacent islands.
The anatomy of P. porteri has been investigated by Hedley (Proc.
Koy. Soc. Queensl. vi, pi. 15) and by myself (see above). The jaw
and teeth of C. chloritoides have been figured by Hedley (Proc.
Linn. Soc. N. S. W. (2), vi, pi. 39, 40). The anatomy of C. argillacea
has been described and figured by Wiegmann, in Webers' Zool
122 CHLORITIS.
Ergebnisse einer Keise in Niederlandisch Ost- Indien, III, p. 171.
Part of his figures are reproduced on pi. 28, figs. 5-9. The epiphallus
bears a short accessory organ (shown below the penis retractor in
fig. 8, above it in fig. 9) of unknown homology and function.
Otherwise the jaw, teeth and genitalia agree with C. porteri.
The section Plecteulota of v. Mollendorff, considered by him to-
be a subordinate group of Eulota, probably belongs here. Its type
Plecteulota goniostoma Mlldff. is shown in pi. 29, figs. 6, 7.
Small, thin-shelled forms, having much the aspect of Eulotella,.
from which they differ in the sculptured apex and the lack of dart-
sack and the associated mucus gland or glands. It is in actual
practice, however, extremely difficult to tell what shells to refer to
Eulotella, what to Chloritis ; and the most experienced conchologists
differ in their treatment of the forms. Most of the shells now
included in Austrochloritis were placed by Pfeiffer in Dorcasia and
Camcena; and v. Mollendorff has expressed the opinion that part of
them are to be referred to Eulota (plus Plecteulota, Eulotella, etc.).
In regard to these conflicting opinions, the writer has only this to
say : the groups Eulota and Austrochloritis, notwithstanding their
superficial similarity, belong to widely different branches of the
Helix stock. Controversy respecting the generic position of certain,
species known by the shells alone is idle ; for the anatomy only can
give a true answer to our questioning.
(Australian species).
C. spinei Cox, vi, 263. C. aridorum Cox, vi, 266.
hysirix Cox, preoc. C. pseudoprunum Pils., viii, 271-
C. porteri Cox, vi, 263. prunum auct. not Fer.
C. mansueta Cox, vi, 264. C. coxeni Cox, viii, 272.
C. blackalli Braz., vi, 264. C. bennetti Braz., vi, 135.
C. buxtoni Braz., vi, 265. C. blackmani Cox, vi, 137.
C. brevipila Pfr., vi, 265. C. coxense Braz., vi, 1 38.
C. mucidaPfr.,vi, 148.
(Species of New Guinea, etc.).
C. occulta Pfr., vi, 266. C. telitecta Mlldff 1 ., viii, 222.
C. chloritoides Pils., vi, 267. C. tenuitesta Mlldff., viii, 273.
C. rhodochila Mlldff., viii, 273. C. argillacea Fer., iii, 210.
C. micholitzi Mlldff., viii, 272. cyclostomopsis Lea.
C. goniostoma Mlldff, viii, 221. C. mendax Martens, iii, 212.
CHLORITIS. 123
Section Trichochloritis Pilsbry.
Triehochloritis PILS., Manual of Conch., vi, p. 267.
Shell depressed, rather thin, the spire low-convex or flat, the base
generally obtusely angled around the umbilicus. Epidermis~not
deciduous; apex and the whole shell hirsute or marked by hair-
scars arranged in regular lines ; lip thin, expanded or narrowly
reflexed. Type C. breviseta Pfr.
Anatomy unknown. Distribution, Southern China to Borneo.
As I have written in this work (vi, p. 242) and von MollendorfF
has emphasized (Nachr., 1892, p. 94), the sections of Chloritis stand
" auf etwas schwachen Fiissen." In other words, the series seems
to intergrade by rather easy stages throughout, not even excepting
Sulcobasis. Disclaiming any desire to supply crutches to a section
which cannot stand upon its own merits, I still retain the name
Trichochloritis for the group of small, thin species having the same
distribution as Camcena, believing it a convenient division. When
enough species are known anatomically to show the true classifica-
tion of Chloritis and the line dividing that genus from Trachia and
Eulotella, I shall be among the first to discard the present arbitrary
system.
The genital system of C. crassula has been figured by Wiegmann
(Zool. Ergebnisse einer Reise in Niederlandisch Ost-Indien. iii, pi.
13, f. 10). It resembles that of C. portei except that the enlarge-
ment at the apex of the penis is long and curved so long that
Wiegmanu calls it a penis gland, although in my opinion, it is not
glandular, but simply a pouch-like enlargement of the penis for the
accomodation of a large imperforate papilla.
The epiphallus bears the retractor, and is continued beyond the
insertion of the vas deferens in a short flagellum. The duct of the
spermatheca is much and abruptly swollen at the base and this
swelling is doubtfully interpreted as a dart-sack and mucus gland
by Wiegmann, who did not open it, however. If his view is correct,
the species must be an Eulotella ; but I prefer to consider the
structure as a mere muscular enlargement of the spermatheca duct,
probably with plicate internal walls, such as is often found in the
Helices. The union of dart-sack with spermatheca duct would be
an extremely unusual character, if confirmed.
124
CHLORITIS-ALBERSIA.
( Continental species').
C. hungerfordiana Nev., iii, 182.
C. miara Mab., vi, 270.
C. herziana Mlldff., vi, 271.
. rhinocerotica Hde., vi, 271.
C. franciscanorum Gred., viii,
[217.
C. seriatiseta Roch., vi, 268.
C. malayana Mlldff., viii, 274.
C. percussa Hde., vi, 111.
C. breviseta Pfr., vi, 268.
C. tenella Pfr., vi, 269.
C. submissa Desh., iii, 182.
C. deliciosa Pfr., vi, 113.
C. remoratrix Mori., viii, 274.
C. lemeslei Mori.
C. balansai Mori., viii, 218.
C. quinaria Pfr., vi, 269.
guinaria Pfr.
C. shanica Bedd., viii, 275.
C. colletti Bedd., viii, 274.
C. bifoveata Bens., vi, 245.
C. nautiloides Val., iii, 212.
C. samuiana Mlldff.
C. tanqueryi C. & F., iv, 64.
C. condoriana C. & F., vi, 269.
C. Dorodomiana Mori., vi, 270.
C. fouresi Mori., J. de C., 1889,
C. propinqua Pfr., iv, 63. [176.
(Species of Borneo, Java, etc.).
C. crassula Phil., viii, 271.
storiana Mouss.
C. cryptopila Marts., iii, 211.
C. hemiopta Bens., vi, 238.
C. meander G.-A., viii, 275.
C. plena G.-A., viii, 276.
v. helicinoides Mouss., iii, 211. C. sibutuensis Sm., Ann. Mag.,
C. everetti H. Ad., iii, 211. [1894, p. 53.
C. tomentosa Pfr., iii, 212.
(Philippine Island species').
C. brevidens Pfr., vi, 272. C. quieta Eve., vi, 271.
C. leytensis Mlldff., Nachr. '90, C. inquieta Dohrn, viii, 273.
C. malbatensis Hid. [203.
Genus ? ALBERSIA H. Adams, 1865.
Albersia H. AD., P. Z. S., 1865, p. 410, type H. granulata
Q. & G. v. MARTENS, Ostasiat. Zool., Landschn. p. 329, 1867.
TAP. CAN., Ann. Mus. Civ. Genov. xix, p. 185, 1883. PILSBRY,
Manual vii, p. 89.
Shell globose, thin ; aperture but slightly oblique, the peristome
hardly thickened, narrowly reflexed ; columellar margin rather
steeply ascending, narrowed below. Unicolored or banded, never
brilliantly colored, the surface dull, granulated or hairy. Type A.
granulata, pi. 41, fig. 30.
ALBERSIA-THERSITES. 125
External anatomy and genitalia unknown.
Jaw arcuate, solid, with 6 strong ribs, denticulating the margins,
and grouped on the median part of the jaw, the ends free from ribs
(pi. 34, fig. 8, A. zonulata). The jaw of A. pubicepa also is
stated by von Martens to be ribbed.
Radula as in Chloritis, etc. ; the central and inner lateral teeth
having a single cusp shorter than the basal plates, the side cusps
represented by slight lateral extensions of the central cusp. Outer
laterals having a long, oblique cusp, which becomes bifid on the
marginals (mesocone+entocone), and on the outer teeth a small
ectocone appears (pi. 34, fig. 9, A. zonulata).
This group should perhaps be considered a subgenus of Chloritis,
but it differs in the thin, capacious form of the shell and the
Cochlostyla-like columella. No just estimate of the rank or posi-
tion of the group can be made until the soft anatomy is investigated.
The jaw and teeth offer no differences from those of Chloritis,
Thersites, etc. Distribution, New Guinea and Moluccas.
A. granulata Q. & G., vii, 90. A. zonulata Fer., vii, 91.
A. pubicepa v. Mart., vii, 90. lemniscata Less.
tortistylis Mouss. v. recluziana Le Guill.
A.pseudocorasiaStrub., viii, 293. A. tenuis Pfr., vii, 91.
Genus THERSITES Pfr., 1855.
Thersites Pfr., 1855, plus Hadra Alb., 1860, plus Badistes Gld.,
1862, plus Sphcerospira Morch, 1867, plus Xanthomelon v. Mart.,
1860, plus Rhagada Alb., 1860, plus Qlyptorhagada Pils., 1890.
Shell narrowly umbilicate or imperforate, varying from globular
to trochoidal or thick lens-shaped and keeled, usually solid.
Whorls 5 or 6, the apex smooth, never granulated or punctate in reg-
ular quincunx ; last whorl varying from smooth to rudely wrinkled,
generally densely granulated or roughened microscopically, but
never bearing spaced liairs or hair-scars in regular oblique series.
Aperture moderately oblique, the outer lip expanded (except in
Glyptorhagada), basal lip reflexed, dilated at the columellar insert-
ion, the ends of the lip rather remote. Type T. richmondiana, pi.
29, fig. 8. (See also all figures on pi. 27).
Animal having the general features of that of Camcena, Chloritis,
etc. ; the sole undivided and without grooves above its margin ;
back with one or few grooves from mantle to head ; sides irregularly
126 THERSITES.
tuberculate ; tail with a slight median longitudinal groove above
(pi. 33 figs. 6, 7. T. gulosa Gld.).
Jaw arcuate, stout, with 5 to 12 unequal, strong ribs (pi. 32,
figs. 47, 48, 50). Teeth having the side cusps of centrals and inner
laterals completely fused with the middle cusps ; marginals having
a long bifid inner cusp (entocone plus mesocone) and a simple or
bifid ectocone (pi. 34, fig. 1, T. mitchellce}.
Genital system having no accessory organs on the female side,
the duct of the spermatheca generally long and swollen below.
Penis enlarged distally, where its cavity contains a solid papilla ;
epiphallus bearing the retractor, and terminating at the entrance of
the vas deferens in a short flagellum (pi. 33, fig. 1, Thersites.
richmondiana, and figs. 2, 3, T. mitchellce. PL 51, fig. 10, T. soloren-
sis). In some species the epiphallus is shortened and the flagellum
very short or absent by degeneration (pi. 32, fig. 52, T. pachystyla,
and fig. 51, T. rainbirdi).
Habits strictly terrestrial. With the exception of a few New Gui-
nea species, and some inhabiting the Timor group, the species of
this genus are confined to Australia, where they are generally dif-
fused, everywhere constituting the most prominent feature in the
Helix fauna.
The various sections assembled under the generic term Thersites
form a very homogeneous group, the extreme forms being well con-
nected by a chain of intermediate species, Xanthomelon and Ehagada
forming outlying or satellite groups of slightly greater systematic
value than the other sections, but still intimately allied. The shell
varies from thin, light forms like corneovirens through a series of
transition species to the solid, richly dyed blomfieldi, mitchellce and
bipartita; and by other chains of almost unbroken continuity, the
globose forms are connected with the keeled richmondiana and
kooringensis. The soft anatomy fully sustains these conclusions.
The genus Thersites is allied to Chloriiis, and might without any
great violence be united to that genus ; but it will probably prove
an aid to clear and correct thinking to retain the two separate.
Thersites never has the depressed earlier whorls, or quincuncially
arranged hairs or spaced points so characteristic of Chloritis, and
the flagellum is shorter or obsolete.
The distribution of the Thersites and Chloritis groups seems to
indicate a hypothesis of two separate times of connection between
Australia and the Papuan tract since the beginning of the Tertiary.
THERSITE8. 127
The first may have been eocene, at which time the Australian land
snail fauna received the ancestors of Thersites (-{-Hadra, etc.), and
of Panda, Pedinogyra, etc. At this time the Hadra group was not
differentiated from Chloritis. Subsequent isolation of Australia
resulted in the spread of the Hadra group and its segregation into
the modern subgenera ; and during this interval the genera Thersi-
tes and Chloritis were differentiated, the one in Australia, the other
in Papua. It is probable that much of the differentiation of
Planispira and Papuina, which are so intimately allied to
Thersites and Chloritis, occurred now, although the bases of these
branches may strike still deeper. At all events, they seem to have
peopled New Guinea during this interval. The second connection
of Queensland with Papua was comparatively recent, although
remote enough to allow specific differentiation (see Hedley in The
Nautilus, March, 1893, p. 124), and at this time, as Hedley believes,
the Chloritis species invaded Queensland from the north, with
Papuina, Atopos and the land operculates. At the same time
Queensland gave to New Guinea its few species of Thersites (Sphce-
vospira broadbenti, etc.), and perhaps some other forms.
THERSITES vs. HADRA. The present group as a whole has hith-
erto been known as Hadra Alb. (See v. Martens, Die Heliceen ;
Semper, Reisen ; Hedley, Proc. Roy. Soc. Q. and P. L. S., N. S.
Wales; Pilsbry, Man. Conch.), but the name Thersites has priority
of five years over Hadra. It has also prior position in Die Heliceen,
where it is diagnosed and restricted. In view of these facts, and of
the further consideration that the nomenclature of Helices is now in
^ transition stage, we cannot refuse to follow the course indicated
by established rules of nomenclature. There is another bar to the
use of Hadra in a generic sense ; it is preceded in the pages of Die
Heliceen by Rhagada, and this would give the latter name priority,
for there can be no doubt that both belong to one genus.
It is now obvious that the use of the name Hadra by German
writers on shells of China and Japan is founded upon a misconcep-
tion of their relationships. Part of the " Hadra" species of these
authors belong to Camcena, part to Euhadra, a group closely allied
to Campylcea, etc.
The subdivisions of Thersites are not very well defined naturally,
but the following may be admitted :
Subgenus THERSITES, in which the shell has rather a conoidal
spire and is yellowish or brown, generally banded, the spermatheca
128 THERSITES.
having a long duct ; containing sections Thersites, Glyptorhagada^
Badistes, Sphcerospira, Hadra.
Subgenus RHAGADA, with small, depressed globose shell, calca-
reous in texture and white or whitish, often multilineate ; the anat-
omy as in the preceding.
Subgenus XANTHOMELON, with a globular shell with wide colum-
ellar lip, the spermatheca duct short.
Subgenus THERSITES Pfr.
Section Thersites Pfr. (restricted).
Thersites PFR. (in part), Mai. Blatter ii, p. 141 (1855 or 1856).
v. MARTENS in Alb., Die Hel. p. 157, type H. richmondiana. PILS-
BRY, Man. Conch, vi, p. 90. Cf. HEDLEY, Proc. Roy. Soc. Queens,
v, p. 62, and vi, 1889, p. 62, pi. 3 (anatomy). Not Thersites Spence
Bate 1857 (Amphipoda), nor Pagenstecher 1861 (Entomostraca).
Shell lens-shaped or trochiform, imperforate when adult, carinated
at the periphery, more or less pinched at the keel, the last whorl de-
scending in front. Aperture sub-triangular) oblique, the outer lip
expanded, sinuous above the outer angle ; basal and eolumellar lips
reflexed. Type T. richmondiana Pfr., pi. 29, fig. 8.
Animal externally like Sphcerospira. Jaw strongly arcuate, with
slightly attenuated, blunt ends, sculptured with about 11 flat ribs,,
broader than their interspaces, and denticulating the cutting mar-
gin (pi. 34, fig. 7, T. richmondiana). Radula as in Sphcerospira.
Genitalia (pi. 33, fig. 1, T. richmondiana) as in Sphcerospira mit-
chellce, etc. The penis is short and dilated distally, evidently for the
accommodation of an internal papilla. Epiphallus long, bearing the
retractor at its middle, terminating in a short flagellum. Duct of
spermatheca very long, its lower portion large and swollen.
As will be seen by the figures, the anatomy of Thersites richmond-
iana offers no departure of more than specific value from that of
Sphcerospira mitchellce and its allies. The group is simply a keeled
form of Hadra, really not more different from the normal Hadras
than Polygyra {Stenotrema} spinosa is from P. stenotrema, or than
Chlorcea thersites is from C. sirena. The development of a keel is
now universally acknowledged to be a character of very slight sys-
tematic value in the Helices, too slight in most cases to be held of
more than specific importance. Scores of sectional groups contain
both rounded and keeled species. The true relationships of Thersites
THERSITES. 129
were perceived simultaneously and independently by Charles Hed-
ley and the writer. Our knowledge of the anatomy is due to
Hedley.
The name Thersites being anterior in date to Hadra, will replace
that term as a generic designation for the entire series. The same'
name has been used in Crustacea and Insecta, but later than Pfeiff-
er's application of it to the present group.
T. richmondiana Pfr. vi, 90. Queensland, northern N. S. Wales.
/. decolorata Pils. vi, 91.
T. novsehollandise Gray, vi, 91. New South Wales, Australia,
depuyana Pfr.
Section Glyptorhagada Pilsbry, 1890.
Glyptorhagada PILS., Man. Conch, vi, p. 191 (Dec. 16, 1890).
Depressed, keeled Badistes, having the surface corrugated by-
strong oblique fold-like wrinkles, the outer lip hardly expanded ;
texture calcareous. Type H. silveri, pi. 27, fig. 19. (See also If.
kooringensis, pi. 27, figs. 7, 8, 9, 10).
This is the South Australian expression of the Badistes type ; the
rudely sculptured, earthy shell responding to the arid condition
prevailing in the interior of South Australia, in accordance to the
well known law governing the modification of desert snails. The
anatomy is unknown. The species were formerly grouped in Rha-
gada, but their affinities are evidently with Badistes.
H. silveri Angas, vi, 191. H. bordaensis Aug., vi, 192.
H. kooringensis Angas, vi, 191. H. howardi Ang., iv, 52.
Section Badistes Gould, 1862.
Badistes GLD., Otia Conch, p. 243, type H. gulosa Gld. PILS-
BRY, Man. Conch., vi, p. 94, 129. For anatomy see HEDLEY, Rec.
Austr. Mus., i, p. 196, pi. 29, 1891.
Shell generally smaller and thinner than that of Splmrospira, the
surface densely microscopically granulated all over ; often with a pe-
ripheral keel. Peristome a little thickened and very natrowly ex-
panded, suddenly dilated at the columellar insertion, closing or
almost closing the narrow umbilicus. Type T. gulosa, pi. 27, fig. 5
(see also pi. 27, fig. 3, T. bitceniata).
The animal has a slight groove on each side, running from lips up-
ward and backward to mantle ; back with a median furrow banded by
two rugse or sets of rugse, on each side owai<j&ere are about six
130 THERSITE8.
ranks of long, narrow tubercles. The rest of the body is covered with
irregular polygonal tubercles which are usually partially subdivided
into minor tubercles ; those on the tail being small, round and en-
tire. There is a small triangular right mantle lappet, and appar-
ently, a long left lappet, which emits two small lobes on the left
side at the origin of the left facial (lateral) groove (pi. 33, figs. 6, 7,
living animal of T. gulosa, after Hedley).
Jaw arched, crossed asymmetrically by 9 stout, flat-topped unequal
ribs, denticulating both margins ; ends smooth (pi. 33, fig. 5, T.
gulosa).
Radula(of T. gulosa) having 180 rows of 39'18-M8'39 teeth.
Central and inner lateral teeth unicuspid ; outer laterals oblique ;
marginals with a long, oblique bifid inner cusp (ento-+naeso-cone)
and a small ectocone.
Genitalia (pi. 33, fig. 4, T. gulosa, after Hedley), having the
penis twisted and swollen near its apex ; retractor inserted low on
the epiphallus, which bears a flagellum at the insertion of the vas
deferens. Duct of the spermatheca long, inserted high on the vag-
ina.
In soft anatomy and dentition, Badistes offers no variation from
the type prevailing in Sphcerospira, Thersites or Chloritis. In dis-
tribution it is more southern than Sphcerospira, occurring mainly in
New South Wales, Victoria and South Australia. The species are
highly polymorphic, and have evidently been moulded by external
conditions into a great number of local forms. There are more than
enough specific names, the only difficulty being which and how
many to discard. The reduction of species in the following list is
mainly made by the advice of Messrs Cox, Hedley and Brazier
Con/. Brazier, Proc. Linn. Soc. N. S. Wales (2), vi, p. 321.
Gould supposed that Helix gulosa travelled like the caterpillar of
a geometric moth, by a series of loops ; but this has been shown to
be an error, probably caused by some confusion in the collector's
notes.
( Group of gulosa.')
T. duralensis Cox, vi, 141. T. laesa Rve., iii, 214.
T. daintreei Braz., vi, 134. T. pliculosa Pfr., iii, 216.
T. patruelis Ang., vi, 131. T. expeditionis Cox, iii, 214.
T. dunkiensis Forbes, iii, 215. T. corneovirens Pfr., vi, 136.
v. mulgo^e Cox, vi, 136.
THERSITES. 131
T. gulosa Gould, vi, 131. T. greenhilli Cox, vi, 138.
lessoni Pfr., not auct. viii, 281. T. liverpoolensis Braz., vi, 141.
coriaria Pfr., vi, 132. T. marcescens Cox, vi, 142.
morosa Morel., vi, 134. T. (?) subgranosa Le Guill. vi, 137.
monacha Pfr., vi, 133. T. (?) plethorica Crse., vi, 138.
mastersi Cox, vi, 133. T. leucocheilus Cox, vi, 139.
scotti Cox, vi, 133. marice Cox, preoc.
Icailleti Crs., iii, 216. T. lismorensis Pils., vi, 140.
T. jervisensis Q. & G., vi, 141 ; T. bellengerensis Cox, vi, 140.
viii, 281. T. yatalaensis Cox, vi, 140.
gilberti Pfr., vi, 142. T. evandaleana Pfr., vi, 142.
grayi Pfr., vi, 130. T. tomsetti Tate, vi, 143.
exoearpi Cox, vi, 139. T. lincolnensis Pfr., vi, 144.
bednalli Braz., vi, 130. T. luteofusca Cox, vi, 144.
? sutilosa Fe"r.
(Group of bitceniata, South, Central and Western Australia).
T. perinflata Pfr., viii, 282. T. bourkensis E. A. Sm., vi, 308.
T. bitseniata Cox, vi, 144. T. angasiana Pfr., vi, 180.
flindersi Ad. & Ang. T. nullarborica Tate, vi, 181.
T. lorioliana Crosse, vi, 145. T. fodinalis Tate, viii, 277.
T. broughami Ang., vi, 146. T. everardensis Bedn., viii, 277.
T. rufofasciata Braz., vi, 146. T. elderi Bedn., viii, 278.
T. sublorioliana Pils., vi, 147. T. oscarensis Cox, viii, 279.
T. cassandra Pfr., vi, 147. T. derbyi Cox, viii, 280.
T. stutchburyi Pfr., vi, 148. T. forrestiana Aug., vi, 182.
Section Hadra Albers, 1860.
Hadra ALB., DieHel. (edit. Martens), p. 165, type H. bipartita.
Cf. SEMPER, Reisen, etc., pi. 17, f. 16, dentition of H. bipartita.
Shell depressed with conoidal spire, narrowly umbilicated,
obliquely striate or hirsute ; unicolored, or brown below, yellow
above, never having many bands ; peristome expanded. Type T.
bipartita Fer.
Dentition (of T. bipartita, pi. 32, fig. 49) similar to that of Sphce-
rospira, etc. ; the central and lateral teeth unicuspid, marginals
with an ectocone. The figure shows a central with one adjacent
lateral tooth, and the 47th side tooth.
Hadra, as restricted, consists of a few north Queensland species,
differing somewhat from Sphserospira in shell characters.
132 THERSITES.
T. bipartita Per., vi, 126. T. forsteriana Pfr., vi, 127.
semibadia Alb. hetcera Pfr.
/. unicolor Cox, viii, 276. /. major Dohrn, vi, 128.
/. minor, vi, 126. T. darwini Braz., vi, 128.
Var. semicastanea Pfr., vi, 126.
funiculata Pfr.
Section Sphcerospira Morch.
Sphcerospira MOERCH, Journ. de Conchyl., 1867, p. 256, for H.
fraseri, lessoni, appendiculata. For anatomy, see SEMPER, Reisen,
p. 160, pi. 14, f. 11 (basalis), and HEDLEY, Proc. Roy. Soc. Queensl.
vi, pi. 7, 8 {fraseri, blomfieldi, rainbirdi), and Proc. Linn. Soc. N.
S. W. (2), vi, pi. 39, 41, 42 (broadbenti).
Shell globose, solid, yellowish, with brown spiral lines and bands
or uniform chocolate-brown by coalescence of the bands; spire ele-
vated, somewhat dome-shaped ; surface smooth to the naked eye.
Peristome broadly expanded. Type H. fraseri. (See pi. 27, fig. 4,
T. blomfieldi var. warroensis Hedl. PI. 27, figs. 1, 2, T. rawnesleyi
Cox).
Animal having the sole indistinctly tripartite ; edges of foot with-
out a foot border ; sides irregularly granulated ; tail convex above,
with an inconspicuous longitudinal impressed line ; back from man-
tle to head with several longitudinal grooves. Mantle bearing a
small triangular right body lappet and a minute left lappet. (Mit-
chellce).
Jaw arcuate, strong, sculptured with broad, rather flattened ribs,
usually 6 to 8 in number, strongly denticulating the cutting mar-
gin. (PL 32, fig. 48, T. blomfieldi. PL 34, fig. 2, T. mitchellce.
PL 32, fig. 50, T. rainbirdi). The jaw of broadbenti has 11 ribs.
Radula having the central tooth smaller than the adjacent later-
als; central and lateral teeth unicuspid, the side cusps represented
by a lateral continuation of the reflection, being completely fused
with the median cusp. Transition teeth and inner marginals hav-
ing a long bifid inner cusp (entocone-f-niesocone) and a simple,
small ectocone. Outer laterals tricuspid (in fraseri, yulei, lessoni,
blomfieldi} or quadricuspid by splitting of the ectocone (incei,
rotteAeo5,pl.34,fig. 1.)
Genitalia lacking all accessory organs on the female side, the
duct of the spermatheca very long (pi. 33, fig. 3, s, s, s), its upper
portion narrow, lower portion stout or swollen. Penis large, club-
THERSITES. 133
shaped, the walls of its cavity granulated, having a large solid,
granulated papilla at the apex, near the entrance of the epiphallus
(pi. 33, fig. 3, papilla indicated by dotted line). Epiphallus
long, the penis retractor inserted at the proximal third of its length ;
ending in a flagellum. Penis retractor short, attached to floo*M>
the lung cavity. Right eye-peduncle retracted between primary
branches of genitalia. PI. 33, figs. 2, 3, H. mitchellce; fig. 2,
reverse view of vagina, showing lower course of uterus and vas
deferens. (From a specimen received from Dr. Cox).
T. mitchellce and broadbenti have the type of genitalia described
above, but in the latter the spermatheca has a shorter stalk. A
second type of genitalia is found in T. basalis (=rainbirdi'), T.
fraseri, T. blomfieldi in which species the epiphallus is extremely
short and the flagellum either extremely short or obsolete, evidently
by degeneration. Only by opening the penis can the true condition
of these organs be ascertained. (See pi. 32, fig. 51, T. rainbirdi,
after Hedley).
In anatomy, Sphcerospira agrees with Badistes and Thersites,
except that in some species the appendages of the penis have under-
gone degeneration resulting in secondary haplogonism. The group
inhabits Queensland with a few forms in New Guinea, being
replaced southward by Badistes, westward by Xanthomelon.
Most of the species of Sphcerospira live under the loose bark of
fallen trees and on the ground, and are gregarious. Some occur
under stones in damp places. No Hadras are arboreal, according
to Hedley; differing totally in this respect from Papuina, but
agreeing with Chloritis.
(Imperforate species').
T. fraseri Gray, vi, 150. T. croftoni Cox, vi, 153.
v. flavescens Hedl., vi, 151. T. blomfieldi Cox, vi, 154.
T. coarctata Fer., vi, 151. v. warroensis Hedl. & Mouss.,
T. zebina Braz., vi, 151. [viii, 281.
T. mossmani Braz., vi, 152. T. mitchella3 Cox, vi, 154.
T. coxi Crosse, vi, 152. T. gratiosa Cox., vi, 155.
forbesi Cox, preoc. T. etheridgei Braz., vi, 156.
cerea Cox, preoc. T. macleayi Cox, vi, 156.
cerata Cox. T. audersoni Cox, vi, 157.
(Umbilicated species).
T. rainbirdi Cox, vi, 157. T. rawnesleyi Cox, viii, 282.
basalis Mouss. T. barney i Cox, vi, 165.
134
THERSITES.
T. oconnellensis Cox, vi, 158. T.
albofilata Mouss. T.
T. arthuriana Cox, vi, 159. T.
T. rockhamptonensis Cox., vi, T.
planibasis Cox, ms. [159.
v. moresbyi Ang., vi, 160. T.
v. pallida Hedl. & Mss. viii, 281.
T. informis Mouss., viii, 282. T.
T. palmensis Braz., vi, 160. T.
v. meridionalis Braz., vi, 161. T.
T. bellendenkerensis Braz., vi, T.
[161.
T. parsoni Cox, vi, 162. T.
T. appendiculata Pfr., vi, 163. T.
T. seminigra Morel., vi, 162. T.
lessoni Pfr., olim., et auct. T.
? =incei var. T.
T. incei Pfr., vi, 166. T.
v. aureedensis Braz., viii, 282. T.
v. bayensis Braz., vi, 166 ; viii, T.
[282. T.
T. thatcheri Cox, vi, 164. T.
T. hilli Brazier, vi, 164. T.
mazee Braz., vi, 165.
hanni Braz., vi, 166.
prsetermissi Cox, vi, 167.
mulgravensis Braz., vi, 168.
mulgravei Braz.
curtisiana Pfr., vi, 168.
WaBraz., vi, 169.
johnstonei Braz., vi, 170.
creedi Cox, vi, 170.
wesselensis Cox, vi, 170.
sardalabiata Cox, vi, 171.
stephensoniana Braz.
whartoni Cox, vi, 171.
mourilyana Braz., vi, 172.
yulei Forbes, vi, 172.
challisi Cox, vi, 173.
nicomede Braz., vi, 173.
beddomse Braz., vi, 174.
bebias Braz., vi, 175.
cookensis Braz., vi, 175.
torasoni Braz., vi, 175.
broadbenti Braz., vi, 176.
hixoni Braz., vi, 177.
Subgenus XANTHOMELON v. Martens, 1860.
Xanthomelon MTS., in Alb., Die Hel., p. 174, type H.pomum;
Mai. Blatter xvi, p. 77, 1869. PILS., Man. Conch., vi,p. 178. For
anatomy, see SEMPER, Reisen, p. 160, pi. 14, and HEDLEY, P. R. S.
Q., vi, p. 250, pi. 14, and p. 121, pi. 8.
Shell large, solid and globular, the spire small, body-whorl large,
globose, descending to the aperture, which is semioval and some-
what oblique. Peristome narrowly expanded, thickened within ;
columellar lip broad, flattened, partly or wholly covering the axial
perforation; surface somewhat roughened, covered with a yellow
cuticle. Type T. pomum, pi. 27, fig. 6.
Jaw stout, arched, with 8 (perinflata) to a dozen (pachystyla) stout
ribs, obsolete toward the ends (pi. 32, fig. 47, pachystyla).
Radula as in SpJuerovpira etc. (pi. 32, fig. 46, pachystyla).
Genital system having the penis rather short and stout, twisted at
its apex, where the retractor-muscle and vas deferens are apparently
THERSITES. 135
inserted. Spermatheca duct short and arising high on the vagina
(pi. 32, fig. 52, pachystyla).
The shell is more globular than that of Hadra s. sir. or Sphcero-
spira, with smaller spire and wider columellar lip. The jaw and
teeth are not different from those of Sphcerospira, etc. The peculi^
arity of the genital system is the apparent obsolescence of the
epiphallus and flagellum, and the shortness of the duct of the sper-
matheca, which is, as a general rule, long in this genus and its
allies. Semper has investigated the anatomy of pachystyla, and
Hedley that of pachystyla and perinflata. The penis should be re-
examined, with a view to finding traces of the missing epiphallus
and flagellum, and the internal papilla.
The species inhabit Queensland, Arnhem land and the adjacent
parts of the northern territory of S. Australia. T. pachystyla is
found on sandy ridges buried a few inches below the surface among
the roots of bushes, in dry weather.
T. pomum Pfr., vi, 178. T. banneri MacGill. vi, 179.
urvillei H. & J. T. lyndi Angas, vi, 183.
pseudomeadei Braz. T. pachystyla Pfr. vi, 184.
? sphceroidea Le Guill. v. daemeli v. Mts. vi, 184.
T. nigrilabris v. Mts., vi, 179. T. jannellei Le Guill, vi, 182.
edwardsi Cox not Bid. pachystyloides Cox.
meadei Braz.
Subgenus RHAGADA Albers, 1860.
Rhagada ALB., Die Hel., 1860, p. 108, type H. reinga Gray.
PILSBRY, Man. Conch., vi, p. 184. WIEGMANN, Weber's Zool.
Ergebnisse einer Reise in Niederl. Ost-Ind. iii, p. 169 (anatomy).
Shell small, compact, globose-depressed, narrowly or covered um-
bilicated, rather solid and cretaceous, whitish, unicolored or rnulti-
lineate with reddish, the supraperipheral band most prominent and
constant ; periphery rounded ; outer lip more or less expanded and
thickened, columella reflexed, partly or wholly closing the umbili-
cus. Type T. reinga Gray, (see pi. 27, figs. 16, 17, 18, T. carcharias
Pfr. PL 27, figs. 11, 12, 13, T. supracostulata Schepm. PL 27,
figs. 14, 15, T. floresiana Martens).
Jaw (pi. 51, figs. 7, 8, T, solorensis) arcuate, with 4 or 5 unequal
and asymmetrically arranged strong ribs.
Radula (pi. 51, figs. 11, 12, T. solorensis') with 126-163 transverse
rows of 31. 1. 31 to 38. 1.38 teeth of the type usual in Chloritis and
136 THERSITES-PAPUINA.
Hadra. Central and inner lateral teeth having the ecto- and ento-
cones completely fused with the mesocones, which attain or project
beyond the posterior edges of the basal-plates. Outer laterals hav-
side side cusps developed, the raeso- and ento-cones forming a long
compound cusp as in Chloritis, etc. Marginal teeth (fig. 11) tricus-
pid, or having the ectocones bifid (figs. 11, 12 show central with two
adjacent laterals, 10th to 13th lateral and transition teeth, 22d, 23d
and 32d to 35th marginal teeth ; after Wiegmann. PL 51, fig. 9,
shows a central and lateral tooth from another individual, in which
the ectocones are developed). In T. convicta the jaw has 7 stout
ribs, dentition as in solorensis (See Binney, Dent. Pulm. Moll. pi. x,
f. G.)
Genital ia (pi. 51, fig. 10, T. solorensis, after a drawing by Mr.
A. Protz) with a short flagellum on the penis, the spermatheca-duct
inserted high on the vagina. No penis retractor is shown in the
sketch, but it is probably present; and it is likewise probable that
the penis proper terminates with the swollen portion seen at about
the middle of its length, and that it contains a papilla there; the
narrower upper part, as far as the entrance of the vas deferens, be-
ing an epiphallus.
The snails of this section have a smaller, more compact and cre-
taceous shell than Hadra, with a different scheme of color. The
anatomy offers no deviation of any importance from that of Hadra
and Chloritis.
(Species of N. Australian coast and adjacent islands').
T. reinga Gray, vi, 185. T. convicta Cox, vi, 187.
T. richardsoniiE. A.Sm., vi,185. T. plectilis Bens., vi, 188.
T. leptogramma Pfr., vi, 186. paleata Rve.
T. dringi Pfr., vi, 186. T. carcharias Pfr., vi, 189.
T. tescorum Bens., vi, 187. T. (?) torulus Fer. vi, 189.
T. elachystoma v. Mts., vi, 187.
(Species of So lor, Flores, and other islands N. of Timor Sea).
T. colona v. Mts., vi, 190. T. floresiana v. Mts., pi. 27, f. 14, 15.
T. solorensis v. Mts., vi, 190. T. supracostulata Schep., viii, 283.
Genus PAPUINA von Martens, 1860.
Papuina MTS., Die Hel. (2d edit.), p. 166, type H. lituus Less.
PILSBRY, Man. Conch., (2), vii, p. 3. Eugenia ALB. Mss. Insu-
PAPUINA. 137
laria TAP. CAN. Ann. Mus. Civ. Genov. xix, p. 115, 138, type H-
lituus, 1883. Pileolus LESSON, Voy. de la Coquille. Zool. ii, p.
313 (preoc.). Cymotropis v. Mart, Die Hel., p. 169, type H.
" vitrea" = antrorsa. Merope ALB., Die Hel., 2d edit., p. 158,
type H.fringilla (preoc.). Geotrochus of BECK and authors, not of
v. Hasselt. Acavus SMITH and TAP. CAN., not of Montf.
Shell turbinate, lens-shaped or trochiform, umbilicated or iraper-
forate, rather thin ; periphery varying from round to acutely keeled.
Surface smoothish, the coloring light or bright. Aperture oblique,
toothless or with a columellar nodule, the peristome thin and gen-
erally expanded, ends of the lip remote. Type P. lituus Less. pi.
29, fig. 12 (see also pi. 29, figs. 14, 15, P. trobriandensis. Fig. 11,
P. splendescens. Fig. 13, P. nortoni. PL 46, figs. 17-19, P. ianthe).
Animal with the foot rather short, sole undivided ; upper surface
densely granulated, with a slight median longitudinal groove above,
the tail densely granulose with no median groove. Mantle with a
triangular right lappet and an elongated low left one, the latter
emitting a lobe on the left side.
Jaw thin and weak, arcuate, its median portion ribbed, ends blunt
-and ribless. (PL 34, fig. 11, P. moseleyi. PI. 34, fig. 1 2, P. vexillaris.
PL 37, fig. 2, P. conscendens. PL 13, fig. 17, P. grata. PL 13, fig.
18, P. taumantias. PL 13, fig. 25, P. louisiadensis. PL 13, fig. 24,
P. boyeri. PL 13, fig. 26, P. brumeriensis. PL 13, fig. 28, P.
macgillivrayi.
Radula of two types. Typically, the transverse rows are nearly
straight ; the central and lateral teeth with wide, blunt mesocones,
shorter than the basal plates, the marginals with three short, wide
cusps (pi. 13, fig. 23, boyeri. PL 13, fig. 29, fringilla. PL 37, fig.
11, conscenden*). In P. moseleyi (pi. 37, fig. 1) the cusps are very
broad, and project beyond the basal plates.
In some divergent species the transverse rows of teeth are v-shaped;
central teeth (pi. 37, fig. 9), with an extremely broad, gouge-like
cusp (united meso- and ectocones), the laterals having the cusp par-
tially divided into entocone and mesocone, an ectocone appearing
on the outer laterals and marginals. The teeth are all of the same
general form, and in all the cusps project over the basal plates.
This type of teeth occurs in P. boivini and in vexillaris (pi. 37, figs.
9, 10), and will probably prove characteristic of the groups those
species belong to, and also of the P. meta group ; the other groups
having the more normal type of teeth. This aberrant type is com-
T7KITEB8ITT
138 PAPUINA.
parable to that of Polymita and Oxychona, and seems to be corre-
lated with arboreal habits. P. moseleyi bridges, to some extent, the
gap between the two types of teeth.
Genital system having no accessory organs on the female side,
the spermatheca on a rather long duct. Penis containing a papilla
at its apex, continued in a long epiphallus which bears the retractor,
and which passes into the vas deferens, having no flagellum or
merely the rudiment of one. (PI. 37, fig. 5, P. trobriandensis ; pi.
37, figs. 3, 4, P. vexillaris ; pi. 37, figs. 7, 8, P.fringilla; pi. 13, fig. 16,
P. grata ; pi. 13, fig. 21, P. yulensis ; pi. 13, fig. 27, P. brumeriensis).
In another series of species the penis is short, the epiphallus very
short, hardly distinguishable, ending in a short flagellum (pi. 13,
fig. 22, P. taumantias ; pi. 37, fig. 6, P. brazieri). Some of these
have the spermatheca duct very short.
The prominent features of the anatomy are the weakness of the
thin jaw, the breadth of the cusps of the teeth, and the lack of a
flagellum upon the epiphallus, or its shortness, the union of epiphal-
lus and vas deferens being indicated only by a slight protuberance
at the end of the former, in most species.
In some species (trobriandensis, woodlarkiana, moseleyi) the penis
is extremely small. In others (boyeri, louisiadensis, fringilla} it is
large and muscular. In one group of forms, taumantias, braziene,
tomasinelliaiia, gestroi, meditata, ridibunda, the epiphallus is reduced
to a very short extent, or even obsolete, and a short flagellum is
developed. There is, therefore, a wide range of variation in the soft
parts, as in the shells, of this genus.
In P.fringilla the papilla is extremely long, and the walls of the
penis cavity are transversely corrugated (pi. 37, fig. 7). In P.
vexillaris the papilla is large but short (pi. 37, fig. 4). The eye-
stalk is retracted between the branches of the genitalia, as usual.
In P.fringilla the left edge of the mantle bears a lobe, at the posi-
tion where two lobes are shown in Thersites (Badistes) gulosa.
The anatomy of many forms is now known : Binney has figured
the teeth of P. fringilla (Ann. N. Y. Acad. Ill, p. 113). Tappar-
one Ganefri has figured the genitalia of P. yulensis, kaiauensis, tau-
mantias, ridibunda, meditata, grata, novoguinecnsis, brazierce, gestroi,.
tomasinelliana (Ann. Mus. Civ. Genov. xix, pi. 6 and 7). Hedley
has illustrated the anatomy of P. brumeriensis, louisiadentif, rollsi-
ana, woodlarkiana, trobriandensis, and boyeri (Proc. Linn. Soc. N.
S. Wales (2), vi, pi. 38-42. Pfeffer has figured the anatomy of P.
PAPUINA. 139
boivini (Monatsber. Berl. Akad. Wissensch. 1877, p. 277, pi. 2, f. 11
-13). The writer has examined the soft parts of P. fringilla, vexil-
laris, moseleyi and conscendens.
Papuina is an exclusively arboreal genus, being strongly con-
trasted in this habit to its allies Thersites and Chloritis. The shell
is of lighter structure and brighter color than in these terrestrial
groups, somewhat approaching that of Cochlostyla a case of con-
vergence of external characters from similar habits. The teeth dif-
fer from those of allied groups, Thersites, Chloritis, Planispira, in
the great breadth and bluntness of the cusps, a structure correlated
with arboreal habits. The jaw is more delicate than in the allied
genera.
The great variation observed in the genitalia and teeth of the
species examined, shows that here lies a wide field for future cultiva-
tion. These features are no doubt characteristic of minor groups in
the genus, and their investigation will lead to valuable results in the
classification of the group, and secondarily may be of use in the
study of its geographical distribution and migrations. The arbo-
real habit has evidently been long established, for otherwise we
should not have so profound a remodeling of the dentition.
The geographic limits of the genus are on the northwest Halma-
heira, on the southeast, the New Hebrides group. There are two
principal centers of specific radiation: New. Guinea and the Solo-
mon archipelago. The former of these has peopled the Moluccas,
Queensland and the Louisiades. The species of Java, Sumatra and
India referred by authors to this genus belong to other groups*
mainly Satsuma.
Subdivisions.
Section Papuina. Shell having the outer lip well expanded,
baso-columellar lip reflexed.
Section Dendrotroehus. Shell trochoid, the columellar lip not in
the least expanded or reflexed.
Section Papuina (restricted).
(Group of P. boivini; Solomon and New Britain groups.)
P. congrua Pfr., vii, 4. P. hargreavesi Ang. vii, 9.
P. chancei Cox, vii, 5. hargravesi auct.
amphizona Pils., vii, 5. P. gamelia Ang., vii, 10.
140
PAPUINA.
P. boivini Petit, vii, 6.
subrepta H. & J.
color uta Mss.
P. ambrosia Ang., vii, 7.
ramsdeni Ang.
P. malantensis Ang., vii, 7.
P. philomela Ang., vii, 8.
P. brodiei Braz., vii, 10.
P. dampieri Ang., vii, 11.
P. walleri Braz., vii, 12.
brenchleyi Ang., not Braz.
P. alfredi Cox, vii, 12.
v. trichroa v. Mart., vii, 12.
P. macfarlanei Cox, vii, 13.
P. guadalcanarensis Cox, vii, 9. P. coxiana Ang., vii, 13.
(Group of P. meta; Solomon Is.).
P. xanthochila Pfr., vii, 15.
P. miser Cox, vii, 20.
beatrix Ang., vii, 15.
P. choiseulensis Braz., vii, 16.
P. spendescens Cox, vii, 16.
brenchleyi Braz., vii, 16.
mendana Ang., vii, 17.
P. meta Pfr., vii, 17.
deidamia Ang.
v. acrnella Pfr., vii, 18.
P. plagiostoma Pfr., vii, 19.
P. guppyi Smith, vii, 19.
P. adonis Angas, vii, 20.
metula Crosse.
P. blandaCox, vii, 21.
P. mendoza Braz., vii, 21.
P. hermione Ang., vii, 21.
biocheana Crosse.
P. migratoria Pfr. vii, 22.
leucophcea Cox.
(Group of P.flexilabris ; Solomons, Louisiades and New Ireland).
P. vexillaris Pfr., vii, 46.
phthisica Pfr.
P. boyeri C. & F., vii, 47.
P. phseostoma Mart., vii, 47.
P. gaberti Less., vii, 48.
trochus Q. & G.
trochoides Desh.
P. lambei Pfr., vii, 48.
lombei Pfr., olim.
P. flexilabris Pfr., vii, 49.
P. coniformis Fer., vii, 50.
turbinata Desh.
v. tuffetii Less., vii, 51.
P. sellersi Cox, vii, 51.
(Group of P. conscendens ; Queensland).
" A small group of Queensland snails seem to differ from the
main body of the genus in their habits. Not the stem or branches,
but the leaves of trees are chosen by these for their favorite abode.
To suit the situation the shell has been modified until the contour
would suggest Partula rather than Papuina. The more conical
shape has probably been adopted for greater safety in the exposed
tree tops ; to the same end every superfluous atom of weight has
PAPUINA.
141
been abandoned, the shell reduced to the thinnest, and the reflected
lip dispensed with." (Hedley, Nautilus, vii, p. 73).
P. fucata Pfr., vii, 14.
P. conscendens Cox, vii, 14.
( Group of P. pileus
P. euchroes Pfr., vii, 23.
P. pileus Mull., vii, 24.
pileata, bifasciata, ambigua
[Gmel.
P. blainvillei Le Guill, vii, 25.
gdrtneriana Pfr.
zoae Pfr.
P. folicola Hedley, Nautilus, I. c.
B. bidwilli Cox not Pfr.
Moluccas and New Guinea).
P. lenta Pfr., vii, 23.
P. canovarii Tap. Can., vii, 26.
P. blanfordiana H. Ad., vii, 26.
blanfordi H. Ad.
turbinata Val., mss.
v. poirieri Tap. Can., vii, 27.
(Group of P. poiretiana ; Night I., N. E. Australia).
P. poiretiana Pfr., vii, 27.
( Group of P. antiq^
P. antiqua Ad. & Kv., vii, 28.
Borneo ?, New Guinea).
P. leonardi Tap. Can., vii, 32.
horderi Sowb., vii, 29.
P. xanthosoma Pils., vii, 28.
(Group of P. pileolus; Moluccas and western New Guinea).
P. pileolus Fer., vii, 29.
/. turrita v. Mart.
/. pyramidata v. Mart.
/. convexa v. Mart.
/. depressa v. Mart.
P. rhynchostoma Pfr., vii, 30.
P. ferussaci Less., vii, 30.
( Group of P. vitrea :
P. vitrea Fer., vii, 33.
albula Le Guill.
vitracea Beck.
P. arrowensis Le Guill, vii, 34
P. chondrodes Strub., viii, 292
P. lanceolata Pfr., vii, 34.
P. grata Mich., vii, 35.
acuta Q. & G., not Lam.
P. leucotropis Pfr., vii, 36.
P. hero Smith, vii, 57.
P. exsultans Tap. Can., vii, 31.
ferussaci Pfr., Novit. Conch.
P. hedleyi Smith, viii, 290.
P. pythonissa Tap. Can., vii, 31.
P. turris H. Ad., vii, 32.
P. fergusoni H. Ad., vii, 32.
P. steursiana Shutt, vii, 33.
Moluccas and New Guinea).
P. ianthe Smith, vii, 58.
P. nodifera Pfr., vii, 37.
P. pelechystoma Tap. Can., vii, 35
P. pennantiana Pfr., vii, 36.
P. carinata Hombr. & Jacq.vii, 36.
P. bevani Braz., viii, 292.
P. ? elisus Hedl., viii, p. 292.
P. ? goldiei Braz., vi, 217.
oxystoma Smith (preoc.).
P. ? tritonensis Le Guill., vii, 88.
142
PAPUINA.
( Group of P. labium : Papuan region).
P. lituus Less., vii, 37.
ardouini Dh.
papuensis Q. &. G.
P. labium Fer., vii, 38.
P. pseudolabium Pfr., vii, 38.
P. multizona Less., vii, 39.
tenuiradiata Q. & G.
multizonata Desh.
spectrum Rve.
P. taumantias Tap. Can., vii, 39.
v. cingulata Hedl., viii, 288.
P. ridibunda Tap. Can., vii, 40.
P. sicula Braz., vii, 45.
meditata Tap. Can., vii, 40.
P. aurora Pfr., vii, 41.
P. serope Smith., vii, 41.
P. novoguineensis Pfr. vii, 42.
v. triumphalis Rve, vii, 42.
v. mysolensis Pfr., vii, 43.
P. waighouensis H. Ad., vii, 43.
P. brazierse Braz., vii, 43.
v. lacteolota Smith, vii, 25.
P. tomasinelliana T. C. vii, 44.
v. anozonata Hedl., viii, 288.
v. agnocheilus Smith, viii, 289.
P. gestroi Tap. Can., vii, 44.
P. maclayana Braz., vii, 45.
(Group of P. louisiadensis : Louisiades, d'Entrecasteaux Is., and
British New Guinea).
P. tayloriaua Ad. & Rv., vii, 58. P. louisiadensis Forbes, vii, 61.
yulensis Braz.
P. strabo Braz., vii, 60.
roseolabiaia Smith.
katauensis T. C.
P. gorenduensis Braz., vii, 63.
P. rollsiana Smith, vii, 63.
P. comriei Ang., vii, 64.
v. millicentse Cox, vii, 62.
v. thomsoni Smith, vii, 62.
P. gurgustii Cox, vii, 61.
P. rhombostoma Pfr., vii, 60.
P. woodlarkiana Souv., vii, 62.
P. trobriandensis Hedl., viii, 290.
P. albocarinata Smith, vii, 59.
(Group of P. brumeriensis: Eastern New Guinea).
P. chapmani Cox, vii, 51. P. zeno Braz., vii, 53.
coraliolabris Smith,
P. brumeriensis Forbes, vii, 52.
v. albolabris Hedl., viii, 289.
P. rangii Less., vii, 53.
extricanda Tap. Can.
(Australian Species').
P. macgillivrayi Forbes, vii, 55. P. bidwilli, Pfr., vii, 55.
P. cerea Hedl. bridwilli Pfr., olim.
latiaxis Smith.
P. diomedes Bras., vii, 54.
P. naso v. Mart., vii, 56.
tapparonei Smith.
P. rhyuchonella Tap. Can., vii, 57.
(Group of P. eddystonensis).
P. eddystonensis Reeve, vii, 64. P. nortoni Braz.
P. motacilla Pfr., vii, 66. P. cserulescens Ang., vii, 68.
PAPUINA-PLECTOPYLIS. 143
P. gelata Cox, vii, 65. P. pudica Pfr., vii, 69.
v. maddocksi Braz., vii, 66. P. lienardiana Crosse, vii, 69.
P. antrorsa Pfr., vii, 67. P. eros Angas, vii, 70.
vitrea v. Mart., olim. P. redempta Cox, vii, 70.
P. sachalensis Pfr., vii, 67. P. nigrofasciata Pfr., vii, 71.
P. leucothoe Pfr., vii, 68. P. donnaisabellse Ang., vii, 71.
(Group of P. moseleyi).
P. moseleyi Smith, vii, 72. P. novsegeorgiensis Cox, vii, 72.
( Group of P. fringilla).
P. fringilla Pfr., vii, 73. P. barnaclei Smith, vii, 73.
Section DENDROTROCHUS Pilsbry, 1894.
Papuina with the shell imperforate, trochiform, with rhombic aper-
ture, the lip thickened within ; columellar lip not expanded or re-
flexed. Type P. helicinoides Hombr. & Jacq.
Soft anatomy unknown. Distribution Solomon Is., New Hebrides,
Admiralty Is. and New Ireland. They are arboreal in habit. Bra-
zier found P. cyrene in hundreds on the under sides of leaves of
small bushes, in Ugi, Solomon Is.
This is quite a well characterized section of Papuina. According
to Hedley the Solomon Islands forms (cleryi, quirosi, zelina, cyrene)
will prove to be varieties of one species (see Man. Conch., viii, p.
290).
P. labillardierei Smith, vii, 75. P. cineracea H. & J., vii, 77.
P. helicinoides H. & J., vii, 76. cinerarea Rouss.
v. cleryi Reel., vii, 76. P. cyrene Crosse, vii, 78.
septentrionalis Sm. P. eva Pfr., vii, 78.
v. ineridionalis Sm., vii, 77. P. layardi Hartm., vii, 79.
v. quirosi Cox, vii, 80. P. pyxis Hinds, vii, 80.
P. zelina Cox, vii, 78. P. crucibulum Pfr., vii, 81.
Genus PLECTOPYLIS Benson, 1860.
Plectopylis BENS., Ann. and Mag. N. H. (3), v, p. 243. STOLICZKA,
Journ. Asiat. Soc. Beng. xl, (2), p. 217, pi. 15 (anatomy). GODWIN-
AUSTEN, P. Z. S. 1874, p. 608.
Shell depressed, with flat or low-eonimi spire and large umbilicus,
dextral or sinistral ; solid or thin, the upper surface generally sculpt-
144 PLECTOPYLIS.
ured with spiral lines, hirsute in the young. Aperture half-round
or lunate, oblique, the lip reflexed, its ends generally joined by an
elevated parietal callus, which usually bears an entering lamella.
Interior of the last whorl obstructed by a barrier composed of a trans-
verse plate or plates on the parietal wall, and several transverse or
longitudinal denticles or plates on the outer wall. Type P. achatina
Gray, pi. 40, figs. 5, 6, 7, 8. (See also pi. 40, figs. 1-4, P. jovia.
PL 40, figs. 9-12, P. ponsonbyi. PI. 40, figs. 13-15, P. fultoni).
Foot short, rarely equalling in length the diameter of the shell ;.
tentacles very short ; eye pedicles of moderate length. Mantle edge
thin, with small right and left body-lappets. Pulmonary cavity
small. Kidney large, triangular.
Jaw very thin, horny, arched, with a small anterior median pro-
jection ; it is marked transversely with a great number of more or
less distant grooves which divaricate in the center (pi. 42, fig. 36. P.
cyclaspis). Radula of moderate width, long, composed of about 100
transverse more or less V-shaped rows of 60-70 teeth. Central tooth
smaller, sometimes much smaller, than the laterals, very narrow, the
reflection small, with three slender cusps. Lateral teeth with a large
inner cusp and simple or bifid outer cusp, and a minute inner cusp
(pi. 42. fig. 35, P. cyclaspis central, 1st, 2d and 12th laterals, and
20th and 25th marginal teeth. In P. pinacis the central tooth is
larger and more similar to the laterals.
Genital system (pi. 42. fig. 34, P. cyclaspis) having the duct of the
spermatheca long. An organ of unknown homology (either a dart
sack, a diverticulum of the spermatheca, or an appendicula) enters
the vagina just above the opening of the spermatheca duct. Uterus
containing few large eggs. Penis simple, receiving the vas deferens
and the penis retractor at its apex, the latter attached distally to the
floor of the lung cavity.
This group differs from Gorilla in having perpendicular internal
lamellae upon the parietal wall of the shell. It is different from Gor-
illa and all other Helices in the converging V like elements of the
thin jaw, which is quite of the goniognathous type found in Gylin-
drella, Orthalicus and Otostomus. The small size of the central
teeth is also au anomalous feature, recalling the Achatinidce. Per-
haps the accessory organ of the vagina (seen between the uterus and
the spermatheca in pi. 42, fig. 34) is really a diverticulum of the
spermatheca duct ; and if this is the case the genital organs will not
differ very much from those of Gorilla, although in that genus the
PLECTOPYLIS. 145
retractor of the penis is attached to the uterus wall (as in Hyalo-
sagda) instead of to the lung floor.
The apex of the shell is rather large, as in Corilla, and usually
somewhat rugose. The grouping of the species is based upon the
form of the internal barrier, which is sometimes simple (pi. 40, fig._
4), sometimes excessively complex by the duplication of the parietal
and palatal barrier (pi. 40, figs. 7, 8, 12). Godwin-Austen found
shells with insects fixed between the teeth, so that there can be little
doubt that this barrier has been evolved for the protection of the
snail from predatory insects which swarm in the regions these forms
inhabit.
P. achatina and cyclaspis are found on limestone hills, the animal
being shy, usually living in crevices and holes, and closely adhering
to the rock, even when moving about.
This genus inhabits India and Farther India, extending north to
central China and south to Ceylon and the Philippine Islands.
Subdivisions.
Pledopylis is herein expanded to contain two Chinese groups of
uncertain affinities, besides the typical group.
Subgenus PLECTOPYLIS. Whorls regular, the last not distorted
nor grooved outside ; having internal transverse barriers within the
last whorl.
Subgenus TRAUMATOPHORA. Latter part of last whorl contracted
outside; throat obstructed by entering palatal lamellae, but having
no internal processes on the parietal wall.
Subgenus STEGODERA. Shell sinistral, the last whorl distorted
straightened, embracing the preceding ; aperture crescentic, tooth-
less ; throat very narrow, but without internal teeth or lamellae.
Subgenus PLECTOPYLIS Bens.
(Parietal vertical lamina double or compound).
P. achatina Gray, iii, 165. P. refuga Old., iii, 164.
trepercussa Gld. P. dextrorsa G.-Aust., iii, 164.
P. anguina Gld., iii, 165. P. leiophis Bens., iii, 163.
P. brahma G.-Aust. iii, 164. P. shiroensis G.-Aust., iii, 163.
P. cyclaspis Bens., iii, 164. P. feddeni Blanf., iii, 163.
catinus Bens. olim. P. brachyplecta Bens., iii, 163.
P. karenorum Blanf., iii, 164. P. biforis Hde., iii, 166.
P. revoluta Pfr., Mon., v, 416. P. ponsonbyi G. A.
10
IJNIVEB8IT1
V
146
PLECTOPYLIS.
(Parietal vertical lamina single).
P. shanensis Stol., iii, 162. P. brachydiscus G.-Aust., iii, 162.
trilamellaris G.-A. P. pseudophis W. Blanf.,iii, 162.
P. perarcta Blanf., iii, 162. P. nagsensis G.-Aust., iii, 161.
P. retifera Pfr., iii, 161.
P. clathratula Pfr., iii, 161.
puteolus Bens.
P. fultoni G.-A., viii, 296.
P. andersoni W. Blf., iii, 161.
P. plectostoma Bens., iii, 160.
prodigium Bens. mss.
P. macromphalus W. Blf., iii, 160.
P. laomontana Pfr., iii, 160. P. munipurensis G.-A., iii, 160.
P. schistoptychia MlldfF., iii, 165. P. pinacis Bens., iii, 159.
P. diptychia Mlldff., iii, 158.
P. polyptychia Mlldff., J. B.
[xiv, 272.
P. trochospira Mlldff. J. B., xiv,
[273.
P. schlumbergeri Mori., iii, 166.
P. joviaMab., viii, 156.
P. villedaryi Anc., viii, 157.
P. phlyaria Mab., viii, 158.
P. fimbriosa v. Mart., iii, 158.
v. emoriens Gred., iii, 158.
v. nana Mlldff, iii, 158.
P. murata Hde., iii, 159.
P. stenochila Mlldff., iii, 159.
P. pettos v. Mart., iii, 156.
P. oglei G.-Aust., iii, 159.
P. serica G.-Aust., iii, 159.
P. coarctata Mlldff. Nachrbl. '94,
[104.
P. pulvinaris Gld., iii, 157.
P. jugatoria Anc., iii, 166.
P. reserata Hde., iii, 166.
P. raultispira Mlldff, iii, 158.
P. cutisculpta Mlldff, iii, 158.
P. invia Hde., iii, 165.
P. secura Hde., Fl. Bleu, 141.
P. laminifera Mlldff, iii, 165.
Subgenus TRAUMATOPHORA Ancey, 1887.
Traumatophora ANC., Conch. Excb., April, 1887, p. 54.
Shell disk-shaped, with low spire and open umbilicus ; granulate.
Whorls 5, regularly increasing, the last constricted behind the -aper-
ture. Aperture lunar, oblique, with reflexed lip, having within
three entering lamellae upon the outer lip, marked outside by grooves,
no parietal processes. Type P. triscalpta, pi. 41, figs. 26, 27.
Anatomy unknown. This group and the next differ from Plecto-
pylism lacking transverse internal barriers, but until their soft parts
are known they had better be grouped in this place.
P. triscalpta v. Mart., vi, 8. Central-southern China.
v. fraterminor Gredl. J. B. xi, 137.
GORILLA. 147
Subgenus STEGODERA v. Martens, 1876.
Stegodera MTS., Novit. Conch., iv, p. 150. PILSBRY, Man. vi, p.
7. Steganodera KOBELT, Illust. Conchylienbuch, p. 236.
Shell sinistral, disk-shaped, with low spire and open, deep umbil-
icus ; solid, opaque, brown. Inner whorls slowly increasing,
regular ; latter half of the last whorl distorted, straightened, covering
the preceding whorl above. Aperture very oblique, crescentic, tooth-
less ; peristome reflexed ; throat very much contracted. Type P. an-
gusticollis, pi. 41, figs. 28, 29.
Anatomy unknown. A single species is known.
P. angusticollis v. Mart., vi, 7. Central China.
Genus CORILLA H. & A. Adams, 1858.
Gorilla ADS., Gen. Kec. Moll., ii, p. 208. SEMPER, Reisen (2),
iii, p. 100 (Anatomy). Atopa ALBERS, Die Hel., p. 90 (in part).
Shell planorboid, with nearly plane spire and broadly open umbil-
icus, the contour subcircular or oblong; rather solid, striated above,
brown or yellow. Whorls 5-5i, the last deflexed in front. Aper-
ture oblique, the lip broadly reflexed or recurved, its ends distant ;
parietal wall smooth or armed with a strong entering lamina. Inte-
rior of the last whorl either without laminse, or obstructed by a
series of blades nearly parallel to the direction oj the whorl, but hav-
ing no transverse barriers. Type C. erronea Alb., pi. 41, fig. 19.
See also pi. 41, figs. 20, 21, 22, C. rivolii Desh. PI. 41, figs. 23, 24,
25, C. charpentieri var. hinidunensis).
Foot (of C. erronea) with undivided sole and without pedal
grooves. No mantle lappets. Kidney very short.
Jaw entirely smooth. Radula with about 79-85 teeth in a trans-
verse row. Central tooth not smaller than the laterals, having a
single cusp, shorter than the basal-plate. Laterals similar but
asymmetrical. Marginals having a large, simple, oblique cusp
longer than the square basal-plate (pi. 42, fig. 37, central, 1st and
24th teeth of C. erronea].
Genital system elongated, with no accessory organs on the female
side. Spermatheca having a long duct, which branches into a very
long flagellum-like diverticulum, containing a cylindrical spermato-
phore, which extended from the end of the diverticulum to the
vagina. Penis short, swollen distally, continued in the vas deferens
upon which the penis retractor is situated, the distal end of the
148 CORILLA.
retractor being inserted on the uterus (pi. 42, fig. 38, C. erronea).
This species is ovoviviparous, the uterus in the individual figured
containing two young, having a membranous shell of about 5 mill,
diam., and more than 3 whorls.
The shell differs from that of Pleetopylis in lacking internal barriers
transversely obstructing the passage. When internal lamellae are
present in Gorilla they run parallel to the sutures or nearly so, as in
Polygyratia. The central teeth are not smaller than the laterals as
in Pleetopylis, and there are further differences in the genitalia. All
of the species are from Ceylon.
( Group of C. erronea : Parietal fold and internal plicae present).
C. erronea Alb., iii, 157. C. anax Bens., iii, 157.
C. rivolii Desh., iii, 156. C. odontophora Bens., iii, 157.
carabinata Fer.
(Group of C. charpentieri : no parietal fold or internal plicae).
C. charpentieri Pfr., iii, 156. C. humberti Brot, iii, 156.
v. hinidunensis Nev.
The nine genera following possess certain features in common,
binding them into a great group which the writer, in 1890, named
MACROON. The literature throwing light upon the anatomy and
affinities of the members of this super-generic group is very re-
stricted, three authors only having discussed them from the broad
standpoint of modern Helicology. SEMPER, in 1873, recognized
the alliance between Aeavus and Panda (with which he also groups
Gorilla and Caryodes) shown in the short kidney, simple genitalia,
smooth jaw and unicuspid teeth. PiLSBRY,in 1890, announced that
Aeavus, Panda, Helicophanta and Stylodonta agreed in having eggs
of extraordinary size, in which the young undergo prolonged ante-
natal development, and the shell actually attains a moiety of its
whorls before the independent existence of the creature begins.
HEDLEY, in 1892, studied the Australian forms, directing attention
to features of their eggs, embryonic shells and anatomy not before
appreciated, and gathering into one assemblage Panda, Pedinogyra,
Caryodes, Anoglypta (and Liparus).
The group, as it is herein understood, contains snails with helicoid
or bulimoid shells, viviparous or with large, hard-shelled eggs ; the
STYLODONTA. 149
jaw strong and ribless ; all of the teeth of the radula unicuspid ; the
genital system without flagellum on penis and with no dart sac or
mucous glands on vagina. To these characters we may add that
the transverse rows on the radula are moderately straight (not V--
shaped), the basal-plates of all the teeth are of the usual quadrate
form, and the large embryonic shell is usually sculptured differently
from the after-growth. The shell never has teeth or folds in the
aperture, although the columella shows often a long spiral, produc-
ing a visible sinuosity or truncation below, which, incidentally, in
some depressed forms, is shortened into a tooth-like columellar pro-
cess.
The affinities of the genera Plectopylis and Gorilla may be with
this phylum, but if so, the connection is so remote or so much ob-
scured by special modifications, that they may better be left isolated
until more fully understood. The Adams brothers, Try on, Fischer
and others, guided by certain analogies in the shells, have classified
these Indian genera with the American Polygyras and the Eur-Asian
Gonostomas, but the group so constructed is shown by a study of
the soft parts to be a house built upon the sand.
Genus STYLODONTA Crist, and Jan, 1832.
Stylodonta DE CRISTOF. et JAN., Catal. p. 2, type H. unidentata.
PILSBRY, Man. Conch., vi, p. 85. Stylodon BECK, Index Moll.,
p. 46. ALB.-MART., Die Hel., p. 149 (in part). Columplica
HARTM. (part) Gast. Schweiz, p. 187. Pachya ALB., Die Hel., p.
107 (in part). For anatomy see W. G. BINNEY, Ann. N. Y. Acad.
Sci., iii, p. 110 (teeth and jaw of Studeriana). VIGUIER, Arch.
Zool. Exper. et Generale, viii, p. 529, pi. 40 (genitalia of Studeriana).
SCHACKO, in Mobius' Beitr. zur Meeresfauna Maurit. u. Seychellen,
p. 342 (anatomy of unidentata}. MARTENS in v. d. Decken's
Reisen in Ost-Afrika, iii, i, p. 56, pi. 1 (varieties of unidentata).
NEVILL, P. Z. S., 1869, p. 61 (conditions of snail life on Seychelles).
Shell depressed- turbinate, solid, with imperforate axis at all stages
of growth ; obtuse apex ; and keeled periphery, at least in the young.
Surface yellowish or dark brown ; whorls 5, the earlier 3 spirally
grooved or decussated, forming an embryonic shell about one-third the
diameter of the adult ; outer whorls finely wrinkled, the last descend-
ing in front. Aperture wide-lunate, quite oblique, the peristome
expanded or reflexed. Columella short, subvertical, its inner edge
with a convex lobe or a sharp tooth-like fold. Type, S. unidentata,
pi. 38, fig. 9.
150 STYLODONTA.
Jaw stout, arched, with no ribs, but having a few coarse, broad
vertical wrinkles (Studeriana), or weak, fine and close striae (uni-
dentata}.
Radula large, composed of nearly straight transverse rows of
teeth. Central teeth with one broadly rounded cusp shorter than the
basal plates ; laterals similar, but the cusp longer and inclined ;
marginals having an inclined broad mesocone and developing a small
ectocone (pi. 48, fig. 9, S. studeriana).
In studeriana the radula measures 12J x 5 mill., and the formula
of teeth is 47.22.1.22.47. In unidentata the radula measures 10 x 4
mill., and the formula is 43.17.1.17.43.
Genitalia imperfectly known by Viguier's figures and description
(see pi. 50, figs. 6, 7, 9, S. studeriana'). The figures show the male
system below, female system above. The penis is large. Vagina
long, bearing a long duct ending in an oval spermatheca. Uterus
large, containing two yoang shells, which are enclosed in membran-
ous pouches, and attached to them by a sort of umbilical cord
formed of the pouch wall (pi. 50, fig. 9). The uterus and pouches
are filled with a glairy substance, probably nutritive, and secreted
by the albumen gland. The organs above the uterus are un-
known.
Large, solid Helices, inhabiting the Seychelles Islands. They are
viviparous, bringing forth one or two young at a time. They live
on bushesor climbing vines,feedupon green leaves,and aBstivate under
the soil or in rock crevices. Dufo supposed that only the lighter
colored individuals were females, but his observations lack anatomi-
cal confirmation. The shells are very dull colored for arboreal forms.
Besides the viviparous reproduction, and the large size of the young
at birth, these species are peculiar in having a small ectocone de-
veloped on the outer marginal teeth, the dentition being otherwise
like that of Aeavas. The large size of embryonal shell, the ribless
jaw, and the peculiar teeth, all forbid the association of these shells
with the genus Camcena. S. studeriana is found upon the island of
Praslin only. It lives upon the leaves and trunk of the Coco-de-mer
tree. S. unidentata occurs on Mahe, Felicite, Silhouette and Curi-
euse Islands. The young shells are acutely keeled, as in Pyrochilus t
not rounded as in Helicophanta, Acavus, Panda, etc.
S. uuidentata Chemn., vi, 86. S. unidentata Chemn.
microdonta Dh. v. exanthematica v. Mts.
uniplieata Hartm. v. militaris Pfr.
normalis Martens. v. globata v. Mts.
HELICOPHANTA. 151
S. studeriana Fer. vi, 87.'
Genus HELICOPHANTA Ferussac, 1821.
Helicophanta FER. (in part), Tab. Syst. des Anim. Moll., p.
xxxii ; Tab. Syst. de la Fam. des Lin^ons, p. 23, 25 (contains,
premier groupe Yitrinoides, H. brevipes, H. rufa and deuxeme
groupe Vesiculse, H. cafra, H. cornu-giganteum, H. magnified).
BECK, Index, p. 46 (except first species). ALBERS, Die He!., p.
110 (in part). ALB. -MART., Die Hel., p. 148, type H. magnified.
Leiostoma SWAINS (in part) Malacol., p. 328. Eurycratera H. &
A. Ad., Gen. Rec. Moll., ii, p. 190.
Shell large, capacious, Helicoid or bulimiform, imperforate or
umbilicate, consisting of 4-5 rapidly enlarging whorls, the several
earlier forming the embryonic shell, the diameter of which exceeds
one-third that of the adult ; the post-embryonic growth consisting of
1J whorls or less, the last very large, deflexed in front. Aperture
large ; lip narrowly expanded or reflexed, the columellar margin
dilated at its insertion. Type H. magnified Fer. (see pi. 38, fig. 4,
H. goudotiana. PI. 38, fig. 5, H. cornugigantewii) .
Animal completely retractile into the shell, having the foot large
and fleshy, sole not divided longitudinally, transversely wrinkled :
sides of foot granular and obliquely deeply grooved down to the sole
edge ; tail rounded behind, smooth above. Mantle margin enor-
mously thickened, having small right and left body-lobes (pi. 49, fig.
23, showing animal of H. magnified completely retracted within the
aperture, the end of tail visible in the mantle cavity).
Jaw strong, slightly arched, smooth ; having no trace of vertical
strise (PI. 49, fig. 19, H. magnified).
Radula resembling that of Acavus, Ampelita, and especially
Panda. Central and lateral teeth with single cusps, shorter than
the basal-plates ; marginals with long, oblique cusps (pi. 48, figs. 12,
.13, H. magnified, central with adjacent lateral, a lateral, and group
of marginals, with outermost marginal tooth).
Genitalia opening near the right eye-tentacle. Penis very large
and flattened, the retractor attached midway its length and inserted
distally on the lung floor ; vas deferens entering beyond the insertion
of the retractor, and continued inside in a vesicular enlargement of
the penis-wall to the apex of penis, where it opens into the large,
smooth-walled penis-cavity. Vagina large, short; uterus large.
152 HELICOPHANTA.
Spermatheca with a long duct, closely bound to the vagina (PI
49, fig. 21, H. magnified showing penis and lower portion of uterus
and spermatheca duct. Fig. 20, reverse side of female side, show-
ing vas deferens, etc. Fig. 22, penis split along line a-b in
fig. 21, showing cavity. Fig. 23, section of enlarged wall of penis
along Mne c-d of fig. 22, showing vesicular structure beyond the
entrance of the vas deferens. PI. 49, fig. 18, genitalia of H. goudo-
tiana, after Brancsik.
The peculiar features of this genus are the very large size of the
shell and its extremely large embryonal or nuclear portion ; the
great thickness of the fleshy mantle-edge, and the peculiar structure
of the penis. The unicuspid teeth of the entire radula, and the
smooth jaw, are characters common to Acavus, Panda, and other
allied genera. The species are restricted to Madagascar, but the
affinities of the genus are entirely with groups of the Seychelles
Islands and Ceylon. It is not yet known whether the young are
brought forth alive as in Stylodonta, or in eggs as in Acavus, Panda
and Borus.
That Ferussac intended his group Helicophanta especially for the
glassy, vitrinoid shells subsequently called Daudebardia, is evident
from his definition, grouping, and the etymology given ; and some
authors have used the name for these forms. But as Beck, Albers
and v. Martens have chosen another course, it seems advisable to
follow the precedent of such high authorities, especially since, by
the " law of elimination," the same result is obtained. The only
other course open to us would be to replace Daudebardia by Heli-
cophanta, and use the term Macroon in a restricted generic sense for
this Madagascar group.
The anatomy of Helicophanta is known by Seinper's description
of H. magnifica (Nachrbl., 1880, p. 60), and by a figure of the geni-
talia of H. goudotiana by Brancsik (Jahresheft des Naturwissen-
schaftlichen Vereines des Trencsener Comitates, 1892-3, p. 209, pi.
6, f. 5. The writer has examined H. magnifica in the flesh, and the
figures on plates 48 and 49 are drawn from this specimen.
Group oj cornugiganteum
H.cornugiganteum Chemn.vi, 60. H. guestieriana Cr., vi, 62.
H. betsileoensis Ang., vi, 61. H. vesicalis Lam.
H. ibaraoensis Ang., vi, 61. bicingulata Smith, vi, 63.
ACAVUS. 153
Group of magnified.
H. magnifica Fer., vi, 65. H. souverbiana Fisch., vi, 66.
polyzonalis Beck. /. audeberti Mouss., vi, 67.
Group of goudotiana.
H. oviformis Grat., vi, 68. H. grandidieri C. & F., vi, 72.
v. phenax Pils., vi, 69. H. partuliformis Bttg., vi, 72.
H. goudotiana Fer., vi, 70. H. oomorpha Mab., vi, 49.
H. echinophora Fer., vi, 71.
Group offarafanga, etc.
H. farafanga Ang., vi, 73. H. gloriosa Pfr., vi, 68.
farafanganensis C. & F. H. (?) follis Fer., vi, 74.
Genus ACAVUS Montfort, 1810.
Acavus MONTF., Conch. Syst., ii, p. 234, type H. hcemastoma.
SEMPER, Reisen, p. 9 (anatomy). SARASIN, Ergeb. Naturwissen-
sch. Forsch. auf Ceylon, i, 1888 (embryology). BINNEY, Ann.N.
Y. Acad. Sci., iii, p. 92 (dentition). Otala (in part) SCHUMACHER.
Oligospira ANCEY, Conch. Exch.,ii, p. 22, 1887, types H.waltoni
and H. skinner i.
Shell imperforate, globose depressed or globose-trochoidal, solid,
bright colored. Whorls less than 5, rapidly increasing, the several
earlier forming the nuclear or embryonic shell, which is about one-
third the diameter of the adult. Last whorl deflexed in front. Aper-
ture very oblique, the lip vividly colored and broadly expanded;
columellar margin long, obliquely descending, broadly flattened, the
columellar lip adnate. Type A. hcemastoma, pi. 38, fig. 1.
Animal with undivided sole, and no pedal grooves ; lung and kid-
ney very short, the latter opening at the base of the kidney. Body-
lobes of the mantle present, of moderate or small size.
Jaw strong, low-arcuate, entirely smooth, without median projec-
tion.
Radula having the teeth all unicuspid (pi. 50, fig. 8, 26, .4. slcinneri.
PI. 50, fig. 5, A. phoenix. PI. 48, fig. 14, A. hcemastoma.
Genital system having no accessory organs. Penis having termir
nal retractor, the interior with two longitudinal pilasters below, with
a very short, imperforate papilla at their base, at the base of which
the vas deferens enters. Spermatheca on a very short duct (PI. 50,
154 PYROCHILUS.
fig. 1, A. skinneri. Fig. 3, A. hcemastoma). Eggs very large, oval,
hard shelled (pi. 50, fig. 4, A. phcejiix, natural size).
The section Acavus comprises Ceylonese Helices of large size and
superb coloring. The shell is capacious, with a broad, polished lip
of vivid red, lilac, or intense black hue. The young shells at the
time of their extrusion from the egg are bright colored, with round
periphery, and are about one-third the size of the adult. The teeth
are all unicuspid, but the marginals have shorter cusps than in Heli-
cophanta or Panda ; and the shell differs from these groups in its
broad columeliar lip and brilliant coloring. They are arboreal in
habit.
Group of A. hcemastomus.
A. hsemastomus L., vi, 78. A. prosperus Alb., vi, 80.
v. melanotragus Born., vi, 79. A. phoenix Pfr., vi, 80.
v. conus Pils., vi, 79. A. superbus Pfr., vi, 81.
v. concolor, Pils., vi, 303. v. roseolabiata Nev., vi, 82.
A. fastosus Alb., vi, 79. v. grevillei Pfr., vi, 82.
Group of A. valtoni (Oligospira).
A. valtoni Rve., vi, 83. A skinneri Reeve, vi, 84.
waltoni auct.
Genus PYROCHILUS Pilsbry, 1892.
Phania ALB., Die Hel., edit. Martens, p. 157, type H. lampas.
MARTENS Landschn. der Ostasiat. Exped., p. 325. PILSBRY, Man.
Conch., vi, p. 193. Not Phania Meigen, Syst. Beschreib. Eur.
zweifliigel. Insekten, iv. p. 218, 1824. Pyrochilus PILSBRY, Proc.
Acad. Nat. Sci., Phila., 1892, p. 391.
Shell large, solid, imperforate, depressed ; keeled at the periphery,
at least in the young', convex above and below, unicolored. Junc-
tion of nuclear shell with the after-growth not distinct. Lip expanded,
bright colored ; columella widened into a flat plate, adnate over the
umbilicus, its inner edge blade-like. Whorls about 4i. Type P.
pyrostoma (see pi. 38, figs. 2, 3, P. lampas).
Jaw of H. pyrostoma smooth, weakly arched, without median pro-
jection. Animal without caudal gland or mantle lobes. Internal
anatomy unknown.
A group of handsome, large helices, all of which are still rare in
collections. The brilliant coloring of the peristome and the widened
AMPELITA. 155
columella, as well as the smooth jaw, are characters which Pyrochi-
lus shares with Aeavus ; but in the present group the embryonic
shell is not differentiated or demarked from the post-natal portion,
as is the oblong, globose nuclear shell of Aeavus ; and the young are
acutely keeled, as in Camcena.
The few species are from Halmahera and Bat j an, Moluccas.
P. lampas Mull., vi, 194. P. pyrostoma Fer., vi, 194.
carina Wood. v. bucculenta Tap.-Can., vi, 195
magna Schum. v. extincta Tap.-Can., vi, 195.
gigas Swains. P. xanthostoma Herk., vi, 197.
P. sulcocinctus Mart, vi, 196.
Genus AMPELITA Beck, 1837.
Ampelita BECK, Index Moll. p. 30 (proposed for zodiaca, xystera,
labrella, lancula, madagascariensis, clotho, aledo*). ALBERS, Die
Hel., 2d edit, p. 163. PILSBRY, Man. Conch., vi, p. 16.
Shell depressed, solid and opaque, varying from broadly openly
.umbilicated to perforate ; spire low, convex ; the periphery rounded
or keeled. Surface smoothish, sometimes malleated. Aperture
very oblique, oblong-truncate ; lip expanded above, reflexed below,
toothless. Type A. xystera Val. (see pi. 41, figs. 31, 32, 33, A. hemi-
oxia).
Foot indistinctly tripartite beneath, the upper surface evenly tuber-
culate, without longitudinal grooves on back or tail. Mantle-edge
unusually thick, the right body-lappet very small, left lappet situ-
ated far to the left, and very low.
Jaw (pi. 51, fig. 5, A. xystera) rather widely arcuate, smooth, its
anterior surface totally lacking ribs or stria?, very minutely denticu-
late in the middle of the cutting edge. The jaw figured measures
1.5 mill. wide.
Radula (pi. 51, fig. 4, A. xystera. PI. 49, fig. 25, A. sepulchralis)
composed of very broadly V-shaped, transverse rows ; all of the teeth
unicuspid. Cusps of all teeth wide and rounded, the centrals and
laterals having the basal plates longer than the cusps, marginals
with shorter basal plates, as usual.
Genitalia without accessory organs. Penis stout and short, the
retractor and vas deferens inserted at its apex ; walls of penis cavity
corrugated, the vas deferens entering through a small papilla (pi.
51, fig. 6). Externally, the lower course of the vas deferens is closely
156 AMPKLITA.
bound to the penis from its base to its apex ; its free portion short.
Duct of spermatheca long. Albumen gland large, the ovisperm duct
imbedded in it nearly its whole length (pi. 51, figs. 1-3, 6, A. xystera.
See also pi. 42, fig. 40, A. loucoubeensis).
Embryonal whorls about 2, indistinctly marked off from the after-
growth ; eggs unknown, but apparently one-fifth to one-seventh the
diameter of the adult shell.
Distribution, Madagascar. The general aspect of the shells is
that of ground snails, but the dentition is more like that of arboreal
forms. The prominent features of this genus, apart from its discoi-
dal and peculiar shell, are (1) that all of the teeth of the radula
have single, simple rounded cusps, even the outermost marginals ;
(2) the ribless jaw; (3) the vas deferens is bound to the penis from
apex to base of the latter, and the lower course of the ovi-sperm duct
is bound to the albumen gland nearly the entire length of that
gland.
The genitalia of A. loucoubeensis have been figured rudely by
Brancsik (Jahresheft der naturwissenschaftlichen Vereines des
Trencsener Comitates, xiv-xv Jahrgang, p. 209, pi. 6, f. 3, 1893).
The anatomy of A. xystera and dentition of A. sepulchralis has been
examined by myself. The species are numerous, and some of them
at least are excessively variable, giving rise to an extensive syn-
onymy. Most of those described without figures by Mabille may
prove synonyms or varietal forms of the well-known species.
Group of A. sepulchralis.
A. sepulchralis Fer., vi, 18, 301. A. subsepulchralis Crse., vi, 22.
labrella Lam. /. obscura C. & F., vi, 302.
f. sganziniana C. & F., vi, 301. /. minor C. & F., vi, 302.
/. prceclara C. & F., vi, 300. sepulchralis Rv., f. 147 b.
f. olivacea Pils., vi, 300. /. nigropurpureaC. &F.,vi,302
f. lethifera C. & F., vi, 300. A. hova Angas, vi, 24.
/. funebris v. Mart., vi, 19. madera Mab., vi, 50.
v. funebris Morel., vi, 301. polydora Mab., vi, 50.
v. eurychila C. & F., vi, 301. A. stragulum C. & F., vi, 23, 302.
cadaverosus Pils., vi, 19. A. lamarei Mke., vi, 25.
/. pallidior C. & F., vi, 301. v. sakalava Ang., vi, 26.
/. excoriata Mart., vi, 22. v. catarella Mab., vi, 49.
A. watersi Angas, vi, 26.
AMPELITA. 157
Group of A. omp ha lodes.
A. omphalodes Pfr., vi, 26. A. basizona Mouss., vi, 29.
v. loucoubeensis Cr., vi, 27. A. guillaini Pet., vi, 30.
lucubeensis Auct. A. consanguinea Fer., vi, 30.
A. calypso Pfr., vi, 28. v. subconsanguinea Pils., vi, 30
v. intensior Pils., vi, 28. A. atropos Fer., vi, 20.
A. chlorozona Grat., vi, 31. A. madagascariensis Lm., vi, 32.
A. vesconis Morel., vi, 31. madecassina Fer.
(y=chlorozona?) A. robillardi Ang., vi, 32.
Group of A. xyster a.
A. novacula Mart., vi, 33. A. cazenavetti F. & B., vi, 35, 302
A. hemioxia Pils., Naut., viii. A. lancula Fer., vi, 36.
A. xystera Val., vi, 33. v. terveriana Grat., vi, 37.
A. shavi Smith, vi, 34. A. fulgurata Sowb., vi, 36.
A. stumpffii Kob., vi, 35. A. (?) testudo Pfr., vii, 89.
/. albina Brancsik. A. unicolor Pfr., vi, 37.
Group of A. lanx.
A. lanx Fer., vi, 38. A. lanciformis Bttg., vi, 39.
v. radama Less., vi, 38. v. nossibeeusis Bttg., vi, 40.
A. suarezensis C. & F., vi, 302. v. carapbelliana Pils., vi, 39.
Group of A. duvallii.
A. duvallii Pet., vi, 41. A. clotho F6r. vi, 42.
A. percyana Smith, vi, 42. A. granulosa Fer. vi, 43.
A. lachesis Fer., vi, 41. A. galactostoma Pfr., vi, 44.
Group of A. covani.
A. covani E. A. Smith, vi, 44.
Unfigured species of uncertain affinities.
A. campelica Mab., vi, 54. A. monacha Mab., vi, 47.
A. cyanostoma Mab., vi, 48. A. omoia Mab., vi, 46.
A. erythromorpha Mab., vi, 51. A. paropta Mab., vi, 55.
A. galactostomella Mab., vi, 53. A. porcaria Mab., vi, 45.
A. gaudens Mab., vi, 54. A. scotina Mab., vi, 46.
A. gaudiella Mab., vi, 55. A. stilpna Mab., vi, 53.
158 PEDINOGYRA.
A. gonostyla Anc., vi, 45. A. subfunebris Mab., vi, 55.
A. lithida Mab., vi, 53. A. thelica Mab., vi, 47.
A. lychna Mab., vi, 52.
Subgenus PCECILOSTYLUS Pilsbry, 1890.
Poscilostylus PILS., Man. Conch., vi, p. 56. Eurystyla ANCEY, not
fital.
Shell compact and globose or globosely-elevated, imperforate or
nearly so, smooth and shining, vividly colored. Peristome blunt,
narrowly expanded, the columellar margin reflexed. Type A.
viridis Dh., pi. 38, figs. 10, 11.
Anatomy unknown. These handsome shells have the appearance
of the Philippine Island Cochlostylas. They are probably arboreal
forms.
A. viridis Desh., vi, 56. A. cerina Morel., vi, 57.
Genus PEDINOGYKA Albers, 1860.
Pedinogyra ALB., Die Hel., p. 162, type H. cunninghami. PILS-
BRY, Man. Conch, vi, p. 13. HEDLEY, Records of the Australian
Mus., ii, 29, and Proc. Roy. Soc. Queensl. vi, p. 63, pi. 3, (anatomy).
Shell large and discoidal, with flattened spire and broadly open
umbilicus, solid, opaque and colored. Whorls 5-6, the last large,
deeply deflexed in front. Aperture oblong-truncate, nearly horizon-
tal, the lip slightly expanded, blunt. Type P. cunninghami, pi. 17,
figs. 5, 6.
Jaw arcuate, ribless, faintly striated transversely and longitudin-
ally, the ends rounded (pi. 17, fig. 2, P. cunninghami).
Radula having the middle cusp only developed, on all the teeth.
Centrals and laterals with the cusp shorter than the basal-plate. Mar-
ginals with a single ovate inclined cusp, projecting beyond the
square basal plate (pi. 17, fig. 4, a central with 1st, 12th and 17th
laterals, and 27th marginal tooth of P. cunninghami).
Genitalia having the penis long, retractor and vas deferens
inserted at its apex, lower course of the latter large. Upper part of
the vagina bearing the long stalked spermatheca, and a long appen-
dicula, and bound firmly to the body-wall at this point. Ovo-testis
imbedded in the digestive gland, as usual (pi. 17, fig. 1, P. cunning-
hami). Eggs globose, white, 9 mill, in diameter, hard, calcareous,
ANOGLYPTA. 159
brittle, coarsely granular outside, smooth within (pi. 17, fig. 3, P.
cunning hami).
Distribution, Queensland and New South Wales, Australia. P.
cunninghami has been found living " under heaps of stones and
drifts of dead leaves, or buried in clusters of from 3 to 6 in the soil^
The sharp edges of broken shells are used by the aborigines of
Port Curtis to polish their spears, boomerangs and waddies."
The more conspicuous characters of this type are its broadly um-
bilicated, quoit-like shell, the presence of an appendicula on the
vagina, and the unicuspid marginal teeth. Both shell and dentition
resemble the South American genus Macrocyclis. Two specific forms
have generally been recognized: a large solid Queensland form,
cunninghami, and a smaller, thinner, keeled form of New South
Wales, muhlfeldtiana ; but Hedley finds that they intergrade. This
difference from north to south is exactly paralleled in other Australian
Helices. Compare Thersites richmondiana of Queensland with T.
novcehollandice of New South Wales ; the solid, highly colored
Sphserospiras, with the thinner, keeled Badistes, etc. It is a well
established rule that as we pass southward from subtropical Queens-
land to the temperate southern regions of Australia, the shells
become thinner, smaller, less richly dyed, and often develop a more
or less obvious peripheral keel.
While the systematic position of this genus in the series cannot
be regarded as unquestionable, I agree with Hedley that it is prob-
ably to be regarded as a depressed manifestation of Panda. It does
not agree with that genus in that Pedinogyra has the ovo testis im-
bedded in the digestive gland. In Pandait is not so imbedded, but
is free as in the Bulimi.
P. cunninghami Gray, vi, 14. v. compressa Mouss.
v. miihlfeldtiana Pfr., vi, 15. v. minor Mouss.
Genus ANOGLYPTA Martens, 1860.
Anoglypta v. MART., Die Hel., p. 312, type H. launcestonensis.
PILSBRY, Man. Conch., vi, p. 92. HEDLEY, Proc. Linn. Soc. N. S.
Wales (2), vi, p. 22 (anatomy) ; and Kec. Austr. Mus., ii, p. 29.
Shell umbilicated, subtrochiform, conoidal above, convex below
the peripheral carina ; lusterless and spirally lirate-tuberculate above,
polished below. Whorls 5 J. the apical ones spirally lirulate, the last
suddenly and deeply deflexed in front. Aperture small, subhor-
160 ANOGLYPTA.
izontal ; outer lip thin, not expanded, having a projecting angle just
above the periphery ; columellar lip slightly thickened and expanded
toward the insertion. Type A. launcestonensis, pi. 29, fig. 16.
Animal having the sole undivided ; upper surface granulated, the
granules arranged in indistinct rows on the back ; facial or lateral
grooves distinct ; tail pointed and flattened. Mantel edge thick,
developing a large left body-lobe in front of the respiratory pore,
and a triangular right one below and behind it. Genital foramen
upon the right lateral groove, below and behind the eye stalk.
Jaw arcuate, with a slight median projection ; very finely, irregu-
larly striated vertically (pi. 47, fig. 6).
Radula having all of the teeth unicuspid. Central and lateral
teeth (pi. 48, fig. 10) having the basal-plates contracted on the outer
margin, forming a sort of socket for a projection on the inner mar-
gin of each succeeding plate. Marginals with long, broad, oblique
cusps, becoming shorter on the outer ones (pi. 48, fig. 11, three
groups of marginal teeth, the right hand group from the outer edge
of r ad ul a).
Genitalia having a very short vestibule ; lower part of vagina
swollen, enlarging again above as it passes into the spermatheca
duct. This duct is very long, slender and closely bound to the uterus
above, ending in a globular receptacle. Below, the lower portion of
the duct is very large, with muscular walls, and bears a short blind
sack, directed downwards. This sack, and the enlarged duct and
vagina together, have strongly ridged internal walls. Uterus hav-
ing a very narrow neck (pi. 47, fig. 5, showing neck of uterus and
its union with vas deferens, below the blind sack of spermatheca
duct). Ovo-testis composed of a very long, straggling series of irreg-
ular clusters of fine follicles, imbedded in the liver along its inner
surface. The penis has the vas deferens inserted below the apex,
above and opposite to the insertion of the retractor muscle, which is
very long, and attached distally far back on the lung floor. Penis
cavity closely and strongly ribbed longitudinally, with no papilla.
Vas deferens firmly bound to the penis its entire length, and also firmly
bound to the vagina PI. 47, fig. 8, showing course of v. d. on penis.
(See pi. 47, figs. 5, 7, 8, A. launcestonensis').
Anoglypta is a monotypic genus created for one of the most
peculiar of all Helices. In its coarse spirally lirate-tuberculate
sculpture A. launcestonensis stands unique; and our knowledge of
CARYODES. 161
the soft parts of the animal throws but a feeble light on the ques-
tions of its origin and affinities. The eggs are like those of Caryodts.
The sculpture of the earlier whorls is almost exactly as in that gemi*.
The perfectly simple, unexpanded edge of the lip, and the basal
color zone are also other points of likeness between Anoglypta ancl "
the Caryodes, Panda, Pedinogyra series. The genital system is
peculiar in having the vas deferens closely bound in the integument of
the penis, as in Ampelita, and in the backward-projecting sack on the
sperrnatheca duct. This may perhaps be interpreted as an appen-
dicula, or it may be an independent development for the reception
of spermatophores, like the diverticulum in the true Helices. The jaw
is not smooth, as in all other genera of the Macroon group, but
finely striated as in Pyramidula. The radu la is altogether similar to>
that of Helicophanta, Panda, Caryodes, etc. On the whole, it seems
that Charles Hedley's estimate of the affinities of Anoglypta is by far
the most probable yet advanced. The position assigned by von Mar-
tens, and those formerly suggested by the writer, are clearly unten-
able.
The only species of this genus, Mr. Hedley writes, is confined
to a mountainous district in north-eastern Tasmania. He found it
plentiful among the fern-tree gullies. " Habits very shy and
timid, crawling very slowly. It frequents damp places under logs-
and decaying stems of tree-ferns. The fire and ax of civilization-
threaten to diminish the already narrow range of this splendid and
interesting species, but its haunts are so rugged and remote that I
do not fear its extinction."
A. launcestonensis Reeve, vi, 93. N.-E. Tasmania.
Genus CARYODES Albers, 1850.
Caryodes ALB., Die Hel., p. 141, type Bulimus dufresnii. MAR-
TENS in Die Hel., p. 228. SEMPER, Reiseu im Arch. Phil., Land
Moll., p. 102 (anatomy). TENISON- WOODS, Proc. Linn. Soc. N. S.
W. iii p. 81 (variation, etc.). HEDLEY, Proc. Linn. Soc. N. S. W.
(2), vi, p. 19 and Rec. Austr. Mus., ii. p. 29 (external anatomy,
systematic position, etc.).
Shell Bulimoid t imperforate, varying from oblong to globose-
ovate ; thin but solid, composed of about 5 whorls, the earlier ones
spirally lirulate, separated by a crenulated suture, apex obtuse, last
whorl punctulate above, encircled just below the periphery by a dark
girdle bordered with light. Aperture higher than wide, subvertical,,
11
102 CARYODES.
the outer lip thin and not expanded, columella somewhat sinuous,
xiibtruncate below, with a closely adherent reflexed urnbilico-parietal
callus. Type C. dufresnii, pi. 46, figs. 15, 16.
Foot undivided and without pedal grooves. Back ornamented
with long, narrow tubercles, arranged in about a dozen longitudinal
rows ; sides and tail divided into irregular polygonal spaces, which
are partially subdivided and finely granulated ; tail tapers slightly,
is rounded behind, and never keeled. Genital orifice behind the
right eye-stalk, just beneath the facial groove. Mantel with a left
body-lobe. Kidney opening at its base.
Jaw arcuate, smooth, with no median projection (pi. 42, fig. 44).
Radula with 81-87 teeth in a transverse row, all of them uni-
cuspid (pi. 49, fig. 24).
Genitalia (pi. 42, figs. 41, 42, 43) partially everted in the example
figured, a short papilla bearing a long thread projecting from the
foramen. Penis sac long and stout, the retractor and vas deferens
inserted at its apex ; within the penis lies an adnate fleshy pillar
'(pilaster), free at its distal end ; its outer walls closely grooved,
covered with thick epithelium, and in the folds lay irregular plates
of lime. In a section the pilaster shows outside the external papilla,
separated by grooves ; then follows a sphincter muscle, then an ir-
regular, apparently spiral muscle (pi. 42, fig. 43, pilaster, papilla
.and thread, Fig. 41. section of same, showing star-shaped cavity,
etc.). Spermatheca having a long duct, near the mouth of which is
^attached a long appendicula.
Eggs hard-shelled, regularly oval, white, shining, minutely gran-
ular, measuring 11 by 8 mill. (pi. 42, fig. 46).
The external appearance of the animal and the form of the jaw,
teeth and genitalia, are very similar to Panda, fully supporting the
classification proposed by Hedley in 1892. The shell resembles that
of Panda in its bulimoid contour, simple lip, and the sinuous sub-
truncate columella. It differs from that Australian genus in the
lobed or crenulated sutures, and the sculpture of the embryonic
whorls, which recall Anoglypta. The embryonic shell of Liparus
differs very much in sculpture from that of Panda, Caryodes or any
other Helix known to me.
The genus contains but one species, the Bulimus dufresnii of
authors, Helix dufresnii Leach. The shell varies from oval to
almost globose. The ground-color varies from light yellow to deep
.maroon or dull olive, but the color-band is permanent. The eggs
PANDA. 163
are disproportionately large for the animal, and deposited under logs
during October and November. The size of the egg probably
varies with that of the mature shell, as is the case with Glandina.
The young, upon emerging, are obliquely orbicular in shape (pi. 42,
fig- 45 )
C. Dufresnii Leach. Tasmania.
Genus PANDA Albers, 1860.
Panda ALBERS, Die Heliceen, edit. Martens, p. 149, type H.fal-
coneri Reeve. SEMPER, Reisen, p. 103 (anatomy). HEDLEY, Rec.
Australian Museum, ii, p. 26 (anatomy and systematic position).
PILSBRY, Nautilus, vi, p. 9, May, 1892 (systematic position). Not
Panda Heyden, Isis, 1826, p. 612 (Aearina).
Shell Bulimoid rather than Helicoid, globose-oblong, higher than
wide, umbilicate or imperforate, thin but strong. Surface smoothish.
Whorls 4J, the earlier two finely beaded, indistinctly marked off
from the smoother or spirally striated after-growth (pi. 46, fig. 12,
P. atomatd] ; apex obtuse. Last whorl very large, hardly descend-
ing in front. Aperture large, subvertical, higher than wide ; outer
lip thin, not expanded', columellar lip reflexed toward its insertion.
Type P.falconeri, pi. 46, fig. 11 (P. larryi, pi. 46, figs. 13, 14).
Animal externally like Caryodes. Sole indistinctly tripartite ;
back with some ill-defined longitudinal granulation ; sides and tail
with flat, irregularly polygonal granulation ; tail rather flat and
sharply pointed. Lung cavity and kidney short. Mantle edge
thick, \\ithout lobes (pi. 46, fig. 13, P. larryi).
Jaw arcuate, smooth, with a slight median projection or none (pi.
47, fig. 2, P. atomata.
Radula having all of the teeth unicuspid. Marginal teeth with
long, oblique cusps (pi. 48, figs. 15, 16, P. atomata. PI. 48, fig. 17,
P.falconeri}.
Genital system having the penis stout, the retractor attached to its
summit, and distally arising from the columellar retractor muscle. At
the base of the retractor is inserted an epiphallus about as long as
the penis, then narrowing into the vas deferens. The epiphallus is
partly filled by a " pilaster," or fleshy cord adnate along one side,
which passes into the penis, and there expands into a peculiar penis
papilla (fig. 3) ; internal walls of penis having having several weak
longitudinal fleshy folds. High on the vagina opens the duct of the
spermatheca, and opposite it enters a long appendicula (pi. 47, figs.
3, 4, P. atomata. PI. 47, fig. 1, P.falconeri).
164 PANDA.
Eggs large, white, hard-shelled.
The special sculpture of the apex is generally worn off in adult
shells. The latter whorls are peculiarly variegated with chocolate
streaks and vermiculate lines on a yellow ground, and usually show
spiral bands of blotches.
This genus is more nearly allied to Caryodes than to any other
group. These two Australian genera resemble Acavus, Helico-
phanta and Ampelita in their smooth jaws, unicuspid side teeth and
comparatively large eggs, but differ from them in the simple lip of
the shell, the presence of an appendicula, the insertion of the penis
retractor muscle on the main columellar retractor instead of on the
floor of the lung, and in the freedom of the ovotestis from the diges-
tive gland. The relationship between the Australian and the Indo-
Madecassine genera is therefore by no means intimate. Hedley, in
the important paper on these snails cited above, brought the Austra-
lian Liparus into the group he composes of Panda, Caryodes, Pedi-
nogyra and Anoglypta, but I am unable to follow his classification
to this extent. Liparus seems to me to belong to a distinct stock
I look to Otostomus, Placostylus, etc., for its kindred.
The generic term Panda Hey den, 1826, has not been used by re-
cent araneologists, and the definition given by Heyden in his
analytical table is not sufficient to rescue it from the status of a
nomen nudum. This antiquated use which can never be revived
should not prevent us from retaining Albers' name for the present
group.
P. falconeri Rve., vi, 75. P. atomata Gray.
v. maconelli Rve., vi, 76. v. kershawi Braz., viii, 293.
v. azonata Hedl., viii, 293. v. elongata Hedl., viii, 294.
v. tigris Hedl., viii, 293. v. azonata Hedl., viii, 294.
P. ponsonbyi Ang., P. Z. S. 1877, P. larryi Braz., P. Z. 8., 1871, p.
p. 170, pi. 26, f. 1. 321*.
* **
The following genera, Macrocyclis, Solaropsis and Chalepotaxis
are intercalated here in the Helix series provisionally, pending the
discovery of their true affinities by the examination of the internal
anatomy. The dentition of Macrocyclis, now for the first time made
known, is excessively peculiar, and comparable only to that ofHeli-
copkanta and its allies. Of Solaropsis there is nothing known suffi-
cient to justify a guess at its affinities. Chalepotaxis has the
MACROCYCLIS. 1 65
highly modified radula of a tree-snail, but so abnormal that it affords,
little ground for conjecture.
The Indo-Chinese group Ganesella is placed here because it was
omitted in its proper place in the Epiphallogonous series, with
Chloritis, Planispira, Papuiua, etc.
Genus MACRO'CYCLIS Beck, 1837.
Macrocyclis BECK, Index Molluscorum p. 24, for H. peruviana
{laxata) and H. cunninghami. ALBERS, Die Hel. p. 128 (restricted
to H. laxata). MARTENS, Die Hel. p. 75 (in part). Not Macro-
cyclis of American authors, =Selenites.
Shell disk or quoit shaped with low, convex spire and widely open
funnel-shaped umbilicus. Whorls 4?-5, the last large, deeply descend-
ing in front ; finely and densely striated; yellowish, not banded.
Aperture very oblique, oval, wider than high, the peristome nar-
rowly expanded throughout, reflexed below, the ends approaching.
Type M. laxata Fer., pi. 22, figs. 11, 12.
Genitalia, jaw, etc., unknown. Radula strap-shaped as usual,
bearing many rows of 33.1.33 teeth, all unicuspid ; centrals with
the single conical cusp projecting beyond the basal-plate ; laterals
similar but asymmetrical ; marginals like the laterals, but the basal-
plates are shorter and the cusps longer, oblique and simple (pi. 51,
figs. 1, 2, central with adjacent 3 laterals, 6th and 9th laterals, 12th
and 13th transition teeth, 16-18 and 25-33 marginal teeth, of M.
laxata}.
The shell in this group, except in being uniformly light colored,
is strikingly like that of the Australian Pedinogyra ; and the denti-
tion is altogether similar to Pedinogyra, Panda, Anoglypta and
Helicophanta in the total absence of side cusps ; the marginal teeth
having long, oblique mesocones as in those Old World genera. In
view of the fact that, although unicuspid marginal teeth are pecu-
liarly characteristic of the Macroon group, they reappear in a few
other Helices, I do not feel justified in associating Macrocyclis with
Pedinogyra and its allies. We may better suspend judgment until
the genitali.a and jaw give their more definite testimony. The
radula is very different from that of Selenites.
The single species inhabits Chili.
M. laxata Fer., iii, 109. Var. banksii Cuming, iii, 109.
peruviana Lam. maxima Beck.
deshayesii Anton. umbilicata Anton.
cincinnus Rve. ? gayi Hupe.
laxata Rve.
166 SOLAROPSIS.
Genus SOLAROPSIS Beck, 1837.
Solaropsis BECK, Index Moll. p. 27 (for heliaca, moricandi, brazi-
liensis, pellisserpentis). Martens, Die Hel., p. 164 (type H. pellis-
serpentis Ch.) ; Ostas. Landsch. p. 7 (jaw). PILSBRY, Man. Conch,
v, p. 177. Solarium SFIX, Test. Brazil, p 23. Helicella SWAINS.
Malacol., p. 333 (1840). Psadara MILLER, Malak. Bl.xxv,p. 162,
1878, (for boetzkesi, selenostoma, iris). Ophiospila ANCEY, Con-
chol. Exch., i, p. 64, 1887 (kuhni, andicola, etc.)
Shell urabilicate, rather depressed, with convex or flat spire, con-
vex below, the periphery rounded or angular. Decorated with a
peculiar pattern of lunate brown spots and streaks in bands on a light
ground. Surface granulate, hirsute or plicate-striate. Last whorl
not deflexed in front. Aperture oblique, lunate; lip expanded or
reflexed, its ends distant. Type S. pellisserpentis. (See pi. 46, fig.
20, S. serpens; fig. 21, S. braziliana).
Animal long and slender ; jaw smooth, without ribs ; anatomy
otherwise unknown.
Distribution, southern Brazil and Peru to Columbia and Guyana ;
one species S. tiloriensis, in Costa Rica. They are forest snails, liv-
ing under stones and bark, etc.
The name Ophidermis Agassiz (Ophiodermis Herrm.),- said to
have been proposed in Charpentier's Catalogue of Swiss Mollusks,
1837, but not mentioned therein, has found its way into the synon-
ymy of this genus, through a guess of Herrmannsen's based on its
suggestive etymology. It was never published except as a nude name,
for the snake skin can hardly be said to cover its nakedness. It is
not now worth the expense of clothing ; especially since it really
pertains to something of the nature of Cyclostoma (see Agassiz,
Nomencl. Zool., Moll., p. 62).
S. pellisserpentis Chemn., v, 178. S. napensis Crosse, v, 188.
constrictor Hupe. S. rosarium Pfr., v, 188.
S. serpens Martyn., v, 178. S. kuhni Pfr., v, 189.
pellisserpentis Hupe et al. S. andicola Pfr., v, 189.
colubrina Perry. S. quadrivittata Hid., v, 190.
S. pellisbose Hupe, v, 180. S. diplogoniaDohrn., v, 190.
boa Hupe. S. nubeculata Desh., v, 191.
S. anguicula Hupe, v, 180. S. catenifera Pfr., v, 191.
S. vipera Pfr., v, 181. S. catenulata Anc., viii, 261.
S. monolacca Pfr., v, 182. S. marmatensis Pfr., v, 191.
CHALEPOTAX IS. 1 67
S. gibboni Pfr., v, 182. S. incarum Phil., v, 192.
magnified Lea not Fer. S. monile Brod., v, 192.
v. amori Hid., v, 183. planorbis Jay.
v. cousini Jouss., v, 183. boetzlcesi Mill.
S. pnestans Pfr., v, 184. S. castelneaudii D. & H. v, 193.___
S. braziliana Fer., v, 184. castelnaudii Hupe".
? moricandi Beck. castelnaui Pfr.
S. heliaca Orb., v, 135. S. selenostoma Pfr., v, 193.
S. pascalia Caill., v, 186. sclerostoma Rv.
amazonica Hupe. S. hians Pfr., v, 194.
S. amazonica Pfr., v, 186. S. tiloriensis Ang., v, 194.
S. feisthameli Hupe, v, 187. S. iris Mill., v, 195.
pundata Wagn. not Mull. S. rugifera Dohrn.,v, 195.
v. planior Pils., v, 188. S. elaps Dohrn v, 196.
Genus CHALEPOTAXIS Ancey, 1889.
Chalepotaxis Ancey, Conch. Exch. Aug. 1887, p. 22, type Nanina
f infantilis Gredl. Cf. SCHACKO, Jahrb. D. M. Ges. XI, p. 157, pi.
3, f. 7-10 (dentition).
Shell small, thin, shining, orbiculate-depressed, with narrow um-
bilicus and low-conic spire ; last whorl scarcely descending in front ;.
aperture lunate, slightly oblique, the peristome simple and unex-
panded except at the columella, where it is slightly dilated. Type
C. infantilis Gredler, pi. 57, fig. 34.
Jaw very delicate. Radula (pi. 57, figs. 35-39, 0. infantilis') with the
formula 25.1.25. Teeth all similar in form and in v-shaped rows.
Middle teeth having the median cusp enormously dilated into an
elliptical gouge projecting far beyond the basal-plate ; neck of the
cusp narrow, bottle-shaped ; side cusps basal, rudimentary and verti-
cal ; basal-plate narrow in front, widening and squared behind.
Lateral teeth similar, but the large cusp bends outward and the
entocone is suppressed. Marginals differ only in becoming smaller,
with the ectocone decidedly longer (pi. 57, figs. 35, 37, group of
middle and lateral teeth; fig. 36 group of marginals; fig. 38, a
lateral in profile, turned 90 ; fig. 39, a lateral turned 45).
This genus is founded upon one species having a shell resembling
an immature If. similar is Fer., or pyrrhozona Phil, and a type of
dentition considerably like Oxyclwna. The jaw is very imperfectly
known, and the genital system is unobserved. I am disposed to
168 GANESELLA.
believe, with Schacko, that it is a modified branch of the Hel
stock. It is probably arboreal in habit. Only species, C. infan-
tili* Gredl., ii, 216. Prov. Kwang-si and Hunan, China; Tonquin.
Genus GANESELLA Blauford, 1863.
Ganesella BLANF. Ann. Mag. Nat. Hist. (3), xi, p. 86, type H.
capitium Bens. (Feb., 1863). Satsuma A. ADAMS, Ann. Mag. (4),
i, p. 463, type H.japonica,patruelis,pecu liaris, (June, 1868). Fru-
ticotrochus KOBELT, Fauna Molluscorum extramarinorum Japonise,
1879, p. 48, same types. Trochomorphoides NEVILL, Hand List
Moll. Ind. Mus. pt. 1, p. 80, type H. acris Bens. (1878).
Shell more or less trochiform, umbilicated (or rarely imperfor-
ate), rather thin ; light-colored, plain or with a peripheral line ;
surface with growth-lines only or densely spirally striate ;
whorls 4^-6, the last a little descending in front. Aperture oval
or angular-lunate, oblique, toothless or with a blunt columellar fold ;
lip expanded, broadly dilated at columellar insertion. Type G.
capitium, pi. 55, fig. 18. See also pi. 64, fig. 7, O.japonica.
Animal (of O.japonica) with the foot very long and narrow, sole
not distinctly tripartite ; upper surface finely and feebly granular,
back with a pair of dorsal grooves, no facial grooves ; tail narrow,
long, with a median longitudinal groove above.
Jaw arcuate, with about 9 ribs denticulating the lower margin
(pi. 60, fig. 1, G.japonica).
Radula of the type usual in ground snails. Middle tooth with
mesocone only developed, shorter than basal-plate, side-cusps repre-
sented by slight lateral extensions. Laterals similar but with the
cusp longer. Marginals with oblique, bifid inner cusp and an ecto-
cone (pi. 60, fig. 2, G.japonica).
Genital system (Frontispiece, figs. 1, 2, G. japonicd) having the
penis long and twisted, ending in a curved blind sack with corru-
gated inner walls (fig. 2, apex of penis and sack opened) ; epi-
phallus long, bearing the retractor, terminating in a flagellum and
the vas deferens. Vagina extremely long, the spermatheca duct
inserted high. Sperrnatheca oblong, on a stout duct, neither duct
nor bulb being bound to uterus. No dart sack or mucus glands.
Distribution, Japan and China to India, southeast to Sumatra
Borneo and Philippine Is.
This genus has the genital system, jaw and radula, as well as the
tail-groove of Chloritis (see PI. 28, figs. 1 to 9), but.the penis-papilla
GANESELLA. 169
is absent, and the spermatheca duct is inserted higher on vagina.
The shell has somewhat the contour of Papuina. The anatomy of
the group has been a complete surprise to me, for I had relegated it
to the Eulota group before dissecting specimens. It is now per-
fectly clear that it belongs in the vicinity of Chloritis and Papuina,
and is the most northern in distribution of that group of genera.
Probably some of the species now referred to Ganesella will prove to
belong to other groups, such as the East Asian Fruticicoloid sec-
tion.
There is much variation in contour, number of whorls, size and
umbilicus among the members of this genus ; and a subdivision of
it into sections will no doubt be made eventually. It is to be hoped
that such division will not be attempted until it can be placed on a
firm footing by the examination of the anatomy of many species;
and anatomical data are also required before the boundary line be-
tween Ganesella, Eulota, Pledotropis and the East Asian Fruticico-
las can be definitely drawn. In some cases the shell alone is not
sufficiently characteristic to base the classification of these groups
upon, even when the relationships of the main types have been
elucidated.
Japanese, Liukiu Is. and Formosa species.
G. papilliformis Kob., iii, 217. G. conella Ad., iv, 56.
G. japonica Pfr., iii, 218. G. lischkeana Kob., iii, 220.
? vitracea Fer., vii, 106. G. peculiaris A. Ad.
G. conospira Pfr., iii, 218. G. gibbosa A. Ad.
G. tabuensis Anc., iii, 218. G. ? serotina A. Ad.
patruelis Ad. not Ang. G. sphserulata Reinh.
G. sphinctostoma Ad., iii, 218. G. largillierti Ph., iii, 218.
G. cardiostoma Kob., iii, 219. immaculata A. & R.
G. hilgendorffi Kob., iii, 219. G. albida Ad., iii, 218.
G. verrucosa Reinh., iii, 219. G. fulvicans H. Ad., iii, 220.
G. macrocycloides Kob., iii, 219. G. sphseroconus Pfr., viii, 200.
G. eumenes West., viii, 199. v. campochilns Pils., viii, 201.
G. goodwini Sm., iii, 219. G. scsevola Mts., vi, 306.
G. conulina Mart., iii, 219.
Chinese species.
G. gradata Mlldff. G. alveolus Hde.
G. brevibarbis Pfr., iii, 221. G. ternaria Hde.
170
GANE8ELLA.
G. pquamosella Hde., iii, 221.
G. micacea Hde., iii, 221.
G. phyllophaga Hde., iii, 221. .
G. dormitans Hde., iii, 222.
G. ? arbusticola Dh., iii, 222.
G. bizona Gredl.
G. squamulina Gredl.
G. trochacea Gredl., viii, 200.
G. microtrochus Moll., viii, 201.
G. lepidostola Hde., iv, 55.
v. trochospira Mlldff.
G. schomburgiana Mlldff.
trochulus Mlldff. not Ad.
G. vitreola Hde.
G. ingloria Hde.
G. subsquamulata Hde.
G. subparasitica Hde.
G. subgriseola Hde.
G. ? galera Hde.
G. ? peneruginosa Hde,
G. radulina Hde.
G. virilis Gredl., iv, 259.
v. subfusca Gredl.
G. laurentii Gredl., iv, 259.
G. editha A. Ad., viii, 204.
Species of India, Toiiquin, etc.
G. capitium Bens., iii, 74. G. phonica Mab., vii, 83
v. hariola Bens., iii, 74.
G. acris Bens., iii, 74.
puellula Bens.
G. perakensis Cr., vii, 82.
v. subperakensis Pils., vii, 82.
G. galea Bens., iii, 75.
G. bouryi Morg., iii, 172.
G. rostrella Pfr., vii, 83.
G. scenoma Bens., vii, 83.
G. hyperteleia Mori., viii, 203.
G. mera Rve., iii, 94.
Species of Sumatra, Java and Borneo.
G. gysseriana Pfr., iii, 75.
? conulm Mart, not Pse.
G. bantamensis Sm., vii, 84.
G. rufofilosa Bock, vii, 84.
G. niahensis G.-A., vii, 85.
G. tigreensisG.-A., vii, 85.
G. subflava G.-A., vii, 85.
G. angulata Iss , iii, 75.
Philippine Island species.
G. trochomorpha Mlldff., viii, 202.G. fernandezi Hid., viii, 202.
microtrochus Mlldff. olim. G. planasi Hid., viii, 202.
v. mimula Mlldff. G. poecilotrochus Mlldff. Nachr.
v. dimidiata Mlldff. ['95, 114.
G. trochus Mlldff, viii, 201.
A section of Oanesella is probably indicated by the lack of flagellum
and the columellar fold of H. ptychostyla (see Semper, Reisen, p..
247, footnote, pi. 16, f. 27). The appendage of penis, figured for G.
japonica, is developed, and somewhat sacculated or feathered. These-
GANESELLA. 171
species were formerly grouped in Pleetotropis, but the lack of dart
sack and mucus glands widely sunders them from that group.
G. ptychostyla Mart., iv, 58. G.'styloptycha Pfr., iv, 58.
goniochila Pfr., iv, 58. ptychostyla Pfr. not Mart.
/. depressior Pfr.
Subgenus (?) BULIMINOPSIS Heude.
Buliminopsis HDE., Notes sur les Moll. Terrestr. de la Vallee du
Fleuve Bleu, p. 146, type H. buliminus. Conf. v. Molldff., Nachr.
d. m. Ges. 1886, p. 195. Rudens HDE., t. c., p. 148. type Funiculus
rudens. Pseudo buliminus Gredl., SCHMACKER & BOETTGER, Nachr.
D. M. Ges. 1891, p, 164.
Shell elevated conic, perforated, the spire acute, 7-8 whorls; aper-
ture small, oblique, peristome expanded. Soft parts unknown.
A middle Chinese group of uncertain position. Mollendorff refers
it to Satsuma, Ancey to Buliminus, while Heude and Gredler cut the
Gordian knot by removing the species from both genera.
G. pseudobuliminus Hde., iv, 31. G. incerta Pfr.
B. macrogonus Anc. taivanica Mlldff., iv, 33.
G. buliminoides Hde., iv, 31. G. quaternarius Hde.
B. tropidophorus Anc. borealis Hde. on pi.
G. buliminus Hde., iv, 32. G. conoidea Hde.
B. helicopsis Anc. G. doliolum Gredl.
v. pinguis Anc. F. rudens Hde.
G. macroceramiformis Dh.
Subgenus (?) COLIO'LUS Tapparone-Canefri, 1887.
Ann. Mus. Civ. Genov. (2), iv, p. 131. Manual of Conchology
(2) vii, p. SI. Not Coleolus Hall, Paleont. N. Y. v, p. 184, 1879.
Shell elevated- conic, many (eleven) whorled, upper whorls
spirally striate, the rest obliquely costulate and setigerous ; apex
obtuse, mamillar ; base depressed ; peristome reflexed below, mar-
gins distant, connected by a callus. Type C. arfakiensis Tap.-Can.
vol. vii, p. 87.
Soft parts unknown. Inhabits New Guinea. This peculiar snail
is considered an ally of Trochomorphoides by Tapparone-Canefri.
Perhaps it may prove to belong to the Charopoid series.
172 DORCASIA.
Genus DORCASIA Gray, 1845.
Dorcasia GRAY, Zeitschr. f. Mai. 1845, p. 87, type H. alexandri ;
P. Z. S. 1847, p. 171. BINNEY, Ann. N. Y. Acad. Sci. iii, p. 106,
pi. 6, f. M (Dentition). Galaxias (part) BECK, Index Moll., p. 42
(preoc.). Of. PFEFFER, Verb. Vereins f. naturwissench. Unterhal-
tung zu Hamburg, vi, p. 118, 1887. Also SIMROTH & BOETTGER,
Berichte d. Senckenb. Gesellsch. 1885, p. 16, pi. 1, f. 2 (as " Bulim-
inus sp.").
Shell rather large and solid, glossy and unicolored ; umbilicated,
globose or depressed with rounded periphery, rather conoid low
spire and deflexed last whorl. Aperture oblique or subhorizontal,
rounded-truncate, toothless ; the lip thickened, and reflexed at least
below. Type D. alexandri. See pi. 38, figs. 6, 7, D. alexandri var.
rotundata. Also D. ylobulus, pi. 38, fig. 8.
Jaw low, wide, slightly arcuate, entirely smooth (pi. 60, fig. 3,
D. alexandri). Foot (of alexandri') short and broad, the sole very
indistinctly tripartite; upper surface coarsely granular, the gran-
ules polygonal, subdivided ; with no trace of pedal grooves ; back
with several longitudinal lines, obsolete toward head ; facial grooves
well marked and continuous from mantle to head, on both sides ;
tail more finely granose, obtuse behind, rounded above, without
median groove. Mantle with small right and left body-lobes.
Right eye-stalk retracted between branches of genitalia. Blind sack
of the foot very long, lying free in body cavity.
Radula (pi. 60, fig. 6, D. alexandri} having mesocones only devel-
oped on middle and inner lateral teeth, the side cusps being repre-
sented by lateral extensions of the mesocones. On the outer laterals
and marginals the ectocone becomes distinct and well developed.
In D. globulus (pi. 51, fig. 3,) both median and lateral teeth are
distinctly tricuspid. Marginals a simple modification of the laterals,
the broad cutting-point trifid.
Genital system (frontispiece, fig. 3, D. alexandri) without accessory
organs of any kind. Atrium very short. Penis long, larger, and
abruptly bent toward the apex where the terminal, short retractor
is inserted, its distal attachment being on the lung floor. The vas
deferens is not terminal, but enters about one and one-half millim.
below apex of penis. Vagina long ; spermathecaon a long branch-
less duct, entering high on vagina.
The specimen of D. alexandri examined by me was kindly com-
munciated by Dr. Simroth, and is the same one which supplied the
data given in Ber. Senck. Ges. 1894. It is a badly preserved
spirit example, and shows signs of immaturity.
DORCASIA. 17$
^
The dentition of D. globulus differs from that of alexandrl in the
development of side cusps, which are represented in the latter by
wide extensions of the mesocones. This is not an unusual variation.
The smooth low jaw recalls Helicophanta, but the egg is apparently
minute in Dorcasia, and we have from the mouth of the animaLik
self an emphatic contradiction of such a relationship, for the teeth
are totally unlike the unicuspid type of the Helicophanta and Am-
pelita group.
The entire simplicity of the genital system shows Doreasia to be-
long to the Euhaplogona, most living members of which are restricted
to America, Polygyra being a leading genus. In this group of
genera the penis bears neither epiphallus nor flagellum, the vagina
or atrium have no dart sack or mucus glands, the duct of the sper-
matheca does not branch into a diverticulum. Dorcasia is, there-
fore, isolated among the Helices of Africa, Asia and Europe. It is
interesting to note that many of its associates in the Cape fauna are
equally so, and mainly belong to a much older fauna than that
occupying these continents : Aerope has its allies only in Aus-
tralia, Tasmania and New Zealand ; Trachycystis (Pella) has the
same geographic alliances ; Peripatus has a similar, though wider,
range ; and many other Cape animals could be named which belong
to an archaic fauna.
With Oriental snails of the type of H. similaris Fer. (Eulotella),
these South Africans have no especial relations.
All of the species are from the South African zoological province,
with the exception of the doubtful D. votiva Cr., from Madagascar,
which differs from all the other species in being banded.
D. rosacea Miill., iii, 213. D. lucana MiilJ., iii, 213.
D. porphyrostoma M. & P., viii, D. inhluzana M. & P.
[262. D. usambarica Crav., iii, 155.
D. globulus Miill., iii, 213. D. kraussi Pfr., iv, 50.
lucana Lam., Fer., Rossm. D. cernua Mts., viii, 263,
D. namaquensis M. & P., viii, D. alexandri Gray, iii, 213.
[262. v. minor Bttg., viii, 261.
D. gypsina Melv. & Pons., viii, v. rotundata Mss., viii, 261.
[262. D. ? bulbus Mke., iii, 213.
D. coagulum Mts., viii, 263. D. ? votiva Crosse, iii, 214.
* * *
Belogona.
The series of genera following are characterized by the possession
of organs wanting in all other Helices, viz. a muscular sack (or
sacks) on atrium or vagina containing a calcareous needle or dag-
174 BELOGONA.
ger like " dart," and a gland or glands inserted upon or above this
sack, the so-called "digitate glands" or mucus gland.
The presence of these organs was early noticed by European mal-
acologists, but their significance has been only recently recognized.
Semper in 1874 made two divisions of rib-jawed Helices, those
genera with no accessory organs on genitalia, and those with such
accessories; and in 1888 the writer used these features of the
genitulia as diagnostic of various groups of Helices, elaborating the
idea in a later paper (1892). Meantime Dr. H. von Ihering issued
a paper of great merit, " Morphologic und Systematik des Genital-
apparates von Helix," in which he proposes to restrict the family
Helicidce to snails with grooved or ribbed jaws and possessing
the dart apparatus, including therein as genera Xerophila,
Fruticicola, Helix (=Pentateenia), Campylcea, Gonostoma, Dorcasia
( Eulota), and Cochlostyla. In the following pages I have adopted
all of these groups as genera (although altering the names of most
of them), and with the exception of Campyfaa and Dorcasia, they
are retained with the limits defined by von Ihering. I need give
no other expression of the high esteem in which I hold v. Ihering's
work, than this use of it. It should be added, however, than many
genera not noticed in von Ihering's paper, are now included in this
group, some of which have ribbed, some smooth jaws. His family
diagnosis of " Helicidse ", therefore, does not cover nearly all the
forms here grouped under Belogona.
The relationship of the Belogona to the Epiphallogona is dis-
cussed in the introductory portion of this volume. It remains to
study the internal affinities of its numerous genera. It has been
seen that the Belogona differ from Epiphallogona only by the addi-
tion of the dart apparatus, the penis having exactly the same mor-
phology in the two groups. Now the simplest type of dart appar-
atus is that found in the genus Helicostyla, consisting of a sack con-
taining a needle-like dart, without crown or blades, and a simple,
mucus gland upon the dart sack, consisting of one layer of secreting
cells arranged radially around a central space or duct (see pi. 54,
fig. 7). This is, there can be no doubt, the primitive type of the dart
apparatus, from which the various elaborate forms of darts and
glands arose. No really primitive Belogona are now known to exist.
Helicostyla is practically so in its dart arrangement, but it is diver-
gent in the loss of the flagellum (present in its Epiphallogonous
ancestors) and in the highly modified shell.
The anatomy of the European types of dart-bearing helices has
been studied by Schmidt, Lehmann, Moquin-Tandon, and many
BKLOGONA. 175
later authors. The American forms have been studied by W. G.
Binney, but as many of his figures are of doubtful accuracy my con-
clusions have been based wholly upon fresh dissections. The West
Indian genera are herein for the first time made known anatomic-
ally ; and the forms of East Asia are partially known by the work
of Semper, but largely by my own dissections. The great mass of
data before me from these sources, has compelled me to reject von
Ihering's phylogenetic scheme, and to offer the following arrange-
ment :
BELOGONA EUADENIA. Mucus gland one, inserted on dart sack
or at its base ; simple or divided, glandular, sacculated, globular or
bulbous.
BELOGONA SIPHONADENIA. Mucus glands usually two or many,
inserted on vagina ; tubular or composed of tubular branches.
Apparent exceptions to this arrangement are seen in Helicigona
quimperiana, where the tubes are shortened into hollow, thin-walled
sacks, and some Fruticicoloid forms with demonstrably degenerate
genitalia. The first of these divisions will now be discussed :
BELOGONA EUADENIA.
This division of the Belogona, characterized by having mucus
glands of typically glandular structure, in contradistinction to the
tube-like glands of the Siphonadenia, is now distributed throughout
Eastern Asia, outlying groups extending to New Guinea and the
Solomon Is., and northward to Japan and Siberia. In America it
occupies the Pacific slope from British Columbia to Argentina, with
genera in the Greater Antilles. It is a significant fact that its area
while in large part coincident with that of the Epiphallogona
(Hadra, Camcena, Obba, etc.) is over stepped on nearly all sides by
the latter. Thus Planispira extends further west in India ; Thers-
ites (+ Hadra) and Chloritis extend beyond it southward to Aus-
tralia ; Papuina has a far greater range throughout the " Melanes-
ian Plateau " ; and Ganesella follows the Euadenia to the confines
imposed by rigorous climate in the north. And in the New World,
again, while both Euadenia and Epiphallogona have a wide range
in South America, the latter are universally dispersed throughout
the Caribbees as well as the Greater Antilles, whilst the former came
too late to follow them to the Caribean chain. The inference is, of
course, that the Kpiphallogona are an older faunal element, and
have had more time to take advantage of the various means of dis-
persal by which islands (especially continental islands) and conti-
nents have been peopled.
176 BELOUONA.
A single European genus, Leucochroa, is herein referred to the
Euadenia ; but it is a degenerate group in genitalia and jaw and
may prove to belong to theSiphonadenia, in the vicinity of Helicella
(Xerophila), which it resembles in the simple-lipped, chalky shell
and the peculiar musculature. The American genus Lysinoe is also
aberrant, differing from all other Euadenia in having three club-
shaped mucus glands inserted on vagina, and in the doubling of the
dart sack ; but it differs from all Siphonadenia as well in having
the mucus glands inserted one behind the others, instead of at the
same level on the vagina. I have considered it a tangent from the
Epiphragmophora circle. Oxychona is still imperfectly known.
The genera of this division may be tabulated as follows :
a. New World genera.
1. Dart sack 1, with subapical constriction, apex attached by a
thread to vagina; mucus gland 2-lobed; jaw smooth; tail not
serrate.
b. Middle and inner lateral teeth 1-cuspid, marginals 3-cuspid,
[CEPOLIS.
bb. All teeth with three subequal cusps, POLYMITA.
2. Dart sacks 2 ; mucus glands 3, on vagina ; tail with serrate keel ;
jaw ribbed.
b. Teeth of normal type; shell subglobose, large, deep colored ,
[LYSINOE.
bb. Teeth with wide middle and minute side cusps ; shell troch-
oidal, OXYCHONA.
3. Dart sack 1 ; mucus glands absent, jaw ribbed ; shell discoidal
with thin, simple and acute lip, GLYPTOSTOMA.
4. Dart sack 1 ; mucus gland single, club-shaped, bifid and bulbi-
ferous, or 2 with flat glandular extremities adnate on vagina or
d. s., EPIPHRAGMOPHORA.
aa. Old World genera.
1. Dart sack 1, well-developed.
b. Mucus gland single, globose, inserted on dart sack,
[HELICOSTYLA.
bb. Mucus gland acinose ; shell bright colored, CHLOR/EA.
bbb. Mucus gland divided, lobes sacculated, elongated,
EULOTA.
2. Dart sack wanting; jaw smooth ; shell strong, chalky and white,
LEUCOCHROA.
CEPOLIS. 177
Genus CEPOLIS Montfort, 1810.
= Cepolis Montf.-f- Eurycampta Mart.-f Jeanneretia Pfr.-j-jHemi-
trochus Swains. -|- Coryda, Dialeuca and Leptoloma Alb. -\-Histrio
and Plagioptyeha Pfr.-f Cysticopsis Morch not Martens.
Shell globose-depressed or globose-conoid, umbilicate or imper-
forate, smoothish, rib-striate or spirally malleated ; lip expanded
(or simple and sharp), reflexed at columella, which is generally
thickened with an oblique callus, sometimes a tooth ; lip otherwise
toothless but occasionally there is a callous fold within the mouth ;
varying from unicolored to conspicuously streaked or banded, the
bands irregularly disposed. Type C. cepa, pi. 25, fig. 9. (See also
pi. 56, figs. 1 to 9, and pi. 58, figs. 54 to 56).
Animal granulated above, without distinct dorsal grooves, facial
furrows or tail-groove, the sole not tripartite except in color ; man-
tle with small right and rudimentary left body-lappets. Right eye
retracted between branches of genitalia.
Jaw high arched, with an obvious or slight median projection and
sometimes a wide, vertical rib-like median convexity ; its surface
smooth or showing slight strise (pi. 57, figs. 41 to 46).
Radula long, with comparatively few longitudinal rows of teeth
(30. 1. 30 to 45. 1. 45). Middle and lateral teeth having long, nar-
row basal plates, and short, broad middle cusps, shorter than the
basal plates, and with no trace of side cusps. Transition teeth
developing the ectocone ; marginal teeth tricuspid, the ento- and
meso-cones short, coalescent at base, ectocone simple or bifid. (PI.
57, figs. 40, 47 to 51).
Genitalia (pi. 52, figs. 12-16, 19, 21) characterized by a long,
slender penis provided with a weak retractor or none, inserted low
on penis and distally on the lung floor ; the apex of penis splitting
into a long flag ellum and the v. d. Low on vagina or on atrium is
borne a long club-shaped dart sack, with constricted head, which is
bound by a string of connective tissue to base of vagina ; at the base
of dart sack the glandular, flat, two-lobed, elongated mucus gland
is inserted. Sperm atheca long, closely bound to upper end of
uterus ; its duct very long, closely adherent to uterus, convoluted on
lower end of same, but free from vagina, near the base of which it
is inserted. Notwithstanding the well developed dart sack, I found
no dart in any of the numerous individuals of this genus exam-
ined.
12
178 CEPOLIS.
Distribution, greater Antilles, Bahamas, Florida Keys.
See under Plagioptycha for notes on the fossil forms.
The prominent features of this group are (1) the smooth, high
arched jaw with median projection, (2) the long radula with few
longitudinal rows, middle and lateral teeth with long, narrow basal
plates and short, broadly rounded mesocones, no side cusps, margi-
nals with short ento-f mesocones, (3) the weak or even lacking
retractor of the long penis, the club-shaped dart sack and two-lobed
mucus gland ; long, unbranched spermatheca duct, etc.
The only near ally of Cepolis is the genus Polymita, which
inhabits the same tract. The latter has the same type of jaw and
geuitalia, but differs in the radula with over twice as many longitu-
dinal rows of peculiarly modified teeth, all of them bearing three
nearly equal cusps. From the Californian and Mexican Epiphrag-
mophora species Cepolis differs in the very characteristic form of
the dart sack, the short inner cusps of the marginal teeth, the ribless
jaw, etc.
Part of the species of this genus are ground snails with dull
brownish shells, but little variegated, as in the sections Cepolis,
Jeanneretia, Euryeampta, Plagioptycha; part are arboreal, and in
these the shell is generally bright in color, often with a rich and
beautiful banded or streaked pattern, Coryda, Hemitrochus and
Dialeuca being of this sort. A parallel series of variations is seen
in the Philippine Island Cochlostylas, where we have also arboreal
and terrestrial forms.
This genus is remarkably homogeneous in characters of the soft
anatomy, which offers no divergence of more than specific value
throughout the entire group. I have given on plates 52 and 57
drawings representing the anatomy of a sufficient number of the
sectional groups to allow any malacologist to judge for himself of
the literal truth of this statement. The shells afford characters for
several sectional divisions, of which it must be said that although
the typical species are quite different, intermediate forms reduce the
diagnostic sectional characters to a minimum. This intergradation
has caused me to disregard the fact that former authors have dis-
tributed the elements of my genus Cepolis far and wide throughout
the Helix series ; and I venture to predict that any one having a
fairly complete collection of the species will endorse the views here
advanced if he will bring the species together and observe the
transition forms uniting the various sections. Cepolis is bound to
CEPOLIS. 179
Jeanneretia by G. squamosa, subtussulcata, etc.; C. exdeflexa is a
transition between Jeanneretia and Eurycampta, and is not far
from some of the Plagioptychas, while nemoralina, filicosta and
maynardi bridge the gap between Plagioptyeha and Hemitrochus.
I am unable to find in Hemitrochus and Polymita any general-
system or plan in the distribution of bands, such as occurs in the
five-banded Helices of Europe or in the epiphallogonous groups of
Asia and Australia. I believe that the color schemes of the arbo-
real West Indian forms have been independently evolved, with the
exception of the supra-peripheral band, which may possibly be
homologous with that of Campylaea, Tachea, etc.
A prominent feature in some species of this genus is the tooth
within the mouth of the shell, marked by an external pit. A simi-
lar structure occurs in Solaropsis, Planispira, Neocepolis, etc., but
it does not seem to be of generic or even subgeneric value in any
group.
The sectional divisions are as follows :
(Cepolia. fPtagioptycha.
T ,. Cysticopsis.
-I Jeanneretia. J ^ . J
-^ Hemitrochus.
Eurycampta.
^Coryda-fDialeuca.
Section Cepolis Montf., 1810.
Cepolis MONTF., Conch. Syst. ii, p. 150 (type nicolsinianum Montf.
=cepa Mull.) ; Cepolum MONTF., 1. c. p. 151.
Shell rather solid and of moderate or large size, imperforate or
umbilicate, compact, globose- depressed, opaque, striate or malleated,
2 or 3 banded, the spire low, conic or convex ; whorls less than 5,
the last abruptly deflexed in front, having a pit below the periphery
a short distance behind the lip, which inside the shell appears as a
callous fold a short distance within the outer lip. Aperture quite ob-
lique, truncate-oval, the lip expanded ; columellar lip reflexed,
armed inside with a compressed or entering tooth. Type C. cepa, pi.
25, fig. 9.
Soft anatomy unknown. Distribution, Hayti.
Differs from Jeanneretia mainly in the stronger columellar tooth
and the constant deep pit behind the lip forming a callous fold
within the mouth.
180 CEPOLIS.
C. cepa Mull., v, 93. C. trizonalis Grat., v, 93.
impressa Blv. v. trizonella Pils., v, 94.
nicolsinianum C. trizonaloides Brown, v. 95.
pimesoma Pils., v, 95.
Section Jeanneretia Pfr., 1877.
Jeanneretia PFR., Mai. Bl. xxiv, p. 7 ; Nomencl. Hel. Viv., p. 116,
PILSBRY, Man. Conch, v, p. 48. Cf. POEY, Memorias, pi. 6, f. 6.
genitalia of parraiana.
Shell imperforate or urubilicate, globose-turbinate, light brown,
generally with darker chestnut bands, two or three in number.
Whorls 5 to 6J, slowly widening, the last deflexed in front and con-
stricted behind the lip ; aperture oblique, rounded-truncate; lip
reflexed and thickened, the columellar margin straightened. Type
(7. multistriata Dh. (See pi. 58, figs. 54, 55, C. parraiana).
Jaw and radula unknown. Genitalia as in Eurycampta (pi. 52,
fig. 21, C. parraiana, after Poey).
Distribution, Cuba ; one species, C. squamosa, is from Porto Rico ;
they live under dead leaves and stones.
The group is allied to Cepolis and Eurycampta, its main distinct-
ive features being the spirally lirate surface and the groove or con-
striction behind the reflexed lip.
C. multistriata Dh., v, 49. C. angulifera Mart.
circumtexta Fer. C. parraiana Orb., v, 50.
vesica Lea. v. parallela Poey, v, 51.
bicincta Mke. C. sagraiana Orb., v, 50.
adjuncta Zgl. C. subtussulcata Wright, v, 51.
v. pityonesica Pfr., v, 49. C. squamosa Fer., v, 95.
C. wrighti Gundl., v, 49. macularia Lm.
C. dermatina Sh., v, 50.
Section Eurycampta Martens, 1860.
Eurycampta MART, in Alb., Die Hel., p. 127, type H. bonplandi.
Shell narrowly umbilicated, orbiculate convex, obliquely rugose-
striate, with a satin like lustre ; brown, uniform or with 1 to 3
bands above, one or none below the rounded periphery. Whorls 5
or less, the last unusually wide, deflexed in front. Aperture large,
transverse, oval; peristome expanded and lipped, reflexed below,
CEPOLIS. 181
the columellar margin often callously thickened within ; ends of lip
somewhat approaching. Type C. bonplandi, pi. 58, fig. 56.
Animal as described for C. alauda, but lighter colored.
Jaw solid, high arched, smooth except for slight striae in places,
(pi. 52, fig. 18, C. bonplandi).
Radula (pi. 52, figs. 20, 22, C. bonplandi) long, the middle and
lateral teeth with long basal plates and short, rounded mesocones,
no side cusps. Transition teeth developing an ectocone (fig. 22,
central with two adjacent laterals and two transition teeth). Mar-
ginals of the usual tricuspid type (fig. 20).
Genital system as in Coryda, etc., but the retractor muscle is
stouter, flagellum and mucus glands longer (pi. 52, fig. 19, G. bon-
plandi}.
C. bonplandi Lam., iv, 82. C. poeyi Petit, iv, 83.
C. supertexta Pfr., iv, 82. staminea Mke.
C. arctistria Pfr., iv, 82. velutinata Bk.
C. exdeflexa Pils., v, 198. C. bryanti Pfr., iv, 83.
deflexa Pfr. not Brauu. C. desidens Rang, iv, 83.
Section Cory da Albers, 1850.
Cory da ALB., Die Hel., p. 100, for alauda and varieties. Histrio
PFR., Mai, Bl., 1855, p. 185 ; 1877, p. 8, for H. dennisoni.Helico-
styla BECK, Index, p. 36, in part.
Shell depressed-globose, imperforate, solid and strong, smooth,
with deeply and abruptly deflexed last whorl, very oblique, trans-
versely oblong aperture, the lip expanded, thickened within, and
having a conspicuous banded, obliquely streaked or dotted color
pattern. Type H. alauda pi. 56, figs. 3, 4.
Animal of H. alauda blue-black, the sole light slate colored in
the middle, not tripartite except in color. Foot long, granulated,
without distinct longitudinal grooves on back and lacking facial
grooves. Tail evenly and more finely granulated, acute behind.
Mantle-edge thin, with a low right body-lappet and a minute left
one. Right eye retracted between branches of genitalia.
Jaw (of H. alauda pi. 57, fig. 45) solid, highly arched, with a
wide median projection, its surface entirely smooth.
Radula (of H. alauda pi. 57, fig. 49) long and narrow, with V-
shaped rows according to the formula 24. 9. 1. 9. 24. Median teeth
with long basal plates and short, broad mesocones, no side cusps.
182 CEPOLIS.
Laterals similar but asymmetrical. Marginals developing a stout
ectocone, and on the outer ones an entocone. The figure represents
a middle tooth with 3 laterals and an inner and outer marginal.
Genital system with vestibule short ; penis (pi. 52, fig. 13) very
long and slender, without retractor, terminating in v. d. and a long
flagellum. Vagina branching low into a very long and much twist-
ed spermatheca duct which ends in a long spermatheca bound
closely to top of uterus. Dart sack very large, dark colored, with
a long fleshy white head, the apical portion separated by a con-
striction and united to base of uterus by a connective thread.
Mucus glands two, long, leaf-like and glandular, uniting at their
bases and inserted on the dart sack near its base. No dart found
on the papilla in several specimens examined, which were of the
"strobilus" variety (pi. 52, figs. 12, 13, C. alauda).
The species are few, and all from eastern Cuba except H. circu-
mornata with its two slight color varieties from western Hayti. H.
dennisoni is hardly more than a variety of alauda. The last named
species is arboreal, and is frequently found living in the cargoes of
bananas brought to Philadelphia and other eastern cities. Through
the kindness of Mr. John Ponsonby I am able to fix at last the
identity of the long lost H. drcumornata, and the status of vigiensis
and stenostoma.
C. alauda Fe*r., v, 42. C. ovumreguli Lea, v, 44.
strobilus Fer. C. circumornata Fer., iv, 222.
avellana Fer. v. vigiensis Weinl., v, 46 !
purpuragula Lea. v. stenostoma Pfr., v, 48 !
mamilla Lea. C. lindoni Pfr., v, 45.
bizonalis Grat. Undent Pfr.
pudibunda Beck. immersa Gundl.
hebe Dh. C. bartlettiana Pfr., v, 45.
C. dennisoni Pfr., v, 44. C. melanocephala Gundl., v, 46.
Juliana Poey. f. perelevata Pils.
C. nigropicta Arango, v, 47.
Section Dialeuca Albers, 1850.
Dialeuca ALB., Die Hel. 1850, p. 114 (for H. nemoraloides) .
Leptoloma ALB.-MART., Die Hel. 1860, p. 136 (type H.fuseocincta).
W. G. BINN., Ann. N. Y. Acad. Sci., iii, p. 96, jaw and dentition
of fuscocincta; p. 107, dentition of gossel.
CEPOLIS. 183
Shell imperforate, rather thin, more or less trochoidal, varying
from high and pyramidal to low trochiform ; lip thin, slightly ex-
panded, a little widened and reflexed at the columella. Type C.
nemoraloides, pi. 56, fig. 5. (See also C. fuscocincta, pi. 56, fig. 6).
Animal light colored or dark as in H. alauda. Jaw, dentition,
etc., also as in Coryda (pi. 57, figs. 43, 48, jaw arid dentition of C.
platystyla).
This section might well be united to Coryda, from which it differs
only in distribution and the somewhat thinner shell. The typical
Dialeucas are from Jamaica, but a few species are from Navassa
(H. gaussoini), and the Cayman Is. (streatori, caymanensis) ; and the
closely allied H. phceogramma, of which I have seen a specimen in
Ponsonby's collection, is not yet located.
C. conspersula Pfr., v, 38. C. nemoraloides Ad., v, 40.
v. fuscocincta Ad., v, 39. v. pulchrior Ad., v, 41.
v. platystyla Pfr., v, 39. gossei (C. B. Ad.), Pfr., Rv.
v. virginea Ad., v. 39. C. gaussoini Tryon, v, 197.
C. subconica Ad., v, 40. C. streatori Pils., viii, 240.
gossei Pfr. ! C. caymanensis Mayn., viii, 241.
C. jacobensis Ad., v, 41. C. phseogramma Pfr., v, 42.
C. blandiana Ad., v, 41.
Section Hemitrochus Swainson, 1840.
Hemitroehus Sw., Malacol., p. 331, type H. hcemastomus=H. var-
ians. BINNEY, Terr. Moll., v, p. 174, and Ann. N. Y. Acad. Sci.,
iii, p. 90 (jaws and teeth of varians, troscheli, gallopavonis, rufoapi-
eata, graminieola, milleri). Polytcenia MARTENS, Die Hel., p. 129,
type H. multifasciata. Phcedra ALB., Die Hel., p. 100.
Shell globose-conoid or globose-depressed, solid, smooth or rib-
striate, opaque, variegated with bands or dots, the umbilicus narrow
or closed ; last whorl slightly descending. Lip blunt, simple or ex-
panded, thickened within, reflexed at columellar insertion. Type C.
varians, pi. 56, figs. 1, 2.
Jaw highly arched, with a median projection and sometimes a
median rib-like convexity (pi. 57, fig. 41, C. varians. PI. 57, fig. 46,
C. milleri).
Radula having long, narrow basal plates and broad, short central
cusps without side cusps on median and lateral teeth. Marginals
184 CEPOLIS.
with a large split inner cusp and a simple or bifid ectocone (pi. 57,
figs. 50, 51, C. varians).
Genital system having a long, slender penis branching into v. d.
and a long flagellum, and with a thread-like retractor attached low,
and inserted distally on the lung floor. Spermatheca duct free
below, but firmly bound to uterus above, with a long spermatheca
and a spiral twist in the duct near base of uterus. Dart sack long ;
mucus glands two, flat and glandular, inserted at base of dart sack-
Eye stalk retracted between branches of genitalia (pi. 52, fig. 14, C.
varians Mke.).
The shell in this section differs but little from that of Coryda and
Dialeuca ; and while quite distinct from the typical forms of Pla-
gioptycha, there are a number of species so intermediate in character
that they may be placed as well in one as in the other group. Ana-
tomically there are no differences of more than specific worth
between these groups, unless the larger (though still very weak)
penis retractor of Remitrochus be considered such. Many of the
species are excessively closely allied.
There are two groups of species: the Cuban, consisting of com-
pact forms of the type of H. cesticulus ; the Bahama group varying
from globose-conical like H. varians to depressed and rib-striate, H.
maynardi.
Species of Florida and Bahamas.
C. varians Mke., v, 24. C. gallopavonis Val., v, 27.
carnicolor Pfr. v. calacaloides Pi Is., v, 28.
submeris Migh. C. troscheli Pfr., v, 28.
rhodocheila Binn. tenuicostata Dkr.
polychroa Binn. v. calacala Weinl., v, 29.
hwmnstomus Sw. C. multifasciata W. & M., v, 30.
C. xanthophaes Pils., viii, 242. f. polytseniata Pils., v, 30.
C. milleri Pfr., v, 25. C. filicosta Pfr., v, 30.
C. constantior Weinl., v, 26. C. brownii Pils., v, 29.
C. caribsea Weinl., v, 26. C. maynardi Pils., viii, 241.
Cuban Species.
C. gilva Fer., v, 31. C. fuscolabiata Poey, v, 34.
corrvgata Pfr. subfusca Poey not Bk.
v. tephrites Morel., v, 31. v. morbida Morel., v, 35.
CEPOLIS. 185
O. lucipeta Poey, v, 32. C. maculifera Gut., v, 35.
picturata Poey not Ad. C. sauvallei Arango, v, 37.
lepida Poey. C. comta Gundl., v, 34.
bellula Poey. C. araplecta Gundl., v, 35.
penicillata Poey not Gld. C. rufoapicataPoey, v, 36.
newcombiana Poey. C. graminicola C. B. Ad., v, 36.
v. velasqueziana Poey, v, 32.
v. cesticulus Gundl., v, 33.
Section Plagioptycha Pfr., 1 856.
Plagioptycha PFR., Mai. BL, 1856, p. 135 (for indistincta, albers-
iana, duclosiana, bahamensis, strumosa, loxodon, monodonta).
MART, in Alb. Die Hel., p. 145 (type H. loxodon}. W. G. BINNEY,
Ann. N. Y. Acad. Sci., iii, p. 95, jaws and dentition of loxodon,
albersiana, monodonta, duclosiana, diaphana, macroglossa.
Shell umbilicate or imperforate, thin, depressed-globose or de-
pressed, the last whorl deflexed in front. Aperture transversely
oblong or lunate ; outer lip expanded or simple ; and either on the
Jloor of the whorl within the mouth, or on the columellar lip, there is
an oblique nodule or fold of callus, sometimes reduced to a slight
trace only. Type C. loxodon Pfr. (See pi. 56, figs. 8, 9, C. duclos-
iana).
Animal light colored, externally similar to Hemitrochus.
Jaw high arched, with a median projection (pi. 57, fig. 42, C. sal-
vatoris).
Kadula (pi. 57, fig. 47, C. sahatoris) similar to that of Hemitro-
chus, but with the cusps rather more acute.
Genital system as described for Hemitrochus (pi. 52, fig. 15, C.
salvatoris).
Plagioptycha is probably nearest to the ancestral forms whence
the modern sections of this genus arose. Its habits are terrestrial
and the dentition is somewhat less abnormal than in Coryda and
typical Hemitrochus. Moreover, characteristic forms of Plagiopty-
cha are found in the Miocene Silex Beds of Tampa, Florida, (jET.
latebrosa Dall, instrumosa Dall), with other species (JET. crusta and
H. diespiter of Dall) which would probably be classed in the mod-
ern section Eurycampta, although it is obvious that these latter
Miocene forms are more intermediate between Eurycampta, Jean-
neretia and Plagioptycha than any living species. In the Miocene
UNIVERSITY
186 CEPOLIS.
we are evidently near the horizon where the paths of the various
sections of the genus Cepolis diverged, although the better defined
forms of the genus no doubt have older roots.
The species of Plagioptycha are numerous and especially charac-
teristic of the Bahamas, extending south to Hayti and the Virgin
Islands. In Miocene times they extended to the (then) island of
Florida, but later became extinct there, for the present Floridian
land shell fauna is not directly descended from that of the Miocene
island. Some forms of Plagioptycha approach Hemitrochus, and
others are near Cysticopsis, so that the grouping, as in many of these
sectional divisions, is somewhat arbitrary.
Umbilicate species, the columellar lip expanded, not adnate to base
(Bahamas).
C. indistincta Fer., v, 14. C. bahamensis Pfr., v, 18.
v. disculus Dh., v, 15. v. holostoma Pils., v, 18.
v. chromochila Pils., v, 15. C. sargenti Bid., v, 18.
C. strumosa Pfr., v, 15. C. duclosiana Fer., v, 19.
C. riisii Pfr., v, 16. v. salvatoris Pfr., v, 19.
C. platonis Pfr., v, 16. v. abacoensis Mts., v, 20.
C. albersiana Pfr., v, 17. C. macroglossa Pfr., v, 20.
C. loxodon Pfr., v, 17.
Imperforate species, with adnate columellar lip (Hayti to Virgin Is.).
C. monodonta Lea, v, 21. C. diaphana Lam., v, 22.
v. acuminata Pfr., v, 21. G. santacruzensis Pfr., v, 23.
haitensis W. & M., v, 21. C. phsedra Pfr., v, 23.
C. nemoralina Pet., v, 22. justi Pfr.
f. intensa Pils., v, 22.
Section Cysticopsis Morch, 1852.
Cysticopsis MORCH, Catal. Yoldi, p. 2 (proposed for cubensis Pfr.
only). Pilsbry, Man. Conch, v, p. 10, Cuban species. Not Cysti-
copsis MARTENS, Die Hel. 1860, p. 144!
Shell globose-depressed, thin, semitranslucent, unicolored or spir-
ally banded and dotted, the last whorl not descending in front;
aperture large, broadly lunate, the lip thin, not in the least expanded
or reflexed, dilated and appressed at the umbilical insertion. Type
C. cubensis Pfr. pi. 56, fig. 7.
i
CEPOLIS-POLYMITA. 187
Animal light colored, otherwise as in Coryda. Jaw high arched,
smooth, with a large median projection (pi. 57, fig. 44, C.
cubensis*).
Radula long. Teeth with long, narrow basal plates, the median
and lateral teeth without side cusps, transition and marginal teeth
with an ectocone, the meso- and ento-cones united into a large bitid
cusp (pi. 57, fig. 40, C. cubensis). The teeth of pemphigodes figured
by Binney are of the same type, but with shorter cusps.
Genitalia (of C. cubensis, pi. 52, fig. 16), as in ffemitrochus, ex-
cept that I see but one lobe to the mucus gland ; the very long penis
is apparently without retractor. Possibly the second lobe of the
mucus gland was inadvertently removed in my dissection.
Morch, in his original publication of this group, mentioned only
one species, H. cubensis Pfr. The authors of the second edition of
Die Heliceen were therefore not justified in naming H. tenerrima as
type of Cysticopsis, and excluding cubensis from the roll of its mem-
bers. On an earlier page of this work (p. 65), the writer has sepa-
rated the Jamaica species formerly referred to this genus, and has
shown them to belong to a separate genus, Zaphysema, near the Sagda
group. The external features of the animal, its jaw, teeth and
genitalia, all support this division.
Cysticopsis is allied on one side to the Cuban band-dotted forms
of Hemitrochus, and on the other to the group of Plagioptychas
like diaphana.
C. cubensis Pfr., v, 10. C. auberi Orb., v, 11.
lanieriana Orb. C. pemphigodes Pfr., v, 12.
trifasciella Beck. pelliculata Poey.
pictella Beck. C. lescaillei Gundl., v, 13.
N.pulchella Beck. C. luzi Arango, v, 13.
penicillata Old., v, 33. C. lassevillei Gundl., v. 14.
ncevula Morel., v, 34. C. pellicula Fer., v, 14.
C. comes Poey, v, 11. 0. jaudenesi Cisn., v, 14.
C. letranensis Pfr., v. 11. C. hjalmarsoni Pfr.,v, 12.
Genus POLYMITA Beck, 1837.
Polymita BECK, Index Moll., p. 44 (picta, globulosa, versicolor,
carnicolor'). GRAY, P. Z. S., 1847, p. 171, type H.picta. MARTENS,
Die Hel., 1860, p. 145, type H. muscarum. W. G. BINNEY, Ann.
188 POLYMITA.
N. Y. Acad. Sci. iii, p. 89 (Jaw and dentition). POEY, Meraorias
sobre la Hist. Nat. Cuba, ii, pi. 7, f. 5. PILSBRY, Man. Conch., v,
p. 52.
Shell subglobular, brilliantly colored, rather thin but solid, imper-
forate ; whorls few (about 4), the last but little deflexed ; aperture
rounded, slightly lunate, the peristome simple, not expanded or re-
Jlexed except at axis, where it is reflexed and adnate over the umbil-
ical region ; axis solid. Type P. picta, pi. 56, fig. 10.
Animal (of P. picta) black above, slaty below; evenly granulated
throughout, without dorsal grooves, facial groove or foot margin,
the tail rounded above, not grooved ; sole not in the least divided,
mantle edge thickened but without lobes.
Jaw arcuate, moderately solid, smooth (PI. 51, fig. 8, P. picta).
Radula short and wide, the teeth all of the same form, and in
v-shaped rows, formula about 85.1.85. Basal plates long and narrow;
cusps situated far backward, and projecting well over the posterior
margin; all teeth tricuspid, the three cusps united into a broad, tri-
dentate gouge-shaped cutting edge. (PI. 51, fig. 5, central with four
lateral teeth; fig. 6, group of outer laterals; fig. 7, two extreme
marginal teeth of P. picta).
Genital system (pi. 51, fig. 4, P. picta) altogether like that of
Cepolis. The vagina is long, with a long stalked spermatheca;
dart sack large, its head marked off by a constriction and united by
connective tissue with the vagina ; at root of d. s. there is a mucus
gland composed of two oval, flat glandular lobes. Penis slender,
with a long flagellum, and apparently no retractor muscle; eye-
stalk retracted between branches of genitalia.
Distribution, Cuba. Habits arboreal.
The shell in this group resembles that of Hemitrochus, except
that the lip is neither expanded nor thickened within. The genital
system is entirely that of Hemitrochus. The radula is excessively
peculiar in having the side cusps as long as the middle cusp and
united with it to form a broad, tridentate gouge, all three cusps be-
ing subequally developed on all the teeth.
This type of radula may be compared with that of Orthalicus t
Oxychona, Papuina, and especially with Amphidromus; all being
arboreal genera, which have independently evolved the same gen-
eral type of teeth.
OXYCHONA.
P. picta Born, v, 53. P. muscarum Lea., v, 54.
venusta Gmel. globulosa Fer.
sulpliurosa Morel., v, 54. carnicolor Orb.
L. tiara Martyn. v. subbrocheri Pils., v, 55.
P. versicolor Born., v, 54. P. brocheri (Gut.) Pfr., v, 55. -
Ipictoria Perry. brocheroi Arango.
? cincta Perry.
Genus OXYCHONA Morch, 1852.
Oxychona MORCH, Cat. Yoldi, p. 14, type H. bifasciata. PILSBRY,
Man. of Conch., v, p. 128. MARTENS, Biol. Centr. Amer., Moll., p.
152. Oeotrochus, Leptoloma, Corasia, Axina and Eurycratera of
authors. Leptarionta CROSSE & FISCHER, Moll. Mex. i, p. 253.
Shell rather shining, thin and light colored, with spiral brown
bands, umbilicate or closed, the spire conic or depressed and merely
convex,the last whorl varying from acutely keeled to subangular.
Surface smoothish, often microscopically striate or granular. Aper-
ture oblique, the lip expanded or reflexed, rather thin, not toothed.
Type 0. bifasciata, pi. 45, fig. 8. (See also pi. 45, figs. 1,2, 0. costa-
ricensis. PI. 45, figs. 3., 4, 5, 0. altispira. PI. 45, figs. 9, 10, 0.
trigonostoma v. stolliana.*)
Animal (of 0. trigonostoma, pi. 45, figs. 9, 10) quite elongated,
the tail surmounted by a conspicuous serrate keel.
Jaw (of 0. bifasciata, pi. 51, fig. 11), rather thin, arcuate, with
no median projection, having about 17 unequal ribs distributed
over its entire extent and denticulating the margin.
Radula (of 0. bifasciata, pi. 51, fig. 10, central with 6 adjacent
laterals, fig. 9, 7th to 13th laterals, with one marginal, and fig. be-
low the latter, a lateral seen in profile) pavement-like, with v-shaped
rows of nearly similar teeth. Centrals with an oblong squarish
basal plate bearing one cusp springing from its middle, spreading
into a spatulate form, and projecting far over the posterior end of
of the basal plate on all sides. Laterals similar, but having the
entocone indicated by a notch in the basal plate near its posterior
angle, and bearing a minute basal ectocone. Outwardly, this ecto-
cone increases in size, and becomes split on the marginal teeth,
which are otherwise like the laterals.
Distribution, Brazil to Mexico. Habits arboreal, as far as
known.
190 OXYCHONA.
The prominent features of this group are the smooth, thin, light-
colored and banded shell which is usually of a markedly trochiform
contour, but sometimes depressed, the periphery angular; the ribbed
jaw ; the extremely peculiar radula, with enormously widened and
enlarged middle cusps, and minute, basal side cusps. When the
radula is torn, the teeth part readily along their lateral faces, but
adhere in chain-like longitudinal rows.
The radula, as well as the jaw and shell, is comparable to that of
Papuina (cf. p. 137, pi. 37, fig. 10), but although the superficial re-
semblance is great, the two are really totally distinct, the broad
cusps of Papuina being formed by the united ento-, meso- and ecto-
cones, whilst in Oxycliona the mesocone only is modified, the side
N cusps becoming obsolete. Polymita also has a slightly similar but
morphologically different dentition. The peculiar type of teeth in
these three genera has evidently been independently evolved in
each, from the usual tricuspid type. It seems to be correlated with
arboreal habits. Compare also the radula of Otostomus.
The affinities of Oxychona are uncertain ; but it will probably
prove to be a member of the Belogona, distinguished from Helix
by its Papuina or Corasia like shell, and the peculiar teeth. Prob-
ably in this group, as in Papuina, some species will be found to
have a more normal type of dentition. The Mexican forms which
have been placed in this group are still unknown anatomically, but
the animal of 0. trigonostoma has been figured with a toothed keel
on the tail, such as occurs in the genus Lysinoe.
Messrs. Crosse & Fischer proposed the section Leptarionta for two
species, bicincta and flavescens ; but as they state that they had not
seen flavescens, their group was evidently founded on bicincta, the
first species described by them.
The first four species are from Brazil ; the others are from
Guatemala and Costa Rica to the province of Vera Cruz, E.
Mexico.
O. bifasciata Burrow, v, 129. O. zhorquinensis Ang., v, 132.
pyramidella Spix, Wagner. O. trigonostoma Pfr., v, 132.
bosciana Fer. /. elevatoconica C. & F.
blanchetiana Moric. lalliana Tristr.
O. lonchostoma Mke., v, 130. /. salleana Pfr.
O. gyrina Val.,v, 131. obscura C. & F.
O. pileiformis Moric., v, 131. /. intermedia C. & F.
LYSINOE. 191
O. trigonostoma Pfr., v, 132. O. guillarmodi Shutt., v, 133.
/. subunicolor C. & F. O. costaricensis Roth, v, 134.
stoliiana Mts. /. virginea Anc.
/. freytagiana Dohrn. /. steiniana Anc.
O. altispira (Dohrn) Mts. /. boucardi Ang.
O. bicincta Pfr., iv, 75. O. adela Ang., v, 135.
Genus LYSINOE H. & A. Adams, 1855.
Aglaja ALB., Die Hel., 1850, p. 107, sole species H. ghiesbreghti.
Aglaia Alb., v. MART., Die Hel., 1860, p. 122, in part, exclusive
of "type" and all but first species. Not Aglaia Renier, 1804,
Eschscholtz, 1825, or Swainson, 1827. Lysinoe, H.& A. AD., Gen.
Rec. Moll, ii, p. 203, for ghiesbreghtii and hogoleuensis (June, 1855).
v. MARTENS, Biol. Centr. Araer., Moll., p. 145. Odontura FIS-
CHER & CROSSE, Miss. Scient. Mex., Moll., pp. 211, 242, 1870, for
ghiesbreghti and eximia. Humboldtiana v. IHERING, Morphol. u.
Syst. des Genitalapparates von Helix, in Zeitschr. f. wissenschaftl.
Zoologie, liv, p. 172, 1892, type H. humboldtiana.
See for anatomy, FISCHER & CROSSE, 1. c. ; W. G. BINNEY,
Bull. Mus. Comp. Zool., v, p. 336, pi. 2, f. i. K. ; v. MARTENS, I. c.,
pi. 8, f. 4.
Shell large, globose or depressed globose, beset with granules and
sometimes hairs, brownish or yellowish with dark spiral bands ; um-
bilicus open or partly covered ; aperture lunate, lip margins more
or less reflexed, at least the columellar margin. TypeL. ghiesbreghti,
pi. 45, fig. 7. See also pi. 58, fig. 75, L. humboldtiana var. badio-
cincta.
Animal large, coarsely granulose, the tail surmounted by a knobbed
or serrate keel (pi. 45, fig. 7, H. ghiesbreghti).
Jaw arcuate, soAid ? with 6-11 spaced ribs (pi. 60, fig. 9, L. eximia\
Radula havingtfifiicuspid median teeth, the stout cusp shorter
than the basal plates. Lateral teeth with a small ectocone. Mar-
ginals haxing a long oblique bifid inner cusp and a small ectocone,
(pi. 60, fig. 5, L. humboldtiana}.
Genital system having the retractor and epiphallus inserted at
apex of the short penis ; epiphallus continued in a long flagellum.
Spermatheca duct varying from moderate to very long, without di-
verticulum. Vagina bearing two equal, symmetrically placed dart
sacks, above the insertion of which three club-shaped mucous glands
192 GLYPTOSTOMA.
are independently inserted on the vagina. Darts unknown. (PI. 60,,
fig. 8, L. ghiesbreghti).
The three species present the same type of genitalia. The shell
is similar to Campylcea or Epiphragmophora, but the doubling of
the dart sack, and the number of the mucus glands, independently
inserted on vagina, are features which can only be compared to the
Xerophiloid and Fruticicoloid groups. The serrate keel of the tail
is a curious feature of this group, but something like it occurs also
in Oxyehona trigonostoma, the internal anatomy of which is un-
known. The jaw and teeth of Lysinoe are of the normal Helix
form. Distribution, Southern Texas (humboldtiana) to Guatemala
and Honduras (ghiesbreghti).
The name Aglaja Alb. is thrice preoccupied in zoology. Lysinoe
was proposed for two species, the second of which, Helix hogoleu-
ensis Le Guill., belongs to the prior genus Rhyssota Alb. We there-
fore follow von Martens' excellent precedent in considering H. ghies-
breghti the type of Lysinoe. Helix humboldtiana agrees entirely
with ghiesbreghti in anatomy, so the sectional name, Humboldtiana
v. Iher., becomes a synonym. Odontura is preoccupied.
L. ghiesbreghti Nyst., iv, 75. L. humboldtiana Fer., iv, 260.
/. subaurantia, v. Mart. v. hegewischi v. Mart.
/. strubelli Bttg. v. hogeana v. Mart.
/. fulvostraminea, v. Mart. v. buffoniana Pfr.
/. bizona, v. Mart. matronuta Uhde.
/. rufozonata v. Mart. v. badiocincta Wiegm.
L. eximia Pfr., iv, 75.
v. stolli, v. Mart.
Genus GLYPTOSTOMA Bland & Binney, 1873.
Glyptostoma BLIX & W. G.B M Proc. Acad. Nat. Sci., Phila., 1873,
p. 244 (June 3, 1873). See for anatomy, BINNEY, t.c.pl 1, f. 1,3 ;
Am. Journ. Conch., vii, p. 190, pi. 17, f. 3, 4; Proc. Acad., Phila.,
1875, p. 219, pi. 16, f. 4, pi. 13, f. 6, and 1876, p. 190, pi. 6, f. H. ;
Terr. Moll., v, p. 373, pi. 14, f. D.
Shall discoidal with slightly convex spire of about 6 whorls and
broadly open umbilicus showing all the whorls; smoothish, dark
and opaque ; last whorl rather tubular, hardly descending in front.
Aperture round-lunar, oblique, the lip simple and acute, neither
EPIPHRAGMOPHORA. 193
thickened nor expanded, margins approaching ; parietal wall densely
spirally striated. Type G. newberryanum, pi. 31, figs. 36, 37.
Jaw low, wide, slightly arcuate, without median projection, with
about 15 ribs extending nearly to the ends. (PL 31, fig. 38.) Rad-
ula long and narrow, formula 30.17.1.17.30 ; basal plates of medran-
and lateral teeth long, the mesocones about the same length, side
cutting points developed. Marginal teeth with shortened basal
plates, the inner cusp rather short and stout, ectocone small (pi. 31,
fig. 40, middle with 1st lateral, 23d, 24th, 25th, 37th and 47th mar-
ginal teeth).
Genitalia (pi. 31, fig. 39) characterized by a stumpy penis with
short, obtuse flagellum ; dart sack obsolete or absent, but one club-
shaped mucus gland, like that of Epiphragmophora fidelis present ;
spennatheca duct long, bound to oviduct, but free above and
below.
A monotypic genus, allied to Epiphragmophora, but differing im
the shell, which has much the form of the typical Zonites or Patula,.
in the wide many-ribbed jaw, and in the genital system, which appar-
ently lacks a dart sack, although the mucus gland is retained. Fur-
ther examination is needed, to ascertain whether any trace of the
dart sack is present, for I suspect this will prove to be the case.
The single species is common around San Diego, on southerly ex-
posed hill-sides under rocks.
G. newberryanum W. G. B., iii, 110. Los Angeles, Cal., to Todos
Santos Bay, L. Cal.
Genus EPIPHRAGMOPHORA Boring, 1875.
Epiphragmophora DC-RING, Bol. Acad. Nacional de Ciencias Cor-
dova i, p. 446, for jEf. hieronymi and H. cuyana. -f Angrandiella
ANCEY, Conchologists' Exchange, i, p. 20, Nov. 1886, type H.
angrandi. -\-Pcecilostola ANC., L c., type H. farrisi (not Pceeilostola
Stal, 1870, Hemiptera, Poecilostola Chaud., 1871, Coleoptera, or Pce-
cilostolus Giinth., 1863, Reptilia) = Pilsbrya ANC., t. c., p. 54, Apr.,.
1887, same type. -\-Helminthoglypta ANC., Conch. Exch., i, p. 76,
June, 1887, type H. tudiculata. + Micrarionta ANC., Le Naturaliste,
Dec., 1880, p. 334, type H.facta. -{-Aglaia of American authors, not
Albers. -\-Arionta of authors, not Leach -\-Lysinoe PILS., check list,
not H. & A. Adams-)- Campylcea (American species) v. IHERING,
Morphol. u. Syst., not of Beck, -f Euparyphaof American authors
not Hartmann.
13
194 EPIPHRAGMOPHORA.
?-|- Ccelospira ANC., Conch. Exch., i, p. 20, type H. maeneili (not
Ccelospira Hall, 1858, HreichioipodeC)=Averellia ANC., 1. e., p. 54,
Apr., 1887, same type, -f- Trichodiscus STREBEL, Beitr. Mex. Land-
und Siissw.-Conch., iv, p. 32, 1880 (not of Ehrenberg, Infusoria)=
Trichodisdna v. MARTENS, Biol. Centr. Amer. Moll., p. 133, March,
1892 ; type H. coactiliata.
See for anatomy W. G. Binney, Terr. Moll., vol. v (figures not
always reliable !). Semper, Reisen, etc., p. 242. Doring, Bol. Acad.
Nac. Sci., Cordova, i, and Estudos Sist. y Anat. sobre los Mol. pulm.
de los poises del Plata, Periodico Zool.,i, 1875, p. 129-204. Strebel
& Pfeffer, Beitr. Mex. Moll., pt. iv. HEYNEMANN, Mai. Bl., xv,
pi. iv, fig. 4.
Shell varying from globose to subdiscoidal, rarely keeled, umbili-
cate or imperforate, with 4 to 7 whorls. Surface generally decus-
sated, malleated or hirsute ; the texture varying from corneous to
chalky; generally variegated, having a supraperipheral band, some-
times 3-banded. Aperture lunar, the lip expanded or reflexed,
dilated at columellar insertion, toothless ; but a few species have a
columellar nodule or internal plicae. Type E. hieronymi Doring,
pi. 59, fig. 77. (See also pi. 58, figs. 57 to 74).
Animal granulated as usual, with a pair of dorsal grooves and
no distinct facial grooves, the tail rounded above, not keeled nor
grooved. Sole undivided (pi. 45, fig. 6, E. fidelis; pi. 59, fig. 76,
E. nickliniana, showing atrium extruded, and the characteristic
granulation of foot, not well shown in the fig. of E. fidelis). Eight
eye retractor passes between branches of genitalia.
Jaw arched, with 3 to 8 stout ribs denticulating the margins (pi.
59, fig. 78, E. semiclausa; fig. 80, areolata ; fig. 83 nickliniana;
fig. 84, fidelis; fig. 85, arrosa; pi. 52, fig. 17, E. eordovana).
Radula characterized by median and lateral teeth without side
cusps ; marginals with the entocone+mesocone forming a long com-
pound cusp, ectocone simple, never bifid. (PI 60, fig. 10, E.fidelis.
Fig. 7, E. nickliniana. Fig. 4, E. areolata}.
Genitalia characterized by a slender penis continued in an epi-
phallus which bears the retractor; flagellum present. Dart sack
short, its apex neither constricted nor attached, containing a two-
edged dart, pi. 59, fig. 82, E. mormonum. Mucus gland either (1)
single and club-shaped, seated on dart sack, pi. 59, fig. 81, or (2)
single but dividing above into two bulb-bearing branches, pi. 59,
EPIPHRAGMOPHORA. 195
figs. 79, 87, or (3) double and glandular with threadlike ducts in-
serted at root of dart sack, one gland bound to dart sack, one to
vestibule or base of penis, pi. 59, fig. 89, glands torn from their
attachments. The spermatheca is globose, its duct often bearing a
diverticulum.
Distribution, British Columbia southward to Argentina, mainly
confined to the Pacific drainage, but spreading to the Gulf slope
in Central America.
The genus Epiphragmophora, while allied to the Helices of Japan,
is distinguished from them by the non-sacculated mucus glands and
some shell characters. It is also allied to Cepolis, a West Indian
genus which is characterized by its flat, two-parted mucus gland,
peculiarly formed dart sack attached at apex to vagina, and ribless
jaw. The middle American genus Lysinoe is similar to Epiphrag-
mophora in features of the shell, but differs widely in the duplica-
tion of the dart sack (elsewhere developed only in Hygromia and
Helicella'), in the three club-shaped mucus glands independently in-
serted on the vagina, and in the serrate keel of the tail. A still
nearer ally of Epiphragmophora is Glyptostoma, characterized by
the simple acute lip of the peculiar shell, and the decadence of the
dart sack.
The diverticulum of the spermatheca duct is present or absent in
closely allied species, just as we find it in other genera. The shell
varies so much that no generic diagnosis can be framed from it
alone, which would cover all forms of Epiphragmophora and still
exclude species of other groups. This difficulty is not encountered
when we diagnose from the soft anatomy, which presents extremely
characteristic and readily recognized features.
Dr. von Ihering, in his essay on Helix, refers this New World
series to Campylcea ; but as the other groups studied by him belong
to the Belogona siphonadenia of my arrangement, he was not aware
of the value of the characters upon which the Belogona are split
into two great groups, and his knowledge of the American forms
was wholly second-hand from figures, not dissections. I feel confi-
dent that if v. Ihering had actually dissected American and East
Asiatic types, he would have taken a different view, and one more
in accordance with the opinions of Semper and the writer.
The members of this genus have hitherto been placed in Arionta,
Euparypha and Aglaia by American authors. Semper, as long ago
as 1880 (Reisen im Archip. Phil. (2) iii, Landmoll., p. 245), emphat
196 EPIPHRAGMOPHORA.
ically stated his belief that the American should be separated from
the European " Arioutas ;" and my own studies have converted me
to the same opinion, although before my dissections were begun, I
had thought otherwise. It is sufficient to say here that in the
American, as in the East Asiatic types of belogonous Helices, the
mucus glands are globular or flat bodies of glandular texture, in-
serted upon the dart sack or at its base; in the European forms
these glands are always tubes of equal diameter throughout their
length, and inserted upon the vagina above the dart sack. If my
division of the belogonous Helices upon this character be correct,
Helix, Helicigona (- Campy lcea-\- Arionta vera), Fruticicola, Xerophila
etc., are all more nearly allied to each other than any European
Helices are to the American Ariontas, so-called.
The American types are closely allied to the large Helices of
Japan and China in anatomical features. The resemblance in shell
characters of the Californian and European species is astonishing,
but I do not doubt that it is due to purely secondary modification,
which has moulded the shells to a deceptive likeness, but left un-
changed the genitalia to tell more faithfully the story of their
lineage.
Synopsis of Sections of jEpiphragmophora.
Although not nearly so homogeneous a group as Cepolis, Helix
or Helicigona in anatomy, this genus exhibits but little modification
in shell characters. The sections here admitted, although natural
groups, have but little systematic rank. Averellia and Trichodiscina
are not sufficiently known anatomically for us to be certain that
they belong here, but the probabilities are strong that they do.
* South American forms.
Section EPIPHRAGMOPHORA Doring. Shell umbilicate, brown,
calcareous, with one supraperipheral band, peristome expanded,
nearly circular. Epiphragm solid, calcareous; jaw four ribbed;
dart sack lengthened, with two globose, stalked mucus glands ; sper-
matheca short-stalked. Type H. hieronymi, pi. 59, fig. 77 ; (see also
E. cuyana, pi. 58, figs. 68, 69.)
Section PILSBRYA Ancey. Shell imperforate or umbilicate,
malleated, similar to that of Helminthoglypta. Epiphragm mem-
braneous, jaw 4-5 ribbed (pi. 59, fig. 78, E. semiclausa') ; median
EPIPHKAGMOPHORA. 197
and lateral teeth without side cusps; marginals tricuspid ; dart bi-
angulate; mucus glands as in Epiphragmophora. Type E. farrisit
pi. 58, figs. 58, 59. See also E. petasemis, pi. 58, figs. 60, 61.
We regret to say that this group is not well distinguished from
the preceding.
Section ANGRANDIELLA Anc. Shell depressed, umbilicated, with
a toothlike prominence within the basal lip, marked by an external
pit. Type E. angrandi, pi. 58, fig. 57.
** Central American and Mexican forms.
Section AVERELLIA Ancey. Shell discoidal with concave spire
and wide umbilicus, hirsute, the last whorl deeply descending in
front ; scrobiculate behind the aperture above and below ; aperture
subhorizontal, biplicate within, peristome narrowly expanded. Type
E. macneili Crosse.
Section TRICHODISCINA Martens. Shell depressed, with open
umbilicus and deflexed last whorl, granulate and hairy ; aperture
small, oblique or subhorizontal, toothless, the lip expanded. Jaw
ribbed (pi. 52, fig. 17, E. cordovana). Type E. coactiliata, pi. 58,
figs. 70, 71.
*** North American forms.
Section MICRARIONTA Ancey. Shell globose or globose depressed,
one or many banded, the bands sometimes interrupted ; substance
rather calcareous. Mucus glands two, with threadlike ducts, one
lengthened, adherent to and spread upon the vagina or base of
penis, its duct entering vagina at root of dart sack ; the other
shorter, usually adherent to dart sack, on base of which its duct
enters. Radula with rather short basal plate and wide mesocones,
no ectocones on middle and lateral teeth. Marginals with a sub-
bifid inner and simple outer cusp (pi. 60, fig. 4, areolata). See
pi. 59, figs. 89, E. areolata. Type E. facia gabbi, pi. 58, figs. 73,
74. (See also E. areolata, pi. 58, figs. 66, 67.)
Section HELMINTHOGLYPTA Ancey. Shell globose or depressed,
its surface either simply striated, decussated or malleated. Mucus
gland one, split into two bulb-hearing branches, and inserted on the
dart sack. (PI. 59, fig. 47, E. arrosa; fig. 87, E. traskii v. cayama-
censis ; fig. 88, E. nickliniana*). Radula with basal plates longer
than cusps of median teeth, middle and lateral teeth without side
198 EPIPHRAGMOPHORA.
cusps, marginals with a bifid inner and simple outer cusp (pi. 60,
fig. 7, E. nickliniana). Type E. tudiculata, pi. 58, figs. 62, 63.
Section MONADENIA Pilsbry. Shell with depressed body whorl,
often more or less keeled or angular, the spire either low or conical.
Surface with growth striae. Mucus gland one, undivided, club-
shaped, its terminal portion glandular, and inserted on the dart
sack. (PI. 59, fig. 81, E.fidelis ; pi. 59, figs. 82, 86, E. mormonum).
Radula with no side cusps on middle, lateral or transition teeth ;
marginals with bifid inner and simple outer cusp (pi. 60, fig. 10>
E.fidelis.') Type E.fidelis, pi. 58, fig. 72. See also pi. 58, figs. 64,
65, E. mormonum.
SOUTH AMERICAN SPECIES. Igp^hragmophora-j-Pilsbrya.
E. alsophila Phil., iv, 78. E. higginsi Pfr., iv, 79.
E. audouini Orb., iv, 81. farrisi Hig., not Pfr.
v. oresigena Orb. E. huancensis Ph., iv, 79.
E. claromphalos Dev. & Hpe., E. jaspidea Pfr., iv, 79.
iv, 80. plaiysoma Pils., vi, 104.
v. mesomphalos Mor. E. macasi Higg., iv, 81.
E. clausomphalos Dev. & Hpe., E. monographa Dor.
iv, 78. E. patasensis Pfr., iv, 81.
E. cuyana Strob., iv, 78. /. minor Pfr.
E. diluta Pfr., iv, 80. E. pelliscolubri Ph., iv, 80.
E. estella Orb., iv, 78. E. semiclausa Mts., iv, 80.
v. tucumanensis Dor., iv, 78. E. trenquelleonis Grat., iv, 82.
E. farrisi Pfr., iv, 77. E. trigrammephora Orb., iv, 80.
E. gueinzii Pfr. E. tschudiana Ph., iv, 77.
E. hidalgonis Dor. E. yocotulana Dor., iv, 81.
E. hieronymi Dor., iv, 78.
(Shell depressed, umbilicate, with a tooth within the basal lip.
Angrandiella.)
E. angrandi Morel., v, 96. Peru.
MIDDLE AMERICAN SPECIES.
(Shell hirsute, depressed, with large umbilicus, 2,-grooved behind lip
and 2-plicate within. Averellia.)
E. macneili Crosse, v, 96. Costa Rica.
(Shell depressed, subdiscoidal, with wide umbilicus ; hirsute. S.-E.
Mexico, Cent. Amer. Trichodiscina.)
EPIPHRAGMOPHORA.
199
\\
E. coactiliata Fer., iii, 49.
tceniata Nyst.
nystiana Pfr.
bridgesi Try., not Newc.
parkeri Tryon.
E. cordovana Pfr., iii, 49.
fobsita Pfr.
E. suturalis Pfr., iii, 49.
v. pressula Morel., iii, 50.
almonte Tristr.
almonteana F. & C.
E. oajacensis Koch, iii, 50.
E. sumichrasti C. &. F., iii,
E. sargi C. & F., iv, 80. [184.
NORTH AMERICAN SPECIES.
(Shell with conic or low spire, often keeled or angular. Monadenia.)
E. mormonum Pfr., iv, 70.
v. hillebrandi Newc., iv, 70.-
E. circumcarinata Stearns, iv, 70.
. fidelis Gray, iv, 69.
nuttalliana Lea.
/. minor W. G. B.
/. flava Hemph.
/. subcarinata Hemph.
v. infumata Gld., iv, 70.
(Shell globose or depressed, smooth, malleated or granose, not keeled.
Helminthoglvpta.^
dupetithoursi Dh,, iv, 71.
oregonensis Lea.
sequoicola Coop., iv, 71.
traskii Newc., iv, 71.
franki Coop., err. typ.
v. proles Hemph.
v. cuyamacensis Hemph.
v. tularensis Hemph.
exarata Pfr., iv, 73.
E. arrosa Gld., iv, 72.
ceruginosa Gld.
J. obscura Pils.
/. holderiana Coop.
/. stiversiana Coop.
E. californiensis Lea, iv, 119.
vincta Val.
v. nickliniana Lea., iv, 73.
arboretorum Val.
/. anachoreta W. G.<B.
r*
. ellipsostoma Pilsbry.
E. carpenteri Newc., iv, 71.
indioensis Yates.
remondii Gabb, not Tryon.
JB. coloradoensis Stearns,viii, 225.
^E. magdalenensis Strns, viii, 226.
E. rowellii Newc., iv, 72.
lohrii Gabb.
* *
E. californiensis Lea.
v. ramentosa Gld., iv, 73.
reticulata Pfr.
/. bridgesii Newc.
parkeri Tryon.
v. diabloensis Coop., iv, 74.
v. contracostse Pils.
E. ayresiana Newc., iv, 70.
E. tudiculata Binn., iv, 74.
/. cypreophila Newc., iv, 75.
200 EULOTA.
E. trypaiiomphala Pfr. E. remondi Tryon.
verrilli Anc. C. Ex., ii, 63.
(Shell globose or depressed, rather cretaceous. Micrarionta.)
E. gabbi Newc., iv, 77. E. ruficincta Newc,, iv, 72.
v. facta Newc., iv, 77.
m% % ^
1 -.^ \A
/E. intercisa W. G. B., iv, 74. '] ,E. stearnsiana Gabb, iv, 119.
crebristriata Newc. E. tryoni Newc., iii, 229.
/. albida Hemph. E. veitchii Newc., iii, 228.
/. callojunctis Pils. E. areolata Sowb., iii, 228.
v. redimita W. G. B., iv, 74. canescens Ad. & Rv., iii, 214.
/. castanea Hemph. /. exanimata Coop.
VA.xE. kellettii Fbs., iv, 119. E. levis Pfr., iii, 228.
/. multilineata Hemph. E. pandora; Fbs., iii, 228.
/. castanea Hemph. damascenus Gld.
Genus EULOTA Hartmann.
= Eulota HARTM. 1842, + Thysanota, Pleetotropis and Aegista
ALB. 1860, -j- Armandia ANC. 1883 and Pseudiberus ANC., -j-
Cathaica MLLDFP. 1884, -f Euhadra PILS. 1890, -f Dorcasia,
Hadra and Camcena of some authors.
Shell usually umbilicated, dextral or sinistral, varying from glo-
bose to depressed or lens-shaped, thin or solid, unicolored or few
banded ; surface striated, often with spiral lines. Aperture lunate
or angular,- the outer and basal lips generally expanded, columellar
lip dilated. Type E.fruticum, pl.j55, figs. 1, 2, 3, 4. (See also pi.
64, all figures except 7, 10-12.)
Animal with feebly tripartite sole, small right and left body-lobes
on mantel, a pair of dorsal grooves, and very weakly indicated
facial grooves or none ; sides of foot granulated as in Helix s. str. t
tail with an ill-defined central line or none. Right eye retractor
passing between branches of geuitalia (or in sinistral species the left
one).
Jaw arcuate, with 3 to 12 ribs dentating the concave margin
(pi. 65).
Radula normal, the mesocones about as long as basal-plates, side-
cusps weakly developed or represented by a lateral continuation of
main cutting-points; marginals with the inner cusp bipid (euto-plus
meso-cone), ectocone simple or bifid (pi. 65).
EULOTA. 201
Genitalia : penis extending into an epi phallus which sometimes
has, sometimes has not, a flagellum. Dart sack containing a round
or flat dart, and either borne on atrium or higher on vagina. Mucus
gland inserted on dart sack, or on an empty accessory sack communi-
cating with dart sack, and consisting of one or many sacculated or
glandular, long or oval branches, bound closely together and to the
dart sack. Spermatheca oval or globose, on rather a long duct,
which lacks diverticulum.
Distribution, Central Europe (one species) to Japan, south to E.
Indian Archipelago. Especially characteristic of Eastern Asia.
This genus differs widely from the European dart-bearing Helices
in having the one (often many-branched) mucus gland inserted on
the dart sack or on an accessory empty sack, and in the structure of
the gland itself. It is more closely allied to the American genera
Epiphragmophora and Cepolis in the structure of the mucus gland.
Eulota, as herein understood, comprises a great variety of shell-
forms and a large number of species; including, indeed, a consider-
able majority of the East Asiatic Helices. As in all other large gen-
era of Helices, the shell varies from globular to lens-shaped (see in-
troduction to this volume) ; and the several stages of contour, each
represented by a numerous progeny of species, have received names
which some writers use in a subgeneric, some in a generic, sense. In
my opinion, the former is the more philosophic view, as the shell
characters fade from one group to another, offering no sharp line of
demarkation throughout the genus, so far as I can see. As to the
anatomy, my dissections (a part of which are shown on plates 65,
66) tell clearly that no grounds for a division of the group into two
or more genera can be based thereon, unless the forms in which the
penis has a flagellum be separated generically from those lacking
this organ ; and I do not think it likely that the examination of
more material will add to the value of this feature. At all events,
I can find no character of shell or soft parts correlated with it, and
we are hardly justified in founding a genus on a single peculiarity,
unless it be one of greater value than this. Like Polygyra, Thersites
or Helix s. sir., the various anatomical divergencies, except as to the
flagellum, are fully covered by intermediate forms ; although, as a
whole, the genus is characterized by well-marked peculiarities which
would enable one to identify any of its members by an inspection of
the genital system alone, without the assistance of shell characters.
The main anatomical divergencies may be tabulated as follows
for the species now known anatomically :
UNIVERSITY
'V
^;* :
202 EULOTA.
Species. Flagellum. Dart sack. Acces. sack. Mucus gl. branches
Eulota fruticum, absent, inserted high, present, 2 to 4.
Eulotella similaris, absent, inserted low, absent, 2 sub-dividing.
Eulotella fodiens, absent, inserted high, present, 2 sub-dividing.
Eulotella duplocin eta absent, inserted low, present, 6 sub-dividing.
Acustatourannensis, absent, inserted high, ? sev'ral br'nchea
Acusta ravida, absent, inserted high, present, sevYl coal'sc'nt
Cathaicapyrrhozona, absent, inserted low, absent, many br'nches.
Cathaica przewalskii, ? inserted low, ? 2-branched.
Plectotropis vulv. present, inserted low, absent, 2-branched.
Mastigeulota kiang., present, inserted low, present, many br'nches.
Euhadra qusesita, present, inserted low, present, many.
Euhadra peliomph. present, inserted low, present, many.
Euhadra brandti, present, inserted low, present, many.
It will be noticed that Plectotropis, Mastigeulota and Euhadra
possess a flagellum ; the other sections lacking it, probably by de-
generation, as this organ was, in all likelihood, present in the primi-
tive Belogonous stock. Among the true Helices Eremina and
Euparypha have also lost the flagellum, by a parallel process.
Section Eulota Hartmann, 1842.
Eulota HARTM., Erd- und Sfisswasser-Gast. Schweiz, p. 179, type
H. fruticum. Helicella and Fruticicola, in part, of some authors.
Bradybcena, in part, BECK. Eulotella MOUSSON (where ?) of some
authors, type H. similaris Fer. Acusta ALB., Die Hel., 1860, p. 56,
type H. ravida Bens.
Shell globose-conoid or globose-depressed, umbilicated, rather
thin, the surface smoothish, generally with minute spiral strise; color
varying from sub-translucent white to light brown or yellowish,
often with a supra-peripheral band (rarely several bands). Whorls
5-6, convex, the last hardly descending in front. Aperture round-
lunate, toothless ; peristome thickened within and expanded, dilated
at columellar insertion. Type E. fruticum, pi. 55, figs. 1, , 3, 4. (See
also pi. 55, fig. 19, E. similaris t and pi. 55, fig. 5, E. duplocincta).
Jaw arched, with 4-11 ribs denticulating the concave margin.
(See pi. 65, fig. 4, similaris ; pi. 65, fig. 5, duplocincta; fig. 2, ravida).
Radula of E. fruticum having the median cusp of middle teeth as
long as basal-plate, side cusps subobsolete. Laterals with longer
mesocones. Marginals with long bifid inner and on the outer ones-
EULOTA. 203
bifid outer-cusps (pi. 65, fig. 3, similaris; pi. 65, fig. 6, duplocincta ;
pi. 65, fig. 1, ravida).
Genitalia (pi. 66, fig. 18, E.fruticum) : penis short, swollen, pass-
ing into a long epiphallus which receives vas deferens and retractor,
but lacks flagellum. Dart sack globose, containing a round, conical
dart (pi. 66, fig. 19), and communicating at base with an empty
accessory sack which bears the mucus glands ; these consist of 2-4
oval glands, closely bound together, and flattened on their adjacent
sides, their ducts uniting into one canal which opens into the acces-
sory sack. Duct of spermatheca long, inserted high on vagina.
(See also pi. 66, fig. 20, E. similaris; figs. 21,22, 23, E. ravida; fig.
24; E.fodiens).
Distribution, middle Europe to China and the East Indies.
Eulota is here used for a considerable number of Oriental snails
having essentially the organization of the European E. fruticum.
The penis lacks flagellum ; the dart sack generally bears an accessory
empty sack into which the many- or few-lobed mucus gland empties ;
and the dart is round in section or but little flattened, the shell
being rather globose with conoidal, though low, spire. Eulotella
Mouss., a sectional name used by von Martens for E. similaris, offers
no distinctive characters of much value, except the obsolesence of
the accessory sack on the dart sack. Acusta differs only in the
thinner shell with simple lip, the mucus glands being either as in E.
similaris (tourannensis) or being more closely bound together into
one compact mass which envelops accessory sack and part of the
dart sack (E. ravida, pi. 66, fig. 21 ; also fig. 22, reverse view of d. s.
with mucus gland, and fig. 23, mucus gland turned back from d. s.,
showing its insertion on accessory sack). The jaw of Acusta (pi. 65,
fig. 2, ravida} has 8 strong close ribs. The radula (pi. 65, fig. 1,.
ravida) is not unlike other Eulotas, but the ectocones of marginal
teeth are not split. Should the Oriental species be held sectionally
distinct from the European E. fruticum on account of their more
elongated and muiti-sacculate mucus glands, they may be separated
under the names Acusta and Eulotella; but v. Mollendorff, certainly
a high authority on Asiatic snails, does not think two names required
for them, uniting the three groups in Eulota.
One species of this group, E. similaris, has an unusually wide
geographic range, extending from middle and southern China to
Penang, Java, Celebes, etc., in which regions it is apparently indi-
genous. By the unconscious intervention of commerce it has be-
204
EULOTA.
come colonized in Japan (Yokohama, Nagasaki, etc.) ; Bengal ;
Reunion, Mauritius, Rodriguez, Seychelles ; Sandwich Is. (Kauai) ;
Ascension Island ; Brazil (Rio Janeiro, Bahia, etc.) ; Barbados, etc.
It has been reported also from Cuba, Porto Rico and Jamaica, but
is not now known to exist in those islands. It has been said to be
found wherever the coffee tree has been carried, but this theory
seems to be unsupported. In many cases I have found that it fol-
lows the cultivation of sugar-cane, also of Oriental origin. This
seems to be the case in Barbados, Brazil, Kauai and the Seychelles,
where E. similaris is commonly found on the borders of cane-fields.
E. arundinetorum Hde., iii, 207. E.
E. assimilaris Gredl. E.
E. assimilis Ad., iv, 48. E.
E. bactriana Hutt., iii, 212. E.
E. billeana Hde., iii, 209. E.
E. bitseniata Mlldff., viii, 221. E.
E. bocageana Cr., vi, 112. E.
E. brardiana Pfr., iii, 210. E.
E. burtiuii Dh., iv, 48. E.
E. cavimargo Mart. E.
E. carinifera Semp., viii, 220. E.
E. cestus Bens., iii, 206. E.
E. cinctoinflata Monss., iv, 47. E.
E. coreanica A. & R., iii, 220.
E. cremata Hde., iii, 207.
E. despecta Gray, iii, 211. E.
E. dichroa Pfr., iii, 208. E.
E. dissimilis Semp., viii, 220. E.
E. duplocincta Mart., viii, 216. E.
E. elatior Mts., iii, 210. E.
E. extrusa T.-C., viii, 218. E.
E. fodiens Pfr., iii, 212. E.
E. fortunei Pfr., iii, 208. E.
v. meridionalis Mlldff. E.
E. frilleyi C. & D., iv, 49. E.
E. fruticum Mull, iii, 200. E.
terrestris Gmel.
cinerea Poir.
lucana Vallot.
carduelis Reib.
/. anderssoni Cless. E.
/. mosellica Bgt. E.
/. aubiniana Bgt. E.
/. lemonia Bgt. E.
/. dumorum B^t. E.
v. insularum West. E.
fuchsi Gredl.
graminum Hde., iii, 207.
hsesitans Hde.
hemisphserica Mlldff., viii, 223.
huberiana Hde., iv, 49.
impatiens Hde.
improvisa Hde., iii, 220.
jourdyi Mori., viii, 219.
laeta Gid., iv, 47.
latrunculorum Hde., iii, 221.
leprosa Hde.
leprosula Hde., iii, 220.
maackii Gerst., iii, 209.
v. depressior Pfr.
conrauxiana Hde., iii, 209.
mabilleiCr.,iii,216.
middendorffi Gerst., iv, 111.
mighelsiana Pfr., iii, 212.
miliaria Gredl.
mirmcula Hde.
mola Hde.
nucleus Dh., iii, 207.
? oenostoma Dh., viii, 223.
oncopila Hde., iii, 208.
oscitans Mts., iv, 47.
paricincta Mart., viii, 217.
/. bisbicincta Mart.
/. ambicincta Mart.
/. incincta Mart.
/. bilaticincta Mart,
peguensis Bens., vi, 113.
phseozona Mts., iii, 205.
phragmitium Hde., iv, 48.
physeta Anc., iv, 50.
pilidion Bens., vi, 114.
plicosa Mts. Nachr., '94, 135.
EULOTA. 205
E. radicicola Bens., iii, 210. squalida Ziegl.
E. ravida Bens., iv, 48. addita Fer.
helvaeea Ph. epixantha Pfr.
v. lineolata Mlldff., iv, 48. /. stimpsoni Pfr., iii, 206.
E. ravidula Hde., iv, 49. genulabris Mart.
E. redfieldi Pfr., iv, 49. /. arcasiana C. & D., iii, 2U
E. rubens Mts., iii, 205. /. borbonica Dh., iii, 206.
/. concolor Mts. /. hongkongensis Dh., iii, 206.
/. finschiana Mts. /. obscura Dh., iii, 206.
/. zeiliana Mts. E. suffodiens Bttg., viii, 219.
/. regeliana Mts. E. straminea Hde., iii, 207.
E. ruppelli Dh., iii, 210. E. striatissima Dh., iii, 207.
E. scalpturita Bens., iii, 21 1 . E. tenimberica Mlldff., viii, 220.
E. schadenbergi Mlldff., viii, 223. E. tourannensis Soul., iii, 209.
E. selskii Gerst., iv, 47. globula Lea.
E. semenovi Mart. clusilis Val.
E. serotina Ad., vi, 106. /. rhodostoma Mlldff.
E. sieboldtiana Pfr., iv, 47. E. transversalis Mss., iii, 210.
E. sirailaris Fer., iii, 205. E. vagoina Gredl., iv, 257.
translucens King. E. weyrichi Schr., iii, 209.
woodiana Lea. E. zoroaster Theob., iii, 211.
Section Armandia Aucey, 1883.
Armandia ANC., II. Nat. Sicil., ii, p. 143, type H. davidi Dh.
(March, 1883).
Shell rather small, quite thin, depressed-convex, the spire low-
conoidal, of few (about 4) rapidly widening whorls; apex obtuse.
Aperture very oblique ; peristome a trifle expanded, acute, much
dilated at the eolumellar insertion, partly closing the narrow um-
bilicus.
Anatomy unknown. Type H. davidi Desh. (See pi. 64, figs. 4, 5,
6, E. calymma Schm. & Bttg.).
Distribution : Thibet ; interior China. The affinities of this group
can only be guessed until the soft parts are made known.
E. davidi Dh., ii, 103. E. plicatilis Dh., ii, 103.
sinica Mts. E. sarelii Mts., iv, 49.
E. moupiniana Dh., ii, 103. nora H. Ad.
E. calyrama Schm. & Bttg.
Section Cathaica v. Mollendorff, 1884.
Cathaiea MLLDFF., Jahrb. D. M. Ges., 1884, p. 339, type H.
pyrrhozona Ph. PILSBRY, Man. Conch., viii, p. 204. Not Cathai-
cus Bates, 1870 (coleoptera).
206 EULOTA.
Shell umbilicated, depressed, sculptured with striae or riblets ;
whorls 5-7, slightly convex, the last usually somewhat angular at
periphery. Aperture oblique; peristome thickened within, the
upper margin unexpanded, outer and basal margins expanded,
columella dilated. Type E. fasciola Dr. (=pyrrhozona Phil.,) pi.
55, figs. 6, 7.
Animal with the tail rounded above, no longitudinal groove. Sole
indistinctly tripartite.
Jaw arcuate, with 3 to 7 weak ribs (pi. 65, fig. 8, pyrrhozona ;
pi. 65, fig. 15, przewalskii).
Radula with blunt mesocones on median and lateral teeth ; mar-
ginals with the ectocone simple or bifid (pi. 65, fig. 7 , pyrrhozona ;
pi. 65, figs. 16, 17, middle, and 1st, 2d, 10th, 14th, 18th, 23d, and
3 marginal teeth of przewalslcii).
Genitalia: penis slender, ending in a long retractor and the ter-
minal vas deferens. Dart sack large, opening into atrium, one
dense cluster of about 10 club-shaped, glandular mucus glands in-
serted near its base. Spermatheca duct long (pi. 66, fig. 32, E.
pyrrhozona).
Distribution, north and middle China and Central Asia.
E. brevispira H. Ad., viii, 208. E. pandynama Mab., viii, 194.
E. buddse Hilb., viii, 208. E. pekinensis Dh., viii, 205.
E. buvigneri Dh., viii, 212. tchiliensis Mlldff.
richthofeni Mts. /. conoidea Dh.
E. confucii Hilb., viii, 213. E. przewalskii Mts., viii, 209.
E. constantise H. Ad., viii, 206. mencii Hilb.
" constantince," viii, 206. E. pulveratricula Mts., viii, 211.
E. desgodinsi Bgt., viii, 194. "pulverella " on pi.
E. giraudeliana Hde., viii, 210. loczyi Hilb.
E. grseseri Mouss., viii, 205. E. pulveratrix Mts., viii, 211.
E. gredleri Hilb., viii, 209. E. fasciola Drap. iii, 208.
stoliczkana Hilb., olim. pyrrhozona Ph., viii, 204.
E. heudei Hilb., viii, 210. faeiola Dr., iii, 208.
E. inopinata Dh., viii, 207. E. schensiensis Hilb., viii, 211.
E. kreitneri Hilb., viii, 211. E. sempriniana Hde., viii, 207.
E. lutuosa Dh., viii, 212. E. siningfuensis Hilb., viii, 211.
" lutosa " Try., iii, 208. E. stoliczkana Nev., iii, 250.
E. magnaciana Hde., viii, 207. E. subrugosa Dh., viii, 211.
E. mongolica MlldfF., viii, 206. v. kalganensis Mlldff.
E. orythia Mts., viii, 210. E. thibetica Dh., viii, 208.
tibetica Mlldff.
EULOTA. 207
Sinistral species.
E. christinse H. Ad., viii, 213. E. filippina Hde., viii, 214.
subchristince Anc. E. dejeana Hde., viii, 215.
v. subsimilis Dh. E. anceyi Mlldff., viii, 215.
? carinifera Ad.
Section Pseudiberus Ancey, 1887.
Pseudiberus ANC., Conchologist's Exchange, i, p. 76 (June, 1887),
types H. tectumsinense , zenonis, etc.
Shell depressed-trochoidal, keeled, narrowly umbilicated, rudely
striated ; heavy, cretaceous and whitish ; whorls about 5, the last
deflexed. Aperture rhombic, oblique, the lip straight above, deeply
arched, expanded and much thickened within, below. Type E.
tectumsinense Mts., pi. 55, figs. 8, 9.
The anatomy of these snails is unknown, but they are probably a
keeled and chalky manifestation of the Cathaica type. They in-
habit interior China and central Asia.
E. tectumsinense Mts., iv, 59. E. plectotropis Mts., iv, 58.
E. zenonis Gredl., iv, 59. E. mataianensis Nev., iv, 59.
Section Platypetasus Pilsbry, 1894.
Shell lens-shaped, acutely keeled, thin, umbilicated ; whorls 4,
the last descending in front. Surface smoothish. Aperture sub-
horizontal, oval ; peristome expanded, reflexed below, the ends ap-
proaching and connected across the parietal wall. Type E. inno-
minaia Hde.
E. mariella H. Ad., viii, 196. E. innominata Hde., viii, 197.
v. submariella Pils. ? aquila H. Ad.
Section Thysanota Albers, 1860.
Thysanota ALB., Die Hel., 1860, p. 63, type H. guerini Pfr.
Shell thin, corneous, narrowly umbilicated, trochiform; whorls
numerous (7 to 8), narrow, with a fringe of hairs at the keeled peri-
phery, extending up the suture. Base flattened ; aperture angulate-
lunar, the lip thin, simple, the columellar margin hardly expanded.
Type E. guerini Pfr.
Distribution, Nilgiri and Anamullay Hills, southern India.
208 EULOTA.
The anatomy is unknown. The group has usually been placed
near Trochomorpha, but I am disposed to consider it near Plectotro-
pis, partly on account of its hairy keel, partly because Blanford
indicates guerini as a species lacking tail-pore. On the other hand,
the simplicity of the lip favors the other view.
E. guerini Pfr., iii, 93. E. tabida Pfr., iii, 94.
crinigera Bens., iii, 94.
Section Plectotropis Martens, 1860.
Plectotropis v. MART., Die HeL, p. 121, type If. elegantissima
ffr. Thea ALB., Die HeL, 1850, p. 118, not Thea Mulsant, 1846.
Shell depressed and carinated, widely umbilicated, dull and
brown, with more or less shaggy cuticle and usually a peripheral
fringe of hairs; whorls numerous (5 to 8), narrow and slowly in-
creasing. Aperture small, angulate-lunar, oblique; lip narrowly
expanded, reflexed below. Type elegantissima, pi. 64, figs. 18, 19.
(See also pi. 64, figs. 16, 17, E. mackensii, typical form from Okin-
awa I., Liukiu group).
Jaw high-arched, with many (10-19) ribs, more or less denticu-
lating the basal margin (PI. 65, fig. 13, E. vulvivaga).
Radula (pi. 65, fig. 14, E. vulvivaga) having the middle tooth
without side cusps, but with a lateral bulging, middle cusps about
the length of basal-plate ; laterals with a small ectocone. Margin-
als with the long inner cusp bifid, ectocone split into two. The
dentition of sumatrana and vulvivaga is practically the same.
Genitalia (pi. 66, figs. 33, 34, E. vulvivaga) : penis rather long,
epiphallus short, strongly bent at the attachment of retractor, con-
tinued in a rather short, blunt flagellum. Dart sack large, contain-
ing a long, slightly curved dart, lens-shaped in section (fig. 34).
Mucus gland inserted high on dart sack, divided into two glandular
branches which are wide, flattened and rather incoherent, the dart
sack and glands bound loosely to vagina. Duct of spermatheca very
long and slender, without diverticulum, bound to oviduct.
Distribution : Japan, China and adjacent islands, south to Su-
matra.
The anatomy of this group is known by Wiegmann's work on
sumatrana and rotatoria, and by my own dissections. E. rotatoria
has much the same form of genitalia as I have found in E. vulvivaga,
EULOTA.
209
except for the dart sack and its appendages, which are absent in
Wiegmann's figure. His specimen was a young one, and the organs
may have been undeveloped ; but I do not think this so likely as that
the species is really no Pledotropis, but a Ganesella. Until adult
examples are examined, I do not venture to transfer the species,
especially since a vast majority of the forms of both groups are still
anatomically unknown, and their systematic position consequently
is only arbitrarily fixed by slight and obscure shell features.
E. mackensii Ad. & Kv., iv, 52. E.
v. rnystagoga Mab., viii, 193. E.
v. vulvivagaSchm.<fe Bttg.,viii, E.
193. E.
E. gerlachi Moll., iv, 52. E.
v. granulosostriata Mts, E.
v. abrupta Mts.
v. hunancola Gredl. E.
E. laciniosula Hde., iv, 53. E.
laciniosa Hde., not Lwe. E.
E. trichotropis Pfr., iv, 53. E.
v. laciniata Hde., iv, 53. E.
v. shanghaiensis Pfr., iv, 56. E.
E. elegantissima Pfr., iv, 52. E.
pretiosa Alb. E.
E. scepasma Pfr., iv, 58. E.
E. ciliosa Pfr., iv, 55. E.
E. lautsi Schm. & Bttg., viii, 193. E.
v. brachylasia S. & B., viii, 194. E.
E. squarrosa Old., viii, 194.
E. granti Pfr.
E. ningpoensis Bttg., viii, 194. E
E. esau Gredl., viii, 158. E
E. patungana Gredl.. viii, 158. E
E. hupensis Gredl., iv, 54. E
orthocheilis Hde.
E. (?) barbosella Hde., iv, 55. E
E. subconella Mlldff., iv, 258. E
E. loufouana Mlldff., iv, 258.
E. visayana Mlldff. E
winteriana Semp.
14
winteriana Pfr., iv, 54.
intumescens Mts., iv, 54.
luzonica Mlldff., Nachr.,'94,1 14
sterilis Hde.
, demolita Hde.
laciniata Hde.
calculus Hde., not Lwe.
sedentaria Hde.
, parasitarum Hde.
, parasitica Hde.
. perplanata ~N"ev.,iv, 57.
, akoutongensis Theob., iv, 57.
emensus Aust., P. Z. S. '88, 242.
ancylochila Cr., iv, 55.
mitanensis G.-A., viii, 195.
grurnulus G.-A., viii, 195.
, pudica G.-A., viii, 195.
. tin ma Pfr., iv, 53.
. huttoni Pfr., iv, 54.
orbicula Hutt., not Orb.
v. savadiensis Nev.
. clarus Aust, P. Z. S., J 88, 242..
. catostoma Bens., iv, 62.
. oldhami Bens, iv, 61.
tapeina Bens., iv, 53.
v. bhamoensis Nev., iv, 54.
(?) rotatoria Busch., iv, 54.
. sumatrana Mts., iv, 56.
v. moussoniana Mts.
. squamulosa Mss., iv, 56.
210 EULOTA.
Section Aegista Albers, 1860.
Aegista ALB., Die Hel., 1860, p. 121, type H. oculus Pfr.
Shell depressed and broadly umbilicated, solid, striated ; brown,
unicolored or with a light peripheral baud; spire low, composed of
many narrow whorls, the last not keeled, descending in front. Aper-
ture round lunar, oblique, the peristome toothless, narrowly ex-
panded, somewhat thickened within, reflexed at base, ends converg-
ing. Type E. oculus Pfr., pi. 64, figs. 13, 14, 15.
External anatomy and genitalia unknown. Jaw arcuate, with
about 6 wide, low, but separated ribs (pi. 65, fig. 10, E.platyomphala).
Radula showing the same characters described for Plectotropis, but
the outer marginals have the ectocone bifid (pi. 65, fig. 9, E. platy-
omphala).
Shells of this section differ from Plectotropis in lacking the peri-
pheral keel and in the smoother surface, but there are some inter-
mediate species. It has the same geographic range, extending north-
ward to Kiusiu Island, Japan. The jaw of the only species examined
has fewer ribs than in Plectotropis, but this is not likely to prove a
constant difference.
(Species of Japan, Liukiu Islands and Formosa.)
E. kobensis Schm. & Bttg., viii, E. oculus Pfr., iv, 59.
196. typinsana A. & R.
E. friedeliana Mart., iv, 61. E. vermis Rve., iv, 60.
E. circulus Pfr., iv, 61. E. subchinensis Nev., iv, 62.
(Species of China and India.)
^E. chinensis Phil., iv, 60. E. subcinctula Hde.
E. pseudochinensis Moll., iv, 60. E. squamosella Hde.
chinensis Hde., not Phil. E. meusalis Hde.
E. platyomphala Moll., iv, 61. E. thoracica Hde., iii, 221.
E. serpestes Hde. E. secundaria Hde.
E. herpestes Hde., i v, 60. E. mellita Hde.
E. furtiva Hde., iv, 60. E. mellitula Hde.
E. aubryana Hde., iv, 60. E. permellita Hde.
E. accresens Hde., iv, 61. E. rebellis Hde.
E. hupeana Gredl., iv, 259. E. languescens Hde.
.E. phayrei Theob., iv, 55. E. vicinella Hde.
EULOTA. , 211
E. gottschei Moll., iv, 62. E. turbo Pils.
E. alphonsi Dh., iv, 61. turbinella Hde., not Morel.
E. aranesetela Hde., iv, 59. E. talifouensis Hde.
E. accedens Hde. taliensis Hde.
v. initialis Hde., iv, 62. E. puberosula Hde., iv, 56.
E. radulella Hde., iv, 57. pulverulenta H., not Lwe.
Section Coccoglypta Pilsbry, 1894.
Shell depressed conoidal with open umbilicus, solid, opaque and
uniform brown ; surface roughened by irregular oblique growth
wrinkles and an uneven granulation ; whorls about 6, convex, the
last inconspicuously angled at periphery, rather tubular, slightly
deflexed in front. Aperture small, round-lunate, oblique ; lip sim-
ple above, expanded outwardly and below, somewhat dilated at
columellar insertion. Type H. dimidiata Hde. (See pi. 64, figs.
20, 21, 22, 23, Kpinchoniana Hde.).
This Chinese group is unknown anatomically, but in my opinion
its species cannot be included in either Aegista, Plectotropis, Eulota
or Cathaica. The general figure of the shell reminds one of such
large American Pyramidulas as P. solitaria or cooperi; but I have
little doubt that Coccoglypta will prove a member of the Eulota
group, conchologically distinguished by its granular shell with non-
reflexed lip. The peripheral angulation is barely mentionable. It
is probable that other species now placed in Aegista or Ganesella will
prove to belong here.
E. dimidiata Hde. E. pinchoniana Hde.
Section Mastigeulota Pilsbry, 1894.
Shell globose-depressed, rather solid, but like Eulota in form and
sculpture. Jaw arcuate, with about 7 convex ribs. Radula much
as in Eulota. Genital system (pi. 66, fig. 26, E. kiang sinensis) like
Eulota in the dart sack and accessory sack, the dart long and
slightly curved, a little flattened ; mucus gland consisting of numer-
ous sacculated branches bound together and to the d. s. (but shown
torn free in figure), as in Euhadra and some Eulotas. Penis end-
ing in a flagellum, and in E. kiangsinensis it is dilated above into a
hollow, thin-walled bulb.
This section is founded upon E. kiangsinensis, which has the
essential features of Euhadra rather than Eulota, the penis bearing
212 EULOTA.
a flagellum. Probably some other Chinese species will prove to belong-
here, which are now placed in Eulota. The natural limits of these
minor groups cannot be defined until more forms are known ana-
tomically, although much can still be done by careful comparisons
of large series of Oriental shells.
E. kiangsinensis Mts., viii, 216. Middle China.
f unizonalis H. Ad.
v. major Mlldff.
v. cerasina Gredler.
Section Tricheulota Pilsbry, 1894.
Chloritis SEMPER, Reisen p. 234, not of Beck.
Shell rather thin, umbilicated, depressed, all over hairy ; aperture
nearly vertical, lunate, the lip well expanded. Type E. spinosissima
Semper.
Genitalia : Penis club-shaped, passing into a long epiphallus
which ends in a flagellum. Dart sack large, the single long, club-
shaped mucus gland inserted upon it (as in the section Monadenia
of Epiphragmophora*). Spermatheca with short duct, less than half
the length of uterus.
Differs from Mastigeulota and Euhadra in the single club-shaped
mucus gland, and the hairy shell. The presence of a flagellum
separates this group from Eulota. The species are from Mindanao,.
Philippines. Conf. Chloritis f brevidens, etc.
E. sanziana H. & J., vi, 272. E. spinosissima Semp., vi, 273.
lituus Rve, figs. 93a, b. boxalli Sowb.
Section Euhadra Pilsbry, 1890.
Euhadra PILS., Man. Conch. (2) vi, p. 94, 95, 305. Hadra and
Camcena of authors.
Shell dextral or sinistral, rather large and thin, capacious, de-
pressed, with low or conic spire and moderate or closed umbilicus ;
whorls about 6; surface striated and typically decussated by micro-
scopic spiral lines; banded or streaked, rarely unicolored. Aper-
ture lunate, oblique, theperistome expanded throughout, subreflexed
at base. Type E. peliomphala Pfr., pi. 64, figs. 1, 2. (See also pi.
64, fig. 3, amalice, and fig. 9, E. qucesita var. perryi Jay).
EULOTA. 213
Jaw wide, arched, with 8 to 14 wide, contiguous ribs, (pi. 65, fig.
12, E. qucesita var.perryi).
Radula having mesocones only on middle and inner lateral teeth;
outer laterals with ectocone. Inner marginals with bifid oblique
inner and simple outer cusp, the outer teeth with bifid ectocone (pi.
65, fig. 11, E. qucesita var. perryi).
Genital system (pi. 66, figs. 27, 28, 29, E. qucesita var. perryi)
having the penis long ; epiphallus long, bearing the retractor,
flagellum rather short ; dart sack enormous, containing a flattened,
longitudinally costate dart, (figs. 27, 28) ; adnate on d. s. is an ac-
cessory sack, upon which a cluster of sacculated mucus glands is
inserted, these glands being closely bound together and to the dart
sack, as in other forms of Eulota, and with fibres connecting with
uterus. Duct of the globose spermatheca long. (In fig. 29 the
mucus glands are shown torn free from each other and from the
dart sack by the removal of their investing membrane, as has been
done with the other figures on plate 66).
Eukadra was instituted to include a group of rather large species
of Japan and China, which had been placed in Hadra and Camcena
by authors. It contains the largest and most conspicuously colored
belogonous Helices of East Asia, excepting, of course, the Philip-
pine Island Helicostylas. The bands of these forms are not homol-
ogous with those of European Helices, excepting possibly the
supraperipheral one, which was probably present in very ancient
Helices, and is retained in many and diverse groups.
The Japanese forms of Euhadra are so variable that after exam-
ining about a thousand specimens from many localities, I am dis-
posed to consider some forms described by Kobelt and others
merely varieties. Thegenitalia of qucesita, peliomphala and brand-
tii are very similar, but the first of these has more ribs on the
jaw.
(Group of Simodcd).
E. simodse Jay, vi, 95. E. herrmannseni Pfr., vi, 98.
E. connivens Pfr., vi, 96. koreana Pfr.
v. phseogramma Ana
(Group of Luhtiand).
E. luhuaua Sowb., vi, 305. v. callizona Cr., vi, 105.
luchuana Auct. v. amalise Kob., vi, 105.
v. peliomphala Pfr., vi, 99. congener Sm.
japonica Dh., not For. v. congenita Sm., vi, 103.
214 EULOTA-CHLORJEA.
E. luhuana. E. herklotsi Mts., vi, 101.
v. eoa Cr., vi, 98. E. senckenbergiana Kob., vi, 102..
v. sandai Kob. E. mi ran da Ad., vi, 104.
v. subatra Pils. E. lewisii Sm., vi, 106.
v. subnimbosa Kob. E. myomphala Mts., vi, 107,
v. nimbosa Cr., vi, 101. daimio Ad.
v. brandtii Kob., vi, 101. E. qusesita Dh., vi, 108.
v. nipponensis Kob. v. perryi Jay, vi, 108.
v. hickonis Kob. montium Mts.
conica Pils., vi, 305.
{Group of Swinlioei).
E. swinhoei Pfr., vi, 115. E. schmackeri Mlldff., vi, 307..
E. caspari Mlldff., vi, 1.15. E. ammiralis Pfr., vi, 117.
E. pantheia Mab., vi, 116. E. cecillei Phil., vi, 109.
E. granulifera Mlldff., vi, 306. E. moreletiana Hde., vi, 110.
E. renaltiana Hde., vi, 307. E. bairdi H. Ad., vi, 111.
( Group of Succincta).
E. succincta H. Ad., vi 118. E. delavayana Hde.
E. friesiana Mlldff., vi, 118. E. hemiclista Schm. & Bttg.
E. stenozona Mlldff., vi, 118. E. mercatoria Gray, vi, 121.
E. hsematozona Hde., vi, 119. E. mellea Pfr., vi, 97.
E. submandarina Pils., vi, 122. E. purpurascens Pfr., viii, 297,
E. nux Mlldff., vi, 307. E. massiei Mori., viii, 223.
E. cremata Hde. E. philippinensis Semp., vi, 123..
E. seguiniana Hde.
(Group of Batanica).
E. latilabris Mlldff., vi, 109. E. batanica A. & R., vi, 111.
E. yaeyamensis Pils. v. pancala S. & B., viii, 224,
E. formosensis Pfr., vi, 112. sinistrorsa Moll., not Dh.
E. bacca Pfr., vi, 112.
Section Mandarina Pils., 1894.
Shell solid, compact, globose-conic ; axis solid, imperforate.
E. mandarina Gray, vi, 124. Benin Is.
Genus CHLORJEA Albers, 1850.
Chlorcea ALB., Die Hel., p. 113 ; v. MART., edit., p. 169, type H.
sirena. Gruppe der bunt-schaligen Chlorceen SEMPER, Reisen, etc.,
p. 226. PILSBRY, Manual vii, p. 93.
CHLORJEA. 215
Shell imperforafe, varying from depressed-globose to lens shaped,
generally solid ; whorls 4-5, the last descending in front or not; lip
decidedly reflexed, at least below. Entire surface showing under
the lens excessively fine close spiral lines'; lacking hydrophanous
markings. Type C. sirena, pi. 55, figs. 15, 16, 17.
Foot, jaw and radula (pi. 54, fig. 3, benguetensis) as in Helico-
styla.
Genital system (pi. 54, figs. 2, 4, benguetensis) ; penis as in Heli-
costyla, without flagellum (benguetensis, hugeli). or having a very
short one (sirena). Dart sack oval, containing a long lance-shaped
dart (pi. 54, fig. 4, benguetensis}. Mucus gland single, lengthened,,
composed of irregularly grouped large follicles. Duct of sperma-
theca moderately long.
Distribution: Luzon, Marinduque, Mindoro, Cebu, Tablas, Min-
danao, Sibuyan, Luban and Guimaras, Philippines; living upon
trees.
This group is more allied to Eulota than to Helicostyla in the
soft anatomy and dart ; but the species have assumed the arboreal
habits and bright coloring of the latter genus.
Group of C. fibula.
C. fibula Brod., vii, 94. C. amoena Pfr., vii, 98.
C. hanleyi Pfr., vii, 95. C. sirena Beck, vii, 98.
v. hugeli Pfr. v. cebuana Mlldff.
bifasciata Lea. v. guimarasensis Pi Is.
C. benguetensis Semp., vii, 96. C. pelewana Mouss., vii, 99.
C. geotrochus Mlldff., vii, 97. C. gmeliniana Pfr., vii, 100.
C. antonii Semp., vii, 97. C. hennigiana Mlldff., Nachr., '93,
173.
Group of C. dry ope.
C. dryope Brod., vii, 100. C. coerulea Mlldff, vii, 101.
prasina Koch. C. cristatella Mlldff, Nachr., '93,
173.
Group of C. paradoxa.
C. paradoxa Pfr., vii, 102. C. constricta Pfr., vii. 103.
/. immaciilata Pils. restricta Pfr.
C. undina Pfr., vii, 102. ttenopsis Moq.
216 HELICOSTYLA.
Group of C. t her sites.
C. thersites Brod., vii, 104. C. malleata Q. & M.. Nachr., '93,
174.
Genus HELICOSTYLA Ferussac.
Helicostyla FER., Tabl. Syst., p. 46, in part. BECK, Index Moll.,
p. 36, 1837, in part. ALBERS, Die Hel., 1850, p. 104. H. &. A.
ADAMS, Gen. Rec. Moll., ii, p. 191. MARTENS, Die Hel., p. 175,
type H. mirabilis Fer. Cochostyla FER., Tab. Syst., p. 47, 1819.
SEMPER, Reisen in Arch. Phil., Land Moll., p. 164. PILSBRY,
Man. Conch., vii, p. 92. Includes the groups Chlonea, Corasia,
Crystallopsis, Axina, Pfeifferia, Calocochlia, Helicostyla, Orustia,
Cochlodryas, Orthostylus, Helicobulinus, Ptychostylus, Phengus,
Eudoxus, Hypselostyla, Canistrum, Prochilus, Chrysallis, etc.
For anatomy see SEMPER, Reisen im Archip. Phil, and PFEFFER,
Jahrb. Dm. Ges., 1878, p. 195.
Shell varying from depressed, helicoid, to elevated and bulimoid,
imperforate, with solid columella (except in Crystallopsis and Chry-
sallis*) ; surface smooth or roughened, usually covered with a thin,
transparent cuticle, often porous in places when it becomes white and
opaque, producing the " hydrophanous" pattern which ornaments
most species, and which disappears upon wetting the shell. Aper-
ture toothless, but sometimes having a columellar twist or trunca-
tion ; the lip reflexed (but simple in Pfeifferia'). Type H. mirabilis
Fer. (see pi. 53, fig. 1-11).
Foot without pedal margination ; a small left body-lappet often
developed ; kidney elongated. (PL 54, fig. 10, H. festiva ; pi. 54,
fig. 5, H. monticula).
Jaw ribbed (pi. 54, fig. 6, H. butl&n).
Radula with bluntly pointed or truncated mesocones on middle
and lateral teeth, without trace of side cusps. Marginal teeth hav-
ing the entocone indicated by a split in the broad inner cusp, a
small simple ectocone being developed. (See pi. 54, fig. 11, H.
aegle; pi. 54, fig. 12, H. pulcherrima.
Genital system : Penis moderately long, passing into an epiphal-
lus which bears the retractor; flagellum wanting. Dart sack short
and globose, seated on atrium or low on vagina, bearing an accessory
sack into which the mucus gland opens. Mucus gland globular or
HELICOSTYLA. 217
oral, with a very short duct, its thick wall composed of radially
arranged follicles (pi. 54, fig. 7, longitudinal section of mucus
gland of H. butleri). Dart short, straight, and round in section.
Spermatheca oval, on a long, branchless duct (pi. 54, fig. 8, H. but-
leri; pi. 54, fig. 9, H. pit hog aster with its dart).
Distribution: Philippine Is., with a few species in the Moluccas,
in New Guinea and the Solomon Islands. Habits mainly arboreal.
As in most large genera of Helices, the shells of Helicostyla
exhibit a very wide range of forms, some being heavy, dark, de-
pressed and keeled, others globose and thin with brilliant green or
variegated coloring, while still other species are of an elongated
Bulimus-like contour. Peculiar air-permeated cream-white epider-
mal bands and patches are characteristic of many but by no means
all species, and in most the columella is solid.
The jaw is of the usual ribbed type. The teeth are like those of
some Papuinas, but in the lack of side cusps on middle and lateral
teeth they resemble Eulota. The genital system is highly charac-
teristic in the globose form of the mucus gland, which as in other
Belogona euadenia is inserted on the dart sack. In the subgenus
Canistrum (q. v.) a variation in this is found. Our knowledge of
the anatomy is due to Semper, whose work upon the group leaves
little to be desired except the examination of those subgenera
which he did not dissect, Prochilus, Chrysallis, Pap'iistyla, Crystal-
lopsis, etc., and the further investigation of Canistrum find allied
forms.
It is clear that Helicostyla is very near in anatomy to the primi-
tive Belogonous stock, retaining early characters in the simple un-
split and nearly sessile mucus gland and needle-like, bladeless dart.
Its differentiation in shell characters is attributable to long isolation
and the assumption of arboreal habits. The Philippine Chlorseas
were probably derived from a later incursion or an early split,
which has not spread through the entire Philippine group. The
(hmcena, Euhadra and Eulota forms are perhaps to be regarded
as a more recent addition to the fauna.
The subgenus Helicostyla was proposed by Ferussac for a hetero-
geneous assemblage of shells including certain Zonitidce, two Sagdas
and a Gastrodonta, the two species of Stylodonta, Cepolis (Coryda)
alaada and H. mirabilis and coniformis. The Sagda and Gastro-
donta were later removed from the group by Ferussac himself
(Tabl. Syst., p. 67) ; and Beck in 1837 eliminated from it most
218 HELICOSTYLA.
other incongruous elements, leaving only H. alauda and its varieties
and H. mirabilis (galactites), and adding smaragdus and roissyana.
Albers in 1850 restricted Helicostyla to Philippine Island snails of
the mirabilis type, erecting for the West Indian H. alauda the new
group Coryda. H. & A. Adams make Helicostyla a genus to include
Calocochlea, Corasia, Axina, Chlorcea, etc., and place the elongated
forms in genus (Cochlostyla under Buliminse.
Coehlostyla was instituted by Ferussac on the page of the
Tableaux after Helicostyla, and contained species belonging to the
groups Helicostyla, Helicobulinus, Orthostylus, Cylindrus, Caryo-
des, Orphnus, Dryptus, Orthalicus, etc. The name dropped out of
nomenclature entirely until 1847, when Gray names it under
Orthostylus, giving metaformis as its type. Later, the Adams
brothers revived it as a genus for elongated Philippine Island forms ;
and in 1860 von Martens uses it in much the same sense. In
enlarging the genus to include both depressed and elongated species,
Semper unfortunately retains the name Cochlostyla for the entire
series, a course which has been followed by subsequent writers.
In conclusion it is evident that for this genus we must use the
name Helicostyla, which not only has prior position in Ferussac's
work, but was restricted in 1837 by Beck and properly limited by
Albers ; while Cochlostyla was later in the original publication, and
remained a heterogeneous mass of Bulimi and Helicostylse until
Gray in 1847 selected metaformis as its type.
Subdivisions.
AVith the exception of Canistrum, the sections of this genus are
practically identical in anatomy; and rest upon such shell charac-
ters as contour, thickness and sculpture. The presence of interme-
diate species renders their arrangement somewhat artificial.
Depressed or globose, thin, without hydrophanous cuticle.
Globose, white, lip simple and sharp, Pfeijferia.
Lip expanded or reflexed, Corasia, Crystallopsis.
Globose, few whorled, with hydrophanous bands, Leytia.
Globose or depressed, solid, lip expanded or reflexed.
No hydrophanous cuticle, lip narrow ; highly colored, Chromato-
sphcera.
Hydrophanous cuticle present, lip wider, Calocochle<t.
Dark, depressed and much roughened forms, Trachystyla.
Dark and smoothish forms, Anixu.
HELICOSTYLA. 219
Globose or elevated-oval, moderate sized or small.
Heavy and thick, without hydrophanous cuticle, Pachysphcera.
Oblong, not especially heavy, color in browns, Helicostyla.
Oblong, suture white-bordered, color vivid, Cochlodryas.
Subconic, with hydrophanous cuticle, Orustia.
Elevated and conical or turbinate, mostly large.
Whorls numerous, equal ; columella very short with a strong fold,
Columplica.
Whorls more rapidly increasing ; columella longer, Orthostylus,
Helieobulinus.
Elongated and bulimoid.
Imperforate, Hypselostyla, Papustyla, Eudoxus, Phengus, Canis-
trum.
Perforated, Procliilus, Chrysallis.
Section Corasia Albers, 1850.
Corasia ALB., Die Hel., p. Ill; second edit., p. 170, type H.
virgo.
Shell imperforate, depressed-globose or flattened and keeled, thin,.
subdiaphanous ; whorls 3j-5, rapidly widening, the last hardly
descending in front ; lip slightly expanded or narrowly reflexed.
No hydrophanous markings. Type H. virgo Brod., pi. 55, fig. 12.
Anatomy as in other Helicostylas. The shell differs from Calo-
cochlia in being thinner without " hydrophanous" decoration. DiV
tribution, Philippine Is., except Palawan. Arboreal.
Group of H. regince.
H. reginse Brod., vii, 116. H. papyracea Brod., vii, 117.
smaragdina Grat. f acutangula Burrow,
v. almteMlldff.,vii,117. H. psittacina Dh., vii, 118.
v. elizabethse Semp., vii, 117.
Group of H. virgo.
H. virgo Brod., vii, 119. H. segrota Rve., vii, 124.
H. dealbata Brod., vii, 119. H. samboanga H. & J., vii, 124..
broderipi Rve. zamboangce Mts.
H. patricia Pfr., vi, 196. v. intaminata Gld.
H. casta Pfr., vii, 120. H. magtanensis Semp., vii, 125.
220 HELICOSTYLA.
H. puella Brod.. vii, 120. H. intorta Sowb., vii, 125.
v. lais Pfr., vii, 121. v. crassa Mlldff, vii, 125.
v. subpuella Pils., vii, 121. v. siquijorica Mlldff., vii, 125.
H. irosinensis Hid., vii, 121. H. limansauensis S., vii, 126.
H. seruginosa Pfr., vii, 122. H. sphserion Sowb., vii, 154.
H. filaris Val, vii, 122. /. intincta Shutt., vii, 154.
v. nympha Pfr. v. nana Semp., vii, 155.
v. tennis Mlldff., vii, 126. v. crassilabris Mlldff.
v. expansilabris Mlldff., vii, v. meridionalis Mlldff.
126. H. saranganica Mlldff., viii, 245.
H. eydouxi Hid., vii, 123. H. globulosa Mlldff., Nachr., '95,
valenciennesii Pfr., not Eyd. 96.
H. broderipi Pfr., vii, 123. H. loheri Mlldff, Nachr., '94, 115.
Section Crystallopsis Ancey, 1887.
Crystallopsis ANC., Conchol. Exch., ii, p. 23, types H. hunteri and
allasteri.
Shell thin, depressed globose or depressed and keeled, translucent
whitish or banded, the axis perforated, at least in the typical forms.
Surface spirally striated ; lip expanded. Type H. hunteri Cox.
See pi. 55, figs. 10, 11, H. tenimberiea Mlldff.
Jaw (of H. conformis) semicircular, perfectly smooth. Radula
as in Helicostyla.
Genital system (Frontispiece, fig. 4, H. conformis) as in Helicostyla.
The shell is similar in general features to Corasia, but the jaw of
H. conformis has been shown by Tapparone-Canefri to be smooth
(Ann. Mus. Civ. Genov., xix, pi. 8, f. 1, 8, 15). Distribution, Solo-
mon Is. to Moluccas.
Group of H. lactiflua.
H. huuteri Cox, vii, 105. H. wisemani Braz., vii, 109.
H. allasteri Cox, vii, 106. H. aphrodite Pfr., vii, 109.
allisteri Pils., typ. err. H. anadyomene Ad. & Ang., vii,
H. aggei Heimb., viii, 244. 110.
H. subvitrea Pfr., vii, 107. H. psyche Aug., vii, 110.
H. cymodoce Cr., vii, 107. H. balcombei Cox, vii, 111.
H. lactiflua Pfr., vii, 108. H. woodfordi Sowb., viii, 243.
isabellensis Souv. H. tricolor Pfr., vii, 111.
H. purchasi Pfr., vii, 108. v. picta Sin., vii, 112.
H. rossiteri Ang., vii, 109. v. transenna Pils., vii, 112.
H. tenimberiea Mlldff, viii, 244.
HELIOOSTYLA. 221
Group of H. extensa.
H. conformis Fer., vii, 113. H. najas Pfr.,vii, 115.
H. leucophthalma Pfr., vii, 113. H. physalis Pfr., vii, 115.
H. extensa Mull., vii, 114. H. obliquata Dh., vii, 116. _
H. coelaxis Pils., vii, 114.
Section Pfei/eria Gray, 1853.
Pfeiffetia GRAY, P. Z. S., 1853, p. 110. type H. micans Pfr.
Shell globose, iiuperforate, the slender axis solid ; thin, brittle?
uniform white and glossy ; whorls 4, the last not descending in front.
Aperture lunar; lip thin, acute and fragile ; columella slightly
thickened, vertical, deeply inserted in the base. Animal as in Heli-
costyla generally, except that the mantle is reflexed over the acute
HP.
H. micans Pfr., vii, 128. Northern Luzon.
Section Leytia Pilsbry, 1892.
Leytia PILS., Man. Conch., vii, p. 129 (Jan. 30, 1892).
Shell imperforate, globose, thin ; whorls few (4), the last angu-
lated at periphery ; surface spirally striate, having hydrophanou&
cream white bands. Aperture very large; lip simple, a little ex-
panded below ; columella simple, thin, vertical, deeply inserted.
Anatomy unknown.
H. fragilis Sowb., vii, 129. Island of Leyte, Philippines.
leytensis Pfr.
Section Chromatosphcera Pilsbry, 1892.
Chromatosphcera PILS., Man. Conch., vii, p. 169 (Jan. 30, 1892).
Shell imperforate, depressed globose, solid and opaque, richly
colored but lacking hydrophanous markings; surface lacking spiral
sculpture; last whorl scarcely descending in front; lip blunt, very
narrowly reflexed throughout; columella subvertic'd], deeply inserted^
the umbilical area covered by a concave white callus. Anatomy
typical. Type H. aurata Sowb.
Distribution, Northern Luzon.
H. aurata Sowb., vii, 170. H. lividocincta Semp., vii, 171.
H. erubescens Semp., vii, 170. H. pudibunda Semp., vii, 171.
v. luteocincta Semp., vii, 171.
222 HELICOSTYLAv
Section Calocochlea Hartmann, 1842.
Calocochlea HARTM., Erd- und Siisswasser Gasterop. Schw., p.
163, type pulcherrima Sovvb. Callicochlias AGASSIZ, 1847, and of
authors.
Shell imperforate, solid, subglobose, generally with varied pattern
and bydrophanous cuticle. Soft anatomy typical. Type H. pul-
cherrima Sowb., pi. 55, fig. 13.
Group of H. cromyodes.
H. cromyodes Pfr., vii, 130. H. obtusa Pfr., vii, 132.
valencienniilZyd. H. amicta Rve., vii, 133.
H. denticulata Jay, vii, 131. H. decora A. & R., vii, 133.
H. albaiensis Sowb., vii, 132. H. ? semirufa Alb., viii, 245.
H. tukanensis Pfr., vii, 132.
Group of H. pulcherrima.
H. pulcherrima Sowb., vii, 133. H. lalloensis Pfr., vii, 136.
H. festiva Don., vii, 134. H. angusta Alb., vii, 136.
luzonica Sowb. H. princeps Rve, vii, 137.
annce O. Semp. H. erythrospira Mlldff., vii, 137.
H. dubiosa Pfr., vii, 135. H. generalis Pfr., vii, 137.
speciosa Jay. H. chrysochila Sowb., vii, 138.
batanica Rve. chrysocheila Sowb.
volubilis Rve.
Group of If. polillensis.
H. polillensis Pfr., vii, 138. H. andromache Pfr., vii, 139.
/. portei Pfr. H. codonensis Hid., vii, 140.
/. ajax Pfr. H. decipiens Sowb., vii, 140.
/. hector Pfr.
/.peraffinis Pils., vii, 139.
Group of H. zonifera.
H. zonifera Sowb., vii, 141. H. cailliaudi Dh., vii, 144.
samarensis var. Semp. ferruginea Lea.
v. circe Pfr., vii, 142. H. microspira Pfr., vii, 145.
purpurascens Mts. H. hemisphserion Pfr., vii, 145.
v. globosa Mlldff. H. samarensis Semp., vii, 146.
v. paraleuca Pils., vii, 142. H. coccomelas Sowb., vii, 146.
H. coronadoi Hid., vii, 142. speciosa Pfr., not Jay.
v. pulchra Pils., vii, 143. H. ponderosa Pfr., vii, 147.
HELICOSTYLA. 223
H. norrisii Sowb., vii, 143. H. luengoi Hid., viii, 245.
H. damahoyi Pfr., vii, 143. H. hidalgoi Mlldff., viii, 246.
samarensis Hid., not Semp.
H. chrysseme Q,.& M. Nachr.,'93, H. streptostoma Mlldff. Nacbr.,^3
175. 176.
H. semperi Mlldff. Nachr., '93, H. trisculpta Mlldff. Nachr., '95,
175. 97.
Group of H. mindanaensis.
H. mindanaensis Sby., vii, 148. H. barfordii Sby., vii, 148.
mindana Rve.
mindorana Hartm.
Group of H. melanochila.
H. pan Brod., vii, 149. H. roissyana Fer., vii, 151.
H. depressa Semp., vii, 149. solida Pfr., vii, 153.
lignaria Pfr., preoc. rossiana Gray.
v. perdepressa Pils. /. lutea Pils.
H. melanochila Val., vii, 150. /. subatra Pils.
brunnea Sowb. H. infuscata Alb., vii, 152.
H. cblorochroa Sowb., vii, 150. H. xanthobasis Pils., vii, 155.
H. matruelis Sowb., vii, 151. H. dimera Jonas, vii, 156.
H. difficilis Pfr., vii, 151. H. crossei Hid., vii, 156.
H. mainitensis Hid., vii, 152. H. retusa Pfr., vii, 156.
H. lignicolor Mlldff., vii, 153. H. libata Rve., vii, 157.
Section Anixa Pilsbry, 1894.
Axina ALB., Die Hel., p. 113, 1850; v. Mart., 2d edit., p. 71,
type H. zebuensis. Not Axina Kirby, 1817 (coleoptera).
Shell imperforate, solid, dark colored ; varying from depressed
and keeled to globose turbinate ; more or less covered with hydroph-
anous cuticle ; smoothish, lacking spiral sculpture; lip narrowly or
broadly reflexed. Soft anatomy as in Helicostyla generally. Type
H. zebuensis Brod. (See pi. 55, fig. 14, H. moreletiana Pfr.).
These are ground snails, mainly confined to Luzon, Cebu and
Siquijor.
H. garibaldiana D.& S.,vii, 159. H. kobelti Mlldff., vii, 160.
H. siquijorensis Brod., vii, 159. H. zebuensis Brod., vii, 161.
v- pallens Mlldff. v. striatissima Pils., vii, 162.
224 HELICOSTYLA.
H. schadenbergi Mlldff., vii, 160. v. gloynei Sowb., vii, 165.
H. pfeifferi Semp., vii, 162. ecarinata MlldfF.
cumingi Pfr., preoc. H. moreleti Pfr., vii, 165.
H. phloiodes Pfr., vii, 163. H. montfortiana Pfr., vii, 165.
H. carbonaria Sowb., vii, 163. H. bruguieriana Pfr., vii, 166.
/. rubeus Mlldff. H. beloni Jouss., Le Nat., '94, 186-
H. magistra Pfr., vii, 164.
Section Trachysiyla Pilsbry, 1892.
Trachystyla PILS., Man. Conch., vii, 166.
Shell solid, depressed, dark colored, with a dull ashen hydrophan-
ous cuticle; spire low; whorls few and rapidly widening ; columella
oblique, straight; surface dull and wrinkle malleuted. Type H.
cryptica.
Species of this group live on the ground under leaves, etc. ; and
while the shell differs widely from arboreal Helicostylas, the soft
anatomy is the same. The few species inhabit eastern Mindanao,
Samar, Bohol, Leyte and Luzon.
H. cryptica Brod., vii, 167. v. panayensis Semp., vii, 168.
v. latitans Brod., vii, 167. v. tumida Mlldff.
fdionacea Dh. v. subglobosa Mlldff.
v. depressa Mlldff, 1893. v. nigricans Mlldff.
v. cretata Brod., vii, 168. v. cineracea Semp., vii, 168.
v. minor Mlldff. H. dataensis Semp., vii, 169.
Section Helicostyla Fer., (restricted).
After the removal of Cochlodryas, Pachysphcera, etc., a consider-
able number of oblong shells grouping around H. mirabilis, meta-
formis and fenestrata remain to represent this section, the type of
which is H. mirabilis Fe"r., pi. 53, fig. 7.
Group of H. mirabilis,
H. mirabilis Fer., vii, 181. f. trichroa Pils.,vii, 182.
formosa Wood. v. fulgens Sowb., vii, 182.
galaetites Lam. H. tephrodes Pfr., vii, 183.
persimilis Dh. H. ? plurizonata A. & R., vii, 183.
Group of H. coUodes.
H. collodes Sowb., vii, 184. H. thomsoni Pfr., vii, 185.
H. indusiata Pfr., vii, 184. H. suprabadia Semp., viii, 246.
HELICOSTYLA. 225
Group of H. metaformis.
H. metaformis Fer., vii, 186. H. fuliginata Mts., vii, 188.
ovularis Mke. fumigata Semp.
H. rustica Mouss., vii, 187. v. nigrolabiata Mlldff.
H. hydrophana Sowb., vii, 187. H. lacera Pfr., vii, 189.
H. butleri Pfr., vii, 188. H. rehbeini Pfr., vii, 190.
H. languida Pfr., vii, 189. H. roebeleni Mlldff., Nachr., '95,
H. unica Pfr., vii, 189. 98.
Group of H. fenestrata.
H. m on tan a Semp., vii, 191. H. curta Sowb., vii, 192.
H. fenestrata Sowb., vii, 192.. v. dilatata Pfr., vii, 193.
Section Cochlodryas Martens, 1860.
Coehlodryas MTS., in Die Hel., p. 176, type H. polychroa. Pce-
eilus ALB., mss.
Elevated, vividly colored species lacking hydrophanous cuticle,
and with a distinct subsutural band which is generally white. Type
H. polychroa, (=viridostriata) pi. 53, fig. 10.
H. florida Sowb., vii, 177. H. orbitula Sowb., vii, 179.
helicoides Pfr. chlorogrmnmica Val.
H. viridostriata Lea, vii, 178. H. tenera Sowb., vii, 179.
f smaragdm Beck. H. ignobilis Sowb., vii, 180.
polychroa Sowb. H. boettgeriana Mlldff., vii, 18L
Section Orustia Morch, 1852.
Orustia MORCH, Cat. Yoldi, p. 15, in part, first species If. mon-
ticula.
Shell imperforate, turbinate-globose, not spirally striated, banded,
with hydrophanous cuticle; lip thin, narrowly reflexed ; columella,
vertical, deeply entering. Anatomy typical. Type H. monticula,
pi. 54, fig. 5 (see also pi. 53, fig. 1, H. versicolor*).
Distribution, Luzon.
H. monticula Sowb., vii, 176. H. versicolor Mlldff., viii, 246.
H. pulchella Mlldff., Nachr. H. strigata Mlldftl Nachr. '95,
['93, 176. [97.
15
226 HELICOSTYLA.
Section Pachysphcera Pilsbry, 1894.
Pachysphcera PILS., Man. Conch, vii, p. 172 (Jan. 30, 1892).
Shell small, globular or globose-elevated, solid, brightly colored ;
lacking hydrophanoas cuticle and spiral striae. Type H. sphcerica
Sowb.
Confined to north-western Luzon.
H. sphaerica Sowb., vii, 172. H. iloconensis Sowb., vii, 175.
H. balteata Sowb., vii, 173. v. heterotseniata Pils.
H. annulata Sowb., vii, 174. v. xanthotseniata Pils.
v. intensior Pils.
Section Columplica Hartmann, 1842.
Ptychostylus MLLDFF., Nachrichtsbl. D. M. Ges. 1888, p. 74.
Not Ptychostylis Gabb, a group of Trochidce. Not Ptychostylus Sand-
berger, Land- u. Susswasser-Conch. der Vorwelt, p. 58 (Melaniidce)
Stylodonta\(in part) of authors. Columplica (in part) HARTMANN,
Gasterop. Schweiz, p. 187, 188 (If. uniplicata and H. dolium=ce-
poides). Hypoptychus PILSBRY, Proc. Acad. Nat. Sci. Phila., 1892
p. 395, footnote.
Shell globose-turbinate, formed of many closely coiled whorls, the
surface having hydrophanous cuticle ; aperture narrowly lunar ;
columella spirally twisted, having a strong tooth-like fold at the base.
Animal externally like Cochlostyla, internal anatomy unknown.
A peculiar and isolated type resembling Stylodonta unidentata
of the Seychelles Islands. It was first classed in Cochlostyla by
Semper. The name Ptychostylus being preoccupied, the writer pro-
posed to substitute Hypoptychus; but it seems best to revert to
Hartmann's Columplica.
H. cepoides Lea, vii, 194. Island of Luban.
dolium Hartm.
Section Helicobiilinus Broderip, 1840.
Helicobulinus BROD., P. Z. S. 1840, p. 123, type H. sarcinosa.
Helicobulimus MOLLENDORFF, Landschn. Cebu, p. 241. Chromo-
cochlea HARTMANN, Gast. Schweiz, p. 137, 1844, type C. turbineides.
Chromatocochlias AGASSIZ.
Shell capacious, turbinate-globose, solid, variegated with green or
brown, covered with a variously patterned hydrophanous cuticle.
HELICOSTYLA. 227
Columella more or less folded. Type H. sarcinosa. (See pi. 53, fig.
2, H. turbinoides).
This section, while closely allied to Orthostylus, presents affinities
with so many groups that it must be regarded as an intermediate or
synthetic type.
H. grandis Pfr., vii, 195. H. turbinoides Brod., vii, 196.
colossea Pfr. H. cinerascens Pfr., vii, 197.
carolus Dh. v. turbo Pfr., vii, 197.
H. sarcinosa Fer., vii, 195. H. bembicodes Pfr., vii, 198.
v. turgens Dh., vii, 196.
Section Orthostylus Beck, 1837.
Orthostylus BECK (in part), Index, p. 49. MARTENS in Alb.
Die Hel. p. 177. v. MLLDFF., Landschu. Cebu, p. 242. Pithohelix
SWAINS., Malacol., p. 166. Pythohelix SWAINS., 1. c., p. 332.
The shell is generally large, solid, ovate-conic, covered with a
variously patterned hydrophanous cuticle. Aperture oblique, ovate
lip reflexed ; columella nearly vertical, more or less obviously folded
below. Anatomy typical. Type H. pithogaster, pi. 53, fig. 3.
The present section is allied to Helieobulinus, which consists of
more inflated shells, and to Hypselostyla, containing more elongated
forms. The limits of Orthostylus are not easy to determine, as there
are species almost or entirely intermediate between this group and
Helicostyla, as well as forms connecting with Hypselostyla. With
the exception of Mindoro and Mindanao, it occurs on all the Philip-
pine group.
H. bicolorata Lea, vii, 199. H. ticaonica Brod., vii, 203.
alberti Brod. B. subglobosus Lea.
v. onyx Brod., vii, 199. /. lutea Pils.
H. imperator Pfr., vii, 199. H. lignaria Pfr., vii, 204.
H. pithogastra Fer., vii, 200. H. gilva Brod., vii, 205.
/. philippinensis Pfr., vii, 201. H. woodiana Lea, vii, 206.
? strigata Mlldff. reevii Brod.
/. bipartita Pils. vii, 201. reevei Pfr.
H. villari Hid., vii, 201. H. portei Pfr., vii, 207.
/. ventricosa Mlldff. portii Pfr. olim.
H. daphnis Brod., vii, 201. H. rufogastra Less, vii, 207.
H. cunctator Rv., vii, 202. B. monozonus Pfr.
H. faunus Brod., vii, 203. H. macrostoma Pfr., vii, 208.
228 HELICOSTYLA.
H. vidali Hid., vii, 208. H. nux Serap., viii, 5.
H. juglaDS Pfr., vii, 208. H. marinduquensis Hid., vii, 209.
/. olivacea Mlldff. woodianus Pfr. not Lea.
/. roseolimbata Mlldff. H. amalise Mlldff., viii, 245.
Elongated Orthostylus.
H. mus Brod., viii, 6. H. ventricosa Chera., viii, 10.
H. leucophsea Sowb., viii, 6. nobilis Rve.
H. glaucophthalma Pfr., viii, 7. /. guimarasensis Rve.
H. senckendorffiana Pfr., viii, 7. /. decorata Fer.
H. solida Pfr., viii, 8. /. frater Fer.
H. pictor Brod., vir, 8. H. satyrus Brod., viii, 13.
B. labropurpureus Grat. v. palawanensis Pfr.
H. solivaga Rve., viii, 9. v. cyanocephala Pils.
H. leopardus Pfr., viii, 9. v. cinerosa Pfr., viii, 15.
H. aplomorpha Jonas, viii, 11. v. graellsi Hid., viii, 14.
H. fulgetrum Brod., viii, 12. v. librosa Pfr., viii, 12.
? Bal. antipodararn Gray. H. csesar Pfr., viii, 15.
H. fischeri Hid., viii, 29.
Section Hypselostyla Martens, 1868.
Hypselostyla Mts., in PFR., Monogr. Hel. Viv. vi, p. 7, 16, for
nympha and dactylus.
Shell imperforate, narrow and elongated, usually rather thin and
somewhat keeled at periphery ; peristome narrowly expanded. (See
pi. 53, fig. 6, H. connectens Mlldff.).
Distribution, central Philippines.
H. cincinna Sowb., viii, 16. /. mollendorffi Hid.
labiozonalis Grat. H. evanescens Brod., viii, 20.
pastorella Val. H. eburnea Rve., viii, 20.
v. gracilis Lea. H. nimbosa Brod., viii, 21.
v. virens Pfr. B. pfeifferianus Rve.
v. spretus Rv. H. elegans Semp., viii, 21.
v. rorablonensis Pfr. siquijorensis Pfr., preoc.
H. groulti JCMSS. Le Nat. '94, 136. H. carinata Lea, viii, 22.
H. succincta Rve., viii, 17. dactylus Brod.
H. cincinniformis Sby., viii, 18. v. nympha Pfr., viii, 23.
H. carneola Grat., viii, 19. H. turris Semp., viii, 23.
H. subcarinata Pfr., viii, 19. H. diana Brod., viii, 24.
f. calista Brod., viii, 24.
HELICOSTYLA. 229
H. calypso Brod., viii, 25. H. hainesi Pfr., viii. 26.
H. camelopardalis Brod., viii, 25. H. accedens Mlldff. Nachr.
v. boholensis Brod., viii, 26. ['95,99.
v. connectens MlldfF.
Group of H. concinna.
H. concinna Sowb., viii, 27. H. incompta Sowb., viii, 28.
v. flammula Semp., viii, 27. H. pyramidalis Sowb., viii, 28.
H. acuminata Sowb., viii, 28. nebulosus Pfr., viii, 29.
Section Papustyla Pilsbry, 1893.
Papustyla PILS., Man. Conch, viii, p. 243, July 1, 1893.
Shell rimate or im perforate, elongated, with slender spire; per-
istome expanded. Distribution, New Guinea, New Britain group.
H. translucida Q. & G., viii, 29. H. hindei Cox, viii, 30.
H. papuensis Hedl., vii, 190. H. heimburgi Branc., viii, 30.
Section Eudoxus Albers, 1850.
Eudoxus ALB., Die He!., p. 137 ; edit. v. Mart., p. 179. type B.
effusus Pfr.
Shell ovate or ovate-conic, imperforate, smooth, shining, very light
colored, generally thin and destitute of hydrophanous cuticle. Col-
umella rather narrow, its face flattened. Type H. effusa Pfr., pi.
53, fig. 11.
Distribution, Marinduque, Luzon, Romblon, Burias.
H. effusa Pfr., viii, 31. H. virginea Lea, viii, 36.
/. fasciata Pils. bullula Brod.
H. halichlora Semp., viii, 32. albinusGrat.
H. leai Pfr. viii, 32. chloroleuca Mart.
H. jonasi Pfr., viii, 32. bustoi Hid.
albersi Pfr. H. hololeuca Pfr., viii, 37.
buschi Pfr. H. smaragdina Rve., viii, 37.
perdita live. v. nigrescens Semp.
breviculus Rve. v. lutea Semp.
? leai Pfr. v. striata Semp.
H. simplex Jonas, viii, 33. v. zonifera Semp.
230 HELICOSTYLA.
H. quadras! Hid., viii, 34. H. straminea Semp., viii, 39
cossmanniana Cr. H. cumingi Pfr., viii, 39.
H. modesta Sowb., viii, 35. H. segle Brod., viii, 40.
B. hindsi Pfr. v. barandse Hid., viii, 40.
B. verecundus Rve. H. oviformis Semp., viii, 40.
H. belcheri Pfr., viii, 35. H. uber Pfr., viii, 41.
B. hindsi Rve. H. phseostyla Pfr., viii, 41.
H. lacerata Semp., viii, 36.
paradoxa Semp., olim.
Section Phengus Albers, 1850.
Phengus ALB., Die Hel., p. 137, for B. opalinus and B. evanes-
cens. v. MART., Die Hel., 1860, p. 180, type H. opalina.
Shell thin, pale green, distinctly trochoidal, and with no hydroph-
anous cuticle. Type H. opalina Sowb., pi. 53, fig. 5.
These shells have the texture and color of Eudoxus, but differ in
their trochiform contour. Anatomically, Phengus forms a transi-
tion to Canistrum, the dart sack being small and without a dart,
and the globular mucus gland is much reduced in size.
H. opalina Sowb., viii, 42. H. dumonti Pfr., viii, 42.
Section Canistrum Morch, 1 852.
Canistrum MORCH, Catal. Yoldi, p. 31. PILS., Manual viii, p.
43.
Shell ovate-conic or oblong, imperforate and solid ; banded ; with
or without hydrophanous cuticle ; surface microscopically spirally
striated ; whorls narrow ; lip well expanded ; columella vertical.
Type H. ovoidea Brug., pi. 53, fig. 4.
Genital system as in Helicostyla except that the dart sack and
mucus glands are absent (pi. 54, fig. 1, H. stabilis).
That the simplicity of the generative system is probably the re-
sult of degeneration of the dart apparatus is shown by the approach
to this condition in Phengus. The anatomy of Canistrum is other-
wise as in Helicostyla. It is very desirable that more species be
examined anatomically, for intermediate stages of development.
When Prochilus and Chrysallis are dissected, their anatomy may
throw light upon the condition of Canistrum.
H. ovoidea Brug., viii, 43. costerii Eyd.
luzonicus Sowb. euryzonus Pfr.
HELICOSTYLA. 231
H. balanoidea Jonas, viii, 44. H. brevicula Pfr., viii, 45.
H. stabilis Sowb., viii, 45. H. velata Brod., viii, 12.
v. elougata MlldfF.
Section Prochilus Albers, 1860.
Prochilus ALB., Die He!., p. 179, type Bui. virgatus.
Shell narrowly perforated, of an elongated ovate-pyramidal long-
spired form, smooth and shining. Aperture less than half the length
of shell; peristome very broadly reflexed. Soft anatomy unknown.
Type H. virgata Jay, pi. 53, fig. 9.
Strikingly handsome elongated shells, confined to Mindoro and
the Cuyos Is.
H. calobapta Jonas, viii, 46. H. virgata Jay, viii, 48.
cuyoensis Rve. labrella Grat.
H. cuyoensis Pfr., viii, 47. v. porracea Jay.
H. fictilis Brod., viii, 47. v. sylvanoides Semp.
v. larvata Brod., viii, 48. v. pulchrior Pils.
H. nigrocincta Semp., viii, 50. H. dryas Brod., viii, 49.
pan Pfr., not Brod. paradoxus Pfr.
H. mercurius Pfr., viii, 50. H. partuloides Brod., viii, 50.
II. calamianica Q. & M., Nachr.,
'95, 99.
Section Chrysalli* Albers, 1850.
Chrysallis ALB., Die Hel., p. 140 (in part). v. Mart., Die Hel.,
1860, p. 179, type B. chrysalidiformis.
Shell perforated, elongated ovate-conic, solid, opaque and not
smooth; aperture ovate, less than half the shell's length; lip
broadly expanded ; columellar margin dilated and reflexed ; hy-
drophanous cuticle usually present. Soft anatomy unknown. Type
H. chrysalidiformis Sowb., pi. 53, fig. 8.
Evidently allied to Prochilus. Inhabits Mindoro only.
H. chrysalidiformis Sby., viii, 51. H. mindoroensis Brod., viii, 52.
v. ustulata Jay. wagneri Grat.
v. antonii Semp. aspersus (part) Grat.
H. electrica Rve., viii, 53. melanogaster Mo'rch.
lichenifer Morch.
/. cailliaudi Petit.
232 LEUCOCHROA.
Genus LEUCOCHROA Beck, 1837.
Leucochroa B., Index Moll., p. 16, in part (keeled Xerophila, etc.,
with L. cariosa, cariosula, candidissima). MORCH, Cat. Yoldi, p. 5,
1852, Mai. Bl. iv, p. 109. KOBELT, Nachrbl. 1875, p. 37 ; Iconogr!
Land- u. Siisswasser-Moll. n. ser., iii, p. 29. Calcarina MOQ.
TAND., Mem. Ac. Toulouse, iv, 1848, and Moll. Fr. ii, p. 69 (not
Calcarina d'Orb.).
Shell solid and strong, chalky, white, subglobose or depressed, and
keeled, at least when young ; axis hollow, often closed in the adult ;
surface smoothish or pitted ; embryonal shell consisting of about 1
smooth whorls ; last whorl generally deflexed in front. Aperture
small, half-round, oblique ; lip blunt and simple (in section Sphinc-
terochila much contracted) the columellar insertion dilated, ends re-
mote. Type L. candidissima, pi. 56, fig. 13 (see also pi. 56, figs.
14, 15, L. cariosa).
Animal with rather small foot ; upper surface coarsely granular,
with a pair of dorsal grooves ; facial grooves indistinct ; no foot
margin, caudal pore or longitudinal line on the tail. Sole distinctly
tripartite, the middle area wide, side areas narrow, meeting at tail.
Mantle-edge rather thick, with very rudimentary right and left
body lappets. Eight eye retractor passing to the left of the genitalia,
not between its branches.
Jaw (pi. 36, fig. 14, L. candidissima) solid, arcuate, with a low
median projection, its surface entirely smooth.
Radula of the type usual in Helicidce. Middle tooth with square
basal plate shorter than the large mesocone ; side cusps absent.
Lateral teeth similar but asymmetrical. Marginals with shorter
basal plate, the inner cusp (ento- plus mesocone) bifid, ectocone
small, simple or bifid (pi. 36, fig. 13, L. candidissima. PI. 36, fig.
16, L. boissieri).
Genitalia (pi. 36, fig. 15, L. bcetica ; pi. 57, figs. 52, 53, L. candi-
dissima') : penis very short, narrowing into a much twisted epiphal-
lus, upon which below, the retractor muscle is inserted ; terminat-
ing in a flagellum and vas deferens. Vagina stout, bearing a
flattened spiral, or an elongated gland upon a slender short duct;
spermatheca duct long, its lower half convoluted upon the base of
the uterus, to which it is closely bound ; bearing a short stouter
diverticulum, the end of which is sunken in the uterus ; upper por-
tion of the spermatheca duct slender, straight, bound to the uterus
LEUCOCHROA. 233
and terminating in a globular spermatheca (pi. 57, fig. 53, duct
dissected away from uterus and straightened). Ovo-iestis very large
and compact, completely occupying the earlier 1 whorls.
Distribution, circum-Mediterranean region. The area occupied
by this genus is the same as that of Otala plus Levantlna,
being coincident with the region where the olive grows. As in
Macularia one species (vermiculata') extends throughout the range
of the group, so in Leucochroa, L. candidissima has an equally wide
distribution, occurring in Palestine (v. hierochuntina) , northern
Africa and westward in Europe to southern Spain. The other spe-
cies are all local. Many of them show not only much individual
variation, but also numerous well-marked local varieties; and the
complete tale of these has not yet been told.
This genus is distinguished by its cretaceous solid shell, conspic-
uously tripartite sole, smooth jaw and the teeth and genitalia of
Helix, except that the dart sack is wholly absent, the mucous ap-
pendages reduced to one straight or coiled sacculated gland, and
the ovotestis not enveloped in the digestive gland.
The group has had a varied literary existence, Moquin-Tandon,
in 1848, removing it from the Helices to Zonites on account of the
smooth jaw ; and later systematists, Martens, Westerlund, Kobelt
and others have adopted this view in their several works. Binney,
upon examining the teeth of L. boissieri, declared it a Helix, and
has been followed by Fischer and Tryon. It only remains to say
that there can be no doubt that Leucochroa belongs to the belogo-
nous Helicidse, and has riot the slightest affinity to the Zonitida3.
It is more nearly allied, in the peculiar position of the eye retractor,
to Helicella than to other genera, but differing in the loss through
degeneration of the dart and its sack, and in the smooth jaw both
of these being purely secondary modifications. I have retained the
genus in Belogona euadenia on account of the sacculated mucus
gland of candidissima ; but a careful dissection of some species with
elongated mucus gland, like bceiica, should be made, with histologi-
cal examination of the mucus gland and the minute spur at its mid-
dle (see pi. 36, fig. 15), to ascertain more certainly the place of the
genus. Probably the spur mentioned is a remnant of the dart sack.
The anatomy is known by Schmidt's figures representing candidis-
sima, bcetica, cariosa and cariosula (= hispanica), and the writer's
dissection of candidissima.
234
LEUCOCHROA.
Two sections are recognized, Leucochroa, with the lip simple,,
type candidissima, and Sp hinder ochila Anc., with the mouth angu-
lar, much contracted by an inward thickening of the lip, and a bifid
nodule in the posterior angle.
Section Leucochroa Beck.
L. candidissima Drap., iii, 10.
/. rimosa C. & J.
v. hierochuntina Boiss.
v. sardoa Malz., viii, 55.
sarda Kob., on pi.
L. isserica Kob., viii, 57.
L. bsetica Rossm., iii, 11.
v. alexandrina Fag., iii, 11.
v. tunetana Let. & Bgt.
L. otthiana Fbs., iii, 11.
v. thayaca Bgt., iii, 11.
v. titanodolena Pch., iii, 11.
v. jeannotiana Terv., iii, 11.
Zonites piestius Bgt.
v. chionodiscus Pfr., iii, 11.
L. spiranomala Bgt., viii, 55.
spelranomala Bgt., in Pech.
L. argia Bgt., iii, 12.
L. adanensis Naeg., viii, 57.
L. prophetarum Bgt., iii, 12.
L. cariosula Mich., iii, 13.
L. fimbriata Bgt., iii, 12.
v. myopa West.
v. illicita Mss.
v. varicosula West.
L. debeauxi Kob., viii, 55.
L. may rani Gass., iii, 13.
v. subcariosula Bgt., iii, 13.
kobeltiana Deb.
L. octinella Bgt., viii, 55.
vet ula West.
L. hispanica West., viii, 56.
L. saharica Deb., viii, 56.
L. cariosa Oliv., iii, 13.
v. amphicyrta Bgt.
v. nazarensis Mouss.
v. crassocarina Mouss.
L. ultima Mouss., iii, 14.
L. pressa Mouss., iii, 14.
L. accola Mouss., iii, 14.
Section Sphinderochila Ancey.
Mima WESTERLUND, Fauna Palaiirct. Binnenconch., i, p. 88 r
1886 (for boissieri a.udfilia'). Not Mima Meigen, Diptera, 1820.
Sphinderochila ANCEY, Conch. Exch., Aug., 1887, p. 23 (for filia
and boissieri).
Shell solid white and chalky like Leucochroa; but the aperture is
contracted by a building inward of the lip on its outer margin and
at the sutural angle. Jaw and teeth as in Leucochroa. Type L*
boixsieri, pi. 56, figs. 11, 12.
Distribution, Palestine and northern Arabia.
L. boissieri Charp., iii, 14.
v. zonata Bgt.
L. filia Mouss., iii, 15.
BELOGONA-SIPHONADENIA. 235
BELOGONA SIPHONADENIA.
Dart-bearing Helices in which the mucus glands are tubes of
equal diameter throughout, inserted directly upon the vagina, never
upon the dart sack.
This definition, while it perfectly distinguishes the group under
consideration from the Euadenia (p. 175), in which the mucus ap-
paratus consists of glandular lobes, flat or globular, and with few
exceptions inserted on the dart sack, will not cover all forms refer-
red to Siphonadenia. The diagnosis-defying process of retrogres-
sive evolution or degeneration has produced forms in which the dart
apparatus and mucus glands have dwindled to a mere vestige, or
been entirely lost ; reverting to the condition found in the Epiphal-
logona, which as I have elsewhere attempted to show, were the stock
whence Belogona arose. In these cases recourse must be had to
such other organs as have not shared the degenerative process; to
less divergent species, and to embryology for clues to the true his-
tory of doubtful forms. We cannot too strongly insist upon the rec-
ognition of that great difference between a primitive structure and
similar structure produced by a reverse process from a more com-
plicated organ. To lose sight of this would be to lose the best mes-
sage these studies can bring us, and reduce systematic zoology to a
mere index.
Fortunately, we have in the recent fauna, a considerable number
of species showing clearly the various stages of degeneration which
have resulted in those simplified forms of the genera Helicella and
Hygrornia which will be found noticed in the account of those groups.
The evidence indicates that such forms as Ciliella, Metafruticicola,
Cochlicella, etc. are recent degenerate groups, quite independently
produced from at least four normal Belogonous types. It is note-
worthy that the penis, jaw, radula and shell show no retrogressive
features in these forms, but retain the characters normal for the
genera they are believed to have descended from. The penis is not
(as vonlhering states) of the Patula type (Haplogona), but is dis-
tinctly of the form normal in Belogona and Epiphallogona.
All recent Helices of Europe (except the Pyramidulas) belong to
this division of the Belogona, and the same is probably true of the
Tertiary fossil forms. Just as anatomical data have enabled us to
eliminate the foreign group Triodopsis from this fauna, so more
philosophical study must cause us to see in the supposed Corasia r
Chloritis, Obba, Pella, Charopa, Mesodon, Coryda, etc. of the tertiary,
236 BELOGONA-SIPHONADENIA.
merely the ancestors of groups now living in Europe, and lateral
branches of those phyla. The presence of snails belonging not only
to modern genera, but to modern subgeneric or sectional groups, as
low as the lower Miocene, indicates that for the roots of even these
weakly characterized divisions, we must look still earlier; and the
large spaces of Eocene time can scarcely be held sufficient for the
differentiation of the genera now occupying the European tract.
The absence of Belogona Siphonadenia from all regions except those
now occupied by that group is negative evidence tending toward
the view that the group developed its special peculiarities in that
quarter of the world; and while this sort of evidence is always in-
conclusive, it has some weight in the total absence of facts making
against it. A provisional hypothesis might be outlined, holding
that the primitive Belogona (with the genital structure like Helico-
styla) spread westward before or at the beginning of Eocene time, and
in the Eur- African tract the stock became modified by the removal
of the mucus glands from the dart sack, and their change into the
tubular form, into the siphonadenious type ; subsequently splitting
into a considerable number of genera. Those genera which have
spread again from this center are mainly minute forms capable of
living in cold regions, such as Vallonia and A canthinula ; but the
presence of Helicodonta and Metodontia in China, and of the East
Asian genus Eulota in Europe, indicates a more southern connection
also. These exchanges between the faunas of the east and west ex-
tremes of the Palsearctic continent are remarkably few, however ;
and we are compelled to believe that since the incursion which
brought Belogona and many other Oriental types to Europe, the
climatic or other conditions prevailing in Central Asia and Siberia
have been unfavorable to the spread of land mollusks.
Of course there is no reason to believe that Helices of the Epiphal-
logonous type did not also reach Europe with or before the Belogona ;
and they may have survived there during Eocene and even Miocene
times ; in fact the genera Dentellocaracolus, Fridolinia, etc., may re-
present such survivors. But to state that this is the case, or that
those genera belong to the Epiphallogona (i. e. are related to Cara-
colus, Obba, Chloritis, etc.) is merely to state one's pleasingly sensa-
tional flights of fancy as scientific truth. The evidence showing the
presence of Ephiphallogona in Europe at any time, rests now upon
the finding of certain rather heavy, rudely sculptured forms; but
they are neither heavier nor more coarsely wrinkled than some Hemi-
BELOGONA-SIPHONADENIA. 237
cyclas, and may as readily have belonged to Belogona as to Epiphal-
logona, for anything now known ; and while we should not at the
present stage of malacology deny the presence in European Eocene
and Miocene of genera allied to Obba, Caracolus (=Pleurodonte),
etc., neither should palaeontologists lightly affirm that " Geotroeku^ 1
Obba, Chloritis, etc., exist in European Tertiary, on the strength of
mere resemblances of contour and sculpture characters of no sys-
tematic value, and now abandoned by all helicologists in studying
recent Helices.
Synopsis of recent genera.
1. Dart sack 1 ; mucus glands 2 or in 2 clusters ; spermatheca on a
very long duct, usually with diverticulum ; shell usually conspic-
uously banded.
a. Jaw with strong, convex vertical ribs ; dentition normal.
b. Dart four-bladed ; diverticulum free when present ;
shell typically five-banded, Helix*
bb. Dart two-bladed ; diverticulum always present, united
by a wide membrane to uterus ; shell none to three
banded, Helicigona.
aa. Jaw with converging flattened ribs ; dentition normal,
Leptaxis*
aaa. Jaw smooth ; teeth all unicuspid and strap-shaped,
Allognathvs.
2. Dart sack 2, 1 or 0, the dart bladeless or two-bladed ; mucus
glands 0, 1, or several, rarely more than 2-branched ; spermatheca
duct short, with no diverticulum ; shell with many or no bands.
a. Right eye-retractor passing between branches of genitalia,
shell unicolored or 1-banded, rather corneous in texture.
b. Shell with well-reflexed and thickened lip, often toothed,
Helicodonta.
bb. Shell with simple or expanded lip, texture corneous,
aperture lunate.
c. Depressed-globose or depressed, not laminate, size
moderate or small, Hygromia.
cc. Conoidal, with costate or lamellar riblets; minute,
Acanthinu/a.
bbb. Aperture round, oblique, toothless; shell minute, de-*
pressed, few-whorled, Vallonia.
"238 GEOMITRA.
act. Right eye-retractor passing to left of gen Italia; shell more
or less chalky ; lip simple or expanded, Helicella.
? Geomitra.
This order of groups is reversed in the following pages. '
Genus GEOMITRA Swainson, 1840.
= Geomitra SWAINS., -j- Plebecula, Helicomela, Lemniscia, Hi
pidella, Spirorbula, Irus, Aciinella, Rimula, Callina, Caseolus t Hyi
tricella, Discnla, Tectula, Plaoentula, Ooronaria and Craspedaria oi
LOWE, 1852-1854, + Ochthephila BECK, 1837, not Fallen, 1823, -f
Heterostoma HARTM., 1841 to '44, -|- Turricula WOLLASTON, 1878,
not of H. & A. Adams, 1856.
Shell generally solid, rather cretaceous, unicolored or from one to
three banded : varying from globular or pyramidal to lens-shaped
or planorboid, the umbilicus open or closed. Aperture half-round
or circular; lip more or less expanded, at least at the columella,
usually thickened within, but having no lip-rib as in Helicella
columella dilated or reflexed. Type G. tiarella W. & B. (See pi.
68, figs. 1-19).
Jaw low, slightly arcuate, with 15 broad, flat, crowded ribs in
tiarella, about 8 broad, separated ribs in lurida. In abjecta there is
a blunt median projection but no ribs.
Radula (pi. 67, fig. 18, G. abjecta Lwe. ; pi. 70, fig. 40, G.
lurida Lwe.) having well-developed side-cusps on middle teeth, the
middle cusp about as long as the basal-plate. Lateral teeth bi-
cuspid. Marginals with the inner cusp long, oblique, and feebly
bifid, outer cusp bifid or even trifid.
Distribution : Madeira group of islands. Only the most unsatis-
factory evidence exists to give ground for believing this genus to
occur outside of the Madeira group, except as occasional immigrants,
unless the occurrence of G. paupercula Lowe in the Azores and
Canaries be owing to natural causes. Those indigenous species of
the Canaries referred to Hispidella, Discula t Ochthephila, etc., may
better be left in Hygromia, Jacosta and other groups, until they may
be shown to actually have some characters of the Madeira forms.
The Canary Island Helix fauna is far more closely allied to that of
northern Africa than to that of Madeira.
It would obviously be quite idle to discuss the origin or genesis of
this genus until its anatomy is made known. We are quite safe in
GEOMITRA. 239
believing it an ancient inhabitant of the Madeira group, and its
peculiarities have probably been developed upon that soil, for neither
in the fossil series of Europe or the recent fauna of Eur-Africa or
the other Atlantic islands are there known forms which may be re-
ferred to the Madeira genus. It is therefore much more restricted
that the Leptaxis group. In this connection con/. WATSON, The
Journal of Conchology. vii, p. 1, 1892.
A large number of subgenera or sections have been founded for
the Madeira Helices, which is not surprising when we consider the
astonishing amount of modification of the numerous minor groups,
altogether unparalled in any other tract of like extent in the world.
Most of the following sectional groups have already been associated
by Martens, Pfeiffer and others ; but Plebecula, Helicomela and His-
pidella are now added to the group for the first time.
The name Ochthephila being preoccupied, I have been obliged to
substitute Swainson's term Geomitra. This has priority over Heteros-
toma Hartmann, as well as over the entire series of names proposed
by Lowe.
Subgenus PLEBECULA Lowe, 1852.
Plebecula LWE.. Ann. Mag. Nat. Hist. (2), ix, p. 114, Feb., 1852,
for giramica, vulgata, canicalensis Lwe. ; P. Z. S., 1854, p. 172, type
H. vulgata Lwe. Helicomela LWE., P. Z. S., 1854, p. 172, type H.
pundulata Sowb.
Shell globose-depressed with conic spire, or subglobular, umbili-
cate or imperforate, solid, above rather rudely striated, granose or
hirsute ; unicolored, or 3-banded above on a brown ground, the base
paler and uniform. Whorls 5-6, separated by deep sutures. Aper-
ture but little oblique, subcircular ; lip hardly expanded, sharp or
thickened within ; columella reflexed. Type G. nitidiuscula Sowb.
(See pi. 43, fig. 26, G.punctulata Sowb.).
(Shell subglobular, imperforate; Helicomela).
G. punctulata Sowb.,iv, 187. G. bowdichiana Fer., iv, 187.
v. avellana Lwe., iv, 187. vargasiana Pfr.
(Shell depressed-globose with conic spire, umbilicate; Plebecula).
240 GEOMITRA.
G. nitidiuscula Sowb., iv, 188. G. nitidiuscula.
vulgata Lwe. v. saxipotens Woll.
v. giramica Lwe., iv, 188. v. canicalensis Lwe., iv, 188.
anaglyptica Rv. G. lurida Lwe.,iv, 188.
v. deserticola Woll. nitidiuscula Woll., not Sow.
v. pulchra Paiva. v. hartungi Alb.,iv, 189.
Subgenus LEMNISCIA Lowe, 1854.
Lemniscia LWE., P. Z. S., 1854, p. 170, type H. michaudii Dh.
Shell barely perforate, nowhere granular, globose- conoid or glo-
bose-depressed, with numerous (6? to 8) slowly widening whorls,
those of the spire striated ; last whorl but little descending, the base
smooth. Aperture half-round, lip blunt, thickened within, ex-
panded toward the columella, its ends remote, parietal wall nude.
Type G. michaudii Dh.,pl. 68, figs. 14, 15.
G. michaudii Dh., iv, 21. G. calva Lwe., iv, 41.
bicolor Lwe. G. galeata Paiva, iv, 41.
Subgenus HISPIDELLA Lowe, 1852.
Hispidella LWE., Ann. Mag. N. H. (2), ix, Feb., 1852, p. 115,
for armitageana, revelata, sericea; P. Z. S., 1854, p. 178, type, If
hispida L.
Shell thin, fragile, brown, not chalky ; perforate, convex-depressed ;
surface bearing flattened cuticular scale-like processes, simulating
the hairs of Trichia. Whorls less than 5, the last angular, slightly
deflexed in front. Aperture half-round, slightly lunate, the peri-
stome slightly expanded, reflexed at columella, ends remote. Type
G. armitageana Lwe.
Has a superficial resemblance to the Fruticicoloid Continental
forms, but the sculpture is like that of Lowe's section Irus, and I
am disposed to consider the group as a member of the present genus.
Lowe was clearly in error in naming as the type of his group a spe-
cies which he had not mentioned in his original publication of the
name. H. horripila, a doubtful member of the group, is from the
Azores.
G. armitageana Lwe., iii, 223. G. horripila M. & D., iii, 222.
Subgenus SPIRORBULA Lowe, 1852.
Spirorbula LWE., Ann. Mag. N. H., Feb., 1862, p. 114 (proposed
for H. latent and obtecta) ; P. Z. S., 1854, p. 175, type H. obtecta
GEOMITRA. 241
Lwe. Irus LWE., Ann. Mag., Feb., 1862, p. 114 (laciniosa,
squalida, depauperata)', P. Z. S., p. 174, type H. depauperata Lwe.
Not Irus Oken, Naturgeschichte fiir Schulen, p. 647 (1821).
Shell perforated, globose-depressed, with conoidal or flattened
spire, the whorls about 5, rounded, sutures deep; surface smoothish
or coarsely wrinkled, sometimes bearing recurved cuticular scales.
Aperture slightly oblique, round or oval, the parietal lip continu-
ous and adnate. Type G. obtecta, pi. 68, fig. 13.
As the name Irus is preoccupied, its species may be merged in
Spirorbula, which offers no very marked difference.
G. obtecta Lwe., iv, 35. G. depauperata Lwe., iv, 36.
G. latens Lwe., iv, 35. G. latinea Paiva, iv, 36.
G. squalida Lwe., iv, 35. G. laciniosa Lwe., iv, 36.
Subgenus ACTINELLA Lowe, 1852.
Actinella LWE., Ann. Mag. N. H. (2), ix, Feb., 1852, p. 118, pro-
posed for stellaris lentiginosa, arcta; P. Z. S., 1854, p. 180, type H.
lentiginosa Lwe. Rimula LOWE, t. c., p. 118, for obserata andfausta;
P. Z. S., 1854, p. 181, not of Defrance, 1827 (see Man. Conch. [I],
xii, p. 269)-}- Callina LWE., t. c., p. 183, sole species H. rotula Lwe.
Shell brownish or variegated, depressed-globose, the periphery
subangular or keeled, umbilicus narrow or closed. Surface scaly,
striate or rather sparsely granulated. Aperture oblique, peristome
expanded and thickened within, its margins not much converging,
parietal callus usually rather slight. Type G. lentiginosa Lwe., pi.
68, figs. 4, 5 (see also pi. 68, fig. 7, G. [Callina] faustd).
Distribution, mainly Madeira.
(Perforate or umbilicate, the callous of basal lip not toothed,
Actinella.)
G. lentiginosa Lwe., iv, 38. G. stellaris Lwe., iv, 38.
G. actinophora Lwe., iv, 40. G. arcta Lwe., iv, 38.
v. descendens Woll. G. arridens Lwe., iv, 40.
(Imperforate or nearly so, compact, granulate, basal callus strong
and truncate, Callina.}
G. arcinella Lwe., G. obserata Lwe., iv, 40.
G. fausta Lwe., iv, 40. v. bipartita Woll.. iv, 40.
v. robusta Woll. G. capsella Lwe., iv, 41.
G. rotula Lowe, iv, 46.
16
242 GEOMITRA.
Subgenus CASEOLUS Lowe, 1852.
Caseolus LOWE, Ann. Mag. N. H. (2), ix, Feb., 1852, p. 115, for
sphcerula, compada, abjecta; P. Z. S., 1854, p. 184, type H. compacta
Lwe. -- \-Hystricella LWE., P. Z. S., 1854, p. 186, type H. bicarinata
So\v.+Discula LWE., t. c., p. 116 ; P. Z. S., 1854, p. 187, type H.
polymorpha Lwe.-f Tectula LWE., t. c., p. 117 ; P. Z. S., 1854, p. 191,
type H. bulveriana Lwe. Turricula]Woi^., Test. Atlant., p. 168,
type H. cheiranthicola Lwe. (1878). Not Turricula (Klein) H. &
A. Ad.= Turris Montf.
Shell perforate or umbilicate, varying from globose-depressed
with rounded periphery, to subdiscoidal or to pyramidal, with
keeled or double-keeled periphery. More or less granulated.
Aperture rounded or oval, the lip blunt, usually a little expanded
at columella. Type G. compacta, pi. 68, fig. 19 (see also G. (Hys-
tricella) bicarinata, pi. 68, fig. 12 ; G. (Discula~) polymorpha, pi. 68,
figs. 8, 9.)
(Perforate ; globose-depressed or globose-conic, periphery rounded
or bluntly angular, parietal lip adnatc ; surface striate and granu-
late above, smooth or granose below ; color whitish. CASEOLUS).
G. consors Lwe., iv, 39. G. sphserula Lwe., iv, 39.
G. calculus Lwe., iv, 39. subcallifera Lwe.
G. compacta Lwe., iv, 39. G. abjecta Lwe., iv, 39.
innominata Gray. v. candisata Mke.
G. commixta.
(Perforate, trochiform, with acutely keeled or double-keeled per-
iphery ; parietal lip raised and freej; surface sharply granose through-
out; color dusky. HYSTRICELLA).
G. echinoderma Woll., iv, 34. G. vermetiformis Lwe.
G. echinulata Lwe., iv, 33. G. turricula Lwe., iv, 33.
G. bicarinata Sowb., iv, 33. v. pererosa Woll.
duplicata Lwe. G. leacockiana Woll., iv, 34.
G. oxytropis Lwe., iv, 33.
( Umbilicate, much banded and varied with brown ; depressed,
subdiscoidal or pyramidal, carinated ; surface smoothish, inconspic-
uously granular. DISCULA).
GEOMITKA. 243
G. tetrica Paiva., iv, 44. v. papilio Lwe., iv, 45.
O. polyraorpha Lwe., iv, 44. calcigena Lwe., iv, 45.
elegantula Jan. v. discina Lwe., iv, 45.
saccharata Lwe. v. gomesiana Pva., iv, 4-5.
tmniata Rv. v. attrita Lwe., iv, 46.
v. salebrosa Lwe., iv, 44. G. cheiranthicola Lwe., iv, 46.
v. poromphala Lwe., iv, 44. v. mustelina Lwe., iv, 46.
v. pittse Pva., iv, 44. G. tabellata Lwe., iv, 46.
v. alleniana Pva., iv, 45. G. testudinalis Lwe., iv, 46.
v. lincta Lwe., iv, 45. bulveriana var., Rv.
v. arenicola Lwe., iv, 45. G. lyelliana Lwe., iv, 41.
cinerea Lwe., iv, 45. G. albersi Lwe., iv, 42.
v. barbosse Pva., iv, 45. G. bulverii Wood, iv, 42.
v. pulvinata Lwe., iv, 45. bulveriana Lwe.
(Umbilicate, lenticular, with a compressed and deflexed keel ;
base coarsely granose ; texture chalky ; white).
G. tectiformis Sowb., iv, 42. . var. ludovici Alb., iv, 42.
Section Disculella Pils., 1894.
Ochthephila BECK, Index Moll., p. 17. A LEERS-MARTENS, Die
Hel., p. 118, type H. maderensis. Not Ochthiphila Fallen, 1823
{Diptera'). Placentula LOWE, Ann. Mag., Nat. Hist. (2),ix, p. 118,
Feb., 1862 ; P. Z. S., 1854, p, 194, type H. maderensis. Not Pla-
centula Lam., 1822.
Shell discoidal, umbilicate, with convex base and spire, the periph-
ery keeled ; solid, brown or whitish banded and maculated with
brown. Surface striate above, smoother below. Whorls about 6,
the last deflexed ; aperture circular, oblique, the lip slightly ex-
panded, narrowly white lipped within. Type G. maderensis, pi.
68, figs. 10, 11.
Distribution, Madeira Is. Distinguished from the very closely
allied section Discula by the smoother, hardly granulate surface,
and the rounder mouth.
G. compar Lwe., iv, 37. G. leptosticta Lwe., iv, 37.
G. tseniata W. & B., iv, 37. G. micromphala Lwe., iv, 37.
G. maderensis Wood, iv, 37. G. dealbata Lwe., iv, 38.
cyclostoma Mke. G. fictilis Lwe., iv, 38.
G. spirorbis Lwe., iv, 41.
244 GEOMITRA.
Subgenus HETEROSTOMA Hartman, 1844.
Heterostoma HARTMAN, Erd- und Siisswasser Gasteropoden der
Schweiz, mit zugabe einiger merkwiirdigen exotischen Arten, p.
177, type H. semitecta Hartm., pi. 62 (vii), f. 1-4, = H. paupercula
Lwe.
Shell small, planorboid, umbilicated, angular or keeled at periph-
ery, with 4 to 5 whorls, the last deflexed, abruptly contracted at the
aperture, the lip-edge thin and slightly expanded, continuous ; basal
lip strongly arcuate, with a heavy callous rib within, which ends in a
tooth within the outer lip ; parietal wall elevated. Type G. pauper-
cula Lowe, pi. 68, figs. 16, 17, 18.
The two species grouped here have been widely separated in for-
mer classifications.
G. paupercula Lwe., iv, 35. G. coronata Desh., iv, 34.
semitecta Hartm. juliformis Lwe.
tracheloides Mke.
Subgenus GEOMITRA Swainson, 1840.
Geomitra S\v., Malacology, pp. 166, 332, type H. tiarella. Coro-
naria LOWE, Ann. Mag. (2), ix, p. 117 for coronula and tiarella; P.
Z. S., 1854, p. 193, type tiarella (preoc.). Craspedaria LOWE, t. c.
p. 117, and P. Z. S., 1854, p. 192, type H. delphinula.
Shell depressed or conoidal, solid, dull brown, rudely sculptured ;
whorls of the spire plicate below the sutures, more or less keeled at
periphery ; base cylindrical, umbilicated, sculptured with granose
spiral cords. Aperture nearly round, very oblique, the peristome
expanded, thin, continuous, solute, the parietal callus raised from
the preceding whorl. Type G. tiarella Webb & Berthelot, pi. 68,
fig. 6. (See also pi. 68, figs. 1-3, G. delphinula}.
Jaw (of tiarellay m \ovf, slightly arcuate, the anterior surface with
about 15 flat, broad, crowded ribs, scarcely denticulating the cutting
margin. (Binney).
Radula with 12. 9. 1. 9. 12 teeth of same character as figured for
Plebecula lurida.
Distribution, Madeira. This group is extremely peculiar in the
coarse spiral sculpture of the^base, and the coronated whorls of the
spire.
GEOMITRA-HELICELLA. 245
Swainson in his first reference to this group figures the H. tiarella,
but does not mention it by name. In his later reference he con-
fuses tiarella with the figure and description of H. bicarinata Sow.,
but his diagnosis shows clearly the species intended.
G. tiarella W. & B., iv, 35. G. grabhami Woll.
G. moniziana Paiva., iv, 34. G. delphinuloides Lwe., iv, 34.
G. coronula Lwe., iv, 34. G. delphinula Lwe., iv, 44.
Genus HELICELLA Ferussac, 1819.
Helicella FEB., Tableau Syst. de la Fam. des Lin^ons, p. 37
(fourth group only). Risso, Hist. Nat. Eur. Merid. p. 67, 1826, in
part.
=Xerophila Held, 1837 and of subsequent authors, with Heli-
eopsis Fitz, 1833, Zenobia and Jacosta Gray, 1821, etc., etc.
Shell umbilicate or perforate, with either cylindrical or keeled
whorls ; opaque and earthy, white or whitish and usually banded,
not hairy ; aperture round-lunate or angular, not very oblique, the
lip acute, hardly expanded, thickened within (See pi. 68, figs. 20 to
30.)
Jaw with 4-11 wide, flattened ribs (pi. 67, figs. 12, 14).
Radula (pi. 67, fig. 13 H. caruance v. gattoi) with teeth of the
type usual in ground-snails. Median teeth with weak ectocones or
none, laterals with small divergent ectocones. On the marginals the
inner cusp is either simple or bifid, outer cusp single or split (see
also pi. 67, fig. 16, H. terrestris}.
Genital system (pi. 69, all figs.) having the penis rather short,
continued in an epiphallus bearing retractor, and ending in a short
flagellum. Dart sack single or paired, with or without accessory
sacks, and containing curved darts which are two-bladed at least
toward the point. Mucus glands simple and tubular, several in
number or numerous, but inserted individually on vagina. Sper-
matheca oval or irregular, borne on a rather short, unbranched
duct. Right eye-retractor passing to the left of genital system, not be-
tween its branches.
An exception to the above diagnosis occurs in Theba, where the
penis lacks retractor-muscle, and the dart sack is empty (pi. 69, figs.
14, 16, 22). In many species a sack-like organ or appendix, of un-
known function is developed on the penis or on the atrium ; and in
246 HELICELLA.
certain forms a spermatophore of unusual size is found, having the
rod-like form and chitinous texture noticed in Leptaxis, but with
serrate edge.
Helicella is allied to Hygromia in the simple-lipped shell, simple
form of dart and frequent duplication of the dart sack. It differs
from Hygromia in having the right eye-retractor pass to the left of
the genital system instead of between its branches, and in the solid,
earthy white shell. Outside of these two European groups the
double dart sack occurs only in the Mexican genus Lysinoe. The
peculiar disposition of the right eye retractor muscle occurs again
in Leucochroa.
The species are very numerous throughout the Mediterranean
countries, and many of them show a considerable range of individ-
ual variation, and also local or geographic racial forms ; but the
number of true species or subspecies is not over one-fourth the
number of nominal species, mostly described by authors of the so-
called Nouvelle Ecole of France. No individual variation is too
slight to be called a " species " by some of these writers ; and a large
list could readily be given of " species " founded merely on young
shells of well-known forms. Unfortunately for science, many of
these worthless names, even some demonstrated to be the young of
other species, have been adopted into works supposed to be author-
itative, such as Westerlund's " Fauna." The result is that in
Europe, where from the number of workers one would expect that
the fauna would be well worked up and understood, the study of
Helices is in a semichaotic condition so far as species work is con-
cerned, and infinitely behind the condition of the science in Amer-
ica, the West Indies or Australia.
The tertiary deposits of Europe have afforded but few members of
this genus ; and although recorded from lower Miocene deposits,
there are few if any undoubted representatives earlier than Pleisto-
cene. This seems to indicate that the group is comparatively new
to middle European soil.
The present group has usually been called Xerophila by recent
authors; but several terms proposed by Risso and Gray precede
Held's publication, besides the still earlier Ferussacian name
Helicella. Even were we to reject Ferussac's term, the next name
in order would be Jacosta Gray, 1821, founded on H. explan-
ata. In no case can one use Xerophila as a generic name without
throwing the rule of priority to the winds. There are, however,
HELICELLA. 247
plenty of writers quite willing to do this. Xerophila has been used
in Ornithology (P. Z. S. 1840, p. 175) but later in date than Held's
publication.
The division of the genus into sections is difficult on account of
the large number of species intermediate in form; and the data at
hand are insufficient for their discrimination on anatomical grounds.
It is further complicated by the number of names proposed for mem-
bers of the group, and their intricate synonymy. Although per-
haps fairly able to distinguish small systematic groups, the writer
claims to be no expert in perceiving the subtle distinctions made by
some authors in this genus. Those who find a greater number of
sections useful should avoid two radical faults in the present Euro-
pean usage : i. e. the use of preoccupied names and the use of old
names for groups containing none of the species on which such sec-
tional terms were originally based. Monterosato has recently pro-
posed an entire new set of no less than forty-one sectional names,
which for uniformity all begin with Xero-. There is much dry
humor in this proposition, for he ignores all previous sectional
nomenclature except Xeroleuca, and the new terms are mostly
Greek X Latin hybrids, hideous in etymology and senseless in mean-
ing.
f Xerocrassa Monts. Jacosta Gray.
Subgenus HELICELLA, | Heliomanes Moq. Xeroleuca Kob.
penis retractor mus- j Helicella s.str. Obelus Hartm.
cle present and well ] Xerocampylsea Kob. Trochula Schluter.
developed. | Candidula Kob. Cochlicella Risso.
[Monilearia Mouss.
Subgenus THEBA, no f
penis-retractor mus- | Theba Risso.
cle. Muscus glands J Lejeania Ancey.
present. No dart Platytheba Pils.
in the empty sack. ^
Section Xerocrassa Monterosato, 1892.
Xerocrassa MONTS., Moll. Terr. Is. adiacenti Sicil., p. 23, type H.
seetzeni.
Shell narrowly umbilicated, thick solid and chalky, varying from
discoidal to turbinate. Type H. seetzeni.
A desert form of Heliomanes characteristic of Palestine and
Arabia, perhaps worth a sectional name.
248 HELICELLA.
H. seetzeni Koch, iii, 223. H. eremophila Boiss.,iii, 242.
sabwa Boiss. cremnophila Boiss. err. typ.
/. fasciata Mouss. v. amuniensis Mts.
/. subiuflata Mouss. H. erkellii Kub., iii, 243.
/. avia West. v. discrepans Pils., viii, 177.
H. beadlei Pils., viii, 176. H. sinaica Mart., viii, 178.
H. psammita (B.) West.
Section Heliomanes Moquin-Tandon, 1855.
Heliomanes (Fer., Tabl. Syst., not used in a generic or subgeneric
sense) MOQ.-TAND., Hist. Nat. Moll. Fr., p. 259, and of subsequent
authors. Xeroampulla MONTS., Moll. Terr. Is. Adi. Sicil. 1892, p.
22 (for aradasii, subprofuga, pellucens, enhalia). Xerofusea MONTS.
/. c. (for luctuosa, benoiti, etc.). Xerolauta MONTS., t. c., p. 23 (for
virgata, variabilis, lauta). Xerolincta MONTS., 1. c. (for arenarum,
astata, euetha). Xerolceta MONTS., 1. c. (for segusse, tuta, edulis, vari-
ata, rufolabris). Xerovaria MONTS., 1. c. (for tergestina, stroniana,
lineata), Xerambigua MONTS., t. c., p. 24 (for dantei). Xerolutea
MONTS., 1. c. (for luteata, luteola, dautezi, melania). Xeromagna
MONTS., /. c. (for cespitum, introducta, marioniana). Xeropicta
MONTS., 1. c. (for krynickii). Xerobulla MONTS., I. c. (for bollen-
ensis, robiniaua, perroudiana). Xeromunda MONTS., t. c., p. 25 (for
turbinata, candiota). Xerocauta MONTS., I. c. (for cretica, cauta).
Xerovera MONTS., 1. c. (for subrostrata, lacertarum, mauritanica, or-
anensis, cyclostoma, sphserita, caruanse, galloi, metabola, rusticana,
fraudulenta). Xerolissa MONTS., /. c. (for acompsia, aconipsiella).
Shell with moderate or small umbilicus, conoidal or low conoidal
spire and rather tubular, unkeeled whorls. Surface nearly smooth ;
solid and chalky, whitish and often banded or striped. Aperture
rounded-lunar, lip acute, labiate. Type H. variabilis Drap., pi. 68,
fig. 20.
Genital system (pi. 69, figs. 1, 2 H. variabilis'} : Dart sack con-
taining a slightly curved dart (fig. 1), and with an accessory sack ;
mucus glands with numerous tubes. In some forms two dart sacks
are developed (see also pi. 69, figs. 3, 4, 5, H. virgata*).
This group contains the largest species of the genus. The whorls
are less tubular than in Helicella s. str. and the spire generally
higher, umbilicus smaller. Some species have one, others two func-
tional dart sacks. The distribution is the same as that of Candidula
middle Europe north to England, and south to northern Africa.
HELICELLA.
249
"Species numerous, but multiplied to an almost inconceivable number
by the " new school " conchologists.
H. sitifiensis Bgt., viii, 165.
H. sphserita Hartm., iii, 249.
H. stiparum Rm., iii, 241.
H. submaritima Desm.
lauta Lwe., iii, 235.
H. subrostrata Fer., iii, 231.
H. terveri Mich., iii, 240.
arenivaga Mab.
luci FJor.
adolice Flor.
H. turbinata Jan., iii, 234.
cyclostomoides Porro.
pilula Mouss.
.H. ungeri Zel.
H. variabilis Dr., iii, 230.
striata Brard.
zonariaDon.
subalbida Poir.
burdigalensis Grat.
lautaretina Bgt.
jussiana Bgt.
grannou.ensis Bgt.
salentina Bl.
mendranoi Serv.
v. turbinata Cafici.
variata West.
H. variegata Friv., iii, 235.
v. pustulosa Parr.
H. vestalis Parr., iii, 240.
? mesopotamica Mouss.
v. foveolata West.
H. virgata DaC.
H. zaccarensis Kob., viii, 168.
.H. krynickii Andr., iii, 247.
babondubii Parr.
theodosice Cl.
radiolata Mss., iii, 240.
\v. candaharica Pfr., iii, 247.
H. lampedusse Kob., viii, 175.
H. laurinse Iss., viii, 166.
H. luteata Parr., iii, 231.
/. subluteata Serv.
/. matronoi Serv.
H. maritima Dr.. iii, 235.
iineata Oliv. not Say.
pseudenhalia Bgt.
canariensis Sh.
H. mauritanica Bgt., iii, 235.
H. millepunctata Bttg., viii, 178.
H. modica Mouss., iii, 236.
H. moesta Parr., iii, 233.
v. luctuosa Caf.
H. moneriana Bgt., viii, 165.
H. oranensis Mori., iii, 249.
H. parva Parr., iii, 232.
H. pellucens Sh., iii, 232.
H. piratarum Kob., iii, 240.
H. richardi Kob., viii, 174.
H. sebkarum Deb., viii, 167.
H. semenowi Mart., iii, 237.
H. simulata Fer,, iii, 232.
? striatula Bk.
H. derbentina Andr., iii, 247.
v. caucasica Parr.
v. depressa Ret.
v. isomera Friv.
v. armeniaca Bay.
v. suprazona Mss.
v. suberrans Mss.
v. constricta West.
H. devauxi Deb., iii, 240.
H. didyma West.
thiessece Mouss., iii, 24.
H. djebbarica Bgt., iii, 236.
H. dragorichi Zel., iii, 249.
H. durieuri Moq., iii, 236.
250
HELJCELLA.
H. erithrocheila Sul., viii, 189.
rufolabris Ben. not Jeffr., iii,
[233.
H. euphorca Bgt., iii, 230.
H. euxina CL, iii, 231.
H. fabriesi Deb., viii, 168.
H. globuloidea Terv.,iii, 243.
arenarum Bgt.
H. gouini Deb., viii, 170.
H. hamilcaris Kob., iii, 233.
H. herbicola Sh.
H. hydruntina Bl., iii, 230.
H. joppensis Roth, iii, 244.
bargesiana Bgt., iv, 7.
/. subkrynickiana Mouss.
/. multinotata Mouss.
? mesopotamica Mss.
H. caruanse Kob., viii, 174.
v. gattoi Kob., viii, 175.
H. cauta West., iii, 240.
H. cespitoides Fisch., viii, 176.
H. cespitum Drap., iii, 241.
carnina Cheir.
eurythmia Hartm.
v. dismathia Nev.
v. alticola Nev.
v. introductaZgl., iii, 242.
H. chalcidica Bl., iii, 24, 231.
H. choretaBgt., iii, 231.
H. cistorum Mor., iii, 236.
H. colomiesiana Bgt., iii, 232.
H. commeata Mouss.
H. cretica Fer., iii, 239.
v. littoralis Mouss.
v. akrotirensis Kob.
H. critica Fer.
H. cyrenaica Mts., iii, 234.
H. danieli Bgt, iii, 230.
H. dautezi Kob., iii, 248.
H. davidiana Bgt., iii, 24.
H. accompsia Bgt., iii, 231,
v. accompsiella Anc.
H. adolphi Pfr., iii, 241.
H. segusse Kob.
H. affinior Deb., viii, 166.
H. agreabilis Zgl., iii, 234.
H. amoma Bgt.
H. aradasii Piraj., iii. 233.
filograna Villa.
H. arcuata Zgl., dii, 234.
H. arigonis Rm., iii, 241.
arigoi Bgt.
H. armoricana Bgt., iii, 242.
H. benoiti Caf., iii, 233.
H. berlieri Mori., iii, 236.
? lacertarum Bgt.
H. bollenensis Loc., viii, 170.
lauracina Fag.
H. breveti Deb., viii, 169.
H. calida Kob., viii, 167.
H. calopsis Bgt., viii, 165.
H. camerata Mouss., iii, 232.
H. candiota Friv., iii, 234.
H. canina Anc., viii, 177.
H. cottyi Mor., iii, 236.
Unfigured, insufficiently known species: H. mayeti, valeryana,
eumona, pachesta, charmesiana, bilottiana, blossura, elithia, arbana,
ionstoma, ianthinostoma, amethysta Let. & Bgt. ; bousqueti (Deb.)
W. ; casertana B. ; tacapica, tabarkana, una, tebourbana Let. &
Bgt. ; therella (Berth) B. ; thera Let. & Bgt. ; foedata (Hagenm.)
B. ; tseniata W. ; dexia, neftana B. ; libertina (Let.) W. ; pompei-
ana B. ; desilvse Serv. ; ogiaca Serv. ; microspila B. ; euxina Cl. ;
HELICELLA. 251
salentina (Bl.) ; privata Galland ; zerguana (Hagenm.) ; philoxera
Caf. ; euetha (B.) ; iimara B. ; erythrsea W. ; halophila (Deb.) ;
xera (Hagenm.) ; mahdarina B. ; didiera (B.) ; nya, latastei Let.;
latasteopsis, fratisiana, tafermica, mezessaria Let. & Bgt. ; sestuosa
Berth. ; inversa, sequa W. ; zemonicensis Stoss. ; naudieri B. ; ^us*
area Anc. ; eusarcomsea Anc. ; occonella Let. & Bgt.; steriolena,
adisana Bourguignat ; psammathsea Let. & Bgt. ; bertina B. ;
eteema Let. & Bgt.; menzelensis Letourneaux & Bgt.; ram-
lensis B. ; comendadori Serv. ; panurga B. ; euphorcella Pech. ;
euphorcopsis, esnorca Let. ; meticulosa Let. & Bgt.; carpensorac-
tensis Fag. ; robiniana B. ; foliorum Fag. ; prinohila Mab. ; perrou-
diana Loc. ; visanica Fag. ; taria B. ; vettonica Serv. ; maxulana
Let. & Bgt. ; entara, zitanica Let. & Bgt. ; rhodochila W. ; loto-
phagorum, meninxica, mesembrica Let. & Bgt. ; mantinica Mab. ;
locardi W. ; panescorsi Bereng. ; dantei, calopsis B. ; eucestella,
eucesta B. ; ammederana, haidrana, birta Let. & Bgt. ; dolomitica
Deb.; rachgonica B. ; kabyliana Deb.; euthymeana Loc.; actia
B. ; actiella, nautica, suberis Loc. ; evenosi B. ; maristorum Flor. ;
cyclostoma W. ; axiotheata B. ; lemoinei Deb. ; ferianica Let. &
Bgt. ; oreta B. ; pedianopsis Hagenm. ; certa B. ; caudefacta Let. &
Bgt. ; leucophora, ingenua, acela, monerea, chioidea B. ; phoebeia
Let. & Bgt. ; spilmenti, catarota, cana, leucesthaB. ; hadrumetorum
Let. & Bgt. ; urbarana Pech. ; eucana Hagenm. ; barrattei, slouguia,
khangetina, artara, burella, boudriesa Let. & Bgt. ; armoricana,
anephela, pediana B. ; ripacurcica Bofill ; ilicis Florence ; mega-
stoma B. ; cselestis Let. & Bgt. ; meteora B. ; suspecta W. ; talepora,
acosmeta B. ; lersiana Fag. ; calographa W.
Section Helicella s. str.
Helicella Fer. t. c. (in part). Risso, t. c., p. 67, in part, and of
authors. Planatella CLESSIN, Deutsche Exc. Moll. Fauna, 1876, p.
143 (for ericetorum and candicans), Mollusken fauna Oesterreich-TJn-
garns und der Schweiz, 1887, p. 180. Xerolaxa MONTEROSATO,.
Moll. Terr, delle Isole adiacenti Sicilia, p. 24, from Atti della R.
Accad. di Scienze, Lettere e Belle Arti, (3), ii, 1892, (for ericetorum,
pamplonensis). Xerofriga MONTS. 1. c. (for nubigena). Xerogyra.
MONTS., L c. (for spadse, bathyomphala). Xerocincta MONTS., I. e.
(for neglecta). Xerolenta MONTS., L e. (for obvia, derbentina).
Pseudoxerophila WESTERL. Aperyu Faun. Mai. Grece, 1879, p. 55
(for bathytera, etc.).
252
HELICELLA.
Shell much depressed and broadly umbilicated, with smoothish,
tubular whorls, rounded at periphery, and of the usual chalky text-
ure and white, banded coloring. Aperture small, round or oval;
but little modified by the preceding whorl, the lip slightly expanded
Type H. ericetorum Mull., pi. 68, figs. 21, 22.
Jaw arched, strongly ribbed. Genital system (pi. 69, figs. 6, 7,
8, H. ericetorum) with short swollen penis, long epiphallus upon
which the retractor is inserted, and very short flagellum. High on
vagina are two symmetrically placed dart sacks (fig. 7), contain-
ing well curved round darts, provided toward the end with two nar-
row blades (fig. 8). Mucus glands numerous.
H. aberrans Mouss., iii, 246.
H. ammonis Schm., iii, 245.
/. Candida Porro.
v. sclerostoma Stef.
v. bononiensis Stef.
v. bonaldi Ad.
H. apollinis Mts.
H. bathytera Bl. & W.
/. affinis Bl.
H. bathyteropsis Serv.
H. enhalia Bgt., iii, 243.
H. ericetella Jouss., iii, 243.
H. ericetorum Mull., iii, 245.
trochlearis Andrz.
kusteri Held.
? itala L. media Gm.
dubia Hartin.
/. devian's West.
erica DaC.
obliterata Hartm.
/. tardyi Bgt.
H. gyroides Parr., iii, 246.
H. instabilis Zieg., iii, 248.
neopolitanaAudr.
spadce Calc.
nubila Charp.
ocellus Villa.
v. nubigena Charp. vi, 84.
v. bathyoraphala Charp.
v. destituta Charp.
v. discrepan? Tib.
v. grseca Mart.
? iphigenice Deb.
H. interpres West., iii, 242.
H. lemoinei Deb., iii, 246.
H. neglecta Drap., iii, 243.
clivorum Hartm.
varians Risso.
H. obvia Mke.
candicans Auct., iii, 244.
v. arenosa Z., Rm.
v. dejecta Z., Rm., iii, 246.
v. renoufi Serv.
v. pullula Parr.
v. dobrudschse Parr.
v. grseca Mts.
H. pamplonensis Schm., iii, 246.
H. spirula Zel., iii, 249.
v. bakowskyana Cl., iii, 248.
H. talmacensis Biz.
H. vukotinovici Hire., iii, 246.
liburnica Stoss.
H. vulgarissima Mouss., iii, 245.
Insufficiently known or unfigured species: H. virgultorum Bgt.,
morbihana Bgt., fagoti West., dysmica West., synerosa Serv., sal-
HELICELLA.
aunica Fag., maladettse (B.) Fag., sabulivaga and marBillyana
Mab., nephseca Fag., homoleuca Parr., tauchoniana Bgt., tennis-
culpta West.
Section Xeroeampylcea Kobelt, 1871.
Xeroeampylcea KOB., Catalog, p. 15, footnote, for H. zelebori.
Shell depressed with horn-colored apex and wide last whorl with
rounded periphery ; rather thin and white with 2 bands (or none),
umbilicus funnel-shaped but very small. Aperture transverse oval,,
the lip dilated at columellar insertion partly covering umbilicus.
Type H. zelebori Pfr., pi. 43, figs. 29, 30.
Genital system as in Helicella, two dart sacks being developed.
Distribution, Bosnia, Servia. Formerly referred to Campylcea, but
now admitted to belong to the Xerophila group.
H. zelebori Pfr., iv, 83.
The following forms or varieties of Zelebori are distinguished by
French new school authors : bortana, adarella, carosina, ottoi, twar-
tkoi, nactara and acaria Servain.
Section Candidula Kebelt, 1871.
Helicopsis FITZ., Syst. Verzeich Weichthiere, etc., 1833, p. 10L
H. striata the sole species. Not Helicopsis Fab., 1808 (Lepidoptera) r
Striatella WESTERLUND, Fauna Eur. 1876, not of Brot (Melani-
idce). Candidula KOBELT, Catalog., 1 871 , p. 22. Xerolena MONTS.,.
Moll. Terr. Isole adiacenti Sicil. 1892, p. 22 (for If. virginalis, ham-
ilcaris, ingoi). Xerotringa MONTS.. 1. c. (for H. tringa, phari, paren-
tina, meridionalis, substtiata), group Cisalpinana Fagot. Xero-
vaga MONTS., 1. e. (for H. caperata, heripensis, gigaxii, andalusicd).
Xeroalbina MONTS., t. c., p. 23 (for candidula,unifasciata, gratiosa,
striata). Xeromicra MONTS., 1. e. (for H. apicina). Xerotricha
MONTS., 1. c. (for H. conspurcata). -Xeroclavsa MONTS., t. c., p. 22
(for meda). Striatella CLESSIN, Deutsche Exc. Moll. Fauna, p. 149
1876, (for H. candidula, H. striata).
Shell rather small, depressed, narrowly umbilicated, solid and
chalky ; the surface striated ; apex corneous or dark ; whorls about
4ii, the last rather wide and rounded. Aperture round-lunate, lip
simple, strengthened by a submarginal rib within. Type H.
candidula, pi. 68, fig. 28.
254
HELICELLA.
Genital system (pi. 69, fig. 10, H. candidula). Dart sack single
and simple; mucus glands consisting of four tubes. Appendix
wanting. Flagellum very short. See also pi. 69, fig. 9, H. caperata.
In some species (pi. 69, fig. 13, H. striata) there are two dart sacks
with two accessory sacks, and about 11 mucus glands.
Distribution, Middle Europe and circum-Mediterranean region.
It is probable that two sections will be distinguished in this group,
the division to be based on the number of dart sacks ; but at present
so few spceies have been dissected that such division is not possible,
H. acutistria Bttg., iv, 10.
H. agrioica Bgt., iv, 9.
H. andalusica Kob., viii, 160.
H. apicina Lam., iv, 5.
cenisia Charp.
hispidula Risso.
cupani Calc.
v. ramburi Mab., iv, 6.
v. requieni Moq.
v. mu'hlfeldtiana Zgl.
v. citharistensis Bgt.
v. psaropsis Loc.
v. marsiana Bgt.
H. armillata Lowe, iv, 15.
lowei P. & M.
eumceus Lwe.
H. arrouxi Bgt., iv, 12.
H. bardoensis Bgt., iv, 7.
H. braidensis Poll.
H. calymnia Mts., viii, 179.
H. camerata Mouss.
H. candidula Stud., iv, 10.
unifasciata Poir.
bidentata Dkr.
graphiea Hartm.
rugellosa Hartm.
striatula Hartm.
azona Andr.
unizona Andr.
radiolata Andr.
elegans Flem.
solitaria Poir.
v. alpicola Stab., iv, 10.
v. thymorum v. Alt. iv, 10.
v. gratiosa Zgl., iv, 10.
strigatula Hartm.
adnumerata Parr,
v. spirilla West,
v. vortex West,
v. albocinctella Colb.
v. namurcensis Colb.
v. lunulata Kryn.
v. mellse Pini.
v. iriana Poll.
v. vincse Paul.
H. cantabrica Hid.
H. caperata Mont., iv, 14.
v. lauta Lwe.
v. barcinensis Bgt., iv, 14.
mirandce Ramb.
iberica Ramb.
v. diniensis Ramb.
H. carascalensis Fer.,^ 103.
H. cisalpina Jan. ^
H. cistorum Morel.
H. codia Bgt., iv, 16.
H. conspurcata Drap., iv, 12.
radiolata Jan.
cetncea Ben.
v. illuviosa Nev.
v. psara Bgt.
H. cyparissias Parr., iv, 11.
HELICELLA.
255
H. derogota Rm., iv, 23.
v. angulata Rm., iv, 23.
murcica Guir.
H. diensis Malz., viii, 162.
H. dohrni Paul., viii, 173.
H. etrusca Iss.
H. eustricta Bgt., iv, 13.
H. fedtschenkoi Mart., iii, 24,
[iv, 9.
H. geryvillensis Bgt., iv, 6.
H. gigaxii Charp., iv, 16.
H. guimeti Bgt.
H. hellenica Bl. & W., viii, 163.
v. contempta Parr.
H. heripensis Mab., viii, 158.
ruida (B.) Gout.
pouzonensis Fag.
v. solaciaca Mab., viii, 159.
H. heynemanni Kob., viii, 169,
H. illibata Parr., iii, 249.
H. improbata Mouss., iv, 12.
H. intersecta Mich., iv, 13.
iqnota Mab.
H.jaylei Pal., viii. 164.
v. rusticula Pal., iv, 14.
H. kotschyi Pfr.
H. lallemantiana Bgt., iv, 6.
H. langloisiana Bgt., iv, 15.
H. letourneuxiana B., iv, 1 2.
H. locheana Bgt., iv, 13.
H. loroglossicola Mab., viii, 159.
H. madritensis Ramb., iv, 16.
H. meda Porro, iv, 17.
subclausa Rm.
turatii Parr.
calypso Parr.
H. meridionalis Parr., iv, 9.
H. mesostena West., viii, 175.
H. metabola West.
H. modica Morel.
v. attafsensis Morel.
H. molinse Hid., iv, 15.
H. moricola Pal., iv, 13.
H. obruta Morel., iv, 6.
H. tuta Paul, viii, 173.
H. vatonniana Bgt, iv, 31.
flor entice Pons., viii, 161.
H. velascoi Hid., vi, 103.
H. ordunensis Kob., viii, 161.
H. paladilhi Bgt., iv, 11.
H. parableta Bttg., iv, 8.
H. penchinati Bgt., iv, 16.
H. perroudiana Loc., iv, 8.
H. profuga Schm., iv, 7.
phari Fagot, iii, 241.
fasciolata Moq.
fimbriata Chier.
apennina Chier.
v. attica Bttg.
v. variegata Mouss.
v. comnena Ret.
H. protea Ziegl., iv, 5.
campestris Zgl.
pustulata Miihl.
H. psiloritana Malz., viii, 162.
H. quisquilise Paul, viii, 164.
H. reboudiana Bgt., iv, 6.
H. rokniaca Bgt., iii, 198.
H. rugosiuscula Mich., iv, 11.
H. sardiniensis Villa, viii, 164.
H. schaufussi Kob.
H. semipicta Hid., iv, 16.
H. striata Mull., iv, 7.
costulosa Zgl.
narbonensis Req.
v. nilssoniana Bk.
v. furedensis Serv.
v. bakonyca Serv.
v. plattenica Serv.
H. subapicina Mouss., iv, 6.
v istera Let. & Bgt.
H. subcostulata Bgt., iv, 9.
256 HELICELLA.
H. submeridionalis B., iv, 14. H. tricastinorum Flor., iv, 9.
H. subprofuga Stab. H. tuta Paul., viii, 173.
H. substriata CL, iv, 9. H. vatonniana Bgt., iv, 31.
H. subvariegata Malz., viii, 163. florentice Pons., viii, 161.
H. trepidula Serv., viii, 171. H. velascoi Hid., vi, 103.
Insufficiently known forms: H. deana and H. pleurestha (Tassy)
Berth., H. mediolanensis Fag., H. grandiscaneusis Fag., H. apruti-
tiana Fag., H. florentina Fag., H. brundusiana Fag. H. muggiaDica
Stoss., H. tringa Fag., H. ingoi Cafic., H. lesiniaca Fag., H. par-
thenia Hag., H. rhytiphora Chemn., H. herbatica Fag., H. kryzeri-
sis Bgt., H. solanoi Serv., H. st;gila Loc., H. xalonica Serv., H.
alluvionum Serv., H. odarsensis Fag., H. montgiscardiana Fag., H.
grannonensis Bgt., H. canovasiana Serv., H. raendranoi Serv., H.
blasi Serv., H. agna Hag., H. cyzicensis Gall., H. ariantina West,
H. tremata Let. & Bgt., H. tritonidis Jon., H. fera Bgt., H. nova
Bgt., H. subneglecta Bgt, H. phthiota West., H. pastorella West,
H. curetum West, H. agreabilis Zgl, H. arcuataZgl.,H. samnitum
and v. pugnax W., H. mehediana L. & B., H. ycaunica Mab., H.
philomiphila Mab., H. vicianica Bgt., H. caturigia Paul., H. arceu-
thophila Mab., JL bardoensis Bgt., H. lecouffei L. & B., H. duvey-
rieriana Bgt, H. herbarum Serv., H. oberthuri Anc., H. incolumis
Bgt, H. codia Bgt, H. subintersecta Bgt, H. strucki Mz., H. pic-
tonum Bgt., H. olisippensis Serv., H. badigerensis Fag., H. monis-
trolensis Fag., H. idiophya Flor., H. callestha Bereng., H. tolosana
Bgt., H. groboni Bgt and v. xenilica Serv., H. lieuranensis Bgt.,.
H. margieriana Fag., H. pauli Bgt., H. valcourtiana Bgt. and v.
veranyi Bgt., H. crouziliana Fag., H. gesocribatensis Bgt, H. phil-
ora Bgt., H. thuillieri Mab. with v. nomephila Bgt, H. coutaguei
Bgt with vars. acentromphala and mauriana Bgt, H. lemesli Mab.,
H. scrupea Bgt., H. siticulosa Fag., H. diniensis Ramb., H. idanica
Loc., H. cahuzaci Bgt., H. vela viana Bgt., H. triphera Bgt, H. jean-
bernati Bgt, H. belloquadrica Mab., H. mouqueroni Bgt., H. leio-
lemma West., H. acosmia Bgt., H. microphana Bgt., H. ilicetorum
Mab., H. garocelianaLoc., H. tarasconensis B., H. simiarum Kob.,
H. alavana Bgt., H. mascarenasi B., H. culmi Fag., H. segetum
Fag., H. lunulata Kryn., H. elimberrisiana Loc., H. aurigerana
Fag., H. lugduniaca Mab., with/, stictica W., H. ussatensis B., H.
arelatensis Loc., H. lusoi Serv., H. saldubensisServ., H. pinii West.,.
H. arganica Serv., H. belernensis Serv., H. taconera Serv., H. mer-
cedesi Serv.,H. ramburi Mab., H. carcusiaca Mab., H. hypseaDu B.,
H. danieli B., H. deferiana B., H. lathraea B., H. melania B., H.
halia B., H. salivosa B., H. barcinonensis Fag., H. madritensia
HELICELLA. 257
Karnb., H. pallaresica Fag., H. salvanse Fag., H. chise Fag., H.
moreri Fag., H. subiberica Fag., H. crisia Let. & Bgt., H. zaragoz-
ensis Serv., H. campoensis Fag., H. tarifensis B., H. specialisB., H.
djebbarica B., H. warnieriana B., H. irrita Berth., H. debeauxi
West., H. micromphalus Let., H. lirouxiana B., H. madida Fag., H,
misara B., H. paladilhiformis Fag., H. romulina Serv., H. noctuella
B., H. arnusi Serv., H. edetanorum Serv., H. ambieliana (Ch.) Pal.,
H. bradybsena L. & B., H. terricola B., H. galeomma B., H. seglia
L. & B., H. argoderma B., H. briarsea B., H. aggarica B., H. eucorea
B., H. amicula B., H. amphibola B., H. ambloxa L. & B., H. anasia
B., H. goniogyraB.. H. concholeuca L. & B., H. vivida Hagenm.,
H. hipponensis Mor., H. issea Hageura., H. irana Hagenm., H. cas-
troiana Serv., H. ademata B., H. avenionensis B., H.tassyana Fag.,
H. diloricata B., H. vafella L. &B., H. propria Gall, H. augustiana
B., H. pisanorum B., H. luteola Serv., H. eucalia Hagenm., H. his-
palina Serv., H. frayssina B., H. crema B., H. hola B., H. hiero-
contina W., H. hierapetrana Mz., H. colosseana Fag., H. romana
Fag., H. fiesolensis Fag., H. membronica Berth., H. artonilla
Hagenm., H. astonara Hagenm., H. pleurabdota B., with v. cacista
B. and v. vaganensis Hag., H. perlutosa Hag., H. syntela B., H.
saharica B., H. ischurostoma B., H. nahrouasselina B., H. honorati
B., H. moricola Pal., H. tenietensis B., H. gibilmanica Serv., H.
polytrichia Anc., H. longipila Mss., H. vestita Rarub., H.dumivaga
Mouss., H. trutatiana Fag., H. renei Fag., H. oreina Fag., H. mon-
tivaga Fag., H. suborcina jFag., H. seirensis Fag., H. bradygyra
Fag., H. carascalopsis Fag., H. esserana B., H. nansoutyana B.,
H. oppidi Fag., H. transfuga Fag.
Section Monilearia Mousson. 1872.
Monilearia Mouss., Rev. Faun. Mai. Canaries, p. 39.
In referring these minutely perforate, mostly well keeled forms to
the Xerophila series, 1 am departing from ordinary usage, which
has associated them with the Maderian group Lemniscia. In general
appearance and sculpture, as well as in the structure of the basal
lip, they do not agree with Lemniscia as well as with Obelus, Cand-
idula, Jacosta, etc.
No type having been nominated for this group, I consider H. pha-
lerata Webb & Berthelot such. The species are confined to the
Canary Islands.
17
258 HELICELLA.
H. monilifera W. & B., iv, 20. H. cjementitia Sh., iv, 20.
H. lancerottensis W. & B., iii, H. tumulorum W. &B., iv, 19.
v. webbii Lwe. [237. ? atomata Mke., iv, 21.
v. bertheloti Lwe. H. oleacea. Sh., iv, 20.
v. adoptata Mouss. v. deusta Lwe.
H. persimilis Shutt., iv, 19. H. woodwardia Tarn., iv, 20.
v. prseposita Mss. H. watsoniana Woll., iv, 21.
v. devia Mouss., iv, 20. H. lemniscata W. & B., iv, 21.
H. phalerata W. & B., iv, 19. H. orbignyi W. & B., iii, 237.
rosetti W. & B. orotavana Tarnier.
nivariensis Sh., Rve. v. mitigata Mouss.
H. umbiculaSh., iv, 21. v. calcarea Mouss.
roseti Pfr. not W. & B. H. phryganophila Mab.
phalerata Pfr. not W. & B. H. dendrophila Mab., iii, 237.
H. aglaonieta Mab., iii, 237.
Section Jacosta Gray, 1821.
Jacosta GRAY, London Med. Repos., xv, March 1, 1821, p. 236,
only species mentioned H. Jacosta albella ~Drap.= explanata Miill.
Numidia ISSEL, Ann. Mus. Civ. Genov. xxii, 1885, p. 8, 9, type H.
idia Bgt. Xerofalsa MONTS., Moll. Terrest. delle Isole adiacenti
alia Sicilia, 1892, p. 21 (for H. idia, eniea, zougitana). Xerosecta
MONTS., L c. (for H. explanata). Xeroplana MONTS., 1. c. (for H.
doumeti, depressula). Xerotropis MONTS., t. c., p. 23 (for gargottce,
jolyi. prietoi, ledereri, milaschewischi, spratti). Xeroamanda MONTS.,
t. c., p. 22 (for amanda, ustieensis). Xeromoesta MONTS., L c. (for
moesta, kaby liana, dormiens, dohrni). Xerocodia MONTS., t. c. p. 23
(for montserratica, penchinati, barneana). Xeroplexa MONTS. I. c.
(for setabulensis, nyelli, coronadoi). Tropidocochlis LOCARD, Ex-
change ix, p. 97, 1893, type H. explanata.
Shell umbilicated, the whorls flattened above, acutely keeled at
periphery, and convex beneath; surface costulate, striate or smooth-
ish ; solid and earthy, whitish with or without bands, the apex black
or light. Aperture angled, the lip rather blunt, not expanded. Type
H. explanata Miill., pi. 68, figs. 23, 24.
Jaw arcuate and ribbed. Genital system (pi. 69, fig. 15, H. ex-
planata) with the flagellum rather longer than usual in the genus ;
dart sack slightly bilobed at apex and containing two darts ; mucus
glands numerous.
HELICELLA.
259
H. explanata shows clearly the transition between the species with
one and those with two dart sacks. The second sack is formed by
splitting of the first. In some other species of this keeled group the
sack is apparently single and simple, as Schuberth has figured it for
H. syrosina.
Jacosta is in all probability a purely artificial group, containing
keeled forms which have arisen from several diverse stocks of un-
keeled Helicellas; but only a thorough study of the shells and
anatomy of many species can demonstrate the true origin of the
several forms.
H. agona Anc.
H. amanda Rm.
limbata Phil,
v. dormiens Ben., iii, 252.
v. insularis Iss.
H. amphiconus Malz., viii, 180.
H. argonantula W. & B., iv, 42.
typica and canariensis Mss.
renati Dautz.
H. arianensis Bgt., iii, 253.
H. barceloi Hid., iii, 257.
H. barneyana Anc., viii, 183.
theodori Anc. ms.
H. biangulosa Mts., iii, 178.
H. boissyi Terv., iii, 254.
v. frater D. & H., iii, 258.
H. brondeli Bgt., iii, 255.
H. cardonse Hid., iii, 258.
H. caroli D. & H., iii, 258.
H. cavannae Paul, iii, 259.
v. scissa Paul.
H. cavimargo Mts.
H. cisternosi Hid., iii, 259.
H. columbina West.
H. corrugata Gmel., iii, 252.
ragosa Chem.
groyana Fer.
gargottce Phil,
v. pleurischurra Bgt.
v. chonomphala Bgt.
H. crenimargo Kryn., iii, 252.
piatigorskiensis Bayer.
H. depressula Parr., iii, 256.
forms globulosa, flammulata,
zonata, fulva, cpmpressa Bgt.,
exserta, murustagensis West.
H. doumeti Bgt.
v. lacosteana Mor., iii, 255.
H. eminens West., viii, 180.
syrensis v. exserta Mts.
H. enica L. & B., viii, 182.
H. eugoniostoma Bgt.
H. explanata Miill.. iii, 255.
albella Dr.
/. minor Bgt.
/. subscalaris Bgt.
H. filimargo (Z.) Rm., iii, 251.
taurica Partch.
cliersonesica Mu'hl.
H. finitima Mor., iii, 241.
H. gradilis Mts., viii, 179.
H. graja West.
v. philesia West.
H. granostriata Mouss., iv, 43.
H. grovesiana Paul, iii, 254.
H. hamudie Kob., viii, 182.
H. hariotiana Bgt.
H. henoniana Bgt., iii, 254.
v. agriuneusis Kob.
H. homeyeri D. & H., iii, 257.
260 HELICELLA.
H. idia L. & B., iii, 256. H. rozeti Mich., iii, 254.
H. ledereri Pfr., iii, 259. v. oxygyra West.
/. regularis Roth. H. rozetopsis L. & B.
H. moraguesi Kob., iii, 255. H. setabulensis Pfr., iii, 256.
H. micropristis Anc. serrula Morel.
/. appressispira Anc. H. siderensis Malz., viii, 181.
H. milaschewitschi Ret. H. sigensis Kob., iii, 256.
H. morata Mouss., iv, 43. /. jolyi Pech., Anc.
H. montserratensis Hid., iii, 257. H. siphnica Kob.
H. multipunctata Mouss., iv, 43. H. spratti Pfr., iii, 253.
H. nummuliformis Ret. siderites Friv.
H. n^elii Mitt., iii, 258. v. perplanata Pils., viii, 181,
H. orophea West. H. syrensis Pfr., iv, 28.
H. parableta Bttg. H. syrosina Bgt.
H. pollenzeusis Hid., iii, 257. H. tineana Ben., iii, 253.
H. ponsii Hid., iii, 257. v. raista West.
H. ponsonbyi Kob., viii, 183. v. kobeltiana W.
H. prseclara Caf., iii, 252. H. tissotiana Bgt.
H. prietoi Hid., iii, 257. H. tristrami Pfr., iii, 253.
H. pulverulenta Lwe., iv, 43. H. usticensis Calc., iii, 254.
H. retowskii Cless., iii, 252. H. zeugitana L. & B., viii, 182.
H. rouvieriana Bgt., iii, 255.
Insufficiently knoivn forms of Jacosta : H. ehola Pech., caficii (Ad.)
West, hierica Bgt., tineiformis Let. & Bgt., soleilleti Bgt., hodnse
Anc., melosina Bgt., sageti Bgt., orphea West., morini, chthamalo-
lena, ablennia, hyperconica and tellica Bgt., eufidana callistoderma
and conicula Let. & Bgt., mitidjana and bibanensis Anc., eup-
hacodes and sphakiota Malz.
Section Xeroleuca Kobelt, 1877.
Xeroleuca KOB., Jahrb. D. M. Ges. 1877, p. 25. CW/.Schepman,
Jahrb. 1877, p. 271, 272, anat. of mograbina and degenerans.
Shell depressed and widely umbilicated, solid, whitish and chalky,
having one, two or three spiral keels ; surface roughly sculptured.
Type H. turcica Chemn.
The species are mostly from Morocco. Genital system (pi. 69,
fig. 11, H. mograbina) substantially as in Helicellaericetorum, but
the dart sacks are smaller.
HELICELLA. 261
H. couopsis Mor., iv, 22. H. mogadorensis Bgt., iv, 22.
H. turcica Chemn., iv. 22. H. cyelostremoides Sby., iv, 256.
cratera Schum. H. mograbina Mor., iv, 22.
v. tetragona Mor., iv, 21. H. darolli (Let.) Bgt.
H. degenerans Mouss., iv, 22. v. djarica Bgt.
/. validior Mouss. H. tunetana Pfr., iv, 21.
Section Obelus Hartmann.
Obelus HARTM., Erd- und Siisswasser-Gast. Schweiz, p. 158, type
O.preauxii. Xeroptyca MONTS., Moll. Terr. Is. adi. alia Sicil.,
1892, p. 25, type H. " ptycodia "=ptychodia.
Shell trochiform, with narrow umbilicus, acutely keeled and serrate
periphery (at least when young), with usually a secondary keel or
series of tubercles midway between periphery and suture ; solid,
whitish and earthy. Aperture basal, angular-oval, the lip simple
but slightly dilated at columella. Type H. despreauxii.
Genital system (pi. 69, fig. 12, H. tuberculosa, typical) ; penis
having a subterminal elongated appendix; other organs as in Hel-
icella ericetorum, but dart sacks small.
Distribution, Canary Is., eastward to Syria. A very natural
group, consisting of one series of species in Morocco and the Canary
Is. and another in Lybia to Palestine. Monterosato's name Xero-
ptyca belongs to the latter, but it seems unnecessary to make any
separation, and the term is etymologically bad.
H. tuberculosa Conr., iv, 25 ; H. pumilio Pfr., iv, 27.
[viii, 184. v. cyclodon W. & B., iv, 26.
v. serrulata Bk., iv, 25. H. despreauxii Orb., iv, 25.
H. philammia Bgt., viii, 185. preauxii Hartm.
H. ptychodia Bgt., viii, 184. v. immodica Mouss.
H. agenora West. H. moderata Mouss., iv, 26.
H. berenice Kob., viii, 185. H. mirandse Lwe., iv, 2b'.
H. lybica Pons., viii, 185. nodosostriata Mouss.
H. hesperidum Mor., iv, 26.
Section Trochula Schliiter, 1838.
Turricula BECK, Index Moll. 1837, p. 10, not Turricula Schum.
Essai, 1817, p. 217 (Pleurotomidce*). ? Oxynota HARTM., Erd- und
Siisswasser-Gast. Schweiz, 1842, p. 159, a nude name ; no species
mentioned. Crenea ALBERS, Die Hel. 1850, p. 77, in part, not
262 HELICELLA.
Crenea Kisso, 1826. ? Trochoida or Trochoidea BROWN, 111. Conch.
G. B. 1827 (publication not seen). Xeroclivia MONTS., Moll. Terr,
Is. adiac. alia Sicil., 1892, p. 25 (for H. pyramidata). Xeronexa
MONTS., /. c. (for H. cumice, calcaratd). Xerocochlea MONTS., /. c.
(for H. caroni, elata). Trochula SOHLTJTER, Syst. Verz., p. 7, 1838,
MORCH, Journ. Conch. 1865, p. 386.
Shell solid and earthy, narrowly umbilicated, trochoidal, with
conic spire, narrow whorls and rather flattened base; periphery
somewhat angular or sharply keeled. Aperture small, lunate or
angular, the lip not expanded, strengthened by a submarginal rib
within. Type H. terrestris Penn., pi. 68, fig. 27.
Jaw with 8 to 18 close, flat ribs (pi. 67, fig. 12, H. terrestris).
Radula (pi. 67, fig. 16 H. terrestris) with mesocone about the
length of basal-plate on median teeth, the side cutting points well
developed. Laterals the same, but lacking entocones ; on the transi-
tion teeth the inner cusps become bifid, and on marginals the ecto-
cone also splits.
Genitalia (pi. 69, fig. 18 H. elegans) ; dart sacks two, very smalL
Mucus glands 6. At base of vagina or on the atrium a large sack-
like appendicula. Other features as in Helicella generally.
Mainly a circum-Mediterranean group, distinguished from Obelus
by the simpler sculpture of the shell and the characters of genitalia
mentioned above.
The name proposed by Brown in 1827, may prove to have first
claim for this group, but I have not seen the original publication
and it is variously spelled by the authors who have quoted it. In
any case, Turricula must be rejected from the nomenclature of Hel-
ices, being preoccupied in Mollusca.
H. newka Dohrn, iv, 29. H. terrestris Penn., iv, 29.
H. newkopsis L. & B. elegans Gmel.
H. majoricensis D. & H., iii, 258. trochulus Hartm.
H. miscella West. H. scitula C. & J., iv, 29.
H. caroni Dh., iv, 29. trochilus Poir.
v. pyramis Phil., iv, 30. depressa Bk.
H. elata F.-B., iv, 29. perdepressa West.
v. dilatata Ben. H. trochoides Poir., iv, 27.
H. trochlea Pfr., iv, 30. algira Chier.
H. sequentiana Ben., iv, 30. solarium Risso.
HELICELLA. 263
v. pyramidella Jan. sabulosa Zgl.
v. sulculata C. & J. spectabilis Zgl.
rugosiuscula Mich. pyramidatoides Orb.
v. vidua West. v. nova Paul,
v. infulata Paul. v. depressa Bgt.
v. conica Drap. v. tarentina Pfr., iv, 24.
H. turritella Parr., iv, 27. H. apiculus Rm., iv, 29.
/. remissa Parr. cumice Calc.
H. verticillata Parr., iv, 27. H. numidica Moq., iv, 24.
H. liebtruti Alb., iv, 28. v. sulliottii Poll.
H. idalise Bgt., iv, 28. H. calcarata Ben., iv, 28.
H. pyramidata Drp., iv, 23. H. schembrii Scac., iv, 28.
agnata Zgl. sehombrii err. orig.
arenaria Zgl. H. cucullus Mts., iv, 28.
littoralis Zgl. H. inops Mouss., iv, 26.
Insufficiently known species : H. crenulata MiilL, H. licodiensis
Cafici, H. pupilla Serv., H. eupyramis, spaellina, kelibiana, spsella,
zitoumica, madana, galactina, veneriana Let. & Bgt, H. tisemsinica,
mactanica, capuana, dyrrachiensis Bgt, H. apiculiformis Anc., H.
subnumidica (Bgt.) West., H. ogygiaca West. See also iv, p. 30.
Section Cochlicella Risso, 1826.
Cochlicella (Fer.) Risso, Hist. Nat. Eur., Merid. iv, p. 77.
MARTENS, in Alb., Die Hel., p. 117. Elisma LEACH, in Turton,Man.
L. and Fr.-W. Sh. Brit. Is., p. 84, 1831. Xeroacuta MONTS., Moll.
Terr. Is. adi. Sicil., 1892, p. 25, proposed for H. acuta, ventrosa. For
anat. see MOQ.-TAND., Moll. Fr., FISCHER, Journ. de Conch., 1856,
p. 121, SCHMIDT, Stylom., p. 41, Moss & PAULDEN, Manchester
Mic. Soc. Trans. 1892, p. 75.
Shell perforated, with elongate, turrited spire, higher than wide ;
opaque and white, usually streaked or banded with brown or waxen ;
whorls rounded, at least the last one. Aperture oval, the lip simple
and acute, expanded toward the columellar insertion. Type H.
barbara L., pi. 68, fig. 29.
Genitalia : penis of the usual form, the retractor inserted in an
obliquely truncated, cylindrical calcareous body at its apex (pi. 69
fig. 21), in which the long epiphallus terminates ; flagellum short.
Vagina without appendages ; but on the atrium there is a long or-
gan of unknown function, either a degenerate dart sack or an " ap-
264 HELICELLA.
pendix" (pi. 69, figs. 19, 20, 21, H. acuta ; fig. 21 upper part of
penis outlined, through which is seen the calcareous ring in which
the epiphallus ends, with attached retractor). The spermatophore
(fig. 20, x 8) is very long, oval in section, chitinoid in substance,
with an elevated closely serrate edge. In ventricosa and conoidea
mucus glands and a small dart sack are present.
Distribution : middle and southern Europe, northern Africa, etc.
H. ventricosa introduced into Bermuda and very abundant there.
The absence of mucus glands in H. barbara is remarkable, and
doubtless the result of degeneration. The spermatophore is similar
to that of Leptaxis which also has a similar boot-shaped sperma-
theca.
H. barbara L., iv, 32. H. ventricosa Drap., iv, 32.
acuta Mull. ventrosa Auct.
meridionalis Risso. bulimoides Moq.
fasciatus Penn. H. pringi Pfr., iv, 32.
v. terveriana Mouss. H. bellucciana Bgt.
H. conoidea Drap., iv, 31. H. duplicata Mouss., iv, 31.
fibula Wood. H. contermina Sh., iv, 31.
turbida Kiist. psammoica Morel.
solitaria Pfr. H. psammsecia Bgt.
v. calaritana Paul. H. psammsecella Let. & Bgt.
Subgenus THEBA Risso, 1826.
Theba (Leach MS.) Risso, Hist. Nat. Eur. Merid. iv, p. 73, in
part. WESTERLUND, Fauna, p. 71, in part. Teba Leach, in TUR-
TON, Man. Land and Fresh-water Sh. Brit. Is., 1831, p. 36 (in part).
Zenobia GRAY, Lond. Med. Repos., xv, March, 1821, p. 239 (for
H. Zenobia corrugata, undescr. and unfig., and H. Zenobia " binar-
ginata "=H. carthusianella Drap.). Not Zenobia OKEN, 1815 (Lep-
idoptera). Euomphalia WESTERLUND, Fauna .Paliiarct Binnen-
oonch., Helix, pp. 31, 92 (1889). Carthusiana KOBELT, Catalog
derim Europ. Faunengebiet lebenden Binnenconchyl. p. 11, 1871.
See for anatomy, HESSE, Jahrb. D. M. Ges. 1884, p. 234, pi. 5.
SCHMIDT, Stylom., pi. 7. MOQ.-TAND., Moll. Fr. pi. 16. ASH-
FORD, Journ. Conch. Leeds, iv, pi. 10. SCHUBERTH, Arch. Naturg-
1892, pi. 2. v. IHERING, Morphol. u. Syst., p. 440.
Shell depressed subglobose, narrowly umbilicate or imperforate,
whitish or banded with white, but rather thin and somewhat translu-
HELICELLA. 265
cent, the surface finely malleated, shining ; last whorl wide, slightly
descending. Aperture wide-lunar, but little oblique ; lip acute,
slightly expanded below, strengthened within by a distinct submar-
ginal rib. Type H. carthusiana Mull., pi. 68, figs. 25, 26.
Jaw with numerous flat, close ribs. Marginal teeth with an-ab-
normally large number of denticles, at least in some species. Penis
short and swollen, passing into an epiphallus which ends in a short
flagellum and the vas deferens ; no retractor muscle. Mucus glands
inserted high on vagina, at root of spermatheca duct, and consisting
of three pairs of tubes. Far below them is a long blind sack with
plicate internal walls, but containing no dart, evidently a degenerate
dart sack. Spermatheca irregular-oblong, its duct long and branch-
less. Right eye retractor not passing between branches of genitalia
(pi. 69, fig. 22, H. carthusiana. PI. 69, fig. 16, H. syriaca. PI. 69,
fig. 14, H. cantiand). The arrangement and number of mucus
glands varies in the different species.
Distribution, middle and southern Europe and Asia Minor.
The group is well distinguished by the whitish but still somewhat
translucent, finely malleated shell, with conspicuous lip-rib, as well
as by the lack of penis retractor muscle (cf. Cepolis) and the lack
of a dart in the lengthened but evidently degenerate dart sack.
Risso's genus Theba contained ten species of several groups, but
after eliminating the first three and last two, which were removed
by subsequent authors to other groups, Euparypha, Cochlicella, etc.,
we have left five forms of the If. carthusiana type. Westerlund
lias made these the basis of Theba; and it seems better to follow
his example rather than to legislate the group completely out of ex-
istence as we would be compelled to do were we to adopt Kobelt's
name Carthusiana, proposed in 1871. The latter name is, more-
over, objectionable from being the duplication of a specific name, al-
though this alone would not bar its acceptance. Gray's Zenobia is
founded upon a species of this group, but is preoccupied. It has
been used hitherto in a totally incorrect manner by European
"writers, evidently from failure to consult the original paper in
which it was proposed. Westerlund's group Euomphalia is founded
on H. strigella and its allies, which are anatomically like Theba.
H. alphabucelliana Paul., iii, 204. dacampi Villa.
H. anconse Iss., iii, 192. /. carfaniensis Stef.
oliviaria, Iss., olim. v. marchetti Stef.
rubella Risso. v. simplicita Parr., iii, 194.
UNIVEBSIIY
266
HELICELLA.
H. apennina Miihl, iii, 202.
H. arpatschaiana Mas., iii, 177.
v. pseudoglobula Mss., iii, 197.
H. berytensis Fer., iii, 194.
v. rachiodia Bgt., iii, 194.
granulata Roth.
v. fourousi Bgt., iii, 194.
H. cantiana Mont., iii, 192.
pallida Jeffr.
/. cantianiformis Bgt.
H. carthusiana Miill., iii, 195.
arenaria Oliv.
olivieri C. Pfr.
binarginata Gray.
gibsi Leach.
/. carthusianella Dr.
rufilabris Jeffr.
innoxia Bgt.
/. fasciata West.
/. leucoloma Stab.
/. claustralis Parr.
v. radiata West.
v. archimedea Ben.
v. ventiensis (B.) Fag.
v. diurna Bgt., iii, 193.
v. leptomphala Bgt.
v. encyse Serv.
v. euscepia Serv.
v. conoidea Branc,
v. glabella Drap., iii, 186.
v. sarriensis Pena, iii, 193.
v. episema (B.) Serv.
v. lamalouensis Reyn.
taurinensis Pini.
/. arvensis Pini.
H. cemenelea Risso, iii, 193.
galloprovincialis Dup.
vars. campanica Paul, oustera
Mab., monerebia, gaude-
froyi, abebaia, apuana Mab.,
riparia Bl., ardesa, sobara,
iadola Bgt., putoniana
(Mab.), Loc., iii, 194, rees-
manni Cless., delacuri Mab~
=delacourti Mab., Bgt.
H. flaveola Kryu.
H. flavolimbata Bttg., iii, 201.
H. globula Kryn., iii, 197.
v. nana Bttg.
H. helvola Friv., iii, 202.
v. martensi West.
H. hirci Cless.
H. holotricha Bttg.
H. martensiana Tib., iii, 203.
lavata Tib.
H. mnenia West.
H. obstructa Fe>., iii, 196.
obstrusa Fer.
/. dilatata West.
/. adpressula Friv.
H. olivieri Fe>., iii, 191.
v. parumcincta Parr,
v. bicincta Ben.
rizzce Arad.
v. cribrata West,
v. gregaria Zgl., iii, 196.
occulta Biv.
H. orsinii (Porro) Villa, iii, 203.
ochracea Zieg.
v. majellse Kob.
v. picena (Tib.) Kob.
H. pachnodes Bttg.
H. pantanellii Stef.
H. parreyssi Pfr., iii, 203.
modesta Parr.
H. phseozona Mts., iii, 205.
H. pisiformis Pfr., iii, 197.
v. atypa Bttg., viii, 187.
H. rothi Pfr., iii, 197.
/. inversa West.
/. draxleri Zel.
H. rubens Mts., iii, 205.
HELICELLA.
26T
/. concolor Mts.
/. finschiana Mts.
/. zeiliana Mts.
/. regeliana Mts.
H. rufispira Mts., iii, 204.
v. albidorsalis Mouss.
H. samsunensis Zel.
H. schotti (Zel.) Pfr.
H. schrenki Midd., iii, 200.
siberica Friv.
H. semenowi Mts.
/. depressa Mouss.
H. septemgyrata Mss., iii, 201.
H. strigella Drap., iii, 202.
sylvestris Alt.
altenana Gartn.
fruticosa Parr.
cornea Hartm.
piligera Andr.
vitrinosa Zgl.
hexagyra Miihl.
peregra Parr,
v. colliniana, lepidophora, rus-
inica, separica, vellavorum,
ceyssoni, buxetorum, neme-
tuna, cussetiensis, mehadise,.
agapeta Bgt., briandi, du-
breili Serv.
H. subobstructa Bgt., iii, 196.
v. distypa West.
H. suborbicularis Mts., iii, 203.
H. syriaca Ehr., iii, 197.
ony china Rm., f. 568.
H. talyschaDa Mts., iii, 195.
H. theobaldi West.
H. transcaspia Bttg., viii, 187.
Section Lejeania Ancey, 1887.
Lejeania ANC., Conchol. Exch. i, p. 75, June, 1887, types H. dar-
naudi Pfr., isseliana Morel., jickeliana Nev. Pella, in part, of
authors.
Shell narrowly umbilicated, depressed-globose, thin, with the tex-
ture of Fruticicola ; broivn with opaque whitish bands, or opaque-
white with dark bands. Lip thin, simple, expanded toward the um-
bilical insertion. Type H. darnaudi Pfr.
Jaw high-arched, with numerous flattened, wide ribs, hardly den-
ticulating the margin (pi. 71, fig. 43, H. scioana). Radula having
the middle teeth tricuspid, median cusp shorter than basal-plate.
Laterals bicuspid ; marginals with the ectocone split (pi. 71, fig. 42,
JET. scioana).
Genital system (pi. 71, fig. 44, H. lejeaniana) with no retractor
on penis, flagellum long. Spermatheca on a short duct. Dart
sack short, swollen, inserted on atrium ; mucus glands in two groups
of 3 or 4 each (lejeaniana\ or wanting apparently (scioana).
Distribution, Abyssinia, southern Arabia. My knowledge of the
anatomy of this group is from Pollonera's work Boll. Soc. Mai. Ital.
xiii, p. 75, pi. 3. The position of the right eye-retractor is unknown.
268 HELICELLA.
No penis retractor is shown in Polloneras's figures or mentioned by
him, so I suppose it is absent ; and it is mainly on this ground that
I place the group next to Theba. The hairy species may belong
elsewhere, but judging from the anatomy of scioana, they are better
placed here. The shell diagnosis given above applies to the typical
forms.
H. isseli Morel., iii, 105. H. jickelii Nev., iii, 230.
darnaudi Jick. part. H. pilifera Mart., iii, 190.
H. lejeaniana Bgt., iii, 104. H. combesiana Bgt, iii, 190.
darnaudi Jick. part. pilifera Jick.
H. achilli Bgt, iii, 105. H. ferretiana Bgt., iii, 190.
H. darnaudi Pfr., iii, 104. H. herbini Bgt., iii, 190.
v. heuglini Mts., iii, 104. H. galinieriana Bgt., iii, 190.
H. hamacenica Raff., iii, 250. H. beccarii Jick., iii, 189.
H. subnivellina Bgt., iii, 250. ciliata Morel.
H. nivellina Bgt. H. scioana Poll., viii, 190.
alexandrina Parr., undescr. H. d'hericourtiana B., iii, 104.
nivea Zgl. not Gmel. H. strigelloides Poll., viii, 190.
H. leucosticta Mts., viii, 190.
Section Platytheba Pilsbry, 1894.
Nummulina KOB., Catalog Eur. Binnenconch., p. 12, 1871. Not
Nummulina d'Orbigny, 1826 (Polyzoa).
Shell narrowly umbilicated, depressed and lens-shaped, acutely
keeled ; thin but rather solid, costulate striate, whitish. Aperture
quite oblique, transverse, and angular outside; peristome simple
above, the basal lip slightly thickened within and dilated at inser-
tion. Type H. nummus Ehr., pi. 68, fig. 30.
Genital system (pi. 69, fig. 17, H. nummus) as in Theba, the
penis lacking retractor, flagellum very short, etc.
These are simply keeled Carthusianas, as von Iheringhas already
maintained, but the shell features demand recognition by name.
The group is characteristic of Syria and the Caucasus region. It
has no alliance with Plectotropis, although the shells are somewhat
similar.
H. nummus Ehr., iii, 199. H. promethus Bttg., iii, 199.
hedenborgi Pfr. H. genezarethana Mouss., iii, 199.
oxygyra Boiss. tiberana Mouss.
H. spiroxia Bgt., iii, 199. H. jasonis Dub., iii, 199.
HYGROMIA. 269
Genus HYGKOMIA Risso, 1826.
=Hygromia Risso, 1826, -\-Bradybcena Beck, 1837 (part),-f Fru~
ticicola Held, 1837,-f MmacAaFitz., 1833,4-THc/waHartm., 1841?
+Petasia Beck, 1837,+Metodonta Mlldff., 1886, etc., etc.
Shell rather thin and subtranslucent, with little calcareous sub-
stance, brown or whitish, unicolored or with a peripheral white
zone, frequently hairy. Umbilicus open or minute ; form globose-
depressed, with convex or conoid spire, and rounded or angular peri-
phery. Aperture lunate, the lip acute, expanded below, usually
thickened within, the basal margin rarely 1 or 2 toothed. Tvpe
H. dnetella (See pi. 55, figs. 20 to 30).
Jaw arched, thin, with delicate low riblets which denticulate the
margins but feebly (pi. 70, figs. 31, 39). Radula as usual in ground-
snails. Median cusps long and acute, the side cusps usually devel-
oped though small on middle teeth. Ectocones well developed on
lateral teeth. Marginals with long simple or bifid inner and small:
simple or split outer cusp.
Genital system (pi. 70, figs. 30-41) ; penis continued in an epiph-
allus which bears the retractor and ends in a short flagellum and
the vas deferens. Dart sack single or repeated, with or without
accessory sacks, the contained dart or darts cylindrical below with
short blades at apex. Mucus glands inserted on vagina above the
dart sack, consisting of several independently inserted or grouped
tubes. Right eye-retractor passing between branches of genitalia-
Dart apparatus sometimes entirely lacking by degeneration.
Distribution, Europe, North Africa and Western Asia.
The genus is not very fully represented in the fossil series as now
known, although a moderate number of forms are found extending
as far down as the Oligocene of middle Europe. I do not know
that any Eocene or earlier species can be referred with certainty ta
the group, but it is not unlikely.
The prominent features of this genus are (1) the thin dull-colored
shell, in which calcareous matter is never predominent, a hairy
coat is often developed, and the lip is not reflexed ; (2) the thin
jaw with many slight riblets ; (3) the normal disposition of the right
eye-retractor, the short flagellum, frequent doubling of the dart sack
and the separation of mucus glands from the latter. These features
separate the genus from Helicella, which is allied in form of the
270 HYGROMIA.
shell and of the genital organs, and from Eulota which has a shell
of much the same form and texture.
In certain forms (H. revelata, H. tiliata, and the section Meta-
Jruticicola) the dart sack and mucus glands are absent ; but as there
are other species showing the gradual steps of this loss, first in the
empty condition of the dart sacks, then their disappearance, and
finally the gradual disappearance of the mucus glands also, we are
compelled to consider these simplified species as degenerate and
secondarily simple lines of descent It is noteworthy that in shell,
jaw and teeth they retain the normal structure of the genus, as well
as in the structure of the penis.
The presence of these forms lacking the cardinal features of the
Belogona might be construed by some as invalidating the premises
upon which the primary groups of Helices are founded ; but this
would be a very short-sighted view. The facts simply show that in
some members of highly organized groups, retrogressive evolution
has taken place, resulting in structures similar to those character-
istic of lower groups. This is a very common phenomenon in many
orders of animals. In the case under discussion, the organs of mas-
tication and the shell have undergone no changes, and the penis and
its appendages retain their normal characters. Compare v. Ihering,
Morphol. u. Syst., p. 450, who supports this view.
In regard to the nomenclature adopted for the group, I have
simply made the changes from current usage demanded by the law
of priority. It is absurd to continue to use " Fruticicola " in a
generic sense when it is everywhere acknowledged that Hygromia is
a dozen years earlier, and is properly diagnosed, etc., in a work
known to and used by all systematic conchologists. Fruticicola is
later than Monacha (type incarnata) and on a par with Bradybcena.
The names Zenobia, Petasia, Trochiscus, Latonia and Triehia are
clearly preoccupied, and can, therefore, have no place in the no-
menclature of Helices. They have hitherto been used in ignorance
of this fact, or in defiance of it.
The sectional scheme proposed below is remodelled from current
European usage, except that Theba and allied groups have been re-
moved to Helicella, as advocated by von Ihering, and the preoccupied
names are dropped. The species herein referred to sections Mon-
acha and Fruticicola require much investigation, and doubtless con-
siderable re-arrangement, and some students may consider it best to
split them into more sections. The other sectional divisions agree
HYGROMIA. 271
in essential points with recent European authorities, and seem to be
well-founded.
Section Hygromia Risso, s. sir.
Shell subconic, narrowly perforated, with convex-conic spire,-
keeled periphery and convex base ; horn-colored or brown, somewhat
translucent, with an opaque white peripheral girdle. Surface
smoothish, not hairy. Aperture oblique, lip expanded below, re-
flexed at columellar insertion, thickened within. Type H. einetella,
pi. 55, figs. 20, 21.
Jaw arcuate, with many wide, flat riblets (pi. 70, fig. 39, H. cine-
tella). Genitalia (pi. 70, fig. 32 H. limbata), having the epiphallus
long, flagellum very small. Four mucus tubes on each side, and be-
low them one small dart sack containing a dart. Duct of sperma-
theca long.
Species few, confined to southern-central Europe.
H. einetella Dr., iii, 18j H. limbata Dr., iii, 189.
? ranzani Orsini. /. sublimbata Bgt.
/. fasciata Paul. /. odeca Bgt.
/. chelydea West. /. hylonomia Bgt.
H. tassyi Bgt. v. delomphala Anc.
Section Monacha Fitzinger, 1833.
Monacha FITZ., I. c., for incarnata only. Not Monachus Kaup,
1829 (Aves). Latonia WESTERLUND (in part), Fauna Palaarct.
Binnenconch., Helix, pp. 30, 68. Not Latonia Mey., 1843 (Repti-
lia), nor Latona Schum. 1817 (Moll.) nor of Strauss, 1817 (Crust.).
Shell covered-perforate or narrowly umbilicated, depressed sub-
globose with 6-7 whorls, the last rounded or subangular ; surface
minutely scaly or hairy. Aperture oblique, widely sublunate, lip
expanded, well thickened within. Type H. incarnata Mull., pi. 55,
figs. 29, 30.
Genitalia (pi. 70, fig. 34 H. incarnata) as in Hygromia s. str ;
penis as usual in the genus ; 4 or 5 mucus tubes on each side ; dart
sack single.
This group is purely conventional, and is retained to contain a
series of species distributed by some authors in Eulota, Carthusiana
and Latonia. Part of the Fruticicolas have the same anatomical
features. See v. Ihering, Morphol. u. Syst. Helix, p. 449, and the
authorities there cited, for characters the soft parts.
272
HYGROMIA.
H. acorta L. & B.
H. andria Mts., viii, 186.
H. aristata Kryn., iii, 201.
H. bidinensis Caf., iii, 189.
v. daphnica Platania.
/. flavida Plat.
H. bifaria West.
H. brigantina Meng.,'iii, 204.
H. caidis Anc.
H. capusi Villes.
H. carascaloides Bgt., iii, 193.
H. catoleia Bgt.
H. cheffiana Bgt.
H. circassica Mss., iii, 195.
colchica Bayer.
H. consona Zgl., iii, 188.
/. panda West.
H. cruzyi Bgt,, iii, 198.
H. dasilepida Bgt.
H. densecostulata Ket.
H. euages Bttg., iii, 201.
/. depressa Bttg.
H. euboeica Kob.
H. faidherbiana Bgt., iii, 189.
H. frequens Mouss., iii, 193.
v. obscura Mouss.
H. freytagi Malz.
H. fruticola Kryn., iii, 200.
v. bourguignati Pfr.
H. grelloisi Bgt.
H. hausknechti Bttg., iii, 193.
H. hiberna Ben., iii, 188.
H. incarnata MiilL, iii, 187.
sylvestris Hartm.
rubra Chier.
/. pallid u la Moq., veprium
[Bgt., silanica Bgt..
v. tecta Zgl., iii, 187.
obtecta West.
v. monodon Villa, iii, 188.
armata Stab.
v. welebitana St.
v. byssina Gredl.
v. juriniana Bgt., iii, 188.
H. inchoata Morel., iii, 200.
H. lenabarica Let.
H. lepidolena Bgt.
H. lurida Zgl.
H. messenica Bl. & W.
v. acaica West.
H. musicola Bgt.
H. nicaisiana Let.
H. nicosiana Mss., iii, 189.
/. pallida Mouss.
H. ovularis Bgt.
H. pirajnea Ben., iii, 192.
H. prserupta West.
H. proclivis Mts., viii, 187.
H. pseudosericea Ben., iii, 196.
H. redtenbacheri Zel., iii, 189.
H. rissoana Pfr., iii, 195.
v. dirphica Blanc., iii, 192.
v. langei Bgt.
H. rusicadensis Let., viii, 188.
H. schuberti Koth, iii, 195.
v. frutis Parr.
H. semirugosa Kob., viii, 188.
H. vicina Rm., iii, 188.
carpatica Friv.
H. villse Desh., iii, 198.
H. zonitomsea Let., viii, 189.
Section Fruticicola Held., 1837 (restricted).
Fruticicola HELD, Isis, 1837, p. 914 (in part). v. MART., Die
Hel., 1860, p. 103, type H. hispida. Not Fruticicola MacGill.,
HYGROMIA. 273
1839 (Aves). " Fruticola" of some writers. Bradybcena BECK,
Index Moll. 1837, p. 18. Not Bradybcenus Dej., 1829 (Coleoptera).
Trichia HARTMANN Erd- und Suss wasser- Moll. Schweiz, p. 41 (for
H. hispida, etc.). Not Trichia de HAAN, Fauna Japonica, Crus-
tacea, p. 109(1840!).
Shell depressed, with convex spire, rounded periphery, and open
or narrow umbilicus ; brown or greenish, the surface generally
hairy. Aperture lunate, slightly oblique, the ends of the thin lip ap-
proaching ; peristome simple, expanded only at the columellar inser-
tion, and with only a weak internal thickening or none. Type If.
hispida L., pi. 55, figs. 27, 28.
Jaw delicate, with 10-18 low[riblets. Genital system (pi. 70, fig.
33, H. hispida) ; penis as usual in the genus ; mucus glands several ;
two dart sacks, each with an accessory sack.
Hartmann's name Trichia has been used for this group, but its
date is uncertain (his book having appeared in parts, from 1840 to
1844), while Trichia de Haan is known to bear date 1840. More-
over it seems best to use H eld's earlier term Fruticicola, which is
well known to all malacologists.
The species are very numerous throughout the European area,
and several have been imported by commerce into America, etc. A
few Chinese species of unknown anatomy are best referred here
until their true position can be ascertained.
H. aclerochroa Bgt., iii. 181. H. clessini Ulic.
H. alsia Bgt, iii, 177. H. Corsica Sh., iii, 180.
H. becasis Ramb., iii, 176. H. crispulata Mouss.
H. bourniana Bgt. H. cynetarum Malz.
H. cselata Stud., iii, 175. H. dieckmanni Mss., iii, 179.
v. coelomphala Loc. H. dussertiana Bgt., iii, 177.
v. vagienna Poll. H. erjaveci Brus.
v. cselatina Loc. /. mortella, tanora, avarica,
H. cedretorum Deb., iii, 179. savinella Serv.
H. chnoodia Bgt., iii, 179. H. filicina Schm., iii, 176.
H. chonomphala Bgt. v. nudata West.
ripularum Lessona. H. fusca Mont., iii, 186.
H. chrysotricha Bttg., viii, 190. corrugata Gray.
H. clandestina Hartm., iii, 175. subrufescens Mill.
gratianopolitana Ramb., iii, H. granulata Alder, iii, 178.
v. isarica Loc. [175. globularis JefFr.
18
274
HYGROMIA.
H. hispida L., iii, 172.
prevostiana Risso.
v. gyrata West, iii, 173.
v. concinna Jeffr., iii, 173.
v. nebulata Mke.
v. septentrionalis Cl.
v. depilata Aid.
v. conica Jeffr.
v. hemisphserica Less.
v. beaudouini Loc.
v. laticensis Loc.
v. morcbii West.
v. hispidosa Mouss., iii, 172.
v. nana Jeffr., iii, 173.
H. kusmici Cless.
H. langsdorffi Mill.
H. lanuginosa Boiss., iii, 180.
flava Terver.
H. lasia Bgt, iii ; 179.
H. latiniacensis Loc.
H. martorelli Bgt., iii, 179.
H. matronica Mab.
H. melaspinse Bgt., iii, 180.
H. mendicaria Pfr.
H. inesoleuca Mts.
H. mongrandiana Bgt.
H. montana Stud., iii, 175.
circinnata Stud.
erecta Hartm.
v. dubisiana Gout.
v. danubialis Cless.
H. rnontivaga West.
salmurina Serv., iii, 181.
H. moquiniana Raym., iii, 181.
/. fradiniana Bgt.
H. multigranosa Mouss., iv, 36.
H. nordenskioldi W., iii, 201.
rufeseens Schrenk.
H. parlatoris Biv., iii, 179.
H. perlevis Sh., iii, 181.
H. plebeia Drap., iii, 174.
/. plebicola Loc.
H. ponsonbyana Pils,, viii, 190.
ponsonbyi West, not Kob.
H. psaturochseta Bgt, iii, 182.
H. ptylota Bgt, iii, 181.
H. reinaB Ben., iii, 187.
H. revelata Fer., iii, 180.
ponentina Dup.
martigena Fer.
/. conimbricensis Silv., veneto-
rum Bgt., nevesiana Silv.,
villula Bgt, platylasia Bgt.
v. occidentalis Reel., iii, 180.
lisbonensis Pfr.
H. roseotincta Fbs.
H. rubiginosa Schm., iii, 178.
v. epirotica Mouss.
H. rufeseens Penn., iii, 175.
/. depressa Tayl., minor Jeffr.,
rubens Moq., albocincta
Ckll., alba Moq., manches-
teriensis Bgt, brittanica
West.
v. striolata C. Pfr.
/. subcarinata Cless.
f. abludens Loc.
v. submontana Mab.
pascali Mab.
v. putoni Cless.
H. saxivaga Malz.
H. sericea Drap., iii, 178.
/. gerstfeldiaua Cless., plana
Mil., caucasica Mouss., fon-
tainei Colb.
v. expansa Cless.
v. corneola Cless.
v. dubia Cless.
v. libertina West,
v. badiella Zgl, Bgt.
v. subbadiella Bgt.
v. subglobosa Jeffr.
'
HYGROMIA.
275
H. sordulenta Morel., iii, 177.
H. stuxbergi West.
sericea Schrenk.
H. subcselata Less., iii, 173.
v. hiaticula West.
H. subplebeia Less., iii, 173.
H. telonensis Mitt., iii, 186.
/. lavandulse Bgt., drueutina
Bgt., disega Bgt., gelida
Bgt., concreta Bgt., crimoda
Bgt., pedemontana Pini,
salassia Poll., pegorarii
Poll., segusina Less., iii, 179.
v. crassilabris Nev.
v. moutoni Mitt., iii, 186.
H. teneitensis Bgt., iii, 180.
H. transsylvanica Biz.
H. umbrosa Partsch, iii, 176.
v. aporata Bgt.
v. urabrosella Jouss.
v. sciraia Bgt.
v. oecoscia Bgt.
v. amella Bgt.
H. urbana Cout.
H. vespertina Morel., iv, 41.
H. villersii Malz., iii, 173.
H. villosa Stud., iii, 177.
hispidula Jan.
pilosa Alten.
/. detrita Hartm.
v. phorochsetia Bgt.
H. villosula ZgL, iii, 176.
pietruskyana Parr., iii, 176.
Oriental species.
H. submissa Dh., iii, 182. H. puberula Hde., iii, 183.
H. subechinata Dh., iii, 182. H. horripilosella Hde., iii, 183.
H. szechenyii Anc. H. nautarum Hde., iii, 183.
H. tchefouensis C. & D., iii, 182. H. rebellis Hde., iii, 183.
munieriana C. & D. H. barbosella Hde.
Unfigured and insufficiently known species : H. hypsellina Loc.,
steneligma Bgt., microgyra Bgt., hispidella Bgt., deobrigana Bgt.,
ataxiaca Fag., vendeana Let., bellovacina Mab., gosseni Mab., ela-
verana Bgt., cavarella Serv., duesmensis Loc., saporosa Mab., astenia
Mab., ferdinandi Serv., bofilliana Fag., alphsea Let., anasina Serv.,
rosai Silv., cularensis Bgt., sarinica Bgt., tumescens West., vendo-
peranensis Bgt., vocontiana Bgt., axonana Mab., sericella Serv., ba-
variana West., aporiua Silva, ischnia Mab., euclastolena Mab., ber-
bruggeriana Loc., baccueti Bgt., challameliana Bgt., bastidiana
Bgt., cotinophila Bgt., tseniata West., inversa West., hierocontina
West., lentiaca Sayn (seeviii, 190).
Section Ciliella Mousson, 1872.
Ciliella Mouss., Rev. Faun. Mai. Canaries, p. 60 (for If. ciliata,
leprosa, lanosa}. Lepinota WESTERLUND, Fauna Palaarct. Reg.
Binnenconch., Helix, 1889, p. 2, 16, type H. ciliata. Not Lepino-
tus Heyd., 1850, (Neuroptera).
276 HYGROMIA.
Shell narrowly umbilicate, subdepressed, keeled or angular at
periphery, thin and brownish ; surface sculptured with short, scale-
like cuticular processes ; aperture oblique, oval ; outer and basal
lips well expanded, somewhat thickened within. Type H. ciliata
Yen., pi. 55, figs. 22, 23, 24.
Animal having the mantle blotched with black as usual in the
genus.
Jaw arched, transparent and yellowish, with numerous, fine,
close, regular but not well defined riblets, hardly crenulating the
margins (H. ciliata). Genital system (pi. 70, fig. 30, H. ciliata)
with short, stout penis, its retractor terminal and with bifid inser-
tion ; epiphallus ending in a short, stumpy flagellum. Duct of
spermatheca moderately short, swollen at base. Dart sack and
mucus glands wanting.
Distribution, south-central Europe, Canary Islands.
H. ciliata Ven., iii, 187. H. guevarriana Bgt.
folliculata Risso. H. stussineri Bttg.
hirsuta Jan. H. leprosa Shutt., iii, 223.
v. biformis Beck. H. lanosa Mouss., iii, 223.
Section Metafruticicola v. Ihering.
Pseudocampylcea HESSE, Jahrb. D. M. Ges. 1884, p. 237, (q. v. for
anatomy) ; TRYON, Manual of Conch. (2), iv, p. 114 (part), and of
other authors. Not Pseudocampylcea Pfeiffer, Mai. Bl., xxiv, p. 8,
1877. Cressa WESTERLUND, Fauna Palaarct. Reg. Binnen-Conch.,
Helix, p. 4, 101, 1889. Not Cressa Bock, 1871 (Amphipoda).
Metafruticicola IHER., Zeitschr. f. Wissensch. Zool. liv, p. 452 (Oct.
4, 1892).
Shell moderately solid, opaque, rather small, depressed-globose,
umbilicated, with convex spire of nearly 5 whorls, the first one
smooth or costulate, the remainder granulate, grano-costulate or
spirally decussated, often with hairs standing in oblique series.
Last whorl rounded at periphery, descending in front. Aperture
subcircular or oval; peristome sharp-edged, slightly expanded
below, strengthened by a scrong submarginal internal rib. Type
H.pellitaFer.
Mantle flecked with dark. Right eye retractor passing between
branches of genitalia. Jaw with many (15 to 20) fine vertical rib-
lets (pi. 70, fig. 31, H. pellita).
HYGROMIA. 277
Genitalia (pi. 70, fig. 37, H. noverca) ; penis short and stout, con-
tinued in a long epiphallus bearing the retractor, and ending in a
long flagellum. Spermatheca oval on a moderately long, unbranched
duct, which is decidedly swollen or enlarged toward its insertion
low on vagina. Dart sack and mucous glands wanting.
The shell has much the aspect of a Helicigona of the cyclolabris
group, but the lip is less expanded and more thickened within, as
in Theba. The group is doubtless correctly placed by Hesse and
v. Ihering, who consider it a Fruticicoloid with the dart apparatus
lost by degeneration. The jaw and other anatomy as far as known,
as well as the structure of the shell, all point to this solution of the
simple genitalia. It will be noticed that in Metafruticicola the loss
of the dart apparatus has left exactly the type of genitalia found
in the group Epiphallogona (Camcena, Chloritis, Planispira, etc.).
No jaw of the delicate Fruticicola type has been found in the Epi-
phallogona.
Distribution, Grecian Archipelago, most species from Crete.
H. pellita Fer., iv, 115. H. lecta Fer., viii, 191.
v. kreglingeri Zel. H. sublecta Malz., iv, 116.
v. graphicotera Bgt. H. zonella Pfr., iv, 110.
H. testacea Mart., viii, 191. notthefigs. cited.
H. dictsea Mart., viii, 191. H. medea West., viii, 192.
H. naxiana Fer., iv, 115. H. giurica Bttg., yiii, 192.
H. westerlundi Bl., iv, 115. H. cerigottana Bttg., Nachr. '94,
H. noverca Friv., iv, 115. 6.
Section Perforatella Schliiter, 1838.
Perforatella SCHLUTER, Syst. Verz., 1838. Petasina MORCH,
Catal. Yoldi, 1852, p. 6 (for edentula Drap.).
Shell low-trochoidal, with rounded-conic raised spire, subangular
periphery and small or minute umbilicus. Whorls numerous and
narrowly coiled. Brownish, often with a light peripheral band.
Aperture basal, narrow ; basal lip expanded, thickened within by a
very strong callus, which is usually more or less truncate or 1-toothed.
Type H. unidentata Drap.
Jaw thin, horn-colored, with 18-25 riblets (leucozona]. Genitalia
(pi. 70, fig. 36, H. leucozona} as in typical Fruticicola. The mucus
278 HYGROMIA.
glands are inserted very high, four on each side ; some distance
below them are two dart sacks, with two accessory sacks, the latter
containing no darts. Penis, etc., as usual in the genus. The geni-
tal system of H. unidentata is the same as in leucozona.
A group of Middle Europe, in which the shell has much similar-
ity to Dibothrion, but the genitalia are as in Frutidcola hispida, etc.
H. unidentata Drap., iii, 171. H. leucozona Zgl., iii, 171.
cobresiana Alt., iii, 171. coadvnata Z.
monodon Fer. /. delopida Jan.
villce Mu'hlf. /. crassilabris Miihlf.
ventricosa Jan. /. rutilans Z.
/. anodonta Tschap. modesta Parr.
/. alpestris Cl. v. ovirensis Rin., iii, 172.
/. subleucozona Fag. v. heteromorpha W., iii, 172,
H. edentula Drap., iii, 171. v. erjaveci Cless., iii, 172.
depilata Drap. H. bielzi Schm.
liminifera Held. v. bosnensis Mlldff.
Section Dibothrion Pfr., 1855.
Petasia BECK, Index, 1837, p. 21. MARTENS, Die Hel., I860, p.
102, type H. bidens. Not Petasia Serv., 1821 (Orthoptera) of
Steph., 1829, (Lepidoptera) or of Morr. 1829, (Polyzoa). Trocliiscus
HELD, Isis, 1837, p. 915. Not Troehiscus Heyden 1826, (Arach-
nida). Dibothrion PFR., Mai. Bl. 1855, p. 128 (for bidens and
bicallosa}.
Shell subtrochoidal, with convex-subconic spire of numerous nar-
row whorls, and closed or nearly closed umbilicus ; arcuate-striate,
translucent brown with light peripheral band. Aperture basal,,
narrow ; lip well expanded, reflexed, with two strong internal teeth
or nodules on the basal margin, marked by pits behind the lip ;
parietal wall toothless. Type If. bidens, pi. 55, figs. 25, 26.
Genitalia (pi. 70, fig. 41, H. bidens) : Dart sack single, cylindrical
and rather large ; two mucus glands on each side. Other organs as
in Hygromia generally, except that the spermatheca duct is unusu-
ally short. Dart needle-like, with four short blades at the point.
Distribution, eastern Europe and Siberia.
HYGROMIA, AULACOSPIRA. 279
H. bidens Chemn., iii, 170. H. bicallosa Friv., iii, 171.
bidentata Gin.
v. diodon Parr., iii, 171.
dibothrion Friv.
Section Metodontia Mollendorff, 1886.
Metodontia MLLDFF., Jahrb. D. M. Ges., 1886, p. 191, type H.
hemipleuris. Tetrodontina ANCEY, Conch. Exch. i, p. 64, May,
1887. gee HILBER, Sitzungsber. k. Akad. Wissensch., Ixxxvi, pi.
1, f. 1-3, development of aperture armature.
Shell perforate, globose-turbinate or subdepressed, brown or
whitish ; whorls numerous and narrow, the last not descending in
front. Aperture lunate, nearly closed by two large lip teeth usually
situated on -a ridge of callus, opposed to two smaller teeth on the
parietal wall. Lip thin-edged, expanded below, reflexed at colu-
mellar insertion. Type H. hemipleuris Mlldff.
Anatomy unknown. The group seems most nearly allied to
Dibothrion of the Eur.-Asian fauna. It has nothing to do with the
American group Triodopsis.
The group is confined to the dry northern half of China, being
about coextensive with the Loss formation.
H. hemipleuris Moll., iii, 149. H. houaiensis Cr., iii, 149.
moltneri Gredl. obstructa Hde. not Fer.
H. yantaiensis C. & D., iii, 149.
v. tetrodon Moll., iii, 149.
Genus AULACOSPIRA Mollendorff, 1890.
Aulacospira MLLDFF., Bericht u'ber die Senckenbergische natur-
forschende Gesellschaft in Frankfurt a. M., 1890, p. 224. Micro-
petasus MLLDFF., I. c.
Shell small, umbilicate, of thin corneous structure; unicolored,
pale brown. Spire more or less raised, from the beginning sub-
scalar and keeled ; whorls 4 to 5, striatulate, flattened or with spiral
concavity, the last whorl keeled or rounded. Aperture oblique,
generally subcircular, with to 5 teeth a short distance within ;
peristome thin, reflexed. Type A. scalatella Mlldff, pi. 64, figs. 10,
11,12.
280 AULACOSPIRA, ACANTHI.NULA.
Anatomy unknown. The species live in clefts and crevices of
limestone rocks, in which the flatness of their shells allows them
ready access, and like Vallonia they seem to be gregarious. The
species now known are fromCebu, Busuanga, Luzon and the Island
Ilin, near Mindoro ; but it probably will prove to have many more
species in other islands of the Philippine group.
In deference to the opinion of Mollendorff I give the group place
herein the vicinity of Fruticicola; but my own course would be to
place it next to Eulota or even as a subgenus within that genus.
Compare the similar group Platypetasus (p. 207). I am now disposed
to consider Pupisoma (see p. 52), as well as Aulacospira, as branches
of the Eulota stock, parallel to, rather than allied to Acanthinula
and Vallonia which seem to be early branches of the Hygromia
phylum. They will probably be found to have the genitalia con-
siderably simplified by suppression of accessory organs, and the
marginal teeth multicuspid ; these changes usually accompanying
such great reduction in the size of snails.
Mollendorff establishes two sections:
Aulacospira s. str. (of which Micropetasus is an absolute synonym).
Keel extending to aperture ; perietome continuous and free.
Pseudostreptaxis Mlldff. (1. c. p. 225). Penultimate whorl dis-
tinctly deviating ; the last whorl not keeled, cylindrical ; aperture
5-toothed, the peristome not continuous. One species A. azpeitice.
A. hololoma Mlldff., viii, 198. A. porrecta Quadr. & Mlldff.,
A. mucronata Mlldff, viii, 198. Nachrbl. 1894, p. 95.
A. scalatella Mlldff, viii, 199. A. azpeithse Hid., viii, 199.
Genus ACANTHINULA Beck, 1846.
Acanthinula BECK, Amtl. Ber. Vers. Kiel, 1846, p. 122. v. MART.
in Die Hel., 1860, p. WO.Euacanthinula WEST., Fauna, p. 16.
Shell minute, pyramidal or globosely-turbinate, thin, brown,
minutely umbilicated. Epidermis raised into lamellae crossing the
whorls. Aperture subvertical, semilunar or subcircular, the lip
acute, expanded toward columellar insertion, the margins remote.
Type A. aculeata Mull., pi. 70, figs. 26, 27, 28.
Animal apparently with even, not crenulated foot-margins ; ovi-
parous? Jaw arched, with numerous flat ribs. Genitalia (pi. 63,
ACANTHINULA. 281
fig. 11, A. lamellata, after Lehmann) apparently lacking mucus
glands, but dart sack present.
Distribution, Palsearctic and Nearctic regions, mainly northward.
The genitalia are very imperfectly known, the figure in Lehmann's
posthumous work representing A. aculeata being very much like a
Buliminus or Pupa, and possibly, as v. Ihering suspects, inserted
through some confusion of drawings. A new investigation is
urgently required. Possibly the group does not belong to the Heli-
cidse. A. lamellata and aculeata have been found in Pliocene
deposits ; and in the lower Miocene of middle Europe the genus is
represented by several species, A. nana A. Braun, paludiniformis
Br., tuchoricensis Klika, plicatella Reuss.
A. aculeata Mull., iii, 53. A. lamellata JefFr., iii, 54.
spinulosa Lightf. scar bur gensis Alder.
granatelli Biv. seminulum Rossm.
delectabilis Sol. A. peracanthoda Bgt., iii, 54.
/. albida JefFr., iii, 53. raffrayi Bgt. not Canef.
albina West. r affray ana Ckll.
v. sublsevis West., iii, 54. A. spinifera Mouss., iii, 54.
A. spermatia Silva. A. monas Morel., iii, 54.
A. harpula Reinh.
Section Zoogenites Morse, 1864.
Zoogenites MORSE, Terr. Pulm. Maine, p. 32, pi. 1.
Shell globose-turbinate, perforate, ornamented with oblique cuti-
cular lamellae. Type Z. harpa Say, pi. 70, figs. 23, 24, 25.
Animal with the foot-edges prominently crenulated ; labial lobes
large ; lower tentacles nearly obsolete ; viviparous, the young at
birth as large as the aperture of the shell. Jaw (pi. 70, fig. 35)
having numerous wide subobsolete ribs. Radula with formula
11.6.1.6.11. Middle teeth tricuspid, the mesocone not reaching edge
of the square basal plate. Laterals bicuspid. Marginals wide,
with many irregular cusps. Genitalia unknown.
A. harpa Say, iii, 54. New England, British America, Sweden,
Xamtchatka, etc.
Syn. : P. costulata Mich., H. amurensis Gerstt.
VALLONIA.
Genus VALLONIA Risso, 1826.
Vallonia Risso, Hist. Nat. Eur. Merid. iv, p. 101, sole species F^
rosalia. Zurama LEACH in Turton's Man. L. and Frw. Sh. Brit.
Is., p. 64, ISBl.Amplexus BROWN, 111. Conch. G. B. 1827 ; Edit.
1844, p. 45. Chilostoma FITZ., in part, 1833. Circinaria BECK,
Index (in part), p. 23. Glaphyra ALB., Die Hel., 1850, p. 87 (in
part). Lucena MOQ.-TAND., Moll. Fr. ii, p. 140, not Oken. See
STERKI, Proc. Acad. Nat. Sci., Phila., 1893, p. 234 (monograph of
genus, jaws and dentition). ASHFORD, Journ. of Conch, iv, p. 198
(dart). LEHMANN, Die lebenden Schneeken u. Muscheln Stettins
u. Pommern, p. 90, pi. 11, f. 30 (genitalia, etc.).
Shell minute, openly and widely umbilicate, depressed, the spire
low-convex, consisting of 3-4? whorls, color light and uniform ;
surface smooth or ribbed ; periphery rounded ; last whorl usually
descending in front. Aperture oblique, circular or short-oval ;
peristome continuous or nearly so, expanded or reflexed, often thick'
ened within. Type V. pulchella Mull., pi. 55, figs. 31, 32.
Foot small, short, with no pedal grooves ; edges of sole somewhat
crenulated; sole undivided ; eye-peduncles cylindrical, not enlarged
distally ; tentacles short ; labial lobes well developed.
Jaw arcuate, with a slight median projection or none, sculptured
with numerous (18 to 25) crowded, low riblets, denticulating the
margins (pi. 70, fig. 29, V. pulchella).
Radula having 23 to 33 teeth in a transverse row. Median teeth
decidedly narrower than laterals, tricuspid, the mesocone not half
as long as basal-plate, side cusps smaller. Laterals with large
square basal plates, the mesocone extending to its edge, ectocone
small. Marginal teeth wide and low, multicuspid (pi. 70, fig 38 V-
pulchella}.
Genitalia (pi. 63, figs. 9, 10, V. pulchella, after Lehmann) having
the penis short, with terminal retractor; epiphallus short, bearing a
flagellum. Dart sack present, single, containing a straight, bladeless
dart (fig. 10, x 100). No mucus glands. Duct of spermatheca
long, branchless.
Distribution, North America South to Texas; Japan and middle
China to Europe and Atlantic Islands. Fossil the group is known
from the lower Eocene ( V. sparnacensis Dh.) ; and in the Miocene
several species, lepida Reuss., subpulchella Sandb., occur.
VALLONIA. 285
This very distinct genus of minute snails occupies the entire
Nearctic and Palsearctic regions, and some species have been intro-
duced (probably with plants) into Australia, Mauritius, etc. They
are gregarious in habit, and live under fragments of wood, stones,
on mossy cliffs and in damp meadows, always avoiding light. The
number of species is very uncertain ; but whether species or varie-
ties, there can be no doubt that a considerable number of recognizable
forms must be distinguished. The arrangement given below is that
of Dr. Sterki, who has made special studies on a far greater mass of
material than any other observer. An alternative to this classifica-
tion would be to make excentrica a variety of pulchella, and unite
adela, declivis and pollinensis under the former of the three names ;
all the forms of the costata group might then fall under eostata as
varieties. Having seen neither mionecton, ladacensis nor asiatica I
do not care to suggest any mode of uniting the forms included in
Sterki's " Group of F cyclophorella."
Group of V. pulchella.
V. pulchella Mull., viii, 248. v. enniensis Gredl.
rosalia Risso (part). v. hispanica Sterki.
paludosa Da Costa. v. persica Rosen.
erystallina Dillw. V. excentrica Sterki, viii, 249.
Icevigata Moq. V. adela West., viii, 251.
nitidula Stud. V. declivis Sterki, viii, 251.
potua Chier. v. altilis Sterki.
minuta Say. V. pollinensis Paul., viii, 252.
Group of F costata.
V. costata Mull., viii, 252. V. albula Sterki.
alexandrce Cox. V. parvula Sterki, viii, 254.
rosalia Risso (pt.). V. tenera Reinh., viii, 255.
crenella Mont. pukhellula Hde.
helicinus Lightf. v. patens Reinh., viii, 257.
v. helvetica Sterki. V. gracilicosta Reinh., viii, 256.
v. amurensis Sterki.
v. pyrenaica Sterki.
v. montana Sterki.
284 HELICODONTA.
Group of V. cydophorella.
V. cyclophorella Anc., viii, 259. V. mionecton Bttg., viii, 260.
V. perspectiva Sterki, viii, 257. v. schamhalensis Rosen.
V. tenuilabris Br., viii, 258. V. ladacensis Nev., viii, 260.
v. saxoniana Sterki, viii, 259. v. asiatica Nev., viii, 260.
Genus HELICODONTA Ferussac, 1819.
Helicodonta FEE. Tabl. Syst. de la Fam. des Limacons, p. 33 (in
part). Risso Hist. Nat. de 1'Eur. M6rid., iv, p. 65, 1826 (re-
stricted to H.obvoluta). Trigonostoma FITZINGER, Syst. Verzeich.,
1833, p. 97, species H. holosericeum, H. obvolutum. Not Trigonostoma
Blainv., 1825, Cancellariidce. Vortex BECK, Index Moll., 1837, p.
29. Gonostoma HELD, Isis, 1837, p. 915 (preoccupied by Rafines-
que in Pisces, 1810.) Anchistoma ("Klein," preLinnsean) H. & A.
ADAMS, Gen. Rec. Moll, ii, p. 205, 1855 (subg. Polygyra and Dre-
panostoma only, the former in part). Euphemia LEACH, teste BECK,
Amtl. Ber. v. Kiel, 1846, p. 122. ? Plieostoma SCHLUTER, Verz. p.
4, 1838, s.-g. Helix, for H. intestinalis Schlu't. (publication not seen
by H. P.^. Chilodon and Helicodon EHRENB. Symb. Phys., 1831.
Drepanostoma PORRO, Mag. de Zool., 1836, classe v, pi. 71, type
D. nautiliformis. Contorta MEG. de MUHLF., test. VILLA, Disp.
Syst. Conch. Coll. Villa, p. 19, 1841.
Caracollina BECK, Index Moll., p. 28, 1837. LOWE, P. Z. S.,
1854, p. 196, type If. barbula Charp. And probably Caracollina
EHRENBERG, Symb. Phys. Evert., Moll., no diagnosis ; no species
mentioned. Caracolina auct.
Aspasita WESTERLUND, Fauna der in der Palaarctischen Region
Lebenden Binnenconchyl., Helix, pp. 18, 26, type H. triaria.
See for anatomy, A. Schmidt, Stylommatophoren p. 34, pi. 8.
Moq.-Tand, Hist. Nat. Moll. France, p. 109-114, pi. 10. Hesse,
Jahrb. d. d. Malak. Ges. xi, 1884, p. 233, pi. 4, f. 5. St. Simon,
Journ. de Conch., 1867, p. 98. Schuberth, Archiv f. Naturg.,
1892, p. 5, pi. 1. v. Ihering, Morphol. u. Syst., p. 475.
Shell depressed, usually umbilicated, rather thin, never cretaceous ;
unicolored brown; striate, granulate, ribbed or hairy. Spire low,
consisting of numerous closely coiled whorls. Aperture triangular,
rhombic or lunate, the lip-ends remote ; peristome expanded and
reflexed, lipped, often toothed. Type H. obvoluta, pi. 56, 'figs. 25,
HELICODONTA. 285
26, 27 (see also pi. 56, figs. 16, 17, R. constricta; figs. 18, 19, H.
lens; figs. 23, 24, H. triaria ; figs. 28, 29, 30, H. biconcava; figs. 31,
32, 33, H. nautiliformis').
Animal rather elongated, with long, narrow foot, the sole undi-
vided ; back with a pair of longitudinal grooves; facial grooves
wanting. Mantle with a small right body-lappet; no left one.
Right eye-retractor passing between branches of genital system.
Epiphragm papery, flat, formed rather deep in the mouth.
Jaw thin and flexible, with numerous (6 to 16) broad, flat ribs,
separated by narrow interstices (pi. 36, fig. 6, H. lenticula ; pi. 36,
fig. 8, H. maroccana).
Radula having the mesocones slightly longer than the basal
plates, ectocones small. Marginal teeth having the ento- and meso-
cones united at base, ectocone developed (pi. 36, fig. 5, jET. lenticula.
PI. 36, fig. 7, H. maroccana).
Genital system: penis long, the retractor median or terminal,
inserted didally on eolumellar muscle; no flagellum. Vagina long,
bearing from one to three cylindrical elongated mucus glands, with
one dart sack at or below their base, sometimes lacking ; the dart,
when present, short and conical (pi. 36, fig. 9, H. lusitanica}.
Spermatheca oval, its duct short, bound to the uterus and without
diverticulum. See pi. 36, fig. 4, H. obvoluta; pi. 36, fig. 10, IT.
lusitanica.
The number of mucus glands varies from one to three, ff. obvo-
luta has one long and one very short gland. The dart sack is en-
tirely absent in some species. The dentition is of the type usually
developed in ground snails. The jaw is uncommonly delicate for
the ribbed type. The union of the penis retractor muscle with the
great eolumellar retractor is a peculiar feature, and it will be inter-
esting to find whether it holds throughout the genus. I have ob-
served it in H. obvoluta only.
The brownish, unicolored shell, with depressed spire, slowly
widening narrow whorls and reflexed, lipped peristome, is very
characteristic ; and no shells of the Palsearctic fauna, except Isog-
nomostoma and Dibothrion can be compared with this genus these
two groups also having toothed apertures, but sufficiently different
in form from those jf Helicodonia.
Von Ihering maintains, I believe with right, that this genus is
more nearly allied to Fruticicola than to Helix or Campy laza, differ-
286 HELICODONTA.
ing mainly in the reflexed, lipped peristome, and more or less degen-
erate dart apparatus. Helicodonta is not in the least allied to the
American or Indian toothed Helices, or to the so-called Gonostoma
of California.
The circum-Mediterranean region is the headquarters of this
genus, although a few forms occupy central Europe, and one, obvo-
luta, has obtained a foothold in southern England. In south-east
Asia it reappears in a number of specific forms comparable to the
European species obvoluta and diodonta, but not readily falling into
the sectional groups established for European forms. The Canary
Islands are nearly the westward outpost of the genus, one species
only occurring in Madeira.
The name Helicodonta was originally proposed for all toothed
Helices ; but was restricted by Risso, in 1826, to H. obvoluta. As
no other name for the group appeared before 1833, there is no ques-
tion as to the propriety of reverting to this one, especially since the
names in common use, Gonostoma and Trigonostoma are preoccu-
pied, and must, in any case, be rejected. Besides two species still
retained in this group, Ferussac included in Helicodonta members
of the prior genera Polygyra, Pleurodonte, Cepolis, and Anostoma,
as well as of the later groups Strobila, Corilla and Petasia. The
term Anchistoma of the Adam's brothers (1855) has been used for
Helicodonta by Kobelt and others. It has been attributed to Klein
{1753), but his "Angystoma" contains none of the European toothed
Helices and, in any case, the genera and species of the Tentamen
methodi Ostracologicoe are not of Linnsean form, and antedate the
Linnsean era. Ehrenberg's contribution to the taxonomy of this
group is of little value. He divides the land snails into two series,
based on the absence or presence of aperture-teeth : Chilogymnus
containing Helix, Caracolla, Bulimus, Pupa, and Chilodon contain-
ing Helicodon, Caracollina, Bulimina, Pupina. The genera of
Chilodon are all new, although he does not so state ; but, as they are
nude names, without a word of diagnosis except what may be
tacitly gathered from the above arrangement, and as no species of
any of them are mentioned, their bearing on nomenclature is nil,
and none of them can be dated from 1831, or adopted at all except
when defined by later authors. The group Vortex of Oken (1815)
contained depressed Helices and Zonitids of many groups, and, as it
is a composite group, and the name was not used in especial connec-
tion with Helicodonta until after the publications of Ferussac and
HELTCODONTA. 287
Risso, it has no claim for adoption, and had better be dropped en-
tirely.
Key to Sections of Helicodonta.
a. Spire deeply sunken, narrower than umbilicus ; aperture narrow-
crescentic, lip simple and retracted above, expanded and lipped
outwardly and below. Nautilus-shaped, Drepanostoma.
aa. Spire wide, nearly level or convex.
6. Aperture triangular or square, lip teeth 2 or obsolete, pe-
riphery rounded.
c. Not ribbed; last whorl wider than preceding; spire
nearly level, Helicodonta s. s.
cc. Small, ribbed above, last whorl narrow as preceding ;
spire convex, Aspasita.
bb. Aperture lunate or rhombic; shell much depressed.
c. Outer edge of parietal callus raised into an erect bar-
rier, Trissexodon.
cc. Parietal callus thin, wholly adnate, Caracollina.
bbb. Aperture lunate, toothless ; shell sub-globose, Klikia.
Species.
DREPANOSTOMA Porro. Nautiloid, biconcave, with crescentic
mouth.
H. nautiliformis Porro., iii, 114. Lombardy.
drepanostoma Bk.
HELICODONTA (Fer.) Risso. Species of middle Europe.
H. angigyra Zgl. Rm., iii, 115. H. obvoluta. "' //J ^
stentzii Partsch /. pallida M.-T.
H. obvoluta Mull., iii, 115. v. bosniaca Bttg., iii, 115.
trigonophora Lam. v. blanci Poll., viii, 150.
bilabiata Oliv. H. holoserica Stud., iii, 116.
holosericea Gmel. diodontostoma Bgt.
/. dentata Held. v. pluridentata Poll., iii, 116.
/. edentata West. H. diodonta Miihlf., iii, 116.
Chinese species of Helicodonta.
H. subobvolutaAnc. J. B.,xi,308. H. diplomphala Moll., iii, 124.
H. molina Hde. H. uninodata Gred.,viii, 150.
H. biconcava Hde., iii, 117. H. binodata Mlldff., iii, 124.
outangensis Crosse. ? bicallosula Hde.
288 HELICODONTA.
ASPASITA Westerlund. Transylvania, Hungaria.
H. triaria (Friv.) Rm., iii, 116. H. trinodis Kim., iii, 116.
ocskayi Stentz. transsylvanica Haz.
v. tatrica Haz. H. triadis Kim., iii, 116.
TRISSEXODON Pi Is. Depressed, umbilicate; outer edge of parietal
callus raised into a transverse lamellar barrier, constricting the
mouth. Pyrennes ; southern Spain.
H. constricta Boub., iii, 121. H. quadrasi Hid., iii, 116.
pittorrii Dup.
CARACOLLINA Beck. Around the Mediterranean.
H. corcyrensis Partsch., iii, 117. barbata Desh.
contorta, tersa Zgl. v. lentiformis Zgl.,iii, 119.
ambliostoma Parr. v. piligera Bl. & W.
v. cephalonica Mouss., iii, 118. abantisorum Serv.
/.minor. v. callojuncta West.
v. octogyrata Mouss., iii, 118. v. aliostoma West., iii, 120.
v. canalifera Ant., iii, 118. v. elia Bttg.
v. girva Friv., Rm.,iii, 118. H. lentina Mart.
H. gyria Roth., iii, 117, H. turriplana Mor., iii, 120.
H. barbata Fer., iii, 118. H. rangiana Fer., iii, 121.
H. lusitanica Pfr.,iii, 117. rangii auct.
H. tarnieri Morel., iii, 118. H. barbula Charp., iii, 120.
H. walkeri Pons., Kob., viii, 149. bituberculata Fer.
H. boscje Hid., iii, 118. guerini Ant.
H. annai Pal., viii, 148. bidentifera\P]iil\ips.
H. lenticularis Mor.. iii, 120. H. barbella Serv.
H. columnse Pons., Kob., viii, 148.H. camerani Less.
H. lenticula Fer., iii, 119. H. gougeti Serv., iii, 121.
H. maroccana Mor., iii, 120. H. tlemcenensis Bgt., iii, 120.
H. calpeana Mor., iii, 120. v. pechaudi B., Anc.
finitima Fer., undesc. H. supracostata Kob., viii, 149.
H. vallisnieri Stef., iii, 117. H. buvignieri Mich., iii, 121.
H. lens Fer., iii, 1 1 9. asturica Pfr.
Canary Island Species.
H. lenticula v. virilis Mouss., iii, H. hispidula Lam., iii, 122.
119. v. subhispidula Mouss.
subtilis Lwe. v. bertheloti Fer.
HELICODONTA. 289
H. parryi Pons. & Sykes. H. everia Mab., iii, 123.
H. afficta Fef., iii, 122. H. marcida Sh., iii, 123.
H. planaria Mouss., iii, 122. H. crispolanata Woll, iii, 123.
H. discobolus Sh., iii, 123. H. beata Woll., iii, 123.
H. fortunata Sh., iii, 123, H. gomerse Woll., 123.
H. pthonera Mab., iii, 123. H. eutropis Shutt. iv., 36.
Section Klikia Pilsbry, 1894.
Shell depressed-globose, narrowly umbilicated, with convex, ob-
tuse spire and round periphery. Surface costulate-striate and
minutely papillose in regular diamond pattern. Last whorl con-
stricted behind the lip, which is well reflexed and thickened. Type
H. osculum Thomae, pi. 71, fig. 49.
This apparently extinct type of Helicodonta is characteristic of
middle European Miocene, where it coexisted -with species of Cara-
collina, such as phacodes Thomae, and with species of typical Heli-
codonta ; H. involuta Thomae being allied to the recent angigyra
and biconcava. The strong differentiation of these sectional groups
at as early a period as the lower Miocene (when they were, in fact,
as strongly differentiated as in the recent fauna), argues a vastly
greater antiquity for the genus as a whole. This group is named in
honor of Gottlieb Klika, author of an excellent memoir upon tertiary
land and fresh-water shells of Bohemia.
Subgenus MOELLENDORFFIA Ancey, 1887.
Mollendorfia ANC., Conch. Exch., May, 1887, p. 64. PILSBRY,
Man. Conch., vi, p. 10. Proctostoma MABILLE, Bull. Soc. Mai. de
France, iv, p. 102, 103,104,1887 (for H. loxotatum^.Polygyra and
Cepolis of some authors. Trihelix ANC., t. c., p. 64 (for H. horrida).
Shell depressed, with low-convex, flat or concave spire of 4%-5?
whorls, rounded or keeled periphery, and convex, umbilicated base.
Surface more or less granular, tubercular or hairy, uniform brown,
dull and opaque. Apical whorl rather large ; last whorl deeply de-
flexed in front, with deep pits or grooves behind the lip. Aperture
very oblique or subhorizontal, trigonal or squarish, the lip expanded
and reflexed, continuous across the parietal margin, sometimes solute ;
basal lip armed with a stout tooth, outer lip with one or two large en-
tering folds. Type H. trisinuata Martens (see pi. 40, figs. 16, 17, 18,
H. hensaniensis Gredl.; pi. 56, figs. 20, 21, 22, H. erdmanniSchmaclu.?
& Boettger).
19
290 ALLOGNATHUS.
Anatomy unknown. Distribution, southeastern China, Tonquin
and Cambodia.
Among Old World Helices this peculiar group can only be com-
pared with the typical Helicodontas (obvoluta, holoserica, diodonta,
etc.) of Europe and China. From these, Mollendorffia differs in the
fewer whorls, continuous peristome, etc. Still, I find no differences
of generic value in the shells; and, unless the anatomical features
prove peculiar, we can hardly accord the group higher rank than
subgeneric. In America, we find analogical shell structures in
Pleurodonte (Labyrinthus} leucodon and its allies, which somewhat
resemble H. trisinuata; and H. horrida may be compared with
Epiphragmophora (Averellia) macneilliCrosse', but these are merely
adventitious resemblances, due to the action of mechanical causes,
which have produced tridentism in many diverse groups of Helices,
H. loxotatum Mab., vi, 13. H. erdmanii Schm. & Bttg.
H. trisinuata Mart., vi, 11. Proc. Mai. Soc. Lond., i, pi. 9.
v. sculptilis Moll., vi, 12. H. faberiana Moll., vi, 10.
H. eastlakeana Moll, vi, 12. H. biscalpta Hde., vi. 9.
H. hensaniensis Gredl., vi, 299. H. horrida Pfr., vi, 9.
Genus ALLOGNATHUS Pilsbry, 1888.
Allognathus PILS., Man. Conch. (2), iv, pp. 121, 149, type H. grat-
eloupi. KOBELT Nachrbl. D. M. Ges., 1891, p. 140. Cf. SCHU-
BERTH, Archiv f. Naturg., Iviii, 1892, pp. 38, 61, pi. 4, f. 10, 11 (an-
atomy).
Shell globose, thin, smooth, imperforate, with low spire of about
4i whorls and obtuse apex ; last whorl descending in front, pale,
with five spotted bands. Aperture round-lunate, oblique ; lip ex-
panded, in the middle of the base closely appressed, the columellar
margin arcuate and rather wide. Type H. graellsiana Pfr., pi. 43,
figs. 39, 40.
Jaw arcuate with a median projection, its surface entirely smooth.
Radula very large (length 7, breadth 4 mill.). Teeth all of the
same form, strap-shaped, bent in a half circle, the cusp single, simple
and blunt (pi. 36, fig. 2, teeth from above, fig. 3, profile of cusps).
Outer teeth similar but somewhat smaller.
Genital system resembling that of Helix but the sacculated
uterus extends far downward; the spermatheca duct is swollen
LEPTAXIS. 291
below, and bears in the middle a diverticulum 27 mill, long, not
bound to the uterus. Dart-sack large (6 mill, long), containing a
a four-bladed dart with expanded funnel-shaped crown ; above the
sack are inserted two mucus-glands, each split into two large, club-
shaped fingers. Penis 16 mill, long, cylindrical, the retractor mus-
cle terminal; flagellum 37 mill, long (pi. 36, fig. 1, A. graellsiana
Pfr.).
This genus is founded upon a single species inhabiting the Bale-
aric Islands. The shell furnishes no characters of more than specific
value, separating it from such species of Otala as beaumieri,
etc. ; and it is by no means impossible that this and some other
species, such as H. quedevfeldti Mts., may prove to belong to Allog-
nathus. Notwithstanding this similarity of shell, we find in the
anatomy profound differences from all other Pentatseniate Helices.
The jaw is smooth, as in Leucochroa; the teeth are very aberrant
in the narrow basal-plates which curve over into a blunt, strap-like
cusp, entirely lacking side cusps, and they are alike over the whole
radula. The genital system presents a resemblance to Cawpylcea in
the two-fingered mucous glands, but is otherwise more like the
Pentatseniate groups. We are disposed to consider Allognathus a
special modification of the Helix stock, comparable to the Poly-
mita off-shoot from Hemitrochus.
A. graellsiana Pfr., iv, 150.
grateloupi Graells not Pfr. tessellata Fer., not Miihlf.
Genus LEPTAXIS Lowe, 1852.
Leptaxis LOWE, Ann. Mag. N. H. (2), ix, p. 164, Feb., 1852 ; P.
Z. S. 1854, p. 164, type H. erabescens Lwe. Katostoma LWE., P. S.
S. 1854, p. 166, type H. phlebophora Lwe. MacAilaria sp. LWE., i.
c.j p. 166. Cryptaxis LWE., t. c, p. 168, type H. undata Lwe.
Campylcea sp. LOWE. Pseudocampylcea PFR., Mai. Bl. 1877, p. 8-
Nomencl. Hel. Viv. 1878, p. 162, types lowei and portosandana.
Lampadia Alb. MSS., LWE., P. Z. S. 1854, p. 197, type H. webbiana
Lwe. (Not " Lampadie/' Montf., French vernacular name for Lam-
pas Montf.). Mitra ALB., Die Hel. 1850, p. 115, type H. webbi-
ana ; (not Mitra Lamarck.)
Shell of moderate or large size, globose, globose-depressed or lens-
shaped and keeled, imperforate (rarely umbilicate), the surface
striate, plicate, granulated or malleated ; rather thin ; uniform
292 LEPTAXIS.
brown, banded or maculated. Whorls 5 to 6, the last wide, deflexed
in front. Aperture transverse-oval, oblique, the outer lip simple or
expanded, columella usually widened. Type H. erubescens Lowe.
(See pi. 43, fig. 41, L. undata; fig. 36, L. lowei; fig. 45, L. web-
biana).
Jaw (pi. 67, fig. 20 L. undata) well arched, strong, bearing very
widely unequally separated linear riblets, converging below, forming
median triangle.
Radula (pi. 67, fig. 19, L. undata) having the cusps of median
teeth about as long as basal plate, side-cusps obsolete; basal plate
with a backward-projecting tongue-like process. Lateral teeth with
a stout ectocone. Marginals having the inner cusp shorter than
usual and obtusely bifid, outer cusp simple.
Genital system (Frontispiece, figs. 8, 9, L. undata) having the
penis continued in an epiphallus which bears the retractor and ends
in a short flagellum and the vas deferens. Dart sack large, seated
on atrium. Mucus glands in two clusters, one composed of 5, the
other of about 10 tubes, which adhere laterally by twos or form larger
palmate groups (fig. 8, d. s. turned downward and groups of
mucus glands spread). Spermatheca very large, rather boot-shaped,
with a basal ccecum embedded in uterus; duct long and without
diverticulum. The penis-retractor is inserted distally on the lung
floor, and the right eye-retractor passes between branches of genita-
lia. Dart of large size, a little curved, with a lateral expansion on
each side. Spermatheca contained a rod-like chitinous spermato-
phore, star-like in section.
The jaw of L. undata is peculiar and unlike that of any allied
form, resembling most the jaw of Plectopylis. The teeth are char-
acterized by the strong development of ectocones on the inner
laterals. The genital system is remarkable for the unusual size and
shape of the Spermatheca which lacks diverticulum unless it be re-
presented by the basal sack figured. The mucus glands are in two
groups, and inserted on the vagina as usual in Helix, but the in-
dividual tubes adhere laterally in a way I have not observed in
other forms. They are not bound together like those of Eulota*
however. The dart had unfortunately been expelled from the in-
dividuals examined, but has been described by Morch (Journ. de
Conch. 1865, p. 390).
I had expected to find in Leptaxis some archaic characters pre-
served ; for its geographic position and the shell-peculiarities argue
LEPTAXIS. 293
for the group an ancient origin ; but the evidence shows that how-
ever remote in the past the type was derived from the continental
fauna, the main anatomical features of modern European Helices
were then well established. If the genetic relationship of Leptaxis
with Oligocene forms of middle Europe claimed by Sandberger and
others be admitted (and this we have no good reason for doubting),
then by implication those fossil forms were anatomically very like
the modern European Helices.
It is a noteworthy fact that in no anatomical feature, whatever,
does Leptaxis approach the West Indian groups of Helices. They
have diverged from different stocks, and since Mesozoic time along
widely separated paths both geographically and structurally.
Distribution, Madeira, Azores and Cape Verde groups.
(Shell depressed, with open umbilicus and expanded peristome.
PSEUDOCAMPYL^A. Porto Santo.)
L. portosanctanaSowb., iv, 199. L. lowei Fer., iv, 200.
(Shell imperforate. LEPTAXIS. Madeira species).
L. undata Lwe., iv, 189. L. psammophora Lwe., iv, 191.
corrugata Sol. ms. L. wollastoni Lwe., iv, 199.
groviana Fer. v. forensis Woll., iv, 199.
scabra Wood. L. chrysomela Pfr., iv, 198.
L. vulcanica Lwe., iv, 190. cenostoma Lwe. not Fer.
L. leonina Lwe., iv, 190. v. fluctuosa Lwe., iv, 198.
L. nivosa Sowb., iv, 190. L. erubescens Lwe., iv, 191.
exalbida Wood. simia Fer.
decolor ata Lwe. v. portosancti Woll.
v. phlebophora Lwe. v. advenoides Lwe.
chlorata Lwe. v. hysena Lwe., iv, 192.
v. planata Lwe., iv, 191. L. furva Lwe., iv, 192.
v. craticulata Lwe., iv, 191. L. (?) exornata Dh., iv, 198.
scrobiculata Lwe.
(Species of the Azores Is.)
L. azorica Alb., iv, 196. L. terceirana Morel., iv, 197.
L. caldeirarum M. & D., iv, 196. L. drouetiana Morel., iv, 197.
L. niphas Pfr., iv, 196. L. vetusta M. & D., iv. 198.
294 FRIDOLINIA, DENTELLOCARACOLUS.
(Species of the Cape Verde Is.)
L. advena W. & B., iv, 192. L. leptostyla Dohrn, iv, 194.
L. serta Alb., iv, 193. milleri Dohrn.
L. fogoensis Dohrn, iv, 193. L. primreva Morel., iv, 195.
L. visgeriana Dohrn, iv, 193. L. atlantidea Morel., iv, 195.
L. myristica Sh., iv, 194. L. subroseotincta Woll., iv, 195.
L.bolleiAlb.,iv,194.
(Imperforate, thin, aeutely keeled, with 3 to 4 rapidly widening whorls.
LAMPADIA. Madeira, Canaries.}
L. webbiana Lwe., iv, 200. L. mernbranacea Lwe., iv, 201.
Vit. bocagei Paiva. L. cuticula Sh., iv, 201.
Genus FRIDOLINIA Pilsbry, 1894.
Shell large, heavy, depressed-turbinate, umbilicate when young,
closed in the adult; surface obliquely coarsely malleated. Last
whorl large, descending in front and strongly constricted behind the
lip, swollen in the middle of the base. Aperture very oblique,
toothless; peristome narrowly reflexed, its remote terminations
joined by a callus, the basal and columellar margins thickened
by a heavy callus within. Type H. lucani Tourn., pi. 71, figs. 55, 56.
This group, the type of which is a Miocene fossil of Dijon, is dis-
tinguished mainly by the tumid base, strong constriction behind the
lip, and coarse sculpture. Its affinities are problematic.
Section Pseudoleptaxis Pilsbry, 1894.
Shell solid, imperforate, globose, sculptured with oblique wrinkles ;
last whorl large, rounded, constricted behind the thickened, expanded
lip. Aperture lunate, oblique; columellar lip dilated. Type H.
corduensis (Noul.) Sandb., pi. 71, figs. 57, 58.
Perhaps to the Oligocene type of this group is to be added the
lower Miocene H. ramondi A. Braun, but that form may really be-
long to Plebecula.
Genus DENTELLOCARACOLUS Oppenheim, 1890.
Dentellocaracolus OPPENH., Denkschr. K. Akad. Wissensch. Ivii,
p. 117.
Shell imperforate or covered perforate, globose-conic, more or
less keeled, the base rather flattened; whorls 4 to 6i, the last
DENTELLOCARACOLUS. 295
suddenly descending in front a third or more the total alt. of shell.
Aperture oval or horse-shoe-shaped, horizontal ; margins thickened
and reflexed, joined by a callus. Type D. damnata A. Brong., pi.
71, figs. 53, 54.
This genus is established for certain heavy, rough sculptured
Helices from the N. Italian Eocene, characterized by the extreme
obliquity of the aperture, the heavy parietal callus, and the aspect
of the West Indian Pleurodonte formosa, or the Canary Island Hemi-
cyclas. Some Obba species are also similar. I am totally unable to
recognize any affinity between these shells and the Antillean Helices,
and regard the superficial resemblance as merely a case of converg-
ence of shell characters, meaningless from a phylogenetic stand-
point. Whether the group is to be referred to the Epiphallogona
or the Belogona is doubtful, and dogmatic assertions are clearly un-
called for.
The species damnata Brong., eoriacea Sandb., amblytropis Sdb.,
hyperbolica Sdb., antigone Oppenh. and mazzinicola Greg, belong
here.
Section Prothelidomus Oppenheim, 1890.
Prothelidomus OPPENH., t. c., p. 120.
Shell imperforate, solid, globose-depressed ; whorls 4J, the last
protracted and sometimes carinated toward the aperture. Aperture
horizontal, oval or horse-shoe-shaped ; peristome thickened, edent-
ulous, the margins joined by a strong parietal callus. Type P.
acrochordon Oppenh., pi. 71, figs. 51, 52.
This group is only feebly distinguished from the preceding, and
as with that, I am obliged to discredit entirely the relationships
implied by its name. It contains //. acrochordon Oppenh. (radula
Sandb. not Pfr.), and H. oppenheimi de Greg. (H. vicentina Op-
penh. not Shaur). The sculpture of the former is like that of Pleur-
odonte lima, but the second species is smooth. This shows how much
dependence is to be placed on a sculpture resemblance, a subject
discussed at more length in the introduction to this volume.
H. declivis Sandb., which Oppenheim places in Eurycratera, may
belong near or in this group or in Dentellocaracolus.
The peculiar minute form described by Stache as Obbinula an-
thracophila (Abh. K. K. Geol. Reichsanst. xiii, p. 119) from the
Stomatopsis Horizon, " Cretaceo-eoceiie " of Carniola, may prove to
296 HELICIGONA.
belong to the Helicidce, but even this is doubtful. It occurs in com-
pany with Stomatopsis, a peculiar genus of Melanopsidce, in beds
considered to lie at the base of the Eocene. The claim of Obbinula
to kinship with Obba is, there can be little doubt, an illusion.
Genus HELICIGONA Ferussac, 1819.
=Helicigona (FER.) Risso,-[- Chilotrema and Arianta Leach,
1831,4- Chilostoma, Latomusand Isognomostoma FITZ., 1833,-f-Oin-
guiifera, Corneola and Lenticula HELD, 1837,+ Campy Icea BECK,
1837,+Sterna ALB., 1850,+ Elona ADS., 1855, etc., etc.
Shell usually depressed-globose, varying to globose-turbinate or
lens-shaped, usually umbilicated, of moderate or large size; surface
smooth, costulate, granulate or hairy, corneous or brown ; unicolored
mottled or streaked, and either with a single supra-peripheral band
or with one above and one below this, or bandless. Aperture
oblique, lunate or oval, the lip expanded, reflexed below and dilated
at columellar insertion. Type H. lapicida L. (see pi. 43, figs. 19
25, 27, 28, 31-35, 42, 46).
Animal externally as in Helix. Jaw strong, with 2 to 16 stout
convex ribs, dentating the cutting edge. Radula as in Helix, the
ectocones sometimes developed on middle and lateral teeth, some-
times represented by lateral continuations of the mesocone.
Genitalia (pi. 62, all figs.) having the penis short, continued in
an epiphallus upon which the retractor is inserted, and ending in a
well developed, spirally twisted flagellum. Dart sack single, inserted
rather high on vagina, containing a curved dart with round shaft
and flat, 2-bladed head ; the base not coronated. Mucus glands 2,
long and tubular, often bifid ; inserted on vagina near base of dart
sack (but in H. quimperiana there are 3 triangular lobes on each
side). Spermatheca small and globose, on a long duct ; diverticulum
long, larger than spermatheca-duct, connected with the uterus
throughout by a broad thin membrane (shown in figs. 16, 18, re-
moved in the other figures on pi. 62). See pi. 62, fig. 16, H. rhce-
tiea; fig. 17, H. cingulata ; fig. 18, H. planospira; fig. 19, H. per-
sonata; figs. 20, 21, H. lapicida; figs. 22, 23, H. arbustorum ; figs.
24, 25, 26, 27, H. quimperiana).
Distribution ; Europe, from the Pyrenees and Greece to Sweden
and England. For geological distribution see under Chilostoma and
the extinct subgenera.
HELICIGONA. 297
The true limits of this genus were first indicated by Schmidt, and
have been confirmed by the researches of numerous later authors.
The two-bladed type of dart, the broad diverticulum bound to
uterus by a wide membrane traversed by blood-vessels, and the two
mucus glands, which are either simple and vermiform or once split,
are all characters unlike Helix or other genera of Belogona. The
shell is never five-banded as it is in Helix, but has either two bands
above, one below the periphery, or only the middle band is retained,
or it is bandless.
The form of the dart is not alone diagnostic, for Eremina and
some Iberus approach the two-bladed type. Helicigona is the only
genus known to me in which the diverticulum is a constant generic
character ; and here it seems to be much more highly developed
than in any other group.
The shell shows a considerable range of mutation in form and
sculpture, sometimes being acutely keeled as in H. banatica, canth-
ensis and lapieida, and again globose or subglobose as in arbustorum,
ehingensis and the Tacheocampylseas. In some forms, such as H.
lepidotricha and ehingensis, the margin of the umbilicus is obtusely
angular, as in some Chloritis ; and high authorities have indeed re-
ferred the former species to this genus. In my opinion such reference
is wholly uncalled for. I would as soon consider Lysinoe ghiesbre-
ghti or Epiphragmopliora remondi species of Chloritis on account of
the similar angled umbilicus and quincuncial sculpture. We must
have better evidence than these unstable and frequently repeated
characters, before admitting Chloritis to the European fauna. The
claim of Metafruticicola (p. 276) to a place in the genus Chloritis
is far better than that of H. lepidotricha and its allies. The variety
of sculpture found in Helicigona is equally remarkable ; some species
being quite smooth, others, as gobanziaud hemonica, heavily ribbed ;
and still other forms, such as lepidotricha, rahtii, setosa, benedicta
have bristles or their papillae arranged in regular quincunx or
oblique sweeps, besides a still more minute granulation of the whole
surface. In some species this regular sculpture extends to the very
apex, being exactly like that of Chloritis, Moellendorffia and a ew
other Helices of very diverse groups.
Synopsis of subgenera and sections.
RECENT GROUPS.
1. Aperture oval or lunate ; lip 1-toothed or toothless.
a. Spire convex or conoidal ; mucus glands tubular.
298 HELICIGONA.
b. Acutely keeled ; lip continuous across parietal wall.
HELICIGONA.
bb. Depressed, not keeled ; or if keeled the lip-ends remote,
Chilostoma, Fruticocampylcea.
bbb. Subglobose, with narrow or closed umbilicus.
c. Baso-columellar lip broadly reflexed, Tacheo campy Icea..
cc. Lip narrow, dilated only at insertion, Arianta.
aa. Spire flat, sunken in middle ; mucus glands short, triangular,
ELONA.
2. Aperture ear-shaped, 3-toothed, ISOGNOMOSTOMA.
TERTIARY GROUPS.
1. Shell with pappillse arranged in oblique series as in H. setosa,
a. An obtuse angle around umbilicus ; periphery rounded,
TROPIDOMPHALUS.
aa. Umbilicus narrow or closed ; periphery acutely keeled,
Metacampylcea.
2. Shell without regularly placed pappillse.
a. Large, depressed, with broad peristome, MESODONTOPSIS.
aa. Subglobose, spire conoidal, peristome narrow,
GALACTOCHILUS.
Section Helicigona (Fer.) Risso.
Helicigona FER. 1. c. (in part). Risso, Hist. Nat. Eur. Me>id. iv,
p. 66, first species H. lapicida. Caracolla TTJRTON, Man. L. and
Fr.-W. Sh. Brit. Is., 1831, p. 66, and of some other authors. Chilo-
trema LEACH in Turton, 1. c., p. 66, and of Beck, et al. Latomus
FITZ., Syst. Verz., 1833, p. 97. Lenticula HELD, Isis, 1837, p. 913.
Shell umbilicate, depressed, lens-shaped and acutely keeled ; surface
minutely granulous, horn-colored or dusky, obliquely streaked with
brown ; last whorl deeply deflexed below the keel ; aperture very
oblique, oval, angled at keel, the peristome reflexed below, thick-
ened within, continuous and raised across the parietal wall, toothless.
Type H. lapicida, pi. 43, figs. 22, 23.
Jaw with four strong ribs ; radula having no side cusps on middle
and lateral teeth ; marginals developing an ectocone, and the large
cusp become bifid. Geuitalia (pi. 62, figs. 20, 21, H. lapicida) with
penis, epiphallus and flagellum as usual. Two long mucus glands ;
dart-sack containing a curved dart with cylindrical shaft and short,.
HELICIGONA. 299
flat, two-edged head (fig. 21) ; spermatheca duct branching into a
diverticuluin about as iorig as itself.
This section contains a single species commonly distributed
throughout middle and northern Europe.
H. lapicida L. iv, 117.
v. medalpedensis Cl. iv, 261.
v. andorrica Bgt.
Section Chilostoma Fitziuger, 1833.
Chilostoma FITZ., Syst. Verz., 1833, for C. corneum (=H. cornea
Drap.), C. zonatum (=foetens Stud.) C. pulchellum (=pulchella
Miill., type of the prior genus Vallonia). CHARP., Cat. Moll. Terr,
et Fluv. Suisse, 1837, p. 8, for cingulata, zonata, fastens, pulchella.
GRAY, A List of the Genera of Recent Mollusca, their Synonyma
and Types, P. Z. S. 1847, p. 172 (type H.fcetens).M.oqv IN-TAN-
DON, Hist. Nat. Moll. Terr, et Fluv. France, ii, p. 131, for fcetens
and cornea. Campylcea BECK, Index Moll. 1837, p. 24. LOWE, P.
Z. S. 1854. ALBERS, Die Hel. 1850, p. 81. MARTENS, Die Hel.
1860, p. 122 (type H. cingulata Stud.). Cingulifera Held, Isis,
1837, p. 911, for ziegleri Schm., intermedia Fer., cingulata Stud., ar-
bustorum L., etc. Corneola HELD, ibid., p. 912, for hirta, feburiana r
setipila, planotspira. fattens, pulchella, etc., etc. Zoniies HARTM.,
Gastr. Schw., p. 161, not of Montf. Eucampylcea PFR., Noinencl.
Hel. Viv. 1878, p. 144. WESTERLUND, Fauna, p. 103.
Shell depressed, openly umbilicated, with convex spire and
rounded (rarely keeled) periphery. Surface unicoloied or 1-3
banded, smooth, costulate or hirsute. Whorls about 5?, the last
deflexed in front ; aperture wide lunate or suboval, toothless or with
a basal tooth; peristome narrowly expanded, reflexed below, dilated
at columellar insertion, rarely continuous across the parietal wall.
Type H. foetens Studer. (See pi. 43, figs. 27, 28, H. planospira
Lam. ; pi. 43, fig. 42, H. setosa Ziegler.)
Jaw strong, with 2 to 10 stout ribs grouped near the middle.
Radula with mesocones only developed on median and lateral teeth ;
marginals with an inclined bifid inner and small outer cusp. Geni-
talia as described for the genus.
In the recent fauna this group is characteristic of the Alpine
Mountain system, extending down the Italian peninsula to Sicily,
and the Balkan peninsula to southern Greece.' A few species occur
300 HELICIGONA.
in south-western France. It is therefore more southern in distribu-
tion than Arianta, Elona or Helicogona s. sir. In the lower mio-
cene deposits of north central Europe, a number of species typical in
form occur, such as H. inflexa Klein, H. exstincta Ramb., H. stand-
festi Penecke. Specimens of infllexa and standfesti before me retain
a distinct trace of the shoulder-band.
The disappearance of the name Campylcea from Helix nomen-
clature is to be regretted, and will probably fail to find many advo-
cates for some years to come. That the course here taken is inevi-
table, will be obvious if the history of the name is considered. Chilo-
stoma Fitz., 1833, and Campylcea Beck, 1837, were both proposed
without diagnosis, and both contained some incongruous elements.
If undefined names are to be rejected, then both of these must give
way to Oingulifera Held, proposed vith an excellent diagnosis in
1837. If, however, the list of species cited be accepted in lieu
of a diagnosis, then Chilostoma must\be accepted on the ground
of four years' priority, as Moquin-Tatidon has recognized. In
either case, Campylsea becomes a synonym! Those who continue
to use a generic or subgeneric name, which is so clearly inadmissable
as this one, must do so in defiance of rules of nomenclature recog-
nized as binding by zoologists generally, for Campylcea is neither the
earliest name for the group, nor the earliest properly defined name.
Neither is it the earliest properly limited group, for Beck's list con-
tains a number of species not belonging to this genus.
H. pouzolzi Desh., iv, 87. v. bosnensis Kob., iv, 88.
savignyana Ehrenb. ? ragusana Fer., undesc.
varronis Cantr. trizona Rve.
brenoensis & macarana Mhl. v. silvestris West.
dinarica Bgt. ! H. soccaliana Let.
? dalmatina Parr., dalmatica H. serbica Mlldff.,iv, 88.
Dh.,gravosaensis Muhl. /. roschiti (Kim.) W.
/.elevatior, depressior, bifasci- J. unitseniata Bttg., iv, 88.
ata Brus. ; unifasciata, uni- H. paucici Mlldff., iv, 88.
color Pfr., obscura Biz., H. banatica Partch, iv, 97.
kuzmici, pellanica, adriat- H. stenomphala Mke., iv, 88.
ica, tschernagorica, diocleti- H. setigera Zgl., iv, 100.
ana, sabljari, horatii, biagioi, /. globulosa Kucik.
brenoica, daniloi, cantrainei H. hoffmanni, Partch, iv, 99.
Bgt., viii, 231. monozona Z.
v. montenegrina Zgl., iv, 88. H. walteri Bttg., iv, 98.
HELICIGONA.
301
H. kleciachi Parr.,iv, 99.
klecaki Pfr.
H. insolita Zgl., iv, 98,
insolida auct.
subcostalis Parr.
H. prsetexta Parr., iv, 99.
prcetextata Kob.
H. narentina Klec., iv, 99.
v. reiseri Branc.
H. denudata Kossm., iv, 98.
H. imberbis Brus., iv, 97.
H. nicolai Klec., iv, 98.
recordera Parr.
H. trizona Zgl., iv, 108.
v. inflata Biz.
v. rumelica Z.
v. dobruschse Cless.
v. balcanica Friv.
v. frauenfeldi Zel.
H. hseterea West.
H. coerulens Mhl., iv, iii.
lacticini Z.
/. hyllica, depressa rugata,
bukowicanica,zrmanja3Brus.
H. cornea Drap.,iv, 110.
v. castanea Rm., iv, 111.
v. squamrnatina Serres.
H. desmoulinsi Far. iv, 111.
moulinsii P. & M.
aerosticha Fisch.
mollerati Morel,
v. crombezi Mill.
H. pterolakse Kob.
langi Pfr., Bttg.
H. phoca3a Roth., iv, 103.
/. ornata Parr.
/. inornata Kob.
v. langi Parr., iv, 102.
H. cingulata Stud., iv, 104.
luganensis Schintz.
v. inornata Rossm.
unicolor West.
rossmassleri Cl.
v. anauniensis de Bett., iv, 105.
v. athesina Paul., iv, 105.
/. pinii Adami, iv, 106.
v. baldensis Villa, iv, 105.
v. bizona Rossm., iv, 106.
v. lucensis Paul., iv, 106.
H. carrarensis Porro., iv, 105.
v. montana Paul., iv, 107.
v. kobeltiana Paul., iv, 107.
H. planospira Lam., iv, 89.
vittata Jan.
v. etrusca Kob.
v. stabilei Paul.
f. illasyaca Adami.
v. ullepitschi West.,iv, 90.
v. kobeltiana Cless., iv, 90.
v. illyrica Stab.,iv, 90.
v. padana Stab., iv, 90.
/. euganea Stab., iv, 91.
v. erjaveci Cless.
v. istriana Stoss.
v. pubescens Tib., iv, 91.
v. casertana Paul., iv, 91.
v. alifaensis Paul., iv, 92.
v. calva Kob.,iv, 91.
depilata Orsini.
v. setulosa Brig.
setipila Zgl.
setosa Costa.
setulosa Auct.
v. cantabrica Paul., iv, 91.
/. depressa Paul.
/. globosa Paul,
v. neapolitana Paul., iv, 91.
/. depressa Paul.
/. luteola Paul,
v. cassinensis Paul., iv, 91.
302
HELICIGONA.
v. occultata Paul,
v. pavelii Haz.
H. tiesenhauseni Gredl., viii, 227.
H. macrostoma Mhl.,iv, 92.
siculina Zgl.
pervia & didyma Mhl.
v. ereta Paul., iv, 92.
v. cryptozona Zgl.
v. confusa Ben., iv, 92.
H. benedicta Kob., iv, 92.
lefeburiana Phil.
setipila Benoit.
v. trichothroa Bgt.
v. choelotricha Bgt.
H. schlserotricha Bgt., iv, 96.
sclerotricha Auct.
H. hirtaMke.,iv,89.
deplana Zgl.
H. lefeburiana Fer., iv, 89.
feburiana Auct.
hirsuta Brumati.
H. sadleriana Zieg., iv,89.
H. mollendorffii Kob., iv,95.
H. hazayana Cless., iv, 89.
H. setosa Zgl., iv, 97.
/. convexior W.
/. litoralis Brus., iv, 97.
H. brusinse Stoss., iv, 98.
v. velebitana Klec.
H. crinita Sandri, iv, 100.
H. preslii Schm., iv, 104.
cingulata Held,
v. nisoria Rm., iv, 104.
intermedia Paul,
v. nicatis Costa iv, 101.
v. affinis Paul., iv, 106.
v. appelii Kob., iv, 105.
v. anconse Gent.,iv, 106.
v. agnata Paul., iv, 107.
v. amathia Bgt.
H. colubrina Jan., iv, 105.
v. nubila Zgl., iv, 106.
v. fascelina Z., Gred.
H. gobanzi Ffld.,iv,107.
v. sigela Bgt.
v. compsopleura B^t.
v. perfecta Bgt.
H. tigrina Ch. & Jan., iv, 107.
v. subtigrina Bgt.
H. frigida Jan., iv, 101.
/. insubrica Jan., iv, 101.
H. cingulellaZgl.,iv, 104.
zinguletta H. & A. Ad.
/. gyrata West.
/. scutellata West.
H. pyrenaica Dr., iv, 94.
v. complanata Bgt., iv, 95.
xanthelcea (B.) Fag.
v. semiclathrata West.
H. faustina Zgl., iv, 95.
v. sativa Z.
v. associata Z.
favirensis Parr.
v. citrinula Z.
v. charpentieri Schol.
v. fortunata Parr.
v. subflava Kim.
H. rossmassleri Pfr., iv, 96.
advena Rm., preoc.
v. bridayi Branc.
H. phalerata Zgl., iv, 100.
v. chamseleon Parr., iv, 101.
H. glacialis Thorn., iv, 109.
v. vesulana Less.
v. chiophila Bgt.
H. alpina F.-B., iv, 100.
v. alpicola West.
v. fontenilli Mich., iv, 100.
tigrina v. michaudiana Rm.
H. schmidti Zgl., iv, 103.
HELICIGONA.
303
H. hessei Kim., iv, 103.
H. hermesiana Pini, iv, 100. H.
v. frigidescens DelPrete, iv,105H.
v. apuana Iss., iv, 105. H.
v. ligurica Kob., iv, 101. H.
v. frigidissima Adami. H.
H. nicolisiana Ad., viii, 227.
H. intermedia Fer., iv, 1 09. H.
catenulata Muhl.
cornea Brum.
H. ziegleri Schm., iv,109.
H. semula Rossm., iv, 109.
ambrosia Strobel. H.
martinatiana de Betta. H.
H. ichthyomma Held, iv, 93.
v. achates Z., iv, 93. H.
cingulina Dh. H.
achatina P. & M.
foetens C. Pfr., iv. 93.
H. zonata Stud., iv, 92. H.
v. flavovirens D. & M. H.
v. monozonata Poll,
v. modesta Moq., iv, 92.
H. foetens Stud. H.
v. millieri Bgt., viii, 228. H.
H. strobeli Less.
H. cisalpina Stab., iv. 94.
gallica Bgt.
sebinensis Kob.
adelozona Parr,
v. debettai Ad. iv, 94.
v. adamii Kob., iv, 94.
v. rhsetica Mouss., iv, 94.
H. argentellei Kob., iv, 94.
H. peritricha Bttg., viii, 230.
v. erymanthia Kob.
kollari Zel.,iv,94.
hemonica Thiesse.
pindicaBttg.,iv, 96.
choristochila Bttg., iv, 102.-
gasparinse Charp., iv, 102.
v. subdeflexa Bttg., iv, 102.
olympica Roth, iv, 101.
thessalonica Mouss.
v. ossica Bttg.,iv, 102.
v. magnesise Bttg., iv, 102.
v. sciara West.
broemmei Kob., viii, 229.
conemenosi Bttg., viii, 229.
v. acarnanica Kob., viii, 220.
oetsea Mart.
subzonata Mouss., iv, 93.
v. distansBl. & W.,iv, 93.
v. depressa Bttg., viii, 228.
brenskei Bttg., iv, 113.
comephora Bgt., iv, 96.
comythophora Bttg.
/. kru'peri Bttg.
eliaca Kob.
cyclolabris Desh.,iv, 114.
v. euboea Parr,
v. arcadica Parr,
v. hymetti Mouss.
v. sphseriostoma Bgt.
lysistoma Shutt.
v. heldreichi Shutt.
v. amorgia West,
v. grelloisi Bgt., iv, 114.
v. bacchica Mart.
Section Fruticocampylcea Kobelt, 1871.
Fruiicocampylcea KOBELT, Catal. Eur. Binnenconch., p. 13.
Shell with moderate or small umbilicus, rather depressed, the sur-
face granulated or spirally striated, generally with a peripheral
304
HELICIGONA.
white band bordered above and below by dark bands ; aperture
oval, basal lip expanded. Type H. ravergiensis Fer. (pi. 43, figs.
24, 25 ; H. narzanemis Kryn.).
Anatomy unknown. This group of Campylsea-like shells is con-
fined to the Caucasus region and adjacent lands to the south. It
may prove to belong to the Hygromia series, but is better left here
until examined anatomically. The distribution of Frtiticoeampy-
Icea is not continuous with that of other Helicigonas.
H. appeliana Mouss., iv, 85.
appelinsi Auct.
v. mediata West.
H. narzanensis Kryn., iv, 84.
hortensis Menetr.
ossetinensis Bayer.
/. bicingulata Bttg.
/. castanea Bttg.
/. subunicolor Bttg.
/. perlineata Mouss.
v. suanetica Bttg.
v. macromphala Bttg.
v. cyclothyra Bttg.
v. olivacea Bttg.
v. kobensis Bttg.
v. depressa Bttg.
H. pratensis Pfr., iv, 85.
bayerii Parr.
/. unicolor Bttg.
/. alutacea West.
v. perforata West,
v. depressa Kob.
v. solidior Kob.
H. joannis Mort., iv, 86.
dumonti Mort.
H. dichrozoua Mart.
H. delabris Mouss., iv, 86.
/. alia West.
H. pontica Bttg., iv, 86.
H. nymphsea Dub.
H. ravergiensis Fer., iv, 85.
raver gii Kryn.
ravergieri Bttg.
limbata Kryu.
caucasica Pfr.
v. persica Bttg., iv, 85.
H. transcaucasica Bay., iv, 85.
/. pygmsea Bttg.
H. phseolema Bttg., iv. 87.
H. eichwaldi Pfr., iv, 86.
v. daghestana Parr., iv, 86.
H. armeniaca Pfr. iv, 86.
airumia Siem.
? nivalis Menetr.
? menetriesii Kalen.
Section Tacheocampylcea PfeifFer, 1877.
Tacheocampylcea PFR., Malak. Bl., xxiv, 1877, p. 8, type H. ras~
paili Payr. See for anatomy, MOQ.-TAND., pi. 12, f. 11-14.
Shell depressed with low spire, the body whorl not keeled ; imper-
forate or partly covered umbilicate ; smoothish, sometimes hairy ;
brownish, yellowish or olive, with two bands above, one below the
periphery. Aperture truncate-oblong, very oblique ; outer lip
HELICIGONA. 305
reflexed, baso-columellar lip straightened or arcuate, its edge broadly
dilated and reflexed. Type H. raspailii, pi. 43, figs. 33, 34, 35.
Jaw (of H. raspaili) arched, with three separated ribs. Epi-
phragm flat, thin and membranous, with some calcareous particles.
Genital system furnished with a curved dart (pi. 6S, fig. 7 H. ras-
pailii) 10-12 mill, long, swollen and channelled at base, then con-
stricted, widening again in the middle, four bladed (?). Mucus
glands four-fingered.
This group has the characteristic shell of Campylcea, but the four-
fingered mucus glands and the apparently quadrangular dart are
characters like Tachea and Otala. It may prove to be a transition
group. A further investigation is needed to demonstrate its affini-
ties, and especially should the dart and the diverticulum of the
spermatheca duct be examined, as these structures afford the only
criterion for the separation of the true Helices from the Campylcea
or tielicigona group. The species are all from Sardinia and Cor-
sica.
H. raspailii Payr., iv, 112. H. cyrniaca Dut., iv, 112.
v. acropachia Mab. revelierii Deb., iv, 112.
v. lenelaia Mab. planospira Payr.
v. pilosa Kob., iv, 112. tachigyra West.
v. garciai Hagenm. v. montigena Hagen.
H. insularis Cr. & Deb., iv, 112. v. faucicola Hagen.
H. brocardiana Dut., iv, 112. H. gennarii Paul., iv, 113.
v. omphalophora Dut. H. carotii Paul., iv, 112.
v. sciaphila Hagenm. /. major, unifasciata, viperina,
v. donata Hagenm. lamarmorse, spectrum, Mal-
zan, iv, 113.
H. melonii Malz., iv, 113.
Unfigured forms : H. vittalacciana Mab., romagnolii Dut., melli-
niana with var. deschampsiana, and arusalensis Hagenmiiller.
Section Arianta Leach, 1831.
Arianta LEACH in Turton's L. and Fw. Shells Brit. Is., p. 35-
(for H. arbustorum). BECK, Index, p. 41 (in part). HARTMANN,.
Gast. Schw., p. 55. Arionta v. MARTENS, Die Hel., 1860, p. 127,
(exclusive of all but type, H. arbustoruvi). Not Arionta of Ameri-
can authors !
20
tJNIVlBSITY
V e-, "_
306 HELICIGONA.
Shell globose or globose-depressed, with convex or conoidal spire,
and narrow or closed umbilicus ; surface shining, spirally striated,
usually with a supraperipheral band, and mottled or dark coloring.
Aperture round-lunate, oblique, toothless ; lip expanded and white-
lipped, reflexed at columellar insertion. Type H. arbustorum L.,
pi. 43, fig. 46.
Jaw with 6-10 strong ribs. Radula having outer side cusps
developed on middle and lateral teeth. Marginals with bifid inner
and simple outer cusps. Genital system (pi. 62, figs. 22, 23, H.
arbustorum} showing the features usual in Helicigona throughout.
The two simple mucus glands are very long; dart sack containing
a curved dart (fig. 22) like that of H. lapicida. Diverticulum bound
to uterus by a wide membrane traversed by blood vessels.
Distribution, middle and north Europe, upper Pliocene and Loess
deposits. In the modern fauna this species or group of species, is
distributed from the northern boundary of the Olive zone (Pyre-
nees and Alps) to Sweden, enjoying a far greater range than any
-other member of the genus Helicigona, especially in its ability to
withstand the cold. The number of local races is remarkable, and
ptheir study is much complicated by the fact that forms with a simi-
lar aspect occur in widely separated localities, probably due to par-
allel development. Typically many of these varieties are very
different, but intermediate forms seem to abolish most boundary
Klines ; so that Kobelt, in his latest contribution on the subject, is not
willing to endorse even the main forms as species (Iconogr. n; F.,
vi, p. 60).
The spelling of this name given above is that of Leach, Beck
and other early authors. Von Martens has changed the name to
" Arionta" on etymological grounds. The single well defined spe-
cies H. arbustorum, is a typical Helicigona in anatomy, having the
diverticulum bound to the uterus by a broad membrane, the two
long, cylindrical mucus glands inserted on vagina, and other feat-
ures diagnostic o that genus. The American species referred to
Arionta by authors present nothing of the sort ; the diverticulum has
#0 membrane ; the mucus glands are bulbiferous and inserted on
xiart sack, etc., etc.
H. arbustorum L., iv, 117. v. picea Zgl., iv, 117.
v. conoidea West., iv, 118. wittmanni Zow.
vv. calcarea Hogb., iv, 118. v. jetschini Ulic., iv, 117.
HELICIGONA. 307
H. arbustorum Linne. subalpina Scholtz.
v. canigonensis Boub. /. costulata Kob.
canigoniea Fag. v. dorise Paul., iv, 117.
v. fagoti Bgt. v. rudis Miihlf., iv, 118.
v. xatarti Far., iv, 118. v. corneoliformisLess.
v. repellini Charp., iv, 118. v. styriaca Ffld., iv, 119.
v. alpicolaFer. H. sethiops Biz., iv, 118.
tris Z., iv, 118. H. camprodunica Kob., iv, 118.
Other named forms of H. arbustorum, some of which have doubt-
less good claims to racial distinction, are: Var, thamnivaga, hypni-
cola, themita Mabille ; var. dravica, vibraiana, musdorfensis, illusana
Servain ; var. sendtneri, excelsa, septentrionalis Clessin ; var. albu-
lana, feroeli, knitteli, nazarina, trachia (Bgt.) Serv. ; var. creticola
Morch. ; var. trochoidalis Roffisen ; var. depressa Held. ; var. baylei
(Lecoq) Moq. ; var. gotlandica; oelandica West. Also forma
flavescens, albina, rufescens, draparnaudia, poiretia, boissieria,
thomasia Moquin-Tandon ; /. ef'asciata Westerlund ; /. lutescens,
luteofasciata, fuscesens D. & M. (=marmorata Taylor) ; /. mor-
bosoalbina Rossm. ; /. nigrescens Locard, /. fusca Fer. ; /. cincta
(=pallida Tayl.), sinistrorsum Taylor; /. minima and major Pfr.
Subgenus ELONA H. & A. Adams, 1855.
Elona H. & A. AD., Gen. Rec. Moll, ii, p. 211, type H. quimperi-
ana (June, 1855). Not Elona Moq.-Tand., 1855. Sterna ALBERS,
Die Hel., p. 93, 1850, same type (preoccupied). See HESSE, Jahrb.
D. M. Ges., xii, 1885, p. 45, pi. 3, f. 1 (anatomy).
Shell um\)i\icsite, planorboid, the spire slightly concave, periphery
broadly rounded ; corneous with a few varicoid white stripes ;
aperture lunar, slightly oblique ; lip white, expanded above, reflexed
below, the ends distant. Type H. quimperiana Fer., pi. 43, figs. 19,
20, 21.
Jaw with 11-16 narrow ribs. Genitalia (pi. 62, figs. 24, 25, 26,
27, H. quimperiana) differing from the typical Helicigonas in hav-
ing the mucus glands shortened into triangular sacks (fig. 26) and
the dart sack is inserted in a sort of calyx at base (fig. 27). Dart
curved at the end, with lens-like section (fig. 24).
This group contains a single French species remarkable for its
{Jhloritis-like shell and the peculiar mucus glands. The latter con-
308 HELICIGONA.
sist of short lobes, somewhat as in Eulota ; but unlike that genus,,
they are inserted on vagina well above the dart sack, so there can be
no doubt that they are merely a shortened form of the finger-like
glands characteristic of Belogona Siphonadenia.
H. quimperiana Fer., iv, 116. Brittany; Spain.
kermorvcmi Coll.
corisopitensis Dh.
Subgenus ISOGNOMOSTOMA Fitzinger, 1833.
Isognomostoma FITZ., Syst. Verzeichniss der in Erzherzogthume
Oesterreich vorkommenden Weichthiere, p. 97, sole species, /. per-
sonatum Eitz., =H. personata Drap. Isognomonostoma TRYON
Triodopsis of modern European authors, not of Rafinesque!
Plicostoma SCHLUTER, Syst. Verz., 4, 1838. See for anatomy Schu-
berth, Archiv f. Naturg., 1892, p. 11, pi. 1, f. 15-18 (Good!)
Shell depressed-globose, with low convex spire, narrow or closed
umbilicus and rounded or faintly angular periphery ; surface smooth,
hirsute in quincuncial pattern. Aperture oblique, ear shaped ;
peristome flatly reflexed, thickened within, toothed on outer and basal
margins; terminations connected by a raised, tongue- like parietal
process. Type H. personata, pi. 43, figs. 31, 32.
Jaw with about 5 strong ribs, dentating the cutting margin ;
radula with large triangular mesocones on middle and inner lateral
teeth ; marginals with a bifid inner and simple outer cusp. Genital
system (pi. 62, fig. 19, H. personata) with penis as usual in the genus;
2 long mucus glands ; an elongated dart sack containing a dart
of typical Helicigone form, base dilated, shaft slender and round,,
spreading into a two-bladed, laurel-leaf shaped head. Diverticulum
bound to uterus by a broad membrane, as usual in Helicigona.
Distribution, middle Europe and Siberia.
The anatomy of this group is typical of Helicigona (Campylcea),
having the diverticulum membrane found in that genus only, (re-
moved in the figure), as well as the characteristic form of mucus
glands and dart. These features of the genital system, as well as
the strongly ribbed jaw, show that the association of H. personata
with " Gonostoma " is entirely illusory. In fact Schuberth, in his
anatomical characterization of "Anchistoma" (" Gonostoma " +
"Triodopsis") was obliged to make an exception of H. personata,.
HELICIGONA. 309
and to compare it with Campylcea; but strangely enough he does
not alter the current classification of the species. The resemblance
of H. personata to the American Triodopsis is merely a case of
incomplete parallelism. The two groups are readily separated by
observing the form of the parietal barrier. Dr. H. von Ihering has
ably discussed the relationships of H. personata, ranking it, of
course, in Campylcea.
H. personata Drap., iii, 147. H. subpersonata Midd., iii, 147.
isognomostomos Gm. pt.
v. debilis West.
Subgenus TROPIDOMPHALUS Pilsbry, 1894.
Shell with the general characters of Chilostoma, but subangular
^around the umbilicus, and quincuncially punctate or papillate (as
in some members of the H. planospira group). Type H. lepidotri-
cha A. Braun, pi. 71, figs. 59, 60.
The lower Miocene forms for which this section is proposed have
the verge of the umbilicus subangular as in most (but not all)
Chloritis (con/, p. 118) and many species of Eulota; and in fact
the group may belong to Eulota rather than to Helicigona. At all
events, the closest resemblance is traceable between H. lepidotricha
and certain southeast Asian Eulotas. On theoretical grounds, how-
ever, I am disposed to believe that Eulota has no extensive past
history in Europe, being a recent straggler from East Asia ; and
this is supported in the main by palseontological evidence.
A thorough study of the Miocene Helices is necesssary to deter-
mine whether the peculiar sculpture which occurs in so many forms,
is a character assumed simultaneously by many subgenera and
genera, or an indication of actual genetic relationship. Not much
evidence can be adduced in favor of the latter view from the recent
fauna, for species of widely different genera exhibit the hairs or
papillae arranged in obliquely decussating series : In HYGROMIA,
H. consona, lanuginosa, etc.; in HELICIGONA, hairy members of the
planospira group; in THYSANOPHORA, T. stigmatica and its allies ;
in EULOTA, numerous oriental species. The list could be indefi-
nitely increased. It will be perceived from this that those authors
who insist upon the presence of Chloritis in the European Miocene
fauna, stand upon narrow and insecure footing.
H. robusta and trichophora Reuss., from the lower Miocene of
Tuchoric, evidently belong to this group.
310 HELICIGONA.
Section Metacampylcea Pilsbry, 1894.
Shell solid, sublenticular, acutely keeled, the spire obtuse-conic.
Aperture oblique, subrhombic ; outer and basal lips reflexed, thick-
ened within, the columellar insertion dilated, partly or wholly clos-
ing the narrow umbilicus. Surface minutely granulate, and with
larger papillae disposed in quincuncial order. Type H. rahtii A.
Braun, pi. 71, figs. 45, 46.
In its acute carination, the lower Miocene H. rahtii is comparable
to H. lapicida or banatica of the recent fauna, but its sculpture is
that of H. setosa Zgl. The lip differs somewhat from that of any
living " Campylsea," but not more than various species of that
group differs from one another. Metacampylcea probably stands in
much the same relation to Tropidomphalus as Helicigona (lapicida}
does to Chilostoma (planospira, etc.).
H. papillifera Klika and possibly H. obtusecarinata Sandb., are to
be referred here, but the latter may belong to the ancestral Tachea
stock.
Subgenus GALACTOCHILUS Sandberger.
Galactochilus SANDB., Land und Siisswasser Conchyl. der Vor-
welt, p. 387 (for H. pomiformis, mattiaca, ehingensis and cornumili-
tare).
Shell subglobose, with low, conoid spire of about 4J whorls, the
last large with rounded periphery, subangular around the narrow,
partly or nearly closed umbilicus, slowly descending in front.
Aperture truncate-oblong, oblique ; lip obtuse, expanded on outer
and basal margins, dilated at columellar insertion, partly closing
the umbilicus. Surface smooth except for growth-strise. Type H.
ehingensis Klein, pi. 71, figs. 47, 48.
This group contains several species from the lower Miocene, H*
pomiformis A. Braun, Ehingensis Kl., mattiaca Stein. I am dis-
posed to believe it an off-shoot from the " Campylsea " phylum.
Some specimens of H. arbustorum exhibit much the same subangu-
lation around the umbilical region.
Subgenus MESODONTOPSIS Pilsbry, 1894.
Shell large, depressed, with convex and very obtuse spire and
covered umbilicus. Whorls 5, convex, the last ornamented with two
CYRTOCHJLUS-HELIX. 311
broad bands above and one below the periphery, deflexed in front.
Aperture half round, oblique ; lip broadly reflexed throughout,
dilated and adherent at the columellar insertion. Surface smooth-
ish. Type H. chaixii Mich., pi. 71, figs. 61, 62.
This group differs from the pentateeniate Helices in having the
lip more reflexed, and not forming a columellar plate. I think
it allied more to the " Campylseas," with which it agrees in the color
pattern (distinctly visible in specimens before me) and the general
features of the aperture. I consider Tacheocampylcea the most
nearly allied group of the recent fauna (conf.pl. 43, figs. 3-35).
The resemblance to Meaodon is merely superficial. It is likely that
H. brocchii Mayer from the upper Pliocene belongs here rather than
to Galactochilus. It is umbilicate and one-banded above ; but I
have not seen that species, nor H. ludovici Noul. and ornezanensis
Noul. from the Miocene freshwater chalk of southwestern France,
which may also find a place in this group. The type, H. chaixii, is
from the middle Pliocene of Hauterive.
Genus (?) CYRTOCHILUS Sandberger.
Cyrtochilus SANDB., Land u. Siisswasser Conchyl. der Vorwelt, p.
386 (for H. expansilabrisSandb.'). Not Cyrtochilus Jak., 1875, or
Meek, 1876, nor Cyrtochila Feld, 1874.
Shell globose-conoid, with 5 convex whorls separated by linear
sutures, the last whorl large, ventricose, broadly constricted behind
the lip; surface of all but first whorl obliquely costulate and deco-
rated with minute papillae arranged in quincuucial order. Aperture
oblique; outer and basal lips expanded, columella narrow, vertical,
closing the umbilicus. Type H. expansilabris Sandb., pi. 71, fig.
50.
The shell has the figure of H. platychela of Sicily, but it is
sculptured like a hairy Chilostoma. The single species is from the
lower Miocene of Hochheim.
Genus HELIX Linne, 1758.
Helix LINNE (in part), Syst. Nat., x, p. 768. LAM., Syst. Anim.
s. Vert., 1801, p. 94, H. pomatia on\y. -\-Pomatia, Tachea, Otala,
Macularia, Iberus, JEremina, Eaparypha, Hemicycla, etc., etc.
See for anatomy A. SCHMIDT, Der Geschlechtsapparat der Sty-
lommatophoren in taxonomischer Hinsicht, in Abhandl. naturwis-
312 HELIX.
genschaftl. Vereins fiir Sachsen u. Thiiringen in Halle, i, pp. 1-52,
pi. 1-5, 1856, and Zeitschr. f. Malak., 1850, vii, p. 1-13, pi. 1
(darts); Ibid, 1849, p. 49. C. BRANCSIK, Sexualapparate einiger
Moll, des Trencsiner Comitates in Jahresheft des naturw. Vereins
der Trenc. Com., Trencsin, 1890, p. 19-22, pi. 1-3. R. LEHMANN,
Die lebenden Schn. u. Musch. der Umgebung Stettins u. in Pom-
meru,1873. MoQ.-TAND.,Hist. Nat. Moll. Terr, et Fluv. Fr., 1855.
O. SCHUBERTH. Beitr. zur Vergleich. Anat. des Genitalapparates
von Helix, in Arch. f. Naturg. Iviii, i, 1892, p. 1-65, pi. 1-6. POL-
LONERA, Bull, della Soc. Mai. Ital. xii, 1885, p. Ill (best figs, of
dentition). ERDL, in Moritz Wagner's Reisenin der Regentschaft
Algier, 1836. PAASCH, Archiv f. Naturg., 1843 and 1845. F,
WiEGMANN,Jahrb. d. m.Ges.iv, 1877, p. 195, pi. 6-8. BAUDELOT.
Ann. Sc. Nat. (4), Zool. xix, 1863. ASHFORD, Journ of Conch.,
Leeds, Vol. iv, 1883-'85. v. IHERING, Morph. u. Syst. des Genital-
apparates von Helix, Zeitschr. f. Wissenschaftl. Zool. liv, 1892, p.
386-520. C. F. JICKELT, Fauna der Land u. Susswasser Moll.
Nord-Ost-Afrika's, in Nova Acta Acad. Cses. Leop.-Carol. Germ.
Nat. Cur. xxxvii, 1875, et al. See for palaeontology of Helix:
SANDBERGER, Land- u. Siisswasser-Conchyl. der Vorvvelt, with the
authorities cited therein ; KLIKA, Tert. Land- und Siisswasser-Conch.
N. W. Bohmen, (cf. BTTG., Verb. K.-K. Geol. Reichsanst., 1891, p.
228) ; PENECKE, Zeitschr. D. geol. Ges. xliii, p. 346 ; OPPENHEIM,
Denkschr. k. Akad. Wissensch. Ivii, p. 113 (cf. TAUSCH, Verb. K.-
K. Geol- Reichsanst., 1891, p. 198, and de GREGORIO, Ann. deGeol.
et de Paleont. 10 e Livr., 1892), etc., etc.
Shell varying from globular to depressed and from rounded to
acutely keeled, imperforate or narrowly umbilicated, rather solid,
with about 5 whorls ; surface striate, ribbed, malleated or granu'
lose. Five-banded, or having fewer or no bands by the absence or
coalesence of some or all ; rarely having more bands by splitting of
bands or interpolation of lines. Lip either expanded, reflexed or
thickened within. Type H. pomatia, frontispiece, fig. 7, (See pi-
44).
Animal with a tough, granulose or reticulate integument, marked
by two or few grooves along back, the tail depressed, with a slight
median line or none', facial grooves well developed on both sides.
Labial processes large; sole undivided. Mantle with a small right
body lappet, and a long left one, usually interrupted across the
HELIX. 313
back. Right eye retractor passing between primary branches of
genitalia. (Frontispiece, fig. 7).
Jaw well arched, stout, with 3-9 strong ribs denticulating both
margins (pi. 67, figs. 1, 4, 7, 8, 9). Radula normal, having the
cusps of median and lateral teeth about as long as the squarish
basal plates, side cusps small or wanting. Marginals with a long,
oblique bifid inner cusp and a small simple or bifid ectocone (pi. 67,
figs. 2, 3, 5, 6, 11).
Genital system characterized by a short penis passing into the epi-
phallus, which bears the retractor (distally inserted on lung floor)
and branches into vas deferens and a flagellum, the latter rarely
wanting. One dart sack present and well developed, containing a
four-bladed dart, with short neck and crenulated base. Mucus
glands two, varying from simple to multifid, but always composed of
smooth, tubular coeca ; inserted on each side of vagina immediately
above entrance of dart sack; both the dart sack and mucus glands
lying free in cavity, not bound together by a stout membrane. Sper-
matheca globose, on a long duct, which usually bears a diverticulum.
Ovotestis compact, imbedded in the side of the liver (frontispiece
figs. 5, 6, H. pomatia).
Uistribution, Europe, North Africa, Asia Minor.
Helix is distinguished from Helicigona mainly by the form of the
dart and the free diverticulum; this being invariably bound to
oviduct by a wide membrane in Helicigona.
The genus Helix contains the most highly organized and complex
snails of the family Helicidce. Like the European type of Homo,
but unlike most highly specialized forms, their specialization has
evidently fitted them for meeting widely diverse conditions of exist-
ence. Their powers of reproduction as well as the ease with
which they adapt themselves to circumstances of climate and envi-
ronment new to them, are remarkable. They love the light, and
for the most part are not exterminated by the destruction of their
native forests, but accepting kindly the new conditions, live and
multiply in vineyards, gardens and around tilled fields. As colo-
nists they rank with man, the dog and the horse. Various species
live and thrive in the United States, Mexico, Cuba, Argentina, S.
Africa, New Caledonia, Australia, etc., etc. None, even of the most
widely distributed Helicoids of other genera such as Eulota similar is,
have so wide a range of climate ; and the species of Helix which
314 HELIX.
have founded colonies in climates foreign to them, outnumber the*
colonized members of all other Helicoid genera together.
The causes of this adaptability are obscure. Perhaps the rather
unusual toughness of the external integument and the unrivalled
complication of the genitalia are factors of importance, the first
allowing them a wider range of station with greater variety and
opportunity of feeding, the second producing more perfect eggs. It
is noteworthy that the dentition is of a very generalized type, show-
ing no tendency toward the specialization seen in the radulre of"
Polymita, Oxychona, Papuina, or the entire series of genera group-
ing around Acavus, Helicophanta and Panda. Such high modifica-
tion of dentition as these genera show, must restrict them to the
special conditions and food which produced it, and would constitute
a bar to their wide dispersal, which is not present in the genus
Helix. The jaw is of high type, but the same efficient odontogna-
thous form has been developed in many genera.
With the exception of Euparypha and Eremina, no divisions of
Helix can be based upon anatomical characters, for the features
intergrade throughout, offering merely specific differences. The
various " sections " of the genus rest wholly upon conchological char-
acters, which though quite appreciable to the eye, are often extremely
difficult to define in words so that they may be distinguished.
The genus Helix is abundantly represented in the Tertiary depos-
its of middle Europe, by species belonging without doubt to the
modern groups, although in many cases they are practically inter-
mediate between some of the latter. The HELICOGENA or Pomatia
group is not known with certainty below Pleistocene deposits,
although it is barely possible that the Oligocene H. globosa Sowb.
belongs here. I do not think this likely ; and the evidence at hand
indicates that the group arose upon non-European soil, and spread
northward or northwest in a few specific forms which have split in
comparatively recent times into numerous species. TACHEA, how-
ever, has an extensive range in time, a considerable number of forms
appearing in lower Miocene deposits, some showing certain features or
Iberus, others with more conoidal spire than usual in normal recent
Tacheas, but still having the characteristic columella and band pat-
tern. H. bohemica, H. moguntina and H. hortulana are examples*
being the " Coryda " of some European authors, so-called on account
of the trifling incident of a raised spire. Such forms as H. crepidos-
toma Sandb., with keeled earlier whorls, are also to be regarded as.
HELIX. 315-
a manifestation of this group. In the upper Miocene, H. sylvana>.
sylvestrina, etc., represent this group. In late Pliocene and Post-
Pliocene times, Tachea was represented by numerous forms, such as
sepulta Mich., tonnensis Sandb., and those described by Nevill from
Mentone. The section OTALA (Macularia Auct.) has a similar his-
tory, appearing at about the same time, in moderately characteristic
forms, many with the malleation of the recent species, as seen in
H. nayliesi, etc. HEMICYCLA, now confined to the Canary Islands,,
seerns to have had a wide range in the Miocene, some species, such
as asperula Dh. being excessively similar to recent forms. The iso-
lation of the Canaries has evidently preserved there this ancient
race. There are a number of Tertiary forms of Europe known to
me by figures or poorly preserved specimens only, which will event-
ually no doubt form new groups.
jET. doderleiniana All. of the Sicilian Pliocene seems to represent
a section distinct from Otala, although allied to that group, which
may be called Allolcemus. It is distinguished by the extraordinary
expansion of the last whorl toward the aperture, after a wide shal-
low constriction, causing the throat to be quite narrow, although the
mouth is expanded and the outer lip flaring. General form globose-
depressed. The specimen before me is from Palermo, collected by
Benoit.
It has not been considered necessary to give varietal names to the
band variations of these five-banded Helices. They may better be
expressed by the well known formula originated by Martens pere
(Ueber die Ordnung der Bander an den Schalen mehrerer Land-
schnecken, 1832), and explained in the Introduction to this vol-
ume.
Synopsis of sections and subgenera.
I. Penis provided with a flagellum.
a. Baso-columellarlip straightened and widened by a callous plate
within.
b. Im perforate, globose-conic, periphery round, smoothish ;
usually yellow or white, banded Tachea..
bb. Imperforate, globose-depressed, periphery round, smooth
or malleated, solid ; uniform, or speckled and banded
Otala.
bbb. Globose-depressed, malleated, ribbed or granulate, dark,
usually banded Hemicycla..
316 HELIX.
bbbb. Globose or depressed, smooth or striate, with spotted
bands Iberus (in part).
aa. Baso-columellar lip concave, not wide or flat.
b. Large, globular ; lip simple or expanded Helicogena.
bb. Depressed, often keeled, ribbed or striate, 0-4 banded
Iberus (part) ; Levantina.
II. Penis without flagellum ; shell chalky.
a. Globose-depressed, heavy, the lip expanded or thickened ; bands
few or none Eremina.
aa. Globose or depressed, decussated above, the outer lip not in the
least expanded, thickened within ; bands many, rarely none
Euparypha.
Section Helicogena Ferussac, 1819.
Helicogena (part) FERUSSAC, Tab. Syst. Fam. Limagons, p. 27.
Risso, Hist. Nat. Eur. Merid., p. 60, first species H. pomatia.
CHARPENTIER, Cat. Moll. Suisse, 1837, p. 5, for H. pomatia only.
HARTMANN, Gastr. Schw., p. 98 (for H. pomatia). Moq.-Tand.,
Hist. Nat. Moll. Fr. ii, p. 179. Pomatia Leach, in TURTON'S Man-
ual of the Land and Freshwater Shells of the Brit. Is., 1831, p. 45.
BECK, Index Moll., p. 43, and of authors generally. f Lucena,
HARTMANN in Syst. Erd- u. Siisswasser Gastr. Eur., p. 40, 1821.
Pomacea PERRY, Conchology, pi. 38, 1811 (in part; but diagnosis
agrees better with Ampullaria species also included).
Cantareus Risso, Hist. Nat. Eur. Merid., p. 64, (Helix naticoides
sole species). MOQ.-TAND., I c., p. 186. Canthareus AGASSIZ,
Nomencl. Zool., 1847. Tapada GRAY, in Turton's Manual L. and
Frw. Sh. Brit. Is. edit. 1840, p. 127, H. aperta sole species.
Cryptomphalus Agassiz in CHARPENTIER, Catal. Moll. Terrest. et
Fluv. de la Suisse, in Neue Denkschriften der allg. Schweizerischen
Gesellsch. fiir die gesammten Naturwissensch. ( Nouveaux Mem-
oires de la Societe Helvetique des Sci. Nat.) i, 1837, p. 5, for arbus-
torum, aspersa, rylvatica, nemoralis. MOQ.-TAND., 1. c. p. 174,
restricted to H. aspersa. Ccenatoria HELD, Isis, 1837, p. 910, for
aspersa, lucana, lutescens, pomatia, etc. Erctella MONTS., Naturalista
Siciliano xiii, No. 9, June, 1894, for H. mazzullii.
Shell globose or globose-conoid, capacious, with about 4* rapidly
widening whorls ; umbilicus narrow or closed ; aperture large, not
HELIX. 317
very oblique, lunate ; outer lip simple or expanded, columella long,,
concave, not thickened within, its edge reflexed. Type H. pomatia
L., Frontispiece, fig. 7; (see also pi. 44, figs. 6, 7, H. asemnisvar.
vetusta).
Animal large with wide fleshy foot, the sole undivided ; above
coarsely gran ose- reticulate ; right and left facial grooves strongly
marked ; labial tentacles well developed ; mantle edge bearing a
bluntly rounded right body lappet and a similar left one, the latter
either continuous or interrupted over the back, but reappearing in
a broad rounded lobe on the left side. Top of tail rounded, with a
subobsolete median line or none ; back from mantle to head with a
pair of longitudinal grooves. (Frontispiece, fig. 7, If. pomatia). H.
aperta, H. aspersa and other species examined show the same exter-
nal characters.
Jaw strong, arcuate, with several stout ribs denticulating both
margins. Radula (pi. 67, fig. 11, H. pomatia) with well developed
ectocones on central and lateral teeth ; marginals with bifid inner
and small simple outer cusps.
Genitalia: Penis short, passing into a short epiphallus upon
which the retractor is inserted, the epiphallus ending in a long flagel-
lum and vas deferens. Dart sack unusually large, containing a
four-bladed dart, the blades long, separated from the coronated base
by a neck or constriction (pi. 61, fig. 11, aspersa; fig. 15, pomatia).
Mucus glands usually divided into several branches subdividing
into very numerous fingers, but sometimes (H. aperta, asemnis) the
number is as low as four or five. Seminal receptacle globular, on a
long, nearly straight duct, which usually bears a diverticulum.
Hermaphrodite duct much knotted ; hermaphrodite gland compact.
(Frontispiece, figs. 5, 6, H. pomatia, Oberau, Bavarian Tyrol. PI.
61, figs. 12, 15, H. pomatia. PI. 61, fig. 9, H. asemnis. PI. 61, figs.
13, 14, 11, H. aspersa).
The typical Helicogenas agree with the types of Otala and with
Tachea vindobonensis in having the mucus glands divided and sub-
divided into many "fingers"; but as in Otala and Tachea, this is
an inconstant feature, the number being 4 or 5 in some species.
The main character of the group is its globose shell, and this offers
a perfectly tangible sectional feature. The dart sack is larger than
in the related sections except Tachea. There is usually a diverticu-
. lum developed on spermatheca duct, but in H. pomatia and some
other species it is generally absent.
318 HELIX.
It has been considered best to revert to Ferussac's name Helico-
gena for this group, as the well known name Pomatia must in any
<jase be abandoned in favor of Cantareus, properly proposed five
years earlier. Pomatia, moreover, is etymologically identical with
Pomatias, the name referring to the calcareous epiphragm, and not
of latin derivation as some have supposed. Strictly speaking, no
sectional name is required for this section, as it is the typical group
of Helix.
The species are European in distribution, but the greater number
occur in Southern Europe, Northern Africa and Asia Minor. H.
aspersa is the most widely dispersed, and has become colonized in
many localities in both North and South America, as well as in
Australia, etc. Most species of this group are edible and used for
food in the latin countries as well as in Turkey, Greece and the
Orient.
{I. Imperforate, solid, malleated,lip expanded throughout; epiphragm
membranous, CRYPTOMPHALUS).
H. aspersa Mull., iv, 235. H. aspersa.
grisea Gmel. secunda Da C.
variegata Gmel. fluminensis Lang.
Thin, plicate or striate, lip hardly expanded, Eretella.
H. mazzullii Jan., iv, 235. H. vermiculosa Morel, iv, 149.
crispata Costa not Fer. /. cretacea Westerl.
retiragis Mke. (undescr.). H. subaperta Ancey.
rugosa Mu'hlf. mazzulopsis Anc., viii, 238.
costce Ben. H. subplicata Sowb., iv, 236.
/. zonata Bgt. H. tristis Pfr., iv, 254.
v. quincayensis (Maud.) Bgt. cerasina Sh.
quincianensis Mauduyt. H. aggerivaga Mab., iv, 255.
II. Imperforate or umbilicated, the lip hardly expanded; epiphragm
rigid, calcareous, HELICOGENA.
Thin, globular, imperforate and unicolored, with large aperture and
dark coloration, Cantareus.
H aperta Born, iv, 254. Southern France to Greece, N.
terrestris Forsk. Africa.
neritoides Ch. v. korsegselia Bgt., Loc.
naticoides Drap. v. kalaritana Prunn., Villa.
HELIX. 319
Shell strong, often umbilicate, usually banded. Helicogena.
H. pomatia L., iv, 236.
antiquorum Lch.
pomaria & sealaris Miill.
v. gesneri Hartm., iv, 237.
pyrgia Bgt.
v. rustica Hartm., iv, 237.
radiata Ulic., iv, 238.
v. pulskyana Haz., iv, 237.
v. sabulosa Haz., iv, 237.
v. hajnaldiana Haz., iv, 237.
v. compacta Haz., iv, 237.
v. solitaria Haz., iv, 237.
eiisarcosoma Serv.
v. piceata Gredl., iv, 237.
brunnea Porro.
v. lednicensis Branc., iv, 238.
v. thessalica Bttg., iv, 238.
v. Iagarina3 Adami, iv, 238.
v. pyrgia Bgt.
v. segalaunica Sayn.
v. promseca (Bgt.) Loc.
v. gratiosa Gredl.
H. buchii Dub., iv, 238.
abichiana Bayer.
H. leucorum L., iv, 239.
mutata Lam.
v. yleobia Bgt.
v. virago Bgt.
depressa Bgt.
v. ryparia Bgt.
v. nigrozonata Bgt.
v. atrocincta Bgt.
v. anaphora West,
v. castanea Oliv., iv, 239.
mahometana Bgt.
v. euphratica v. Mts., iv, 240.
v. rumelica Mouss., iv, 240.
v. onixiomicra Bgt., iv, 240.
v. elongata Bgt., iv, 240.
v. straminea Brig., iv, 240.
/. straminiformis Bgt.
v. taurica Kryn., iv, 241.
H. radiosa ZiegL iv, 241.
H. schlseflii Mouss., iv, 241.
v. pra?stans Bl. & W.
H. secernenda Rm., iv, 242.
v. insignis Branc.
H. valentini Kob., viii, 239.
H. ligata Miill., iv, 242.
f annularis Perry.
decussata Parr.
melissophaga Costa.
varians Ziegl.
/. pomatella Tib., iv, 243.
/. prsetutia Tib., iv, 243.
/. campana Tib., iv, 243.
/. delpretiana Paul., iv, 243.
/. truentina Masc., iv, 243.
/. pseudopomatia Bl., iv, 244.
/. rupicola (Bl.) West,
v. albescens Jan., iv, 244.
v. interamnensis Bgt.
v. dichromolena Bgt.
v. gussoneanaSh., iv. 243.
H. ambigua (Parr) Mss. iv, 244.
cyrtolena Bgt.
/. clathrata West,
v. aetolica Kob., viii, 239.
acarnanica Kob.
v. thiesseana Kob., iv, 244.
H. anctostoma Mts., iv, 244.
beilanica West.
H. cincta Miill., iv, 245.
lemniscata Brum.
dalmatica Miihlf.
v. pollinii DaC., iv, 245.
320
HELIX.
v. calabrica Kob., iv, 243.
albescens Adami.
v. trojana Kob.
v. anatolica Kob.
v. cypria Kob.
H. asemnis Bgt., iv, 245.
solida Ziegl.
v. homerica Mart., viii, 239.
v. venusta Mart., pi. 44, f. 6,
7.
H. moabitica Goldfuss.
H. melanostoma Drap. iv, 246.
f. pachypleura Bgt.
f. vittata Rm., iv, 245.
pronuba West.
f. Candida Rm., iv, 247.
rugosa Ai)t.
f. nupta West.
v. nucula Parr., iv, 247.
v. cathara West.
v. giulise Bgt.
v. uticensis (Bgt.) Pech.
v. melanonixia Bgt.
H. figulina Parr., iv, 247.
v. pomacella Parr., iv, 247.
H. pachya Bgt., iv, 248.
v. texta Mouss., iv, 248.
v. dehiscens Westerl.
pachya Kob., Icon., f. 1031,.
not Bgt.
H. pathetica Parr., iv, 248.
H. socia Rm., iv, 248.
H. philibinesis Friv. Rm., iv, 249
philibensis Pfr.
H. nilotica Bgt., iv, 249.
H. vulgaris Parr., iv, 249.
obtusata Ziegl., preoc.
obtnsalis Mouss.
f. vallionis Ret.
v. bicincta Dub., iv, 250.
H. lutescens Ziegl., iv, 250.
cinerascens Andr.
H. nordmanni Parr., iv, 257.
H. raddei Bttg., iv, 251.
H. christophi Bttg., iv, 251.
H. prasinata Roth, iv, 252.
H. cavata Mouss., iv, 252.
H. engaddensis Bgt., iv, 253.
H. pycnia Bgt., iv, 253.
H. godetiana Kob., iv, 253.
latecava Mouss.
H. malzani Kob., iv, 254.
H. equitum, luynesiana, jauberti, edroea, schahbulakensis Bgt.,.
(iv, p. 256) are unfigured and insufficiently known forms.
Section Tachea Leach, 1831.
Tachea Leach, TURTON, Manual of the Land and Freshwater
Shells of the British Is., 1831, p. 33 (nemoralis and hortensis').
HARTMANN, Erd- u. Siisswasser-Gasterop. Schweiz, pp. 24, 189, 212,.
1840. Helicogena (part) FERUSSAC, BECK, etc. Cepcea HELD.,
Isis, 1837, p. 910. Archelix (second section) ALB., Die Hel., 1850,
p. 98.
Shell imperforate, globose or subdepressed, with low-conoid spire
and rounded periphery ; bright colored, usually yellow, with five
dark bands, any or all of which may be absent. Whorls 5, the last
HELIX. 321
deflexed in front, tumid. Aperture wide-lunar, oblique ; lip ex-
panded and thickened within, the baso-columellar margin straight,
widened by a blade-like callus within, flattened and adnate. Sur-
face smoothish. Type H. nemoralis L., pi. 44, figs. 4, 5.
Animal showing a pair of longitudinal grooves on back and indis-
tinct right and left facial grooves, elsewhere coarsely granular ; sole
very indistinctly tripartite; mantle-edge with small right and
minute left body-process on each side of breathing pore.
Jaw (pi. 67, fig. 1, H. nemoralis, Wurzburg) solid, arcuate, with
4-6 strong ribs grouped in the median part and denticulating either
margin. Radula having the middle cusps only developed on cen-
tral and lateral teeth, the side-cusps represented by a slight lateral
wave, but in some forms they are present and minute. Marginal
teeth having the inner cusp long, oblique and bifid, outer cusp small,
simple (pi. 67, figs. 2, 3, H. nemoralis').
Genitalia: penis long, bearing a long flagellum ; duct of sperma-
theca very long and usually with a diverticulum. Dart sack mus-
cular, containing a four-bladed dart, with conspicuously coronated
base and long head, the blades split in some species. Mucus glands
two, inserted on opposite sides of vagina immediately above d. s.>
each subdivided into several long, slender cylindrical finger-like
caeca (pi. 63, fig. 12, H. nemoralis).
Distribution, entire Europe.
Tachea is one of the most conspicuous and characteristic forms of
European snail life. They live on bushes and walls, in gardens,
vineyards, etc., and, while avoiding the direct rays of the sun, are
light-loving creatures. They colonize freely, H. nemoralis increases
rapidly where introduced in America. H. hortensis inhabits many
of the islands off the New England coast, and being found in pre-
Columbian kitchen-midding deposits, cannot be regarded as a recent
immigrant. Possibly it may be the sole survivor of that Viking
incursion in the eleventh century.
The variations of the Tacheas are mostly in coloring, and it has-
not been considered worth while to give below the multitude of
names founded on band-variations, etc. There is considerable vari-
ation in the size of dart sack, and in the darts of various species, as
well as in the number of fingers of the mucous glands, which are
generally quite long (15-16 mill, in nemoralis, splendida, coquandi^
29 in vindobonensis), and vary from four in each group in nemoralis,
to from 16 to 30 in vindobonensis.
21
322 HELIX.
H. atrolabiata Kr., iv, 124.
calligera Dub.
v. stauropolitana Schmidt,
v. leucoranea Mousson.
v. pallasii Dubois.
v. decussata Boettger.
v. intercedens Retowski.
v. nemoraloides Martens.
H. vindobonensis Fer., iv, 124.
austriaca Miihl.
mutabilis Hartm.
arvensis Kryn.
H. subaustriaca Bgt.
H. nemoralis Miill.,iv, 122.
/. pura West, Verb. k. k.
z.-b. Ges., '92.
v. erjaveci Kobelt.
v. lucifuga Hartm.
appenina Stabile.
genuensis Porro.
H. hortensis Mull., iv, 123.
subglobosa Binn.
H. sylvatica Dr., iv, 125.
v. montana Stud.
v. rhenana Kob.
?v. litturataPfr.,iv,126.
H. coquandi Mor., iv, 125.
/. nemoraloides Kob.
/. nahoni Kob.
/. ellioti Kob.
/. depressa Kob.
H. spleudida Drap., iv, 147.
/. roseolabiata Rm.
v. cossoni Let., iv, 148.
v. calseca Fag.
H. aimophila Bgt., iv, 126.
v. tchihatcheffi Kob.,iv, 126.
v. aimophilopsis Villes, iv, 126.
H. vicaria West.,Nachr.,1894,168
(Quaternary fossil species from Mentone.)
H. paretiana Issel, iv, 130.
monaecensis Ramb.
H. oedesima Nev.
v. colorata Nev.
v. crassior Nev.
H. mentonica Nevill.
H. bennetiana Nev.
H. williamsiana Nev.
v. subuemoraiis Nev,
v. spanias Nev.
Two species described by Deshayes are referred to this group by
Pfeiffer. H. gibbosula Desh., iv, 126, and H. filosa Desh., iv, 126.
Their localities are unknown, and subsequent authors have not
identified them.
Section Otala Schumacher, 1817.
Archelix ALB., Die Hel.,^1850, p. 14, 21, 98 (exclusive of section
). Maeularia MARTENS Die HeL, 1860, p. 132, and of authors
.generally, not Maeularia Albers, 1850. Helicogena in part of
Ferussac, Risso, Beck, et al. Otala SCHUM., Essai d'un nouv. Syst.,
p. 191 (for hcemastoma, atomaria = lactea, sulcata = Plieadomus).
MOQ.-TAND. (in part), Hist. Nat. Moll. Fr., ii, p. 142. Not Otala
Beck, 1837, Index, p. 35.
HELIX. 323
Shell depressed-globose, imperforate, solid, somewhat cretaceous ;
white, wnicolored or banded, a five-banded pattern usually trace-
able. Surface usually finely malleated. Last whorl rounded at
periphery, deeply deflexed in front. Aperture very oblique, trun-
cate-oval, the outer lip expanded and thickened within, baso-
columellar lip straightened, reflexed and adnate, widened by an
internal callus. Type H. lactea Mull., pi. 44, fig. 11 (see also pi. 44,
figs. 9, 10, H. vermiculata).
Animal externally like Tachea.
Jaw arcuate with blunt ends and 4 to 7 strong, convex ribs den-
tating both margins (pi. 67, fig. 4, H. vermiculata). Radula similar
to that of Tachea, side cusps being developed in some species, absent
in others (pi. 67, fig. 5, H. vermiculata).
Geuitalia (pi. 63, fig. 8, H. vermiculata') similar to Tachea, but in
the typical species the mucus glands are split into a great number
of caeca, as in Pomatia. Dart coronated at base, with four blades,
which may be either simple (H. alonensis, pi. 63, fig. 13) or divided
(H. vermiculata, pi. 63, fig. 5). In the group of H. alonensis, balear-
ica, minoricensis, etc., the mucus glands have few fingers, as in H.
( Tachea) nemoralis.
Distribution, southern Europe, Northern Africa, Canary Islands.
This section differs from Tachea in the more compact, solid shell
with generally a more deflexed last whorl and irregular color-
pattern. It presents no constant anatomical difference from Tachea,
but in most species the fingers of the mucus glands are more numer-
ous.
The name Otala was proposed for three species, placed in two
sections. Section a contained hcemastoma (which, being the type of
a prior genus, must be eliminated, see ant. p. 153), and atomaria, a
new name for lactea Mull. Section b contained the Helix sulcata of
Miiller, a form which Swainson, in 1840, made the type of his group
Plicadomus. These eliminations leave H. lactea the valid. nucleus of
Schumacher's group, and this name should have been adopted by
Albers in 1850 ; but, instead, he coined a new one Archelix. This
name was dropped in Marten's edition of Die Heliceen, 1860, and
the species placed in Macularia, a group originally proposed by
Albers for the spotted and unkeeled Iberus, and which did not
originally contain the species vermiculata, which Martens names as
its type! As the the type of Macularia had been expressly said to
324
HELIX.
be nieiensis by Lowe iu 1854, Martens action clearly cannot be
sustained ; and, unless we use the name Otala for this group, it must
be called Archelix. Beck's use of Otala has no bearing upon the
case, as he included none of Schumacher's species in his group..
H. vermiculata Mull., iv, 128.
muraloides Chier.
v. thalassina Porro.
v. grimaldiensis Nev.
v. uticensis Kob.
v. gaidurina Bl. & W.
v. saharica Kob.,iv, 128.
v. linusina Ben.
linusce Calc.
v. subangulata Iss.,iv, 129.
v. pelogosana (Stoss.) West.
H. punica Morel., iv, 129.
H. constantinse Fbs.,iv, 129.
cirtw Terv.
v. fleuratiBgt,iv,129.
H. boghariensis Deb., iv, 129.
H. lactea Miill., iv, 130.
irrorata Say.
atomaria Schm.
/. bertheri Bgt. (albino).
v. ezquerriana Bgt.
v. turturina (Guirao) Rm.
v. maura (Guirao) Rm.
simocheila (Bgt.) Serv.
v. sevillensis Serv.
v. sevilliana (Grat.) Mss.
v. murcica Rm.
v. axia Bgt.
v. malacensis Anc.
v. bleicheri Palad. iv, 132.
stomatodcea Bgt.
v. ibrahimi Bgt.
v. sphseromorpha Bgt.
v. plesiasteia Bgt.
v. bathylsema Bgt.,iv, 130.
v. alybensis Kob., iv, 130.
v, tagina Serv., iv, 130.
H. gibbosobasalis Woll.,iv, 131.
H. atavorum Mab., iv, 131.
H. ahmarina (B.) Mab., iv, 131.
H. punctata Mull., iv, 131.
myristiqmcea (Bgt.) Pech.
f. galena (Bgt.) Pech.
v. punctatissima Jen.
v. bredeana Deb.
v. apalolena Bgt., iv, 132.
H. tingitanaPal.,iv, 132.
H. lucasiDh., iv,132.
H. ghazouana Deb., iv, 133.
H. hieroglyphicula Mich., iv,|133.
oranica Bgt.
/. integrivittis Anc.
H. alabastrites Mich. ,iv, 134.
soluta Mich.
v. pycnochilia Bgt.
H. atlasica Mouss., iv. 134.
H. alcyoneKob.,iv,134.
H.juilleti Terv., iv, 134.
chottica Anc.
saidana Deb.
v. marguerittei (B.) Pch.
v. heliophila (B.) Pch.
H. bailioni Deb., iv, 135.
H. denansi Kob., iv, 135.
H. beguirensis Deb., iv, 135.
beguirana\A.uct.
H. wagneri Rossm., iv, 136.
H. charieia Pech.,iv, 136.
H. jourdaniana Bgt.,iv, 136.
H. arichensis Deb., iv, 137.
v. crassidens Deb.,iv, 137^
v. catodonta (B.) Pech.
HELIX.
325
v. lobethana Deb.,iv, 137.
H.zaffarina Terv., iv, 137.
v. zelleri Kob., iv, 138.
/. doubletti Bgt.
H. anoterodon Pech., iv, 138.
H. dupotetiana Terv., iv, 138.
H. brevieri Pech., iv, 139.
dupot. v. aspera Gass.
v. rugosa Kob.
euglyptolena Bgt.
v. subbrevieri Bgt.
H. xanthodon Ant., iv, 139.
v. ema Bgt.
v. pseudoernbia Bgt., iv, 141.
H. arabica Terv., iv, 139
v. abrolena Bgt.
H. odopachya Bgt.,iv, 140.
H. bonduelliana Bgt.,iv, 140.
v. asteia Bgt.
H. leucochilops Pils., iv, 240.
leucocheila W., not Cox.
H. senilis Morel., iv, 140.
H. subsenilis Cr.
H. embia Bgt, iv, 140.
H. burini Bgt., iv, 141.
H. tigri Gerv.. iv, 141.
tigriana Bgt.
maresi Cr.
v. stereodonta Bgt.
v. dicallistodon Bgt.
H. surrodonta Bgt., iv, 142.
H. dastugui Bgt., iv, 142.
H. subjobseana Kob., iv, 142.
H. jobaeana Cr., iv, 142.
H. beaumieri Mouss., iv, 149.
H. raymondi Moq., iv, 149.
desfontanea Morel.
H. efferata Mousss., iv, 145.
H. moussoniana Woll., iv, 145.
adonis Mouss., not Ang.
H. alonensis Fer.,iv, 146.
/. lorcana Rossm.
v. carthageniensis Rossm.
v. campesina Ezq.
v. bajoi (Bgt.) Serv.
v. loxana Rossm.
H. alcarazana Guir., iv, 147.
H. guiraoana Rossm., iv, 147.
v. augustata Rossm., iv, 147.
H. ebusitana Hid.
H. marmorata Fer., iv, 147.
exornata Parr,
v. menobana (Bgt.) Pech.
v. violacea Rossm.
pulchella Rm.
paftschii Bgt.
H. balearica Ziegl., iv, 148.
hispanica Partsch.
speciosa Ziegl.
/. valdemusana Bgt.
/. eustrapa Bgt.
v. companyonii Aler. iv, 148.
companyoi West.
pyrenaica Rossm.
v. palmana (Berth.) Bgt.
H. minoricensis Mitt., iv, 148.
minorica (Berth.) B.
/. sampoli (Bgt.) Pech.
H. massylsea Morel., iv, 144.
v. zenatia Kob., iv, 144.
H. prsedisposita Mss.,iv, 145.
H. rereyana Mss.,iv, 145.
H. codringtoni Gray, iv, 143.
ferussaci C. & J.
eucincta Bgt.
euchromia Bgt.
eupcecilia Bgt.
v. pseudoparnassia Mss.
v. lycica Mart,
v. callirhoe Rolle.
326 HELIX.
subsp. parnassia Roth., iv, 1 43. bland Bgt., mss.
subsp. oetse Kob., iv, 143. v. pantocratis Broem.
/. alba Kob. v. coracis Kob.
v. setolica Bttg., iv, 143. subsp. intusplicata Pfr. viii,
subsp. crassa Pfr. iv, 144. v. subangulata Kob. [240.
Unfigured and imperfectly known species or forms of Otala.
H. miloni, parisotiana, hermieri, chydopsis, of (Bgt.) Pechaud. H.
ramisi, catharolena, toukriana, galiffetiana, eugastoria, bandotiana,
agenna. lucentumensis, acanonica, nitefacta, sticta, azorella, lampri-
mathia, takredica, romalea, brocha, seignetti, Bgt. H. cantse
chorista, tiranoi (Bgt.) Serv. H. secouria, mattarica Let & Bgt.
H. seguyana, acatergastra, speiratopa, bouthyana, alabalstra Pechaud.
H. duriezi Deb. H. flattersiana Anc.
Section Hemicycla Swainson, 1840.
Hemicycla SWAINSON, Malacology, p. 331, type H.plicaria Lam.
Mycena Alb., Die Hel., 1850, p. 123. Cochlea (part) ADANSON
etal
Shell imperforate or umbilicate, globose-depressed, solid and
opaque ; 5-banded, but the number frequently reduced by the ab-
sence of band v or the coalesence of bands ii and iii, sometimes all
bands obsolete. Surface strongly striate, decussated or malleated*
Whorls 4 to 6, the last deflexed in front. Aperture very oblique ;
lip reflected throughout, thickened within, the baso-columeller mar-
gin wider, usually flattened and appressed, often obliquely toothed.
Young shells angular or keeled. Type H plicaria Lam., pi. 43, fig.
43 (see also pi. 43, fig. 44, H. saulcyi Orb.). Anatomy unknown.
Distribution, Canary Islands. Although the anatomy of this
group is still unknown, the close correspondence of its shell to Otala
renders its systematic position moderately certain. The soft parts
will probably prove the same as in other pentatseniate snails, unless
an earlier stage of development be retained in fewer- branched mucus
glands. The Canary Island fauna is much less individualized than
that of the Madeira, Azores and Cape Verde groups, and in its Helices
it seems much more nearly allied to that of the Mediterranean tract.
The number of species will probably be somewhat reduced by more
critical study of their variations.
HELIX.
327
H. plicaria Lam.,iv, 151. H.
plicatula Lara. H.
orbieulata Wood. H.
planorbula Gray. H.
H. chersa Mab., iv, 153.
H. benthencourtiana Sh., iv, 151. H.
H. sarcostoma W. & B., iv, 152. H.
v. thaumalea Mab., iv, 152.
H. paeteliana Sh., iv, 152. H.
H. bathycoraa Mab., iv, 153. H.
H. eucalypta Mab., iv, 154.
H. sabiniana Mab.. iv, 154.
H. zelotaMab.,iv,154. H.
H. ephedrophila Mab., iv, 155.
H. themera Mab.,iv, 155. H.
H. riproclri Mab., iv, 155. H.
H. janthiuaMab., iv, 156. H.
H. gravida Mouss.,iv, 156. H.
H. bathycampa Mab., iv, 157.-
H. subgravida Mab., iv, 157. H.
H. barckeriana Mab., iv, 157. H.
H. cacopista Mab.,iv, 158. H.
H. cateucta Mab., iv, 158. H.
H. justini Mab.,iv,159. H.
H. helygaia Mab., iv, 159. H.
H. cacoplasta Mab.,iv, 159. H.
H. callipoua Mab., iv, 160.
H. perrieri Mab., iv, 160.
H. verneaui Mab., iv, 1 61. H.
H. idrytaMab.,iv,161. H,
H. hedonica Mab.,iv, 161.
H. galdarica Mab., iv, 162. H.
H. ledrui Mab., iv, 162. H.
H. ethelema Mab., iv, 163. H.
H. agaetana Mab.,iv, 163. H
H. saulcyi Orb., iv, 164.
v. temperata Mss., iv, 164. H
H. baiaMab.,iv, 164. H
H. embritha Mab., iv, 165.
H. crypsidoma Mab., iv, 165.
stulta Mab., iv, 165.
carta Mab.,iv, 166.
retrodens Mouss., iv, 166.
pouchet Fer., iv, 167.
adansoni W. & B.
desculpta Mouss., iv, 167.
modesta Fer., iv, 167.
paivana Lwe.
idiotrypa Mab.,iv, 168.
malleata Fer., iv, 168.
bidentalis Lam.
v. nivarise Woll., iv, 169.
, glasiana Sh,,iv, 169.
pellislacerti Rv.
, glyceia Mab., iv, 170.
empeda Mab.,iv, 170.
fritschi Mouss., iv, 170.
, consobrina Fer., iv, 171.
v. vetusta Mouss.
, evergasta Mab., iv, 17L
. cacopera Mab., iv, 172.
, bathyclera Mab., iv, 172,
. thanasima Mab.,iv,173.
, ephora Mab., iv, 173.
. cardiobola Mab., iv, 173.
. guamartemes Grass., iv, 174.
manriquiana Lwe.
guartemes Martens.
, invernicata Mouss., iv, 174.
. maugeana Sh., iv, 175.
gaudryi Rv.
. distensa Mouss., iv, 175.
. hedybia Mab., iv, 176.
. perraudierei Grass., iv, 176.
. hierroensis Grass., iv, 176.
valverdemis Lwe.
. indifferens Mouss., iv, 177.
. gaudryi Orb., iv, 177.
v. evergeta Mab.
v. gaudryopsis Mab.
328 HELIX.
H. granomalleata Woll., iv, 178. H. quadricincta Morel., iv, 182.
H. vermiplicata Woll., iv, 178. H- berkeleii Lwe., iv, 186.
H. amblasmodon Mab.,iv, 179. H. saponacea Lwe.. iv, 183.
H. zorgia Mab., iv, 179. H. psathyra Lwe., iv, 183.
H. planorbella Lam., iv, 180. H. thespesia Mab., iv, 183.
villiersii Orb. H. bituminosa Mab., iv, 184
v. incisogranulata Woll. H. merita Mouss., iv, 185.
H. inutilis Mouss., iv, 181 H. harmonica Mouss., iv, 185.
H. plutonia Lwe., iv, 181. H. gomerensis Morel., iv, 185.
H. semitecta Mouss., iv, 181. H. thoryna Mab., iv., 185.
H. paivanopds Mab.,iv, 182. H. hedeia Mab., iv, 186.
paivana Morel., not Lwe. H. digna Mouss., iv. 186.
Section Iberus Montfort, 1810.
Iberus MONTF., Conch. Syst. ii, p. 146, type /. gualterianus.
Euiberus WESTERL. Fauna Paliiaract. Binnenconch., Helix, p. 367,
1889, same type. Macularia ALBERS, Die Hel. 1850, p. 80.
LOWE, P. Z. S. 1854, p. 166, type H. nwiensis Fer. H. & A. AD.,
Gen. Rec. Moll, ii, p. 210. Not Macularia of v. Martens and sub-
sequent authors. Murella PFR., Mai. Bl. xxiv, p. 8, proposed for
H. serpentina, surrentina, therescv, strigata, carseolana, melitensis,
provincialis, muralis. MONTS., Moll. Terrest. della Isole adiacenti
alle Sicilia, p. 32, 33, restricted to group of H. serpentina. Trans-
iberus MONTS., Moll. Terrestri, etc., (in Atti della R. Accad. di Sci-
enze, Lettere e Belle Arti (3) ii), p. 33, 1892 ; proposed for Sicilian
Iberus.
Shell rather cretaceons, imperforate or partly covered perforate,
varying from depressed or lens-shaped to globular; solid, smooth or
wrinkled, with to 5 spiral bands. Last whorl rounded or keeled,
deflexed in front. Aperture very oblique, subovate. Lip expanded
on outer and basal margins and thickened within ; columellar lip
reflexed, dilated toward insertion. Type H. gualtierana L., pi. 44,
fig. 8. See also pi. 44, fig. 15, H. scabriuscula ; figs. 16, 17, H. mur-
alis ; fig. 18 H. sicana.
Jaw (pi. 67, fig. 7 H. serpentina) with 3-6 ribs, denticulating the
margins. Sometimes the ribs are almost obsolete. Radula char-
acterized by the lack of side cusps on central and lateral teeth, the
middle cusps being about as long as the basal plates. Marginals
with a long bifid inner cusp and small simple or bifid ectocone (pi.
HELIX. 329
-67, fig. 6 H. serpentina). Genital system (pi. 61, fig. 8 H. gualtier-
ana; pi. 61, figs. 6, 7, H. muralis; pi. 63, fig. 1, 2, 3, H. serpentina).
Penis rather short, the retractor and epiphallus inserted at its apex,
epiphallus ending in a moderately long flagellum. Spermatheca
globular, on a long duct which bears a diverticulum. Dart sack as
in Otala. Mucus glands two, simple, or each dividing into two or
three branches. Dart four-bladed, with expanded, feebly cren-
ulated base (pi. 63, figs. 1, 2, H. serpentina).
The anatomy of Iberus is like that of Otala except in the fewer
fingers of the mucus glands. In this respect, both Tachea, Helico-
qena and Otala exhibit great variation ; and the same is true of
Iberus. It is perfectly clear that no characters whatever, for the
separation of these groups, can be obtained from the soft anatomy.
They rest wholly upon conchological characters. The dart is not
of the typically four-bladed type in some species, although it is in
H. gualtierana. In serpentina it seems more like a modified two-
bladed form.
The group Iberus was originallyp roposed for H. gualtierana only,
so that Westerlund's name Eaiberus seems to me wholly superfluous.
The next published name for the group was Macularia of Albers,
proposed for the species with rounded periphery and shotted bands.
Albers selected no type from his list, but in 1854 Lowe named H.
niciensis as the type. The name Macularia was used for the species
of Albers original list by Morch (Catal. Yoldi, 1852), by H. &
A. Adams (Gen. Kec. Moll. 1855) and others, so that both by the
formal selection of a type from the original list of species, and by
usage in well-known publications, the name became fixed. It was,
therefore, directly contrary to the fundamental principles of nomen-
clature for Albers-Martens in the second edition of Die Heliceen
(1860) to shift the name to the group of H. vermiculata ; and
although this unlawful usage has been followed by all later authors
to this day, it is too obviously unjustified to stand longer uncor-
rected. As to the synonymy of the other sectional names, it would
seem that in the absence of characters thei* discussion is not worth
the ink it would cost.
Bourguignat believes that the north African group of globose
Iberus, such as If. sicanoides, is a modification of the H. raymondi
stock (Bull. Soc. Mai. Fr. i, p. 7).
I. Iberus s. str. Keeled and depressed, coarsely latticed.
Mucous glands several-branched.
330
HELIX.
H. gualtierana L., iv, 202.
obversa Born.
v. laurentii Bgt. Spain*
II. Murella Pfr. Mucus glands simple.
Keeled species : Group of H. sultana ; Algeria, Tripoli,.
Morocco.
H. sultana Morel., iv, 202. H. leachii Fer., iv, 203.
subscabriuseula Bgt. tripolitana Wood.
H. culminicola Pons., viii, 234. H. quedenfeldti Mart., viii, 235.
H. viola (Pons.) Kob., viii, 234.
Keeled species : Group of H.
H. scabriuscula Dh., iv, 203.
erycina Jan.
v. verrucosa Monts., viii, 233.
v. selinuntina Ph., iv, 203.
v. segestana Ph., iv, 203.
v. explanata Ben., iv, 204.
v. demissa Ben., iv, 204.
v. drepanensis Huet., iv, 204.
scalariformis Ben.
scabriuscula ; Sicily, Sardinia.
H. nadorrica West., viii, 233.
tumidosa Monts.
H. paciniana Ph., iv, 204.
vieta Rm.
/. eulasia Westerl.
H. grohmanni Phil., iv, 205.
H. sardonia v. Mts., viii, 236.
v. dorgaliensis Mts., viii, pi.
[26, f. 38.
H. buelowi Malz., viii, 236.
Globose species: Group of H. sicanoides ; Morocco, Algeria.
H. weberi Kob., viii, 236. H. tetuanensis Kob., iv, 213.
platyclieloides Kob., iv, 211. H. sicanoides Kob., iv, 213.
H. boettgeri Kob., iv, 211. H. sollieri Bgt.
H. gyrostoma Fer., iv, 212.
Globose or globose-depresssed :
H. sicana Fer., iv, 213.
soluta Zgl.
H. platychela Mke., iv, 209.
prcetexta Jan.
v. rosalise Ben., iv, 209.
v. iparia Ben., iv, 209.
v. connexa West. Rm. f. 593.
H. aschera? Kob., iv, 210.
H. ragusse Kob., iv, 208.
H. provincialis Ben., iv, 208.
Group of H. muralis. Sicily, etc.
H. melitensis Fer., iv, 207.
/. caruanse Pils., iv, 208.
H. ridens v. Mts., viii, 236.
/. splendens Malz., iv, 211.
f. minor Malz., iv, 211.
H. globularis Ziegl., iv, 206.
frivaldskyi Calc.
/. conspicua Ben., iv, 207.
/. cossurensis Ben., iv, 207.
/. undulata Kob., iv, 207.
HELIX.
331
/. achatina Ben., 207.
/. saracena Ben.
v. tarentina Kob., iv, 207.
v. latilabris Westerl.
H. rollei Malz., viii, 236.
H. arista Wester!.
H. caltabellotensis Kob., viii,
H. talamonica Kob., viii, 231
H. tiberiana Ben., iv, 206.
H. muralis Mull., iv, 205.
abromia Bgt.
v. rugosa Ziegl., iv, 205.
v. costulata Ben., iv, 205.
v. crispata Ben., iv, 206.
v. alutacea Paul., iv, 206.
v. insularis Ben., iv, 206.
v. undulata Mich., iv, 206.
communis Ben.
orgonensis Philb.
v. abrsea (Bgt.) Mab.
232. v. ciofaloi Cafic.
v. messanensis Suliotti.
v. substrigata (Bgt.) Mab.
H. eugenia Pfr., iv, 221.
mgenia Parr.
calypso Ben.
v. huetiana Ben., iv, 221.
hueti Pfr.
III. Macularia Alb. Mucus glands two or three-branched at ends ;
dart peculiar. Depressed species: mainly Italy, Sardinia and
Corsica.
H. niciensis Fer., iv, 214.
nicceensis Rissor.,
v. faudensis Sulliotti.
v. clairi Bgt.
H. oberndoerferi Kob., iv, 217.
? halmyris Mab.
? v. tylota Westerl.
H. serpentina Fer., iv, 214.
v. isilensis (V.) Paul., iv, 215.
v. jaspidea Mab., iv, 215.
marmorellata Mab.
v. trica Paul.
v. velanicia Mab.
H. isane Paul., viii, 236.
H. magnetti Cantr., viii, 236.
hospitans Bon., Paul., iv, 215
v. alabastrina Paul., iv, 216.
H. carse Cantr., iv, 216.
v. orites Westerl.
v. adjaciensis Paul., iv, 216.
H. cenestinensis Cr. & Deb., iv,
[216.
H. suburbana Paul., viii, 236.
H. pudiosa Paul., iv, 217.
v. austera W.
forsythi Kob. Icon., n. f., 508.
H. villica Paul., iv, 218.
H. tetrazona Jan., iv, 218.
picvena Tib., iv, 109.
/. ascoliensis Bgt.
H. strigata Mull., iv, 218.
/. posidoniensis Tib., iv, 219.
/. corrugata Z., Rm. Icon., 229.
f polita Paul., iv, 219.
v. sicula Ben., 219.
v. umbrica Charp., iv, 218, 224
/. moltenii Ad., West.
. v. fuscolabiata Rm., iv, 219.
H. theresse (Ben.) Mts., iv, 224.
H. surrentina Schm., iv, 218.
/. lucana Bl., West.
H. saxetana Paul., iv, 224.
H. mariaunse Kob., iv, 222.
circumornata Kob., olim.
332 HELIX.
v. peucetana Kob., iv, 122. carseolana Auct.
v. apula BL, iv. 219. marrucina Tib.
H. forsythi Paul., iv, 223. v. recondita West., iv, 220.
/. orta Paul., iv, 224. v. contaminata Paul.,iv, 220.
H. argentarolse Paul., iv, 223. v. uzielliana Paul., iv, 220.
H. signata Fer., iv, 223. v. persianii Tib., iv, 220.
circumornata Mts. not Fer. v. uniarmata Paul., iv, 220.
fauum Miihlf. v. milettiana Paul., iv, 220.
/. virginea Blanc. H. nebrodensis Prj., iv, 220.
H. scherzeri (Zel.) Pfr. v. silvestrii Cafici, iv, 226.
H. carsoliana Fer., iv, 219.
Section Levantina Kobelt, 1871.
Levantina KOB., Catal. Eur. Binnenconch., p. 19. See for anat-
omy SCHMIDT, Stylom., pi. 4, f. 21, and SCHUBERTH, Arch. Naturg.,
1892, pi. 5, f. 9, 10.
Shell solid, rather cretaceous, large and depressed, the earlier
whorls acutely keeled, rendering the sutures rather superficial, last
whorl wide, rounded, umbilicate or imperforate, ornamented with 4
or 5 bands of arrow-like spots, or unicolored whitish, deflexed in
front. Aperture ovate-rounded, very oblique; lip expanded, the
baso-columellar margin reflexed and dilated. Type H. guttata.
(See pi. 44, figs. 1, 2, 3, H. guttata var. sesteri.)
Jaw with 4-6 ribs denticulating the margins. Radula with the
middle cusp of the median and lateral teeth wide, about as long as
basal plates, no side cusps. Marginals with a long bifid inner cusp
and a small ectocone.
Genital system (pi. 61, fig. 10, H. hierosolyma) with penis and
flagellum as usual in Otala. Dart sack rather small ; mucus glands
3-branched. Spermatheca duct long, with a long diverticulum.
Dart four-bladed.
Distribution, Island of Rhodes and Palestine to the Mesopota-
miau Desert and Persia.
The anatomy offers nothing separating this group from Otala or
Iberus, but the characteristic shell and the distribution render it
convenient to retain the group.
HELIX. 333
H. spiriplana Oliv., iv, 226. H. guttata Oliv., iv, 228.
v. malziana (Parr.) Pfr., iv, v. ergilensis Galland.
[227. v. sesteri Gall.
v. gallandi Bgt. v. michoniana Bgt., iv, 229.
H. hierosolyma Boise. baschkira Pfr.
/. masadse Tristr., iv, 227. H. mazenderaneusis Nev., iv, 229.
/. lithophaga Conr., iv, 228. H. kurdistana Parr., iv, 230.
H. cresareana Parr., iv, 227. H. ceratomma Pfr., iv, 231.
/. maxima Bgt. H. escheriana Mss., iv, 230.
/. carinata Bgt. /. diarbekirana Gall.
/. albidula Bgt. /. euthyomphala Gall.
/. nana Mouss. H. bellardii Mouss., iv, 231.
v. globulosa Bgt. /. occlusa Mouss.
H. werneri Rolle, viii, 236. H. ghilanica Mss., iv, 231.
H. dschulfensis (Dub.) Bgt., iv, H. urmiensis Naeg., viii, 237.
228. H. ninivita Gall., viii, 238.
dschulfensii Dubois. H. lapithcensis Rolle.
djulfensis Mouss. H. gertrudis Rolle.
H. chrysostomi Rolle.
Subgenus (?) PARACHLOR^EA Sandberger.
Parachlorcea SANDB., Conch. Vorwelt., p. 292, type H. coquandiana
Matheron.
Shell iraperforate, lens-shaped, acutely keeled. Last whorl de-
flexed in front. Aperture very oblique, angulate-oval, the margins
remote; upper margin of lip slightly expanded, basal margin
reflexed and appressed. Type H. coquandiana Math.
The type of this group is from the French " Palseotherium chalk "
of Oligocene age, but similar forms are found in Eocene and in
lower Miocene deposits. The group, whether rightly limited or not,
is probably a side branch of the Helix phylum. For any one to
connect it with Chlorcea, on account of the keel, seems unjustified
in view of the vast variability of this character. The development
of a keel in Helix, with the consequent modification of shell contour,
is a character of the most trivial import. A few of these forms
which I have seen, seemed to be keeled manifestations of the Tachea
group.
334 HELIX.
Subgenus EREMINA Pfeiffer, 1855.
Eremina PFR., Mai. BL, 1855, p. 139, sole species H. desertorum.
Eremophila KOBELT, Katalog Eur. Binnenconch., p. 19, 1871 ;
Iconographie iv, p. 13. Erinna MORCH., Journ. de Conch., 1865,
p. 387 (desertorum). Eremia Auct. Conf. JICKELI Moll. N.-O.-
Afrika's, pi. 1, f. 7-9 (dentition, jaws and darts) ; SCHUBERTH,
Archiv f. Naturg., 1892, pi. 5, f. 11, 12 (teeth and dart). SEMPER,
Reisen im Arch. Phil. Landmoll., pi. 14, fig. 14 (genitalia).
Shell iraperforate or narrowly umbilicate, solid, chalky, with
rudely striate surface; white with reddish bands or suffused and
streaked with tawny. Whorls about 5, the last somewhat or not
descending. Aperture slightly oblique, wide lunate ; lip a little
expanded and blunt or greatly thickened, the columellar margin
reflexed, arcuate, not thickened by an internal plate of callus. Type
H. desertorum Forsk., pi. 44, figs. 12, 13 ; also fig. 14, var. ehrenbergi
Roth, (chilembia Bgt.)
Jaw stout, arcuate, with blunt ends; having grouped near the
middle two to four strong ribs dentating both margins (pi. 67, fig.
9, H. desertorum ; fig. 8, H. desertella). Radula having basal plates
rather short; cusp attaining edge of basal plate, the side cusps ob-
solete; laterals bicuspid, with a small outer cusp. Marginals
having the larger cusp bifid, the ectocone also splitting on the outer
ones (pi. 67, fig. 10, H. desertorum).
Genitalia: Spermatheca duct long (45 mill.) ivith no diverticulum.
Dart sack small, containing a two-bladed dart with serrate crown,
and a longitudinal rib or sometimes a blade on one or each side
(pi. 63, fig. 6, H. desertorum). Mucus glands 2, long pediceled,
branching into 5 to 6 fine tufted fingers. No flagellum on penis, the
retractor and vas deferens terminal (pi. 63, fig. 4, H. desertorum).
The solid, cretaceous shell and lack of flagellum on the penis
ally this group to Euparypha; the long-stalked pair of digitate
mucus glands and the tendency to be five-banded are points of like-
ness to Tachea. The dart is slightly coronated at base, the head
long and two-bladed, but with side ribs which sometimes develop
into blades, and it, therefore, is mainly of the type common in Pen-
tata3nia. A diverticulum on spermatheca duct is wanting, but this
feature varies greatly even among closely allied species.
The species are desert forms, inhabiting northeastern Africa.
HELIX. 335
H. desertorum Forsk., iv, 127, H. desertorum.
[261. v. hasselquisti Ehr.
maculosa Born. /. ehrenbergi Roth.
irregularis Fer. chilembia Bgt.
forskalii Ehr. v. hemprichi Ehr.
arabica Roth. v. aschersoni Reinh.
psamitus Bgt. H. desertella Jick., iv, 127.
gemellarii Ben. H. duroi Hid., iv, 128.
rhodia Chemn. /. minor Kob., iv, 261.
depressa Mts. haploa West.
pachytoichea West.
kobelti West.
dillwyniana Pfr.
Subgenus EUPARYPHA Hartraaun, 1842.
Euparypka HARTM., Erd.- u. Siisswasser Gasterop. Schw., p. 204,
for H. rhodostoma =pisana. Conf. for anatomy, SCHMIDT, Sty-
lommat., p. 22. pi. 5, f. 23 ; MOQ.-TAND., Moll. Fr,, p. 259, pi. 19, f.
9-19, SCHUBERTH, Arch. f. Naturg., 1892, p. 55, pi. 6,f. 1-3.
Shell narrowly umbilicate or subimperforate, compact, solid, creta-
ceous, subglobose or depressed and keeled, white or light, usually
decorated with many dark irregularly placed bands and lines. Sur-
face striated, the striae decussated by spiral mcised lines, at least on
the spire. Suture superficial ; last whorl descending or straight ;
aperture little oblique, lunate, the lip sharp, not expanded, thickened
by an internal callus rib, columellar end triangularly reflexed.
Type H. pisana, pi. 43, figs. 37, 38.
Jaw well arched, with two or three strong ribs denticulating both
margins. Radula as in Pentatsenia, Helicigona, etc. ; the middle
cusp of middle tooth is longer than the short, square basnl plate,
side cusps small ; laterals similar but without inner cusp, outer cusp
larger; marginals with a long bifid inner cusp and smaller outer
cusp, the latter split on the outer teeth.
Genital system (pi. 61, figs. 1,2, 3, 4, 5, H. pisana) characterized
by the lack of flagellum on penis, the retractor and vas deferens ter-
minal ; spermatheca duct long, without a diverticulum (or accord-
ing to Schmidt with a short one) ; mucus glands two, long, simple
and tubular. Dart sack small, containing a straight, four-edged,
coronated dart (fig. 2, 4, 5) like that of Tachea splendida.
336 HELIX.
Distribution, Mediterranean region, Canary Is., Madeira, etc.; JET_
pisana extending to southern England and throughout the whole
range of the genus. H. macandreiviana is confined to the tiny
Atlantic Islands known as the Salvages; H. impugnata and the
beautiful varieties geminata, grasseti, hierophanta of H. pisana, are
Canarian.
The cretaceous, many-banded shell, with non expanded, inwardly
thickened lip, is more like that of Helicella than the Pentatseniates ;
and the lack of a flagellum is also a strong differential feature,
allying Euparypha to Eremina.
The single dart sack with its four-sided dart is a structure char-
acteristic of the five-banded group; and the right eye retractor
passes between the branches of the genitalia, as in normal Helices ;
these features at once removing Euparypha trom the Xerophiloid
stock. The simplicity of the two mucus glands is a character in
common with Helicigona and Iberns ; and evidently represents an,
earlier stage of development than the digitate type, which has been
retained in these groups. I suppose the lack of flagellum to be a
degenerative change. Euparypha is, therefore, a curious mixture,
the mucous glands being of antique character, the male organs
degenerate, and the dart modernized. Teeth and jaw offer nothing
characteristic.
H. pisana, the typical species, is an abundant snail from northern
Africa to southern England and the Atlantic islands. It has been
split by "new school" authors into a multitude of alleged species,
some of which may prove worth retention as local varieties. The
principal names are as follows : thusuroi, subpisana Bgt., brysce,
radesiana (Mares) Bgt., chambardi, salemensis, gergisensis, carpiensis,
hamadanica, djerbanica, zitanensis Let. & Bgt., maculata Mke.,
anonyma W., donatii, levesquei (Berth.) Bgt., pisanella, pisanopsis,
dermoi Serv. Other synonyms are zonaria Penn., petholata Oliv.
rhodostoma Dr., eigenda Mont., slrigata Dillw., leucostoma Risso, and
doubtful varieties are perruginea, Mke., pundella M.-T., subzonata
Bgt., sigarellina Charp., alboranensis (Webb. & Berth.) Mab. (not
Pfr. et Auct.), catocyphia Bgt., iii, 256, hyperplatceaServ., etc.; etc.
H. macandrewianaPfr., iii, 224. v. pisanopsis Serv., iii, 225.
ustulata Lwe. v. aegusse Kob., iii, 225.
H. pisana Miill., iii, 224. v. sardoa Ziegl., iii, 224.
/ decorata Pfr., iii, 225. v. graphica Morel.
ADDITIONS AND CORRECTIONS. 337
cestivalis Bgt. callio4oma A. & R.
v. geminata Mouss., iii, 224. H. dehnei Rm., iii, 225.
alboranensis Pfr., iii, 224. epigtottidea Bk.
v. hierophanta Mab., iii, 225. f. erythronixia Bgt.
v. grasseti Tarn., iii, 225. f. thlipsa West.
pisanoides Orb. H. subdentata Fer. iii, 226.
H. impugnata Mouss., iii, 226. subcarinata Mke.
/estiva Lwe., Mss. cince Kl.
v. subgeminata Mouss. H. pisaniformis Bgt., iii, 227.
H. planata Chemn., iii, 226. H. comaliana Bgt., iii, 227.
helieella Wood. v. tiani Bgt., iii, 227.
v. arietina Rossm. v. tohenica Bgt., iii, 227.
v. erythrostoma Ph.
ADDITIONS AND CORRECTIONS.
Genus TROCHOMORPHA (p. 1).
To species of first group on page 4, add :
T. horiomphala Pfr., iv, 51. T. cathcart% Reeve, iv, 51.
From list of Philippine I. species omit T. conomphala Pfr., which
is a young Obba parmula, teste Ponsonby in litt., and T. radula Pfr.
a species of Bensonia ; and add :
T. crassula Mlldff. T. gracilis Mlldff.
T. pseudosericina Mlldff. T. suturalis Mlldff.
T. morongensis Mlldff. T. heptagyra Mlldff.
T. alticola Mlldff. T. sericina Mlldff.
T. schmackeri Mlldff. T. splendidula v. carinaria Mlldff
T. intermedia Mlldff. T. boholensis Semper.
Mollendorff (Ber. Senck. Nat. Ges. 1893, p. 74) considers the
well-distinguished local races sibuyanica Hid., boettgeri Mlldff., with
qvadrasi Hid. as referable to metcalfei Pfr. if it be advisable to under-
stand the species in such wide limits. Is Tr.stenozona Mlldff., men-
tioned as a new species from Luzon, but still undescribed, in Ber.
Senck. Ges. 1890, p. 213, another form of this species?
T. natunensis Smith, Ann. Mag. N. H. '94, 455. Natuna Is.
T. partunda (not " partunga," p. 5) Angas.
T. hidalgoana Crosse (p. 6) is reported by Brazier from N. Georgia,
Solomon Is.
22
338 ADDITIONS AND CORRECTIONS.
Genus PUNCTUM Morse (p. 6).
Add : P. massoti Bgt., iii, 29, shown by Pollonera to belong here.
Genus LAOMA Gray (p. 8).
Add the following species described in Trans. N. Z. Inst. xxvi.
L. ciliata Suter.
And to section Phrixgnathus, these from New Zealand :
L. murdochi Suter. L. cheesemani Suter.
And the following from Tasmania: (see Ann, Mag. N. H., Jan.,
1894, p. 64).
L. weldii Tenison-Woods. L. pipaensis Petterd.
L. furneauxensis Petterd. L. halli Cox.
L. hobarti Cox.
Genus FLAMMULINA Mart. (p. 10).
Suter in Ann. Mag. Nat. Hist. Jan., 1894, p. 64, gives a classifica-
tion of Tasmanian Flammulinas from the examination of the denti-
tion, as follows:
Sect.: Flammulina: F. jungermannise Pett., sitiens Cox, luck-
manni Braz.
Sect.: Gerontia: F. albanensis Cox, stanleyensis Pett., legrandi
Cox, marchiannse Cox, diemenensis Cox, gadensis Cox, tasmanise
Cox, subrugosa Braz., mathinuae Pett., macdonaldi Cox, bassi Braz.,
tamarensis Pett.
Sect. : Phacussa : F. savesi Pett., stephensi Cox, hamiltoni Cox.
Sect.: Allodiscus: F. limula Cox.
Sect.: T/talassohelix : F. fordei Braz.
Some of these may prove to be Charopas, however.
P. 13, add after tranquilla Cox, iii, 26. After hamiltoni Cox, iii,
87. ccepta Cox should stand coepta.
Section Allodiscus Pils. (p. 14).
F. smith! Suter, Tr. N. Z. Inst. xxvi. F, rustica Suter, t. c. p. 135.
Section Pyrrha Hutt. (p. 15).
F. subincarnata Suter, Tr. N. Z. Inst., xxvi, p. 133.
Section Phenacolielix Sut. (p. 16).
F. pilula v. unicolor Suter, Tr. N. Z. Inst. xxvi, p. 134.
ADDITIONS AND CORRECTIONS.. 339
Section Flammulina Mart. (p. 18).
The species novarce proves to belong to Zonitidce, and should be
removed from list on p. 18.
F. pilsbryi Suter, Tr. N. Z. Inst., xxvi, p. 133.
Section Carthcea Hutton, 1884.
Carthcea HUTTON, Trans. N. Z. Inst., xvi, p. 189, type H. kivi.
These smooth, subtrochiform shells, with conspicuously streaked
color-pattern, prove to belong to Flammulina, the typical species
having been investigated by Mr. Suter. F. kivi is from New Zea-
land, floscu lus from Norfolk Island.
F. kivi Gray, iii, 37. F. flosculus Cox, viii, 77.
irradiata Gld.
radiaria Pfr.
Genus ENDODONTA Alb. (p. 20).
P. 24. E. sculptilis Pease should be a synonym under E.frater-
cula Pse. the former name being preoccupied in Helix. On p. 27
add iii, 39, after E. lamellicosta Garrett. Add to list :
E. mariannarum Quadr. & Mild if., Nachr. D. M. Ges. 1894, p. 14.
E. heptaptycha Quadr. & Mlldffi, Nachr. D. M. Ges. 1894, p. 15.
Mr. Suter in Proc. Linn. Soc. N. S. Wales, viii, p. 494, adopts the
etymology "hunnaensis" for E. (Ptyehodon) hunuaensis of his several
previous publications. As I do not find the Hunua or Hunna
Range on the maps accessible to me, I do not know which spelling
is correct.
Add on p. 28 these from St. Helena (Conchologist ii, pp. 164,
165):
E. sexdentata Smith. E. perarmata Smith.
Section Charopa Alb.
On p. 31, " Pterotropis" was a pen-error for Pterodiscus.
Omit " E. raricostata " from list on p. 33 ; place E. coma var. beta
as a synonym under v. globosa Suter ; add after E. ostiolum Cr., ii,
180. E. serpentinula Suter is a variety of bitccinella Rve. E. muta-
bilis Suter is a synonym of tau Pfr.
E. anguiculus Reeve (p. 32). E. pseudocoma Suter.
v. raontivasra and v. fuscosa Sut.
340 ADDITIONS AND CORRECTIONS.
Add the following Marianne and Natuna Island species of Charopa r
Nachr. D. M. Ges. 1894, p. 13, 14 :
E. fusca Quadr. & Mlldff. E. quadrasi Mlldff.
E.rottila Quadr. & Mlldff. E. persculpta Sm., Ann. Mag. '94,
The Helix (Helicella) australis of Menke, from Mt. Eliza, Swan
River, may possibly belong to Charopa if it is really Australian
but it certainly does not look like one. See vol. iii, p. 103.
On p. 33, E. microscopica Cox (not Krauss) must be dropped iiv
favor of E. microcosmos Cox.
On p. 34, E. " cupera " Cox=cuprea. For " retepora " and " re-
teporoides," read reiipora and retiporoides.
Genus PHASIS Alb., and Trachycystis Pils. (p. 37).
Suter (Ann. Mag. N. H. 1894, p. 60) believes a caudal pore to
be present in Pella burnupi, the dentition of which he figures. None
was visible in the badly preserved specimens of P. rariplicata ex-
amined by me. On p. 39 the following corrections should be made i
P. inclara Morel. P. zanguebarica Crav., iii, 105.
H. inops Morel, not Mouss.
Genus SCULPTARIA (p. 39).
Possibly this may prove to be a genus of Protogona. Ponsonby
writes that Ancey's S. chapmanni (subsequently changed to S. mel-
villiana, Brit. Nat. 1892, p. 126) has been compared with the type
of damarensis H. Ad., and found to be absolutely the same.
Genus AMPHIDOXA (p. 41).
A. chiliensis Muhl. (not chilensis*).
A. tenuistria Phil, (not tenuistriata).
Genus PYRAMIDULA (p. 42).
Section Microconus Strebel & Pfeffer, 1880.
Microconus STREBEL & PFEFFER, Beitr. Mex. Land- und Suss-
wasser-Conchyl. iv, p. 29, type M. ivilhelmi Pfr.
This name is proposed for a small species of eastern Mexico resem-
bling Pyramidula rupestris in contour, but ribbed as in P. rotundata
or perspective It can hardly be regarded as more than a " section "
of Pyramidula. The position of hermanni and mazatlanica is doubt-
ful. '
ADDITIONS AND CORRECTIONS. 341
P. wilhelmi Pfr., iii, 53. P. Hermann! Pfr., iii, 22.
P. mazatlanica Pfr., ii, 204.
Section Patulastra Pfr. (44).
P. massoti Bgt. has been shown to be a Punctum. P. luseana
Paiva becomes a synonym of P. placida Shuttl., an earlier name.
The name luseana is repeated by error at foot of p. 47. P. tenuicos-
tata Shuttl. being preoccupied in Helix, Servain calls the species
shuttleworthi.
Section Gonyodiscus (p. 45).
Add to list of species, p. 47, the following Palsearctic forms :
P. machadoiMilne-Edw. P. kompsa Mabille.
rotundata Morel. concinna Lwe. not Jeffr.
v. azorica Mouss. P. rotundata Mull.
P. scutula Shuttlw. v. infracostata Westerl.
P. omalisiana Bgt. abietina Paul., non Bgt.
omalisma, err. typ., p. 47. P. chaperi West. Verh. k.-k. zool.-
P. putrescens Lowe. bot. Gesell. Wien. 1892, p. 27.
P. ganoda Mabille.
Section Lyrodiscus (p. 48).
The type of this group is believed by Wollaston to be a Zonitid.
This is not unlikely, but the shells before me are dull, and the
animal is unknown.
Section Lynda Wollaston, 1878.
Lyrula WOLLASTON, Testacea Atlantica p. 382, type H. loweana.
Wollaston is disposed to class this elegant, spirally laciniate Pat-
uloid snail with the Madeiran H. lentiginosa. The single species is
from Lanzerote, Canary Is.
P. loweana Wollaston: (Not lowei Fer.)
torrefacta Lowe not C. B. Ad.
usurpans Furtado, iv. 40.
Section lulus Woll., 1878,
This (preoccupied) name is proposed by Wollaston (Test. Atlant.
p. 326) for thePatulagarachicoensis Woll. (H. agrestis Lwe. in litt.\
a minute form found in Tenerife, Canary Is. It is said to have rela-
tions with P. putrescens Lwe. of Palma, and P. bertholdiana Pfr., of
the Cape Verdes. P. garachicoensis has not been figured, and I have
seen no specimens. A variety submarmorata is described by Woll-
aston.
342 ADDITIONS AND CORRECTIONS.
Genus PARARHYTIDA Anc. (p. 52).
Mr. T. D. A. Cofkerell has called my attention to the fact that
the name Saissetia (p. 53) is preoccupied in entomology (Coccidce),
see Zool. Rec. 1865, and Proc. Amer. Ent. Soc. 1893, p. 54. He
proposes to call the m oil u scan group PLATYRHYTIDA, the species-
saisseti being its type.
Genus THYSANOPHORA.
P. 58, add to synonyms of T. boothiana, H. mauriniana Orb.,
(" lavalleana " on plate). The reference after T. incrustata should
stand : ii, 204.
Genus POLYGYRA (p. 68).
Add to list on pp. 76, 77, P. thyroides v. pulchella Ckll., (Journ.
of Conch. 1892, p. 39), and
P. sanburni W. G. Binn., iii, 145. Kingston, Idaho.
Genus POLYGRATIA (p. 81).
Cancel the sectional name Entodina, (p. 83) and the species rey-
rei Souv., as a radula received from Prof. Gwatkin proves it to belong
to Streptaxidce. The other species are quite different, and may as
well be placed in Sysirophia, from which they differ only in the
parietal lamina.
Genus PLEURODONTA, Section Isomeria (p. 93 ).
P. meyeri Strubell, Conchyl. Cab., p. 693.
Genus CA1VLENA (p. 101).
C. stolidota Quadr. & Mlldff. Paragua, Philippines.
The radula of C. cicatricosa is figured on pi. 34, fig. 10, from a
mount kindly lent me by Prof. Gwatkin.
Section Pse,udobba Mlldff. (p. 105).
C. brunonis Kobelt, Conch. Cab., p. 681. Halmaheira,
Genus OBBA (p. 107).
Add H. conomphala Pfr. to synonymy of 0. parmnla (p. 109).
O. viridiflava Mlldff. O. marginata v. nan a Mlldff.
G. subhorizontalis Mlldff. v. pullescens Mlldff.
O. flavopicta Mlldff. O. moricandi v. radiata Mlldff.
O. basidentata v. grandis Mlldff. O. scrobiculata v. conoulalis
Mlldff.
ADDITIONS AND CORRECTIONS. 343
Genus PLANISPIRA Beck (p. 110).
Add to species of Cristigibba, (p. 113).
P. parthenia Kobelt. Conch. Cab., pi. 200, f. 9,10.
And to list on p. 114, the following from W. Australia, Proc. Mai.
Soc. i, p. 93.
P. bathurstensis Smith. P. gascoynensis Smith.
Genus CHLORITiS (p. 117).
Add these Australian species, which may belong in the preceding
group :
C. millepunctata Smith. C. rectilabrum Smith.
v. cassiniensis Smith.
C. subsulcata MlldfF. Cuban, Calamianes Is.
. latecostata Kobelt. C. (Sulcobasis ?) djamnensis Kob.
C. buxina Heude. China.
Genus THERSITES (p. 125).
Section Badistes Gld. (p. 129).
Add below T. OSCARENSIS Cox (p. 131) the synonym H. (Rhagada)
inconvicta Smith, Proc. Mai. Soc. i, p. 90. Add to T. DERBYI Cox (p.
131) the synonym H. (Trachia*) derbyana Smith, t. c., p. 92. And
the following species, described in the same place, all from N. W.
Australia :
T. obliquerugosa Smith. T. sykesi Smith.
T. prudhoensis Smith. T. imitata Smith.
T. burnerensis Smith. v. cassinieusis Smith.
T. montalivetensis Smith.
Subgenus RHAGADA (p. 135).
Smith in Proc. Malac. Soc. i, p. 89, suggests that H. torulus Fer.
is the same as j reinga Gray, and places H. elachystoma Mart, as a
synonym under richardsoni Sm. He queries the subgeneric refer-
ence of plectilis and carcharias, but it seems to me unnecessarily, for
the specimens of both examined by me are very close to typical
Rhagada.
344 ADDITIONS AND CORRECTIONS.
Genus PAPQINA (p. 136).
Col. Beddome writes me that Helix plurizonata Adams & Reeve,
described evidently in error from Borneo, is really the same as torn-
asinelliana Tap. -Can. (p. 142) and agnocheilus Smith, which, there-
fore, become synonyms. Also that the "Group of P. antiqua" (p.
141) consists of one species, antiqua, of which the other names are
synonyms. Add the following:
P. divaricata Kobelt. P. lintschuana Kobelt.
The latter much like P. goldiei Braz. (p. 141), and like that, of
doubtful generic position.
Genus PLECTOPYLIS (p. 143).
P. quadrasi Mlldff. Nachr. 1893, p. 172.
P. azona Gredl., viii, 158. P. vallata Hde., viii, 158.
Genus PYROCHILUS (p. 154).
P. pyrostoma vars. lucernalis and nigrescens Kobelt.
Genus EPIPHRAGMOPHORA.
P. 197, for petasensis read patasensis.
Genus EULOTA (p. 200).
Add to Plectotropis, p. 209 : E. luzonica Mlldff., Nachr. '94, p.
105.
Genus HELICOSTYLA (p. 216).
Col. Beddome (in Hit.) informs me that Cochlostyla belcheri is a
bleached velata ; C. andromache is a color-variety ofpolillensis.
Another name for Orthostylus is BULINA Lesson, Illustr. de Zool.
1831, pi. 22, for Helix (Bulina) rufogaster. Perhaps it is meant for
a spelling of Bulimus.
Genus LEUCOCHROA (p. 234).
Add : L. debeauxi v.hypophysa West., Verh. k.-k zool.-bot. Ges.
Wien, '92, p. 26.
The list on pp. 249, 250, was intended to be alphabetical, but
through wrong paging of the mss. this end was defeated.
REFERENCE TO PLATES
FRONTISPIECE.
FIGURE. PAGE.
1. Ganesella japonica Pfr. Genitalia. Pilsbry, del. . . . 168
2. Ganesella japonica, end of penis laid open. Pilsbry, del. . 168
3. Dorcasia alexandri Gray. Genitalia. Pilsbry, del. . . 172
4. Helicostyla (Crystallopsis) conformis Fer After Tap.-Can. 220
5. Helix pomatia, penis, dart sack and vagina opened. Pilsbry,
del.
^6. Helix pomatia L. Pilsbry, del. ..... 317
a.gl. albumen gland; air. atrium; d.s. dart sack; epi. epi-
phallus; fl. flagellum; h.gl. hermaphrodite gland; m.gl.
mucus gland ; p. penis ; r. penis retractor ; sp. spermatheca ;
sp.d. spermatheca duct; ut. uterus; vag. vagina; v.d. vas
deferens.
7. Helix pomatia, showing mantle lobes, etc. Pilsbry, del. 317
8. Leptaxis undata Lowe, dart sack and mucus glands. Pilsbry,
del. 292
9. Leptaxis undata Lowe. Pilsbry, del. 292
PLATE 1.
1. Laoma leimonias Gray. Pilsbry, delin. ... 8
2. Laoma acanthinulopsis Sut., teeth. After Suter, . . 8
3,4. Laoma glabriuscula Pfr., teeth and jaw. After Suter, . 8
5. Punctum cryophilum Mart., jaw. After Jickeli, .
6. Punctum pygmseum Dr. After Schako, Mai. Bl., xx, . 7
7. Punctum pygmseum, one plate of jaw x 1000. Ibid. . 7
8. Punctum conspectum Bid. Teeth. Pilsbry, del. . . 7
9. 'Punctum conspectum Bid., jaw. After Bin ney, . . 7
10. Laoma (Phryxgnathus) celia Hutt. Pilsbry, del. . . 8
11-13. Punctum pygmseum Drap. Pilsbry, del. ... 7
14, 15, 16. Amphidoxa (?) hookeri Rve. Anat. Monatsber.
Berl. Akad., 1877, .... .40
PLATE 2.
1, 2, 3. Flammulina (Phacussa) hypopolia Pfr. Suter, Tr.
N.Z. Inst. xxiv, ... ... 12
4, 5. Flammulina (Gerontia) pantherina Hutt. Suter, Tr.
N. Z. Inst. xvi, 14
(345)
346 REFERENCE TO PLATES.
FIGURE. PAGE,
6, 7. Flamnmlina (Phenacohelix) pilula Reeve. Suter, Tr.
N. Z. Inst. xvi 16
8. Flammuliua (Suteria) ide Gray. Suter, Tr. N. Z. Inst.
xvi, 17
9, 10. Flammulina (Pyrrha) cressida Hutt. Suter, Tr. N. Z.
Inst. xiv, ........ 15
11. 12. Flammulina (Allodiscus) tullia Hutt. Suter, Tr. N.
Z. Inst. xxiv, . . . . . . .14
13, 14. Flammulina(Allodiscus)godeti Sot. Suter, Tr. N. Z.
Inst. xxiv, 14
15, 16. Flammulina corneofulva Pfr. Suter, Trans. N. Z.
Inst., xxiv, ........ 18
17, 18. Flammulina chiron Gray. Suter, Trans. N. Z. Inst.
xxiv, .18
19, 20. Flammulina (Therasia) decidua Gray. Suter, Trans.
N. Z. Inst., xxiv, 16
21. Flammulina (Therasia) thaisa Hutt. Hutton, Trans. N.
Z. Inst, xvi, ........ 16
PLATE 3.
1-3. Flammulina (Gerontia) pantherina Hutt. Shepperd, del. 14
4-6. Flammulina (Allodiscus) planulata. Hutt. Pilsbry, del. 14
7-9. Flammulina (Monomphalus) rossiteriana Cr. J. de
Conch., 1873, 19
10, 11, Flammulina (Phacussa) hypo polia Pfr. Shepperd, del. 12
12. Flammulina (Allodiscus) tullia Gray, apexx 24. Pilsbry,
del ' . 14
13. Flammulina (Phenacohelix) pilula Reeve. Pilsbry, del. . 16
14-16. Flammulina (Therasia) thaisa Hutt. Type. * Suter,
del 15
17-19. Flammulina (Pyrrha) cressida Hutt. Type. Suter,
del '. . . 15
20-22. Flammulina (Calymna) costulata Hutt. Type. Suter,
del 18
23. Flammulina zebra Le Guill. Conch. Icon. . * 17
24-26. Flammulina (Suteria) ide Gray. Shepperd, del. . . 17
27. Flammulina (Thalassohelix) zelandise Gray. Hutton,
Tr. N. Z. Inst . .13
28. Flammulina (Thalassohelix) ziczac Gld. Pilsbry, del. . 13
29. Flammulina (Thalassohelix) zelandise Gray. Shepperd,
del 13
PLATE 4.
30-32. Endodonta (Ptychodon) aorangi Suter. Pilsbry, del. . 28
33, 34. Endodonta (Thaumatodon) derbesiana Cr. Pilsbry, del. 2fr
REFERENCE TO PLATES. 34T
FIGURE. PAGE.
35-37. Endodonta (Thaumatodon) multilamellata Grt. Pils-
bry, del 26
38. Internal palatal lamellae of same, x 100. Pilsbry, del. . 26
39. Endodonta obolus Gld. Pilsbry, del 25
40. 41. Endodonta lamellosa Fer. Pilsbry, del. . . 25
42, 43. Endodonta (Helenoconcha) polyodon Sowb. P. Z. S.,
1892, 28
44. Endodonta (Pterodiscus) alata Pfr. Pilsbry, del. . . 36
PLATE 5.
45-48. Endodonta (Libera) subcavernula Tryon. Pilsbry
del
49, 50, 51. Endodonta ? (Brazieria) velata H. & J. Pilsbry
del
52, 53. Endodonta fabrefacta Pse. Pilsbry, del. .
54. Endodonta (Diglyptus) pagodiformis Sm. Pilsbry, del.
PLATE 6.
23
29
25
22
55, 56. Endodonta (Paratrochus) dalbertisi Braz. Hedley,
P. L. S. N. S. W 31
57-59. Endodonta (Charopa) coma Gray. Pilsbry, del. . 31
60. Endodonta (Phenacharopa) novoseelandica Pfr. Pilsbry,
del 29
61, 62. Endodonta (Tropidotropis) trichocoma Cr. J. de
Conch., 1868, 36
63-65. Endodonta (Charopa) tapirina Hutt. Pilsbry, del. . 31
66. Endodonta (Nesophila) tiara High. Aperture. Pilsbry,
del. . 27
67, 68. Endodonta (Aeschrodomus) stipulata Rve. Pilsbry,
del . 30
69, 70. Flammulina (Rhytidopsis) chelonites Crosse. J. de
Conch., 1868, 20
71-73. Endodonta (Acanthoptyx) acanthinula Cr. Pilsbry,
del 36
PLATE 7.
1-3. Trochomorpha quadrasi Hid. Shepperd, del. . . 1
4-6. Trochomorpha merzianoides Grt. Shepperd, del. . 1
7. Trochomorpha meleagris Pfr. Shepperd, del. . . 1
8, 9. Trochomorpha trochiformis Fer. Shepperd, del. . 1
10-12. Amphidoxa marmorella Pfr. Conchyl. Cab. . . 40
13-15. Trochomorpha planorbis Less. Wiegrn., Webers'
Zool. Erg 1, 2
16-18. Amphidoxa (?) hookeri Rve. Monatsber Berl. Akad., 40
348 REFERENCE TO PLATES.
FIGURE. PAGE.
1877, . 40
19-21. Amphidoxa (Stephanoda) dissimilis Orb. Voy. Am.
MSrid ... 40
22-24. Pararhytida (Platyrhytida) saisseti Mont. Shepperd,
del 54
25-27. Pararhytida dictyodes Pfr. Shepperd, del. . . 53
PLATE 8.
1, 2. Endodonta (Phenacharopa) novoseelandica Pfr. Suter
Tr. N. Z. Inst., xxiv, . .
3, 4. Endodonta (Aeschrodomus) barbatula Rve. Suter, Tr
N. Z. Inst., xxiv,
5, 6. Endodonta (Ptychodon) microundulata Suter. Suter
Tr. N. Z. Inst., xxiv, .....
7, 8. Pararhytida (Platyrhytida) astur Sow. W. G. Binn
Dent. Pulm. Moll. . . . .
9-12. Trochomorpha assimilis Grt. Pilsbry, del.
13, 14. Trochoraorpha beckianaPfr. After Semper, .
15, 16. Trochomorpha metcalfi Pfr. After Semper,
16. Base of uterus of same, showing high insertion of v. d.
17. Trochomorpha troilus Gid. After Semper,
18, 19. Trochomorpha subtrochiformis. After Semper,
PLATE 9.
29
30
28
54
1
1
1
1
1
1
20, 21. Endodonta (Charopa) sylvia Hutt. After Suter, . 32
22. Endodonta huaheinensis Grt. After Binney, . . .25
23, 24. Endodonta (Charopa) coma Gray. Pilsbry, del. . 32
25. Endodonta (Acanthoptyx) acanthinula Cr. Pilsbry, del. 36
26. Endodonta (Libera) tumuloides Grt. After Binney, . 23
27-29. Flammulina delta Pfr. After Hedley, . . .19
30-33. Trochomorpha timorensis Mart. After Stoliczka, . 2
34. Endodonta recedens Grt. Pilsbry, del 23
35-37. Pararhytida dictyodes Pfr. After Fischer, . . 53
PLATE 10.
1-3. Phasis menkeana Pfr. Shepperd, del., . . .37
4. Sculptaria sculpturata Gray. Conch, Icon., . . .39
5, 6. Phasis (Trachycystis) bisculpta Bens. Pilsbry, del. . 37
7. Phasis (Trachycystis) bisculpta, apex. Pilsbry, del. . 37
8, 9. Phasis (Trachycystis) browningii Bens. Pilsbry, del. 37
10, 11. Pyramidula (Planogyra) asleriscus Morse. After
Morse, 45
12, 13. Pyramidula (Gonyodiscus) rotundata Mull. Pilsbry,
del. 46
REFERENCE TO PLATES. 349
FIGURE. PAGE.
14. Pyramidula (Gonyodiscus) solaria Mke. Pilsbry, del. . 46
15, 16. Pyramidula rupestris Drap. Pilsbry, del. . 44
PLATE 11.
17, Pyramid ula strigosa Gld. Pilsbry, del 49
18, 20, 23. Pyramidula alternata Say. Pilsbry, del. . . 49
19, 22, 26. Pyramidula perspectiva Say. Pilsbry, del. . 46
21. Pyramidula asteriscus Morse. After Binney, . . .45
24. Pyramidula (Helicodiscus) lineata Say. After Binney, . 51
25. Pyramidula rupestris Drap. Pilsbry, del. . . .44
27. Pyramidula strigosa Gld. After Binney, . . .49
28. Pyramidula strigosa Gld. Pilsbry, del. . .49
PLATE 12.
1,3,7. PlanispirazonariaL. After Taparone-Canefri, , 110
2. PlanispirazonariaL. After v. Martens, . . . 110
4-6. Planispira zonaria L. Shepperd, del. . . . 110
8. Planispira (Cristigibba) plagiocheila T.-C. After Tap.-Can. 113
9, 10, 12. Planispira (Cristigibba) dominula T.-C. After
Tapperone-Canefri, . . . . . .113
11, 13, 15. Planispira (Cristigibba) macgregori Hedl. After
Hedley, . .... 112
PLATE 13.
16, 17. Papuina grata Mich. Tapparone Canefri, Ann. Mus.
Civ. Genov. xix, . 137
18, 19, 22. Papuina taumantias T.-C. Tapparone- Canefri,
Ann. Mus. Civ. Genov. xix, ..... 137
20, 21. Papuina yulensis Braz. Tapparone-Canefri, Ann.
Mus. Civ. Genov. xix, 137
23, 24. Papuina boyeri F. & B. Hedley, P. L. S. N. S. W. . 137
25. Papuina louisiadensis Forbes. Hedley, P. L. S. N. S. W. 137
26, 27. Papuina brumeriensis Forbes. Hedley, P. L. S. N. S. W. 137
28. Papuina macgillivrayi Forbes. From orig. sketch by
Hedley, . 137
29. Papuina fringilla Pfr. After W. G. Binney, . . .138
PLATE 14.
29-31. Pyramidula (Helicodiscus) lineata Say. Pilsbry, del
32, 33. Pyramidula (Atlantica) semiplicata Pfr. Pilsbry, del
34-36 Pyramidula alternata Say. Terr. Moll, iii,
37-39. Pyramidula strigosa Gld. Terr. Moll, iii, .
40, 46. Pyramidula alternata Say. Pilsbry, del. .
51
51
48
48
48
350 REFERENCE TO PLATES.
FIGURE. PAGE.
47, 48. Pyramidula (Helicodiscus) lineata Say. After Morse, 51
41, 42. Pupisoma lignicola Stol. Journ. As. Soc. Beng., 1870, 52
43, 44. Pupisoma philippinica Mlldff. Landschn. Cebu, . 52
45. Pyraraidula perspectiva Say. Pilsbry, del. . . .46
PLATE 15.
1. Pyramidula lineata Say. Pilsbry, del 51
2. Pyramidula balmei Pot. & Mich. Pilsbry, del. . . 46
3. 4. Trachycystis bisculpta Bens. Pilsbry, del. . . 37
5, 6. Thysanophora turbiniformis Pfr. After Binney, . . 55
7-10. Thysanophora peraffinis Ad. Pilsbry, del. . . 55
11, 12. Camsena (Pseudobba) quoyi Pfr., jaw. Mai. Bl. xx, 105
13, 14. Cama3na (Pseudobba) quoyi Pfr., teeth. Mai. Bl. xx, 105
PLATE 16.
1, 2. Thysanophora turbiniformis Pfr. Shepperd, del. . 55
3. Thysanophora conspurcatella Morel. Pilsbry, del. . 55
4. Thysanophora caeca Guppy. Pilsbry, del. . . .55
5-7. Thysanophora hypolepta Shuttlew. Pilsbry, del. . 55
8-10. Thysanophora stigmatica Pfr. Shepperd, del. . . 55
11-13. Sagda cookiana Gmel. Shepperd, del. . . .59
14, 15. Sagda alligans C. B. Ad. Shepperd, del. . . .60
16, 17. Sagda connectens C. B. Ad. Shepperd, del. . . 60
18-20. Sagda (Hyalosagda) similis C. B. Ad. Shepperd, del. 64
21. Zaphysema tenerrima C. B. Ad. Shepperd, del. . . 65
PLATE 17.
1-4, Pedinogyra Cunningham i Gray. After Hedley, . . 158
5. 6. Pedinogyra Cunningham! Gray. Shepperd, del. . 158
PLATE 18.
1-7. Camsena xanthoderma (?) Mlldff. Pilsbry, del. . .101
PLATE 19.
8. Camsena cicatricosa Mull. After Heude. . . .101
9. Camsena (Pseudobba) mamilla Quoy. Shepperd, del. . 105
10. Camsena (Camamella) platydon Pfr. Jahrb. D. M. Ges. 106
11. Obba (Oreobba) codonodes Pfr. Shepperd, del. . .109
12. Camsena monochroa Sowb. After Hidalgo, . . .104
13. Camoena (PlKenicobius) arata Sowb. Shepperd, del. . 104
14. 16. Obba planulata Lam. After Hidalgo, . . .107
15. Obba planulata Lam. Shepperd, del 107
i- 1 1 1 1- 1 < i PO I-I,\TKH. :;.M
VMM,
17. oi.i.:. i.;i ideatatfl I'd Aft. , Hidalgo . , . 107
I8,i!. Planiipira (Tracblella) tuckeri Prr Pil-Uy, d-i. . 104
20-22. PhiniHpira (Angasella) c;yrt,pkuMi I'd Sl.<-ppcl,
d-l ...... . .114
2:;, 21. Planfopira (TnwW) vittati Mflll Bhepprd,Sl !.'. : .
25. plum j.ini (Tnujhia) atperelli i'iv Coocb, i-'.n. . . 116
I'l.ATI, 20.
20, 27. Pniticoklla griseola Pfr. Uiol. Onlr. A KM T. . .07
28. PraticoleUa bermndicriaoa Morie* sii-j.|.rrl, del. . . (\l
29-">l. l'r;itKv,l<-lla fluvescens I'll. I'.iol. C.M.t. A mT. . . 67
'{2 .">4. Poly^yrella ( AmmoniUlla) rateti Coop- Pilsbry, del. 81
la (Odontoiagda) l.iii'-i ^undl. Pilwbry, del. . o. ; ,
.",7, ."18. I ynila Born. Sli-|>|i-rd, del. . . 81
!'>. Entodina reyrd Bow, 8hepperd,deL . 342
n.-i'!. Polygyratia ftenogyra Pfr- sii -|,p crd, del. . . 8.'}
44- Pi ' ( - f '-.iM-liyl. Cat. . . .83
PLATI; 21.
1-5. I'ruti(!"l-ll;i '.'-;ur i|. i !irn|>l;i 1'fV. After Htrebel and
87
:;, pcni.s ;i.nd :i|i|,-ndix '.(,< n-d : Hg. 4, talon).
0. Praticotolla ffrUeola Pfr Uter W, G, Binoey, . 87
7, 8. Sagda cookiaoa OoieL rilwiiry, d^l. .
i;i cookini linin above, appendix on left
side, HpiplmlliiH lrfiri^hin^ in riglit, where retractor
JH in-' ( ('--I , ^a deferent deicending i>ni.wn by
Pilibnr from fpfcimeo collected ly.J. !' n -nfUirKon 50
i') Appendix of ame partly uncoiled,
11-14. Polygyra albolabrii &ay- I'H-l.ry, d-l. . . .70
!.'. A i.rium ;ind p< ni <,\' ;in.< '.[.-ried, showing pilaster, . 70
16. Section of iwollen bate of spermatbeca duct of iao . 70
l'i A-II, 22.
'p.
2. I'l'Mirodont,*-. f I';. i t -,pc -td, '!<).
i'i-.urf,d'.rit- rPafthena) dominicenfii Pfr. sii'j>p<-rd d-i.
4. Pif.iirodonu: cZacbrrfia) trinitarU GuodL sii-ppTd, d-i. 'H;
IM'-.iir'idont" ''I 'li-.lid'rriii-'.; linci ' ! I . 96
<',. PleurodoDte (Eurycratera) jam n Shepperdi
del. . . .100
7,8. Pl-iir')d'int<: M/ihy rinl hug) iev r i \T;ut Concll
Mit.thfril ..... . 96
!>, io. PletirodoDte(Polydontei)iniperatorMoDtf Sbepperdi
352 REFERENCE TO PLATES.
FIGURE. PAGE_
11, 12. Macrocyclis laxata Fer. Shepperd, del. . . .165
PLATE 23.
13-17. Pleurodonte (Parthena) dilatata Pfr. Pilsbry, del. . 99
18. Pleurodonte (Parthena) angulata Fer. After Binney, . 99
19-23. Pleurodonte (Zachrysia) auricoma havanensis. Pils-
bry, del 96
24. Pleurodonte (Thelidomus) lima Fer. After Semper, . U6
25. Pleurodonte (Thelidomus) aspera Fer. After Binney, . 96
PLATE 24.
1. Pleurodonte schroeteriana Pfr. After Binney, . . 88
2. Pleurodonte (Caprinus) Josephine Fer. After Binney, . 90
3. Pleurodoute (Caprinus) nuxdenticulata Fer. After
Binney, .90
4. Pleurodonte acute Lam. Pilsbry, del 88
5. Pleurodonte invalida C. B. Ad. After Semper, . . 88
6. 7. Pleurodonte. acuta Lam. Pilsbry, del. . . . 8&
8-10. Pleurodonte (Caprinus) orbiculata Fer. Pilsbry, del. 91
11, 12. Pleurodonte dentiens Fer. After Binney, . . 91
PLATE 25.
1. Pleurodonte (Caracolus) caracolla. \V. Shepperd, del. . 92
2, 3, Pleurodonte (Isomeria) faunus v. ritchieana. W. Shep-
perd, del . . 93
4, 5. Pleurodonte (Labyrinthus) labyrinthus Chem. W.
Shepperd, del 95
6, 7. Pleurodonte sloaneana v. vendryesi Ckll. W. Shep-
perd, del. ... .... 88
8. Pleurodonte acuta v. nobilus Ad. W. Shepperd, del. . 88
9. Cepolis cepa Mull. W. Shepperd, del 179
10. Pleurodonte (Caprinus) nuxdenticulata Chem. W. Shep-
perd, del 90
11. Pleurodonte (Caprinus) Isabella Fe"r. W. Shepperd, del. 90^
12. 13. Pleurodonte (Gonostomopsis) auridens Rang. W.
Shepperd, del. ... .92
PLATE 26.
/
1. Pleurodonte (Caracolus) marginellaGmel. After Bin ney, 92
2. Pleurodonte (Caracolus) marginella v. semiaperta. After
Binney, 92
3. Pleurodonte (Caracolus) marginella Gmel. After Binney, 92
4. 5, 6. Pleurodonte (Caracolus) rostrata Pfr. Pilsbry, del. 92
REFERENCE TO PLATES. 353
FIGURE. PAGE.
7,8. Pleurodonte (Caracolus) caracolla Linn. After Semper, 92
9. Pleurodonte (Labryinthus) labyrinthus Chemn. After
Semper, 95
PLATE 27.
1, 2. Thersites (Sphserospira) rawnesleyi Cox. W. Shep-
perd, del . . . 132
3. Thersites (Badistes) bitseniata Cox. W. Shepperd, del. . 129
4. Thersites (Sphserospira) blomfieldi v. warroeutis. After
Tap.-Can 132
5. Thersites (Badistes) gulosa Old. W. Shepperd, del. .129
6. Thersites (Sphserospira) pomurn Pfr. W. Shepperd, del. 134
7. 8. Thersites (Glyptorhagada) kooringensis Ang. \V.
Shepperd, del 129
9, 10. Thersites (Glyptorhagada) kooringensis Ang. After
Cox, 129
11-13. Thersites (Rhagada) supracostulata Schep. After
Schepman. . . . . . . . .135
14, 15. Thersites (Rhagada) floresiana Mart. Weber's Zool.
Ergeb. 135
16-18. Thersites (Rhagada) carcharias Pfr. VV. Shepperd,
del 135
19. Thersites (Glyptorhagada) silveri Ang. W. Shepperd,
del 129
PLATE 28.
1-4. Chloritis porteri Cox. Pilsbry, del., . . .121
5-9. Chloritis argillacea Fer. After Wiegmann, . . 121
10. Chloritis dinodeomorpha Tap.-Can. After Tapp.-Can. . 119
PLATE 29.
1-3. Chloritis ungulina L. E. Shepperd, del. . . . 117
4, 5. Chloritis porteri Cox. E. Shepperd, del. . . .121
6, 7. Plecteulota goniostoma Mlldff. Nachr. 1892, . .122
8. Thersites richmondiana Pfr. E. Shepperd, del. . . 125
9. 10. Chloritis (Sulcobasis) sulcosa Alb. Novit. Conch. . 120
11. Papuina splendescens Cox. Shepperd, del. . . .137
12. Papuina lituus Less. Conchyl. Cab 137
13. Papuina nortoni Braz. E. Shepperd, del. . . . 137
14. 15. Papuina trobriandensis Hedl. E. Shepperd, del. . 137
16. Anoglypta launcestonensis Reeve. E. Shepperd, del. . 160
PLATE 30.
1-3. Polygyra cereolus septemvolva Say. Shepperd, del. . 71
4. Polygyra auriculata Say. Shepperd, del. . . .71
5. Polygyra septemvolva Say. After Binney, . . .71
23
354 REFERENCE TO PLATES.
FIGURE. PAGE.
6. Polygyra troostiana Lea. After Binney, . . 71
71
74
74
74
74
74
74
74
74
74
74
7. Polygyra postelliana Lea. After Binney,
8. Polygyra tridentata Say. After Binney,
9. 10. Polygyra appressa perigrapta Pils. Shepperd, del.
11, 12. Polygyra tridentata Say. After Binney, .
12, Polygyra inflect a Say. After Binney,
13, 14. Polygyra albolabris maritima Pils. Shepperd, del.
15. Polygyra clausa Say. Bfter Binney,
16. Polygyra albolabris Say. After Binney,
17. 18, 19. Polygyra sayi Binney. After Binney,
20. Polygyra clausa Say. After Binney,
21. Polygyra kiawaensis Simpson. Pilsbry, del. .
PLATE 31.
22-24. Polygyra (Stenotrema) monodon v. alicise. E. Shep-
perd, del. ........ 77
25. Polygyra (Stenotrema) monodon, jaw. After Binney, . 78
26. Polygyra (Stenotrema) hirsuta Say. After Binney, . 78
27. Polygyra (Stenotrema) spinosa Lea. After Binney, . 77
28-30. Polgyrella polygrella Bid. & Coop. Shepperd, del. . 80
31, 32. Polygyrella polygyrellla Bid. & Coop. After Binney, 79
33-35. Polygyrella harfordiana Coop. Pilsbry, del. . . 80
36-40. Glyptostoma newberryanum W. G. B. After Binney, 193
41. Polygyrella polygyrella Bid. & Coop. After Binney, . 80
PLATE 32.
42, 43. Chloritis leei Cox. After Hedley, .... 119
44, 45. Planispira delibrata Bens. After Stoliczka, . . 115
46, 47, 52. Thersites (Xanthomelon) pachystyla Pfr. After
Semper, 134
48. Thersites (Sphserospira) blomfieldi Cox. After Hedley, . 132
49. Thersites (Hadra) bipartita Fer. After Semper, . . 131
50. 51. Thersites (Sphserospira) rainbirdi Cox. After Hedley, 132
PLATE 33.
1. Thersites richmondiana Pfr. Hedley, Proc. K. Soc. Q'ld. 128
2. 3. Thersites (Sphserospira) mitchellse Cox. Pilsbry, del. 133
4-7. Thersites (Badistes) gulosa Gld. Hedley, Rec. Austr.
Mus. . . 130
PLATE 34.
1, 2. Thersites (Sphserospira) mitchellse Cox. Pilsbry, del. 132
3. Planispira zonaria L. Pilsbry, del. . . . .110
4. Planispira (Trachia) asperella Pfr. Pilsbry, del. . .115
REFERENCE TO PLATES. 355
FIGURE. PAGE.
5, 6. Planispira (Trachia) trochalia Bens. Pilsbry, del. .115
7, Thersites richraondiana Pfr. After Hedley, . . .128
8, 9. Albersia zonulata Fer. Pilsbry, del ..... 125
10. Camaena cicatricosa Mull. Pilsbry, del ..... 342
11. Papuina raoseleyi Smith. Pilsbry, del ..... 137
12. Papuina vexillaVis Pfr. Pilsbry, del ..... 137
PLATE 35.
1. Sagda (Hyalosagda) haldemaniana Ad. After Binney, . 59
2-8. Sagda (Hyalosagda) sirnilis Ad. Pilsbry, del. . . 59
9, 10. Zaphysema tumida Pfr. After Biuney, . . 66
11,12. Zaphysema tenerrima Ad. Pilsbry, del. . . 66
PLATE 36.
1-3. Allognathus graellsiana Pfr. After Schuberth. . . 290
4. Helicodonta obvoluta Mull ....... 285
5, 6. Helicodonta lenticula Fer. Pilsbry, del. . . 285
7, 8. Helicodonta maroccana Mor. After Schuberth. . 285
9, 10. Helicodonta lusitanica Pfr. After Schuberth. . . 285
11, 12. Polygyrella(Ammonitella) yatesi Coop. After Binney, 81
13. 14. Leucochroa candidissima Drap. Pilsbry, del. . . 232
15. Leucochroa boetica Rossm. After Schmidt. . . . 232
16. Leucochroa boissieri Char p. After Binney, . 4 . . 232
PLATE 37.
1. Papuina rnoseleyi Smith. Pilsbry, del. . . . 138
2. Papuina conscendens Cox. Pilsbry, del. . . . 138
3. 4. Papuina vexillaris Pfr. Pilsbry, del. . . .138
5. Papuina trobriandensis Hedley. Hedley, Proc. Linn. Soc.
K S. W ......... 138
6. Papuina brazierse Braz. Tapparone-Cauefri, Ann. Mus.
Civ. Genov ......... 138
7. 8. Papuina fringilla Pfr. Pilsbry, del ..... 138
9, 10. Papuina vexillaris Pfr. Pilsbry, del. . . 138
11. Papuina conscendens Cox. Pilsbry, del. . . .138
PLATE 38.
1. Acavus hsemastomus L. Pilsbry, del. .... 153
2, 3. Pyrochilus lampas Mull. Kobelt, Conch. Cab. . . 154
4. Helicophanta goudotiana Fer. Shepperd, del. . . 151
5. Helicophanta cornugiganteum Chemn. Pilsbry, del. . 151
6. 7. Dorcasia alexandri var. rotundata. Journ. de Conch.
1887, ......... 172
tJNIVlBSJTY
356 REFERENCE TO PLATES.
FIGURE. PAGE>
8. Dorcasia globulus Mull. Shepperd, del. . . .172
9. Stylodonta unidentata Chemn. Shepperd, del. . . 149
10, 12. Ampelita (Poecilostylus) viridis Desh. Moll. Madag. 158
PLATE 39.
1-5. Camsenella platyodon Pfr. Pilsbry, del. . . .106-
6. Obba basidentata Pfr. After Semper, .... 107
7, 8, 11, 12, 13. Obba planulata Lam. After Semper, . 10&
9, 10. Neocepolis merarcha Mab. Shepperd, del. . . 107
PLATE 40.
1-4. Plectopylis jovia Mabille. Pilsbry, del. . . . 144
5, 6. Plectopylis achatina Gray. Conchy!. Cab., pi. 6f>, . 144
7. Plectopylis achatina Gray. P. Z. S. 1874, . . .144
8. Plectopylis achatina Gray. Pilsbry, del. . . . 144
9-11. Plectopylis ponsonbyi Godw.-Aust. Shepperd, del. . 144
12. Plectopylis ponsonbyi Godw.-Aust. Pilsbry, del. . 144
13-15. Plectopylis fultoni Godw.-Aust. Shepperd, del. . 144
16-18. Mollendorffia hensaniensis Gredl. Ann K. K. Mus.
1887, 289
PLATE 41.
19-22. Gorilla rivolii Desh. Shepperd, del 147
23-25. Gorilla charpentieri v. hinidunensis Nev. Shepperd,
del 147
26,27. Traumatophora triscalpta Mart. Novit. Conch. . 146
28, 29. Stegodera angusticollis Mart. Novit. Conch. . . 147
30. Albersia granulata Quoy. Voy. Astrol. . . .124
31-33. Ampelita hemioxia Pils. Shepperd, del. . . . 155
PLATE 42.
34-36. Plectopylis cyclaspis Bens. J. A. S. Beng. xl. . .144
37, 38. Corilla erronea Alb. Keisen Phil 147
39. Trachia penangensis Stol. J. A. S. Beng. xlii, . .115
40. Ampelita loucoubeeusis Crse. After Brancsik. . . 156
41-46. Caryodes dufresnii Leach. After Semper, Hedley
and Ten .-Woods, 162
PLATE 43.
19-21. Helicigona (Elona) quimperiana Fer. After Hidalgo, . 307
22, 23. Helicigona lapicida Linn. After Hidalgo, . .298
24, 25. Helicigona (Fruticocampylsea) narzanensis Kryn.,
after Kobelt, .... .304
REFERENCE TO PLATES. 357
FIGURE. PAGE.
26. Geomitra (Plebecula) punctulata Sowb. Shepperd, del. 239
27, 28. Helicigona (Chilostoma) planospira Lam. After Kobelt 299
29,30. Helicella (Xerocampylsea) zelebori Pfr. After Kobelt, 253
31. 32. Helicigona (Isognomostoma) personata Drap. Shep-
perd del. . . 308
33, 35. Helicigona (Tacheocampylsea) raspailii Payr. Moll.
Nouv.,Litig. . . ' . . ! . . 305
36. Leptaxis (Pseudocampylsea) lowei Fer. Shepperd, del. 292
37, 38. Helix (Euparypha) pisana Miill. After Hidalgo, . 335
39, 40. Allognathus grsellsiana Pfr. After Hidalgo, . . 290
41. Leptaxis undata Wood. Shepperd, del 292
42. Helicigona (Chilostoma) setosa Ziegl. After Rossm. . 299
43. Helix (Hemicycla) plicaria Lam. Shepperd, del. . . 326
44. Helix (Hemicycla) saulcyi Orb. Shepperd del. . . 326
45. Leptaxis webbiana Lowe. Shepperd del. . . . 292
46. Helicigona (Arianta) arbustorum L. After Kobelt, . 306
PLATE 44.
1-3. Helix (Levantina) guttata v. sesteri Gall. Bull. Soc.
Mai. Fr 332
4, 5. Helix (Tachea) nemoralis L. After Hidalgo, . . 321
6, 7. Helix (Helicogena) asemnis v. vetusta Mts. Archiv
Naturg., 1889, 317
8. Helix (Iberus) gualtierana L. After Hidalgo, . . 328
9, 10. Helix (Otala) vermiculata Miill. After Hidalgo, . 323
11. Helix (Otala) lactea Mull. After Hidalgo, . . .323
12,13. Helix (Erernina) desertorumForsk. After Rossm. . 334
14. Helix (Eremina) desertorum v. chilembia. After Bgt. 334
15. Helix (Iberus) scabriuscula Desh. After Rossm. . . 328
16. 17. Helix (Iberus) muralis Miill. Shepperd, del. . . 228
18. Helix (Iberus) sicana Fer. After Rossm. . . .328
PLATE 45.
1, 2. Oxychona costaricensis Roth. Shepperd, del. . .189
3-5. Oxychona altispira Mart. Biol. Centr. Amer. . . 1 89
6. Epiphragmophora fidelis Gray. Shepperd, del. . .194
7. Lysinoe ghiesbreghti Nyst. Biol. Centr. Amer. . .191
8. Oxychona bifasciata Burrow. Viag. al Pacif. . .189
9, 10. Oxychona trigonostoma v. stolliana Mts. Biol. Centr.
Amer 189
PLATE 46.
11. Panda falconeri Reeve. Conch. Icon 163
12. Panda atomata Gray, apex. After Hedley, . . .163
13. 14. Panda larryi Braz. After Cox, . . . .163
15. Caryodes dufresnii Leach. After Hedley, . . .162
358 REFERENCE TO PLATES.
FIGURE. PAGE.
16. Caryodes dufresuii Leach. Shepperd, del. . . . 162
17-19. Papuina ianthe Smith. Shepperd, del. . . . 137
20. Solaropsis serpens Martyn. Shepperd, del. . . . 166
21. Solaropsis braziliana Fer. Ostas. Conch. . . .166
PLATE 47.
1. Panda falconer! Reeve. After Semper, . . . .163
2. 4. Panda atomata Gray. After Hedley, . . . .163
3. Panda atomata Gray, opened penis. Pilsbry, del., . . 163
5. Anoglypta launcestonensis Rve., baseofsp. d. Pilsbry, del. 160
6, 7. Anoglypta launcestonensis Rve. Pilsbry, del. . . 160
8. Anoglypta launcestonensis Rve., penis. Pilsbry, del. . 160
PLATE 48.
9. Stylodonta studeriana Fer. Pilsbry, del. . . .150
10, 11. Anoglypta launcestonensis Rve. Pilsbry, del. . . 160
12, 13. Helicophanta magnifica Fer. Pilsbry, del. . . 152
14. Acavus hsemastomus L. Pilsbry, del. .... 153
15, 16. Panda atomata Gray. Pilsbry, del. .... 163
17. Panda falconeri Reeve. After Semper, . . . .163
PLATE 49.
18. Helicophanta goudotiana Fer. After Brancsik, . .151
19-23. Helicophanta magnifica Fer. Pilsbry, del. . . 151
24. Caryodes dufresnii Leach. After Semper, . . .162
25. Ampelita sepulehralis Fer. Pilsbry, del. . . . 155
PLATE 50.
1,8,26. Acavus skinneri Rve. After Semper, . . . 153
3. Acavus hsemastomus L. After Semper, . . .153
4. Acavus phoenix Pfr., egg. Shepperd, del. . . .153
5. Acavus phoenix Pfr., teeth. After Binney, . . .153
6. 7, 9. Stylodouta studeriana Fer.. (penis below). Arch.
Zool.Gen.etExper 150
1, 22, 57, 62, 69. Stylodonta studeriana Fer. After Binney, 150
PLATE 51.
1,2. Macrocyclis laxata Fer. Pilsbry, del. . . . 165
3. Dorcasia globulus Miill. After Binney, . . .172
4-8. Polymita picta Born. Pilsbry, del. * .
9-11. Oxychona bifasciata Burrow. Pilsbry, del. . .189
REFERENCE TO PLATES. 359
PLATE 51a.
FIGURE. PAGE.
[By error this number was duplicating in preparing the plates.
As the figures are entirely different, there need be no confusion in
actual reference from the text].
1-6. Ampelita xystera Val. (from no. 63,879 Acad. coll.
Pilsbry, del 155
7-12. Thersites (Khagada) solorensis Mart. After Wiegm. 135
PLATE 52.
12. Cepolis (Coryda) alauda Fer. Pilsbry, del. . . .182
13. Cepolis alauda, penis with flagellum. The thread-like
retractor should pass over v.d. and insert on penis, 182
14. Cepolis (Hemitrochus) varians Mke. Pilsbry, del., . 184
15. Cepolis (Plagioptycha) salvatoris Pfr. Pilsbry, del. . 185
16. Cepolis (Cysticopsis) cubensis Pfr. Pilsbry, del. . .187
17. Epiphragmophora cordovana Pfr. After Strebel, . . 197
18. 19, 20, 22. Cepolis (Eurycampta) bonplandi Lam. Pils-
bry, del. . .... 181
21. Cepolis (Jeanneretia) parraiana Orb. After Poey, . 180
PLATE 53.
1. Helicosyla (Orustia) versicolor Mlldff. Shepperd, del. . 225
2. Helicostyla (Helicobulinus) turbinoides Brod. Shepperd,
del 227
3. Helicostyla (Orthostylus)pithogaster Fer. After Hidalgo, 227
4. Helicosyla (Canistrum) ovoidea Brug. Shepperd, del. . 230
5. Helicostyla (Phengus) opalina Sowb. After Pfeiffer, . 230
6. Helicostyla(Hypselostyla)connectens Mlldff. After Mlldff. 228
7. Helicostyla mirabilis Per. After Hidalgo, . . . 224
8. Helicostyla (Chrysallis) chrysalidiformis Sowb. After Eve. 231
9. Helicostyla (Prochilus) virgata Jay. Shepperd, del. . 231
10. Helicostyla (Cochlodoyas) viridostriata Lea. Shepperd,
del. . 225
11. Helicostyla (Eudoxus) effusa Pfr. After Keeve, . . 229
PLATE 54.
1. Helicosyla (Canistrum) stabilis Jonas. After Semper, . 230
2-4. Chloneabenguetensis Semp. After Semper, . . 215
5. Helicostyla (Orustia) monticula Sby. After Semper, . 216
6-8. Helicostyla butleri Pfr. After Semper, . .' .216
9. Helicostyla (Orthostylus) pithogaster Fer. After Semper, 217
10. Helicostyla (Calocochlea) festiva Don. After Semper, 216
11. Helicostyla (Eudoxus) segle Brod. After Semper, . 216
12. Helicostyla (Calocochlea) pulcherrima Sowb. After Semper, 216
360 REFERENCE TO PLATES.
PLATE 55.
FIGURE. PAGE.
1, 2. Eulota fruticum Miill. After Hartmann, . . .202
3, 4, Eulota fruticum Miill. After Dupuy, ... . 202
5. Eulota duplocincta Martens. After Martens, . . . 202
6, 7. Eulota (Cathaica) fasciola Drap. After Philippi, . 206
8, 9. Eulota (Pseudiberus) tectumsinense Mts. Shepperd, del. 207
10, 11. Helicostyla (Crystallopsis) ten imb erica Mild tf. Nach.
1892," 220
12. Helicostyla (Corasia) virgo Brod. After Hidalgo, . 219
13. Helicostyla (Calocochlea) pulcherrima Sowb. Shepperd,
del 222
14. Helicostyla (Anixa) moreletiana Pfr. Shepperd, del. . 223
15-17. Chloraea sirena Beck. Shepperd, del. . . .215
18. Ganesella capitium Bens. Shepperd, del. . . .168
19. Eulota similaris Fer. Shepperd, del 203
20. 21. Hygromia cinctella Drap. After Dupuy, . .271
22-24. Hygromia (Ciliella) ciliata Ven. After Dupuy, . 276
25,26. Hygromia (Dibothrion) bidens Chem. Shepperd, del. 278
27,28. Hygromia (Fruticicola) hispidaL. After Dupuy, . 273
29, 30. Hygromia (Monacha) incarnata Miill. Shepperd, del. 271
31, 32. Vallonia pulchella Miill. Pilsbry, del. . . 282
PLATE 56.
1, 2. Cepolis (Hemitrochus) varians Mke. After Binney . 183
3, 4. Cepolis (Coryda) alaudaFer. Shepperd, del. . . 181
5. Cepolis (Dialeuca) nemoraloides Ad. Shepperd, del. . 183
6. Cepolis (Dialeuca) fuscocincta Ad. Shepperd, del. . 183
7. Cepolis (Cysticopsis) cubensis Pfr. Shepperd, del. . 1 86
8. 9. Cepolis (Plagioptycha) duclosiana Fer. Shepperd, del. 185
10. Polymita picta Born. Shepperd, del. .... 188
11, 12. Leucochroa (Sphincterochila) boissieri Charp. After
Kobelt, . . . 234
13, Leucochroa candidissima Drap., large var. After Kobelt, 232
14,15. Leucochroa cariosa Oliv. After Bgt. . . . 232
16, 17. Helicodonta (Trissexodon) constricta Boub. After
Kobelt, 285
18,19. Helicodonta (Caracolina) lens Fer. After Kobelt, . 285
20-22. Moellendorffia erdmanni S. & B. After Boettger, . 289
23,24. Helicodouta (Aspasita) triaria Friv. After Kobelt, 285
25-27. Helicodonta obvoluta Drap. After Dupuy, . .285
28-30. Helicodonta biconcava Hde. After Heude, . . 285
31-33. Helicodonta (Drepanostoma) nautiliformis Porro.
After Rossm. . . . 285
PLATE 57.
34-39. Chalepotaxis infantilis Gred. Jahrb. D. M. Ges. xi, 167
40, 44. Cepolis (Cysticopsis) cubensis Pfr. Pilsbry, del. . 187
REFERENCE TO PLATES. 361
FIGURE. PAGE.
41, 50, 51. Cepolis (Hemitrochus) varians Mke. Pilsbry, del. 188
42, 47. Cepolis (Plagioptycha) salvatoris Pfr. Pilsbrv, del. 185
43, 48. Cepolis (Dialeuca) platystyla Pfr. Pilsbry, del. . 183
45,49. Cepolis (Cpryda) alaudaFer. Pilsbry, del. . . 182
46. Cepolis (Hemitrochus) milleri Pfr. After Binney, . 188
52. Leucochroa candidissima Drap. Pilsbry, del. . . 232
53. Spermatheca of same with duct and diverticulum, . 232
PLATE 58.
54. 55. Cepolis (Jeanneretia) parraiana Orb. Shepperd, del. 180
56. Cepolis (Eurycarapta) bonplandi Lam. Shepperd, del. 181
57. Epiphragmophora (Angrandiella) angrandi Morel. Ser.
Conch 197
58,59. Fpiphragmophora (Pilsbry a) farrisii Pfr. Novit.Conch. 197
60, 61. Epiphragmophora (Pilsbrya) patasensis Pfr. Shep-
perd, del., 197
62, 63. Epiphragmophora (Helminthoglpta) tudiculata Binn.
Terr. Moll. Ill, 198
64, 65. Epiphragmophora (Monadenia) mormonum Pfr.
Shepperd, del. .... .198
66, 67. Epiphragmophora (Micrarionta) areolata Sowb.
Shepperd, del. 197
68,69. Epiphragmophora cuyana Strob. Novit.Conch. . 196
70, 71. Epiphragmophora (Trichodiscma) coactiliata Fr.
Shepperd, del 197
72. Epiphragmophora (Monadeuia) fidelis Gray. Shepperd,
del . .198
73, 74. Epiphragmorphora (Micrarionta) gabbi Newc. Shep-
perd, del 197
75. Lysinoe humboldtiana var. Biol. Cent. Amer. . . 191
PLATE 59.
76. Epiphragmophora nickliniana Lea, atrium everted. Pils-
bry, del 194
77. Epiphragmophora hieronymi Doring. After Kobelt, .196
78. Epiphragmophora semiclausa Mart. Mai. Bl. xv, ., . 196
79. Epiphragmophora (Helminthoglypta) arrosa Old. After
Semper,. . . . ' . . . . . 194
80. Epiphragmophora areolata Sowb. Pilsbry, del. . . 197
81. Epiphragmophora (Monadenia) fidelis Gray. Pilsbry, del. 198
82. 86. Epiphragmophora (Monadenia) mormouum Pfr. Pils-
bry, del 194
83. Epiphragmophora nickliniana Lea. Pilsbry, del. . . 197
84. Epiphragmophora fidelis Gray. Pilsbry, del. . . 198
85. Epiphragmophora arrosa Gld. Pilsbry, del. . . 197
362 REFERENCE TO PLATES.
FIGURE. PAGE.
87. Epiphragmophora (Helminthoglypta) traskii v. cayama-
censis Hemph. Pilsbry, del. .... 197
88. Epiphragmophora (Helminthoglypta) nickliniana Lea.
Pilsbry, del. . . , . . . . .197
89. Epiphragmophora (Micrarionta) areolata Sowb. Pilsbry,
del 197
PLATE 60.
1, 2. Ganesella japonica Pfr. Pilsbry, del. . . . 168
3,6. Dorcasiaalexandri Gray. Pilsbry, del. . . . 172
4. Epiphragmophora areolata Sowb. Pilsbry, del. . . 197
5. Lysinoe humboldtiana Fer. After Binney, . . . 191
7. Epiphragmophora nickliniana Lea. Pilsbry, del. . . 198
8. Lysinoe ghiesbreghti Nyst. After Fischer, . . . 191
9. Lysinoe eximia Pfr. After Fischer, . . . .191
10. Epiphragmophora fidelis Gray. Pilsbry, del. . . 198
PLATE 61.
1-5. Helix (Euparypha) pisana Mull. After Schuberth
and Ashford, . 335
6. 7. Helix (Iberus) muralis Mull. After Schuberth, . 329
8. Helix (Iberus) gualtierana L. After Schmidt, . . 329
9. Helix (Helicogena) asemnis Bgt. After Schuberth, . 317
10. Helix (Levantina) spiriplana v. hierosolyma. After
Schmidt, . . , 332
11. 13, 14. Helix (Helicogena) aspersa Mull. After Ashford, 317
12. Helix (Helicogena) pomatia L. After Schmidt, .
15. Helix (Helicogena) pomatia L. After Ashford, . . 317
PLATE 62.
16. Helicigona rhsetica Mouss. After Schuberth, . . 296
17. Helicigona cingulata Stud. After Schmidt, . . . 296
18. Helicigona planospira Lam. After Schuberth, . . 296
19. Helicigona (Isognomostoma) personata. After Schmidt, 308
20. Helicigona lapicida L. After Schmidt, . . . 298
21. Helicigona lapicida L., dart. After Ashford, . . 298
22. Helicigona (Arianta) arbustorum L., dart, after Ashford, 306
23. Helicigona (Arianta) arbustorum L. After Schmidt, . 306
24-27. Helicigona (Elona) quimperiana Fer. After Hesse, 307
PLATE 63.
1-3. Helix (Iberus) serpentina Fer. After Wiegmann, .329
4. Helix (Eremina) desertorumForsk. After Semper, .334
5, 8. Helix (Otala) vermiculata Mull. After Wiegmann, . 323-
REFERENCE TO PLATES. 363
FIGURE. PAGE.
6. Helix (Eremina) desertorum Forsk. After Jickeli, . 334
7. Helicigona (Tacheocampylsea) raspaili Payr. After
Moq.-Tand 305
9. Vallonia pulchella Miiller. After Lehmann, . . 282
10. Vallonia pulchella Miiller, dart. After Ashford, . . 282
11. Acanthinula lamellata Jeffr. After Lehmann, . .280
12. Helix (Tachea) nemoralis Mull. Pilsbry, del. . . 321
13. Helix (Otala) alonensis Fer. After PfefFer, . . 323
PLATE 64.
1, 2. Eulota (Euhadra) peliomphala Pfr. After Kobelt, . 212
3. Eulota (Euhadra) amaliseKob. After Kobelt, . .212
4-0. Eulota (Armandia) calymma S. & B. After Boettger, 205
7. Ganesella japonica Pfr. After Kobelt, .... 168
8. Eulota (Acusta) ravida Bens. After Heude, . . . 203
9. Eulota (Euhadra) qusesita v. perry i Jay. Shepperd, del. 212
10-12. Aulacospira scalatella Mlldff. Pilsbry, del. . . 279
13-15. Eulota (Aegista) oculus Pfr. Shepperd, del. . . 210
16,17. Eulota (Plectotropis)mackensii A. &R. Shepperd, del. 208
18, 19. Eulota (Plectotropis) elegantissima Pfr. Shepperd,
del. 208
20-23. Eulota(Coccoglypta)pinchonianaHde. After Heude, 211
PLATE 65.
1, 2. Eulota (Acusta) ravida Bens. Pilsbry, del. . . 203
3, 4. Eulota (Eulotella) similaris Fer. Pilsbry, del. . . 203
5, 6. Eulota (Eulotella) duplocincta Mart. After Schacko. 203
7, 8. Eulota (Cathaica) fasciola Dr. (pyrrhozona). Pilsbry,
del 206
9, 10. Eulota (Aegista) platyomphala Mlldff. Pilsbry, del. 210
11, 12. Eulota (Eubadra) qusesita perryi Jay. Pilsbry, del. 213
13, 14. Eulota (Piectotropis) vulvivaga S. & B. Pilsbry, del. 208
15-17. Eulota (Cathaica) przewalskii Mart. After Schacko. 206
PLATE 66.
18, 19. Eulota fruticum Drap. After Schuberth, . . 203
20. Eulota similaris Fer. Pilsbry, del 203
21-23. Eulota (Acusta) ravida Bens. Pilsbry, del., . . 203
24. Eulota fodiens Pfr. After Semper, . . . .203
25. Eulota (Cathaica) przewalskii Mart. After Schacko. . 206
26. Eulota (Mastigeulota) kiangsinensis Mart. Pilsbry, del. . 211
(vas deferens omitted by lithographer).
27. 29. Eulota (Euhadra) qusesita v. perryi Jay. Pilsbry,
del. . 213
364 REFERENCE TO PLATES.
FIGURE. PAGE .
30,31. Eulota duplocincta Mart. After Schacko. . .203
32. Eulota (Cathaica) fasciola Drap. (pyrrhozona). Pilsbry,
del . . . .206
33, 34. Eulota (Plectotropsis) vulvivaga S. & B. Pilsbry, del. 208
PLATE 67.
1-3. Helix (Tacbea) nemoralis L. Pilsbry, del. . 321
4, 5. Helix (Otala) vermiculata Miill. Pilsbry, del. 323
6, 7. Helix (Iberus) serpentina Fer. After Wiegmann, 329
8. Helix (Erernina) desertella Jickeli. After Jickeli, 334
9, 10. Helix (Eremina) desertorum Forsk. After Jickeli, 334
11, Helix pomatia L. Pilsbry, del. .... 317
12, 16. Helicella (Trochula) terrestris Penn. After Binney 262
13, 14. Helicella galloi Kobelt. Pilsbry, del. . . 245
15. Helicella (Theba) cantiana Mont. After Binney, . 265
17. Helicella ericetorum Miill. After Scbuberth, . 252
18. Geomitra abjecta Lowe. Pilsbry, del. . . . 238
19. 20. Leptaxis undata Lowe. Pilsbry, del. . . 292
PLATE 68.
1-3. Geomitra delpbinula Lowe. Shepperd, del. . . 244
4, 5. Geomitra (Actinella) lentiginosa Lwe. Pilsbry, del. . 241
6. Geomitra tiarel la Webb & Berth. Pilsbry, del. . . 244
7. Geomitra (Callina) fausta Lowe. Pilsbry, del. . , 241
8. Geomitra (Discula) polymorpha v. barbosse Pva. After
Paiva, 242
9. Geomitra (Discula) polymorpha Lwe. Pilsbry, del. . 242
10, 11. Geomitra (Disculella) maderensis Wood. Pilsbry, del. 243
12. Geomitra (Hystricella) bicarinata Sowb. Pilsbry, del. . 242
13. Geomitra (Spirorbula) obtecta Lowe. Pilsbry, del. . 241
14. 15. Geomitra (Lemniscia) michaudi Desh. Pilsbry, del. 240
16-18. Geomitra (Heterostoma) paupercula Lowe. Pilsbry,
del. . . 244
19. Geomitra (Caseolus) compacta Lowe. Pilsbry, del. . 242
20. Helicella variabilis Drap. After Moq.-Tand. . . 248
2b, 22. Helicella ericetorum Miill. After Rossmaessler, . 252
23, 24. Helicella (Jacosta) explanata Mull. After Hidalgo, 258
25,26. Helicella (Theba) carthusiana Miill. After Moq.-Tand. 265
27. Helicella (Trochula) terrestris Penn. After Hidalgo, . 262
28. Helicella (Candidula) candidula Stud. After Moq.-Tand. 253
29. Helicella (Cochlicella) acuta (barbara L.). After Moq.-
Tand 263
30. Helicella (Platytheba) nummus Ehrenb. After Bgt. . 268
REFERENCE TO PLATES. 365
PLATE 69.
EIGURE. PAGE.
1, 2. Helicella variabilis Drap. After Schuberth, . .248
3-5. Helicella virgata Da Costa. After Ashford, . . 248
6, 7. Helicella ericetorura Mull. After Ashford, . ._252
8. Helicella ericetorum Mull. After Schuberth, . . 252
9. Helicella caperata Mont. After Ashford, . . . 254
10. Helicella candidula Studer. After Schmidt, . . 254
11. Helicella mograbiria Mor. After SchepmaD, . . . 260
12. Helicella tuberculosa Conr. After Schmidt, . . .261
13. Helicella striata Mull. After Schmidt, . . . 254
14. Helicella cantiana Mont. After Ashford, . . . 265
15. Helicella explanata Mull. After Moq.-Tand., . . 258
16. Helicella syriaca Ehrenb. After Schuberth, . . . 265
17. Helicella nummus Ehrenb. After Schmidt, . . .268
18. Helicella elegans=terrestris Penn. After Schmidt, . 262
19-21. Helicella acuta (barbara). After Moss & Paulden,
Trans. Manch. Mic.Soc., 1892, . . . .263
22. Helicella Carthusian a Miill. After Schuberth, . .265
PLATE 70.
23-25. Zoogenites harpa Say. After Morse, . . .281
26-28. Acanthinula aculeata Miill. After Dupuy & Drouet. 280
29. Vallonia pulchella Miill. After Sterki, . . 282
30. Hygromia ciliata Ven. After Moq.-Tand.
31. Hygromia pellita Fer. After Hesse,
32. Hygromia limbata Drap. After Moq.-Tand
33. Hygromia hispida Linn. After Schmidt,
34. Hygromia incarnata Mull. After Schmidt,
35. Zoogenites harpa Say. After Morse,
36. Hygromia leucozona Ziegl. After Schuberth
37. Hygromia noverca Friv. After Hesse, .
38. Vallonia pulchella Miill. After Sterki, .
39. Hygromia cinctella Drap. After Moq.-Tand
40. Plebecula lurida Lowe. After Binney, .
41. Hygromia bidens Chemn. After Schmidt,
PLATE 71.
276
276
271
273
271
281
277
277
282
271
239
278
42, 43. Helicella (Lejeania) scioana Poll. After Pollonera, 267
44. Helicella (Lejeania) lejeaniana Bgt. After Pollonera, . 267
45, 46. Helicigona rahtii A. Brn. After Sandberger, . . 310
47,48. Helicigona ehingensis Klein. Ross del. . . .310
49. Helicodonta osculum Thorn. After Klika, . . . 289
50. Cyrtochilus expansilabris Sdb. After Sandberger, . 311
51. 52. Prothelidomus acrochordon Oppenh. After Oppenh. 295
53, 54. Dentellocaracolus damnatus Sandb. After Oppenh. 295
55,56. Fridolinia lucani Tourn. J. de C., 1869, . . 294
366 REFERENCE TO PLATES.
FIGURE. PAGE.
57, 58. Pseudoleptaxis corduensis. After Sandberger, . . 294
59, 60. Helicigona lepidotricha Braun. After Sandberger, 309
61, 62. Helicigona chaixii Mich. After Sandberger, . . 311
SUMMARY. This volume contains 561 figures, illustrating over
300 species of shells, and 571 figures illustrating the anatomy of
Helices; a total of 1132 figures. Of these, 330 figures were drawn
by the author; 203 other original figures were drawn by Messrs.
Win. and Edw. Shepperd, and about a dozen are from unpublished
drawings supplied by friends. The figures of Flammulina cressida,
thaisa and costulata Hutton, were drawn by Mr. H. Suter from the
type specimens, these species being here for the first time illustrated.
The figures of Plectopylis ponsonbyi (pi. 40, fig. 9-12), and Papuina
ianthe (pi. 46, fig. 17-19) are from examples kindly loaned by Mr.
John Ponsonby from his collection. The other original figures are
from specimens in the collection of the Academy, where also most
of the author's dissections over 500 in number are preserved.
PLATE 1
HELICID>e.
80
PLATK 2
y? ^
10.
00?
20.
21.'
17.
j&p.
trnt ~\
UNIVERSITY
HELICID>e.
PLATE 3
S-frg/V
27
28
rf
29
XJNIV1BSIT*
HELICID^E.
PLATE 4
*
44
UNIVEB8ITI
HELICID^E
PLATE 5
HELICID>G.
PLATE 6
HELICID^E.
PLATE 7
'' 21
PLATE 8
HELJCID^G.
tJHIVEBSIIl
HELJCID^E
PLATE 1O
13
14
HELICID^E.
PLATE 11
HELJC1D.4S.
PLATE 12
HELICID^C
PLATE 13
C
HELJCID./E:
PLATK 14
44 45
46
47
48
UBRj/^v
Of THf ^ \
EBSril J
PLATE 15
PLATE 16
10
: 18
; 19
20
UKITBBBIffTl
HELICID^.
PLATE 17
HELICID^E.
PLATE 18
XJSIVBB8I"
5
HELICID^E.
PLATE 49
HELJCID^E.
PLATE 2O
26
27
30
31
HELICIDvC.
PLATE 21
J
14
HELICID^E
PL-ATE 22
tJSIVBBSIIl
PLATE 23
HELICID>E.
PLATE 24
PL- ATE
tJNIVEBSITT
HELICID^E.
PLATE 26
PLATE 27
PLATE 28
PLATE 29
TJNIVEB8IT1 ]
HELICID^E.
PLATE 3 O
HELICID^E.
*
PLATE 32
IDHIVBBBUt
HELICID^E.
PLATE 33
HELICID^E.
PLATE 34
HELICID^E
PLATE 35
HELICID^E
PLATE 36
PLATE 37
10
II
CXLIFOW**:
PLATE -38
HELICID>E
PLATE 39
tJNIVEBSMl
PLATE 4O
PLATE 41
HELICID^E
43
PLATE 43
46
HELICID>E.
PLATE. 44
13
HELICID>.
RLATE 45
XJNIVEBSITl
HELICIDvE.
PLATE 46
iff
i
Of TH
UNIVEBBITT
HELJCID^G
HELICID^E
PLATE 48
IS
17
UNIVEBBIlt
.C4Ufbw%
HELJCID^E.
PLATE 49
HELJCID^E
PLATE 5O
HELICID>e
PLATE 51
PLATE 51
HELICTFJ^E
PLATE 52
2\
HELICID^E.
PLATE 53
PLATE 54
HELICID^E.
PLATE 55
\
30
OSIVEBSITl 1
HELICID>
PLATE 56
31
30
HELICiD^E.
PLATE 57
PI-ATE 58
HELJCID>e
PLATE 59
87
TH
UNIVEB8IM
HELICID^E
PLATE 6O
HELICID^.
PLATE 61
PLATE 62
HELICID^E.
PLATE 63
13
PLATE 64
^
UNIVBBSItl]
X
HELICiD>E.
PLATE 65
HELJCID^E.
PLATE 66
33
HELICID/E.
PLATE 67
HELICID>e.
PLATE 68
30
PLATE 69
19
Of THE
TJNIVBBSIT1
HELICID^E.
PLATE 7O
HELICID^E.
PLATE 71
62
.v. .?'*.
U.C.BERKELEY LIBRARIES
TVNews
OpenLibrary