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Full text of "Manual of conchology; structural and systematic. With illustrations of the species. Second series: Pulmonata"

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Vol. IX. 


3?iablislied by Concliologioal Section 




*. Of 




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- 


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- 

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. 


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 

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. 





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. 







11 Campylcea" 



hora calif or niensis, 












" Mesodon" 



Tliei sites 
















The list is capable of indefinite extension ; and even those minor 
groups called " sections " often show the same series of changes in 
form, thus: 



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 


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, 

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. 


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 


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, 

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. 



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. 


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- 


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^ 

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 


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 


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. 


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 - 

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 

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- 


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. 


General considerations: Helicidse, like all pulmonates, are her- 
maphrodites, the male and female genitalia uniting below in a com- 


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 


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). 


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. 


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 

IH, 1819-1837. FERUSSACIAN EPOCH. The Tableaux Systema- 
tique de la Famille des Limacons presented the first consistent attempt 


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 

subgenus Helicella [depressed, mostly simple 

lipped Helices and Zonitidce]. 
subgenus Helicostyla [^elevated Helices, not 

Evolutse, Cochloides, subgenus Cochlostyla [=Bulimoid forms, im- 

perforate, with entire mouth], 
subgenus Cochlitoma [ Liguus, Achatina]. 
subgenus Cochlicopa [ Glandina, Stenogyra, 

subgenus Cochlicella, [=Cochlicella, Rumina, 

subgenus Cochlogena [=Limicolaria, Bulimus, 

Achatinella, etc.]. 

subgenus Cochlodonta [=Pupa, Strophia, Gib- 
bus, etc.], 
subgenus Cochlodina ^Cylindrella, Clausilia, 


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 


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. 


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 

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 


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. 


Oenus SAGDA Beck (Teleophallogona). 

Hyalosagda, Proserpinula, Odontosagda. 
Genus LEUCOCHROA Beck (Belogona). 


' Acanthinula, Belogona. 
Vallonia, Belogona. 
Petasia, Belogona. 
Frutiricola, Belogona. 
Dorcasia,Protogon a & Belogona. 

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. 



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~ 


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 

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- 

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 


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, 

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, 
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^. 

Although founded upon the arrangement of Morch, this classifica- 
tion exhibits a distinct advance, not only in the recognition of the 


-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 

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 


\ ** OF 


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. 


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. 


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- 


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. 


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- 

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, 


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 

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. 


Synopsis of genera of Endodontidce. 


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. 


Jaw smooth, S. African, Dorcasia, p. 172. 

f No blind sack 
I on vagina \ 


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. 


Keeled; lip and columella wide, colored, Pyrochi- 
Eggs of mod- lus, p. 154. 

rate size Columella narrow, lip not bright, Atnpelita, p. 

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. 


f Thysanophora, p. 54. 
Teleophallogona -< Sagda, p. 58. 

{Zaphysema, p. 65. 


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. 


Tail with a longitudinal groove above. Shell somewhat Zoni- 
toid, unicolored, with sharp simple lip, neither thickened or 

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 

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. 


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, 


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. 


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 


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. 


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. 


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. 


a. Epiphallus developed, flagellum usually present, but short. 

b. Penis retractor inserted at apex of penis ; Ameri- 
can Pleurodonte, p. 84. 


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. 


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. 


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 



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 

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 


214, Eulota 200, Vallonia 282, Helicodonta 
284, Helicigona 296, Helix 311, Plectopylis- 
143, Gorilla 147. 


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 

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- 


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 


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 


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 

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 


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- 

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 


the glands into the tubular or finger-like form, is characteristic of 

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. 


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 


as the Chinese H. binodata is to certain living and tertiary European 

* * * 

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, 


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. 



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. 


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 


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" 



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. 


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" 




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. 


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 

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 


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. 


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 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. 


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. 


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. 


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 


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. 



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- 

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. 


. y 


( 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 

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 


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 

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. 


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 


(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. 

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. 


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- 

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. 


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 

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 

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. 


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. 


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. 

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. 


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 


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. 


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. 


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 

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 

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. 



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, 

aa. Aperture lacking teeth, internal folds or lirse. 

b. Spire elevated, the alt. nearly equalling or exceeding the 

c. Shell cylindrical, pupiform, Phenacharopa. 

cc. Shell convex-conoidal or thimble shaped, hairy, 


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 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 

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 


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- 


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 


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. 


E. huaheinensis Pfr. iii, 61. 
aranea Behn. 


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. 


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 


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. 


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. 

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. 


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). 


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. 


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- 


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 

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. 



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. 





E. albanensis Cox, ii, 209 ; viii, 
[pi. 37, f. 43-46. 

eastbournensis Beddome & 

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, 


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. 


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 


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 ; 


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. 


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. 


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, 


Subgenus ? SCULPTARIA Pfr., 1856. 

Sculptaria PFR., Malak. Blatter ii, p. 135, type If. sciilpturata 

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. 


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- 


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 . 


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 

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 


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, 


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, 

bb. Internal teeth lamellar ; surface obliquely sculptured, 


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 

Section Pyramidula Fitz., s. sir. 

Pyramidula FITZ., Syst. Verz., p. 95. 

Shell minute, openly umbilicated, with pyramidal spire and obtuse 


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. 


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 

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 

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, 


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- 

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. 


Radula bearing crowded teeth (i n P. perspectives, arranged accord- 
ing to the formula 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. 


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). 


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 

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, 

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. 



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. 


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 


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. 


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. 


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. 

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 


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. 


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 


Euclasta Lederer, Verh. Zool.-bot. Vereinsin Wein, v, p. 252, 1855, 
and Weiner En torn. Monatschr. vii, p. 423, 1863 (Microlepido- 

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. 


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- 

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- 


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 

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. 



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. 

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. 


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 

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. 


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. 


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., 




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 

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. 


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 


r^.-" / 


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- 

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 

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 


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 & 

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, 


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, 


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- 


|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, 


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 

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 


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. 



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. 


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 


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. 

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- 


(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 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 



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- 


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, 
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, 

f. castanea Hemph. 
* * 

v. traversensis Leach. 
t^^T. major Binn., iii, 150. 



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). 


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- 


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. 


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. 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. 


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, 

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. 

Radula long and narrow; teeth after the formula 
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. 


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. 

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. 


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 

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. 


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). 


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- 

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. 


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- 


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. 


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. 


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- 

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, 

* 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., 


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. 



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. 


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. 


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, 


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. 


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). 


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 

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 


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. 


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 

Shell umbilicate, depressed, carinated, microscopically granulated, 
not transversely dilated. Whorls less than 6, the last descending in 


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. 


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. 


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). 


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 

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,, 


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 

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). 


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- 


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 


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 


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. 


C. cicatricosa Mull., vi, 198. v. inflata Mlldff., vi, 199. 

senegalensis Fer. v. ducalis Anc., vi, 199. 

chinensis Voigt. 



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. 


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. 


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 


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 


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- 

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 

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. 


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 

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 


-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. 


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. 


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 

(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. 


(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 


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. 


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, 


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) (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 


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. 


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 


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 

Chloritis may be divided into several sectional groups Chloritis, 
Sulcobasis, Austrochloritis, Trichoehloritis probably natural, but 



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, 


( 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. 


(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. 


(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 

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 

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 


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. 


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 

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. 



( 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, 


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. 


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 


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. 


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 


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 


were perceived simultaneously and independently by Charles Hed- 
ley and the writer. Our knowledge of the anatomy is due to 

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 


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. 

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- 

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 

( 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. 


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. 


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 

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- 

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- 


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. 



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. 


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 


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 


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- 


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. 

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- 



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. 


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 

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. 


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. 



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 



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 


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. 



( 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. 


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. 

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- 


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 


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 

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. 


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. 





(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, 


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, 


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. 


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 


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 


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- 

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. 


Jaw stout, arched, with no ribs, but having a few coarse, broad 
vertical wrinkles (Studeriana), or weak, fine and close striae (uni- 

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 In unidentata the radula measures 10 x 4 
mill., and the formula is 

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- 

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. 


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. 


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, 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- 

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, 


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 

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 


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- 

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 


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 

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. 


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. 


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 

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 

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, 


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- 


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 


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- 

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,, 


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, 

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 

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 


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. 

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. 


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. 


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 


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 


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- 

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 

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. 



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- 


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. 


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. 



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. 

* * * 


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- 


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 


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 

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 : 


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. 


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 

b. Middle and inner lateral teeth 1-cuspid, marginals 3-cuspid, 

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 , 


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 

aa. Old World genera. 

1. Dart sack 1, well-developed. 

b. Mucus gland single, globose, inserted on dart sack, 


bb. Mucus gland acinose ; shell bright colored, CHLOR/EA. 
bbb. Mucus gland divided, lobes sacculated, elongated, 


2. Dart sack wanting; jaw smooth ; shell strong, chalky and white, 



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- 



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 


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 

The sectional divisions are as follows : 

(Cepolia. fPtagioptycha. 

T ,. Cysticopsis. 

-I Jeanneretia. J ^ . J 

-^ Hemitrochus. 



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. 


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, 


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- 

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. 


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. 


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. 

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 


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. 

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. 


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- 

Animal light colored, externally similar to Hemitrochus. 

Jaw high arched, with a median projection (pi. 57, fig. 42, C. sal- 

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. 

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 



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 

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. 



Animal light colored, otherwise as in Coryda. Jaw high arched, 
smooth, with a large median projection (pi. 57, fig. 44, C. 

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. 


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. 


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 


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. 

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. 


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- 

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 


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, 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 


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 ; 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 

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. 


?-|- 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, 


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 

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 


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 

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 


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 


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. 

E. angrandi Morel., v, 96. Peru. 


(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.) 




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. 

(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. 

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. 


. 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 

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 : 


^;* : 

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- 



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. 
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- 

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. 

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- 

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, 



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. 


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- 

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 

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 

(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. 


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. 


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- 

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 

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, 


Group of C. dry ope. 

C. dryope Brod., vii, 100. C. coerulea Mlldff, vii, 101. 

prasina Koch. C. cristatella Mlldff, Nachr., '93, 


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. 


Group of C. t her sites. 

C. thersites Brod., vii, 104. C. malleata Q. & M.. Nachr., '93, 


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 


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 

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 


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. 


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- 

Hydrophanous cuticle present, lip wider, Calocochle<t. 

Dark, depressed and much roughened forms, Trachystyla. 

Dark and smoothish forms, Anixu. 


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, 


Whorls more rapidly increasing ; columella longer, Orthostylus, 

Elongated and bulimoid. 

Imperforate, Hypselostyla, Papustyla, Eudoxus, Phengus, Canis- 

Perforated, Procliilus, Chrysallis. 

Section Corasia Albers, 1850. 

Corasia ALB., Die Hel., p. Ill; second edit., p. 170, type H. 

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. 


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 

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. 


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 


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. 


Section Calocochlea Hartmann, 1842. 

Calocochlea HARTM., Erd- und Siisswasser Gasterop. Schw., p. 
163, type pulcherrima Sovvb. Callicochlias AGASSIZ, 1847, and of 

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. 


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 

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. 


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. 

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. 


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- 

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. 



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 

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. 


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. 


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. 


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. 


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. 

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. 


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. 


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 


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. 



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. 



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, 


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- 


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, 

aaa. Jaw smooth ; teeth all unicuspid and strap-shaped, 


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, 

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, 


bbb. Aperture round, oblique, toothless; shell minute, de-* 
pressed, few-whorled, Vallonia. 


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 


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). 


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 

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 


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, 


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. 


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). 


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. 


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 

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 


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 

=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 

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 


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, 


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 

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- 

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. 


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. 


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. 



"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. 
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. 



H. erithrocheila Sul., viii, 189. 

rufolabris Ben. not Jeffr., iii, 

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. ; 


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.). 



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. 
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- 


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. 



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. 



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. 


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 


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. 


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. 



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. 


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. 


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 


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. 


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- 


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- 

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- 


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. 




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. 



/. 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 

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. 


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 

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 

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. 


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 


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 


in essential points with recent European authorities, and seem to be 

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. 



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., 


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. 



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 

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. 




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). 


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 

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 

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). 


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- 

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 


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 

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. 


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, 


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, 


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 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. 


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- 

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. 


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. 


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, 


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. 

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- 


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 


Risso, it has no claim for adoption, and had better be dropped en- 

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. 


DREPANOSTOMA Porro. Nautiloid, biconcave, with crescentic 

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. 


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. 


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). 


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- 

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 


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 

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 


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- 

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 


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. 


(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. 

(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 


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 

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 


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. 


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. 

1. Aperture oval or lunate ; lip 1-toothed or toothless. 
a. Spire convex or conoidal ; mucus glands tubular. 


b. Acutely keeled ; lip continuous across parietal wall. 

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, 

2. Aperture ear-shaped, 3-toothed, ISOGNOMOSTOMA. 


1. Shell with pappillse arranged in oblique series as in H. setosa, 

a. An obtuse angle around umbilicus ; periphery rounded, 


aa. Umbilicus narrow or closed ; periphery acutely keeled, 


2. Shell without regularly placed pappillse. 

a. Large, depressed, with broad peristome, MESODONTOPSIS. 
aa. Subglobose, spire conoidal, peristome narrow, 


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,. 


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 


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. 



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, 

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. 



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. 



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 



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 


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- 

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 ! 


V e-, "_ 


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. 


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- 


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,. 


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. 


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 

Subgenus GALACTOCHILUS Sandberger. 

Galactochilus SANDB., Land und Siisswasser Conchyl. der Vor- 
welt, p. 387 (for H. pomiformis, mattiaca, ehingensis and cornumili- 

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 


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 ( 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. 

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- 

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 

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- 

Synopsis of sections and subgenera. 
I. Penis provided with a flagellum. 

a. Baso-columellarlip straightened and widened by a callous plate 

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 


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 


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. 

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 

{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. 



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, 


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. 

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- 

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 



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. 

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. 



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 

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. 



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 

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. 



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,. 

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. 



/. 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 

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, 


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 

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 

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 

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 

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. 


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. 


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. 



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. 


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, 

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. 


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. ' 


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 

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- 


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. 


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 



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 


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. 

P. 197, for petasensis read patasensis. 

Genus EULOTA (p. 200). 

Add to Plectotropis, p. 209 : E. luzonica Mlldff., Nachr. '94, p. 

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. 



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, 

^6. Helix pomatia L. Pilsbry, del. ..... 317 albumen gland; air. atrium; d.s. dart sack; epi. epi- 
phallus; fl. flagellum; hermaphrodite gland; 
mucus gland ; p. penis ; r. penis retractor ; sp. spermatheca ; 
sp.d. spermatheca duct; ut. uterus; vag. vagina; v.d. vas 

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 


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 


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 




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 


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 


30-32. Endodonta (Ptychodon) aorangi Suter. Pilsbry, del. . 28 
33, 34. Endodonta (Thaumatodon) derbesiana Cr. Pilsbry, del. 2fr 


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 


45-48. Endodonta (Libera) subcavernula Tryon. Pilsbry 

49, 50, 51. Endodonta ? (Brazieria) velata H. & J. Pilsbry 

52, 53. Endodonta fabrefacta Pse. Pilsbry, del. . 

54. Endodonta (Diglyptus) pagodiformis Sm. Pilsbry, del. 




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 


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 



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 


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, 





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 



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. . 





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 


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 


:;, 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. 

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 



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 



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 




6. Polygyra troostiana Lea. After Binney, . . 71 


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 



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 




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 



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 



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 


PLATE 51a. 


[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 


PLATE 55. 


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 



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 



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- 



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 



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 


PLATE 69. 


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. 


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 



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. 





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