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SECOND SERIES: PULMONATA

MANUAL

OF

CONCHOLOGY

VOL, XXVIII

Geographic Distribution of Pupillidae; Strobilopsidae,
Valloniidae and Pleurodiscidae

BY

HENRY A. PILSBRY

Review of the anatomy of Pupillidae and related groups
By H. Burrington Baker

^'^^»^

PHILADELPHIA :

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT

ACADEMY OF NATURAL SCIENCES /^

OF. PHILADELPHIA i*^f^

^927-1935 LuJLIBRAfi

Dates of Publication, Vol. 28.

Part 109, pages 1 to 48, plates 1 to 8, November, 1927.
Part 110, pages 49 to 96, plates 9 to 12, April, 1931.
Part 111, pages 97 to 160, plates 13 to 23, June 14, 1934.
Part 112, pages 161 to 226, plates 24 to 31, November, 1935.

PREFACE

In this final volume treating of the Pupillidae and allied
families their geographic distribution has been discussed, with
lists of the species of the several zoogeographic provinces,
given for the convenience of those concerned with local faunas.

The classification of Pupillidae and their allies, formerly
based almost wholly upon the shells, has been in large part
placed upon a more satisfactory basis by the admirable ana-
tomical work of C.-M. Steenberg, Hugh Watson and others,
which appeared during the progress of the present mono-
graph. Further data contributed by H. Burrington Baker
are now presented in pages 191-209, plates 26-28. A revision
of the classification of the Pupillidae into subfamilies, accord-
ing to the present views of the author, will be found in the
Introduction, pages vii-xii.

I am indebted to Dr. E. Degner, of the Zoologische Staats-
institut u. Zoologische Museum, Hamburg, for compiling
numerous names omitted from the indices of volumes 24-26.
These have been supplied in the index of vol. 27.

H. A. PiLSBRY.

Philadelphia, October 10, 1935.

(iii)

CONTENTS

PAGE

Introduction: The division of the Orthurethra into families;

The family Pupillidae and its subfamilies . . . vii

Description of Genera and Species: —

Family Strobilopsidae 1, 193

Genus Strobilops Pils 12, 194

Family Pupillidae, Appendices 63, 169

Genus Gastrocopta WoU 63, 117, 169, 203

Genus Gyliotrachela Tomlin 73

Genus Fauxulus Schauf 73, 121

Genus Abida Leach 76, 122, 201

Genus Granopupa Bttg 77

Genus Chondrina Reichenb 78, 122, 201

Genus Pupoides Pfr 80, 137, 200

Genus Microstele Bttg 137

Genus Pupilla Leach 81, 135, 199

Genus Lauria Gray 86, 138, 170

Genus Orcula Held 88, 123, 172

Genus Vertigo Miill 93, 208

Genus Truncatellina Lowe 105, 171, 208

Genus Columella Westerl 109, 172, 208

Genus Sterkia Pils 109

Genus Nesopupa Pils 110, 204

Genus Bothriopupa Pils 114, 207

Genus Pupisoma Stol 114

Geographic Distribution of Pupillidae 139

Family Valloniidae 173

Genus Vallonia Risso 174, 195

Genus Acanthinula Beck 174, 196

Genus Zoogenetes Morse 175

Genus Spermodea Westerl 176

Genus Planogyra Morse 176, 197

Genus Spelaeodiscus Brusina 176

Family Pleurodiscidae 177

Genus Pleurodiscus Wenz 178, 198

Genus Pyramidula Fitz 182

Review of the Anatomy of Pupillidae and Related Groups, by

H. Burrington Baker 191

Explanation of Plates 210

Index .... - 221

(v)

INTRODUCTION

The division of the Orthurethra into families seems to be
largely a matter of expediency. Hugh Watson (1920), after
showing that the various anatomical characters occur in a
great number of combinations, suggested that the entire series
be merged into three families, Achatinellidae, Partulidae and
Pupillidae, the latter with numerous subfamilies. Steenberg
(1925), with more analytical tendencies, proposed to recognize
16 families. The failure to find exclusive character-combin-
ations for families has concerned me since 1900, when the
snails composing the group Orthurethra were first brought
together. The division into ten families in this Manual had
its basis in (1) the recognition of features of the shell as im-
portant family characters, and (2) the idea that penial com-
plications are fundamental, apparently present in the group
as it was before breaking up into most of the existing families,
and thus inherited in nearly all of them. Forms with simple
genitalia are therefore relatively evolved and have arisen
de novo in various families — a view which has been already
either admitted or assumed by several authors. To believe
that similar penial complications have arisen in several
obviously distinct lines seems to stretch the probabilities of
homoplasy too far.

Apertural teeth or lamellae in the shell appear to have
evolved after the initial divergence of the family stocks, as
they do not seem to be traceable to any common type. The
Pupillidae, Cochlicopidae, Tornatellinidae, Partulidae, etc.
apparently each evolved its special type of apertural arma-
ture independently. In some families, such as Pupillidae and
Tornatellinidae, this evolution appears to have been very
early, and many members now show degeneration of teeth.
In others, such as Partulidae and the genus Spelaeodiscus in
Valloniidae, the evolution of teeth seems to have been one of
the latest developments.

The family Pupillidae and its subfamilies. — The Pupillidae
are characterized by the oval to eylindric shape of the shell,

(vii)

vm

INTRODUCTION.

with typically five laminae or teeth within : angular, parietal,
columellar, upper and lower palatal, no marginal teeth on the
peristome. The shape of the shell is changed in some genera,
but not especially in the direction of other families of
Orthurethra. From the strict homologies of the five primary
teeth throughout the family it seems likely that the original
pupillid stock possessed them; but in all of the derivative
lines of descent (subfamilies) there are forms with the full
tooth formula, together with others, in which there has been
simplification or total loss of teeth.

In the collateral family Valloniidae there is no evidence of
a five-plicate aperture in any of the living or fossil forms,
though two of the leading genera are known in numerous
species from the Paleocene to the present time. The char-
acter of being toothless from the earliest appearance has not
I think, been sufficiently appreciated by those who would
unite the Valloniidae with Pupillidae or Strobilopsidae. The
only dentate genus, Spelaediscus, has teeth of a type wholly
different from Pupillidae, and which may reasonably be re-
garded as newly arisen in that genus. The Strobilopsidae
have a complicated system of internal laminae, already de-
veloped in the Eocene and persisting with little change to the
present. I have elsewhere mentioned the possibility of homo-
logizing the principal elements of this armature with pupillid
teeth.

The interrelations of the subfamilies of Pupillidae, as I
understand them, may be represented in a diagram, thus :

Gastrocoptinae Vertigininae

Pupillinae

Orculinae Nesopupinae

Ancestral
Pupillidae-

-Strobilopsidae

Valloniidae-

INTRODUCTION. IX

The upper subfamilies in this diagram are relatively evolved
groups which have lost penial accessories. In Pupillidae I
fail to trace any relation whatever between penial complica-
tion and the development of apertural armature, since four of
the five subfamilies contain forms with highly complicated,
together with others having toothless apertures.

Subfamily Gastrocoptinae.

Gastrocoptinae Pilsbry, 1918, p. x. Includes Chrondrinidae
Steenberg, 1925, p. 201.

Shell varying from minute to the largest of the family,
typically with 5 teeth, the angular and parietal more or less
converging or concrescent; but varying to forms with many
accessory teeth or with no teeth.

Animal oviparous, without penial accessory organs. Jaw
as in Pupillinae;

This subfamily is retained in the limits assigned in Manual
1916-1918 except for the genus Bothriopupa, which was re-
moved to the Vertigininae near Nesopupa in 1926 (vol, 27,
p. 228).

The genera were originally classified in four series. Those
of my fourth group had already been brought into sequence
by Pfeiffer, 1878. Steenberg (1925) formed the family
Chondrinidae for Chondrina, Ahida and Sandahlia, which he
dissected, but H. Burrington Baker in this volume has shown
that Oastrocopta agrees essentially with the European genera
mentioned.

In genera starred the genitalia have been examined.

1. Gibbulina, 24:5. Chaenaxis, 24:1. *Gastrocopta, 24:6.

2. Hypselostoma, 24:175. Boysidia, 24:192. Gyliotrachela,
24:210.

3. Aulacospira, 24:220. Systenostoma, 24:224.

4. Fauxulus, 24:234. Odontocyclas, 24:254. *Sandahlia,
24:250. *Abida, 24:262. Granopupa, 24:332. *Chondrina
(Modicella), 25:1; 27:283.

Dr. H. B. Baker contends that the name Chondrina is in-
admissible under the rules, and should yield to Modicella;
see page 203.

X INTRODUCTION.

Subfamily Pupillinae.

Pupillinae Pilsbry, 1918, vol. 24, p. x ; p. 106 + Orculinae,
part. Pupillidae Steenberg, 1925, p. 201.

The shell is cylindric or tapering, of medium size for the
family, with teeth varying from fully developed {Leiostyla)
to wanting (Pupoides).

Animal either oviparous or viviparous. Penis bifid, with
a long appendix and a forked retractor; spermathecal duet
short or medium, often with a diverticulum. Jaw of very
narrow, conerescent plates, producing a finely striate surface.
Radula with central teeth tricuspid, as wide as the laterals or
narrower; laterals bicuspid; marginals wide, with numerous
narrow cusps.

The limits of this subfamily were materially extended by
Steenberg 's papers of 1925 and 1929, in which Lauria and
Agardhia were shown to be like Piipilla in genitalia. Genera
starred have been dissected.

*Pupoides, 26:103. *Pupoidopsis, 26:106: Microstele,
26:147. Microcerion, 26:151. *Pupilla, 26:152. *Lauria,
27 :43. *Agardhia, 27 :129. Paracoryna, 27 :132, 320. Enneo-
pupa, 27 :222. ?Boysia, 26 :225.

Subfamily Orculinae.

Orculinae Pilsbry, 1918, p. x. Orculidae Steenberg, 1925,
p. 201. See also Hesse- Wiegmann, Archiv. Molluskenk.
56 :1, and Soos, 1. c. 57 :94, for descriptions and figures of
genitalia.

Animal oviparous. Penis terminating in a long blind sac
beyond entrance of the epiphallus and retractor insertion;
no appendix in Orcula and Pagodulina, but it is present in
Orculella. Oviduct provided with a short cul-de-sac. Sper-
mathecal duct long in Orcula, but short or medium in
Orculella. Jaw and teeth about as in Pupillinae.

*Orcula, 27 :1. *Orculella, 28 :92. Pagodulina, 27 :166.

This group is used in the limits established by Steenberg,
who has also pointed out the close resemblance of Moquin-

INTRODUCTION. XI

Tandon's figure of the penis of Pagodulina to that of Orcula.
The homologies of the penial appendages of Orcula and
Orculella have been discussed by H. Burrington Baker, p. 201,
whose conclusions appear well founded.

Orculella, distinguished from Orcula by the possession of
an appendix, and a shorter spermathecal duct, does not appear
to have any conspicuous differential shell characters. It is
this group which approaches the Pupillinae closely, but has
not the bifid penis or forked retractor.

Subfamily Nesopupinae.

Nesopupinae Steenberg, 1925, p. 201. For anatomy see
H. B. Baker, p. 204.

Vertigininae, part, Pilsbry, 1920.

Shell and animal substantially like Vertigininae except that
the penis bears an appendix and the retractor muscle is
forked. Part of the genera are viviparous.

The genera composing the Nesopupinae had been grouped
in a continuous series in my monograph of Vertigininae.
Further anatomical work is needed to confirm the constancy
of the differential characters of the two subfamilies, which
are clearly more closely allied than either is to any other.

It is an old group of world-wide distribution in tropical
and subtropical zones ; it comprises a number of extinct genera,
and is now largely replaced in northern continents by the
Vertigininae, which were apparently derived from nesopupoid
ancestors by simplification of the male organs. Starred
genera have been dissected.

Campolaemus, 25:364. Staurodon, 25:224. Ptychalaea,
25:273; 27:225. *Lyropupa, 25:226. *Nesopupa, 25:274.
Costigo, 25 :336. *Pronesopupa, 26 :1. *Bothriopupa, 24 :226 ;
27 :228. *Pupisoma, 26 :19. Cylindrovertilla, 26 :43. Sterkia,
26:49. Paracraticula, 27 :221. Glandicula, 25 :221. Pseudelix,
25 :222.

The fifth to eighth genera of the above list are evidently
closely allied, and whether all are generically distinct seems
open to question. Ptychalaea may belong to the Vertigininae.

Xii INTRODUCTION.

Subfamily Vertigininae.

Vertigininae Pilsbry, in part, vol. 25, p. 68.

Vertigininae and Truncatellininae Steenberg, 1925, p. 201.

Shell small or minute, usually brownish, compact, oval or
cylindric ; aperture with many teeth or none.

Animal oviparous, usually without tentacles. Penis simple,
without appendages, the retractor muscle simple, attached to
epiphallus. Hermaphrodite gland divided into two groups of
short, wide acini. Male organs are frequently absent. Jaw
formed by the concrescence of relatively few, wide plates
separated by grooves. Central tooth tricuspid, laterals bi- or
tricuspid with interstitial cusplets, the ectocone often about
as long as the mesocone. Marginals usually tricuspid with
numerous interstitial cusplets. The genera fall into two series.
Genera starred have been dissected.

1. *Vertigo, 25:68.

2. •Truncatellina, 26:59. *Columella, 27:232.

MANUAL OF CONCHOLOGY

Family STROBILOPSIDJE.

Helicidcc and Pupidce of various authors.

StrohilidcB Jooss, Jahrb. Nassauisclien Vereins f, Natur-
kunde, LXIV, 1911, p. 61.

StroMlopsincB Pilsbry, Man. Concli. XXIV, 1918, p. x. —
Wenz, Fossilium Catalogus, Gastropoda Extramarina Ter-
tiaria, pars 20, 1923, p. 1041.

Strobilopsidce Hanna, Nautilus XXV, 1922, p. 91. — Steen-
BEBG, fitudes sur I'Anat. et la Syst. des Maillots, 1925, p. 202.

Shell trochiform, dome-shaped or discoidal, umbilicate, of
4l^ to 6 slowly-enlarging whorls. The aperture is small, ob-
lique, with armature of 2 or 3 parietal lamellae and several
deeply-placed basal folds, all growing continuously from an
early neanic stage. Peristome more or less thickened and
expanded, the ends of the lip remote, joined by a parietal
callus.

Urethra lies very near the last part of the intestine. Ovo-
testis forms two groups of follicles. Penis is continued in a
long epiphallus and bears a long appendix, with swollen basal
and distal divisions, the penial retractor bifurcate, one branch
inserted on the epiphallus, the other on the base of the ap-
pendix (distally it attaches to the right ocular retractor,
according to Hanna). The jaw has numerous ribs. Radula
"v^dth tricuspid central tooth with square basal plate, as large
as the bicuspid laterals, the marginals multicuspid (PL 4,
fig. 10).

By the structure of the male organs Strohilops resembles
Vallonia; Papilla, Lauria, the Achatinellida3, and some other
groups are similar in having a bifurcate penial retractor and
a long, tripartite appendix. If Hanna is right in stating that
the penial retractor is a branch of the right ocular band, this
is an important difference from any known orthurethroua

2 STR0BIL0PSlDi3E.

group. The mouth parts do not differ materially from some
Pupillidae.

The shell, aside from its helieoid shape (not a character of
great importance), differs from all Pupillidse in the arrange-
ment of the lamellae and baso-palatal folds. In multidentate
Pupillidae the five primary teeth are always recognizable (see
diagram in Vol. XXV, p. vii) while in Strohilops only the
main parietal lamella and the columellar lamella can certainly
be said to correspond, and these are found in so many other
land shells that their occurrence is not especially significant.
It is possible, however, that upper and lower palatal folds of
Pupillidas are represented by teeth 5 and 2 of fig. 1, and the
basal fold by tooth 1.

Lamellae -{

' Parietal
Interparietal
Infraparietal
Columellar

I

-- — Palatal 1

folds.

Basal

Fig. 1. Section of last whorl of Strobilops at the internal barrier,
showing terminology of teeth.

By the accelerated lamellse and folds of the shell, which
appear early in the neanic stage, Strohilops resembles various
Tomatellinidae (Manual XXII). In that family both pari-
etal and palatal folds or laminae are sometimes present in the
neanic stage. Various Pupillid genera also, such as Orcula
and Lauria, have apertural armature during the neanic stage,
described in this work, Vol. XXVII. Orcula has spiral pari-
etal and columellar lamellae but no basal or palatal folds.
Lauria has basal folds, but they are spaced, transverse bar-
riers, wholly unlike the adult basal or palatal armature of the
species, and differing equally from the folds of inmiature
Strohilops, which from their inception appear to develop con-
tinuously into those of the adult shell. It appears likely that

STROBILOPSIDiE. 3

the acceleration or early appearance of apertural armature in
Tomatellinidae, Orcula, Lauria and Strohilops has been inde-
pendent in the four groups, and is not indicative of direct
relationship between any of them.

On the whole, I am inclined to rank the Strobilopsidse as a
family, distinguished chiefly by characters of the shell.

In the course of my work on the super-family Orthurethra
in this Manual, the problem of defining family groups has
constantly been before me. The partial solutions offered have
not been wholly satisfactory to myself. The difficulties have
been stated lucidly by Hugh Watson (Proc. Malac. Soc. Lon-
don XIV, pp. 20-27; diagram on p. 25). He has called atten-
tion to the fact that a division of the Orthurethra based on
any single character (such as the presence or absence of penial
appendix or flagellum, single or forked penial retractor, pres-
ence of a diverticulum of the spermathecal duct, degree of
elongation of the shell, etc.) would not accord with a division
based upon any other characters. At present the relative
value of these characters appears in most cases to be uncer-
tain. That some forms have been simplified secondarily seems
highly probable. Watson proposed that the Partulidse and
Achatinellidee (including Tomatellinidaj as a subfamily) be
retained as families, all the rest of the Orthurethra to be in-
cluded in Pupillidffi under a number of subfamilies. He does
not recognize characters of the shell as significant for family
grouping, though the construction of his diagram appears to
show that he relied upon them for his subfamily grouping.
Steenberg, whose work, fitudes sur 1 'Anatomic et la Systema-
tique des Maillots, 1925, is the most important document we
have on PupiUid anatomy, would divide the Orthurethra into
sixteen families.

Although the meaning of characters of the shell may be-
come vague or illegible by degenerative changes, convergent
evolution and the like, I am inclined to believe them somewhat
more stable than the details of genitalia in these Orthurethra,
and to utilize them in grouping the genera into families. To
rank the Amastridse, the Cochlicopidge or even the Strobilop-
sidse as subfamilies of Pupillida does not seem to me to clarify

4 STR0BILOPSID.5:.

our conceptions of this intricate group, though it is admitted
that they are not strongly characterized families.

Paleontology.

Strobilopsidae appeared in the Upper Eocene of western
Europe in several species having all the external characters
of the genus Sirohilops, and though the internal structure has
not been worked out, it is safe to assume that they are closely
related to the well-known Oligocene forms following them.
In Europe this genus continued in numerous species into the
Pliocene, the last one in the Upper Pliocene (Astian stage)
of Piedmont.

In late Cretaceous beds there are various forms described
as Helix, or under the names 01)'biniila and PseudostroMlus,
which certainly have some of the characters of Strohilops.
The poor preservation of the very small number of specimens
yet found does not admit of a definite reference to this family ;
their position can only be cleared up by further material.
All are larger than any Strohilops. Notes and references, with
figures of the type species of these groups, follow the list of
Tertiary species of Strohilops.

Genus Strobilops Pilsbry,

The following list is taken from that of W. Wenz, Fossilium
Catalogus, Pt. 20, III, 1923, pp. 1041-1061, which must be
consulted for full references to previous literature. A few
minor alterations and additions have been derived from other
sources.

Strohilops appeared in the upper Eocene in three species.
Oligocene species are more numerous and their structure has
been fully investigated. Some, such as S. headonensis and
S. pseudolahyrinthica, carefully worked out by L. R. Cox,
approximate rather closely to recent species of the typical
labyrinthica group, having the interparietal lamella developed,
the series of internal plicae extending above the periphery, and
the exterior costate. This group continued into the Pliocene,
represented by several species, S. romani (pi. 12, figs. 11-13),
S. lahyrinthicula and others.

STROBILOPSID^. O

Two other series of species appeared first in the Oligocene,
one, Eostrohilops, in which the surface is finely striate, begin-
ning with S. diptyx, continuing to the Pliocene, S. duvali.
The other series, Discostrohilops, contains S. uniplicata (pi. 8,
figs. 10-13), which lived from the upper Oligocene to upper
Miocene in various varietal forms, and is interesting for its
great similarity to the living American S. huhhardi.

In Europe at its first appearance as now known, the com-
plex structure of the genus was apparently about as fully
evolved as in any modern species. In the Oligocene, the sub-
divisions Strohilops proper, Eostrohilops and Discostrohilops
appeared in central Europe, typically developed, probably by
migration from somewhere eastward.

Strohilops doubtless had a long Tertiary history in North
America, still to be recovered. At present it is not knoAvn
here below the Pleistocene. ^S'. lahyrinthica has been identified
from several Pleistocene deposits, the earliest an Iowa deposit
believed to be of Aftonian age. S. affinis and S. Idbyrinthica
occur from the Sangemon on to the Recent, according to
records assembled by F. C. Baker. All of the Pleistocene
localities are v/ithin the areas still inhabited by these species.
In Bermuda S. huhhardi occurs in Pleistocene lime rock (con-
solidated dune formation) and later cave deposits, up to
Recent.

The list of Tertiary species follows. For their classifica-
tion see p. 16.

Strobilops boettgeri (Andreae). Strohilus hoettgeri An-
DRE.VE, Mitt. a. d. Roemer Museum, Hildesheim, No. 18, pp. 5,
10, 26, text-fig. 5. — Strobilops hoettgeri Wenz, N. Jahrb. f.
Min., Geol. u. Pal., 1915, II, p. 80, pi. 4, f. 4a-c; text-fig. 8.
Upper Miocene, Tortonian : Oppeln, Silesia.

Strobilops costata (Clessin). Strohilus costatus Clessin,
Corresp.-bl. Zool. Min. Ver. Regensburg, 1877, p. 'il.— Stro-
hilops costata Wenz, N. Jahrb. Min., Geol. u. Pal. 1915, p. 79,
pi. 4, f. 15a-c; text-fig. 7. — Strohilus hilamellatiLS Clessin,
Malak. Blatter, n. F. VII, 1885, p. 79, pi. 7, f. 10.— Strohilus
curdoofensis Clessin, Bericht Naturwiss. Ver. Regensburg,
XIII (1912), p. 104 = Strohilus xmdorfensis Clessin, t. c,

6 STROBILOPSID^:.

p. 112. Miocene, Tortonian : Undorf bei Regensburg ; Oppeln,
Silesia.

Strobilops desmarestina (Brong.), Helix desmarestina
Brongniart, Ann. du Mus. d'Hist. Nat. XV, 1810, p. 380, pi.
23, f. 10. — Strohilops desmarestina Wenz, Foss. Cat., pt. 20,
III, 1923, p. 1042. Upper Oligoeene, Chattian : Palaisean and
Montmorency; Etampes. Insufficiently known.

Strobilops diptyx (Boettger). Helix diptyx Boettger,
Beitrag paleont. geol. Kenntnis Tertiarform. in Hessen, 1869,
p. 24, pi. 1, 1 5. — Strohilops diptyx Wenz, N. Jahrb. Min.
Geol., Pal. 1915, II, p. 75, pi. 4, f . 6a-c ; text-fig. Z.—StroUhis
clyptyx Klebs, Jahrb. Kon. Preuss. Geol. Reichsanst. 1885
(1886), p. 378. Upper Oligoeene, Chattian: Offenbach a. M.;
Hochheim-Florsheim (Ilessen-Nassau) .

Strobilops diptyx suprema Wenz. Strohilops diptyx var.
suprema Wenz, Jahresber. u. Mittheil. d. Oberrhein. Geol.
Ver., n. f. V, 2, p. 175. Foss. Cat., 1923, p. 1045. Upper
Miocene, Aquitanian: Donanrieden, Wiirttemberg.

Strobilops dollpusi (CoUot). Fitys dollfusi Collot, Feu-
ille Jennes Naturalistes (5), XLI, 1911, p. 93, f. 1. — Stro-
hilops dollfusi Yv^ENZ, Foss. Cat. 1923, p. 1045. Miocene,
Vindobonian: Manthelan (Dep. Indre-et-Loire).

Strobilops duvali (Michaud). Helix diwalii ]\Iichaud,
Joum. de Conchyl. X, 1862, p. 65, pi. 3, fig. U-IG.— Strohilops
duvali Wenz, N. Jahrb. Min., Geol., Pal., 1915, II, p. 82, text-
fig. 11 ; pi. 4, f . 2a-c. Middle Pliocene, Plaicensian : Celleneuve
near Montpellier and Montpellier (Dep. Herault) ; Hauterive
(Dep. Drome) ; Trevoux (Dep. Ain).

Strobilops elasmodonta (Reuss). Helix (Conidus) elas-
modonta Reuss, Sitz.-Ber. Akad. Wissensch. Wien, Math.-Nat.
CI., XLII, 1860, p. 66, pi. 1, f. 2. — Strohilops elasmodonta
Wenz, N. Jahrb. Min., Geol., Pal., 1915, II, p. 77, text-fig. 5,
pi. 4, f . la-c. Lower Miocene, Burdigalien : Kolnsoruk, Lipen
and Tuchorshitz, Czechoslovakia.

Strobilops fischeri Wenz. Strohilus diptyx Klika, Archiv
naturwiss. Landesdurchforschung Bohmen, VII, 1891, p. 34,
text-fig. 26a-c. Not of Boettger. — Strohilops fischeri Wenz,
Jahrb. Nassau. Ver. Naturk. in Wiesbaden, LXVII, p. 107;

STROBLLOPSID.T]]. 7

N. Jahrb. Min. GeoL, Pal., 1915, p. 78, text-fig. 6, pi. 4, f. 5a-c.
Lower Miocene, Burdigaliaii : Tuchorshitz, Czechoslovakia.

Strobilops gedanensis ( Klebs) . Strohilus gedanensis Klebs,
Jahrb. kon. Preuss. geol. Landesanst., 1886, p. 374, pi. 17, f.
Za-d. — Strohilops ged-anensis Wenz, Fossilium Catalogus, 1923,
p. 1047, Lower Eocene : amber of eastern Prussia.

Strobilops HEADONENSis (Edwards). Helix headonensis F.
Edwards, Monogr. Eocene Moll. Palgeontographical Soc. 1852,
p. 70, pi. 11, f. 5a-d. — Strohilops headonensis Wenz, N. Jahrb.
Min., Geol., Pal., 1915, II, p. 73.— L. R. Cox, Proc. Malac. Soc.
London, XVII, 1926, pp. 50, 56, figs. 1-6. Upper Oligocene:
Bembridge Limestone of Headon Hill and Whitecliff Bay.

Strobilops joossi (Gottschick). Strohilus costatus K. Mil-
ler, Jahresh. Ver. vaterl. Naturk. in Wiirttemberg, LVI, 1900,
p. 396, pi. 7, f . 8, Not of Clessin, — Strohilus joossi Gott-
schick, same serial, LXVII, 1911, pp. 502, 533, pi. 7, f. 16a-c.
— Strohilops joossi Wenz, N. Jahrb. Min., Geol., Pal., 1915,
II, p. 80, test-fig. 9; pi. 4, f. 14a-c. Upper Miocene, Sarma-
tian : Steinheim am Albuch, Wiirttemberg.

Strobilops labassetieri (de Morgan). Helix (Anchistoma,
Strohila) La Bassetieri J. de Morgan, Bull. Soc. Geol. France
(4), XIX, Nov. 1920, p. 313, fig. 8. Miocene: Faluns de Tou-
raine, Charenton pres de Pont-Levoy.

Strobilops labyrinthicula (Michaud). Helix lahyrinth-
icula Michaud, Actes Soc. Linn. Lyon II, 1854-5, p. 43, pi. 5,
f . 4, 5. — Strohilops lahyrinthicula Wenz, N. Jahrb. Min., Geol.,
Pal., 1915, II, p. 82, text-fig. 10, pi. 4, f. lla-c. Middle Plio-
cene, Plaicensian: Celleneuve and Montpellier (Dep. Herault),
Hauterive -(Dep. Drome), etc.

Strobilops menardi (Brong.). Helix menardi Brongniart,
Ann. du Mus. d'Hist. Nat. XV, 1810, p. 380, pi. 23, f. 11.—
Helix (Strohila) pseudolahyrinthica Cossmann, Cat. lUustr.
Coq. foss. FEoc. Env. Paris, IV, 1889, p. 360, pi. 12, f. 24-26.
Not of Saudberger. — Strohilops cossmanni Wenz, N. Jahrb.
Min., Geol., Pal. 1915, II, p. 74, pi. 4, f . 18a-c. — Strohilops
menardi Wenz, Foss. Cat., 1923, p. 1050. Middle Eocene,
Bartonian : Sables moyens de Bauchamp : Ducy, Montjavoult
etc. (Dep. Oise) and other localities in Dep. Seine-et-Oise,
Sarthe, Seine-et-Marne.

8 STROBILOPSID/E.

Strobilops monilia (Deshayes). Helix monilia Desh., De-
script. Anim. s. Vert. Bassin Paris, II, 1863, p. 816, pi. 54, f.
4-7. — Helix monile Sandberger, Land- u. Siisswasser-Coneh.
de Vorwelt 1873, p. 258, pi, 14, f. 15. — Helix nionilis Harris
and Burrows, Eocene and Oligoeene Beds Paris Basin, 1891,
p. 100. — Strobilops monilia Wenz, N. Jahrb. Min. Geol., Pal.,
1915, II, p. 73, pi. 4, f. 17a-c. — Strobilops monile Wenz, Foss.
Cat. 1923, p. 1051. Middle Eocene, Bartoniau: Auvers and
Beauchamp (Dep. Seine-et-Oise), etc.

Strobilops patuliformis (Sacco). Helix {Gonostoma)
patuliformis Sacco, Mem. R. Accad. Sc. Torino, XXXVII,
1886, p. 191, pi. 1, f . 19a-6. — Strobilops patuliformis "Wenz,
Foss, Cat. 1923, p. 1052. Upper Pliocene, Astian: Fossano
(Prov. Cuneo), Italy.

Strobilops pseudolabyrinthica (Sandberger). Helix pseu-
doUibyrintliica Sandberger, Die Land- u. Siisswasser-Conchyl.
der Vonvelt, 1873, p. 277, pi. 14, f . 25. — Helix labyrinthica
S. V. Wood, London Geol. Journ. I, 1847, p. 118.; Edwards,
Mouog. Eocene Moll., II, 1852, p. 67, pi. 10, f. la-e, and of
many other authors. Not of Say. — Strobilops pseudolaby-
rinthica Wenz, N. Jahrb. Min., Geol., Pal., 1915, p. 73, pi. 4,
f. 19a-c, 20c.— Cox, Proc. Malac. Soc. London, XVII, 1926, p.
51, 57, f. 7-9. Upper Oligoeene, Tongrian: Lower Headon
Beds, Totland Bay, Hordle; Headon Hill?; Melanienkalk at
Brunnstadt, Flaclanden etc., Alsace.

Strobilops romani Wenz. N. Jahrb. Min., Geol., Pal., 1915,
II, p. 83, text-fig. 12, pi. 4, f. 12a.-c. Middle Pliocene, Plai-
censian: Hauterive (Dep. Drome). Figured on pi. 12, figs.
11-13.

Strobilops sandbekgeri (Lomnicki). Strobilus lenticvlaris
Sandberger, Verhandl. k.-k. Geol. Reichsanst., Wien, 1885, p.
76, name only. — Strobilus sandbergeri Lomnicki, same Ver-
handl., 1886, p. 421. — Strobilops sandbergeri Wenz, Fossilium,
Catal., 1923, p. 1054. Upper Miocene, Tortonian : Folwarki,
Steniatyn and Wyczolki, eastern Galicia. "Perhaps identical
with S. costatus" (Wenz).

Strobilops subconoidelv (Jooss). Strobilus siibconoideus
Jooss, Nachrbl. d. Malak. Ges. XLIV, 1912, p. 34, pi. 2, f. 4.—

STROBILOPSID^. 9

Strohilops siibconoidea Wenz, N. Jalirb. Min., Geol,, Pal.,
1915, II, p. 81, pi. 4, f. 3a-c. Upper Miocene, Sarmatian:
Steinheim am Albiich, Wiirttemberg.

Stbobilops tiarula ( Sandberger ) . Strohilus tiarula Sandb.,
Verhandl. k.-k. Geol. Reiclisaiistalt, Wien, 1886, p. ddl.— Stro-
hilops tiarula Wenz, Fossilium Catal., 1923, p. 1056 ; Sencken-
bergiana III, 1921, p. 32, fig. 4. Lower Pliocene, Pontian:
Leolobersdorf, Kottingbruun, Austria.

Strobilops uniplicata (A. Braun). Helix uniplicata A.
Braun, in Walclmer, Handbuch der Geognosie, 2 Aufl., 1851,
p. 1138. — Strohilops imiplicata Wenz, N. Jahrb. Min., Geol.,
Pal., 1915, II, p. 76, fig. 4, pi. 4, f . 8a-c ; Fossilium Catal. 1923,
p. 1057. Upper Oligocene, Chattian: Hochheim-Florslieim,
Hesse-Nassau. Lfower IVIiocene of Germany and Tuchorshitz,
Czechoslovakia. Middle Miocene, Helvetian : Dettighofen near
Eglisau, Canton Ziirich.

Strobilops uniplicata plana (Clessin). Strohilus planus
Clessin, Malak. Blatter, n. F. VII, 1885, p. 80, pi. 7, f. 8.—
Strohilus diezi Clessin, Bericlit naturwiss. Ver. Regensburg,
XIII (1912), p. 103. — Strohilops uniplicata var. depressa
Wenz, N. Jahrb. Min., Geol., Pal., 1915, II, p. 77, pi. 4, f.
lOa-c. — Strohilops uniplicata plana Wenz, Fossilium Catal.,
1923, p. 1059. Upper Miocene, Tortonian: Undorf near
Regensburg, Steiermark; Hohenmemmingen, W^iirttemberg.

Strobilops uniplicata sesquiplicata (Boettger). Strohilus
unipUcatus var. sesquiplicata 0. Bttg., Bericht Sencken-
bergische Naturf. Ges., Frankfurt a. M., 1884, p. 259.— Stro-
hilus unipUcatus var. semiplicata K. Fischer, Bericht
Senckenb. naturf. Ges. 1904, p. 51. — Strohilops uniplicata
sesquiplicata Wenz, N. Jahrb. Min., Geol., Pal., 1915, II, p. 77,
pi. 4, f . 9a-c ; Fossilium Catal. 1923, p. 1060. Lower Miocene,
Aquitanian : Budenheim near Mainz, Wiesbaden. Upper Mio-
cene, Tortonian : Frankfurt a. M.

Genus PSEUDOSTROBILUS Oppenheim.

Helix sect. Pseudostrohilus Oppenh., Zeitsch. d. Deutschen
Geol. Gesellschaft, XLIV, 1892, p. 778.

10 STROBILOPSID^.

In the Upper Cretaceous of Austria and Hungary several
snails have been found which, though imperfectly known,
appear to have some characters of Strobilops. The name
Pseudostrohilus has been applied to them.

The shell is depressed, fragile, umbilicate, of 5 slowly in-
creasing, costulate w^horls. The lip is thickened, and on the
parietal wall there are three strong entering lamellae in the
best known species and type of Pseudostrohilus, P. riethmul-
leri, pi. 12, figs. 7, 8. Diam. 5 mm. Nothing is known of in-
ternal teeth or folds ^\dthin the basal lip.

PsEUDOSTROBiLus RiETHMULLERi (Tauscli). Helix 7'ieth-
miilleri Tausch, Abhandl. k.-k. Geol. Reichsanstalt, XII, 1886,
p. 14, pi. 2, f. 10, 11. — Helix {Pseudostrohilus) riethmiilleri
Oppenheim, Zeitseh. Deutschen Geol. Gesellsch., XLIV, 1892,
p. 778. Non-marine beds of the Upper Cretaceous of the
Csinger Valley near Ajka, Bakony, Hungary.

PsEUDOSTROBiLus AiGENENSis (Tausch). Helix oigeneiisis
Tausch, Abhandl. k.-k. Geol. Reiehsanst., XII, 1886, p. 26, pi.
2, f. Vla-h. Cretaceous: Gosau marl of Aigen near Salzburg,
Austria.

Genus OBBINULA Stache.

Ohhinula Stache, Abhandl. k.-k. Geol. Reichsanstalt, XIII,
1889, p. 119. Monotype 0. anthracophila (Stache).

Thin, imperforate, flattened-conic, of 5 narrow, slowly in-
creasing whorls, with fine, sharp growth-lines, with stronger
ones at regular intervals; the earlier whorls somewhat more
steeply conic. Aperturp with two or three entering ridges.
Type, 0 anthracophila, pi. 12, fig. 10. Diam. 4 mm.

Obbinula anthracophila (Stache). Helix anthracophila
Stache, in Sandberger, Land- u. Siisswasser-Conchyl. Vor-
welt, 1871, p. 129, pi. 19, f. 7 a-h. — Ohhinula anthracophila
Stache, Abhandl. k.-k. Geol. Reichsanst., XIII, 1889, p. 119,
pi. 1, f. 26. — cf. Oppenheim, Zeitsehr. Deutsch. Geol. Ges.,
XLIV, 1892, p. 779. "Cretaceo-Eocene" of Cosina, Dalmatia,
in a coal layer of the Stomatopsis beds.

As Oppenheim has remarked, the accounts of this little
fossil are conflicting; it seems to have apertural armature

STROBILOPSID^. 11

somewhat like PseudostrohUus, but it differs by being imper-
forate. Only a single, very much compressed example is
known.

This genus and the preceding may possibly be related to
Dimorphoptychia.

Mollusks erroneously referred to Strohilopsidcs, and

non-molluscan forms.

Dimorphoptychia Sandberger.

Dimorphoptychia Sandberger, Land- und Siisswasser-Con-
chyl. der VorAvelt, 1871, p. 149. Monotype Helix arnouldi
Michaud. E-illy, Eocene.

H. arnouldi, pi. 12, figs. 1, 2, 3, has been considered an
endodontid, a strobilopsid or a proserpinid snail by different
authors. Berthelin, in Bull. Soc. Geol. France (3), XV, 1887,
p. 61, found that there was complete removal of the internal
partitions, and on that account he referred the genus to the
Proserpinidag.

Because of its blunt, expanded peristome I am inclined to
\dew Dimorphoptychia as a member of the Helicinidae, near
the genera Calyhium L. Morlet and Heudeia Crosse.

KANiVBOHELix, new genus.

The shell is low trochiform, with rounded periphery and
broadly expanded lip, within which there are four narrow
folds (represented by linear grooves in the cast), subequally
spaced, the lower one shortest, more prominent; a parietal
tooth present ( f). Type Helix kanahensis, pi. 12, figs. 4, 5, 6.
Diam. about 12 mm.

K.VNABOHELix KANABENSis (White). Helix kanahensis
White, in Powell's Rep. on (^-eol. of the Eastern Portion of
the Uinta Mountains, 1876, p. 120; Review non-marine fossil
Mollusca of N. A., p. 48, pi. 25, figs. 12-14, in Ann. Rep.
Director U. S. Geol. S\irv. 1881-2 (1883). Cretaceous, Lara-
mie : Point of Rocks Group, Upper Kanab, Utah. Figured on
pi. 12, figs. 4, 5, 6.

In America no Tertiary or old fossils referable to Strohilops
have been found. Helix 'kanahensis White was referred to

12 STROBILOPS.

Strohila in this author's later paper (1883). The cast shows
impressions of a series of baso-palatal teeth or short folds, and
the general form is not inconsistent Avith the Strohilops group,
but the folds are close to the lip within, and the shell is so
much larger (diam. about 12 mm.), that the relationship
claimed appears highly problematic. At present I am inclined
to consider K. kanahensis an helieid snail.

StroUla leiodus Hutton, 1883. Trans. N. Z. Inst. XV, p.
135. Belongs to Ptychodon in Endodontidse.

Strohila octoradiata Sars, 1835. Beskrivelser etc., p. 16,
Belongs to the Acalephas.

Helix siihlabijrinthica Edwards, 1852 (Monogr. Eocene
Moll., p. 69, pi. 11, f. 4), referred to StroMlus by Sandberger,
1873, and to Strohilops by Wenz, 1915, 1923, has been shown
by L. R. Cox to belong to Acanthimda (Proc. Malac. Soc. Lon-
don, XVII, 1926, p. 53, fig. 10). Upper Oligocene. Cox sug-
gests that this species may prove identical with Acanthinula
paludincrformis Sdbg.

Helix lautricensis Noulet, 1867, Bull. Soc. Hist. Nat. Tou-
louse, I, p. 151, has been considered a sjnionym of the pre-
ceding. It occurs in various localities in Dep. Tarn, France.

Helix (Patula) recurrecta Oppenheim, Denkschriften K,
Akad. Wissensch., Wien, LXVII, 1890, p. 123, from the Vin-
centine Eocene, Altissimo, is possibly a Strohilops, but the
aperture is filled up so that teeth, if present, would not show.

Genus STROBILOPS Pilsbry.

Strohila Morse, Terrestrial Pulmonifera of Maine, in Joum.
Portland Soc. Nat. Hist., I, 1864, p. 26, for Helix lahyrinthica
Say. Not Strohila M. Sars, Beskrivelser og lagttagelser over
nogle maerkelige eller nye i Havet ved den Bergenske Kyst
lebende Dyr, 1835, p. 16. Not Strohila Sodoffsky, Bull. Soc.
Imp. Nat. Moscou, 1837, No. VI, pp. 92, 97, a name substituted
for Coccyx of Ochsenheimer and Treitschke, Sehmetterlinge
von Europa, VII, 1829.

Strohilus Sandberger, Land- und Siisswasser-Conchyl. der
Vorwelt, 1872-1875, pp. 258-726, and of many other authors.
Not Strohilus Anton, Verzeichniss der Conchylien, etc., 1839,
p. 46; see Man. Conch., XXIII, p. 188 (TomatellinidaB).

STROBILOPS. 13

StroUlops PiLSBRT, Proc. A, N. S. Phila. for 1892, p. 403
(Feb. 7, 1893), a substitute for Strohila Morse; Nautilus, VI,
1892, p. 56; XXII, 1908, p. 78. — Hanna, Nautilus, XXV,
1922, p. 91 (anatomy). — ^Wenz, Neues Jahrbucli fiir Miner-
alogie, Geologic unci Palaontolo^e, 1915, II, pp. 63-88 (synop-
sis of Tertiary species) ; Nachrichtsblatt d. Deutschen Malako-
zoologischen Gesellschaft, 1916, pp. 178-192 (synopsis of
Eecent species) .

The shell is small, perforate or umbilicate, trochiform to
subdiscoidal, with rounded, angular, or carinate periphery, of
4% to 6 closely-coiled whorls. Cavity of the last whorl is
obstructed by two or three long parietal lamellse, the upper
one emerging to the edge of the parietal callus, the lower one
weaker, emerging or immersed, the intermediate one when
present, smallest and remote from the aperture; a series of
two or more short folds on the basal wall of the cavity deep
within the last whorl. These lamellas and folds appear very
early in life, grow at the forward end and are absorbed be-
hind. Peristome expanded, usually thickened, the insertions
of the lip remote, connected by a parietal callus.

The sole is short and broad, in movement showing two or
three advancing muscular waves at one time ; no pedal grooves.
The integument has a network of impressed lines. Eyes arei
well-pigmented, on eye stalks a little swollen distally. Ten-
tacles short but moderately well-developed (pi. 4, figs. 9, 11,
S. lahyrinthica) .

Anatomy according to Hanna. — The kidney is long and nar-
row, passing anteriorly into the long urethra (ur.) which lies
close to the intestine (g) and opens near the breathing pore
(pi. 4, fig. 8). Genitalia (pi. 4, fig. 7, S. lahyrinthica) : Ovo-
testis (ot) composed of follicles in two groups, imbedded in
the liver. The lower portion of the spermoviduct {h. d.) is
thick and strongly convoluted. Albumen gland with finely
granulose surface. The prostate forms a ''series of lamellar
pouches." Vas deferens is long, passing below into a rather
thick and very long epiphallus {ep.). Penis {p) is rather
stout in the lower half, abruptly becoming thin in the upper
(or possibly this thin portion is part of the epiphallus; a ques-
tion to be settled by opening the organ) . It bears a very long

14 STROBILOPS.

appendix (ap) which is thickened at the base and in the distal
half. The penial retractor (pr) is bifurcate, one branch in-
serted at the summit of the penis (or on the epiphallus), the
other on the swollen basal part of the appendix; posteriorly
the retractor joins the right ocular muscle. The oviduct is
"thin-walled, slightly pouched." The spermatheca {sp) on a
duct of medium length without diverticulum.

The animal is oviparous.

The digestive tract is composed of the usual elements f
buccal mass, salivary glands, oesophagus, stomach and intes-
tine. Two features seem to be noteworthy. The oesophagus
is not a slender duct as usual, but the walls are "knotty" or
slightly convoluted throughout. Also on the stomach there
appears to be an accessory gland, closely appressed to the
walls of that organ. The salivary glands are united into one
but they seem to discharge into the buccal mass at the usual
two points.

Type, S. labyrinthica (Say).

Distribution : Humid eastern half of North America from
Quebec, Ontario and Manitoba, N. Lat. 52°, to Guatemala;
Cuba and Jamaica; South America from Venezuela, north-
eastern Brazil and the Galapagos Islands; Japan, Korea,
China and the Philippines. Eocene to Pliocene of central and
western Europe.

Strohilops comprises small snails living on decaying logs
and dead leaves in moderately humid forest. The genus is
not known in America west of the 100th meridian, but east-
ward its distribution is probably almost continuous, from
James Bay in the north to Guatemala, although at present
there are few Mexican records. It has not been turned up in
Costa Rica or Panama, the best known Central American
faunas, but appears again in Venezuela, Para and the Gala-
pagos. These three remote localities in South America may
indicate a wide distribution in that continent, and perhaps
many species to be discovered.

In eastern Asia species have been found in Uzen Province
of northern Nippon, in Korea and China, The peculiar sub-
genus Entcroplax is confined to the Philippines.

STROBILOPS.

15

In Europe about twenty species are known running from
the Eocene to the Upper Pliocene.

The remarkable discontinuity in the distribution of Stro-
bilops, and even of its subgenera, will be seen by the accom-
panying map upon which the occurrence of the genus is
roughly indicated.

Subgenera

1. atrobilops s.str

2. Eostrobilops

3. Discostrobilops

4. Enteroplax

Fig. 2 World in North Polar projection. Tertiary distribution of

Strobilops represented by dots; Recent by black spots, and Discostrobilops
by horizontal lines.

These small snails attracted attention more than a century
ago, the first (Eocene and Oligocene) species having been de-
scribed by Brongniart in 1810. Say discovered the first living
species in 1817. The genus appears to have been a favorite
one with paleontologists; notices of the species, especially in

16 STROBILOPS.

Oligocene and Miocene beds, are very numerous. All of the
early authors referred the species to Helix. E. S. Morse, 1864,
was the first to demonstrate the intricate internal structure of
the shell, and to recognize that it belongs to a special genus,
whicli he named Strohila — a name which unfortunately proved
to be a homonym. Sterki (Nautilus, VI, 1892, pp. 3, 6) re-
moved the genus from the Helicidse to the family Pupidse
(Pupillidae). Hanua (1922) described and figured the anat-
omy, and proposed the name Strobilopsidae. Several articles
by the present writer (1893-1909) enlarged our knowledge of
American forms, pointed out the occurrence of the genus in
eastern Asia, and indicated the relationship of the Philippine
forms of the subgenus Enteroplax to Strohilops. Finally, W.
Wenz has reviewed the species in two excellent articles:
"Die fossilen Arten der Gattung Strobilops Pilsbry und ihre
Beziehungen zu den lebenden," 1915, and "Zur Kenntnis der
Gattung Strobilops Pils.," 1916.

Of the twenty-eight Recent forms now known — nineteen
species and nine subspecies — I have not seen specimens of the
following four : Strohilops veracruzcnsis crossei, 8. salvini, 8.
diodontina and 8. trochospira. The type specimens of 13
species and 6 subspecies have been examined; the remaining
three species are known to me by specimens from the original
collectors.

Classification.

The first attempt to classify the species of Strohilops into
natural groups was made by W. Wenz (1915, pp. 84-86).
Considering the fossil species only, he proposed three groups.

1. Group of 8. diptyx, for the species diptyx, fischeri, hoett-
geri, suhconoidea, duvali. This line begins in mid-Oligocene
time and runs to mid-Pliocene, in central and western Europe.
Perhaps 8. elasmodonta belongs here: it differs from all the
others by possessing an interparietal lamella.

2. Group of 8. costata, with the species costata, joossi,
tiarula, romani, lahyrinthicula. Probably 8. monilia and 8.
ynenardi, and evidently 8. headonensis and 8. pseudolahy-
rinthica are to be added. This series ran from Eocene prob-

STROBILOPS. 17

ably, and certainly from Lower Oligocene to mid-Pliocene
times.

3. Group of 8. uniplicata, for S. unipUcata and its sub-
species. Upper Oligocene to Upper Miocene.

A few of the Tertiary species are not known fully enough
to decide upon the group. Nearly all of the living species
appear to be referable to one or other of Wenz's three groups,
though the definitions given by him require some modification
to provide for them, as would be expected.

The number and relative size of the basal folds can hardly
be considered a subgeneric character, being rather variable
among closely allied species. Probably forms with a columel-
lar lamella and three basal folds, with none above the peri-
phery, represent a rather primitive stage, this number being
found in various Tertiary species as well as in the peripheral
South American forms. At the same time, some early forms,
such as 8. headojiensis and siihlahyrinthica possessed more
numerous basal folds and also some above the periphery (pala-
tals), just as in the Recent 8. lahyrinthica. It appears that
there has been some parallel evolution in these folds, and an
increased number may appear in different phyletic lines, if
the multilaminate forms are actually the later products of
evolution.

The interparietal lamella may be either present or wanting
in the group of 8. hub'hardi, but it is certainly absent in most
of the fijiely striate species (group 1 of 'Wenz=: Eostrol}ilops),
and it is present in the costate ones (group 2 = typical 8tro-
hilops). Probably this lamella was present in the ancestral
stock, but degenerated in some groups. Thus, it is developed
in 8. unipUcata, but very weak or usually absent in the modem
8. huhhardi. The degree of minute sculpture of the edges of
the parietal lamellaa varies rather widely ; Wenz did not find
prickly knots on the lamella of his group 2, but they are
present, though inconspicuous, in the Pliocene ;S^. romani (pi.
12, figs. 11, 12, 13), and are more or less conspicuously de-
veloped in all living species of the same group.

The following arrangement is here adopted :

18 STROBILOPS, CANADA AND UNITED STATES.

C ^ r. , M ( Sect. Strobilops s. s.

StrobOo s i Strobilops.... | g^^^_ Eostrobilops.

^ 1 S.-g. Discostrobilops.
I S.-g. Enteroplax.

Key to Subdivisions of Strobilops.

1. Parietal callus having a thickened, raised edge, prominent

where the parietal lamella joins it ; parietal lamellae with
smooth or microscopically serrate edges, not nodiferous ;
shell thin, carinate. Philippines.

Subgenus Enteropl^vx, species Nos. 16-19.
Parietal callus not noticeably thickened or raised at the
edge. (2)

2. Form conoidal; umbilicus narrow; parietal lamellae gener-

ally with prickly or rugose nodes. (3)
Form subdiscoidal ; umbilicus wide; parietal lamellae with
sparsely prickly or smoothish edges, not nodiferous.

Subgenus Discostrobilops (type ^S'. huhbardi),
species No. 15.

3. Upper sui'face finely striate ; no interparietal lamella.

Section Eostrobilops (type 8. hirasei), species
Nos. 11-14.
Upper surface costulate (except in S. morsel, which is
nearly smooth) ; an interparietal lamella present.

Section Strobilops proper, species Nos. 1-10.
The species are grouped geographically as follows :

Canada and the United States, species 1 to 4&, 15.

Mexico and Central America, species 4c to 7, 15.

"West Indies, species 15.

South America, species 8 to 10.

Japan, Korea and China, species 11 to 14.

Philippine Islands, species 16 to 19.

Section Strobilops proper.
The shell is troehiform or dome-shaped with the upper sur-
face or the ivhole shell ribbed (except in S. morsei). One or
two parietal lamellae emerge; deep within, their edges have
rugose or prickly nodes (in the Recent species, at least) ; an

STROBILOPS, CANADA AND UNITED STATES. 19

interparietal lamella is developed deep within. Internal bar-
rier consisting of a columellar lamella, several basal folds and
often other palatal folds above the periphery. Type, S. lahy-
rintJiica.

All living American species except 8. hubbardi belong to
this group. Also the European Tertiary Gruppe der Stro-
bilops costata of Wenz (1915, p. 85). In the latter the knots
of the lamellge appear to be very weak or wanting; "von
einer Zahnelung der Lamellen habe ich nichts bemerkt, "
Wenz observes ; but in the Pliocene ^S^. romani (PI. 12, figs. 11,
12, 13) roughened nodes are present though inconspicuous.

There are two groups of species : first, that of S. Idby-
rinthica, iu which there are numerous basal folds and one or
two above the periphery; this group including also S. texas-
iana and 8. affinis. Second, the group of 8. strebeli, in which
there are three (or in 8. cenea, four) basal folds and none
above the periphery ; the Mexican and South American species
belong here. Both of these groups appear to be represented
in the Tertiary fauna of Europe.

8pecies of Canada and the United 8tates.

1. Trochiform or dome-shaped species, in which the height

much exceeds half the diameter; umbilicus narrow or
moderate, contained 6 to 12 times in the diameter. (2)
Subdiscoidal species, the height about half of the diameter ;
umbilicus contained between 3 and 4 times in diameter.

^S'. hubbardi, No. 15.

2. Spire convexly conic or dome-shaped; elevated; usually 5

or more basal and palatal folds. (3)
Spire rather low conic, its outlines but slightly convex ; less
elevated; periphery angular; 3 or 4 basal folds, none
above the periphery. 8. cenea, No. 4.

3. Diam. 2.3-2.5 mm.; basal folds strongly unequal. (4)
Diam. 2.75-2.9 mm.; basal folds short, subequal, disposed

in a regular curve. 8. affinis, No. 3.

4. Finely ribbed. 8. laiyrinthica, No. 1.
Coarsely ribbed ; southern. 8. texasiatia, No. 2.

20 STBOBILOPS, CANiVDA ANT) UNITED STATES.

1. Strobilops labyrinthica (Say). PL 1, figs. 1-11.

The shell is narrowly umbilieate, the -width of umbilicus
contained about 11 (9 to 12) times in the diameter of the
shell; very convexly conic or dome-shaped, the periphery ob-
tusely subangular. Whorls 5i/4, convex, very slowly widen-
ing, the first iy2 smooth, pale, the rest chestnut-brown, sculp-
tured with narrow obliquely radial ribs narrower than their
intervals, passing over the periphery but weakening at the
base, the first half of which is typically nearly smooth. The
aperture is semi-lunar. Peristome brown, expanded, thick.
The parietal lamella emerges to the edge of the parietal callus
and penetrates inward a little more than half a whorl. The
infraparietal lamella is much smaller, only shortly emerging,
the end visible in a basal view; inside it penetrates as far as
the parietal lamella. There is a low and slender interparietal
lamella between these lamellsB deep within; all three are
strongly nodose at the edge, the nodes armed with minute
prickles directed towards the aperture (PL 1, fig. 11). "Within
the basal and outer waUs, at the last third of the base, there
is a low, rather blunt columellar lamella and a forwardly
curving series of five (or six) unequal basopalatal folds; first
and second folds are large and high, the second longer; two
or three following folds are low and thin, the one immediately
above the periphery usually longer, and there is sometimes
another fold above it (PL 1, fig. 6, a topotype; also figs. 7, 9).

Height 1.7, diam. 2.3 mm. (Philadelphia).

Height 1.8, diam. 2.3 mm. (Philadelphia).

Maine and Quebec west to Manitoba, Minnesota, Kansas and
Arkansas, south to Georgia and Alabama. Type locality,
Philadelphia, Pa.

Helix labyrinthica Say, Journ. Acad. N. S. Phila., I, 1817,
p. 124. — Strohila labyrinthica Say, Morse, Terrestrial Pul-
monifera Maine, Journ. Portland Soc. N. H., I, p. 26, figs. 61-
67, pi. 8, f. 68.— BiNNEY, Terr. MolL, V, 1878, p. 259, text-
figs. 149-152, pi. 17, f. 3; pi. 5, f. o (teeth).— Strebel, Beitrag
Mexikanischer Land- und Siisswasser-Conch., IV, 1880, p. 43,
pi. 4, f. 6a (Ohio).— F. J. Ford, Naut. Ill, p. 106 (Sedgmck
Co., Ks.). — Sampson, Naut. VII-, p. 33 (Missouri). — Prime,

STROBILOPS, CANADA AND UNITED STATES. 21

Naut. VIII, p. 70 (Long Island, N. Y.).— Price, Naut. XIV,
p. 76 (Kentucky).

StroMlops labyrinthica Pilsbrt, Proc. A. N. S. Phila. 1892,
p. 404; Naut. VII, p. 57.— Pleas, Naut. VII, p. 69 (Henry Co.,
Ind.).— Walker, Naut. VII, p. 126; XI, p. 122 (Michigan).—
Nylander, Naut. VIII, p. 126; XIV, p. 103 (Aroostook Co.,
Me.). — Schick, Naut. VIII, p. 137 (Philadelphia). — Smith,
Naut. XIII, p. 35 (Michigan).— Ferriss, Naut. XIV, p. 103
(Cove, Ark.).— CoLTON, Naut. XVII, p. 100 (Mt. Desert).—
Van Hyning, Naut. XVII, p. 131 (Des Moines, Iowa). —
K-HO.VDS, Naut. XVIII, p. 65 (Delaware). — Blaney, Naut.
XVIII, p. 46 (Sowards I., Me.).— D.vniels, Naut. XIX, p. 63
(Pleistocene, New Harmony, Ind.). — Hinkley, Naut. XX, p.
35 (Ala., Miss.); XXXIII, p. 14 (Dubois, 111.). — Wheat,
Naut. XX, p. 101 (Cayuga L.). — Jackson, Naut. XXI, p. 144
(North Haven, Me.). — F. C. Baker, Naut. XXIV, p. 69
(Unionville, Conn.) ; XXV, p. 67 (St. Lawrence R. islands) ;

XXVIII, p. 10 (Michigan) ; Mollusca Chicago Area, Chicago
Acad. Sci. Bull. Ill, 1902, p. 225, f. 58-60 (jaw and teeth) ;
Life of the Pleistocene, Univ. Illinois BuU. XVII, 1920, p. 388
(Aftouian Pleistocene to Recent). — Wheeler, Naut. XXV, p.
124 (Ala.) ; XXXI, p. 114 (Clark Co., Ark.).— Sterki, Naut.

XXIX, p. 123 (Geneva, 0.).— Vanatta, Naut. XXXIII, p. 97
(Maine). — Walker, 111. Cat. MoU. Michigan, 1906, p. 505,
figs. 116-118. — Dall, Alaska: Land and Fresh Water Mol-
lusks, Harriman Alaska Exped. XIII, 1905, p. 27, f. 7-9 (Car-
berry, Manitoba; Moose Factory; James Bay). — Wenz,
Nachrbl. D. M. Ges. 1916, p. 179.— Zetek, Trans. Illinois State
Acad. Sci. XI, 1918, p. 170 (near Urbana, in woods).

StroMlops affinis Vanatta, Nautilus XXXIII, p. 97 (Se-
bago, Cumberland Co., Maine).

Strohilops floridanus Vanatta, Naut. XXX, p. 72 ; XXXIII,
p. 68 (New Jersey).

Strobila labyrinthica var. virgo Pilsbry, Nautilus VI, p. 94,
Dec. 1892 (Sebec Lake, Piscataquis Co., Me.). — S. virgo Sar-
gent, Nautilus IX, p. 89 (Clearwater Lake, Wright Co.,
Minn.).— G. H. Clapp, Naut. XIV, p. 64 (Mt. Agamenticus,
Me.).— Berry, Naut. XXIV, p. 63 (Windemere, Waldo Co.,
Me.).— ? Wheeler, Naut. XXXI, p. 114 (Arkadelphia, Ark.).
—Walker, 111. Cat. Moll. Michigan, 1906, p. 506, fig. 120
(abundant in northern Michigan; southward in Oakland,
.Saginaw and Kalamazoo counties) . — Sterki, Proc. Ohio State
Acad. Sci. IV, pt. 8, 1907, p. 378 (New Philadelphia, 0. ;
seems to prefer elevated and dry places, whereas ;S'. labyrin-
thica prefers damp places).- — Daniels, 27th Ann. Rep. Dept.

22 STROBILOPS, CANADA AND UNITED STATES.

Geol. and Nat. Res. Indiana, 1902, p. 585 (near Cannelton,
Perry Co. ; New Harmonj^, Posey Co. ; Princeton, Gibson Co. ;
Wyandotte, Crawford Co., and Lake Maxinkuckee, Marshall
Co.; beneath logs and rubbish in ravines and moist places).

Many of the older references to Helix or Strohila laby-
rinthica have been omitted, as it is impossible to tell whether
they refer to this species, ;S^. affinis or 8. cenea.

The back, eye-stalks and tentacles are blackish gray, darker
streaks running from the collar to the eye-stalks ; sides of the
foot and the tail are clear whitish gray (PI. 4, figs. 9, 11).

S. lahyrinthica differs from S. affinis by the longer, more
conspicuously unequal basal folds, the first (inner) two much
larger than the others, and the series does not form an even
curve as in affitiis; the infraparietal lamella generally emerges
more; the shell is smaller and generally less elevated, the
spire with more strongly convex outlines. In ;S^. miea the
whole shell, and especially the last whorl, is lower, the out-
lines of the. spire are less convex ; the basal folds are less
numerous, and there are none above the periphery; the color
is brighter and more transparent than in S. lahyrinthica.

No locality was mentioned by Say, but his cotypes, four
specimens, are labelled "Penna. ; Hyde and Mason," and
probably came from the immediate vicinity of Philadelphia,
which place I have selected as type locality. One of these
shells is drawn in PI. 1, figs. 3, 4.

Its usual stations are "under loose bark of logs, in half-
decayed wood, among dead leaves and in sod at bases of
trees. ' '

The umbilicus may be narrow, enlarging but little in the
last whorl, or it may enlarge to double the fonner -uddth in
the last half-whorl. The infraparietal lamella emerges more
or less conspicuously. The degree of angulation at the peri-
phery varies somewhat, and the sculpture of the base is also
variable, often nearly smooth in front of the aperture as in
the type, but sometimes distinctly ribbed throughout (the ribs
are represented somewhat too strong in PI. 1, fig. 8). The
parietal lameUa in rare instances penetrates as much as three-

STROBILOPS, CANADA AND UNITED STATES. 23

fourths of a whorl (var. parietalis). The number of baso-
palatal folds outside of the second one (the third from, the
columellar), varies from three to four in adult shells, and
there may be either one or two of them above the periphery.

The color of the shell varies from dark to light brown, and
albino shells occur (Neosho Co., Kansas) which are similar to
S. I. virgo in color, but have a wider umbilicus and a less
emerging infraparietal lamella than typical northern virgo.

In a young shell of 1.7 mm. diameter the internal barrier
consists of a rudimentary columellar lamella, four long basal
folds and one above the periphery. In some individuals the
number of basal folds is increased in the late neanic stage.
In one young Kansas specimen, nearly ready to form the lip,
there are ten, including the columellar lamella (PI. 1, fig. 10) ;
but so large a number is very exceptional, and I have never
seen a fully adult shell with more than a columellar, four
basal and a palatal fold — six in all. The supernumerary folds
are evidently absorbed at the inception of the adult stage.

Morse's figure of the spines on the parietal lamellge of 8.
lahyrinthica represents them as longer and stronger than in
any of the many shells I have opened. I conclude that his
figures of the internal structure are rather diagrammatic.

la. S. lahyrinthica form virgo Pils. PI. 4, figs, 3, 4.

Shell whitish with a faint green or yellow tint (or pale
brown), the lip and lamella white. Umbilicus small, con-
tained 10 times in the diameter. The infraparietal lamella
emerges somewhat more strongly than usual in lahyrinthica;
otherwise the lamellse and folds are the same. Height 2, diam.
2,5 mm.

Maine to Minnesota and Iowa ; reported also from Arkansas ;
described from Sebec Lake, Piscataquis Co,, Maine,

The albino shells are found associated with pale brown
specimens in the type locality, all agreeing in lamellas and
other characters except color. As a color-form it is recog-
nisable, but I believe of very little significance racially. It
appears to be chiefly a Canadian Zone form.

24 STROBILOPS, CANADA AND UNITED STATES.

1&. Strohilops lahijrinthica form parietalis, new form. PL 9,
figs. 10, 11.

Similar to iS^, Idbyrinthica in the eonvexly conic, ribbed shell
and weakly emerging infraparietal lamella, but the lamellee
penetrate more deeply, being between tivo-thirds and three-
fourths of a whorl long (fig. 10), There is a blunt columellar
lamella and four basal folds, four within the side wall (fig.
11, oblique view of baso-palatal folds). The whole base is
ribbed in some examples, or smoothish, merely finely striate
in others. Height 1.75, diam. 2.35 mm.

The type locality of this form is Ardsley, Montgomery Co.,
Pennsylvania (Bayard Long) ; but very similar examples are
before me from Gray Bluff, Marion Co., Tennessee; Lookout
Mountain, near Valley Head, and Blount Spring, Alabama
(H. H. Smith) ; Talahassee, Florida (C. W. Johnson) ; and
Catahoula Parish, Louisiana, 1.55 x 2.15 mm. (C. B. Moore).

Most of the southern specimens are rather small, but in a
lot from "Woodville, Ala. (H. E. Sargent), the largest meas-
ures 2 x 2.7 mm. While it approaches S. texasiana and
fioridana by the long parietal lamella, it differs by the de-
cidedly less coarse ribbing. The status of this form is doubt-
ful; the sporadic distribution, as now known, seems to indi-
cate that it is not a subspecies. It is introduced here to induce
further investigation,

2, Strobilops texaslvna Pils, & Ferr. PI, 2, figs. 5 to 11,

Shell moderately elevated with dome-shaped spire; light
bro^vn ; umbilicus contained about 8 times in the diameter.
Whorls 514, the first l^/^ smooth, pale-corneous, the rest regu-
larly ribbed obliquely, the last whorl rounded peripherally or
a trifle and obtusely subangular in front, the riblets passing
over undiminished upon the base, which is as strongly sculp-
tured as the upper surface (or sometimes smoothish just in
front of the aperture) , Aperture with slightly expanded, well
thickened, whitish peristome and a strong parietal callus.
Parietal lamella emerging to the edge of the callus, pene-
trating fully three-fourths of a whorl, Infraparietal lamella
less emerging, being very shortly and weakly visible in a basal

STROBILOPS, CANADA AND UNITED STATES. 25

view, penetrating about as deeply as the parietal. There is a
quite short and very low interparietal lamella, situated near
the inner ends of the others. A third of a whorl within there
is a very short, low lamella on the axis and five baso-palatal
folds: near the axis two large basal folds, the inner tongue-
shaped, the outer one longer ; outside of these there is a minute
and low third fold, often wanting, leaving a space ; the fourth
and fifth folds long and low (fig. 8). Just above the peri-
phery on the outer wall a very weak, low, long sixth fold may
often be traced, but in fully adult shells the smaller folds are
usually reduced in number (fig. 10), or even wanting (fig. 9) ;
the full number described above being rare except in the late
neanic stage.

Height 2, diam. 2.4 mm. Type.

Height 2, diam. 2.3 mm. Alexandria, La.

Texas : drift of the Guadalupe river about four miles above
New Braunfels (type loc. ; Pilsbry and Ferriss, 1903) ; Austin
(Pilsbry) ; San Marcos (Pilsbry and Ferriss) ; New Braunfels
(Ferriss, Pilsbry, and Singley) ; Guadalupe river bottom, Vic-
toria county, and Lavaca river, Jackson county (J. D. Mit-
chell) ; Lee county (Singley) ; Calhoun county (E. "W. Hub-
bard) ; Gainesville (J. B. Quintard) ; Colorado river near
Travis, Bastrop Co. (Julia Gardner). A smaller form, diam.
2 mm., was taken in drift debris of the Hondo river about two
miles north of Hondo, Medina county (Ferriss and Pilsbry).

Oklahoma: Limestone Gap and Wyandotte (Ferriss and
Pilsbry) ; Fort Gibson (B. W. Hubbard) ; Pottawatomie Co.
(J. B. Quintard).

Arkansas: Ashley and Calhoun counties (C. B, Moore).

Louisiana : Sycamore and Wilson landings, Morehouse Par-
ish; Alexandria, Rapides Parish (C. B. Moore).

Strohilops lahyrmthica texasiana Pilsbry and Ferriss,
Proc. A. N. S. Phila. 1906, pp. 147, 557.— Wheeler, Nautilus
XXXI, p. 114 (Arkadelphia, Ark.). — Strecker, Naut. XXII,
p. 65 (McLennan Co., Tex.).

This form differs from the closely related 8. lahyrinthica by
the stronger, more spaced ribs, continued over the base, and
by the more deeply penetrating parietal lamella.

26 STROBILOPS, C.ySTADA AND UNITED STATES,

2a. Stroiilops texasiana floridana (Pils.). PL 2, figs. 1-4.

The shell is conic with convex outlines, almost dome-shaped ;
the periphery only weakly angular, the base convex; rather
solid; brown, the summit whitish-corneous. Umbilicus con-
tained about 8 times in the diameter. Whorls 5i/^, the first
two smooth, the rest sculptured with narrow, rather widely
separated ribs (about 30 on the last whorl). These ribs con-
tinue on the base, which is radially ribbed. Aperture semi-
lunar, the peristome thick, narrowly reflected, brown or
whitish; parietal callus rather thick at the edge. Parietal
lamella emerging to the edge of the callus, penetrating inward
fully a whorl (PI. 2, fig. 3). Infraparietal lamella scarcely
emerging, penetrating as far inward as the parietal lamella.
The inner half of this lamella and of the parietal is nodose,
the nodes minutely asperate. Interparietal lamella very low,
about a half-whorl long, nodose, penetrating as deeply as the
parietal lamella. There is one small, short axial lamella and
four basal folds, the outer one peripheral in position ; a single
palatal fold is generally developed. These folds form a
curved, very obliquely radial series, the middle of which is
about a half-whorl back of the aperture. The two inner basal
folds are much stouter and higher than the others, the second
from the axis (or third, counting the axial) being the longest
and highest of the folds, and somewhat sinuous (PL 2, fig, 4) .

Height 2, diam. 2.4 mm, (Miami),

Height 1.9, diam. 2.2 mm. (Orlando).

Florida: Miami, Dade Co., type locality (S. N. Rhoads, M.
Hebard, Pilsbry). Throughout the State, from Sugarloaf,
Boca Chica and No-name keys to Clay Co. on the St. Johus
River and Jackson Co. on the Chipola.

StrohUops floridanus Pilsbry, Nautilus XXIII, Dec. 1909,
p. 90. — Vanatta, Naut. XXVI, pp. 21, 33 (No-name Key,
Sugarloaf Key) .— Wenz, Naehrbl. D. M. Ges. 1916, p. 184.—
Walker, Naut. XXXI, p. 56 (Palm Beach). — ;S^. labyrinthica
Rhoads, Naut. XIII, p. 45. — Johnson, Naut. XIII, p. 142,

This form differs from ;S^. labyrinthica by its widely-spaced
ribs and long parietal lamella. This is typically a full whorl
long, or somewhat more, thus differing from S. texasiana in

STROBILOPS, CANADA AND UNITED STATES. 27

which the lamella is a little short of one whorl long, as will be
seen on comparing pi. 2, figs. 3 (floridana) and 5 (texasiana).
However, on examining long series I fiiid some examples of
floridana in which the parietal lamella is not over three-
fourths of a whorl long, so that the distinction between
floridana and texasiana is not a complete one, though obvious
in most examples. There are also shells which approach the
long-lamellate parietalis form of la'hyrinthica rather closely,
though the coarser ribbing of the base in floridana is generally
a differential character.

It is sometimes quite elevated, as in the second measure-
ment above, but the high examples occur in lots "with lower
specimens.

3. Strobilops affinis Pilsbry. PI. 3, figs. 1-5.

The shell is convexly conic with obtusely angular peri-
phery; the base moderately convex, becoming rather strongly
so in its last third ; glossy, brown, with pale apex ; narrowly
umbilicate, the umbilicus contained about 1^2 to 8 times in
the diameter. There are 6 moderately convex wiiorls, in-
creasing very slowly, the first two smooth, the rest sculptured
with narrow, somewhat retractive ribs. The first half of the
base is smooth, the ribs barely passing over the peripheral
angle and reappearing within the umbilicus, but they continue
weakly over the last half. The aperture is semilunar, oblique.
Peristome is well expanded, thickened within, its face convex
and fleshy-bro^\^l in color. Parietal callus moderately strong.
The parietal lamella emerges to the edge of the callus and
penetrates inward about two-thirds of a whorl. The infra-
parietal lamella is low and weak, deeply immersed, not visible
in a front or basal view. Interparietal lamella short and very-
weak. A third of a whorl within there is an obliquely radial
series of about 8 folds : a short low lamella on the columellar
axis, follow^ed by two folds larger and higher than the rest,
and an oblique series running to the suture, composed of 4 to
7 short subequal folds (fig. 4).

Height 2.5, diam. 2.75 mm.

Massachusetts to Minnesota and Kansas, south to northern

28 STROBILOPS, CANADA AND UNITED STATES.

New Jersey and west of the Alleghanies to northern Alabama
and Oklahoma. Localities of specimens seen follow. Massa-
chusetts: New Bedford (J. H. Thomson). New York: Syra-
cuse, Pleistocene (Burnett Smith) ; Poughkeepsie (G. Van
Ingen) ; Upper Red Hook, Duchess Co. (W. S. Teator, type
loc.) ; Staten Island (E. W. Hubbard, T. Bland) . New Jersey :
Wliite Pond, Warren Co. (Pilsbry). Ohio: Kent (G. W.
Dean). Michigan: Pine Lake, Oakland Co. (B. Walker),
Minnesota: Dallas Lake, Wright Co. (H, E. Sargent). Illi-
nois: Athens (E. Hall).

Strobilops affinis Pilsbry, Nautilus VII, Sept. 1893, p. 57.
—Walker, Illustr. Cat. Moll. Michigan, 1906, p. 507, f. 121
southern Michigan, not reported north of Saginaw-Grand val-
ley).— Daniels, Nautilus XIX, p. 63 (Posey Co., Ind.). —
Ferriss, Naut, XX, p. 17 (Oklahoma City). — Nylander,
Naut. XXII, p. 143 (Aroostook Co., Me.). — H.wna, Naut.
XXIII, p. 95 (Douglas Co., Kan.).— Sterki, Naut. XXV, p.
116 (Glenrose, Texas) .—Hinkley, Naut. XXXIII, p. 14 (Du-
bois, 111.). — Sterki, Prelim. Cat. Land- and Fresh-water Moll,
of Ohio, Proc. Ohio State Acad. Sci. IV, 1907, p. 378 (Sum-
mit Co.). — Zetek, Trans. Illinois State Acad. Sci. XI, 1918,
p. 170 (Urbana, 111., under old logs). — Chadwick, Notes ou
Wisconsin Moll., Bull. Wisconsin Nat. Hist. Soc, IV, 1906, p.
75 (Lake bluffs south of Whitefish Bay; Menomonee valley;
Cudahy; Mishicot, Manitowoc Co., under conifers). — Daniels,
27tli Ann. Rep. Dept. Geol. and Nat. Resources of Indiana,
1902, p. 586 (near Lake James, Steuben Co.; Tippecanoe
Lake, Kosciusko Co, ; Mitchell, La^\Tenceburg, Wyandotte and
Huntingburg in southern Indiana). — G. H. Clapp, in litt, :
Thayer, Kansas (Ferriss) ; Evanston, 111, (Baker) ; Culver,
Ind, (Daniels) ; Cowan, Tenn, and Stevenson, Ala, (H, H,
Smith), — F, C. Baker, Life of the Pleistocene, Univ. Illinois,
Bull. XVII, 1920, p. 388 (Sangemon Pleistocene to Recent).

This shell is somewhat larger than 8. lahyrinthica, thinner,
with the infraparietal lamella more deeply immersed and the
baso-palatal folds less unequal, all being rather short and
fonning a regular curve across the base and up the outer wall.
The outlines of the spire are some^^hat less convex than in
8. IdbyrintJiica but more convex than in 8. miea.

STROBILOPS, CANADA AND UNITED STATES. 29

4. Strobilops ^nea Pilsbry. PL 3, figs. 6-12.

The shell is narrowly lunbilicate, the width of umbilicus
contained about 6I/2 times in the diameter of the shell, low-
conic, with obtuse, rounded summit, the periphery distinctly
tut hluntly angular. The base is somewhat fattened helow
the periphery, elsewhere moderately convex. Whorls 5%,
convex, slowly increasing, the first 1% smooth, corneous, the
rest dark brown with a red-gold gleam ; sculptured with nar-
row riblets which are somewhat oblique, retractive, rather fine
and close. The base is smoothish, marked with growth-striae
only, except on its last third, where the riblets of the upper
surface continue over the base. The aperture is semilunar,
low but wide. Outer and basal lips brown, well expanded,
somewhat thickened, the columellar margin dilated. The
parietal lamella emerges to the edge of the parietal callus,
penetrating inward a half whorl, Infraparietal lamella weakly
emerging. Midway between the lamella there is a very weak,
low, deeply-placed interparietal lamella. These lamellae are
nodose far within, the nodes roughened, shortly prickly (PL
3, fig. 10; pi. 4, fig. 2). The internal barrier, situated one-
third of a whorl from the aperture, is radial, but slightly
oblique (fig. 12) ; it consists of a short, weak columellar fold
and four basal folds, visible through the shell ; the second and
fourth folds from the axis are long, the first short, the third
fold weak or sometimes wanting; there is no fold above the
periphery.

Height 2, diam. 2.7 mm. Paratype.

Height 1.9, diam. 2.75 mm. Type, fig. 6.

Height 1.5, diam. 2.4 mm. Randolph Co., Ala.

Massachusetts to Michigan, Illinois and Arkansas, south to
southern Florida and Louisiana. Type loc, Cazenovia, N. Y.
Localities of specimens examined follow: Massachusetts: Mt.
Holyoke (A. D. Bro\\ai). New York: Cazenovia (Henderson,
Walker, Clapp and Pilsbry, 1905) ; Lanesville and Llarble-
town (Van Ingen). New Jersey: Burlington Co. Bayard
Long). Pennsylvania: Philadelphia; Chester Co. (B. Long) ;
Berks Co. (Pilsbry) ; York Furnace (W. Stone). Delaware:
Delaware City and Dover (S. N. Rhoads). Maryland: Great

30 STROBILOPS, CANADA AND UNITED STATES.

Falls (also on the Virginia side) (Pilsbry). Ohio: Kent (GT.
W. Dean) ; Garrettville (R. Walton). Illinois: Edwards Co.
(C. S. Hodgson); Sheridan (Ferriss). Tennessee: Cade's
Cove, Blount Co., 2000 ft. (Ferriss) ; Cumberland Plateau,
South Pittsburgh and Dove (H. H. Smith). North Carolina
Tuskeegee Mts., Graham Co. (Ferriss). South Carolina
Manning (W. Stone); Columbia (J. B. Clark). Florida
Manliattan, Lake Co. (Pilsbry and Johnson) ; Imri, Hamilton
Co. (E. B. Chope) ; Fort Lauderdale (C. T. Simpson) ;
Gainesville (J. B. Clark). Alabama: many places throughout
the northern half of the state (H. H. Smith, H. E. Sargent).

Strobilops lahyrinthica streheli Pfr., Pilsbry, Nautilus VII,.
Sept. 1893, p. 57 ; Proc. A. N. S. Phila. 1900, p. 133 ; 1902, p.
429 (North Carolina). — Ferriss, Naut. XIV, p. 56 (Cade's
Cove, Tenn.). Not Helix streheli Pfeiffer. — Strobilops lahy-
rinthica (Say) Walker, 111. Cat. Moll. Mich., 1906, p. 506,_f.
119 (not the synonymy) (Michigan). — StrohiUi, lahyrinthica
D.U.L, Proc. U. S. Nat. Mus. VIII, 1885, p. 262 (Archer, Fla.).
— Strohilops cenea Pilsbry, Nautilus XL, Oct. 1926, p. 69.

This species is well characterized by being more depressed
than others of its region, with a decidedly angular periphery,
the base flattened just below the angle ; by the fine, thin ribs,
the dark color and the wider umbilicus ; moreover, there is no
internal fold above the periphery. It was recognized as dis-
tinct over thirty years ago, but at that time it was confused
with the Mexican species 8. streheli Pfr., which differs by its
far narrower umbilicus.

8. conea is often found associated with 8. lahyrinthica, these
two being the most abundant and widely distributed species
of 8trohilops in the states east of the Mississippi River. West
of the Mississippi 8. ccnea has been found only in Arkansas
and Louisiana. It is the common Strobilops of the Gulf
coastal plain.

4a. 8trohilops ccnea form ynicromphala, new form. PL 9, figs.
8, 9.
Lighter colored than 8. cenea; between cinnamon and cinna-
mon-brown; outlines of sjpire more convex; umbilicus nar-

STROBILOPS, CAN^yDA AND UNITED STATES. 31

rower, contained about 8 times in the diameter ; last third of
base ribbed. The infraparietal lamella does not emerge.
Tliree folds visible thi'ough the base. Height 1.7, diam. 2.5
mm. ; 5i/4 whorls.

Arkansas: Menard Landing, Arkansas Co., Ouachita Co.
(C. B. Moore). Louisiana: localities in Morehouse, Richland,
Franklin, Catahoula and Rapides parishes (C. B. Moore) ;
Grand Cane (Williamson) and Frierson (L. S. Frierson), De
Soto Parish; type loc. Boeuf River above Charlieville, Rich-
land Parish. Mississippi: Yazoo and Jackson counties (C. B.
Moore). Alabama: Wetumpka (H. H. Smith); Simpson
Island, Mobile River, Baldwin Co. (C. B. Moore). "Washing-
ton, D. C. (E. Lehnert). New Jersey: Barton Run, Burling-
ton Co. (Bayard Long). Pennsylvania: Tulpehocken Valley,
near Germantown (B. Long).

This form occurs in the lower Mississippi Valley further
southwest than the typical form of the species has been found.
By its small umbilicus it approaches the Mexican 8. streheli,
in which the ribs are more slender. I considered it a south-
western race of ^S'. cBnea until lots from "Washington, D. C,
Pennsylvania and New Jersey turned up, in which the um-
bilicus is almost as small as in Louisiana specimens — ^being
contained 7% times in the diameter. Further collections are
required to fix the status of this form, which appears quite
distinguishable in the series now available.

4&. Strohilops cenea spiralis, new subspecies. PI. 9, figs. 5, 6, 7.

The shell is somewhat less depressed than 8. cenea, light
brown ; ribs rather distinct on the base. The parietal lamella
is much longer, forming a full whorl in the type (but slightly
shorter, over three-fourths of a whorl, in specimens from
"Wyandotte, Indiana). The hasal harrier is situated more
deeply than in cenea, and consists, as in that species, of four
basal folds and a small, short one on the columella.

Height 1.9, diam. 2.6 mm. Type.

Height 2.0, diam. 2.6 mm. Paratype.

North side of the summit of Magazine Mountain, Logan Co.,
Arkansas {tyipe loc. ; F^rriss and Pilsbrj-, 1903). "Wyandotte,

32 STROBILOPS, MEXICO, CENTRAL AMERICA.

Indiana (L. E. Daniels, 1905). Koanoke, Randolph Co., and
Lookout Mountain near Valley Head, Alabama (H. H. Smith).

Species of Mexico and Central America.

All of the Mexican species except S. huhhardi belong to the
group of S. streheli, in which there is a small columellar
lameUa and three hasal folds; na folds above- the periphery,
such as occur in the lahyrinthica group. Tliis Mexican group
includes also the northern 8. conea and all of the South Amer-
ican species now known, except perhaps S. helleri.

In 8. veracruzensis crossei the internal barrier has not been
examined ; probably it conforms in this respect to other conic
Nootropic species. In 8. csnea there are usually four basal
folds, a minute one appearing between the second and third
larger folds.

Key to 8pecies.

1. Form subdiscoidal, the height about half the diameter;

umbilicus contained 3 to 4 times in diameter.

8. huhhardi, No. 15.
Form low-conic (2).
Form elevated, trochiform, the height 80 to 90 per cent of

the diameter (3).

2. Umbilicus contained 5 to Si/o times in diam.

8. (vnea mexicaim, No. 4c.
Umbilicus contained 10 to 12 times in diameter.

8. streheli, No. 5.

3. Umbilicus large, contained 3 to 4 times in diameter; shell

carinate, diam. 3 mm. 8. salvini. No. 7.

Umbilicus small, about 10 times in diameter; shell smaller,
subangular, with rather widely spaced ribs (4).

4. Diameter about 2 mm. ^S'. veracruzensis. No. 6.
Diameter 2.5 to 2.8 mm. 8. v. crossei, No. 6a.

4:C. 8trohilops cenea mexicana new subsp. PI. 5, figs. 9-13.

Umbilicus contained 5 to 5i/2 times in the diameter, thus
far larger than in 8. streheli, and somewhat larger than in
8. ccnea. There are tliree basal folds, a short one and two

STROBILOPS, MEXICO, CENTRAL AMERICA. 33

long, as in S. streheli. The color of shell and lip is cinnamon
in the type lot, all dead shells, but chestnut brown in the fresh
shells from Necaxa. The glossy base is largely smoothish, the
last third ribbed more or less.

Height 1.8, diam. 2.7 mm. ; 5% whorls. Type.

Height 1.95, diam. 2.75 mm. ; Si/o whorls. Necaxa.

Height 1.8, diam. 2.6 mm. Necaxa.

Eastern and northeastern Mexico: Diente, near Monterey,
Nuevo Leon (type loc. ; S. N. Rhoads, 1899) . Around Necaxa,
northern Puebla (H. Burrington Baker, 1926).

This form appears to be abundant in both of the localities
known, which are in a higher zone than that inhabited by S.
streheli. I have separated it from S. cunea with some doubt,
as the difference is slight, but the umbilicus in cenea seems
always to be a little smaller, the color of fresh shells is dif-
ferent, and adult individuals of cenea from the United States
generally have four basal folds.

5. Strobilops strebeli (Pfr.). PI. 5, figs. 1-4.

"Shell perforate, depressed-conoid, closely costulate above,
brown ; spire convexly conoidal. Whorls 5I/2, a little convex,
slowly increasing, the last not descending, with a crenulate,
subcarinate periphery, nearly smooth below the angle. Aper-
ture oblique, ear-shaped, furnished with two parietal lamellse
(the outer one reaching forward, the other more deeply
placed) ; peristome simple, the upper margin straight, the
basal a little reflected towards the insertion. Greater diam.
22/3, lesser 21/0, height II/2 mm." (Pfr.).

Helix streheli Pfeipfer, Malak. Blatter VIII, 1862, p. 71,
pi. 1, f . 5-8 ; Monographia Hel. Viv. V, p. 222.— Strohila laby-
rinthica Say (specimens from Mirador), Strebel, Beitrag
Mex. Land- und Siisswasser-Conch. IV, 1880, p. 43, pi. 4, f. 6
(only) ; pi. 15 fig. 7 ?. — Von Martens, Biologia Centr. Amer.,
Moll., 1892, p. 173.

Mirador specimens, which are doubtless part of the original
lot, are drawn in pi. 5, figs. 1 to 4. They agree fully with
three examples, 14554 Strebel collection, coll. by Berendt,
kindly lent by the Zoologische Staatsinstitut und Zoologische
Museum, Hamburg. Two of these measure :

34 STROBILOPS, MEXICO, CENTRAL AMERICA.

Height 1.65, diam. 2.5 mm. Umbilicus slightly more than
11 times in diameter.

Height 1.6, diam. 2.4 mm. Umbilicus 11 times in diameter.

Strebel's figures are reproduced in pi. 4, fig. 5. Pfeiffer's
description and figures appear to have been from a specimen
scarcely mature. The small umbilicus is contained about 11
to 12 times in the diameter. The rather strong ribs pass over
the very bluntly subangular periphery, but then gradually
disappear, leaving the greater part of the base merely striate ;
but under a high power, some specimens show faint traces of
fine, close, weak spiral striae. The shell is rather strong,
opaque and cinnamon-brown, the slightly expanded and thick-
ened lip of the same color. There is a slight spiral groove in
the latter part of the last whorl, just outside of the umbilical
suture. The white parietal lamella is high near its anterior
termination, penetrates inward exactly half of a whorl;
towards the inner end its edge shows strong nodes which are
roughened by microscopic, rather blunt asperities. The infra-
parietal lamella is visible in a basal view but does not emerge
to the edge of the parietal callus; it penetrates as far inward
as the parietal; between them, near the inner ends, is a very-
low, nodose interparietal lamella. There is a low columellar
lamella and three basal folds, that near the axis short. (In
one specimen, pi. 5, fig. 2, there seems to be two short and
two longer folds, seen faintly through the base, but the indi-
cations of folds are so weak in this shell that the appearance
drawn is not trustworthy; its visibility was exaggerated by
the draughtsman).

Height 1.8, diam. 2.6 mm. ; 5i/4 whorls.

Height 1.8, diam. 2.4 mm. ; 5 whorls.

5a. Strohilops strebeli guatemalensis Hinkley. PI. 5, figs. 5,
6, 7, 8.

The shell is more sharply ayigular at the periphery than 8.
stredeli, with a somewhat smoother base, microscopically stri-
ate spirally, and a less thickened lip. The width of the um-
bilicus is contained about 10 times in the diameter of the
shell. There are three basal folds.

STROBILOPS, MEXICO, CENTRAL AMERICA. 35

Height 1.55, diam. 2.4 mm. ; 5 whorls. Paratype.

Height 1.5, diam. 2.2 mm. ; 4% whorls. Type.

Guatemala : near Jocolo, in beach debris of Lake Izabal (A.
A. Hinkley).

Strohilops streheli guatemalensis Hinkley, Nautilus, vol. 34,
Oct., 1920, p. 52.

There is an obvious error in the measurement of width as
printed in Hinkley 's description. The type is figured.

6. Strobilops veracruzensis, new species. PI. 4, fig. 6 ; pi. 9,
figs. 1, 2, 3.
The shell is pyramidal, the spire with weakly convex out-
lines, periphery bluntly subangular, base moderately convex.
Umbilicus narrow, its width contained about 10 times in the
diameter of the shell. Surface slightly glossy, the initial l^/o
whorls smooth, next whorl with some fine striation, after
which there are narrow, oblique ribs, mdely and unevenly
spaced, passing over the periphery but then disappearing ex-
cept around the margin of the umbilicus and behind the lip,
leaving the greater part of the base smooth. The whorls are
moderately convex. The aperture is oblique ; peristome blunt,
expanded, thickened within, dull orange-cinnamon, the pari-
etal callus rather thin. The parietal lamella penetrates in-
ward about two-thirds of a whorl, its edge inwardly having
numerous low nodes. The infraparietal lamella emerges, but
not to the edge of the callus, and inwardly it is shorter, length
about half a whorl, and more conspicuously nodose than the
parietal, the nodes rugose. Between the inner ends of these
lamellaB there is a short, low interparietal lamella, consisting
of a series of about half a dozen rugose tubercles strung
together. Nearly a half-whorl inward there is a small blunt
columellar lamella, a larger inner basal fold, followed by a
much larger, high and long basal, and at some distance
peripherad a long, rather low and laminar third basal fold
(PL 9, fig. 2).

Height 1.9, diam. 2.1 mm. ; 5^ whorls.

Mexico: neighborhood of Vera Cruz (H. Strebel).

StroiUa labyrinthica, (Umgegend von Vera Cruz), Strebel,

86 STROBILOPS, MEXICO, CENTRAL. AMERICA.

Beitrag Fauna Mex. Land- und Siisswasser-ConchyL, IV, 1880,
p. 44, pi. 4, fig. 66. Not II. labyrinthica Say.

This species stands close to S. hraziliana of Para. It is
smaller, with the spire more straightly conic. By the high
contour and widely-spaced ribs it differs from other Mexican
species.

Strebel had two specimens of this species, both figured by
him, the figures copied photographically on my PI. 4, fig. 6,
and both are drawn in PI. 9, figs. 1-3. These specimens are
No. 14581 of the Strebel collection in the Zoologisches Institut
und Zoologisches Museum, Hamburg.

Strebel suspected that this form might be worthy of varietal
separation from S. labyrinthica, in which species he included
8. streheli, but it certainly differs widely from both. No com-
parative study of the different Strobilops forms had been made
at the time he wrote. ^S^. veracruzensis belongs to the strebeli
group, by the arrangement of internal folds, but it differs
strongly by the high contour and the widely-spaced ribs.

6a. Strobilops veracruzensis crossei, n. subsp. PI. 9, figs. 4, 4a.

The shell is narrowly umbilicate, conoidal, quite thin, with
well-spaced, slightly oblique riblets; pale brown. Spire con-
vexly conic, the apex rather obtuse ; suture impressed. Whorls
5%, very weakly convex and slowly increasing, the last whorl
not descending, obtusely subcarinate peripherally, and be-
coming nearly smooth below the carina and over all the basal
surface. The aperture is oblique, auriform, colored like the
rest of the shell, provided with and narrowed by two slightly
diverging parietal lamellge, of which the first is noticeably
stronger and more projecting, while the other, nearer the
columella, is less developed. Peristome simple, the margins
united by a tliin callous deposit, columellar margin slightly
reflected, the basal and outer margins weakly thickened,
upper margin straight. Height 2.25, greater diameter 2.5,
lesser 2.25 mm. {Fischer and Crosse).

Mexico: Plantation "Mirador," near Soledad, in the State
of Vera Cruz (Dr. Bereudt).

Helix strebeli Fischer and Crosse, Miss. Sci. au Mexique,

STROBILOPS, MEXICO, CENTRAL AMERICA. 37

Moll. terr. et Fluv., I, 1872, p. 267, pi. 12, f. la-h. Not H.
streheli Pfeiffer. — Strohila lahyrinthica (in einer hochgewun-
denen Form), Strebel, Beitrag etc., IV, 1880, p. 44.

This is a form which I have not seen, but, since it has been
figured by Fischer and Crosse, I introduce it here in order to
complete the account of Mexican Strohilops. It appears to
differ from 8. veracruzensis by the larger size and the more
prominent infraparietal lamella, if we may trust the figure,
which is possibly somewhat diagrammatic. Fischer and Crosse
state that it differs from lahyrinthica by the spaced instead
of close riblets, also they emphasize the perfect visibility of
two parietal lamellie, the infraparietal emerging more promi-
nently than in lahyrinthica. Their criticism of Pfeiffer 's fig-
ures of streheli was due to the fact that they did not have that
species before them, but a much more highly conic form.

Strebel stated that he had the Mirador plantation Strohila
in "a depressed and a high-coiled form, which last resembles
the North American form. Transitions are not present. ' ' He
gave the dimensions: height 2.4, greater diam. 2.8, lesser 2.6
mm. ; 6 whorls.

7, Strobilops salvini (Tristram). PI. 4, fig. 1.

Shell deeply umbilicate, conic, trochiform, rufous-corneous,
acutely carinate ; spire conic, the summit glossy ; suture deep ;
whorls 7, a little convex, regularly increasing, regularly orna-
mented above with transverse, acute and very conspicuous but
not continuous lirse ; the last whorl delicately striate beneath,
peristome somewhat reddish, polished, reflected; aperture
semilunar. Diam. maj. 3, miu. scarcely 3, height 2% mm.
( Tristram) .

Shell rather widely umbilicate, trochiform, angular, tawny,
obliquely ribbed above, smoothish below, showing by trans-
lucence three spiral laminae. Spire conic, of 51^2 slightly con-
vex whorls. Aperture rather oblique, lunar, the peristome
thickened, shortly reflected, the parietal wall provided with
an entering fold. Greater diam. 3, lesser 23/2, height 1^ to
2 mm. {v. Martens).

Northern Guatemala.: mountain forests of Vera Paz ( Salvin) .

38 STROBILOPS, MEXICO, CENTRAL AMERICA.

Helix salvini Tristram, Proc. Zool. Soc. London, 1863, p.
411. — Pfeipfer, Monogr. V, p. 334. — Strohila salvini von
JVLvRTENS, Biol. Centr.-Amer., Moll., p. 174, pi. 10, f. 1-lc.

I have not seen this species, which appears to be more ele-
vated and more distinctly angular than 8. strebeli, with a
•vsdder umbilicus than any North American species of its group.
The measurements given by von Martens from the type do not
agree well with his figures, and are given differently in his
table (Biologia, p. 173), perhaps partly transposed with those
of 8. labyrinthica. According to his figure, the umbilicus is
contained about 3.7 times in the diameter. Von Martens
writes as follows :

" An inspection of the typical specimens, kindly lent me
by Canon H. B. Tristram, enables me to give a fuller de-
scription of this species, which proves to be nearly allied to
8. labyrinthica (Say). The riblets of the upper surface cross
the angle of the periphery and extend a little way on the
lower face of the shell. The umbilicus occupies about one-
third of the diameter of the shell, whereas in 8. lahyriiithica
it occupies only about one-fifth of it.

' ' The internal lamellae are in this species about the same as
in 8. lahyrintliica, according to the description given by E.
Morse and the figures contained in Binney's works. Two
whitish spiral lamellae are visible from outside at the base,
shining through the shell, more distinctly in young shells than
in full-grown ones, because in those the shell is thinner; they
are situated really on the upper side of the lower wall of the
last whorl. Two others, situated on the lower side of the
upper wall or ceiling (parietal lamellae), are to be seen by
looking into the aperture of a not full-grown shell ; the upper-
most, or that nearest the suture, is the longest, extending
even somewhat beyond the limit of the aperture in the ob-
served specimen, the following one is shorter and lies more
beneath — it corresponds to the third of 8. labyrinthica in the
above-mentioned figures. A corresponding lamella to the
second of 8. labyrinthica is not visible in the specimens of 8.
salvini, probably only because it does not come so near to the
aperture, and I would not break the only shell which exhibited

STROBILOPS OP SOUTH AMERICA. 39

clearly those lamellse by seeking it. Under an ordinary lens
these lamellae show also in S. salvini the repeated swellings
described by Morse. In fragments of S. lahyrintJiica from
Florida, examined for comparison by myself and Dr. Hilgen-
dorf, under a higher magnifying power, the swellings appear
to be somewhat less regular, and composed of more scattered
and shorter points than in the said figures."

South American Species.

Strohilops morsel and 8. hraziliana appear to be closely re-
lated to species of Central America and Mexico, with which
they agree in internal armature.

By its smooth surface S. morsei is unique, but some speci-
mens show very weak traces of ribs, showing descent from a
ribbed ancestral stock.

8. helleri is a very distinct species, possibly forming a sep-
arate subgenus, but I have not opened the single specimen
found; its internal structure is not known.

Key to South American Species.

1. Shell elevated, trochiform, of 5I/2 to 6 whorls, the last with

angular periphery and weakly convex base ; umbilicus
narrow, contained 11 to 13 times in the diameter; diam-
eter about 2.5 mm. (2)
Shell rather low, of 4I/2 whorls, with dome-shaped spire,
carinate periphery and strongly convex base ; umbilicus
large, contained about 4 times in the diameter ; diameter
about 3 mm. Galapagos Islands. ;8'. helleri, No. 10.

2. Surface nearly smooth. Venezuela. S. morsei, No. 8.
Surface strongly ribbed. Brazil, ;S^. hrasiliana, No. 9.

8. Strobilops morsei (Dall). PI. 6, figs. 4, 5, 6.

The shell is trochiform with nearly straight lateral outlines,
obtuse apex, strongly angular periphery and weakly convex
base, perforated by a very small umbilicus, contained between
12 and 13 times in the diameter. The whorls are weakly
convex. Dead but rather fresh specimens are cinnamon-
colored without much gloss; nearly smooth, but there are

40 STROBILOPS OP SOUTH AMERICA,

some low growth-wrinkles, and in a few places the faintest
traces of a few ribs may be recognized. The base is smooth
except for some obliquely radial wrinkles in the umbilical
region. The peristome is slightly expanded and moderately
thickened. The parietal lamella emerges to the edge of the
parietal callus, the infraparietal nearly to the edge. There is
a short columellar lamella immersed about one-third of a
whorl, and three basal folds, the inner one short, the others
rather long ; the middle fold is arcuate, the outer one straight-
ened (fig, 5).

Height 2.3, diam, 2,5 mm. ; 6 whorls. Type,

Venezuela: Puerto Cabello (Sumichrast),

[Strohila labyrinthica] var. Morsel, and 8. Morsei Dall,
Proc, U, S, Nat, Mus. VIII, 1885, p, 263,

This species is well distinguished by its almost smooth
upper surface, the strongly angular periphery and the long
emergence of the infraparietal lamella. The basal plicae are
similar to those of the S. streheli group. Figs. 4, 6 represent
the type specimen, 24013 IT. S. N. M. ; fig. 5 is from one of
the shells collected by Ralph Tate, No. 22923 collection of the
Academy. The faint ribs traceable on some specimens are
apparently indicative of a costate ancestral stock.

9. Strobilops brasiliana F. Baker. PI. 6, figs. 7, 8, 9, 10, 11.
The shell is trochiform with somewhat convex lateral out-
lines, obtuse apex, angular periphery and weakly convex base ;
the very small umbdicus contained between 11 and 12 times
in the diameter. The whorls are weakly convex. Cinnamon-
brown, the surface without much gloss, last three whorls sculp-
tured with widely, unequally -spaced narrow ribs, the base
smoothish but with obliquely radial WTinkles in the umbilical
region. The lip, colored like the shell, is somewhat expanded
and moderately thickened. The parietal lamella emerges to
the edge of the parietal callus, the infraparietal nearly to the
edge; both penetrate about a half- whorl. Between their
inner ends there is a weak trace of an interparietal lamella.
All have nodose edges, the nodes bearing blunt, forwardly-
directed tubercles (fig. 11, edge of parietal lamella). There is

STKOBILOPS OP SOUTH AMERICA. 41

a short columellar lamella and three basal folds (fig. 8), the
inner fold short, the second long, arcuate, the third long and
straightened.

Height 2.3, diam. 2.6 mm. ; 53-4 whorls. Type.

Height 2.1, diam. 2.55 mm. Paratype.

Brazil : Para, under the bark of a decaying tree in the dense
forest surrounding the water works of the city (Fred Baker).

Strohilops hrasiliana Fred Baker, Proc. A. N. S. Phila.,
1913, p. 647, pi. 21, f. 8, 9 (Jan. 28, 1914).

This species stands near S. morsei, differing by the well-
developed ribs and the more convex lateral outlines of the
spire. The ribbing is represented as nearly regular in the
original figure and in our fig. 9 ; but it is noticeably irregular
in all of the eight specimens found, which are now No. 109310
of this collection.

10. Strobilops helleri (Dall). PI. 6, figs. 1, 2, 3.

The shell is solid, much wider than high, dome-shaped
above, with obtuse apex, convex beneath, with an acutely cari-
nate median periphery and a widely-open umbUicus contained
about four times in the diameter. The whorls are convex,
with a concavity just above the keel. Cinnamon-colored, with
little gloss; first 1% whorls smooth, after w^hich they are
finely, sharply costulate, the riblets becoming coarser and
more spaced on the last whorl, weak on the projecting keel,
rather close and somewhat irregular on the base (fig. 1).
The lip is expanded, thickened, a little angular at the outer
extremity, reflected at the columellar insertion. Parietal callus
heavy. The parietal lamella is rather low, enlarging near its
end, which does not quite reach the edge of the parietal callus.
Very deep in the throat a low, infraparietal lamella can be
seen in an oblique view in the mouth. At the last third of
the base two basal folds can be seen by transparence through
the base.

Height 1.9, diam. 3.1 mm., umbilicus 0.8 nun. ; 414 whorls.

Galapagos Islands : near Iguana Cove, Albemarle Island, at
2000 ft. elevation (Snodgrass and Heller).

42 STROBILOPS OF JiVPAN, KOREA AND CHINA.

Endodonta helleri D.vll, Proc. A. N. S. Phila., 1900, p. 94,
pi. 8, figs. 7, 8, 9.

The low contour, the acute, margined keel, the wide um-
bilicus and deeply immersed iufraparietal lamella, as well as
the small number of whorls, strongly characterize this species.
It appears to be most like the Central American S. salvini,
which however has more numerous whorls.

As 8. helleri is known by only one specimen (no. 108515 U.
S. N. M.), I have not examined the interior. Only two basal
plicae are visible through the shell, wliich is rather thick;
probably there are three, in addition to a columellar lamella.
The last two whorls are somewhat more convex above than the
draughtsman has represented them in fig. 3.

Section Eostrobilops, n. sect.

Gruppe der StroMlops diptyx Wenz, Neues Jahrb. Min.,
Geol. u. Pal., 1915, II, p. 84.

The shell is convexly conic, finely striate, with a rather
small umbilicus. The parietal and iufraparietal lamellae
emerge, and deep within their edges have prickly or rugose
knots or nodes (and in recent species flare outward) ; there is
no interparietal lamella. Internal barrier consisting of a low
columellar and two to four basal folds, sometimes also a palatal
fold. Type S. hirasei.

All known species of Japan, Korea and China belong to this
group. Some of the European Tertiary species are more de-
pressed than Recent forms.

Key to Species of Japan, Korea and China.

1. Larger species, diam. 3 to 3.5 mm.; umbilicus contained

about 5% to 7I/2 times in diameter of the shell (2).
Smaller species, diam. 2.5 to 2.7 mm. (3).

2. Internal barrier consisting of a columellar, 3 or 4 basal

folds and one above the periphery. Japan.

8. nipponica. No. 14.

Barrier consisting of a columellar and two basal folds.

Quelpart Island. S. hirasei, No. 13.

3. Umbilicus small, contained about 9 times in the diameter ; '

STROBILOPS OP JAPAN, KOREA AND CHINA. 43

internal barrier consisting of a columellar, 3 basal folds
and one above periphery. Korea. S. coreana, No. 12.
Umbilicus larger, contained about 5% times in the diam-
eter; internal structure unknown. China.

8. diodontina, No. 11.

11. Strobilops diodontina (Heude). PI. 10, figs. 1, 2, 3.
Shell very small, moderately umbilicate; fulvous, with a

silky gleam ; trochoidal, spire conic ; whorls 5, cylindric, the
last indistinctly depressed-angular, convex below; with an
epidermis provided with thick spiral striae and folds. Aper-
ture oblique, semilunar, obstructed by two parietal lamellae;
peristome expanded, reflected, umbilicus wide, penetrating.
Height 1.5, diam. maj. 2.5, min. 2 mm. {Heude).
China: Tchen-k'eou (Heude).

Helix diodontina Heude, Notes sur les Moll. terr. de la
vallee du Fleuve Bleu, in Mem. concernant I'Hist. Nat. de
I'Empire Chinois, 1880, p. 112, pi. 29, f. 11.

This species is known only from Heude 's incomplete ac-
count. His figures are reproduced. It appears to be similar
in size and shape to S. coreana, but from the description and
figures it should have a larger umbilicus, contained about 5%
times in the diameter of the shell. That of the Korean species
is much smaller. S. diodontina differs from 8. Jiirasei by its
smaller size.

12. Strobilops coreana Pilsbry, n. sp. PI. 10, figs. 8, 9, 10.
The shell is rather depressed with dome-shaped spire,

rounded periphery (very weaklj^ subangular in front) and
convex base ; umbilicus small, contained 9 times in the diam-
eter. Cinnamon-brown, rather glossy, with weak, rather fine
growth-wrinkles above, weaker on the base. The whorls are
rather strongly convex. The aperture is lunate, lip expanded,
heavily thickened within, russet. The parietal lamella is some-
what higher than the infraparietal, both emerging to the edge
of the parietal callus; they penetrate about one-fourth of a
whorl, then diminish to a low thread which continues some
distance further. They flare outward and are strongly nodose

k

44 STROBILOPS OF JAPAN, KOREA AND CHINA.

at the free edges. Nearly a fovirth of a whorl behind the aper-
ture there is an internal barrier composed of a short, blunt
columellar lamella, a series of three basal folds (the second
one longest, the others short), and a small fold within the
outer wall above the periphery. Some of these folds are visible
in an oblique view in the aperture.

Height 1.85, diam. 2.7 mm. ; barely 5 whorls.

Korea : Pyong-Yang, in the northwestern part of the penin-
sula (T. Kuroda).

Strohilops coreana Pilsbry, Proc. A. N. S. Phila., vol. 78,
p. 470, name only.

This is a smaller shell than S. hirasei, with smaller um-
bilicus, fewer whorls and four baso-palatal folds; differing
further by the infraparietal lamella, which emerges further
than in S. hirasei, about as far as the parietal lamella. The
internal armature of this species is practically identical with
that of S. nippomca, which is a larger shell with relatively
larger umbilicus.

The relation of 8. coreana to 8. diodonUna must be con-
sidered when specimens of that species become available.
I do not understand Heude's description of the sculpture of
diodonUna — " epidermide strids plicisque crassis spiralihus
donato ' ' — terms which could not be applied to the very weakly
obliquely striate Korean species. S. diodontinu is said to be
about the size of 8. coreana, with five whorls and two emerging
parietal lamellae, but the umbilicus is described as moderate or
wide, and figured as contained about 5i/^ times in the diam-
eter, thus differing from 8. coreana.

13. Strobilops HIRASEI Pilsbry. PI. 10, figs. 4, 5, 6, 7.

The shell is rather depressed, with dome-shaped spire, the
periphery mainly rounded but indistinctly subangular in front
of the aperture, the base rather strongly convex; umbilicus
small, widened in the last half-whorl, contained about 5l^
times in the diameter of shell (in some examples smaller, 6
times or 71/0 times in diameter). Opaque, cinnamon-brown,
without much gloss, smoothish, with low growth- wrinkles.
Whorls are strongly convex and increase slowly. The aper-

STBOBILOPS OP JAPAN, KOREA AND CHINA. 45

ture is oblique and lunate, peristome russet, expanded and
well thickened; parietal callus moderately heavy. The pari-
etal lamella is rather strong and emerges to the edge of the
callus. The infraparietal lamella is relatively strong though
much lower than the parietal, and emerges nearly to the edge
of the parietal callus. Both lamellae penetrate inward about
one-third of a whorl, being conspicuously nodose at the edges,
and there is a very weak continuation to about half a whorl
inward. At a point about one-fourth of a whorl inward there
is a low, short and blunt columellar lamella and two short
basal folds. All or part of these are visible in an oblique view
in the aperture, but owing to the opaque texture of the shell
they are not visible through the base in specimens examined.

Height 2.2, diam. 3.2 mm. ; 51/2 whorls. Type.

Height 2.1, diam. 3 mm. ; 5l^ whorls.

Korea: Cheju, Quelpart Island (T. Kuroda).

Strohilops hirasei Pilsbry, The Conchological Magazine, II,
No. 8, August, 1908, p. 39, fig. 1; Nautilus, XXII, p. 79.

This relatively large, solid Strohilops is distinguished from
the preceding and the following species by the number of
internal basal plicse (two). It is somewhat larger than S. dio-
dontina, in which the internal armature is unknown.

The growth wrinkles or striae are rather fine and somewhat
sharp below the suture, but are not regular in development in
the peripheral and basal parts.

14. Strohilops nipponica Pilsbry, n. sp. PL 10, figs. 11, 12,
13, 14.

The shell is depressed, with low dome-shaped spire, rounded
periphery (very obtusely subangular in front of the aperture)
and convex base, the rather wide umbilicus contained about
5I/2 times in the diameter; rather thin, the base somewhat
translucent. Cinnamon-colored, rather glossy, finely and
lightly striate, the stria; stronger below the suture. The aper-
ture is lunate, lip russet, slightly expanded, rather wide at
basal and columellar margins but less thickened than in the
Korean species. The parietal lamella emerges to the lip edge,
is high and flares outward. Infraparietal lamella is lower and

46 STROBILOPS, S. G. DISCOSTROBILOPS,

does not quite attain the edge. Both penetrate one-third of a
whorl and have strongly nodose edges (fig. 14). About one-
fourth of a whorl behind the aperture there is a barrier com-
posed of a short acute columellar lamella, three basal folds,
of which the second from the axis is largest, sometimes a
minute fourth fold at the periphery, and there is a short fold
mthin the outer wall above it (fig. 11 ; in this example the
third basal fold is longest, an exceptional condition). All of
these folds are usually visible in an oblique view in the aper-
ture, and also by transparence through the base.

Height 2.2, diam. 3.5 mm, ; 5 whorls.

Japan: Yonezawa, Uzen (Y. Hirase).

This beautiful little snail is larger than others known from
Asia — in fact, after the Endoplaces, the largest recent species
of Strobilops. It is more depressed than S. hirasei and has
more numerous basopalatal folds. It is closely related to the
Korean S. coreana, but is much larger, with a relatively wider
umbilicus. There is some variation in the basal folds (as in
fig. 11) among the ten specimens examined, but most of them
agree with the type.

Subgenus Discostrobilops, n. subg.

Gruppe der Strohilops uniplicata Wenz, Neues Jahrb. Min.,
Geol. u. Pal., 1915, II, p. 86.

The shell is thin, strongly depressed, the height less than
half the diameter, subdiscoidal with well-opened umbilicus;
finely eostulate or rib-striate above, smoother beneath. Pari-
etal lamella emerging; infraparietal lamella either wholly im-
mersed or weak and emerging; deep within, the edges of the
lamella are either smoothish or sparsely pi'ickly, without knots
or nodes; interparietal lamella either present or absent. There
is no columellar lamella. Basal folds three or four. Type S.
huhhardi.

Distribution : warm temperate to tropical parts of North
America and the West Indies; Upper Oligocene and Miocene
of Germany and Czechoslovakia.

This group comprises a single variable living species in
America and one species, S. uniplicata (Sandberger), with two

STEOBILOPS, S. G. DISCOSTROBILOPS. 47

subspecies, in central European Tertiary. In all of its char-
acters the Tertiary species agrees so completely with the living
S. Jiuhhardi that the relationsliip appears to be well estab-
lished. See PI. 8, figs. 10-13, S. uniplicata (Sclbg.), Hydro-
bienschichten, Budenheim bei Mainz ; Lower Miocene.

The parietal lamella of S. huhhardi generally appear to be
smooth, but under a high power the rounded edge is seen in
the most perfect specimens to bear irregularly and sparsely-
placed little points directed towards the aperture and some
minute granulation, but without a trace of nodes.

15. S. HUBBARDi (A. D. Brown). PI. 7, figs. 1-3 (type).

The shell is subdiseoidal with very low spire, rounded peri-
phery and convex, broadly umbilicate base ; width of umbilicus
contained 3% times in that of the base. The whorls are
strongly convex, increase slowly, and the last descends a little
to the aperture. Surface light brown (opaque in the "dead"
type, but when living doubtless glossy and somewhat trans-
lucent). Initial 1% whorls pale, minutely granulose, the rest
with slightly irregular sculpture of close, retractive riblets,
about as wide as their intervals, weaker and partly obsolete on
the base, where close, weak, microscopic spiral lines are seen.
The aperture is rounded-lunate, the lip well expanded and
thickened within. The parietal lamella is somewhat elevated
and triangular, reaching the edge of the parietal callus. Infra-
parietal lamella very low, inconspicuous, weakly emerging, but
not to the edge. Both enter slightly further than one-third
of a whorl; between them near their inner ends there is a
thread-like interlamellar lamella. At about a third of a whorl
within there is a series of four basal folds: the first one sit-
uated where the basal curves into the columellar floor; the
second, stout and erect, in the middle of the basal wall; the
third one small; the fourth near the periphery and longer
than the others.

Height 1.2, diam. 2.6 mm. ; 4i/2 whorls. Type.

Texas: Indianola, Calhoun Co., in the coastal plain; forms
of the species occur also in northeastern Mexico, Jamaica,
Cuba, Bimini Islands on the western edge of the Bahamas,
Bermuda, Florida, Georgia and Mississippi.

48 STROBILOPS, S. G. DISCOSTROBILOPS.

Helix huhhardi A. D. Brown, Proc. A. N. S. Phila., 1861,
p. 333, text-fig. — Strohila huhhardi Binxey, Terrest. Moll. V,
1878, p. 260, fig. 153; pi. 5, n. (dentition of a Georgia speci-
men).—W. Wenz, Nachrbl. D. M. Ges. 1916, p. 1S5.— St rohilus
huppardi Clessin, Malak. Blatter, n. F. VII, p. 79.

Brown's type is a "dead" shell, drawn in PI. 7, figs. 1-3.
After these figures were drawn it was opened and found to
have basal folds as described above, and like those of PL 8,
fig. 9. The anterior part of the infraparietal lamella is very
low. The interparietal lamella is delicate and thread-like but
perfectly distinct, and by possessing this lamella, the type
from the Texas coastal plain differs from all other huhhardi
from Mexico, the West Indies and Florida which I have
opened. The significance of this lamella is uncertain, in the
absence of series from the coast of Texas. It may turn out to
be merely occasional, and to be regarded as an atavistic muta-
tion. If it is a constant character in this region, a separate
subspecies is indicated. Temporarily, and in order to provoke
further investigation, I am taking this view.

15a. S. huhhardi vendryesimia (Gloyne). PI. 7, figs. 7, 8, 9
(topotype) ; figs. 4-6, 10-12. PI. 8, figs. 1-9.

General shape as in S. huhhardi; light brown, glossy, finely
costulate above, striate beneath. Infraparietal lamella either
wholly immersed (as in PI. 8, figs. 3, 4) or having a very low
extension forward (as in PI. 7, figs. 5, 7). No interparietal
lamella. Basal folds typically three (PI. 7, figs. 4, 7; pi. 8,
fig. 7), but frequently four (PL 8, figs. 1, 8, 9). Lip narrow,
but slightly thickened, more or less brown-tinted.

Height 1.05, diam. 2.6 mm.; 4% whorls. Topotype.

Jamaica: Bellevue, St. Andrew, type locality (Glo^Tie,
Bland and others).

Cuba: El Abra, Vinales (J. B. Henderson).

Bermuda: Admiral's Cave, Church Cave near Tuckerstown,
"Whitby Cave, Bailey's Bay, quarry between Tuckerstown and
Walsingham (A. Gulick, S. Brown, A. Haycock, H. C. Hoyt).

Florida: Sugarloaf Key, Big Pine Key (Pilsbry) ; Loss-
man's Key (C. B. Moore) ; Miami (S. N. Rhoads) ; LaCosta

STROBILOPSIDiE

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STROBILOPSID^

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A R Y

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STROBILOPS, S. G. DISCOSTROBILOPS. 49

Island, Demorey Key, Fikahatchee Key, Josselyn Key, Dismal
Key, all in Lee Co. (C. B. Moore) ; Volusia Co. at Tick Island
(Pilsbry and Johnson) ; and Lake Helen (G. W. Webster) ;
near Gainesville (T. Van Hyning) ; St. Au^stine (C. W.
Johnson); Imri, Hamilton Co. (E. B. Chope). Mississippi:
mouth of West Pascagoula River, Jackson Co. (C. B. Moore).
Georgia : Bonaventure Cemetery near Savannah, according to
W. G. Binney.

Mexico: Tampico and Valles, S. L. Potosi (A. A. Hinkley).

Helix vendryesiana Gloyne, Joum. de Conchyl. XIX, 1871,
p. 333. — Strohila huhhardi C. W. Johnson, Nautilus III, p.
20. — G. W. Webster, Naut. VII, pp. 84, 94 (variation in
teeth). — Strohilops huhhardi Gulick, Proc. A. N. S. Phila.
1904, p. 413 (Bermuda, fossil). — Verrill, Trans. Conn. Acad.
Sci. XII, p. 168, f. 53 (not uncommon living, Bermuda). —
G. H. Clapp, Naut. XXVII, p. 64 (Bimini Is.).— J. B. Hen-
derson, Naut. XXVII, p. 41 (many localities in Havana and
Pinar del Rio provinces, Cuba) . — Hinkley, Naut. XXI, p. 76
(Mexico).— Walker, Naut. XXXI, p. 56 (Palm Beach, Fla.).
— Vanatta, Nautilus XXI, p. 100; XXVI, pp. 12, 19, 32;
XXXIII, p. 18 (distribution in Florida). — Strohilops huh-
hardi stevensoni Pilsbry, Proc. A. N. S. Phila., 1899, p. 404. —
Rhoads, Nautilus XIII, p. 46 (under bark of dead limbs in a
mangrove swamp, Biscayne Bay near State Agricult. Sta.,
Miami, Florida).

Specimens from Cuba (PI. 7, fig. 6, El Abra, Vinales, Prov-
ince of Pinar del Rio) are small, as Henderson has remarked.
The one figured measures 2 mm. diameter with 41/^ whorls.
Sculpture weak.

The form found in Bermula (PI. 7, figs. 10, 11, 12) is large
with the lip somewhat more thickened than usual. There is
no interparietal lamella. The umbilicus is wider than in other
lots, but somewhat variable ; in a specimen 2.85 mm. diameter
the umbilicus is contained only 2% times.

In Florida (PL 8, figs. 1-6, Miami; 8, 9, Lake Helen) the
species appears to be generally spread. The parietal lamella
is low, the infraparietal generally not emerging though it be-
comes high far within. There are generally three basal folds
(PI. 8, fig. 7), but often four (PI. 8, fig. 9), and in rare cases

A(.

50 STROBILOPS, S. G. ENTEROPLAX.

five folds are seen in immature shells (PI. 12, fig. 9, Lake
Helen, Florida).

In a young shell from Miami 1.2 mm. diameter (PI. 8, fig. 5)
there are no basal folds, but the two parietal lamellae are well
developed. Folds are present but small in a shell 1.5 mm.
diameter, and at 2.2 mm. (PI. 8, fig. 6) they are longer than
in an adult of the same lot (PI. 8, fig. 7) .

Mexican specimens from Tampico (PL 7, fig. 5) and from
Valles, San Luis Potosi (PI. 7, fig. 4) are like those of Jamaica;
the few seen have three basal folds.

The form I described as S. huhhardi stevensoni (PI. 8, figs.
1-7) is identical with vendryesiana. It was distinguished from
huhhardi by the smooth base, somewhat wider umbilicus and
three basal folds — the type of huhhardi having more basal
sculpture and four folds. Large series which I have examined
and opened appear to show that the characters relied upon are
inconstant, and moreover, are those of vendryesiana.

Subgenus Enteroplax Gude.

Enteroplax Gude (as a section of Plectopylis) , Science
Gossip VI, October, 1899, p. 149. Type by original designa-
tion P. quadrasi.

The shell is thin, low conoidal to almost discoidal in form;
the edge of the parietal caUus thickened and raised, prominent
where the parietal lamella joins it; infraparietal lamella
emerging, both lamellfe with smooth or serrate (not nodose)
edges, and penetrating a half whorl; usually there is a low
radial callus across the parietal wall at their inner ends.
There is no columellar lamella. A radial series of few or many
basal folds is immersed from a fourth to a third of a whorl
back from the peristome. Soft anatomy unknown.

Distribution : Philippine Islands.

This little group is well characterized by the raised parietal
callus, somewhat like that of the typical Polygyras, and tlie
absence of a columellar lamella, as in S. huhhardi of the Sec-
tion Discostrobilops. The parietal lamellae are either smooth
at the edge, as in some Tertiary species of Strohilops, or the

STROBILOPS, S. G. ENTEROPLAX. 51

rounded edge is set with many low microscopic teeth pointing
towards the aperture, sometimes subobsolete, but evenly dis-
tributed, not gathered upon nodes as they are in typical Stro-
bilops. This is also a character of Discostrohilops, which on
the whole seems to be the most nearly related group. No
closely related fossil species have been found, and as Wenz
remarks, they are hardly to be expected. The group may very
likely have been evolved locally in its present area. All the
known species are quite thin shells having a delicate peripheral
carina over which the riblets of the upper surface do not pass.
These forms were first found by Dr. 0. von Moellendorff
when on a collecting trip in Cebu in company with 0, Koch,
a compatriot settled there and working up the butterfly fauna.
They ascended Mt. Licos, estimated to reach a height of about
500 meters, a peak circled by precipitous crags and a girdle
of high forest springing from a slope strewn with great blocks
of rock and rubble. Here in clefts and crevices of the rocks,
often reached by standing insecurely on the shoulders of his
guide, von Moellendorff found the Enteroplaces, Aulacospiras
and many other new snails. "None of my finds," he writes,
"have given me such joy as this one and the following {S.
polyptychia and trochospira) , secured almost at the peril of
my life."

Key to Species of the Philippine Islands.

1. Form subdiscoidal, with weakly convex spire and a series

of about 10 short basal folds. Cebu.

8. polyptychia, No. 16.
Form low-trochoidal ; fewer basal folds (2).

2. Four or five basal folds (3).

Three basal folds ; diam. 3.4 to 3.7 mm. Northern Luzon.

8. quadrasi, No. 19.

3. Diam. 4 mm. ; a low interparietal lamella extending inward

beyond the two parietals. Cebu. ;S^. trochospira, No. 17.
Diam. 3.2 to 3.7 mm. ; a low radial callus extending across
the wall at the inner end of the two parietals. Bohol.

8. hoholensis, No, 18.

52 STROBILOPS, S. G. ENTEROPLAX.

16. Strobilops polyptychia (Mlldff.)- PI- 12, figs. 14, 15, 16.
Shell openly umbilicate, diseoidal, closely costulate above,

striatulate beneath, browoi. Spire slightly raised ; whorls 51/0
to 6, nearly flat, the last subacutely angular at the periphery
and somewhat so around the umbilicus. Aperture diagonal,
obliquely heart-shaped, the peristome continuous, free, ex-
panded, a little reflected and thickened within ; parietal margin
deeply sinuate. Two parietal lamellse, one extending to the
edge of the peristome, the other falling a little short of it,
both penetrating inward to the middle of the whorl. Nine or
ten short palatal lamellae situated at the third part of the last
whorl. Height II/4, greater diam. 314, lesser 3 mm. (Mlldff.).
Cebu : peak of Mount Licos (Moellendorff ) .

Pledopylis polyptychia Moellendorff, Jahrb. D. Malak.
Ges. XIV, 1887, p. 272, pi. 8, f. 8-8c.; Abh. d. naturf. Ges.
Gorlitz, 1897, p. 147. — Gude, Science Gossip, IV, September,
1897, p. 102, fig. 55a-d.

The parietal and infraparietal lamellae enter fully a half
whorl; between, above and below their inner ends is a low
white callus forming somewhat triangular lobes with inwardly
directed angles in the intervals, its surface only weakly gran-
ular. The edges of the parietal lamellae appear under a high
power to be smooth and rounded. The ten basal folds within
the last third of the last whorl are short, slightly unequal, and
form a close series from the foot of the columellar wall to the
periphery. Two specimens measure:

Height 1.8, diam. 4.3 mm. ; 5% whorls ; umbilicus contained
2% times in diameter.

Height 1.5 diam. 4 mm., o^/^ whorls; umbilicus contained
about 3 times in diameter.

It is much lower than other species, with a more broadly
open umbilicus and more numerous baso-palatal folds (fig.l4).

17. Strobilops trochospira (Mlldif.). PI. 11, figs. 5a-c, 6a-e.

Shell openly and perspectively umbilicate, depressed-conoid,
costulate above, striatulate below, corneous, with conoidal
spire. Six slightly convex whorls, the last rather acutely

STROBILOPS, S. G. ENTEROPLAX. 53

carinate at the periphery, obtusely angular around the um-
bilicus. Aperture diagonal, lunar, the peristome thickened
within, expanded and slightly reflected, margins joined by a
callus. Two parietal lamellae, one reaching the edge of the
parietal callus, the other stopping a little short of it; both
penetrating deeply inward; at a third of a whorl within are
5 or 6 thin, rather long lamellae on the palatal margin oppo-
site (Mlldff.).

Height 2%, diam. 4 mm. (Mlldff.).

Cebu: peak of Mt. Licos (Moellendorff).

Plectopylis trochospira Moellendorff, Jahrb. D. Malak.
Ges. XIV, 1887, p. 273, pi. 8, figs. 9-9c ; Abh. d. naturf . Ges.
Gorlitz, 1897, p. 148.— Gude, Science Gossip, IV, March, 1898,
p. 285, fig. 73.

I have not seen this species, of which Gude writes :

''The aperture is diagonal, lunate; the peristome white, a
little thickened and reflexed, the margins being slightly con-
vergent and united by a scarcely raised sinuous ridge at the
parietal callus. The parietal armature consists of two long,
parallel, horizontal folds, which revolve over nearly half a
whorl, the upper one being the stronger and united to the
ridge at the aperture, while the lower one is thinner and
terminates at a short distance from the ridge; a very thin,
short, horizontal fold occurs posteriorly a little below the
upper fold (see fig. 6e, which shows the parietal wall of the
shell with its folds).

' ' The palatal armature is composed of five short, thin, hori-
zontal folds, which descend a little anteriorly (see fig. 6d,
which shows both the parietal and the palatal armature from
the posterior side). The specimen figured is in the collection
of Professor Oscar Boettger, of Frankfort, by whom this shell
— which measures : major diameter, 4 millimeters, minor diam-
eter, 3.5 millimeters; altitude, 2 millimeters — was obligingly
lent to me.

"Plectopylis trochospira is allied to P. quadrasi, but it is
larger and much lighter in color; there are also certain dif-
ferences in the armature" {Gude).

54 STROBILOPS, S. G. ENTEROPLAX.

18. Strobilops boholensis (Gude). PI. 11, figs. 1-4, 7-10.

Differs from the type {S. trochospira) in being smaller and
having a narrower umbilicus. Major diameter 3.25 mm., minor
diameter 3 mm., latitude 1.75 mm. The armature is nearly
identical, but the palatal folds are connected at their posterior
terminations by a very slight transverse sinuous ridge, plainly
discernible externally through the shell wall (Gude).

Philippine Is.: Bohol. Type in Mr. Ponsonby's collection
(now contained in that of Bryant Walker).

Plectopylis trocJiospira var. boholensis Gude, Science Gossip,
IV, March, 1898, p. 285, fig. 74.

In the Ponsonby Collection, now contained in that of Bryant
Walker, there are two specimens. No. 62067, one of which,
measuring height 1.95, diam. 3.3 mm., barely 6 whorls, I pre-
sume to be the type. The slight discrepancy in size — one-
twentieth of a millimeter in the diameter, two-tenths in the
height — might easily be due to differences in method of meas-
uring, especially in the height, where the points measured are
far out of alignment. The umbilicus is contained 3i/4 times
in the diameter; the ribs are well developed within its open-
ing. The two lamellge of the parietal wall slightly exceed half
a whorl in length; near the inner ends a low radial callous
band connects them. Their edges are serrate, as described
below. There are five basal folds, the one nearest the colu-
mella small, the second and fourth larger; there is a callous
ridge connecting their inner ends. The parietal callus is less
raised than in other species of Enteroplax.

In this form there is no interparietal lamella between and
beyond the inner ends of the parietals, such as Gude figured
for S. trochospira. If the latter normally has this structure,
the Bohol form must be specifically distinct.

In other lots from Bohol (A. N. S. P. No. 78457 and B.
Walker Coll. No. 32117, Sierra Bullones), out of the Quadras
collection, the shell is larger (PI. 11, figs. 8, 9, 10).

Height 2.2, diam. 3.7 mm. 78457.

Height 2.25, diam. 3.5 mm.

Height 2.3, diam. 3.6 mm. ' ^^^^'^'

STROBILOPS, S. G. ENTEROPLAX. 55

The umbilicus is contained 3I/2 times in the diameter. The
internal structure is like the type of hoholensis. The parietal
lamellae are slightly more than a half-whorl long, with an
irregular, smooth, radial callous band across the parietal wall
at their inner ends. Under a high power the rounded edge
of each lamella is seen to be closely set with short prickles
directed toward the aperture, so that in profile view the ap-
pearance is that of a fine but irregular saw (PI. 11, fig. 7).
These disappear towards the aperture, and are well developed
only near the inner ends of the lamellae.

19. Strobilops quadrasi (MUdfe.). PL 11, figs. 11, 12, 13, 14.

Shell rather openly umbilicate, conoid-depressed, thin, cos-
tulate above, striatulate beneath, corneous-brown ; spire little
elevated, with slightly convex lateral outlines; whorls 6, a
little convex, separated by an impressed suture, the last rather
acutely carinate at the periphery, obtusely angular around
the umbilicus. Aperture diagonal, irregularly heart-shaped;
peristome moderately expanded, strongly lipped within, brown,
the margins connected by a strong callus which is somewhat
deeply sinuate at the insertion of the outer margin and lamel-
larly raised. There is a high upper parietal lamella joined
to the callus and a lower one not reaching the callus, both
entering deeply, and opposite them, a third of a whorl within,
three rather strong and long palatal lamellae. Height 1.75,
diam. 3.5 mm. (Mlldff.).

Northern Luzon: village of Siamsiam (J. Quadras) ; Sitio
Cobayo, Palanan (Quadras).

Plectopylis quadrasi Moellendorpf, Nachrichtsblatt D.
Malak. Ges XXV, December, 1893, p. 172 ; Abhandl. naturf.
Ges. Gorlitz, XXII, 1898, p. 148.— Gude, Science Gossip, lY,
August, 1897, p. 71, fig. 54a-e. — Strohilops (Enteroplax) quad-
rasi Mlldff. and var. 'brunescens Mlldff., Wenz, Nachrbl. d. m.
Ges. 1916, p. 189.

The parietal lamellee enter half a whorl, and towards the
inner end their edges are set with low, rather blunt asperities
or granules, less developed than in 8. hoholensis. Across the
parietal wall and connecting the inner ends of the parietal

56 STROBILOPS.

lamellae there is a low whitish callus, irregularly trilobed in-
wardly, and on the apertural side having some tubercles along
V-shaped sinuses, or sometimes this pattern is not distinct.
The three thin, short and rather high basal folds, the middle
one largest, have their inner ends united by a low and nar-
row, irregularly-festooned radial callus. The size of the shell
is somewhat variable.

Height 2.4, diam. 3.7 mm. ; 6 whorls ; umbilicus contained
314 to 314 times in diameter.

Height 1.9, diam. 3.4 mm.; 5^ whorls; umbilicus contained
about 3^5 times in diameter.

Wenz mentioned a var. hnmescens Mlldff., which, so far as
I know, has not been described. Specimens before me so
labelled do not appear distinguishable from those received as
typical quadrasi.

APPENDIX TO STROBILOPSIDAE.

Species discovered since the publication of the classification
of Strobilops necessitate an extension of the synopsis of sub-
genera and sections given on page 18 and of the key to Mexican
and Central American species on page 32.

Key to Subdivisions of Strobilops.

1. Parietal callus having a thickened, raised edge, prominent

where the parietal lamella joins it ; parietal lamellae with
smooth or microscopically serrate edges, not nodiferous;
shell thin, carinate. Philippines.

Subgenus Enteroplax, species Nos. 16-19.
Parietal callus not noticeably thickened or raised at the
edge. (2)

2. Spire conoidal. (3)

Subdiscoidal, the spire low, periphery rounded, umbilicus
wide; parietal lamellse with sparsely prickly or smooth-
ish edges, not nodiferous.

Subgenus Discostrobilops, species 15.

3. Umbilicus large, contained 3 or 4 times in the diameter (4)
Umbilicus decidedly smaller. (5)

STROBILOPS. 57

4. Spire rather highly conic, of 5i/^ to 6 closely coiled whorls ;

parietal lamellae with nodes.

Section Coelostrohilops, species 7, 22.
Spire low conic, of 4^/^ whorls; lamellae of parietal wall
without nodes; no columellar or interparietal lamella.

Section Nesostrohilops, species 10.

5. Upper surface finely striate; no interparietal lamella.

Section Eostrobilops, species 11-14.
Upper surface costulate (except in >S^. morsel, which is
nearly smooth) ; an interparietal lamella present.

Section Strobilops proper, species, 1-9, 20, 21.
The following grouping will replace that given at the top
of page 18.

r Section Strobilops s. s.
/- Q Q+ v,"i J S^^^^^^ Coelostrobilops.

„ , „ I rs* -r^- 1 -1 * 1 Section Nesostrohilops.

Strobilops ^ S.-g. Discostrobilops. 1 g^^^.^^ Eostrobilops.
y S.-g. Enteroplax. ^

Species of Mexico, Central America and West Indies.
The following key is to be substituted for that on page 32.

1. Form approaching discoidal, the height about half the

diameter ; umbilicus contained 3 to 4 times in diameter.

8. hubbardi. No. 15.
Higher, the spire conic or conoidal (2)

2. Umbilicus large, nearly one-third the diameter (3)
Umbilicus decidedly smaller (4)

3. Periphery strongly angular. Guatemala. S. salvini. No. 7.
Periphery rounded. Grand Cayman I. S. wenziana. No. 22.

4. Umbilicus small, contained 10-12 times in the diameter (5)
Umbilicus contained about 5-6 times in the diameter; 3 or

4 basal folds (6)

5. Shell low conic S. strebeli. No. 5.
Spire elevated, the height 80 to 90 percent of the diameter

8. veracruzensis. No. 6.

6. Outer basal fold weakly developed (or sometimes want-

ing?) ; Socorro Island 8. hannai. No. 21.

Outer basal fold well developed (7)

it I B R A F

58 STROBILOPS.

7. Low conic; 3 basal folds. Mexico.

S. aenea mexicana, No. 4c.

Rather high conic; 3 basal folds and one above periphery.

Old Providence Island S. piratica, No. 20.

20. Strobilops piratica, Pils. PL 13, figs. 8, 9, 10, 11.

The shell is conic with the lateral outlines of the spire some-
what convex; umbilicate, the width of umbilicus contained 5
to 6 times in the diameter of the shell. There are 51/2 whorls,
the first 11/2 smooth, the rest with sculpture of retractive
riblets, which are about half as wide as their intervals on the
penult whorl; on the last whorl they are somewhat unevenly
spaced, partly closer. They pass over the periphery and con-
tinue over the base. The periphery is weakly subangular in
front of the aperture, elsewhere rounded. The aperture has
a blunt, weakly expanded lip and a moderately strong parietal
callus. The parietal lamella is strongly developed and pen-
etrates inward slightly more than a half whorl. Its slightly
thickened edge has swollen nodes bearing minute points and
granules. The infraparietal lamella penetrates inward nearly
as far as the parietal, is similarly nodose and emerges very
weakly. There is a short interparietal lamella consisting of a
series of tubercles weakly connected. The palato-basal barrier,
situated nearly a half whorl inward, consists of four folds:
three in the basal wall, the middle one much the higher, the
outer one narrow and low, and one small, very narrow fold
above the periphery.

Height 2 mm., diam. 2.2 mm.

Old Providence Island : on the summit of the ridge north of
High Peak, in leaf debris of forest. Type No. 150860 ANSP,
paratypes No. 150862.

StrohUops piratica Pilsbry, Proc. A. N. S. Phila., vol. 82,
p. 256, pi. 19, figs. 9-11. July 18, 1930.

This species has a higher, more couvexly conic shape than
S. veracruzensis, with a wider umbilicus. By the possession
of a palatal fold above the periphery and of ribs on the base it
differs from the Mexican forms, and resembles some northern

STROBILOPS.

59

species, such as 8. texasiana. It differs from that species by
the weaker ribs and wider umbilicus among other features.

I obtained no living specimens of this little snail, and indeed
did not see it in the field. Three entire and three broken
examples were found in leaf debris gathered on the ridge
running north from High Peak. {Piratica, belonging to
pirates — who used this island as a base after Morgan wrested
it from the Spaniards.)
21. Strobilops hannai, new species. PI. 16, figs. 3, 3a, 4.

The shell has a dome-shaped spire of closely coiled, moder-
ately convex whorls, a bluntly but distinctly angular periphery
and moderately convex base. Umbilicus contained about 614
times in the diameter. Color deep olive-buff, rather dull
above, glossy at the base. Sculpture: the first 11/2 whorls
smooth, the rest with retractive ribs about half as wide as
their intervals. Most of these ribs disappear after passing
over the periphery, but some continue diminished over part
of the base, and a few weakly to the umbilicus. The aperture
is lunate, oblique. Lip is white, moderately expanded, thick-
ened within. The parietal callus is thick. The parietal
lamella is strong, emerges to the edge of the callus, and
penetrates half a turn. Infraparietal lamella also penetrates
as deeply as the parietal, is rather high at the inner end but
weakens further forward, not emerging in the aperture. Both
of these lamellae have strong nodes which are profusely, shortly
spinose. There is a low interparietal lamella between the
inner ends of the large lamellge. There appears to be no
columellar lamella. Basal folds three: the inner or fold 1 is
well developed but short; fold 2 is larger; the third fold is
very small and thin (it apparently represents either No. 4
or 5 of the diagram on page 2 ; in a second specimen it could
not be seen, but possibly was broken away in opening the
shell).

Height 1.7, diam. 2.4 mm. ; 5i/^ whorls. Type.

Height 1.5, diam. 2.2 mm; 5i/^ whorls.

Socorro Island, off western Mexico, at 2000 feet (G. D.
Hanna).

60 STROBILOPS.

Strohilops lahyrinthica Say, Dall, Proe. Cal, Acad, Sci, (4),

vol. 15, No. 15, 1926, p. 480. ? Strobilops strebeli Pfeiffer,

Dall, I. c.

This species differs from S. lahyrinthica (Say) by the
smaller number of basal folds and absence of any above the
periphery of the cavity, among other differences. It is closely
related to ^S*. strebeli and 8. aenea. The former has a smaller
umbilicus, and both differ from ;S^. hannai by the strong de-
velopment of the outer basal fold, which is very weak in the
Socorro Island form. S. strebeli guatemalensis, which re-
sembles S. hannai by its angular periphery, has a far smaller
umbilicus, contained about 10 times in the diameter.

It is named for Dr. G. D. Hanna, who collected! the
specimens.

I have not seen the single specimen which Dall identified
as 8. strebeli, but I suspect that it wiU prove to belong to the
present species. DaU apparently did not open the shells for
examination of the internal armature.

Strobilops strebeli guatemalensis Hinkley. Page 35,
2d line.

The dimensions of the type should be : Height 1.4 mm., diam.
2.4 mm.

Section Coelostrobilops, n. sect.

Conic strobilops with broadly open umbilicus, contained 3
to 4 times in the diameter. Type 8. wenziana.
Strobilops salvini (Tristr.), p. 37, belongs to this group.

22. Strobilops wenziana Pilsbry. Plate 13, figs. 1-7.

The shell is conic with blunt apex, nearly straight lateral
outlines and rounded periphery, below which it is flattened
between periphery and the strong basal convexity around the
umbilicus; openly umbilicate, the width of umbilicus con-
tained nearly three times in that of the shell. There are 6
convex whorls, the first two smooth, the rest sculptured with
narrow retractive ribs, separated by spaces averaging on the
last whorl three times as wide as the ribs. Most of the ribs

STROBILOPS. 61

stop at the periphery, but a few on the last half whori continue
over the base ; they reappear in the umbilicus. The aperture
is lunate, the outer and basal margins about equally curved,
expanded a little, strongly thickened within. The parietal
lamella is strong and penetrates inward almost an entire whorl.
The infraparietal lamella emerges and is broad and united
with the parietal by a callus at the end ; it penetrates as far as
the parietal. Under a high power the edges of both lamellaa
show very indistinct swellings or nodes, which are slightly
roughened. The inner end of the parietal has more distinct
serration. There is no interparietal lamella. Within the base,
slightly more than a half whorl behind the aperture, are two
basal folds, corresponding to those numbered 1 and 2 in the
diagram of strobilopsid teeth (see p. 2, fig. 1)., the inner one
is lower than the strong outer one, and only about half as long
(fig. 5). No columellar lamella seen.

Height 2 mm., diam. 2.7 mm. ; umbilicus 1 mm. Type, figs.
1-3.

Height 1.9 mm., diam. 2.4 mm. Figs. 6, 7.

Heights 2.1 mm., diam. 2.8 mm.

Grand Cayman Island, about midway between North Sound
and Eed Bay. Type 150861 A. N. S. P. (Pilsbry).

StroMlops wenziana Pilsbry, Proc. A. N, S. Phila., vol. 82,
p. 238, pi. 19, figs. 1-7, July 18, 1930.

This species can be compared only with Strohilops salvini
(Tristram) of the mountain forests of Vera Paz, Guatemala,
which has a similarly wide umbilicus. I have not seen that
species, but it is said to be angular peripherally, of 51/^ whorls,
with only one lamella emerging on the parietal wall (the in-
fraparietal being shorter) . The shape of the aperture is quite
different, according to the figure of the type given by von
Martens. Unfortunately, the internal armature of S. salvini
has not been fully described. I have seen all other American
species of the genus, and none is at all like S. wenziana.

This is the only West Indian Strohilops of the conic group.
Its presence on Grand Cayman is the most surprising occur-
rence in the West Indian land snail fauna which has turned

62 STROBILOPS.

up in a long time. I am naming it in honor of my friend
Dr. W. Wenz of Frankfurt a.M., whose work on the Tertiary
Strobilops of Europe has contributed materially to our under-
standing of this ancient group.

Section Nesostrobilops, n. sect.

Strobilops with conic spire of few (4i/^) whorls, the parietal
lamellae without nodes ; no interparietal lamella and no colum-
ellar. Basal folds 3 or 4. Type S. helleri.

The only species known of this insular group resembles Dis-
costrohilops in having no colum.ellar lamella, fewer whorls
than in other Strobilops, and with the internal barrier less
deeply immersed, so that the folds can be seen in an oblique
view in the aperture. Also, there are no nodes on the parietal
lamella, though it is microscopically roughened.

10. Strobilops helleri (Dall). PL 16, figs. 1, la, 2, 2a.

See this vol., p. 41, where the type was described and
figured. The California Academy Expedition collected speci-
mens at the following places: Albemarle Island near Cowley
Mountain on moist ground at 350 to 500 feet ; on Narborough
in grass and bushes at the rim of the crater at 4000 feet; and
on James Island, in moist grass and bushes at 1800 to 2000
feet (W. H. Ochsner), in Dall and Ochsner, Proc. Cal. Acad.
Sci. (4), vol. 17, p. 176, 1928.

Tlirough the courtesy of Dr. G. D. Hanna of the California
Academy of Sciences I have obtained specimens of Strobilops
from Narborough and James Islands. The Narborough shell
(pi. 16, figs. 1, la) is barely mature, being still somewhat
transparent, with the lip somewhat expanded, but little thick-
ened. The base is finely striate, not ribbed, but ribs reappear
within the umbilicus. The peripheral carina is not distinctly
bordered above and below, as in the type specimen from
southern Albemarle. There are four basal folds, the fourth
(nearest the periphery) being longest, the second next, the
first and third shorter and about equal in length, though the
first is stouter (PL 16, fig. 1). The parietal lamella extends
inward very little beyond the basal armature; its inner half

APPENDIX TO PUPILLIDAE. 63

accompanied by a short infraparietal lamella, about as figured
for S. huhhardi in pi. 8, fig. 3. These lamellae have no nodes
or knots, but there is a weak microscopic roughening of the
edge about as described for Discostroljilops on page 47. I
could not get the objective near enough to determine the exact
structure without breaking the whole base. Height 1.7 mm.,
diam. 2.9 mm., umbilicus 0.75 mm.

The James Island specimen (PI. 16, figs. 2, 2a) is fully-
mature and opaque. It also is striate basally, but the striae
in part are somewhat thread-like, with ribs within the
umbilicus and a few near the lip. The expanded lip has a
white thickening. There is no little fold between the second
and outer basal folds. The parietal lamella, as in the Nar-
borough shell, does not penetrate more than a third of a turn.
The infraparietal reaches weakly almost to the aperture.
Height 1.9, diam. 2.9 mm.

There appear to be some small differences in form and
sculpture between the shells from different islands, but until
longer series are at hand for comparison it is not certain that
they have racial significance.

APPENDIX TO PUPILLID^

GASTROCOPTA (Vol. 24, p. 6).

Vol. 24, p. 8. The status of Leucochilus Boettger (1881,
type Pupa armifera Say) was left uncertain in the place cited.
This case has now been considered by the International Com-
mission on Nomenclature in Opinion 115, and the following
conclusion reached.

' ' The Commission herewith suppresses Leucochilus, 1881, in
favor of LeucocMla, 1860. Any other course would involve
risk of lasting and constant confusion in two rather closely
allied genera."

Vol. 24, p. 117, add to fossil species: Gastrocopta fissidens
infrapontica Wenz, Senckenbergiana vol. 9, 1927, p. 47, pi.
2, f . 8. Pliocene : Pontian beds of Leobersdorf , Austria.

64 GASTROCOPTA.

Gastrocopta gemenidens (Pils.). Vol. 24, p. 228; vol. 27,
p. 202.

A new section, Geminidens Pilsbry, 1930, was proposed for
Gastrocopta geminidens (Pils.) in Proc. A. N. S. Phila, 1930,
p. 351 (Dec. 13). It is characterized by the microscopically
granulose surface of the conic shell of few whorls, and
apertural dentition of Gastrocopta, the palatal folds not
immersed.

Gastrocopta pilsbryana (Sterki). Vol. 24, p. 36.

Pupilla stoneri Chamberlin and Jones (Bull. Univ. Utah,
xix, No. 4, June, 1929, A Descriptive Catalog of the
Mollusca of Utah, p. 83, fig. 32), of which I have examined the
holotype (PI. 15, fig. 3) through the courtesy of Professor
Chamberlin, is in all respects a typical G. pilshryaiia. The
type is a fresh specimen, spermaceti-wliitish, showing the
pinkish dried animal faintly through the shell. It measures
1.7 mm. long, 0.8 wide above the aperture; 5 whorls. It is
from Cedar Canyon about 10 miles from the mouth, east of
Cedar City, Utah.

The pale color of this specimen is not due to weathering, as
supposed to the authors, but is proper to the species. The
' ' fourth fold, back of the parietal tooth ' ' which they mention,
was evidently an effect of light, as there is no tooth in that
position.

The several forms described as paratypes by Chamberlin
and Jones are partly not conspecific. One sent me is Vertigo
coloradensis arizonensis P. & V. (No. 150656 A. N. S. P.).

GiS^TROCOPTA PAZi (Hidalgo). PI. 18, fig. 1.

Vol, 24, p. 102. The localities "Guayaquil" and "Panama"
are to be deleted. Also delete the phrase in the description
"or joined at the base and appearing like a bifid tooth".

Florentino Azpeitia Moros has discussed this species fuUy
in Revista Real Acad, de Ciencias de Madrid, vol. 22, 1925, p.
177-180, and has given a figure (p. 178, fig. 8) of an Amancaez
(Ecuador) topotype, which is reproduced in our PI. 18, fig. 1.

GASTROCOPTA. 65

It appears that G. wolfii and perhaps other forms of the
servilis group have been confused with the true G. pazi, which
appears to belong near G. harhadensis. I have not seen this
species.

Gastrocopta duncana, new species. PL 17, figs. 1, 2.

The oblong shell tapers slowly from the last whorl to the
obtuse apex, the lateral outlines being convex, the diameter
nearly equal at the last two whorls. Color translucent cin-
namon, the aperture tawny. Surface glossy, very weakly,
finely marked with growth lines. The whorls are moderately
convex, the last without wave or crest behind the outer lip.
The aperture is shortly oval. Peristome reflected, strongly
thickened within, the outer lip becoming narrower above, as
usual ; the terminations of lip are connected by a rather short,
somewhat heavy parietal callus, thin at the edge. Three or
four teeth, as follows: angulo-parktal lamella strong, simple
and straight. Columellar lamella rather strong, horizontal;
lower palatal fold rather strong, entering, the upper-palatal
minute and tubercular (often wanting). The summits of the
teeth are whitish.

Length 2.75 mm., diam. above aperture 1.2 mm. ; 51-2
whorls.

Galapagos : Duncan Island, on the south side, from about
500 feet to near the summit (Pinchot Exped.) Type 152689
A. N. S. P., coll. by Pilsbry and Cleaves, 1929.

This species differs conspicuously from other Galapagos
gastrocopts by the simple^ completely concrescent, angular and
parietal lamellsB, the absence of a basal fold, and the small
size or absence of the upper-palatal fold. It has some re-
semblance to the North American G. quadridens in teeth.
There is sometimes a very slight median depression in the
crest of the angulo-parietal lamella where the two elements
meet, but others do not show this.

All of the living examples were partly covered with an
irregular incrustation of black dirt, differing from G. munita
and G. clausa, which are generally clean. The seventeen speci-
mens found were picked out of lots of G. clausa, of Avhich

66 GASTROCOPTA.

there were over 1000 examples. G. duncana was much more
abundant at the summit of the island than lower down, but
it was found also in the lowest lot taken, at about 500 feet.

Nine of the specimens have no upper-palatal fold, though
apparently fully adult. This is a very distinct species.

Gastrocopta munita (Reibisch). PI. 17, figs. 3-9.

Vol. 24, pp. 96, 357. Originally described from Albemarle,
among bushes near the seashore (Th. Wolf), it was taken by
Ochsner near Tagus Cove and near Iguana Cove; on Nar-
borough at 50 feet ; Charles Island at 750 feet, and on Tower
Island. The Pinchot Expedition obtained it at Chatham
Island, from Wreck Bay to the lower edge of the grassy zone.
Charles Island at Black Beach, a mile inland. Barrington
Island. Daphne Island, near Indefatigable. Albemarle
Island in numerous places on the road from Villamil to San
Tomas, at a low elevation, and at Tagus Cove. Tower Island
around Darwin Bay. Often found in great abundance, in some
places associated with G. clausa. Several thousand specimens
collected by Pilsbry, 1929.

This a variable species, in which extreme forms might be
considered distinct if there were not so many transitional
examples. There are three main forms:

1. Corneous, ovate, typically with an interpalatal fold and
a subcolumellar callus ; about 2.4 x 1.2 mm. Typical G. munita
(PL 17, figs. 7, 8, 9).

2. Brown, subcylindrie, often lacking an interpalatal fold
and subcolumellar callus ; length 2.4 to 3 mm. Forma hrunnea,
n.f. ; chiefly from Kicker's Rock, Daphne Island and Tagus
Cove, Albemarle (figs. 3-6).

3. Brownish, subcylindrie, sometimes lacking an interpalatal
fold and subcolumellar callus ; small, length 2 to 2.2 mm.

Albemarle Island. — On the lava plain back of Villamil and
along the trail inland to San Tomas, only the typical form of
munita was found, the shell stout, ovate, corneous, with white
aperture (PI. 17, figs. 7-9; length 2.4, diam. 1.2 mm). This
is rather more likely to be the type locality than Tagus Cove,
as the latter is a rather out-of-the-way place.

GASTROGOPTA. 67

In a low, moist spot on the trail behind Villamil, a place not
covered by the last lava flow, brownish, partly more cylindric
shells were found, one of the smallest measuring 2.15x0.95 mm.

At Tagus Cove, Albemarle, I found "corneous", that is,
pale buffy gray examples (the typical color), of stout contour
with a white lip, such as I figured, vol. 24, pi. 19, fig. 1 (figure
too brown in the colored copies), together with more cylindric,
corneous and decidedly cylindric brown examples, all living
together. While a majority of the examples can be assorted
into two groups by color and shape, many intermediate shells
occur, so that the distinction of " Pupilla reihischi " Dall
appears to me impracticable. The data upon this supposed
species are given below.

Pupilla reihischi Dall, 1928, not 1917. — " Shell subcylin-
dric, blunt, five-whorled, of a dark brown color, whorls moder-
ately inflated, suture distinct; aperture with a wide reflected
margin which in well developed individuals is often of a
reddish color ; pillar lip reflected over a well marked umbilical
chink; teeth after Sterki's dental formula but substituting
figures for dots after his numeration : A B 3 D 4 5. The
parietal tooth (A) when fully developed is bifid anteriorly
and somewhat produced behind into the whorl ; the other teeth
appear short, and none of them are white. Length of shell
2.5 mm.; diameter 1.0 mm. {Dall).

" This species is easily distinguished from both P. mimita
and P. clausa by its darker color. It is more slender and
cylindrical than the former and larger than the latter. It has
been submitted to Dr. Sterki who pronounces it distinct from
munita. The accessory lamellse are rather deeply ensconced
in the aperture. The species seems to be considerably rarer
than P. munita. P. clausa, by the results of collections made,
is rare " (Dall).

Galapagos: Albemarle Island, near Tagus Cove, under
rotten wood at a height of 250 to 300 feet, mixed -with P.
munita (W. H. Ochsner).

Pupilla (Gastrocopia) reihischi Dall, in Dall and Ochsner,
Proe. California Acad. Sci. (4), XVII, June 22, 1928, p. 174.
Not Pupilla reihischi Dall, Proc. Biol. Soc. Washington XXX,
1917, p. 10; cf. Pilsbry, Man. of Conch. XXIV, 1918, p. 357.

The tooth formula (A B 3 D 4 5) used in the above descrip-
tion calls for the following teeth : parietal lamella, columellar

68 GASTROCOPTA.

lamella; infrapalatal, lower palatal, interpalatal and supra-
palatal folds. This cannot be correct; he apparently means
that there is a well united but bifid an^o-parietal lamella, a
columellar lamella, basal, upper and lower palatal and inter-
palatal folds; or according to Sterki's formula (Proe. U. S.
Nat. Mus. 1888, pi. 42, fig. 5) he should have written
1ABCD4E.

I have already (1918), explained that Ball's original
Pupilla reihischi of 1917 was based upon an error ; it was not
described, but the new name was proposed for '' Pupa wolfii "
as figured by Eeibisch. This being the case, the name becomes
a synonym of P. wolfii. The type of P. reihischi Ball, 1917,
is the specimen figured by Reibisch on his plate 2, fig. 11,
Abhandl. Nat. Ges. Isis, 1892 (see Man. Conch. XXIV, pp.
95, 357), which appears to be what Ball referred to in his
original proposition of the name. This figure represents the
Ecuadorian G. wolfii, which is hardly distinguishable from G.
servilis (Gld.).

The Ball and Ochsner paper of 1928 was published after
the death of both authors, and was thus not subject to their
final revision.

One of the original lot of reihischi, 1928, kindly given me
by Br. Ball, is figured, pi. 17, figs. 5, 6. Length 2.4 mm.,
diam. 1 mm. The color is between cinnamon and cinnamon-
bro'WTi on the last whorl, paler above, the lip and teeth light
russet vinaceous. The teeth represented are: a bifid angulo-
parietal lamella, a strong, simple and horizontal columellar,
and upper and lower palatal and basal folds. This specimen
has no interpalatal fold, such as Ball described ; but this small
fold is often lacking in P. munita. There is no swelling or
crest behind the outer lip. In any large lot of G. munita this
crest may be either present, weak or wanting.

I collected many specimens at Tagus Cove in 1929, the lot
including typical G. munita distinguished by its lighter corne-
ous (brownish or buffy gray) color, and the presence of an
infraparietal nodule and more or less callus under the colum-
ellar lamella. With them are darker, more cylindric shells
referable to the present race; but there are specimens of

GASTROCOPTA. 69

intermediate character, making a division arbitrary. All were
taken at one spot on the rim of the crater, directly inland
from the head of Tagus Cove.

Tower Island shells are rather small, 2.3 to 2.6 mm. long.
They are rather cylindric, the color corneous to light brown.

In the crater of Daphne Island, north of Indefatigable, both
brown and corneous forms occur. The shape is rather cylin-
dric. The infraparietal tubercle and the interpalatal fold
are often absent : 2.95 x 1.25 mm. Others are smaller, 2.6 x
1.15 mm.

On Kicker's Rock, off Chatham Island (PL 17, figs. 3, 4,
forma hrunnea) the shells resemble those of Daphne, reach-
ing a large size, up to length 3, diam. 1.2 mm., 5i/) whorls.
The form is subcylindric and the color brown. The infra-
parietal tubercle is lacking and the interpalatal fold often
absent. These two islands are arid breeding places of sea
birds, blue-footed boobies on Daphne, man-o'-war birds on
Kicker's Rock. The station of corneous and brown Gastro-
copta on the crater rim at Tagus Cove is similarly arid and
dusty.

Small or very small brown shells with the apertural char-
acters of munita have been noticed from a moist spot back of
Villamil, Albemarle. They also occur on Chatham Island,
where there seem to be transitions to small cylindric forms of
clausa, as noticed under that species.

Gastrocopta clausa (Reibisch). PI. 17, figs. 10-19.

Vol. 24, p. 99. This species is not distinguishable from 0.
munita by any characters of shape, size or color, all being
variable in both species. The number of teeth also varies
somewhat, but I have found the only really distinctive char-
acter to be the presence of a well developed suhcolumellar,
tooth in G. clausa. G. munita has merely a sloping callus
below the columellar lamella, sometimes very weakly tuber-
cular, or there is none. In typical G. clausa there is a nodule
peripherad of the inner end of the parietal lamella (PI. 17,
fig. 12) not present in G. munita.

Typically developed specimens of each, such as I figured in

70 GASTROCOPTA.

Man. Conch, vol. 24, pi. 19, figs. 1-3 (munita) and figs. 12-14
(clausa), are very readily distinguishable.

Distribution: Originally described from Indefatigable
Island (Theodor Wolf), it was taken on Abingdon by Snod-
grass. During the Pinchot Expedition I found it on Chatham,
from near shore around Wreck Bay to the lower limit of the
grassy zone on the road to Progreso. Charles Island at Post-
ofifiee Bay and Black Beach. Barrington Island. Indefatig-
able Island at Academy Bay, Seymour Bay and Conway Bay.
Duncan Island, from about 500 feet to the summit. Albemarle
Island inland from Villamil. The localities are all in the
arid zone, except on Chatham and Duncan where it extends
to the lower edge of the humid region. It is a very abundant
species near shore, around the roots of shrubs and among
rocks.

In many lots there are two main forms: a larger, about
2.6 X 1.3 mm., 5 whorls, ovate-conic, the color very light
brown, aperture white (PL 17, fig. 10) ; and a smaller form,
about 2.1 X 1.0 mm., 4% whorls, cylindric, of a darker color,
between cinnamon and cinnamon-brown, the aperture colored
(PI. 17, fig. 11). These forms agree in teeth and in having a
low crest behind the outer lip. There are some intergrading
specimens in both shape and color of shell and aperture ; albino
individuals of both forms occur. Those figured are from an
aa ridge on the trail between Villamil and San Tomas, Albem-
arle, in the arid zone.

At Academy Bay, Indefatigable Island, small and large
forms measure : 2.2 x 1.1 mm. and 2 x 0.95 mm.

Duncan Island at about 500 feet on the southern side : 2.6 x
1.15 mm., 2.2 x 1.15 mm. and 2.2 x 0.95 mm. (PI. 17, figs. 16,
17,19).

In the most fully developed examples of G. claitsa there are
no less than eleven teeth (PL 17, fig. 18, a mile inland from
Black Beach Road). Usually however, there are nine.

On Chatham Island back of Wreck Bay typical G. clausa
occurs, though the teeth are less strongly developed than in
many specimens from islands westward. There are also
small brown examples in some of the lots, not often over 2.25

GASTROCOPTA. 71

mm. long, and down to 2x1.05 mm. (PL 17, figs. 13, 14, 15).
In these the infraeolumellar tubercle varies from well de-
veloped to absent, and the interparietal fold is often wanting.
I at first tried to divide the lots into two species, clausa and
munita, and it may be that such a division is possible, but I
am more inclined to believe the Chatham form a variable,
relatively unspecialized population. These small, ill-character-
ized clausa with munita-like forms occur together near the
shore and also as high as the upper edge of the wooded zone
at the border of the grassy zone, on the road to Progreso.

Gastrocopta thomasseti Pilsbry. PL 14, figs. 5, 6.

The shell is rimate, thin, cylindric-oblong, the last two
whorls almost equal in diameter, those above tapering to a
very obtuse summit; pale brown fading to nearly white near
the summit, composed of extremely convex whorls, the first
two microscopically granulose, narrow riblets beginning then
very weakly, on the last two whorls becoming conspicuous,
rather widely, unevenly spaced, some ripples between them.
There is no crest or ridge behind the outer lip. The aperture
is somewhat squarish with rounded angles and a semicircular
basal outline, the peristome thin, expanded. The angulo-
parietal lamella is more or less distinctly bilobed in a front
view, there being usually a slight depression between the sum-
mits of the angular and parietal elements. The columellar
lamella is rather strong, subhorizontal. The lower-palatal
fold is moderately elongate. No other folds are present
typically, but in one example a very small upper palatal
nodule is developed.

Length 2.15, diam. above aperture 0.95 mm. ; 5 whorls.

Length 2.25, diam. above aperture 1.05 mm. ; 5 whorls.

Kopje on Block Island, Weenen, Natal. Type No. 145474
ANSP., paratypes in South African Museum. Collected by
Mr. H. P. Thomasset, 1923-1926.

Gastrocopta thomasseti Pilsbry, Ann. Natal Museum VI,
part 2, p. 301, pi. 20, f. 5, 6.

This species belongs to the little group comprising Gastro-
copta klunzingeri (Jickeli), var. senegalensis (v. Maltzan)

72 GASTROCOPTA.

and G. hermosa (Jousseaume) from the northern part of the
Ethiopian Region in Abyssinia, Eritrea, Senegal, and British
East Africa. The discovery of a closely related form in Natal
extends the range of this group of gastrocopts, which is
characterized by the great convexity of the whorls. It prob-
ably occurs in places all over the Ethiopian Region.

Pupa klunzingeri var. senegalensis von Maltzan appears to
agree rather closely with G. thotnasseti except perhaps in the
sculpture, which is described as more densely and delicately
costulate than G. klunzingeri. It has not been figured, and I
have not seen specimens. Moreover, the name of that Senegal
form is preoccupied by Pupa senegalensis Morelet. G. klun-
zingeri of Abyssinia and the Kenya Colony tapers more
rapidly than G. thomasseti, the last whorl being wider, the
costulation is slightly stronger and more regular, and both
upper-palatal and basal folds are developed. In the speci-
mens of G. thomasseti seen, a very small upper-palatal fold
was noticed in one only.

While the summit of the angulo-parietal lamella is generally
distinctly bilobed, as in the figured type, there are a few
examples in which it is level, the angular and parietal com-
ponents being more completely united.

Gastrocopta dahli Thiele. PL 18, fig. 2.

Two colorless shells from Ralum are rather cylindric, with
weakly, obliquely striate, convex whorls. Apex rather obtuse.
The upper whorls increase distinctly; the penult is about as
wide as the last but not so high. The aperture is rather
small ; a parietal lamella is bifid, its outer part passes in a
curve into the outer lip; in the right side there is a tooth
some distance from the margin, and a second, somewhat
longer, where the outer passes into the basal margin. On the
columella a denticle on the lower part of a more inwardly
placed thickening is visible. The peristome is widened but
not expanded; the umbilical fissure is hardly perforated.
Height 2.25, diam. 1.1 mm. {Thiele).

Bismarck Archipelago: Ralum (Dahl).

FAUXULUS. 73

Gastrocopta dahli Thiele, Zoologische Jahrbucher, Abth.
f. Syst. etc., Bd. 55, p. 124, pi. 5, figs. 8, 8a. 1928.

Gastrocopta semiclausa Thiele. PI. 18, fig. 3.

Two shells, also from Ralum, differ from the preceding
species by the somewhat larger and more ventricose shell with
the same number of whorls, and especially by the dentition
of the aperture. This has on the right side 3 teeth, a small
denticle near the parietal lamella and a rather broad, lamelli-
form projection of the columella obliquely in the interior of
the terminal part of the last whorl. Height 2.6, diam. 1.2
mm. (Thiele).

Bismarck Archipelago: Ralum (Dahl).

Gastrocopta semiclausa Thiele, Zoologische Jahrbiicher,
Abth. f. Syst. etc., Bd. 55, 1928, p. 125, pi. 5, figs. 9, 9a.

Belongs in the kindred of G. macdonelli (Brazier), from
which it may be considered specifically distinct ( Thiele ) .

GYLIOTRACHELA Tomlin.

Gyliauchen Pilsbry, Manual Vol. 24, p. 174, 210, 1917. Not
of Nicoll, 1915. — Gyliotrachea Tomlin, Proc. Malac. Soc,
Lond., Vol. 19, March, 1930, p. 24.

FAUXULUS (Vol. 24, p. 234).

Fauxulus burnupianus Pilsbry. PL 14, figs. 1, la (type),
2,3.

The shell resembles F. capensis in form, being cylindrie with
a shortly conic summit, sinistral, with a deep, comma-shaped
umbilicus ; thin ; opaque whitish with a faint pink tint and
some sparsely scattered purplish-brown dots and small spots.
The surface has but little gloss, and is closely sculptured wdth
fine, rather sharp, curved striae, somewhat weaker on the
last whorl and wanting on the first 1% whorls, which are
slightly roughened by a close, superficial, somewhat uneven
microscopic pitting. The early whorls are convex ; later ones
are slightly convex, the last whorl compressed and tapering

74 PAUXULUS.

do^vllwards, forming a narrowly rounded base; it ascends to
the aperture and is somewhat excavated within the umbilicus,
which becomes a mere perforation beyond the last whorl.

The aperture is shaped like that of F. capensis, ochraceous
within, and has three lamellae as in that species. The angular
lamella is rather long and a little impressed or sinuate in the
middle. Parietal lamella enters deeply. The columellar
lamella is strong, horizontal, and enters about a half whorl.
There are three folds a short distance within the outer lip :
the upper palatal fold is very long, its inner end curving
downward; interpalatal fold small (often twinned) ; lower
palatal fold long, its inner end not visible in a directly face
(ventral) view. The peristome is white, thin and well ex-
panded, the margins connected by a thin parietal callus.

Length 5.9, diam. above aperture 2.8 mm. ; 9I/2 whorls.
Type.

Length 5.2, diam. above aperture 2.9 mm. ; 8I/2 whorls.

Length 5.2, diam. above aperture 2.6 mm. ; 9 whorls.

Length 7.1, diam. above aperture 3.1 mm. ; 10 whorls.

South Africa: Seal Rock, opposite Dyer Island, Cape
Province. Type 144264 ANSP., collected by Herbert Lang.

Fauxidus hurnupianus Pilsbry, Nautilus, vol. 41, Jan.,
1928, p. 108; Ann. Natal Museum VI, part 2, 1929, p. 300,
pi. 20, f. 1-3.

This snail has the shape, texture and color of F. capensis
(Kstr.), and is about the size of the small race of that species
from Port Elizabeth. It differs by the larger apertural
lamellse and folds, which have more the form of those teeth
in F. pamphorodon (Bens.), the upper-palatal fold being
laminar, entering deeply and descending, and the lower-
palatal is prolonged inward.

The shell is whitish to the eye but under a lens is seen to
be delicately tinted pink or bluish, variously but sparsely
maculate, and sometimes having an ochraceous band below
the suture, such as occurs in certain shells of F. capensis. Of
the 18 examples seen, only one exceeds 6 mm. in length.

This species was found with specimens of F. capensis 7 to

FAUXULUS. 75

9 mm. long, and F. layardi. It is named in honor of the late
Mr. H. C. Bumup, author of several excellent papers on South
African snails.

Subgenus Anisoloma Ancey.
Fauxulus falconianus Pilsbry. PL 14, figs. 4, 4a.

The shell has a small perforation and long rimation, is
ovate-conic, tapering from the last whorl, the spire attenuate
above, where the outlines are lightly concave ; base compressed
in its last half. The color is light cinnamon, paler at the
summit. The first two whorls are strongly convex and smooth ;
following whorls are nearly flat, with sculpture of delicate
oblique striae closely placed but much narrower than their
intervals and bending forward as they approach the suture
below. On the penult whorl the striae are coarser and more
strongly curved forward near the anterior suture. The last
whorl has less oblique riblets nearly as wide as their intervals.
The aperture is somewhat triangular, there being an impres-
sion or bay in the middle of the left margin, which projects in
a semicircle above it. The microscope shows a dense, irregular
papulation over the interior of the aperture as well as upon
the thin parietal callus covering the riblets above it. The
angular and parietal lamellge are about equal in height, the
former emerging to the outer lip near its insertion. Colum-
ellar and supracolumellar lamella about eciual, long, emerging
to the edge of the columellar lip. Infracolumellar lamella
represented by a small nodule (or in other specimens, absent).
Basal fold small, short, receding. Lower-palatal fold is sinu-
ous, emerges to the lip-edge and enters deeply. The upper-
palatal fold is long but much shorter than the lower, towards
which it converges inwardly. The suprapalatal fold is short
but in front view about as prominent as the upper-palatal
fold.

Length 4.4 mm., diam. 2.6 mm. ; 9 whorls. Type.

Length 4.5 mm., diam. 2.6 mm. Paratype.

Natal : Zimbaba Bush, Krantzkop, collected by W. Falcon,
1923. Type No. 145477 ANSP., cotype in coll. Natal Museum.

76 ABIDA.

Fauxulus falconianus Pils., Ann. Natal Museum VI, May,
1929, p. 299, pi. 20, f. 4, 4a.

By its attenuate spire this species resembles Fauxulus per-
eximius (Melvill and Ponsonby), but that is a far larger shell,
said to be 7 ram. long. The figure given by Melvill and
Ponsonby is not sufficiently exact to admit of any comparison
of the teeth ; if it is to be trusted their species must be quite
unlike F. falconianus in the shape of the outer lip. None of
the other Fauxuli of the subgenus Anisoloma now known have
the spire attenuate as in these species.

ABIDA (Vol. 24, p. 262).
Abida variabilis sillarensis (Piersanti). PI. 18, figs. 5, 6, 7.

Shell brown or cinereous, alabastrine, elongate, cylindric,
not swollen in the middle, the peristome expanded, generally
aorthic, sometimes with a white epistomatic callus. 10-13
whorls.

Length 11.2 to 14.3 mm., diam. 2.9 to 3.2 mm. (Piersanti) ;
86 per cent of the specimens are from 2.9 to 3 mm. in diameter,
and 178 out of 200 are from 11 to 12 mm. long.

Italy : Appenines of the upper Sillaro valley, Prov. Bologna.

Pupa variahilis Drap. v. sillarensis Piersanti, Boll, dell'-
Istituto Zoologico della R. Univ. di Roma, V, 1927, p. 129-134,
figs. 5-7 (fig. 5, variation orthostoma, fig. 6, variation plagio-
stoma, p. 134).

A less convex, more cylindric shell than A. variahilis, differ-
ing also from the cylindric A. v. polita, and a descendant of
var. producta, found by Strobel on the Tidone and the Secchia
( northwestward ) .

The locality of this race is a wind-beaten treeless terrace
with southern exposure, at an elevation of about 400 meters,
having scant vegetation of Artemesia, broom etc., reached by
a mule trail which ascends from a spring of sulphur water
on the left bank of the upper Sillaro. There is a fauna of
mullusks numerous in individuals but few in species, among
them this race of Abida variahilis. It lives in families of 20
or 30 individuals on small di-y bushes or broom. Even when

GRANOPUPA. 77

they are covered with snow and ice it does not seek refuge
by burrowing or under stones. It is capable of fasting an
entire year.

Two forms are named: orthostoma (PL 18, fig. 5), and
plagiostoma (PL 18, fig. 6). These are merely individual
variations in the inclination of the axis of the aperture to
that of the shell.

Piersanti, from whose paper the above is taken, considers
A. V. sillarensis a recent immigrant, as it was not mentioned
by Tassinari, who collected in the Sillaro valley. It is con-
sidered the furthest outpost of one of the two streams of
migration of variabilis into Italy.

Abida pyrenaeablv (Mich.). Vol. 27, p. 277.

Ninth line from bottom, in place of " pi. 24, figs. 7, 8, 9 "
read pi. 24, fig. 10.

Fourth line from bottom, in place of " pi. 24, fig. 10 "
read pi. 24, figs. 7, 8, 9.

On p. 328, lines 15-17 from bottom: the names P. saxicola
M.T. and P. clausilioides Boub. were transposed.

Abida frumentum gigantea (Schroeder). Vol. 24, p. 303.

Light horn-colored, the whorls, especially the middle ones,
very finely and obliquely striate transversely. 15-17 mm.
long, 4-5 mm. thick, with 12-13 whorls. {Schroeder) .

Italy: Salo in Monte S. Bartolomeo, Lake Garda, also at
Gargnano.

Pupa {Torqiiilla Stud.) frumentum Drap., var. gigantea
Schroeder, Beitrage zur Konchylienkunde von Tirol und
italienischen Grenzgebieten, 1910, p. 40.

The name is preoccupied by Moquin-Tandon, see Vol. 25, p.
32 ; but Schroeder 's form need not be renamed until its status
can be investigated anew.

GRANOPUPA (Vol. 24, p. 332).
Granopupa marmouchana (PaUary). PL 18, fig. 4.

Shell elongate, conic, of a deep brown tint, provided with
an umbilical crevice. Spire acuminate, tapering, terminating

78 CHONDRINA.

in a smooth, glossy protoconch, 9 convex whorls of regular
increase, separated by a not very deep suture, and regularly
sculptured with fine striae oblique from left to right, the
closeness of the striae emphasized on the lower whorls. Penult
whorl well rounded, last whorl ascending a little to the in-
sertion. Aperture very little oblique, a little reflected at the
outer margin, with two oblique internal lamellae which do not
reach the outer margin. Columella vertical, well rounded
at base and ornamented with a small denticle (angular
lamella) on its upper part. Finally the parietal wall carries
two internal folds of which the upper (parietal) is more
prominent than the median; these two lamellse opposite the
two palatal lamellae (Pallary).

Length 7, greatest diam. 214, smaller diam. 2 mm.

Morocco: Talzent, in the gorges of Meskedal and Arourirt
on cliffs of Jurassic limestone.

Pupa marmouchana Pallary, Journ. de Conchyl., vol. 72,
August, 1928, p. 13, fig. B.

This species seems to be near Pupa rhodia Roth of Syria,
but our form has less convex whorls and the size is smaller.
No North African species is comparable (Pallary).

Granopupa granum (Drap.). Vol. 24, p. 335.

Pallary (Mem. I'lnst. d'Egypte XI, 1926, p. 40) thinks
that Pupa lamarckii Aud. (Vol. 26, p. 238) is a repaired
Granopupa granum.

CHONDRINA (Vol. 25, p. 1).
Chondrina avenacea (Brug). Vol. 25, p. 10.

Var. elangata Stossich. Form long cylindric, of 8 whorls.
Ospo, territory of Trieste. (Torquilla avenacea var. elongata
A. Stossich, Boll. Soc. Adriatica di Sci. Nat. in Trieste, vol.
19, 1899, p. 32).

Paul Ehrmann (Archiv f. Molluskenkunde, vol. 63, pp. 1-
28) believes that two species are commonly comprised under
the name avenacea. The true Pupa avenacea he believes to
be identical with P. cereana (" Miihlfeld " Kiister). For

CHONDRINA.

79

the other species, usually known as avenacea, he uses the name
clienta Westerlund, contrasting them thus.

C. avenacea (Brug.)

1. Shell surface irregularly
striate.

2. Both principal palatals and
the infrapalatal almost
equally strong, the parietal
and columellar lamellae placed
opposite them.

3. Only the upper whorls
strongly convex, the middle
and lower laterally com-
pressed.

4. Last whorl usually strongly
flattened or even impressed
below the shoulder-like swell-
ing behind the aperture.

5. Peristome little expanded,
weakly lipped, light brown-
ish ; interior of aperture more
or less reddish brown.

6. Color of shell usually light-
er or darker reddish brown.

C. clienta (West.)

1. Shell regularly, finely
striate.

2. Only the two principal
palatals strongly developed ;
super- and infra-palatals, if
present, very short and
receding.

3. Middle and lower whorls
are also quite convex.

4. Last whorl flatly convex,
but little flattened.

5. Peristome distinctly ex-
panded, lipped, whitish; in-
terior light grayish brown.

6. Color of shell usually
corneous brown.

In distribution, according to Ehrmann, avenacea is mainly
western, from the Maritime Alps through the Swiss and
French Alps to the Alpenrhein. C. clienta is mainly east, as
far as the Caucasus, where a new race which is called [Chon-
DRiNA clienta] caucasica Ehrmann occurs (I.e. p. 19, foot-
note 1, pi. 1, fig. 4). It differs from the type by the more
elongate form, less regular striation, and an infrapalatal,
which is often stronger than the supra-palatal (PI. 20, fig. 4).
Type locality Letschghum, Caucasus. Unfortunately the var.
suhcereana West, of the western Crimea could not be
compared.

80 PUPOIDES.

While I do not question the zoological observations in-
volved in this discussion, I may venture to suggest that the
nomenclatorial questions require further examination, Bru-
guiere and Geoffroy, whom he quotes, dealt with shells from
the environs of Paris. It is not at all certain that Draparnaud,
far to the south in Montpellier, had the same form. Paris
specimens, which I do not possess, are required in order to
determine just what the typical P. avenacca is. Also, the
long list of synonyms and varieties must be gone through to
determine whether there is an earlier name for the species
Ehrmann has segregated as Ch. clienta.

PUPOIDES (Vol. 36, p. 108).

The genotype, P. nitidulus, was first designated by Kobelt,
1880: Illustr. Conchylienbuch, p. 267.

Add to synonyms: Themapiipa Iredale, The Victorian
Naturalist vol. 47, Nov. 1930, p. 120, mt. Pupa heltiana Tate.

Mr. Iredale states that " it has no real relationship " with
Piipoides, but he does not attempt to mention any differential
character in the half page treating of "Themapupa" heltiana.
Until such characters are indicated, there seems little reason
to regard Themapiipa seriously.

PuPOiDES MARGINATUS (Say). Vol. 26, p. 111.

A sinistral specimen was found by Dr. Julia Gardner of
the U. S. Geol. Survey in Texas, on the Colorado River, li/^
miles below the Travis-Bastrop Co. line.

PUPOIDES KARACHIENSIS Pcilc. PI. 18, fig. 8.

The shell is subperforate with six very convex whorls, apex
rather flat. Color (evidently faded) whitish buff. Growth
lines faint on spire but a little stronger on back of body whorl.
Aperture sub-circular, labrum expanded but not reflexed.
Parietal denticle weak. Length 4.5 mm. Diameter of pen-
ultimate whorl, above aperture, 1.5 mm. Length of my
smallest specimen, 4.1 mm. ; diameter, 1.4 mm. The shell
somewhat resembles P. hryantwalkeri Pilsbry, but has more
convex whorls and more rounded aperture (Peile).

PUPILLA. 81

India: Karachi (God win- Austen and Peile).

Pupoides karachiensis Peile, Proc. Malac. Soc, London,
vol. 18, p. 139, fig. ; March, 1929.

" Among the undescribed material in the collection of the
late Col. H. H. Godwin-Austen, now in the British Museum,
was found a single specimen labelled ' Buliminus, sp. nov.
Karachi '. The shell agrees with half a dozen specimens in
my own collection which were found in sand from the Karachi
beach in company with P. coenopictus, Hutton, and a multi-
tude of interesting small marine shells. The Museum speci-
men has been selected as the type. ' '

Ptjpoides hoggaeensis (Pallary).

Shell elongate, of very convex whorls with very fine striae,
visible only under a lens, 6I/2 swollen whorls separated by a
slightly oblique suture, of very regular increase; the last
whorl a little descending. Aperture very little oblique, with
the outer margin very little reflected. Aperture oval, pro-
duced to the right, with the outer margins subparallel and
rounded at the lower part. Umbilicus narrow. Height 4.5
to 4.75 mm., greatest diameter 2 mm. (Pallary).

Hoggar: Tihaliouine at 2150 meters; Imegha; Tiniker
(Tifidest) up the Aguelman.

Leiicochiloides hoggarensis Pallary, Bull. Soc. d'Hist. Nat.
de I'Afrique du Nord, vol. 20, 1929, p. 6.

This species differs from L. senaaricus by the more slender
contour, the whorls more convex and especially by its aperture
produced towards the right. The little denticle which appears
at the junction of the outer lip in L. senaaricus is wanting in
our species (Pallary).

Pfeiffer called his species sennaarieiisis, not senaaricus.

PUPILLA (Vol. 26, p. 152).

Mr. B. B. Woodward has called attention to the first pub-
lication of the name Pupilla Leach MS. It was in Fleming:
British Animals, 1828, p. 268. The name occurs in the text
under Pupa marginata Drap. thus: " This species was sent

82 PUPILLA.

me many years ago by Dr. Leach, from Battersea, under the
title of Pupilla marginata."

PUPILLA OERSTEDH (Morch).

Shell umbilicate, cylindrie, obliquely striatulate, glossy,
brown. Whorls 6i/^, convex, with impressed suture, the last
equal to one-fourth the length. Peristome thickened subre-
flected. Aperture triangular, the columella wide, entering,
provided with a punctiform white tooth; parietal tooth sub-
compressed; lips joined by a funicular callus. Length 3.5
mm., diam. 2 mm. {Moerch).

Nicaragua (A. S. Oersted).

Pupa (Pupilla) oerstedii Moerch, Malak. Bl. VI, 1859, p.
111.— Pfr., Monogr. VI, p. 306.

This species comes from further south than any other
American Pupilla, if it belongs to that genus. It is known
by the original account only. Moerch notes that ' ' The outer
lip has two or three notches which give the aperture an
angular appearance. The parietal tooth is somewhat removed
from the curving lip-margin, and equally removed from the
funicular callus. Epidermis is weathered on the oldest whorls.
The habitus and color are quite as in our European species,
such as Pupa dolium, muscorum etc."

Pupilla sterkl\na (Pilsbry). Vol. 26, p. 156.

Add the locality: San Martin Island, Lower California (G.
D. Hanna). A few beautiful albino shells were among many
of the normal brown color {Pilshry, Proe. Cal. Acad. Sci. (4),
vol. 16, 1927, p. 186).

Pupilla goniodon Pilsbry. PL 15, fig. 12.

The shell resembles P. sterkiana of the Lower Californian
mainland in shape and sculpture. It is thin, cylindrie, with
blunt, rounded ends, cinnamon-colored, with sculpture of
strongly retractive, widely spaced riblets, which are more or
less irregular or in places dislocated, sometimes with short,
twig-like branches ; they are about one-fourth as wide as their
intervals or less. The initial 11/2 wliorls have irregularly an-

PUPILLA.

83

astomosing net-like sculpture and pale grayish color. Sub-
sequent whorls are moderately convex, and the last one rises
somewhat and becomes flattened laterally towards the aperture.
The aperture is about as wide as long, somewhat squarish with
rounded angles. The peristome is expanded, the outer and
basal margins thickened with a cinnamon callus within, upper
external margin thin ; columellar margin dilated. There is a
transverse white nodule on the parietal wall within the angle
of the lip, united with the latter, and continued in a thin
callus across to the columellar lip-insertion. Length, 4 mm.;
diam., 1.9 mm. ; 6% whorls.

Guadalupe Island, Lower California, at Northeast Anchor-
age at an elevation not greater than 100 ft., in the canyon
back of the buildings, abundant (G. D. Hanna).

Pupilla goniodon Pils., Proc. Cal. Acad. Sci. (4), XVI,
1927, p. 172, pi. 7, fig. 3.

The presence of a strong angular nodule and the more deli-
cate riblets differentiate this species from P. sterkiana. In ex-
ceptional specimens there is the barely perceptible trace of a
parietal tooth, rather deep within ; in none do I see any colu-
mellar tooth or truncation, such as P. sterkiana usually shows
in an oblique view in the aperture. The species was collected
alive in some numbers.

Pupilla guadalupensis Pilsbry. PL 15, figs. 10, 11.

Shell cylindroid, slowly tapering in the upper half, cinna-
mon colored, only slightly glossy, very evenly sculptured with
retractive riblets paler than the ground color, from a half to a
third as wide as their intervals and about 14 to 16 in one milli-
meter on the face of the last and penult whorls; the usually
paler 1% embryonic whorls have the net-pitted sculpture of
Striopupilla. Subsequent whorls are rather strongly convex.
Aperture rounded below, straightened with rounded angles
above; no trace of a crest or contraction behind outer lip.
Peristome expanded, heavily thickened within except at the
posterior-lateral curve, continuous in a slightly free or adnate
callus across the parietal wall. Aperture typically four-

84 PUPILLA.

toothed : angular lamella in form of a callous pad within the
posterior angle, parietal lamella short, stout, deeply placed,
columellar lamella low and broad, palatal fold a rounded,
deeply placed tubercle. (Other forms of the species may
lack all but the angular pad.)

Length 2.8 mm.; diam. 1.5 mm., 51/2 whorls. (Holotype.)

Length 3.1 mm., diam. 1.5 mm. ; 5% whorls.

Guadalupe Island, Lower California, 1000 ft. above North-
east Anchorage ; also from 2 miles north of south end, on the
east side, near sea level at N-E. Anchorage and on the crest
of Pine Eidge at an elevation of 3000 ft. (G. D. Hanna).

Pupilla guadalupensis Pilbry, Proc. Cal. Acad. Sei. (4),
XVI, 1927, p. 173, pi. 7, f. 1, 2.

The close, regular, evenly developed ribs distinguish this
species from other American Pupillse, the other two species of
the subgenus Striopupilla having the ribs more uneven and
more widely spaced. It resembles the Asiatic Pupilla annan-
dalei, which, however, belongs to a different subgenus.

As in most species of this genus, the parietal, columellar
and especially the palatal teeth are variable, either present or
absent. The angular pad appears in all adult individuals, and
is sometimes the only tooth present, as in fig. 11. The four-
toothed form, selected as typical on account of its status as the
most primitive form, is exceptional in some lots seen, but
common in other lots.

Five specimens taken at random from the type lot have
teeth as follows :

1. Angular, parietal, columellar, palatal.

2. Angular, parietal, columellar, palatal.

3. Angular, parietal,

4. Angular, columeUar,

5. Angular,

In No. 2 the parietal is very small ; in No. 4 the columellar
can be seen only in an oblique view in the aperture.

A similarly unselected lot of ten, from two miles north of
the south end of the island, has teeth as follows:

PUPILLA. 85

1. Angular, parietal, columellar, palatal.

2. Angular, parietal, columellar, palatal.

3. Angular, parietal, columellar, palatal.

4. Angular, columellar, palatal.

5. Angular, parietal, columellar,

6. Angular, parietal, columellar,

7. Angular, parietal, columellar,

8. Angular, parietal, palatal.

9. Angular, parietal,

10. Angular,

PuPiLLA cuPA (Jan). Vol. 26, p. 185.

The type of Vertigo eumicra Bourguiguat (Vol. 26, p. 233)
from near Lucerne, is stated by G. Mermod (Revue Suisse de
Zool. Vol. 33, July, 1926, p. 568) to be a specimen of PupiUa
sterri Voith {=cupa). Vertigo eumicra Clessin is considered
by Mermod to be a toothless V. arctica Wallenb.

On page 188 of Vol. 26 the first two paragraphs should
follow the account of P. cupa turcmenia. They were accident-
ally transposed.

PUPILLA TRIPLICATA (Stud.). Vol. 26, p. 189.

Rosen (Mittheil. Kaukasischen Museums VI, 1914, p. 192,
193) gives Caucasus localities for var. luxurians Reinh, ; var.
inops Reinh. {cylindrata Bttg.), which was placed under
muscorum in Manual Vol. 26, p. 180, following Boettger, prob-
ably a mistake ; and var. s^iiconiformis Bttg., the latter a pen
or typographical error for suhoviformis Bttg., Jahrb. D. M.
Ges. VI, 1879, p. 26 ; Manual Vol. 26, p. 192 (where Vol. VII
of the Jahrbiicher is erroneously cited).

PUPILLA SIGNATA (MOUSS.). Vol. 26, p. 194.

This species has been reported from Alaska by Westerlund
(Vega-Exped. Vetenskap. lakttagelser, IV, 1887, p. 164) and
from the Rocky Mountains by Sterki (Nautilus VI, 1892, p.
3) but both records seem to have been based upon erroneous
identifications.

Pupa diecki Gredler (Vol. 26, p. 196) is thought by Lind-
holm to be identical with signata (Ann. Mus. Zool. Acad. Sci.
Russ., 1922, p. 309).-

86 LAURIA.

PupiLLA PUPULA (Desh.). Vol. 26, p. 214.

Colonel A. J. Peile writes me that he suspects that Desh-
ayes' figures referred to this species were drawn by an artist's
error from Lauria bourboneiisis, not from the specimen
Deshayes described. Deshayes himself noticed that his figures
imperfectly represented his species. My figures in vol. 26,
pi. 17, figs. 12, 13 are the true P. pupula.

PuPiLixiV l'eprevieri Pallary. PI. 18, figs. 10.

Shell turriculate, cylindrie, with slightly convex whorls, the
summit obtuse. T^/o whorls ; suture well marked. Last whorl
noticeably ascending to the insertion. Aperture vertical,
semicircular. Outer margin doubled by a whitish external
crest. Columellar wall ornamented with a very small lamella.
Color varying from ashy brown to uniform deep brown
(Pallary).

Length 3.5 to 4, diam. 1.5 mm.

Morocco: Laghouat, dep. d 'Alger. (Messrs de I'Eprevier).

Pupilla VEprevieri Pallary, Journ. de Conchyl. vol. 71,
March, 1928, p. 245, pi. 3, f. 36-38.

Pallary remarks that this species, common at Laghouat, is
perhaps approached by P. muscorum and u-mMlicata, but
differs by the more numerous whorls in examples of equal
size and the more cylindrie form. It has much the appear-
ance of the Pleistocene form of Orcula of the northern Sahara,
from which it scarcely differs except by being of half the size.

Laurlv zonata (Bttg.). Vol. 27, p. 79.

Boettger's Pupa zonata, 1883, is a homonym of Pupa zonata
Gassies, 1869, and may be changed to Lauria albina Bttg., a
name given to a greenish-white mutation. If the usual banded
form requires a name it may be called forma zonifera.

Laurlv wouramboulchiensis Connolly. PI. 18, fig. 9.

Shell very small, ovate, rimate, rather solid and opaque,
which may be due to weathering, glossy, pale brown, with
darker oblique stripes on the striae. Spire moderately pro-

LAURIA. 87

duced, sides somewhat convex, apex obtusely rounded. Whorls
5, convex, regularly increasing, first li/^ smooth, next II/2
very closely, microscopically, transversely and slightly
obliquely striate in the lines of growth, last 2 sculptured with
regular, flattish, somewhat distant, slightly oblique striae,
darker than the intervals between them, there being about 12
and 13 visible on the 4th and 5th whorls respectively ; suture
simple, well defined. Aperture quadrate, rounded at base,
peristome glossy-white, expanded and reflexed, outer lip
receding only very slightly in profile, columella erect, margin
broadly triangularly reflexed, callus thin, but well marked
and continuous; the only dental process is an inrunning
parietal plait, inclined downwards to the right, three-quarters
of the distance from the columella to the outer lip, but there
is a slight inflation of the inner margin one-third way down
the outer lip and another, inset and hardly noticeable, half-
way up the columella. Long. 3.3, lat. 2.1 ; apert. alt. 1.3, lat.
0.8; last whorl 2.0 mm. (Connolly).

Abyssinia: Serpent Lake, "Wouramboulchi (Omer-Cooper).

Lauria wouramboulchiensis Connoll"s^, Proc. Zool. Soc.
London, 1928, p. 168, pi. 1, f. 6.

' ' The shell is smoother and squatter than L. bruguierei and
comparatively wider, while the dark transverse stripes impart
to it a very distinctive appearance. I find nothing resembling
it closely in the fauna of continental Africa, but from the
respective figures it must be closely allied to L. bourhonicensis
Pilsb. The peristome, however, in the Abyssinian race is
white and somewhat broadly reflexed, instead of brown and
very narrowly so, as emphasized in the case of bourbonicensis,
which, too, should have stronger sculpture. It has not been
proved hitherto that any of the South or Central African
Pupillidae are actually conspecific with those of adjacent
lands, and even if wouramboulchiensis eventually proves to be
merely a variety, it will be entitled to full varietal rank on
account of its remarkable striped coloration,

I have not been able to examine the young stage, but it
appears hardly likely to belong to a genus other than Lauria."

88 ORCULA.

Section Senilaueia Pilsbry

Petrarca Pils., 1922, Man. Conch. Vol. 27, p. 45, 122, not
Petrarca Fowler, 1899 (Cirripedia) — S&nilauria Pils., Nauti-
lus Vol. 41, Jan., 1928, p. 108, tod. Lauria fasciolata (Morelet).

AGARDHIA (Vol. 27, p. 129).
Agardhia macrodonta Hesse. (Vol. 27, p. 156).

Has been figured by Dr. Antoni Wagner in Ann. Zool. Mus.
Polonici Hist. Nat. VI, Dee. 1927, p. 306, pi. 15, figs. 115, 116.

A. m. rumelica Hesse, I.e., p. 307, pi. 15, fig. 117.
OECULA (Vol. 27, p. 1).

Sajphus ' ' Monts. ' ' with the type Pupa doliolum Brug. was
first published, I believe, by Cocconi, 1908, Bol. Mus. Zool.
ed Anat. Comp. R. Univ. Torino vol. 23, No. 583, p. 7, footnote.
Scyphus doliolum var. singularis Monts., n. var., is reported
from southern Italy and Tremiti Island by Cocconi, I.e., with-
out description.

Orcula dolium pseudogularis Wagner. PI. 19, fig, 5.

Vol. 27, p. 11, In comparison with the typical form the
shell appears more slender and cylindrically drawn out, with
weaker gro\\i;h striae, so that the surface is smoother and
glossier. The 9 whorls increase more rapidly and are re-
latively higher; the last ascends only a little to the aperture.
The lip callus is thinner, the upper columellar lamella, though
it is often weaker, yet is always present. Height 8, diam.
3-3.4 mm. {Wagner).

Orcula dolium pseudogidaris A. J. Wagner, Ann. Zool.
Mus. Polonici Hist. Nat. I, 1922, p. 121, pi. 6, fig. 39.

On dry clifs, covered only with shrubs and low plants, at
Gleissenfeld (Tiirkensturz) in the Pittental, and similar places
in this valley. Such slim, cylindric forms of 0. dolium Drap.
are to be found sporadically, together with typical examples,
in other places, especially at high elevations, as on the
Schneeberg by Wiener-Neustadt ; but at the Gleissenfeld

ORCULA. 89

locality in the Pittental no typical examples, or at most tran-
sitions toward such, are to be found, showing this to be a
stable local form by the action of local conditions {Wagner).

Orcula dolium tatrica Wagner. PI, 19, fig. 6.

This is the smallest form of 0. dolium Drap, yet known to
me, showing only 7I/2 whorls, fine growth striae, and in fresh
examples a light silky luster. Moreover, the lip-callus is very
thin, the lamella on the parietal wall very low, and an upper
columellar fold is lacking. Alt. 5, diam. 2 mm. {Wagner,
Ann. Zool. Mus. Polonici Hist. Nat. I, 1922, p. 121, pi. 6,
f. 40).

Collected in the Tatra by Slosarski, without nearer local-
ization ; from the occurrence of conspicuously small specimens
of Pirostoma tumida A. Schm,, P. plicatula Drap. and
Clausilia orthostoma Mke. with the same label, a considerable
elevation may be presumed.

Orcula puchsi Zimmermann. PI. 20, figs. 5, 6, 7.

Shell strikingly slender, gradually tapering from the last
whorl to the apex, finely and rather regularly striate, dark
corneous-brown to reddish brown, somewhat glossy. Umbili-
cus very narrow, almost punctif orm ; neck rounded. The 8I/2
to 9I/2 whorls increase slowly and regularly, are relatively
strongly convex, and separated by a markedly deep suture,
the last whorl forming about one-fifth the length of the shell.
Aperture semiovate, rather vertical. Columellar margin
deviating little from the longitudinal axis of the shell ; outer
margin nearly parallel to it, only a little produced up on the
penult whorl. Peristome distinctly expanded, not thickened;
whitish; a palatal callus, showing through outside, is always
present; its development is irregular, but in the majority of
specimens it is well developed though not so strong as in
0. spoliata Rossm. ; as in this species, advancing towards the
interior and thus narrowing the cavity in a vertical view from
below ; there is no fold-like projection inward. The columellar
margin is straight and without any indicatio^i of a fold;
further inward also it is smooth as seen in a broken shell.

90 ORCULA.

The parietal lamella stands midway between columellar and
outer lip insertions, and is low and short, ending immediately
beliind the aperture in the last third of the last whorl.

Length 6.6 mm., width 2.3 mm. {Zimmermann) .

Austria : at the foot of the so-called ' ' Turmauer, ' ' northern
spur of the Goeller, at an elevation of about 750 meters, in a
botanieally well-known subalpine enclave, which here extends
from the Goeller nearly into the valley (Arnold Penther;
Anton Fuchs). Later found near the first locality on the
** Waldhuttsattel ", on the road from Kernhof, at the upper
limit of the subalpine forest at about 1200 meters.

Orcula fuchsi Zimmermann, Archiv f. Molluskenkunde, vol.
63, p. 44, pi. 4, f. 3-5. Jan. 1, 1931.

At the higher locality it is smaller, 6.3 x 2.2 mm. (PI. 20,
fig. 7). In both places it lives with 0. dolium and the lower
Austrian race of 0. spoliata, but without intergradation.
Figures from Zimmermann.

Orcula gularis pseudodolium Wagner. PL 19, figs. 3, 4.

Vol. 27, p. 14. The shell is larger and wider than the
typical form, and tapers in an acute cone above; in habitus
recalling 0. conica Rm. and 0. dolium Drap. As a form of
0. gularis Rssm. it is characterized by the features of the
aperture. Of the two columellar lamellae, here the upper is
also well developed and emerging (weak in the typical form).
The palatal callus is strongly developed as in 0. gularis
spoliata, yet with the pliciform appendages characteristic of
the type. The peristome is, like that of 0. dolium Drap.,
more broadened and expanded. Alt. 7, diam. 3 to 3.5 mm.

Feuchtenauer Aim on the Hochengsengebirge at Wind-
ischgarsten. Upper Austria (Wagner, Ann. Zool. Mus. Polonici
Hist. Nat. 1, 1922, p. 122, pi. 6, f. 37 a, h.

Orcula gularis tolminensis Wagner. PL 19, figs. 7, 8.

The shell is smaller throughout than the typical form, with
only 7 to 9 whorls, thin lip-callus and conspicuously lower,
often obsolete columellar folds, which are not visible in a

ORCULA. 91

direct view in the aperture. The strong palatal callus has
excessively developed, very long appendages, which extend as
far as over the umbilical slit in the throat. Alt. 5.5 mm.,
diam. 2.3 mm.

Neighborhood of the waterfall of Perichnik at Tolmein,
Isonzo Valley.

Orcula gularis tolminensis A. J. Wagner, Ann. Zool. Mus.
Polonici Hist. Nat. I, 1922, p. 122, pi. 6, f. 38 a, h.

The excessively long palatal folds in an otherwise weakly
developed apertural formation appear remarkable.

Orcula kaznakovi (Rosen). PI. 19, figs. 1, 2.

Shell with a curved umbilical crevice; ovate-cylindric ;
smoothish, sculptured with remote, obsolete lamellee ; yellowish,
subpellucid. Apex obtuse. Whorls 91/0, slowly increasing,
the upper 5 a little convex, the rest flat, the last whorl rather
rapidly ascending to the aperture, constricted. Aperture
truncate-oval, with a compressed, very high and deeply enter-
ing parietal lamella, columella within biplicate above. Peris-
tome somewhat dilated, with a white thickening, the margins
converging, joined by a lamelliform white callus. Length 8,
diam. 5% mm., aperture 2i/^ mm. high and wide (Rosen).

Caucasus: Ardanuch, with Buliniimis fiorenskii.

Pupa kaznakovi Rosen, Mittheilungen des Kankasischen
Museums VI, 1914, p. 190, pi. 2, figs. 11 a, h.

It is distinguished from Pupa doliolum Brug, var, hatu-
mensis Ret., which stands nearest the species under considera-
tion, by the shape of the aperture, the conspicuously strong
spiral lamella, the columellar teeth, dimensions, shape of the
shell and the strong callus. I allow myself the pleasure of
giving it the name of the Director of the Caucasian Museum
(Rosen).

Orcula tingitana Pallary. PI. 19, fig. 9.

Vol. 27, p. 256. Pallary 's figures (Journ. de Conchyl., Vol.
65, 1920, p. 133, pi. 2, fig. 15) are copied. The original locality
was banks of the Oued Querrha (Lt. Brunot). Said to be

92 ORCULA.

the only living Orcula of north Africa, where however there
are two fossil species, Pupa ectina Bgt. (Vol. 26, p. 225) and
P. amhUja Bgt. (Vol. 27, p. 4).

Orcula turcica (Let.). Vol. 27, p. 23.

Orcula scyphus graecus Pils,, Vol. 27, p. 36, appears to be
practically identical with 0. turcica Let., which is no doubt
the inadequately defined Pupa critica Zel. of Pfeiffer, collected
on Syra by Zelebor. In the Hesse collection I find specimens
from the Piraeus, Lykabettos near Athens, Syra and Tinos
Islands.

Orcula wagneri ljubetenensis Sturany.

By a typographical error the name was spelled " Ijube-
tensis " in Vol. 27, p. 257.

Subgenus Orculella Steenberg.

Orculella Steenb. Videnskabelige Meddelelser fra Dansk.
naturh. For, i Kobenhavn, vol. 80, 1925, p. 176, tod. Orcula
orientalis Pfr.

This subgenus apparently includes the species No. 9 to No.
13 of the monograph in Manual vol. 27, pp. 27-36.

Not much can be done towards stabilizing the nomenclature
of the Orculellae until the internal lamellae of 0. sirianocori-
ensis of Cyprus, 0. orientalis, type in Pfeiffer 's collection, and
0. scyphus from the type locality, Brusa, are examined, as
these are the early names. The internal characters are also
unknown in part of the named local forms.

Orcula orientalis (Pfr.). Vol. 27, p. 29.

The form from Lebanon, which Westerlund called var.
cedrorum, has a quite weak lower-palatal fold half a whorl
inward. The inner end of the parietal lamella flares out-
ward, and there is a low cord-like lamella near and outside
of the parietal lamella near its inner end.

The form common in the Sarus river debris at Adana,
Cilicia, has no trace of a lower-palatal fold ; otherwise similar
to the preceding, the inner end of the parietal lamella flaring

VERTIGO. 93

outward and accompanied by a low callous cord. Size
variable.

Length 11.3 diam., 4.6 mm. ; 10 whoris.

Len^h 8.5 diam., 4 mm. ; 9i/^ whoris.

Var. antiochensis, n. v. A broad form from Antioch
(Hesse coll., from Berlier) which Hesse has identified as var.
nitida Mouss. (Archiv f. Molluskenkunde, vol. 59, 1927, p.
177) has no lower-palatal fold within. In two of the three
examples there are two cord-like lamellas between the parietal
lamella and the suture (pi. 19, figs. 10, 10a). In the third,
only one lamella is there (PI. 19, fig. 11).

Length 9.8, diam. 4.6 mm. (Fig. 10, 10a, type).

Length 10.5, diam. 5.0 mm.

Length 9.5, diam. 4.8 mm. (Fig. 11).

Whether this is really var. nitida of Mousson is not material,
since that name cannot be used, being a homonym.

0. ORiENTALis MOussoNi Reinh. PI. 19, figs. 12, 13, 13a.

Vol. 27, p. 36. In this form there is a very small lower-
palatal fold, which is visible externally as a short white mark-
ing above the umbilical rima in fresh shells (fig. 13a), and
internally appears as a very low callous fold (fig. 12). The
specimens figured are from Aleppo, Hesse coll., and were
recorded as Orculella orientalis in Archiv f. Molluskenkunde,
vol. 59, p. 176, 1927.

Vertigininae (Vol. 25, p. 68).

Vertiginidae Stimpson, Shells of New England, 1851, p. 53,
including Vertigo and Columella simplex (Gld.).

VERTIGO (Vol. 25, p. 69).
American Species.
Vertigo elatior (Sterki). Vol. 25, p. 95.

I am inclined to rank this as a distinct species, differing
from V. ventricosa by the constantly longer spire. The follow-
ing appears to be a synonym.

Vertigo gouldii loessensis F. C. Baker (PI. 15, fig. 2) was

94 VERTIGO.

thus described: " Shell differing from recent gouldii in being
more ventricose, especially on the last whorl, having the apical
whorls wider and more obtuse, the outer lip more auricled
causing the aperture to have its longest diameter more
diagonal than in gouldii; there are five denticles placed as in
gouldii, but more delicate than in the typical form ; the palatal
denticles are long and subequal.

" Length 2.00; diameter 1.3 mm. Holotype.

" Length 2,00; diameter 1.3 mm. Paratype.

" Length 2.1; diameter 1.1 mm. Paratype." (Baker).

Illinois: 31/2 miles east and 1% miles south of Lewistown,
Fulton County (Liverpool township), in loess of Peorian age.
Types: Museum Natural History, University of Illinois, No.
P2366. Paratypes, Acad. Nat. Sci. Phila., No. 145107.

Vertigo gouldii loessensis F. C. Baker, Nautilus, Vol. 41,
April, 1928, p. 135 ; Trans. Illinois State Acad. Sci. Vol. 21,
1929, p. 297 (Aftonian to Early Wisconsin loess).

" This Vertigo has been listed as goiddii but is different
from the typical species as living today. It has also been
listed as ventricosa, and some small, wide specimens do re-
semble this species. Loessensis strongly resembles Vertigo
elatior Sterki, and Sterki has referred certain forms found in
loess at New Harmony, Indiana to this species. They lack
the strong palatal callus so characteristic of elatior and are
the same as the Illinois variety here differentiated. It is
probable that the Vertigo listed as goiddii by Hanna (Kansas
Science Bull. VII, p. 120, pi. 18, fig. 4) is also this variety.
It is found in the Pleistocene from Yarmouth to Early Wis-
consin time. The form is here considered a marked variety of
gouldii but it might be advisable to consider it a distinct
species." {Baker).

A paratype of loessensis is figured. It resembles V. elatior
in having a longer spire than V. vemtricossa. The palatal
folds of elatior stand on a more or less thick but variable
callus. It is very thin in some Maine examples, heavier in
those from Ohio. This palatal callus is distinctly developed
in paratypes of loessensis, which does not appear separable
from elatior.

VERTIGO. 95

In V. gouldii the striation, especially of the penult whorl,
is much stronger, and the basal tooth is situated further to the
left.

Vertigo gouldii paradoxa Sterki. PI. 15, fig. 5.

Vol. 25, p. 99. The specimens from Newfoundland referred
to V. coloradensis and V. coloradensis hasidens by Vanatta
(Nautilus Vol. 40, p. 113, 1927, and 43, p. 134, 1930, seem to
me to be V. g. paradoxa. None of them has a basal fold.
The lower-palatal fold stands distinctly deeper than its fellow.
In V. gouldii and V. coloradensis it is not quite so deeply
immersed. The distinction is admitted to be rather finely
drawn, but it appears to be valid. A Newfoundland specimen
from Hannah's Head, Humber River, is figured, PI. 15, fig. 5.

On going over the series I conclude that the Rocky Mountain
shells, V. coloradensis and its subspecies, are not specifically
distinct from gouldii. The differences are trivial, and while
absolute continuity of the goiddii and coloradensis areas is
not proven, and seems improbable, there is no great gap
between the ranges as recorded in Manual Vol. 25, pp. 98,
115-118. I would therefore write Y. gouldii coloradensis, V.
g. arizonensis, etc.

Vertigo gouldii paradoxa appears to be generally spread in
Newfoundland, the specimens being from Tucker's Head, east
arm of Bonne Bay; Hannah's Head, Humber River, Bay of
Islands; Ha Ha Cape, Ha Ha Bay; Bard Harbor Hill, high-
lands of St. John, on the Straits of Belle Isle. I have seen
specimens of V. g. paradoxa from Anticosti Island also, in
Museum of Comparative Zoology.

Junius Henderson (Nautilus Vol. 44, p. 9, 10) has raised
the question whether the occurrence of V. coloradensis in New-
foundland " really means a survival of the species at the
extreme eastern end of its pre-glacial range ". This ques-
tion is hardly affected by the change in name of the New-
foundland specimens, since the Rocky Mountain and north-
eastern races, though not absolutely identical, seem to be
barely distinguishable.

96 VERTIGO.

Vertigo wheeleri Pilsbry. PL 15, fig. 1.

The shell is rather obesely oval, the diameter about two-
thirds of the length, cinnamon colored. The first 11/2 whorls
are pale and smooth; the rest are closely and finely striate,
the striation rather strong, about as in V. coloradensis ; it is
somewhat coarser on the penult than on the last whorl. The
whorls are strongly convex, the last near the outer lip becom-
ing a litlte flattened peripherally, and having a weak, wide
swelling or crest behind the outer and basal lips. The rather
small aperture is broadly pear-shaped, obstructed by five
teeth : the angular lamella is tuberculiform, the parietal
lamella higher and rather long; columellar lamella hori-
zontally entering, but rather short; the two palatal folds are
short. The lip is slightly expanded, the outer margin notice-
ably straightened in the middle, and in a profile view seen to
be weakly arched forward there.

Length 1.6, diam. 1.05 mm. ; 4i/^ whorls.

Alabama: Monte Sano, near Huntsville, collected by H. E.
Wlieeler.

Vertigo wheeleri Pilsbry, in Walker's Terrestrial Shell-
bearing Mollusca of Alabama, Univ. Michigan Misc. Pub. No.
18, 1928, p. 146, fig. 224. — Vertigo concinnula Wheeler,
Nautilus XXV, 1912, p. 124, c/. Pilsbry, Man. Conch. XXV,
p. 121.

This species has the broad, ventricose figure of V. hehardi
Van., but is a larger, less fragile and more strongly striate
shell, the columellar lamella differing in form. It is decidedly
more ventricose than V. goiddii which, with five teeth as in
wheeleri, has generally a second columellar and but one tooth
on the parietal wall; however these two teeth are variable in
V. goiddii; the main distinction is in the shape of the shell.
These two species appear to be the nearest relatives of the new
form. V. rugosula and oralis have much more strongly de-
veloped teeth and an outer lip of different shape.

Specimens presumably those now under consideration were
shown me by Mr. H. E. Wheeler many years ago and were
identified as V. concinnula Ckll., a Rocky Mountain species

STROBILOPSID^

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VERTIGO. 97

(Nautilus XXV, 1912, p. 124). That is a larger shell, less
inflated, with longer palatal folds, but very similar to the
present form in sculpture.

Vertigo columbiana Sterki. Vol. 25, p. 109, 18th line.
Eeference to figure of type of V. c. utahensis should stand:
PL 12, fig. 13.

Vertigo californica catalinaria (Sterki). PI. 15, fig. 8.

The specimens from Guadalupe Island, off Lower Cali-
fornia: at Northeast Anchorage and 1000 ft. above, and 2
miles north of the south end on the east side (G. D. Hanna),
are darker than the Santa Catalina Island form, but no other
difference was detected. There is some variation in diameter
among them, also in the spacing of the ribs, the specimen
figured having them more widely spaced than some others.

Length 1.95, diam. 1 mm.

Vertigo californica catalinaria (Sterki), Pilsbry, Proc. Cal.
Acad. Sci. (4), XVI, 1927, p. 174, pi. 7, f. 5.

Vertigo californica guadalupensis Pilsbry. PI. 15, fig. 7.

The differential characters of this form are that it possesses
a distinctly developed angular lamella standing about midway
of the length of the long parietal lamella, and the striation is
quite fine, there being about 30 striae in 1 mm. on the face
of the last whorl. Length 2 mm. ; diam. 1.1 mm.

Guadalupe Island, Lower California; collected about 1000
feet above the landing at Northeast Anchorage by G. D.
Hanna.

Vertigo californica guadalupensis Pilsbry, Proc. Cal. Acad.
Sci. (4), XVI, 1927, p. 175, pi. 7, fig. 4.

In most groups of Vertigo the presence or absence of an
angular lamella has little significance, but in the V. californica
group I have never seen this lamella in hundreds of shells
examined. However, Dr. V. Sterki has mentioned seeing
*' traces " of an angular lamella in some specimens of V.
californica, which is a larger, more coarsely sculptured form
than V. c. guadalupensis. I have thought it best to name this

98 VERTIGO.

island form in order to stimulate the attention of future
collectors. It was found associated with V. c. catalinaria.

Vertigo degeneris Pilsbry. PI. 15, fig. 9.

The oblong, brown shell tapers from the last whorl to the
obtuse apex. The whorls are strongly convex, almost smooth,
but on the penult whorl some spaced wrinkles along growth
lines are noticeable; though weak, they evidently correspond
to the riblets of V. calif ornica. The rounded aperture has no
teeth. Columella concave, the columellar lip being rather
broadly expanded. Length 1.9 mm., diam. 1.1 mm., 41/2
whorls.

Guadalupe Island, Lower California: collected about 1000
feet above the landing at Northeast Anchorage by G. D.
Hanna.

Vertigo degeneris Pils., Proc. Cal. Acad. Sci. (4) XVI,
1927, p. 175, pi. 7, fig. 6.

A strongly characterized form, which, however, may prove
to be connected with a similarly toothless form found at North-
east Anchorage in which the riblets are well developed, length
1.8 mm., diam. 1.1 mm. As only broken examples of this
ribbed form were taken, its status is left in suspense for the
present.

Vertigo torrei Aguayo & Jaume. PL 24, fig. 1.

The shell is rimate, very small, ovate-conic, corneous, trans-
lucent, with a waxy luster, finely striate axially, the striation
more distinct on early whorls and obsolete on the last. Suture
deej). Whorls 4iA, convex, the last well rounded ; apex obtuse.
Aperture transverse, semicircular, provided with seven ivory
white teeth in all specimens examined, as follows: a small
angular lamella ; a high and thick parietal lamella, about twice
as large as the angular and slightly oblique ; a large columellar
lamella which is slightly curved toward the apex of the shell ;
a basal lamella, well developed but more compressed, and
three-fourths the length of the preceding ; a lower-palatal fold,
high, deep, and curving towards the columella; an upper-
palatal fold, parallel to the preceding and of about the same

VERTIGO. 99

size ; and a siiprapalatal tooth, well developed in all specimens
examined. Peristome ferruginous, expanded, very broad in
the eolumellar margin, the outer margin with a deep inflection
in the middle, and with three impressions behind, of which the
shortest is above, the longest below. Length 1.75 mm., diam.
1.05 mm., aperture 0.6x0.73 mm. {Aguayo & Jaiime).

Cuba: "La Lisa", Havana, under dead leaves in damp,
shady places, collected by M. L. Jaume and A. Bierig. Holo-
type in Aguayo collection, paratypes in collections of Jaume,
the Academy of Natural Sciences of Philadelphia, Mus. Com-
parative Zoology, Harvard, and U. S. National Museum.

Vertigo torrei Aguayo & Jaume, Univ. de la Habana
Memorias de la Sociedad Poey, Vol. 8, no. 1, p. 11, text-fig.,
March 10, 1934.

Aguayo and Jaume add that this species resembles V. ovata
Say in general appearance, but differs by the constantly
smaller size, the more expanded eolumellar margin of the
peristome, as well as by the number and form of the apertural
teeth. In V. ovata an infraparietal lamella is often found,
which appears in none of the examples of V. torrei observed.
In the latter a suprapalatal tooth is always present, this being
infrequent in V. ovata. V. neglecta Poey resembles our torrei
in size and proportions, but lacks parietal teeth, yet it is pos-
sible that it was described from an imperfect example which
had lost these teeth by wear. We were unable to find the type
of V. neglecta in the Academia de Ciencias de la Habana.
Probably it is lost. At present it appears that we have to con-
sider V. neglecta as unindentifiable.

Vertigo gouldii hubrichti, n. subsp. PL 22, figs. 12, 13, 14.

The shell is subcylindric, larger than V. nylanderi Sterki,
with a similar long and deep impression over the palatal folds.
The lower-palatal is deeply immersed. There is no angular
lamella. The basal fold is well developed. The intermediate
whorls are strongly, sharply striate as in other forms of
y. gouldii. It differs from V. g. paradoxa by the deep ex-
ternal impression over the palatal folds.

100 VERTIGO.

Length 2 mm., diam. 0.9 mm. ; 5 to 5y2 whorls.
Loess of Mona, St. Louis Co., Missouri. Type and para-
types 160362 A. N. S. P., collected by Mr. Leslie Hubrieht.

Vertigo iDxVhoensis, n. sp. PL 22, figs. 9, 10, 11.

The shell is ovate, rimate, with eonvexly conic spire and
obtuse summit ; cinnamon-brown, the spire paler ; glossy, with
extremely weak and rather sparse lines of growth and a faint
microscopic granulation. The whorls are strongly convex,
the last having an impression at the lip-point, a very low,
hardly noticeable crest, and behind it a deep pit over the
lower-palatal fold. The peristome is distinctly bent in above
the middle of the outer margin, well expanded below the point.
Columellar margin reflected. There are four teeth : a well
developed parietal lamella, a somewhat oblique columellar
lamella, and two palatal folds rather near together, as in
y. ventricosa and its allies, the lower-palatal being decidedly
larger and a little further in. There is no palatal callus.

Length 2 mm., diam. 1.2 mm. ; 4i/^ whorls.

Meadows, Adams Co., Idaho, along a creek east and north-
east of the old town. Type and paratypes 158670 A. N. S. P.,
collected by H. Burrington Baker, Sept. 12-15, 1929.

The whorls are more convex and the suture deeper than in
y. ventricosa, V. columhiana or V. andrusiana, and it differs
from all of these by the much deeper external pit over the
lower-palatal fold, the other species mentioned having the
whorl only somewhat flattened there. The teeth are much the
same in this whole ventricosa series, the palatal folds being
nearer together than in the V. modesta series. The develop-
ment of angular lamella and basal denticle is variable in some
forms of the ventricosa group, but neither is present in speci-
mens of V. idahocnsis seen.

Vertigo modesta sculptilis, new mutation. PL 24, figs. 2, 3, 4.

The shell has rather sharp sculpture on the penult and ante-
penult whorls of the spire, as in V. m. insculpta, but it differs
by the reduced size of the teeth ; small parietal, columellar and
lower-palatal being present in the specimen taken as type ; in

VERTIGO. 101

others there may be minute traces of the upper-palatal fold
and an angular lamella, but still other examples have only
two teeth, no palatal folds being developed. Type 162884
A. N. S. P., from 2 to 10 miles up Rock Creek, northeast of
Garrison, Montana, collected by H. Burrington Baker. It was
taken also near the head of Wallowa Lake, Wallowa Co.,
Oregon, and at Aneroid Lake, about 8 miles south.

Vertigo milium Gld., var. Plate 21, figs. 16, 17.

Vol. 25, p. 146. Specimens from Rio Guayubin at crossing
of Sabaneta Road, northwestern Santo Domingo, are figured
to show the slightly different race of that island. The shell
differs from V. milium by the decidedly shorter lower-palatal
fold, which is wholly visible in a direct front view (but in
milium longer, the end passing out of view), and by the heavy,
calloused columellar lamella. In milium the inner end turns
downward, but in antillarum it is brought forward almost as
far as the upper end, the space between the two ends being
largely filled with callus. The shell is of a compact, oval
shape, finely striate, especially the penult whorl. The speci-;
mens figured are No. 160406 A. N. S. P., collected by A. A.
Olsson.

Chinese Species.

Vertigo teilhardi Ping and Yen. Plate 24, figs. 7, 8, 9.

"Shell minute, rimate, ovoid. Apex very small, slightly
higher than first whorl. Whorls about 4i/^, increasing gradual
in first two and gradually in last two. Body whorl more than
% of the total length of the shell. Surfaces of the first two
whorls moderately convex, while that of the last two more
convex, body whorl with its base much reduced. Apex and
first three whorls look free from striae under ordinary magni-
fying lens, and body whorl with extremely fine striae, which
become a wrinkle or crease close to the aperture. Sutures well
developed, distinct owing to convex surfaces of whorls. Aper-
ture somewhat cardiform, if the inwardly bent outer lip be
taken as its base. It bears 6 teeth within : the parietal tooth
slightly larger than the angular which is small but not obsolete.

102 VERTIGO.

The coliimellar about as large as parietal. The basal rather
small and quite near the posterior end of columella. The two
palatal teeth or lamellae well developed which are not smaller
than either parietal or columellar, although they are of com-
paratively slender ridges. The upper one of the palatal teeth
situated on the inner side of the concavity of the outer lip
looking slightly thicker than the lower one. Outer and inner
lips slightly thick, simple and smooth, with a little tendency
of being expanded. Umbilicus very narrow.

''The apex and second whorl are grayish white, the rest of
the shell is rufous brown, and the aperture, apertural teeth
and the peristomal region pale or whitish. When the shells
are comparative worn, the surface is almost uniformly grayish
white" (Ping and Yew).

Alt. 2.2 mm., diam. 1.2 mm., length of aperture 0.5 mm.,
width of aperture 0.75 mm.

N. W. China : Aksu, in a dry river bed. Type No. 545 Fan
Mem. Inst. Biol.

Vertigo teilhardi Ping and Yen, Bull. Fan Memorial In-
stitute of Biology, vol. 4, p. 277, fig. 14; March 15, 1933.

" This species resembles in some respects Vertigo eogea
Pilsbry, 1919, described from Kashima, Harima of central
Japan. The present species is, however, differentiated from
the latter by its less broad outline, its palatal and basal teeth
separated from each other without ' a strong buff callus', and
its slightly larger size, with but one volution less" {Ping
and Ye7i).

European Species.

Vertigo pusilla godetiana Piaget.

Shell normal except that the peristome is thinner, subcon-
tinuous, the right margin not arched in, the parietal wall one-
toothed. Length 1.5 mm., diam. 1 mm. (Piaget).

Switzerland: Pierrabot (Godet) ; Quaternary sandy alluvial
deposit in front of Champion (Gampelen), near the road to
Infini.

VERTIGO. 103

[Vertigo pusilla] var. godetiana J. Piaget, Mittheil. Naturf.
Ges. Bern, Jahrg. 1913 (1914), p. 156.

Vertigo lilljeborgi West. (Vol. 25, p. 167; vol. 27, p. 218).

According to Esmark (Skrifter Norske Videnskaps-Akad. i
Oslo, 1925, p. 50), Vertigo pachygaster Jensen (Vol. 25, p.
182) is only a light-colored shell of V. lilljchorgi, deviating a
little in form.

Vertigo moulinsiana (Dup.) Vol. 25, p. 178.

Pupa desmoulinsiana JeflEreys, 1855 (Vol. 25, p. 179, line
19) and Vertigo moulinsiana Jeffr., Brit. Conch. 1, 1862, p.
255 ; V, pi. 15, f . 6, are referable to Vertigo lilljehorgi accord-
ing to Kennard and Woodward, Syn. Brit. Non-Marine Moll.,
1926, p. 123.

On p. 179, 23d line, the reference to the synonym. '' Pupa
laevigata ", is erroneous. The paragraph should read:
Vertigo laevigata Kokeil, von Gallenstein, Karntens Land-
und Siisswasser-Conchylien, in Jahrb. naturhist. Landes-
museums von Kamten, 1852, p. 80. — Pupa laevigata Kokeil,
Pfeiffer, Monographia IV, 1859, p. 681.

Vertigo arctica Wallenb. Vol. 25, p. 189.

G. Mermod (Revue Suisse de Zoologie, Vol. 33, July, 1926,
p. 568-571) examined the type lot of Vertigo " eumicra
Bgt." of Clessin (Vol. 26, p. 233), one of which he figured
(t.c. p. 570, fig. 7, shell, p. 571, fig. 8, radula). It is a tooth-
less form of V. arctica, in his opinion.

Vertigo zschokkei Butikofer (Vol. 27, p. 219) was also ex-
amined and submitted to other authorities. The large form
of V. zschokkei is considered a bidentate form of V. arctica
(Mermod, t. c, p. 571-576, figs. 9). His figures are copied in
my PI. 18, figs. 11, 12. The smaU form of '' F. zscJwkkei "
is said to be identical with V. parcedentata Al. Braun.

Vertigo substriata Jeffr. Vol. 25, p. 172.

The occurrence of this species in the Netherlands, and its
Btatus as a " glacial relict " as supposed by some authors, is

104 VERTIGO.

discussed by Dr. A. L. J. Sunier in Zoologische Mededeelingen,
Rijks Museum van Nat. Hist., Leiden, IX, 113-178.

Vertigo genesh Gredler. PI. 18, figs. 13-16.

The characters and distribution have been discussed by A.
W. Stelfox and R. A. Phillips in Journ. of Conch., Vol. 17,
July, 1925, p. 236-240, with figures of a topotype from Salten
(reproduced in my PI. 18, fig. 14), and of Irish specimens
(reproduced in PI. 18, fig. 13, Fancraft marl bed; fig. 15,
marl at Golden Grove; fig. 16, marshy pasture near Mount-
mellick. Queen's Co., Ireland).

V. parcedentata Al. Braun, received from Geyer, is said by
Stelfox to be " more cylindric and alpestris-shaited " than
the Irish shells.

See also this Manual, Vol. 27, p. 216, and Geyer, Arcliiv fiir
Molluskenkunde, Vol. 57, 1925, p. 99-102, where Sandberger's
figures of parcedentata are discussed, and a grouping of
European dextral Vertigo species is proposed.

Vertigo genesii geyeri Lindholm. PL 18, fig. 17.

Shell dextral, minute, rimate, ovate, obtuse, deep brown or
chestnut, lightly striatulate, glossy. Whorls 4I/2, convex, the
first three slowly, last two rapidly increasing and somewhat
swollen, separated by a deep suture which ascends slightly in
front. Aperture semioval with the margins nearly straight,
the right regularly arcuate or slightly subconstricted, the
columellar vertical, dilated and reflected above ; four-toothed,
one remote, subcompressed strong parietal, one conic, strong,
lamella in the middle of the columella, two unconnected
palatals, the larger nodiform or pliciform below the middle of
the margin, a very minute denticle above the middle ; palatal
and cervical calluses wanting. Alt. 2 mm., diam. 1 mm.
{Lindholm).

WUrttembergischen Allgau; Tirol; Poland and European
Russia. Tirol: Salten (D. Geyer). U.S.S.R. : Wlachernskaja,
Gouv. Moskau; Ishora R. debris, Gouv. St. Petersburg (Lind-
holm) ; shore meadow of Tseheremenstzkoje Lake, Gouv. St.
Petersburg (I. K. Wislouch). Fossil in peat-bog of Pfohren
near Donaueschingen (D. Geyer).

TRUNCATELLINA. 105

Vertigo genesii Gredler subsp. gei/eri Lindholm, Arehiv fiir
MoUuskenkunde, LVII, 1925, p. 241-251, fig.

The passages quoted from Geyer in Manual Vol. 25, p. 205,
evidently refer to V. g. geyeri. According to Lindholm, in all
three Russian localities this form occurs without mixture with
the typical toothless V. genesii. Grown specimens differ from
V. antivertigo and V. lilljehorgi (with which they agree well
in luster, sculpture and color) by the entire absence of the
cervical crest. In V. pygmaea var. callocarens the two palatal
folds are united by a palatal callus, not developed in V. g.
geyeri.

TRUNCATELLINA (Vol. 26, p. 58).

Vol. 26, p. 63, third line: for '' Truncatella " read Trun-
catellina.

TRUNCATELLINA CYLINDRICA (Fcr.) (Vol. 26, p. 65).

Odhner states that the types of Pupa odontostoma Wester-
lund (Vol. 26, p. 78) in the Museum at Gothenburg are in all
respects identical with T. cylindrica. The sculpture is of
similar riblets, not striae as stated in the description, and the
aperture shows no trace of teeth. It is not even a variety.
Esmark agrees with this synonymy, (c/. Arkiv f. Zoologi, vol.
16, No. 29, 1924, p. 4; Skrifter Norske Videnskaps-Akad. i
Oslo, 1925, p. 118).

TRUNCATELLINA CYLINDRICA COSTIGERELLA Lindholm.

Differs from the type by the costulate (not striate) shell, as
in T. costidata Nilss. Size, color and aperture as in the type.

Crimea: widely distributed, for instance at Kisil-Koba;
Nishnjaja Massandra; Merdwen, and other places.

Truncatellina cylindrica (Fer.) var. costigerella Lindholm,
Archiv f. MoUuskenk. LVIII, 1926, pp. 166, 176.

" Together with specimens of T. cylindrica Fer. typical in
sculpture, size and coloring, a second form occurs in the
Crimea, not specifically separable from T. cylindrica, but dis-
tinguished distinctly from German, French and Russian ex-

106 TRUNCATELLINA.

amples in ray collection by the ribbing being double as coarse.
In this it approaches T. costulata Nilss., but naturally has no
apertural armature whatever, and a thin peristome, in the
full grown stage.

" The var. costigerella predominates alone in a few places,
in others being mixed with typical specimens, and connected
therewith by transitions which stand about midway in sculp-
ture " (Lindholm).

Truncatellina tauricola Lindholm.

Shell rimate, rather small, cylindric, slightly attenuate near
the apex, under the lens seen to be very delicately striate ;
deep reddish brown, at the apex ashy. "Whorls 6, a little
convex, slowly and regularly increasing, separated by an
impressed suture, the last whorl slightly ascending. Aperture
oblong-subcircular, toothless, the peristome a little reflected,
thin.

Length 2 mm., width 0.8 mm. (Lindholm) .

Crimea: Tschatyr Dagh (Lindholm).

Truncatellina tauricola Lindh., Archiv f. Molluskenk.
LVIII, 1926, p. 177.

This form, of which two grown, living specimens were taken,
differs from T. cylindrica by the dark reddish-brown color and
the extremely fine striation, so that it does not fall within the
hitherto known variation amplitude of T. cylindrica. By the
two characters mentioned it recalls T. monodon Held {T.
striata Grdl.). But a direct comparison of Tyrolese specimens
of this species with the Crimean snail shows that the latter
has conspicuously more convex whorls and a more deeply
impressed suture than in T. monodon, which moreover is one
of the toothed truncatellinas (Lindholm) .

Truncatellina sundleri Odhner. PL 21, figs. 1-3.

Shell cylindric (with the contours of the right and left
sides parallel from the fourth to the last whorl), roundly
truncate above, glossy, reddish horn-brown, striate, very
narrowly umbilicate. Whorls 6, well rounded, with deep.

TRUNCATELLINA. 107

simple suture. Protoconch (II/2 upper whorls) smooth,
whitish, the following two whorls rapidly increasing in width,
horn-brown, rather coarsely, irregularly and very obliquely
striate; the three following whorls of equal width, colored
and striate like those above, progressively increasing in height,
the lower always higher than the preceding; last whorl li/o
times as high as the penult. On the lowest whorls are in-
distinct traces of a microscopic spiral striation. Suture
ascends a little to the aperture. The aperture is small,
rounded triangular, with simple, not thickened lip. Colum-
ellar margin gradually widening above, reflected, white.
Parietal wall with a thin white callus. On the parietal wall is
a projecting, short (not entering) lamelliform white tooth;
on the straight columella a longitudinal callus within, form-
ing a little brownish eminence, under it a distinct sinus.

Height 2.25 mm., breadth 1 mm. (Odhner).

Sweden: Schonen, near Krageholm, on stone wall at the
south end of the garden, under moss (A. Nilsson).

Truncatellina sundleri Hj. Odhner, Arkiv for Zoologi,
XVI, No. 29, p. 1, figs. 1-5. 1924.

Up to this time only two examples of this apparently
endemic species are at hand, which moreover are of somewhat
different forms. One is a little higher, though with the aper-
ture not fully developed, as it has no trace of the parietal
tooth and also a weaker columella callus. In order to clear
up the affinities of the new species the radulae of T. monodon,
T. cosiulata and T. cylindrica are repeated here. It appears
that the differences between T. sundleri and the other two
Swedish species relate only to minor details. The radula of
T. monodon, on the other hand, differs considerably from those
of all Swedish species, as the accessory cusps of the lateral
teeth are very unequal in length; the inner is about double
the size of the outer.

The conchological affinities of the new species are the
reverse : it differs significantly from known Swedish species
by the larger shell, darker color and finer sculpture. In the
latter respects, as well as in the presence of a parietal tooth.

108 TRUNCATELLINA.

T. sundleri agrees well with T. monodon. The new species
however has a larger shell, a much smaller columellar callus,
no impression on the neck and no parietal tooth standing far
within the aperture, such as is distinctly observable in the
last named species (Odhner).

Truncatellina strobeli (Gredler). Vol. 26, p. 71, Vol. 27,
p. 229.

Lothar Forcart has discussed the nomenclature in Archiv
fur Molluskenkunde LX, 188 (May, 1928). He quotes Gred-
ler's description from III Programm des K. K. Gymnasium
zu Bozen, which appeared prior to September, 1853, therefore
prior to Benson's publication of Pupa rivierana.

Having examined Hartmann's specimens of P. minutissima
preserved in the Natural History Museum of St. Gallen, For-
cart states that he could not confirm the presence of teeth in
any of the 33 examples, notwithstanding Hartmann's state-
ment that the aperture has one tooth.

Forcart does not accept the earliest name, callicratis Scacehi,
because Scacehi stated that his species was toothless.

Gredler 's description appears to be the first recognizable
definition of the species, though several earlier names are con-
jectured to pertain to it with more or less probability. I am
now disposed to agree with Forcart in brushing aside the in-
sufficiently diagnostic early work, noticed in Vol. 26, pp. 73-
75, in favor of T. strobeli Gredler.

Perhaps "Pagodina" bourguignati Gout. (27: 179) is really
T. strobeli.

Truncatellina clavella (Reinh.) Vol. 26, p. 81.

Rosen (Mittheil. Kaukasischen Mus. Vol. 6, 1914, p. 194)
treats this as specifically distinct.

Truncatellina micra (Ping). Pupa micra Ping, 1929.
Palaeontologia Sinica, Ser. B, VI, fase. 5, p. 18, f. 4a, b ; pi. 2,
f. 10a, &. Chou Kou Tien, Hopei Prov., China. "Polycene"
(Lower Quaternary) red clay. This genus, which was not
known from China before, seems indicated by the terms of
Ping's description.

COLUMELLA, STERKIA. 109

COLUMELLA (Vol. 27, p. 232).

Columella edentula (Drap.). Vol. 27, p. 236.

Vertigo cylindrica Colbeau (Vol. 25, p. 214) is considered
by Major Paul Dupuis to be an abnormal form of Columella
edentula (Ann. Soc. Roy. Zool. de Belg. LV, 1924, pp. 50, 51,
fig. L 1925).

Columella pygmaeorum Pils. & Ckll. PI. 22, fig. 1.

The minute shell is sinistral, deeply rimate, subcylindric,
the upper third distinctly tapering to the obtuse apex, of a
pale, light brown tint ; the surface dull, without evident sculp-
ture under a lens. Outlines of the spire are somewhat convex.
Whorls strongly convex and very gradually enlarging, the last
ascending slightly to the aperture. The aperture is longer
than wide, toothless. Peristome thin, the outer margin not
expanded, arching forward slightly; columellar lip somewhat
dilated. The umbilical chink is deep and rather long, with
deep axial pit, but we believe no actual perforation beyond
the last whorl.

Length 2.4 mm., diam. 1.1 mm., aperture 0.75 x 0.6 mm. ;
81/4 whorls.

Belgian Congo at Tshibinda, west of Lake Kivu, at about
6000 ft. Type 159672 A. N. S. P., paratype in Brit. Mus.
(Cockerell-Mackie-Ogilvie Exped.)

Columella pygmaeorum Pilsbry and Cockerell, Proc. Zool.
Soc, London, 1933, p. 374, pi. 2, fig. 3.

This species has the essential shell-structure of Columella,
but is peculiar in its narrow form, long taper and sinistral
coil, as well as by its diminutive size. It is the first Ethiopian
species of the genus.

STERKIA (Vol. 26, p. 49).

The species were numbered incorrectly in the key on pages
50, 51. Probably S. rhoadsi and S. antillensis would be better
treated as subspecies of 8. eyriesi.

110 NESOPUPA.

Sterkia calamitosa martiniana Pils. PI. 15, fig. 4.

The shell is very similar to S. calamitosa but differs by the
larger size and longer whorls, and the presence of a supra-
palatal tubercle (rarely wanting).

Length 1.95 mm., diam. 0.9 mm. ; fully 5 whorls. Type.

Length 1.75 mm., diam. 0.9 mm. ; 5 whorls. Paratype.

Length 1.60 mm., diam. 0.9 mm. ; smallest.

San Martin Island, off Lower California, under stones
among plant debris (G. D. Hanna).

Sterkia calamitosa martiniana Pilsbry, Proc. Calif. Acad.
Sci. (4), vol. 16, p. 186, pi. 12, fig. 2. April 22, 1927.

There are a few beautiful albino specimens in the lot.
While very close to mainland calamitosa, the small differences
seem to indicate a slightly differentiated insular race.

NESOPUPA (Vol. 25, p. 274).

Nesopupa corrugata (Preston). Vol. 25, p. 362, 5th line
from bottom : for Fig. 14 read Fig. 12.

Nesopupa rodriguezensis Connolly. PI. 21, fig. 8.

Shell minute, acuminate ovate, rimate, smooth, semi-trans-
parent, moderately glossy, dark corneous brown. Spire
moderately produced, sides convex, apex bluntly rounded.
Whorls 414, rather convex, gradually increasing; apex
smoothly granulate; remainder, under fifty-fold magnifica-
tion, similarly granulate and sculptured with regular, nearly
straight, fine, slightly oblique transverse striolae, a little less
than half as broad as the space between them ; suture deep and
simple. Aperture subquadrate, rounded at the base; per-
istome whitish, very slightly expanded, outer lip nearly
straight and perpendicular in profile, but conspicuously sinu-
ate in plane just below the suture, dental process five-fold : a
short blunt plait at the angle of the paries and outer lip, and
a much longer one, parallel to it and much more deeply inset,
in the centre of the paries; a conspicuous denticle, equally
deeply inset, running out towards the point of the latter from
the base of the siuulus; a longer denticle, more deeply inset

NESOPUPA. Ill

and slightly inclined to the left, just to the right centre of the
base, and a shorter one, nearly as deeply inset and slightly
inclined toward the base, a little more than halfway up the
columella, which is weak, erect, slightly concave, with margin
broadly expanded, but not obscuring the rima (Connolly).

Long. 1.3, lat. 0.75 mm.

Rodriguez Island (Snell and Thomasset). The type has
been presented by H. P. Thomasset to the British Museum.

Nesopupa rodriguezensis Connolly, Journal of Conchology
XVII, Oct., 1925, p. 262, fig. 4.

In view of the numerous insufficiently described and poorly
figured species of Pupillidae recorded from the Mascarene
Is., I am very chary of adding to their number. However,
careful study of Pilsbry's Manual, vol. XXV, seems to show
only one, N. mhmtalis (Mori.) from Mayotte, which can be
compared with the Rodriguez shell. The following points
from Pilsbry's description of minutalis should be found on
comparison with mine of rodriguezensis to establish what must
surely be specific difference between them.

In minutalis: —

(a) The surface is dull, with some irregular striation.

(b) The angular lamella is quite low, forwardly diverging
from the parietal and joining, or almost joining, the outer lip.

(c) The parietal lamella is high.

(d) There are two quite short palatal folds; the upper
palatal is sometimes very small.

N. hisulcata rhodesiana Pilsb., from the Victoria Falls,
appears to resemble more closely the new species, but is larger,
with stronger sculpture and longer denticles {Connolly).

Nesopupa vengoensis Connolly. PI. 21, fig. 9.

The minute shell is almost exactly a replica of N. griqual-
andica (M. & P.), but is slightly less strongly striate, and
differs clearly and constantly in dentition, the lower palatal
fold being absent and the basal tooth so much more deep set
that in some specimens it is hardly visible (Connolly).

Length 1.6, width 0.8, aperture alt. 0.5, width 0.5, last whorl
1 mm.

112 NESOPUPA.

Portuguese East Africa: Mount Vengo, Lorenzo Marques,
5500 ft. (Bernard Cressy).

Nesopupa (Afripupa) vengoensis Connolly, Trans. Roy.
Soc. S. Africa, vol. 12, 1925, p. 165, pi. 4, f. 23.

I would have been inclined to regard this race as merely a
subspecies of griqiialandica were it not that the exact number
of palatal folds is now considered of sectional importance in
the genus Ptychotrema, so should presumably be of at least
specific importance in the Pupillidae (Connolly).

Nesopupa armata (Pease). Vol. 25, p. 327.

' ' Last July Mrs. Cockerell visited the Island of Rarotonga,
and collected there a. Nesopupa which can by no means be separ-
ated from N. armata, described from Borabora, Society Islands.
The aperture is fully armed with the characteristic lamellae,
as figured by Pilsbry, only the infraparietal lamella is larger,
quite half as long as the parietal. The shell is about 2.3 mm.
long, with the characteristic sculpture, the upper whorls with
remote cuticular riblets. This is the first record of a Neso-
pupa from Rarotonga, since it appears that N. dentifera
(Pease), supposed to be from there, came from Aitutaki.
Our shell does not agree with Pilsbry 's figure of N. dentifera,
but it may be that the latter is a varietal form of armata.
Since the fauna of the Polynesian Islands is in general so
extremely different from that of the Hawaiian group, and
the sculpture of typical Nesopupa is so distinctive, it would
seem that N esopupilla and other groups should be regarded
as generically distinct" {T. D. A. Cockerell, Jour, of Conch.
XVII, 1924, p. 168).

Nesopupa novopommerana Rensch. PL 21, fig. 5.

Shell ovate, translucent, dark brown, the apex paler; per-
forate ; 5 whorls. Suture deep. Aperture rounded, the peris-
tome expanded, whitish, with 5 lamellae : angular lamella low
and short; parietal lamella long and higher; the palatal and
columellar lamellae as opposed to the basal lamella [lower-
palatal fold] smaller. The last whorl with a quite weak im-

NESOPUPA. 113

pression at the place of the palatal lamella. After some
delicate growth striae sharp ribs, unevenly spaced, appear,
and the whole surface is sculptured besides with a finely
tubercular structure (Rensch).

Height 1.9 to 2 mm., diam. 1.1 to 1.2 mm.

Bismarck Archipelago: Karlei, New Pommerania (Josef
Schneider) .

Nesopupa novopommerana Rensch, Zool. Anzeiger, vol. 98,
p. 35, fig. 7. March, 1932. Type in Zool. Mus. Berlin.

This species is evidently an hisulipupa, close to N. harrack-
porensis, N. malayana, etc., not nearly related to N. cocosensis.

Rensch remarks that with this species the genus Nesopupa
is recognized from New Pommerania for the first time. . . .
The New Pommeranian species is most similar in shape, color,
dentition and sculpture to N. cocosensis Dall. N. cocosensis is
larger throughout, the peristome is corneous to reddish-brown
(the New Pommeranian perhaps older and therefore white
lipped), the aperture is relatively larger, and the superficial
sculpture lacks the ribs, therefore the tubercular structure
appears more strongly developed than in N. novopommerana.

Nesopupa quadrasi transaequatorialis Rensch.

Differs from N. quadrasi (Mldff.) of Guam [see vol. 25,
p. 335] by the less deep suture and therefore less strongly
convex whorls, by the somewhat stronger impression of the
palatal fold and by the surface structure: the tuberculation
of the surface is very strongly expressed in the New Pommer-
anian form, while the ribs are less prominent, just the opposite
of the Guam specimens before me.

Height 1.5 to 1.8 mm., diam. 1 to 1.1 mm. The specimens
from Guam are uniformly somewhat higher; the type is 1.7
mm. high, 1 wide (Rensch).

Bismarck Archipelago : Malkong, Wide Bay, New Pom-
merania.

Nesopupa quadrasi transaequatorialis Rensch, Zool. An-
zeiger, vol. 98, p. 36. March, 1932. Type in Zool. Mus.
Berlin.

114 BOTHRIOPUPA, PUPISOMA.

BOTHRIOPUPA (Vol. 24, p. 226; Vol. 27, p. 228).
BoTHRiopUPA PERUVIANA, new specics. PI. 16, fig. 6.

The ovate-conic shell has a deep, comma-shaped axial
crevice. Color cinnamon to snuff brown. The rather glossy
surface is minutely, irregularly pitted throughout, with some
wrinkles along growth lines on the intermediate whorls. The
whorls are strongly convex, joined by a deep suture. The
peristome is white, expanded, slightly thickened within.
Parietal lamella strong. Upper and lower palatal folds small,
subequal, the latter slightly further in. Columellar lamella
median, receding.

Length 1.7 mm., diam. 1.2 mm.; 4^/^ whorls.

Peru: Quebrada Boca Pan, below Trigal, Dept. of Tumbez
(A. A. Olsson). Type and paratypes No. 152673 ANSP.

The teeth are decidedly smaller than in B. tenuidens (C. B.
Ad.). It differs from T. breviconus Pils. by the more elevated
conic shape and the longer aperture. Three specimens were
collected.

PUPISOMA (Vol. 26, p. 19).

PupisoMA DioscoRicoLA (C.B.Ad.). Vol. 26, p. 36.

This widely spread species has been reported from Serra da
Baturite, State of Ceara, Brazil (Fred Baker, Proc. A.N.S.
Phila. 1913, p. 632). I found it with the following species at
the upper limit of the wooded zone of Chatham Island, Gal-
apagos (Pinchot Exped.). It is one of the most widely spread
snails of tropical America.

PUPISOMA GALAPAGORUM, nCW SpCcicS. PI. 16, figS. 7, 8, 9.

The shell is globosely conic, or turbinate, the diameter about
87 per cent of the height, with obtuse apex; umbilieate, the
umbilicus contained about five times in the diameter; cinna-
mon-brown. Sculpture of fine, close wrinkles, irregularly
developed, being weak or nearly effaced in places, and absent
on the embryonic li/^ whorls; also a weak and very minute
pitting, which is also quite irregularly developed, and in some
places replaced by granulation. There is no spiral striation.

PUPISOMA. 115

The whorls are strongly convex, joined by a very deeply
impressed suture; the last whorl rounded basally around the
open umbilicus. The aperture is rotund, slightly oblique.
Lip thin, the columellar margin broadly reflected, half cover-
ing the umbilicus in a front view, but scarcely impinging upon
it in a basal view.

Length 1.56 mm., diam. 1.36 mm. ; 3% whorls.

Length 1.6 mm., diam. 1.31 mm. ; 3% whorls.

Galapagos Is. : Chatham Island, on a wooded hill on the
right side of the road from Wreck Bay to Progreso, near the
point where the road emerges into the grassy zone (Pinchot
Exped., July 4, 1929).

This form stands close to P. macneilli Clapp, of Alabama.
It has a noticeably larger, more open umbilicus (compare PI.
16, fig. 8, galapagorum, and fig. 5, macneilli) and a higher
spire, and the growth wrinkles are decidedly stronger in the
Galapagos species.

PuPisoMA coMicoLENSE H. B. Baker. Plate 15, fig. 6.

Shell : small, turbinate ; epidermis light brownish with a
satin luster. Whorls: 4i/^, globose; suture well impressed;
last whorl descending. Embryonic whorls: 1% to 2, darker
in color than later ones; decorated with minute, raised tes-
sellation, which gives surface the appearance of fine-graiued
leather; growth-wrinkles emerge near end of second whorl.
Later whorls: growth-wrinkles weak and low over most of
shell, but slightly intensified at suture, giving latter a puck-
ered appearance (occasional specimens develop a few higher
riblets) ; entire surface vermiculate-granose, without definite
spirals. Umbilicus: small, rimate (about 14 times in raaj.
diam.). Aperture: well rounded and oblique (about 40° to
long axis of shell). Peristome: very weakly expanded in basal
and palatal regions and weakly arcuate in latter; broadly
reflected in columellar, so that half of umbilicus is hidden
(H. B. B.).

Alt. 2.19 mm., maj. diam. 85 per cent of alt. (1.87 mm.),
min. diam. 80(1.76 mm.), aperture, alt. 46(1.00 mm.), diam.
96 (.96 mm.) ; 4l^ whorls.

116 PUPISOMA, ACANTHINULA.

Mexico, Estado Puebla : Necaxa, 2625-3800 ft. ; mainly below
the falls, common on leaves of shrubs and trees (H. B. Baker).

Pupisoma comicolense H. H. B., Proc. Acad. Nat. Sci. Phila-
delphia 79 : 224, pi. 15, f . 4-7, 1927, anatomy only ; Occ. Pap.
Mus. Zool. Univ. Mich. No. 220, Oct. 15, 1930, p. 3, pi. 7, f. 1,
2, shell.

P. comicolense is about the size of P. dioscoricola (C. B.
Ad.), but with one more whorl, is relatively and actually
higher. Its sculpture is most like that in P. minus Pilsbry,
but P. comicolense is far larger and has a relatively much
smaller umbilicus and higher whorls with better impressed
suture. Although it occurs with P. mediamericanum, the
latter is most abundant in lower vegetation and at higher
altitudes {H.B.B.).

Pupisoma bailyi, n. sp. PI. 24, fig. 15.

The shell is umbilicate, globose-conic, the apex very obtuse,
cinnamon colored, of 3I/2 strongly convex whorls, the first
minutely and weakly pitted, the rest with some wrinkles of
growth, unevenly developed, and a very shallow and minute
pitting. Aperture rotund-tnincate, the peristome thin, colu-
mellar margin broadly reflected. Length 1.95 mm., diam.
1.8 mm.

Mexico: Cuernavaca, State of Moreles; type 156099 A. N.
S. P., collected by Joshua L. Baily, 1931.

This equals in size the largest P. dioscoricola, and is of a
more conic shape. It is larger, darker colored and more conic
than other Mexican species.

Pupisoma orcula (Bens.). Vol. 26, p. 31.

Reported from the island of Makatea by C. M. Cooke, Jr.,
Occ. Pap. B. P. Bishop Mus. vol. 10, No. 11, pp. 4, 8. He con-
siders it one of several species introduced by man.

ACANTHINULA (Vol. 27, p. 188).
Acanthinula aculeata (Miill.). Vol. 27, p. 191.

The original reference ioY Helix nucleata (p. 192) is Turton,

ANTHRACOPUPA, GASTROCOPTA, 117

A General System of Nature, etc., vol. 4, 1802, p. 520. He
referred to accounts of aculeata by Muller and Chemnitz. Cf.
Cockerell, Nautilus, vol. 40, July, 1926, p. 31.

ANTHRACOPUPA (Vol. 27, p. 316).

Add: AntJiracopupa hritannica L. R. Cox, Ann. Mag. Nat.
Hist. (9), vol. 18, 1926, p. 168. Carboniferous, Keele Beds,
Clent Hill, Worcestershire.

Ptychospira deloplecta Slavik, Vol. 27, p. 264, is said to
be a scalariform shell of Planorhis declivis Al. Braun, which
Wenz ranks as a synonym of Gyraulus trochiformis applanatus
(Thomae). (Petrbok, Acad. Tcheque des Sciences, Bull.
Internat. XXVI, 1925, p. 192, figs. 4-6. 1926). Ptychospira
thus becomes a synonym of Gyraulus (Planorbidae).

GASTROCOPTA Woll. (Vol. 24, p. 6).
Gastrocopta pellucida hordeacella (Pils.). PI. 22, fig. 7.

Specimens from Mojave Mt., California, are long and cylin-
dric, with the lower-palatal fold of unusual length, and the
basal fold either small or wanting; crest obsolete.

Length 2.6 mm., diam. 0.95 mm.

Length 2.25 mm., diam. 0.9 mm.

S. S. Berry has figured similar specimens from Palm Canyon,
San Gabriel Mts., California (Proc. A. N. S. Phila. 1922, p.
97, figs. 1-4).

Gastrocopta v^olfii (Miller). Vol. 24, p. 94.

Add to synonymy : [Pupilla] reihischi Dall, 1917, Proc. Biol.
Soc. "Washington, vol. 30, p. 10. Not Pupilla {Gastrocopta)
reihischi Dall, 1928, = G. munita hrunnea Pils.

Gastrocopta ashmuni imperfecta Pils. & Ferr.

The columellar lamella is simply curved within, and out-
wardly remains horizontal, as in G. cochisensis, not passing
into the position of an infraparietal lamella. Peristome more
or less free or continuous. The very large angulo-parietal

lis GASTROCOPTA.

lamella and the very deeply immersed lower-palatal fold
remain as in G. ashmimi.

Arizona : Huaehuca and Mustang Mountains, Type 162445
A. N. S. P., coll. by J. H. Ferriss.

Gastrocopta ashmuni imperfecta P. & F,, Proe. A. N. S.
Phila. vol. 75, 1923, pp. 62, 63.

Gastrocopta pilsbryana amissidens, new subsp. PI. 24,
figs. 5, 6.

The shell is like the typical form except that there is no
basal fold. Sometimes the upper-palatal fold also is wanting.
Type 161437 A. N. S. P., from the San Francisco Mountains,
Arizona, collected by J. H. Ferriss. It also occurs at Mahan
Mt. near Mormon Lake, about 20 miles south of Flagstaff,
Arizona, and eastward at the Betatakin ruins, and in New
Mexico at Golden, Santa Fe Co. It is therefore rather widely
distributed in northern Arizona and New Mexico.

Gastrocopta pediculus (Shuttl.). Vol. 24, p. 158.

Bifidaria bannertonensis C. J. Gabriel, 1930, Proc. Roy. Soc.
Victoria, vol. 43, p. 64, pi. 3, figs. 9, 10, appears, from topo-
types received from Mr. Gabriel, to be completely identical
with G. pediculus ; but his record extends the known range of
the species far southward. The original description follows,
with photographic copies of the original figures. Plate 21,
fig. 13.

" Shell minute, white, dextral, attenuate, narrowly umbili-
cate. Apex obtuse. Whorls 5, convex, ornamented by
numerous impressed growth-striae. Sutures deeply impressed.
Aperture roundly oblong, armed with five white teeth; one
situated about center of parietal wall, comparatively large and
unequally bifid; three placed within the basal and outer
margin, the center of which is most prominent; the fifth on
the columella. Peristome expanded ; the columellar expansion
partly concealing the narrow umbilicus. Size of type : length
2.6, breadth 1.3 mm." {Gabriel).

Bamierton [Victoria, Australia] {A. C. Nilson).

GASTROCOPTA. 119

*' Few species have been described from Australia, and the
present is the first representation of the genus in Victoria. In
general arrangement the dentition is fairly constant, but speci-
mens have b^en examined showing the teeth a trifle stronger.
As regards measurements the species is subject to variation,
the paratype figured being 2.3 x 1 mm. Pupa larapinta Tate
and P. nworeana Smith, from Central Australia, bear some
resemblance, but may be distinguished by their broader con-
tour and more convex whorls" {Gabriel).

In view of its wide distribution, my suggestion (Vol. 24,
pp. 158-9) that G. pediculus was imported into New South
Wales within the human period, seems improbable though not
impossible. It may be a very old and conservative species
which reached Australia in the Pliocene. If Pupa margaretae
Cox, from Wallaroo, South Australia, is also pediculus, its
range will be further extended. New collections at that place
will be required to settle this point, as the type of margaretae
seems to be lost.

In 1923 I found pediculus abundant at Tweed Heads, Rous
Co., N. S. Wales, and the var. queenslandica in Brisbane,
Queensland.

Gastrocopta niobe (Fulton). PI. 22, fig. 2.

Man. Conch, vol. 24, p. 153 ; vol. 27, p. 209. Tenimber Is.

Mr. Fulton has been so good as to send a paratype specimen
which he compared with the type in Brit. Mus., and which is
here illustrated, pi. 22, fig. 2. As I noted in vol. 27, it is
identical with the later G. moellendorffiana Pils., 1917, from
Bohol, M. C. vol. 24, p. 145. The description given on p. 145
applies fully to this paratype, which measures, length 1.9 mm.,
diam. above aperture 1 nun. It will be seen that it bears little
resemblance to the original figure and description of P. nioie ;
the species could never have been identified by them; but it
serves to illustrate the difficulties which beset a monographer
of the Pupillidae. Many of the descriptions and figures of
minute species are misleading.

G. niohe belongs to the subgenus Sinaliimda.

120 GASTROCOPTA.

Subgenus Cavipupa, n. subg.

The shell is broadly and deeply umbilicate, of few, strongly
convex whorls; apertural teeth as in Gastrocopta s. sir., the
basal fold subcolumellar in position. Type G. euryomphala.

In the last whorl the diameter of the columellar axis is
contained about three and a half times in that of the shell.
It tapers rapidly upwards (PI. 22, fig. 3). By the open
umbilicus this group resembles the American genus Chaenaxis ;
both were doubtless derived from the typical group of Gastro-
copta, but of course independently.

Gastrocopta euryomphala, new species. PI. 22, figs. 3, 4, 5, 6.

The shell is thin, whitish (dead), widely and deeply
umbilicate, conic, composed of 4^/2 very convex whorls, the
penultimate whorl bulging very conspicuously, the last whorl
becoming flattened peripherally in its last half, and marked
with a short groove over the position of the lower-palatal fold ;
without swelling or crest behind the outer lip. Suture very
deep. Sculpture of microscopic granulation on the first whorl,
the later whorls with fine, somewhat uneven growth striae.
The aperture is shortly oval, with rather thin, broadly ex-
panded and continuous peristome, the upper margin either
free or shortly adnate to the preceding whorl. Parietal lamella
straight and rather strong; angular lamella short, uniting
with the parietal at the front third of the length of the latter.
Columellar lamella median, strong, turning down a little at
its inner end. Lower-palatal fold strong, elongate, the upper-
palatal and basal folds tuberculiform.

Length 2.2 mm., diam. 1.3 mm. Type.

Length 2.1 mm., diam. 1.3 mm.

Philippine Islands : Bintuan, Busuanga. 160431 A. N. S. P.
(Quadras).

The large umbilicus and conspicuously convex whorls dis-
tinguish this from all other known Philippine species. The
specimens were received through Mr. "Walter F. Webb, with-
out name, but I think that they are without much doubt tlie
species which von Moellendorff called Leucochilus eury-

PAUXULUS. 121

omphalum (Abhandl. Naturf. Ges. Gorlitz, vol, 22, p. 153;
1898) from Tangat, Busuanga, and which, as I noted in vol.
24, p. 141, seems not to have been described.

FAUXULUS Schauf. (p. 73).
Fauxulus fryanus (Bens.). PI. 21, fig. 14.

Vol. 24, p. 246. "As this almost unknown species has never
been figured, I illustrate an example in my collection gathered
by Layard at Bredasdorp. The aperture is tubular and par-
ticularly remarkable for the basal canal, which continues as a
sharp, strong keel around the wide, deep umbilicus, while,
owing to the apertural prolongation, two of the lamellae are
so deepset as to be hardly visible ; the sculpture also is remark-
able, consisting after the first two whorls of close, coarse,
curved oblique costae, which are cut from the sixth whorl
onward by particularly strong, close, spiral grooves. The pale
lilac hue of the lower whorls is another unusual feature"
(Connolly, Ann. Mag. N. H., 10 ser., vol. 8, p. 308, pi. 10,
fig. 11).

Fauxulus barnardi Connolly. PI. 21, fig. 4.

"Shell sinistral, rather small, subfusiform, rimate, slightly
silky, rather glossy, corneous reddish-brown. Spire produced,
sides convex, apex mammillate. Whorls 8, not very convex,
increasing very gradually until the last, which is narrower
than the one before it; the first smooth, remainder sculptured
with close, fine, slightly curved, oblique striae ; suture simple,
well defined. Aperture subquadrate, peristome just free,
expanded and continuous; lips vertical in profile; dentition
7-fold : a strong, slightly incurved, inrunning parietal lamella,
arising in the central of the paries ; a similar angular lamella
arising on the edge of the labrum; two blunt denticles, the
lower slightly larger, close together half-way down the labrum ;
a strong, inrunning lower-palatal fold ; a small tubercle about
the center of the base, and a very prominent, descending, in-
running columellar lamella" (Connolly).

"Long. 4.7, lat. 2.6; apert. alt. 1.7, lat. 1.7; last whorl 2.2
mm."

122 ABIDA, CHRONDRINA.

South Africa: Keurboom River Bush, Cape Province
(Barnard).

Fauxulus harnardi Connolly, Ann. Mag. N. H. 10 ser., vol,
8, p. 309, pi. 10, fig. 5. Oct., 1931. Type in South African
Museum.

"Three examples, all mature and alike. While bearing no
resemblance whatever to F. capensis, hurniipiamis, pamphoro-
don, fry anus, or the dextral layardi and crawfordianus, they
are easily distinguishable from falconianus, mcheanianus,
glanvilleanus, ponsoribyanus and pereximius by having only
a single process on the columella, whereas the others have two
or more" (Connolly).

ABIDA (Vol. 27, p. 267).

J. B, de Aguilar-amat has given a sketch of the life of Artur
Bofill i Poch, with portrait and a bibliography of his mala-
cological works, prefatory to his catalogue of Pupillidae in the
Museum of Barcelona, in Treballs del Museu de Ciencies
Naturals de Barcelona, vol. 10, No. 4, 1932. The following
forms, named but not published by Bofill, are illustrated, the
figures reproduced photographically on our Plate 20. No
descriptions are given.

Abida polyodon montserratica forma monsjovica Bofill,
MS. PL 20, fig. 1. Montjuic, Prov. de Barcelona, Spain.
(D 'Aguilar-amat, 1. c, p. 18, pi. 1, f. 1).

Abida polyodon montserratica forma vidali Bofill, MS.
Pi. 20, fig. 3. Hostel Roig, Prov. de Lleida, Spain (D 'Aguilar-
amat, 1. c, p. 18, pi. 1, f. 2),

Abida brauniopsis Bofill, MS. PL 20, figs. 8, 8a. Gosal,
Prov. de Lleida, Spain (D 'Aguilar-amat, 1. c, p. 19, pi. 1, f. 5).

Abida catalonica v^vlida Bofill, MS. PL 20, fig. 2. Mont-
grony, Prov. de Barcelona, Spain (D 'Aguilar-amat, 1. c, p. 22,
pLl,f.2).

CHONDRINA (Vol. 27, p. 283).

Chondrina avenacea subcornea Bofill, MS. PL 20, fig. 12.
Montf errand, Aude, France (D 'Aguilar-amat, 1. c, p. 27,
pLl, f. 6).

ORCULA. 123

Chondrina bigorriensis tenuimarginata forma macro-
CHiLus Bofill, MS. PL 20, fig. 11. Hospitalet de I'lnfant,
Prov. de Tarragona, Spain (D'Aguilar-amat, 1. c, p. 32, pi. 1,
f. 9).

Chondrina bigorriensis tenuimarginata forma flaccida
Bofill, MS. PI. 20, fig. 10. Castejon de Sos and Pont de Neu
d'Astos, Prov. d'Osea, Spain (D'Aguilar-amat, 1. c, p. 32,
pl.l,f.8).

Chondrina dertosensis FxVrta Bofill, MS. PI. 20, fig. 13.
Hifac, Prov. d'Alacant, Spain (D'Aguilar-amat, 1. c., p. 33,
pi. 1, f. 4).

SOLATOPUPA SIMILIS MONTEROSATOI Bofill, MS. PI. 20, fig. 9.

Castel Gandolfo, Massa Carrara, Italy (d'Aguilar-amat, 1. e.,
p. 35, pi. 1, f. 7).

Solatopupa thieuxi Locard, from Correns, Var, France, was
mentioned by d'Aguilar-amat, 1. c., p. 35, without description.
So far as I know it has not been defined.

ORCULA (continued from p. 93).

Dr. Stephan Zimmermann has published a critical study of
great merit, "Ueber die Verbreitung und die Formen des
Genus Orcula Held, in den Ostalpen" in Archiv fiir Natur-
geschichte, n. ser. 1, 1932, pp. 1-56. In order that all names
proposed up to the end of 1933 may appear in tlie Manual, the
following much abbreviated notes are compiled ; his figures are
reproduced. Those concerned with Alpic Orculae will of
course consult Zimmermann 's paper.

In the discrimination of forms below the rank of species
great weight is placed upon dimensions of the shell; but this
is never the measurements of single specimens, but the aver-
age of an entire population. In ascertaining this the length
and breadth of 20 or 30 shells from one place are measured
and the mean taken ; the maximal and minimal records being
also noted.

A synopsis of the forms treated follows. It will be noticed
that four categories are recognized : species, subspecies, forms
and morphs. The distinction between forma and morpha in

124 ORCULA.

this connection is not clear to me ; but I presume that morplia
is applied to populations having characters not thought to be
of racial significance, but denoting changes presumably re-
sulting from environmental influence, such as are correlated
with elevation, aridity and the like. In Zimmermann's
descriptive text the names of both forms and morphs are
written as trinomials, as in the abstract following the table.

Orcula dolium Drap. morpha dolimn, s. str.

morpha infima, n. m.

morpha edita, n. m.

morpha oreina, n. m.

forma pseudogidaris A. J. W.

forma gracilior, n. f.

Orcula dolium Drap. intermediate with gularis Rssm.
(0. gularis pseudodolium A. J. Wagner)

Orcula gularis Rssm. subsp. gularis Rssm.

morpha gidaris s. str.
** ** " " morpha oreina, n. m.

" " " " restituta West.

Orcula dolium intermediate with tolminensis A. J. Wagn.

Orcula tolminensis A. J. Wagner.

Orcula spoliata Rssm. subsp. spoliata Rssm.
" " " AUSTRIACA, n. subsp.

morpha austriaca s. str.
morpha oreina, n. m.

Orcula fuchsi St. Zim.

Orcula conica Rssm. forma conica s. str.

*' " forma minor, n. f.

Orcula dolium DOLIUM (Drap.). Text-figs. 1-6.

The fundamental form has as a rule two equally strong and
nearly equally emerging columellar lamellae. Among them
there are almost always single examples with the upper
lamella somewhat weaker. A third, middle columellar lamella,
somewhat weaker (aberration triplicata) occurs not rarely
as a purely individual hyperplasy, almost entirely in large

ORCULA. 125

shells. The diam. is from 3 to 3.4 mm., length 6.8 to 8 mm. ;
some specimens occur above and below these dimensions.

Orcula dolium infima St. Zim. [1. c. p. 14). Text-fig. 7,

The largest of the species, up to length 9.1 diam. 4.1 mm.,
with 9 to 10y2 whorls. Southern and eastern parts of the
Wiener Wald. It is very similar to the Carpathian (Bad
Trencsin-Teplitz) 0. d. titan Brancsik [vol. 27, p. 9].

Orcula dolium edita St. Zim. (l. c. p. 17). Text-fig. 8.

Smaller, more cylindric than the typical form, diam. be-
tween 2.8 and 3 mm., length 6.7 to 7.3 mm. Palatal callus
and generally the peristome strongly developed. TJie upper
colmnelUir lamella is always weaker than the lower, sometimes
almost obsolete. This is the form of the middle heights of the
subalpine region to the limit of trees. The most characteristic
state of this morph is from the southern slopes of the Schnee-
berg massif ; also found in some heights of the northern Tirol
and Bavarian Alps, and in some places far down in the valleys.
0. d. uniplicata Pot. and Mich, apparently belongs in the
formenkreis of this morph, but no recent author has seen the
type. 0. d. plagiostoma Sandb. (Man. Conch, vol. 27, p. 11)
is also to be considered in this connection. ' ' I would confirm
that the plagiostoma of the Danube valley, which I know from
the loess at Passau, from Pielachberg at Melk, in the "Wachau,
and from Nussdorf and Heiligenstadt near Vienna (fig. 9),
agree in measurement, external shape and apertural characters
with the forms which live today in heights between 1000
and 1600 meters in the Niederosterreichisch-Steirischen
Kalkalpen."

Orcula dolium oreina "A. J. Wagn." St. Zim. {I. c, p. 20).
Text-fig. 10.

The shells are still smaller throughout than the morph edita.
The average dimensions are : diam. 2.6 to 2.8 mm., length 6.2
to 6.7 mm. ; whorls 8 to 9. They are thick-shelled and strongly
rib-striate ; the aperture is small and the palatal callus gener-
ally stronger than in edita. The reduction of the columellar

126 ORCULA.

Explanation op Figures 1 to 20.

Fig. 1, Orcula dolium Drap., Reutte, Nordtirol, 860 meters.

2, Unteres Barenthal bei Feistritz, Karawanken, about 530 m.

3, Emberg bei Kapfenberg, Nordoststeiermark, about 500 m.

4, R. Pfannberg bei Frohnleiten, Steiermark, 530 m. 5,
Merkenstein bei Baden, Niederosterreich, 500 m. 6, conic
shells, Mariaschutz bei Schottwien, Semmeringgebiet, about
700 m.

Fig. 7, Orcula dolium infima St. Zim., Kierling bei Kloster-
neuberg, Niederosterreich, 220 m.

8, 0. d. edita St. Zim., Schneeberg: eug bei Reichenau,
Niederosterreich, 1100 m.

9, 0. d. plagiostoma Sandb., Loess von Pielachberg bei Melk,
Niederosterreich, 230 m.

10, 0. d. oreina St. Zim., Rax : Heukuppe, Niederosterreich-
ischsteirische Grenze, 2009 m.

11, 0. d. pseudogularis Wagn., R. Tiirkensturz, Pittental,
Niederosterreich, 480 m.

12, 0. d. gracilior St. Zim., Adlitzgraben bei Schottwien,
Semmeringebiet, 660 m.

13, 0. dolium -\- — gularis {gularis pseudodolium A. J.
Wagn.), Unteres Reichramingbachtal bei Reichraming, Ober-
osterreich, 400 m. Fig. 14, Hange der Wildgrabenklause bei
Reichraming, 600 m. Fig. 15, Lainautal, Siidfuss des Traun-
stein bei Gmunden, Oberosterreich, 800 m.

16, 0. gularis Rssm., Klein Hollenstein, Ybbstal, Nieder-
osterreich, about 490 m. 17, Ennstal bei Hieflau, Steiermark,
490 m. 18, Nordseite des Loiblpasses, Karawanken, about
1000 m.

19, 0. gularis oreina St. Zim., Grosser Buchstein, Gesause,
Stiermark, 1800 m.

20, 0. gularis aberration reducta St. Zim., Nordwesthange
des Spitzkofel bei Lienz, Osttirol, about 1700 m.

ORCULA.

127

77

14

10

16 17 IS 19 og

Fig. 3 (1 to 20). Orcula in the Eastern Alps (after S. Zimmermann) .

128 ORCULA.

lamellae has gone further : the lower lamella is weak and not
rarely represented by a short, low little ridge far within,
occasionally wanting. As a rule the upper lamella is wanting ;
only seldom a small vestige remains. The smallest examples
of this form I have from the Heukuppe, 2009 meters, the
highest point of the Rax. Width 2.5 mm., length 5.6 to 5.7
mm. (fig. 10). 0. d. oreina is confined to the subalpine zone
of the highest elevations of the northern Dolomites and reaches
about to the tree limit, thus upward generally to 1600 meters,
at its lower limit gradually merging into the form edita.

Orcula dolium pseudogularis a. J. Wagner (vol. 27, p.
11). Text-fig. 11, is a local form of the metamorphic Mesozoic
rocks of the Tiirkensturz, Gleissenfeld in the Pittental, lower
Austria. Wagner's measurements are erroneous. The aver-
age length is 7.2 mm. single specimens up to 8 mm., none over
3 mm. wide. The medium breadth is 2.7 mm., the slenderest
2.6 mm., only a few reaching 3 mm. wide.

Orcula dolium gracilior St. Zim. Text-fig. 12.

The dimensions of this form are strikingly constant, its
medium being for length 6.3 to 6.8 mm., for width 2.8 to 3
mm. ; the diameter being that of edita, but the length some-
what less. A conspicuous difference from edita is in the
eolumellar lamellae which wholly resemble those of typical
dolium and are even more regular than in that, strong and
equally developed. Thus out of 40 from one place in the
Adlitzgraben on the Semmerling, only three have the upper
lamella weaker than the lower (fig. 12). It is a miniature of
dolium. It inhabits a restricted district of the eastern border
of the Alps, from the slopes of the Sonnwendstein and the
Semmering in lower Austria in a northwestern direction about
to Gloggnitz.

Ziramermann is disposed to view pseudogularis and gracilior
as modified subalpine enclaves of 0. d. edita.

Orcula dolium Drap. intermediate with gularis Rssm.
(0. gularis pseudodolium A. J. Wagner, vol. 27, p. 14).
Text-figs. 13, 14, 15.

ORCULA. 129

This form is one of the most remarkable of the genus Orcula.
Its morphologic peculiarity lies in the manifold combination
of the characters of two species, 0. dolium and 0. gularis.
Its relation to these can be interpreted as due to a secondary
mixture, or to the survival of an undifferentiated stock in this
part of the northern Dolomites, representing a stage at the
beginning of the evolution of the two species, which elsewhere
show no intergradatiou. The former hypothesis is more prob-
able, that is, that these forms are the progeny of an actual
mixture of the two species. It inhabits part of the upper
Austrian Alps, bounded on the west by the Traunsee and in
the east by the valley of the Enns.

Orcula gularis (Rssm.). (Vol. 27, p. 13). Text-figs. 16, 17, 18.

This species is confined to the eastern Alps; forms so
labelled from the Carpathians seen are nothing else than slen-
der forms of 0. dolium. ' ' True montane forms are before me
from few places. Those collected on the northern slopes of
the Duerrenstein near Lunz (lower Austria) in about 1500
meters afford average measurements of from 5.7 x 2.45 mm. ;
on the Grossen Buchstein in 1700-1800 meters, 5.8 x 2.5 mm.
The shells are in general somewhat thicker and more strongly
rib-striate, the folds of the aperture, especially the parietal,
weaker. This is forma 0. gularis oreina St. Zim. (fig. 19).

Orcula gularis restituta (Westerl.) (Vol. 27, p. 14). Text-
figs. 21, 22.

This form does not differ from 0. gularis proper in external
form or dimensions, — width 2.7 to 2.8 mm., length 6.3 to 6.5
mm. The columellar lamellae are of equal strength, lie rather
close together, and reach equally far, quite to the sharp edge
of the peristome. The palatal callus is mainly very delicate,
not rarely lacking entirely, but it is also often quite strongly
developed, and then shows, near its outside limit, a little
prominence, which however has nothing in common with the
rudiment of a palatal fold ; the latter being ivanting in all
cases (figs. 21, 22). St. Annatale south of the Loiblpasse, etc.

130 ORCULA.

Explanation of Figures 21 to 42.

21, Orcida gularis restituta West., Feistritztal bei Stein,
Steiner Alpen, 390 ra. 22, Siidhang der Velka Planina ober-
halb St. Primus, Steineralpen, 950 m.

23, Orcula gularis -j tolminensis A. J. Wagn., Daehstein-

Siidseite : Rabenkogel, Steiermark, 1270 m. 24, Zaghalswande,
Steiermark, 1850 m.

25, 0. tolminensis A. J. Wagn., Strasse zwischen Hallstatt
und Gosaumiihl, Oberosterreich, 500 m. 26, Kuppitzklamm
bei Eisenkappel, Karawanken, about 700 m,

27, 0. spoliata Rssm., Valdagno bei Vicenza, Lessinische
Alpen, about 270 m. 28, Fennberg bei St. Margreid im
Etsehtal, Siidtirol, 1000 m. 29, Oberhalb S. Romedio im Val
de Non, Siidtirol, about 950 m.

30, 0. spoliata austriaca St. Zim., Lilienfeld, Niederoster-
reich, 400 m. 31, Hohe Wand bei Wiener Neustadt: Sprin-
gelsteig, Niederosterreich, 700 m, 32, Schwartzau im Gebirge,
Sehneeberggebiet, Niederosterreich, 620 m.

33, 0. spoliata austriaca oreina St. Zim., Goller, Nider-
osterreich, 1750 m.

34, 35, 0. fuchsi St. Zim., Turmmauer, Nordhang des Goller,
750 m. 36, Waldhiittsattel siidlich von Kernhof, Nieder-
osterreich, 1200 m.

37, 0. conica Rssm,, Koranatal bei Plitvice, Kapellenger-
birge, Siidkroatien, 520 m. 38, Loibltal, Karawanken, about
600 m. 39, Oelgraben im unteren Lavanttal, Kamten, about
450 m.

40, 0. conica minor St. Zim., Gurnitzer Schlucht, Satnitz,
Kamten, about 490 m.

41, 0. dolium -\- — conica ? Grosser Suhagraben bei
Maria Elend, Karawanken, about 800 mm. 42, Windisch-
bleiberg, Karawanken, about 950 m.

ORCULA.

131

24

25

26

27

2S

29

31

32

38 39 40 41 42

Fig. 4 (21 to 42). Orcula in the Eastern Alps (after S. Zimmermann).

132 ORCULA.

Orcula tolminensis a. J. Wagner. (Vol. 27, p. 16). Text-
figs. 25, 26.

Orcula SPOLiATA (Rssm.) (Vol. 27, p. 14). Text-figs. 27, 28, 29.
Orcula spoliata austriaca St. Ziin. Text-figs. 30, 31, 32.

The habitat of this species, of which I have material from
more than 60 localities, comprises the eastern part of the
northern Dolomites. It differs from 0. spoliata spoliata in
dimensions ; it is constantly more slender. The average length
lies between 6 and 7 mm., the breadth between 2.6 and 2.8
mm.; a further difference is that the whorls are less convex
and the suture shallow; only the last whorl is somewhat
swollen. Whorls 9-10%. The columellar lamellae are often
weaker than in the typical race, the upper always weaker
than the lower, and not rarely indicated by only a delicate
ridge of the columellar wall. The strongly entering palatal
callus is common to both forms, but in austriaca is rather
more strongly developed (figs. 30, 32).

At high elevations it is smaller. On the Goller in lower
Austria at an elevation of 1750 meters, as well as on the Rax
plateau (Gsollhirn) at height of 1540 meters there is a true
high region form, 5.5 x 2.5 mm. The shells are thicker, palatal
callus rather stronger; parietal and lower columellar lamellae
are weaker, while the upper columellar lamella is not per-
ceptible. This is 0. s. austriaca morpha oreina, Text-fig. 33.

Orcula fuchsi St. Zim. (This vol. p. 89) . Text-figs. 34, 35, 36.
Orcula conica (Rssm.) . (Vol. 27, p. 15) . Text-figs. 37, 38, 39.

The distribution of this species comprises the extreme south-
east of the Alps and the contiguous Dinaric mountains, there-
fore chiefly the Karawanken Julischen and Steiner Alpen and
the elevations southeastward of these mountains including the
Kapellen mountains. Nearly all localities are upon Mesozoic
limestone.

The average dimensions are : length 6 to 7 mm., diam. 3 to
3.5 mm. Though smaller specimens occur anywhere in its
area, they are constant in a few places in the Satnitz

ORCULA. 133

(Gurnitzer Schlueht and in a Schlueht of the Windachwald).
The shortly conic shells there have average size of 5.4 x 2.95
mm. This is Orcula co-nica minor, new local form, H. von
Gallenstein in schedis (fig. 40).

Among typical conica from two places in the Karawanken
(Grosser Suhagraben bei Maria Elend and Windischbleiberg)
there are specimens standing midway between the two species
in being more cylindric than typical conica and more conic
than typical dolium, with a weakly labiate peristome and two
thin, equally strong columellar lamellae reaching to the edge
of the peristome (as occurs exceptionally in 0. dolium) ; so
that it seems impossible to refer them with certainty to either
species (text-figs. 41, 42). The number in each place is very
small. Can it be that there are transitions between 0. dolium
and 0. conical

Orcula doliolum (Brug.) var. (Vol. 27, p. 17).

Valley of the Vantsch River, Darvaz, Pamir (Kaznakov).
"These specimens agree with those of middle Europe in size
and sculpture and differ from them only in that the shell is
quite cylindric (not broader above), and that the two colu-
mellar lamellae are more weakly developed and not visible in
a direct front view in the mouth. Wholly similar are many
specimens from Transcaucasia. The species has been reported
for Central Asia (Rosen, 1901) and is known from various
places in the mountainous part of Turkestan. It has been
recorded from Kopet-Dag in Transcaspia, Baron 0. von Rosen,
and from north Persia (0. Boettger 1898), so that it is easy
to make the connection with its occurrence in Transcaucasia
and Asia Minor. It is well known to extend thence over cis-
Caucasia (Retowski, 1914) and the Crimea (Lindholm 1926)
to southeastern and middle Europe. It is a pronounced moun-
tain dw^eller" (Lindholm, 1931, Abhandlungen der Pamir-
Expedition 1928, VIII, p. 43).

Orcula tomlini Connolly. PI. 21, fig. 12.

"Shell small, cylindrical, rimate, thin, silky, translucent,
corneous brown. Spire produced ; sides straight and parallel,

134 ORCULA.

summit (4 whorls) bluntly conic. Whorls 8, almost flat,
gradually increasing; protoconch (1% whorls) engraved to the
apex with close, regular, microscopic spiral grooving, re-
mainder sculptured with fairly close and regular, slightly
curved and oblique transverse striae. Suture simple, shallow.
Aperture subquadrate, rounded at the base ; peristome white,
glossy and reflexed. Dentition 7-fold: an acute, deeply in-
running lamella on the center of the paries, with two large,
rounded tubercles, thickening outwards in the direction of,
but not touching, the ends of the peristome; two prominent
inrunning lamellae, arising on the outer margin of the colu-
mellar lip, one-half and three-fourths way up the columella,
and two narrower ones deepset above them; columella erect,
margin rather narrowly reflexed" (Connolly).

"Long. 6.3, lat. 2.7; apert. alt. 1, lat. 1; last whorl 3.5 mm."
Tripoli: Cyrenaica north of Merj (/. W. Oregory).

Orcnla tomlini Connolly, Ann. Mag. N. H., 10 ser., vol. 8,
p. 305, pi. 10, fig. 1. Type in Tomlin collection.

"Two specimens, the second measuring 6 mm. in length
and lacking one of the smaller immersed lamellae at the top
of the columella. The feature which appears to separate this
species from all other known Orculae is the presence of the
small tumuli on the paries; they are exactly similar in both
examples, and can hardly represent an ordinary callous pro-
cess, since, despite their size, they neither connect with the
parietal lamella nor with either end of the peristome. Some-
thing of this kind is present on 0. orientalis var. nitida Mouss.
and 0. moussoni Reinh., both from Aleppo, but the tubercles
are smaller and are connected with the margins. In Orcula
tomlini the two lower columellar lamellae are extremely con-
spicuous.

"I have much pleasure in naming this beautiful species in
honor of its owner, in grateful recognition of the courteous
and ever-ready help and advice in many branches of science,
for which countless students besides myself are indebted to
him" {Connolly).

PUPILLA. 135

Of uncertain position.
"Pupa" lata C. B. Adams. PI. 21, fig. 15.

See vol. 27, p. 246, where the original description of this
species is repeated, and it was referred with a mark of doubt
to the genus Columella. The unique type is here figured. It
is imperforate, yellowish corneous, slightly translucent, very
shortly cylindric with conic summit. The high first half-
whorl is smooth, the rest delicately costulate, the riblets low,
very narrow with wide intervals, stronger on the upper part
of the spire, on the last whorl stopping abruptly at the weak
angulation which bounds the smooth, rather flattened base.
There are nearly 4 rather strongly convex whorls. The aper-
ture is oblique. Peristome is not in the least expanded and
is thick for so small a shell, its edge smoothly rounded. Colu-
mella with an indistinct, closely adnate reflection above.
Length 1.9 mm., diam. 1.35 mm.

It differs from Pupiwma and Columella by the smoothly
rounded finish of the lip, and the columella. The riblets are
not euticular. I take it to be the embryonic stage of some
larger shell, but I have not been able to trace the species. It
is not a pupillid snail, in my opinion. No locality further
than ' ' Jamaica ' ' is known.

PUPILLA (Vol. 26, p. 152).
PuPiLLA cuPA (Jan). Vol. 26, p. 185.

A study of the specimens in the Charpentier collection has
been made by G. Mermod (Revue Suisse de ZooL, vol. 33, No.
17, July, 1926, pp. 577-582).

The types of Pupa alpicola Charp., two specimens from
Mont Gietroz near Fiomay, valley of Bagnes, Valais, are fig-
ured, his figures reproduced in our PI. 24, fig. 11.

Pupa halleriana Charp. agrees with P. cupa, the only dis-
tinction mentioned by Charpentier being the presence of fine
granulation on the shell. In certain lights such microscopic
granulation occurs in the majority of the shells of muscorum,
sterri and triplicata.

136 PUPILLA.

M. Mermod concludes that ' ' the specific value of P. alpicola,
halleriana, madida and muscorum var. praten^is seems to me
very doubtful. I think that, without risk of gross error, all
these names can be put in synonymy and considered to form
simply a variety of P. muscorum.^' The synonymy, in this
view, will stand as follows:

PupiLLA MUSCORUM var. CUPA (Jan).

Pupa alpicola Charp. [Vol. 26, p. 183].

Pupa halleriana Charp. [Vol. 26, p. 185],

Pupa madida Gredler [Vol. 26, p. 184] .

Pupa muscorum var. pratensis Clessin [Vol. 26, p. 178] .

PuPiLLA STERRi (Voith). Vol. 26, p. 186.

According to Mermod (i^. c, p. 581) this differs from P. cupa,
with which it was synonymized by 0. Boettger, by the strong
epidermal striae, often anastomosing, the excentric aperture
with a strong cervical crest, and the more widely open um-
bilicus. See also, Manual, vol. 27, pp. 253, 254.

PupiLLA PSEUDOCYLiNDRiCA (Ping & Yen). Plate 24, fig. 10.

''Shell small, cylindrical in outline, shallowly umbilicated.
Apex rounded, fairly broad. Whorls 6i/2- First whorl widen-
ing not so much as the second, which almost equals any of the
following whorls in width. Then third, fourth, fifth, even
sixth, which constitutes the body-whorl, all equal to one an-
other in width, thus making the shell cylindrical. Apex and
first whorl smooth, and surface of each of the following whorls
marked with very fine oblique striae. Surface of each whorl
moderately convex. Suture between each two whorls fairly
deep. Aperture ovoid. Outer and inner lips continuous and
somewhat expanded, with a thin callus on the parietal surface,
which bears a very small, inconspicuous tooth. Umbilicus
shallow and slit-like. Along the outer lip the peristomal
region thickened, forming a blunt keel. The general color is
yellowish white, and the apical region and the lips are purely
white" (Ping and Yen).

Alt. 4 mm., diam. 1.7 mm., length aperture 1 mm., width
0.9 mm.

PUPILLA, MICROSTELE, PUPOIDES. 137

N. W. China : Aksu, in a dry river bed. Type 547 Fan Mem.
Inst. Biol.

Truncatellina paseudocylindrica Ping and Yen, Bull. Fan
Memorial Institute of Biology, vol. 4, p. 278; Truncatellina
pseudocylindrica on p. 279, fig. 15.

"This species resembles in some way Truncatellina cylin-
drica (Fer. 1821), but that its size is much larger, with one
more whorl, the striae on its whorl surfaces are much finer,
and its parietal surface has a small nodule or tooth, although
it is very inconspicuous, make it different from the latter"
(Ping and Yen).

This species and Pupilla cupa Jan, described by Ping and
Yen from the same place (1. c, p. 279), require comparison
with Pupilla miiscorum asiatica and P. cupa turcmenia. I
have not seen them, and it is a very difiicult group.

Pupilla ( ?) inermis Eussell, 1931. Trans, Roy. Soc.
Canada, (3), vol. 25, p. 15, Probably Paleocene. Alberta.

Pupilla grabaui (Ping). Pupa grahaui Ping, 1929, Palae-
ontologia Sinica, Series B, VI, fasc. 5, p, 10, f, da-c, pi. 1, f. 3).
Shueh Hua Shan, Hopei Province, China, in the supposed
Hipparion red clay ; if so, of Pliocene age, A sinistral species.

Pupilla hopeiensis (Ping), Pupa hopeiensis Ping, 1929,

1, c, p. 11, f. 5a-c, pi. 1, f. 5a-c. Same locality and horizon.

MICROSTELE (Vol, 26, p. 147).

Microstele pingi, n. n. Pupa suhconica Ping, 1929, 1. c,
p. 11, f . 4a, h ; pi. 1, f , 4, Not Pupa suhconica Sandberger,
1858, Shueh Hua Shan, Hopei Prov., China, in the supposed
Hipparion red clay; if so, of early Pliocene (Pontian) age.
I have not seen this fossil, but its characters seem to be wholly
those of Microstele.

PUPOIDES Pfr, (Vol, 26, p, 108) ,

Pupoides "Fer." Hermannsen, 1848, Ind. Gen. Malac, vol.

2, p. 364, type Helix carinula Gmelin (Syst. Nat. (13), p.
3655, based upon Chemnitz, Conch. Cab. IX, p. 190, pi. 136, f.

138 LAURIA.

1263, nos. 1-4). Being without generic definition this has
only the status of a nude name, since the genotype is not a
determinable species. The name therefore does not preoccupy
Pupoides Pfr. 1854. "Pupoides" was not used in a generic
or subgeneric sense by Ferussac.

Pupoides hordaceus (Gabb). Vol. 26, p. 116.

Add to synonymy: Pupoides eupleura Chamberlin and
Berry, 1931, Proc. Biol. Soc. Washington, vol. 44, p. 7, fig. 1
(Henry Mts., Utah, west of King's Kanch, type loc. ; also
Cannonville, Utah). Topotypes received from Prof. Cham-
berlin agree in every respect with P. hordaceus.

PAGODULINA Clessin (Vol. 27, p. 166).

Add to synonymy of the genus: Pagodinella Thiele, 1917,
Nachrichtsblatt D. M. Ges., vol. 49, p. 24, new name for
Pagodula Hesse. Type Pupa pagodida Desm.

LAURIA (Vol. 28, p. 86).

Lauria bequaerti, new species. Plate 24, figs. 12, 13, 14.

The shell is perforate and openly rimate; ovoid, tapering
from the last whorl to the rounded, obtuse summit; chestnut
colored. The surface is glossy, with fine, retractive striae of
growth, more prominent below the suture. The whorls are
moderately convex ; suture impressed. The peristome is light
brown, narrowly expanded, thickened within, the thickening
excised in the upper third of the outer lip. Columellar margin
dilated. The angular lamella tapers inward and is rather
short, not penetrating deeply, outwardly curving towards and
strongly connected with the insertion of the outer lip. Colu-
mella convex but without a superposed lamella or tooth.

Length 4.35 mm., diam. above aperture 2.2 mm. ; 6I/2 whorls.

Length 3.65 mm., diam. above aperture 2.1 mm. ; 5^/2 whorls.
Cotype.

Length 3.65 mm., diam. above aperture 2.2 mm. ; 5l^ whorls.
Cotype.

Belgian Congo: Lukafu River, Katanga, near the falls, in

DISTRIBUTION OP PUPILLIDAE. 139

moss dripping wet from spray of the falls, collected by Michael
Beqiiaert. Cotypes 162443 A. N. S. P. and in Mus. Comp. Zool.

A young shell of fully four whorls has a long parietal
lamella and four radial folds in the base (fig. 13) . Three folds
persist in a shell of fully 4i/^ whorls, but they are weaker.

It is more robust and darker colored than any of the South
African Lauriae. L. hruguierei of Abyssinia is paler and
more cylindric. L. desiderata (Preston), from Mt. Kenia at
9-10,000 ft., is apparently the species most nearly related, as it
is the nearest geographically; from the description it is a
smaller shell, said to measure 3x2 mm.

By its angular lamella entering only shortly, L. bequaerti
differs from the majority of the typical section of Lauria.
Like many of the Madeiran species, it lives in very wet places,
A few of the shells are partially coated with hard calcareous
material such as one sometimes sees deposited on fresh water
shells.

Lauria zonifera n. n. (This vol. p. 86) .

New name for Pupa zonata Bttg. 1883 (see vol. 27, p. 79),
not Pupa zonata Gassies, 1869. Boettger proposed the name
mut. alhina for albinistic examples, but this name had been
used in Pupa several times, as by Moquin-Tandon, 1855.

Geographic Distribution of the Pupillidae.

The family Pupillidae is essentially a group of the northern
continents. The data now at hand indicate Eurasia as the
main area of evolution and radiation. All of the major groups
(subfamilies) occur in this continent. Of about 50 genera
recognized in the family, 38, or about 75 percent, are repre-
sented in Eurasia, either living or as Tertiary fossils.

The southern continents and islands have, in addition to
northern genera which extend into them, only about 8 endemic
genera :

Gihhulina, South America.

Fauxulus, South Africa.

Canipolaemus, St. Helena.

Costigo, Indo-Malayan and Melanesian islands.

140 DISTRIBUTION OP PUPILLIDAE.

Cylindrovcrtilla, Melanesia and tropical Australia.
Lyropupa, Hawaiian Islands.
Pronesopupa, Polynesia, Hawaii.
Pupoidopsis, Polynesia, Hawaii.

All of these are more or less closely related to genera occur-
ring in Eurasia. There is no trace of Antarctic elements sug-
gesting dispersal via Antarctica.

Of the 38 genera of Eurasia, 30 genera (18 of them Recent)
occur in the western division of the Palaearctic Region, and
about 15, all Recent, in the Oriental Region; 8 of these not
in the Palaearctic. It is evident that old centers or areas of
evolution are in both southeastern Asia and in the European
subregion; but the presence of such Oriental genera as
Nesopupa, SinalUnida and Microstele in European Tertiary,
and Truncatellina in Chinese Tertiary and in the Riukiu curve,
is evidence that Tertiary exchanges of genera were more ex-
tensive than have taken place under the strongly differentiated
climates of post-Tertiary time.

Few if any of the extinct Tertiary genera now known are
generalized or synthetic types. The main evolution of the
group seems to have been in the Paleocene and Cretaceous,
and still remains to be recovered. The European Oligocene
and Miocene pupillid fauna appears to be about as specialized
and mature as the Recent, and contains many genera and sub-
genera still widely spread.

In no other family of land snails are the genera so widely
distributed as are some of the genera of Pupillidae, such as
Oastrocopta, Pupilla, Pupoides and Nesopupa. The data may
be found in the lists on following pages, but the ranges and
probable lines of dispersal of several genera are illustrated in
figures 5 to 8, pages 142 and 143. These nearly world-wide
ranges are an evidence of the antiquity and the great con-
servatism of the groups. Many of the genera of this family,
as of most others, are restricted to single zoological regions
or to smaller faunal divisions.

Figures 5 and 7 represent the distribution and probable
routes of dispersal of two genera of northern origin, the dotted

DISTRIBUTION OP PUPILLIDAE. 141

areas on tlie map being more or less dubious, Pupilla proper
has not spread far to the south, but the earlier subgenus
Gibhulinopsis^ has reached South Africa, Australia and
Tasmania. The subgenus Striopiipilla in western North
America must be due to a much earlier migration than that
of Pupilla s. str.

Fig. 5, the northern group of Gastrocopta, has about the
same range as Pupilla, but it is still present in many islands
between Asia and Australia, and it has been differentiated into
several regional subgenera in its secondary areas. North
America and Australia.

Fig. 8, the routes of dispersal of the typical group of Gas-
trocopta are problematic. The rather high stage of evolution
shown by the complete concrescence of angular and parietal
lamellae, and the great similarity of American and Ethiopian
species are points in favor of recent evolution and dispersal
of the stock; but on the other hand, its distribution in the
tropics of both hemispheres, and tiie presence of strongly
differentiated derivatives, such as Immersidens, Chaenaxis
and probably Gihhulina, argue for considerable antiquity. It
may be a genus which matured early in the Ethiopian Region
and reached America via a transatlantic land bridge, but this
would put it back into the Cretaceous or at least Paleocene,
with practically no change in part of the stock since. Further
paleontologic discoveries must be awaited before deciding be-
tween an Atlantic or the northern route ; but in any case, the
extreme similarity of African and American species is
remarkable.

The very extensive distribution of some species of Gastro-
copta due to commerce is not represented on the maps.

Fig. 6, Pupoides has much in common with Gastrocopta
s. str. in distributional peculiarities, but unlike that group,
Pupoides reached Australia. The problem of dispersal is the
same in both genera. The presence of Microstele in European
Miocene and apparently in Chinese Pliocene is interesting in
this connection, as this genus appears to stand in an ancestral

1 This is a prior name for Primipupilla; see vol. 27, p. 254.

142

DISTRIBUTION OF PUPILLIDAE.

Distribution of Northern Group
of Gastrocopta

1. Alb inula

2. Sinalb inula

3. Vertigopsls

4. Privatula

5. Australbinula

o

(^\-[

Di6T;ribution of Pupoides.
At M in Florida, Microcerion.
M in central Europe, Africa and Asia, Microstele.
At I the subgenus Ischnopupoides.

Fig. 5 (upper). Distribution of Northern Group of Gastrocopta.
Fig. 6 (lower). Distribution of Pupoides and its Allies.

DISTRIBUTION OF PUPILLIDAE.

143

1. Pupilla s. str.

2. Gibbulinopsis

3. Striopupilla.

• 2

Distribution of Pupilla,

Distribution of Gastrocopta s. str.

r.-a

Fig. 7 (upper). Distribution of Pupilla.

Fig. 8 (lower). Distribution of Gastrocopta s. str.

144 DISTRIBUTION OF PUPILLIDxNJ].

relation to Pupoides. Microcerion, in the Upper Oligocene of
Florida, is evidently a minor modification of Microstele. As
living snails, Microstele has a few widely scattered species in
Ceylon, East and South Africa, fragments of its once wide
range.

Palaearctic Region.

Atlantic Islands. — In the archipelagoes of the north Atlantic
the Leiostyla group of Lauriae has been widely deployed, with
many endemic species forming several special subgenera and
sections, and occupying stations from arid to subaquatic.

Madeiran species.

Besides a few obviously introduced species, Lauria cylin-
dracea and L. c. anconostoma (27:51), Madeira has four
genera of Pupillidae. Staurodon and several subgenera of
Lauria are endemic. Staurodon stands close to Nesopupa,
which was present in the European Oligocene and Miocene.
The Lauriae are partly near European Oligocene and Miocene
Leiostylas, and in part are intensely specialized derivatives
from Leiostyla. The other genera are Palaearctic groups,
here represented by endemic species. The fauna seems to have
been derived from Europe not later than Miocene, and per-
haps earlier.

Lauria cylindracea anconostoma Lwe. 27 :51. Introduced.
L. fanalensis Lwe. 27 :58.

(S.-g. Scarahella) L. s. simulator P. 27 :101.

L. cassida Lwe. 27 :88. L. wollastoni Paiv. 27 :102.

(8.-g. Leiostyla) L. laevigata Lwe. 27:103.

L. eheilogona Lwe. 27:90. L. recta Lwe. 27:104.

L. vincta Lwe. 27 :91. L. r. macilenta Lwe. 27 :104.
L. V. watsoniana P. 27:92. {Beet. Craticula)

L. irrigiia Lwe. 27:92. L. fusca Lwe. 27:105.

L. loweana Woll. 27:93. L. millegrana Lwe. 27:106.

L. 1. transiens Woll. 27:96. L. abbreviata Lwe. 27:107.

L. cassidula Lwe. 27 :96. L. comeocostata W. 27 :108.

L. concinna Lwe. 27 :97. L. c. resticula W. 27 :109.

L. laurinea Lwe. 27:98 L. relevata Woll. 27:110.

L. sphinctostoma Lwe. 27 :99. L. ferraria Lwe. 27 :110.

L. s. arborea Lwe. 27 :100. L. degenerata Woll. 27 :111.

L. s. heterodon P. 27:101. L. monticola Lwe. 27:112.

DISTRIBUTION OF PUPILLIDAE. 145

L. m. pumilio W. 27:113. L. gibba Lwe. 27:115.

L. ealathiscus Lwe. 27:113. (S.-g. Mastula)

{S.-g. Wollastanula) L. lamellosa Lwe. 27:116.

Staurodon saxicola Lwe. 25 :224.
S. s. seminulum Lwe. 25 :225.

Truncatellina linearis Lwe. 26 :62.

Columella mierospora Lwe. 27 :234.
C. limnaeana Lwe. 27 :235.

Azores Archipelago.

As in Madeira, the Leiostyla stock is the most abundant
group, and has been notably modified to form the subgenera
Senilauria (28:88) and Azoripiipa (27:127). L. c. ancono-
stoma and C. mierospora are apparently introduced forms.

Lauria cylindrica anconostoma Lwe. 27:51.

L. (Senilauria) fasciolata Morel. 27:123; 28:88.

L. (S.) f. hortanaPils. 27:123. Fayal.

L. (Leiostyla) fuscidula Morel. 27 :124.

L. (L.) f. hexodon Pils. 27:125. San Miguel.

L. (L.) rugulosa Morel. 27:126. Pico.

L. (L.) vermiculosa Morel. 27:126. San Miguel.

L. (Azoripupa) tesselata Morel. 27:128. Santa Maria.

Columella mierospora Lwe. 27 :234.

Acanthinula azorica Pils. 27 :193. San Miguel.

Canary Islands.

The occurrence of a large Gastrocopta on Tenerife is re-
markable, but its affinities are unknown, and the record has
not been confirmed by any later collector. Lauria c. ancono-
stoma and Columella mierospora, common to Madeira, the
Azores and Canaries, are the only non-endemic species.

Gastrocopta moreletiana Grasset. 26 : 228. Tenerife.
Granopupa granum bulimiformis Mss. 24:340.

Lauria c. anconostoma Lwe. 27 :51. Introduced ?

L. fanalensis pyramidata P. & S. 27 :60. Tenerife.

L. (Leiostyla) castanea Sh. 27:118. Tenerife and Palma.

L. (L.) pythiella Mouss. 27:119. Tenerife, Palma, Hierro.

L. (L.) taeniata Sh. 27:120. Tenerife and Palma.

146 DISTRIBUTION OF PUPILLIDAE.

Truncatelliiia atomus Shuttl. 26:63. Tenerife.
Columella microspora Lwe. 27:234. Tenerife, Palma.
Aeanthinula spinifera Mouss. 27 :193. Palma, Grand Canary.

Cape Verde Islands.

While the Helices of these islands appear to be generically
related to those of the Azores and Madeira, the few Pupillidae
have a decidedly Ethiopian aspect. Three of the genera are
Ethiopian, the other two, Lauria and Truncatellina, are Palae-
arctic genera which have spread widely, being common to
Europe, the Atlantic islands and the Ethiopian Region, from
Abyssinia to the Cape.

Gastrocopta acarus Bens. 24:122.

Pupilla fontana gorgonica Dhn. 26 :206.

Lauria cylindracea anconostoma Lwe. 27 :51.
dohrni Pfr. 27 :53. S. Antao.

Pupoides gemmula Bens. 26 :137.

Truncatellina molecula Dohrn. 26:63. S. Antao.

Pupillidae of European and Mediterranean suhregions.

Four circumpolar genera, Pupilla, Vertigo, Columella and
Zoogenetes extend to the Pacific the eastern limit of the
Palaearctic Region, and into America. Truncatellina has
species in China and the Riukiu Islands, and the single Recent
species of Ptychalaea occurs in the Ogasawara (Bonin) group.
Otherwise the European Pupillid fauna has nothing in com-
mon with the rest of the Palaearctic Region. The relation
was more obvious in the middle Tertiary, when Sinalbinula
and Ptychalaea, now living in China and the Japanese islands,
were represented in the central European fauna.

Europe, North Africa and nearer Asia possess by far the
richest fauna of Pupillidae in the world. There are 18 Recent
and 12 additional Tertiary genera, of which no less than 11
Recent and 7 fossil genera are endemic. The largest Pupil-
lidae, comprised in the subfamilies Orculinae and Abidinae
(the Abida Group, vol. 24, p. xi), are, with the exception of
the genus Fauxulus, all European.

DISTRIBUTION OF PUPILLIDAE, 147

Eastward, in central Asia, the Palaearetic Region has only
a scanty fauna, listed on p. 148. In China and Japan there
is a somewhat richer fauna, resembling the Miocene of Europe
in the copious development of Gastrocopta and Gibiulinopsis.^
Vertigo also is well represented.

In the following list of the genera of the European system,
those restricted to that region are starred (*). References
are added to volume and page where the genera and species
are described. The geological age of first appearance is added.

Odontocyclas* 24:254.

Sandahlia* 24:258.

Abida* 24 :262 ; 25 :370 ; 27 :212, 269 ; 28 :122. Lower Eocene.

Granopupa* 24 :332 ; 27 :214.

Chondrina* 25 :1 ; 27 :212, 283 ; 28 :122.

Pupilla 26 :152 ; 27 :253, 254 ; 28 :135. Upper Oligocene.

Lauria 27:43, 258, 319. Upper Oligocene (Subg. Leiostyla),

Orcula* 27 :1, 256 ; 28 :88, 123. Paleocene.

Orculella* 28 :92.

Pagodulina* 27 :166, 320. Upper Pliocene.

Spelaeodiseus* 27 :180.

Agardhia* 27 :129, 319. Miocene.

Vertigo 25 :69 ; 27 :216 ; 28 :93. Eocene.

Truncatellina 26 :58 ; 27 :229 ; 28 :105. Upper Oligocene.

Columella 27:232; 28:109.

Spermodea* 27 :186. Upper Oligocene, Lower Miocene.

Acanthinula 27:188. Paleocene.

Zoogenetes 27 :195.

The following genera occur fossil in the European Tertiary
but are now either extinct, or are no longer living in the
European fauna. Genera starred (*) are extinct and endemic.

Gastrocopta. 24 :112.
Alhinula. Middle Oligocene to Pliocene.
Sinalhinula. Late Oligocene to Pliocene.

Microstele 26:147. Miocene.
Enneopupa* 27 :222. Upper Oligocene.
Paracoryna* 27 :132, 320. Miocene.

1 It is rather a pity that the inappropriate name Gibbulinopsis has to
be used in place of Primipupilla. See vol. 27, p. 254.

148 DISTRIBUTION OP PUPILLIDAE.

Ptychalaea. 25 :220 ; 27 :225. Miocene to Pliocene.

Paracraticula* 27:221. Eocene.

Glandieula* 25 :221. Upper Oligocene to Miocene.

Pseudelix* 25:222. Upper Oligocene.

Nesopupa 25 :274. Upper Oligocene to Miocene.

Acmopupa* 26:101. Upper Oligocene.

Negulus 26 :101 ; 27 :231. Upper Eocene to Pliocene.

Aeanthopupa* 27:198. Upper Oligocene.

Central Asia, Transcaiicasus and Turkestan to N. China.

Gastroeopta (Sinalbinula) theeli West. 24:118. Trans-
caucasus.

Pupilla muscorum asiatica IMlldff. 26 :179. Turkestan,
N. China.
P. m. lundstromi West. 26 :179. Turkestan, northward.
P. triplicata inops Eeinh. 26 :180. Transcaucasus.
P. t. luxurians Reinh. 26 :192. Transcaucasus.
P. cupa turkestanica West. 26 :187. Turkestan.
P. c. turcmenia Bttg. 26 :188, 238. N. W. China, etc.
P. (Gibbulinopsis) armeniaca Issel. 26:193. Armenia.
P. (G.) signata Mouss. 26:194. Turkestan, etc.

diecki Gredl. 26 :196.
P. (G.) interrupta Reinh. 26:196. Transcaucasus.
P. (G.) eurina Bens. 26:203. Nepal, Kashmir.
P. (G.) annandalei Pils. 26:202. Upper Burma.
P. (G.) chinensis Hilb. 26:198. Kansu, China.
P. (G.) richthofeni Hilb. 26:198. Kansu.
P. (G.) aeoli Hilb. 26:199. Kansu.
P. (G.) heudeana graudis Mlldff. 26:201. Kansu.
P. pseudocylindrica P. & Y. 28 :136. Aksu, N. China.

Orcula doliolum tereticollis West. 27 :23 ; 28 :133. N. Persia.

Vertigo thibetica Bens. 24 :138 ; 25 :378. Thibet.
V. teilhardi P. & Y. 28 :101. Aksu, N. China.
V. regularis West. 25 :188. Turkestan.

Maritime Provinces of Siberia.

Vertigo denudata Mouss. 25:156. Vladivostok.
V. modesta Say. 25 :123. Kamchatka.
V. krauseana Reinh. 25:136. E.Siberia.
V. alpestris Alder. 25 :197. Vladivostok.

Zoogenetes harpa Say. 27 :196. Kamchatka.

DISTRIBUTION OF PUPILLIDAE. 149

Japan, Korea and China.

North China Pupillids are enumerated above. The fauna
of south China is included here for convenience ; hence such
Oriental Region genera as Hypselostoma and Anauchen.
Boysidia and Pupisomu enter the lower border of the Palae-
arotic Region. The Gastrocopts belong to subgenus Sinal-
iinula. Microstele (28:137) appears to be represented in the
Chinese Pliocene, but is now confined to the Oriental and
Ethiopian Regions. Truncatellina, living in the Riukiu
Islands, is only known as a fossil on the Chinese mainland
(28 :108). Vertigo seems to be rare except in northern Japan.
Ptychalaea, a genus of the European Miocene, is represented
by a single living species on the Bonin Islands.

No Pupillidae are yet known from Fonnosa or Hainan.

Gastrocopta chichijimana Pils. 24 :105. Ogasawara.

G. c. ogasawarana Pils. 24:106. Ogasawara (Bonin) Is.

G. boninensis Pils. 24:107. Ogasawara (Bonin) Is.

G. armigerella Reinli. 24:107. Japan, China.

G. a. hachijoensis Pils. 24:109. Hachijo-jima.

G. coreana Pils. 24:110. Korea.

G. hirasei Pils. 24:110. China.

G. monadicula Heude. 24:111. China.

G. meridionalis Mlldff. 24:111. China.

Hypselostoma insularum Pils. 24 :182 : 27 :211. Riukiu Is.
Anauchen angulinus Gredl. 24:191. China.

Boysidia hunana Gredl. 24 :194. Yangtse Valley.

B. h. conspicua Mlldff. 24 :195. Near Canton.

B. dorsata Anc. 24 :195. Lake Poyang.

B. hangchowensis Pils. & Hir. 24 :196. Prov. Chekiang.

B. strophostoma Mlldff. 24 :197. Guang-dung Province.

B. plicidens Bens. 24 :198. China, Japan.

B. hupeana Gredl. 24 :206. Prov. Hupeh.

Vertigo eogea Pils. 25 :151. Hokkaido.

V. hydrophila Reinh. 25 :152. Hakodate.

V. hachijoensis Pils. 25 :153. Hachijo-jima.

V. kushiroensis Pils. 25 :153. Hokkaido.

V. hirasei Pils. 25 :154. Kiushiu.

V. japonica Pils. & Hir. 25 :155, Okinoerabushima.

V. j. coreana Pils. 25 :156. Korea.

V. j. tosana Pils. 25 :156. Tosa, Shikoku.

150 DISTRIBUTION OP PUPILLIDAE.

Ptyehalaea dedeeora Pils. 25 :158, 273. Ogasawara-jima.
Truncatellina insulivaga Pils. 26 :84. Riukiu Is.
Columella edentula Drap. 27 :242. Saghalien I.

Pupisoma japonica Pils. 26 :25. Hirado, Hizen.
P. harpula Reinh. 27 :198. Tokyo.

Ethiopian Region.

The relations of this Pupillid fauna are mainly with the
Palaearetic Region, but also with the Oriental, and by the
presence of Gastrocopta s. str., Pupoides and Microstele, with
the middle American. The only strongly characterized en-
demic group is Fauxulus, which by conchological features
appears to belong to the Abidinae ; but it is evidently an old
group in South Africa. Afripupilla {Pupilla tetrodus) is
another endemic form, but of uncertain affinities. Gastrocopta
is represented in Abyssinia by one or two species of 8inal'
hinnla (a Palaearetic group), but the tropical and South
African species belong to the typical group of Gastrocopta,
and are closely related to those of South America and the West
Indies, the Mascarene Islands and India. The following
genera appear to be of Palaearetic origin :

Fauxulus Negulus

Orcula Truncatellina

Pupilla Columella

Lauria Acanthinula

Elements common to the Oriental Region are :

Gastrocopta s. str. Pupisoma

Pupoides Nesopupa

Microstele

Two of these, Microstele and Nesopupa, are represented also
in European Miocene.

Gastrocopta pleimesi Jick, 24:119. Abyssinia.
G. haggenmacheri Jick. 24:358. Abyssinia.
G. klunzingeri Jick. 24 :120. Abyssinia.

insidsa Prest. 24 :359.
G. k. senegalensis Malz. 24 :121. Senegal.
G. hermosa Jouss. 24 :121. Eritrea.
G. thomasseti Pils. 28 :71. Natal.

DISTRIBUTION OF PUPILLIDAE. 151

G. damarica Anc. 24 :125, 359. S. and S.-W. Africa.

microbus Dtz. 24 :123 ; 27 :209. Senegal.
G. annobonensis Gir. 24 :124, 27 :208. Anno-Bom.
G. flocculus Mor. 24 :124. Angola.
G. duplicata Prest. 24 :127. Tanganyika Territory.

Fauxulus capensis Kiist. 24 :236. Cape Province.
F. bumupianus Pils. 28 :73. Cape Province.
F. pamphorodon Bs. 24 :240. Cape Province.

(S.-g. Tomigerella)
F. crawfordianus M. & P. 24 :242. Cape Province.
F. layardi Bs. 24 :243. Cape Province.
F. 1. stoaphora Bs. 24 :245. Cape Province.
F. fryanus Bs. 24 :246, 28 :121. Cape Province.

(S.-g. Anisoloma)
F. pereximius M. & P. 24 :247. Cape Province.
F. falconianus Pils. 28 :75. Natal.
F. barnardi Conn. 28 :121. Cape Province.
F. mcbeanianus M. & P. 24 :249. Natal.
F. glanvillianus Anc. 24 :249. Cape Province.
F. g. novenarius Pils. 24 :250. Cape Province.
F. g. darglensis Bp. 24 :250. Natal.
F. g. tomlini Bp. 24 :252. Cape Province.
F. pousonbyanus Mor. 24 :253. Cape Province and Natal.

Pupilla antinorii Pal. 26 :193. Aden.

P. f ontana Kr. 26 :207 ; 27 :255. S. and S. W. Africa.

P. (Afripupilla) tetrodus Bttg. 26 :216. S. Africa.

Lauria bruguierei Jick. 27 :61. Abyssinia.

L. wouramboulchiensis Conn. 28 :86. Abyssinia.

L. desiderata Prest. 27 :62. Mt. Kenia, Tanganyika Terr.

L. bequaerti Pils. 28 :138. Katanga, Belgian Congo.

L. farquhari M. & P. 27 :63. Cape Province.

L. dadion Bs. 27 :64, 259. Cape Prov., Natal.

L. cryptoplax M. & P. 27 :65. Cape Province.

L. tabularis M. & P. 27 :65, Cape Province.

Pupoides coenopictus Hutt. 26 :123. S. Africa.
P. senegalensis Morel. 26 :136. Goree I., Senegal ; Angola.
P. senaariensis Pfr. 26 :136. Eritrea to Senaar ; Egypt.
P. cerealis Palad. 26 :132. Aden.

hoggarensis Pall. 28 :81. Hoggar.
P. reboudi Bgt. 26 :132, Prov. Constantine, Algeria.
P. aethiopicus Bgt. 26 :132. Hamacen Plateau.
P. fabianus Gredl. 26 :133. Soudan.

152 DISTRIBUTION OF PUPILLIDAE.

P. bawriensis Tayl. 26:134. Zanzibar channel.

P. zanguebaricus Tayl. 26 :134, 27 :251. Zanzibar.

P. chanlerensis Prest. 26 :135. Tanganyika Territory.

P. gaziensis Prest. 26 :135. Tanganyika Territory.

P. eonsanguineus Prest. 26 :135. Tanganyika Territory.

P. ealaharicus Bttg. 26 :138. Rhodesia, south and west.

P. minuseulus Mor. 26 :139, 27 :252. S. W. Africa.

P. bryantwalkeri Pils. 27 :251. Cape Province.

P. samavaensis Pal. 26 :128. Aden.

P. maharasicus Bgt. 26 :128. Arabia.

P. vermiformis Pal. 26 :129. Aden.

P. marebiensis Bgt. 26 :129. Arabia.

P. kursiensis Bgt. 26 :130. Aden.

P. ragius Jouss. 26 :130. Aden ; Abyssinia.

Mierostele noltei Bttg. 26 :150. Bechuanaland ; Kalahari.

ohlonga Bttg. 26:150. Damaraland.
M. iredalei Prest. 26 :148. Tanganyika Territory.

Orcula imbricata Jick. 27 :25. Abyssinia, Eritrea.

Nesopupa bisulcata Jick. 25 :339. Abyssinia.
N. b. rhodesiana Pils. 25 :360. Victoria Falls.
N. griqualandica M. & P. 25 :357. S. Africa.
N. farquhari Pils. 25 :358. Cape Province.
N. iota Prest. 25 :361. Tanganyika Territory.
N. corrugata Prest. 25 :361, 28 :110. Victoria Falls.
N. vengoensis Conn. 28 :111. Portuguese E. Afr.
N. bandulana Conn. 27 :226. Portuguese E. Afr.
N. megalomastoma Malz. 25 :362. W. Africa.

Costigo (?) nobrei Gir. 25:368. San Thome.

Truneatellina lardea Jick. 26 :86. Abyssinia.

T. schilleri Jick. 26 :87. Abyssinia.

T. similis Jick. 26 :88. Abyssinia.

T. blanfordi Jick. 26 :89. Abyssinia.

T. naivashensis Prest. 26 :89. Tanganyika Territory.

T. mutaudaensis Prest. 26 :90. Tanganyika Territory.

T. perplexa Bp. 26 :91. South Africa.

T. pretoriensis M. & P. 26 :92. S. Africa.

T. dysorata M. & P. 26 :94. S. Africa.

T. quantula M. & P. 26 :95. S. Africa.

T. iota M. & P. 26 :95. S. Africa.

T. i. livingstonae Bp. 26 :97. S. Africa.

T. sykesi M. & P. 26 :97. S. Africa.

T. s. inconspicua Bp. 26 :98. S. Africa.

DISTRIBUTION OF PUPILLIDAE. 153

Negulus reinhardti Jick. 26 :102. Abyssinia.
N, abyssinicus Jick. 26 :103. Abyssinia.
N. kenianus Prest. 26 :103. Mt. Kenia.

Columella pygmaeonim Pils. & Ckll. 28 :109. Tshibinda,
Belg. Congo.

Pupisoma orcula Bs. 26 :31. S. Africa.

steudneri Jick. 26 :35. Abyssinia.
P. japonica Pils. 26 :25. S. Africa.

Acanthinula peracanthoda Bgt. 27 :194. Abyssinia.
A. expatriata Prest. 27 :194. Mt. Kenia.

Saint Helena.

This island has an African fauna of evidently ancient date.
Lauria cylindracea anconostoma Lwe. 27 :51. Introduced.
Negulus obliquecostatus Sm. 26 :104.
Nesopupa (Helenopupa) turtoni Sm. 25:363.
Campolaemus perexilis Sm. 25 :365.

Mascarene Islands, Madagascar, Comoros.

The Gastrocopts and Nesopupae are related to those of both
the Ethiopian and Oriental Regions. Pupilla and Lauria to
the Ethiopian forms.

Gastrocopta seignaciana C. & F. 24 :128, 27 :208. Comoro Is.

G. tripunctum Mor. 24 :130. Comoro Is.

G. microscopica Mts. 24 :131. Seychelles.

G. lienardiana Cr. 24 :132. Rodriguez.

G. 1. eudeli Pils. 24:133. Reunion.

G. madagascariensis B. & S. 24 :210. Madagascar.

Pupilla pupula Desh. 26 :214, 28 :86. Reunion.

Lauria bourbonensis Pils. 27 :62. Reunion.

Nesopupa gonioplax Pils. 25 :351. Mauritius.

N. micra Pils. 25 :351. Mauritius.

N. minutalis Mor. 25 :352, 27 :228. Comoro Is.

N. rodriguezensis Conn. 28 :110. Rodriguez.

N. comorensis Pils. 25 :353. Comoro Is.

N. incerta Nev. 25 :353. Reunion.

N. ventricosa Ad. 25 :354. Mauritius.

N. praslinensis Nev. 25 :355. Seychelles.

154 DISTRIBUTION OF PUPILLIDAE.

Costigo ( ?) borbonica H. Ad. 25:367. Reunion.
C. ( ?) desmazuresi Cr. 25:368. Rodriguez.

Oriental Region — India and Ceylon.

Related to the Ethiopian Region by Gastrocopta s. sir.,
Pupoides, Microstele and Nesopupa, and to the Indo-Chinese
fauna by the presence of Sinalhinula and Boysidia. Trun-
catellina and Columella are Palaearctic elements barely enter-
ing the Oriental borders, but included here for convenience.

Gastrocopta mimula Bens. 24 :134. Ceylon.
G. serrula Bens. 24 :135. India.
G. (Sinalbinula) bathyodon Bens. 24:136. India.
G. (S.) huttonianaPfr. 24:137. India.

Boysidia plicidens Bens. 24 :198. India etc.

B. landurensis Pils. 24:204. India.

Pupilla ( ?) eurina Bens. 26 :203. Nepal, Kashmir.

P. ( ?) seriola Bens. 26:204. Cuttack, Darjiling etc.

P. (?) diopsis Bens. 26 :205. Nerbudda Valley.

Pupoides coenopictus Hutt. 26 :123. India.

P. lardea Pfr. 26 :126, 237. India.

P. karachiensis Peile. 28 :80. India.

P. doriae Issel. 26:122. Persia.

P. tutulus Rve. 26 :122. India.

Microstele muscerda Bens. 26 :148. Ceylon, India ?

Nesopupa barrackporensis Gude. 25 :348. India.

N. brevicostis Bens. 25 :346. Bengal.

N. salemensis Blf . 25 :347. Kalryen Hills.

N. cinghalensis Gude. 25 :347. Ceylon.

Pupisoma evezardi Blf. 26 :24, 235. Bor Ghat.

P. miccyla Bens. 26 :26. Ceylon.

P. longstaffi G.-A. 26 :27. Ceylon.

P. eacharicum G,-A. 26 :29. Silchar.

P. orcula Bens. 26 :31. India.

Boysia boysii Pfr. 26 :226. India.

Truncatellina hiraalayana Bens. 26 :71. Simla etc.

Columella giitta Bens. 27 :241. Spiti Valley.

Burma, Indo-China, Malay Peninsula.

The Gastrocoptas belong to the subgenus Sinalbinula. The
extraordinary development of endemic genera of several col-

DISTRIBUTION OF PUPILLIDAE. 155

lateral lines of Gastrocoptinae characterizes this and the
adjacent insular subregion. Pupisoma is abundant, but ex-
cept for Nesopupa, the Ethiopian genera are lacking.

Gastrocopta ejecta Bav. & Dautz. 24:111. Bay of Along.
G. avanica Bens. 24 :138. Burma.
G. palmira Stol. 24 :139. Penang.

Hypselostoma tubiferum Bens. 24:178. Burma.

H. annamiticum Mlldff. 24 :178. Annam.

H. a. altius Pils. 24:181.

H. laidlawi Cllge. 24 :181. Malay Peninsula.

Anauchen messageri Bav. & Dautz. 24:189. Tonkin.
A. gereti Bav. & Dautz. 24 :189. Tonkin.

A. rochebruni Mab. 24 :190. Tonkin.

Boysidia robusta Bav, & Dautz. 24 :201. French Indo-China.

B. lamothei Bav. & Dautz. 24 :202. French Indo-China.
B. paviei Bav. & Dautz. 24 :203. French Indo-China.
B. dayana Stol. 24 :205. Burma.

B. salwiniana Theob. 24:206. Burma.

B. kelantanensis Sykes. 24:208. Malay Peninsula.

Gyliotrachela ^ bensoniana Blf . 24:211. Ava.

G. hungerfordiana Mlldff. 24 :212. Perak.

G. transitans Mlldff. 24 :214. Samui Is.

G. striolata Mlldff. 24 :215. Samui Is.

G. crossei Mori. 24 :215. Tonkin.

G. c. endodonta Mlldff. 24 :217. Tonkin.

G. c. brevituba Mlldff. 24 :217.

Systenostoma pulverea Bav. & Dautz. 24 :225. Tonkin.
S. pauperrima Bav. & Dautz. 24 :225. Tonkin.
S. defixa Bav. & Dautz. 24 :226. Tonkin.

Pupilla annandalei Pils. 26 :202. Burma ?
Nesopupa filosa Theob. & Stol. 25 :345. Burma.

Pupisoma lignicola Stol. 26 :23. Burma,

P. 1. unidentata G.-A. 26 :24, 235.

P. constrictum G.-A. 26 :28. Andaman Is.

P. orcella Stol. 26 :29. Penang.

P. orcula Bens. 26 :33. Generally distributed.

1 Mr. Tomlin evidently meant to write " Gyliotrachea ". By a pen or
typographic error the name was so spelled in this volume, p. 73.

156 DISTRIBUTION OF PUPILLIDAE.

Indo-Malayan Islands, Philippines, Java, Borneo, etc.

These islands are faunally similar to the foregoing mainland
area. Aidacospira and the gastroeopt subgenus Cavipupa are
endemic.

Gastrocopta servilis Gld. Introduced in Philippines.

lyonsiana Anc. 24 :141.
G. (Sinalbinula) capillacea Dhn. 24:144. Philippines.
G. (S.) pediculus Shuttl. 24:145.
G. (S.) p. ovatula Mlldff. 24:149. Philippines.
G. (S.) niobe Fult. {moelloidorffiana Pils.). 28:119.
Philippines.

G. (Cavipupa) euryomphala Pils. 28:120. Philippines.

Boysidia boettgeri ]\Illdff. 24 :208. Java.

Hypselostoma luzonicum Mlldff. 24:183. Philippines.

H. 1. major Mlldff. 24 :360. Philippines.

H. 1. imbricata JVDldff. 24 :360. Philippines.

H. sibuyanicum Mlldff. 24 :184. Philippines.

H. polyodon Mlldff. 24 :185. Philippines.

H. quadrasi Mlldff. 24 :185, 360. Philippines.

H. pusillum Mlldff. 24 :186. Philippines.

H. roebelini Mlldff. 24 :186. Philippines.

H. edentulum Mlldff. 24 :187. Philippines.

Gyliotrachel a fruhstorferi Mlldff. 24:207. Java.
G. everetti Smith. 24 :218. Kalao I.

Aulaeospira mueronata IVUldff. 24 :222. Cebu.

A. hololoma ]\01dff. 24 :222. Cebu.

A. porrecta Hid. 24 :222. Ilin.

A. triptycha Q. & M. 24 :223. Masbate.

A. scalatella Mlldff. 24 :223. Luzon.

A. rhombostoma Mlldff. 24 :223. Tablas.

A. (Pseudostreptaxis) azpeitiae Hid. 24:224. Busuanga.

Nesopupa moreleti Bm. 25 :339. Borneo.

N. moellendorflfi Bttg. 25 :341. Philippines.

N. nannodes Mlldff. 25 :341. Philippines.

N. malayana Iss. 25 :342. Philippines, Borneo.

N. proscripta Sm. 25 :344. Christmas I.

Costigo calamianica Mlldff, 25 :367. Philippines.

Pupisoma orcula Bens. 26 :31. Java.
P. pulvisculum Iss. 26 :30. Borneo.
"Acanthinula" perpusilla Mlldff. 27:195. Java.
" A " tiluana Mlldff. 27 :195. Java.

DISTRIBUTION OF PUPILLIDAE. 157

Austro-Malayan and Melanesian Islands.

Gastrocopta (Sinalbinula) pediculus Sh. 24:145.
G. (S.) dahli Thiele. 28:72. Bismarck Archipelago.
G. (S.) semiclausa Thiele. 28:73. Bismarck Archipelago.
G. (S?) microsoma T.-C. 24:152, 27:209. Am Is.
G. (S.) recondita T.-C. 24:153, 27:209. Am Is.
G. (S?) neocaledonica Pils. 24:154. New Caledonia.
G. (S.) macdonnelli macrodon Pils. 24:164. New Guinea.
G. (S.) niobe Fult. 24:153, 28:119. Tenimber Is.
moellendorffiana Pils. 24 :145. Bohol, Philippines.

Gyliotrachela dohertyi Fult. 24 :219. Tenimber Is.

Nesopupa novopommerana Rensch. 28:112. Bismarck
Archip.

N. quadrasi transaequatorialis Rch. 28:113. Bismarck
Archip.

N. moluccana Bttg. 25 :338. Moluccas.

N. selebensis T.-C. 25 :343. Celebes.

N. norfolkensis Sykes. 25 :333 ; 27 :227. Norfolk I.

N. lif ouana Gass. 25 :333. Lifu, Loyalty Is.

N. mariei Crosse. 25 :334. New Caledonia.

Costigo saparuana Bttg. 25 :366. Moluccas.

Cylindrovertilla fabreana Crse. 26 :47. New Caledonia.
C. paitensis Crse. 26 :48. New Caledonia.

Pupisoma vimontiana Crse. 26 :35. New Caledonia.
(=P. orculaBs. ?).

Pupillidae of Australia and Tasmania.

Australia is not rich in Pupillidae. Of the six genera rep-
resented, Pupisoma is apparently a recent importation on
plants. Cylindroveriilla, elsewhere found only in New Cale-
donia, and Gyliotrachela, a genus of the Malay Peninsula and
East Indies to the Tenimber islands, are confined to tropical
Queensland. Gastrocopta, Pupilla and Pupoides are widely
distributed.

Gastrocopta is represented by species of the subgenus Sinal-
bimda, a Palaearctic group which extends from Japan and
China through the tropics to Australia, and by Australhinida,
special to Australia, but evidently a derivative of Sinalbinula.

158 DISTRIBUTION OF PUPILLIDAE.

A key to the species of Gastrocopta may be found iu vol. 24,
p. 155.

Pupoides appears iu several species not differing materially
from the typical forms of the northern continents. Glypto-
pupoides, referred to Pupoides as a subgenus, is endemic in
Australia.

Pupilla is represented by species of Gihhulinopsis, also
traceable to the Palaearctic Region, but like Pupoides it is
here very widely separated from its present-day range in the
other continents, as neither genus has occurred in the East
Indies. These genera are doubtless emigrants from Asia
which reached Australia by a chain of early Tertiary con-
nections through the East Indies, where they appear to have
become extinct.

'^Pupa'" scotti Braz. of Fitzroy Island, is not sufficiently
known for generic reference.

Gastrocopta pediculus Sh. 24 :158. Eastern Australia.

rossiteri Braz. 24 :159. New South Wales.

hannertonensis Gabriel. 28:118. Victoria.
G. p. queenslandica Pils. 24 :159. Queensland.
G. strangei Pfr. 24 :157. N. S. Wales.
G. mooreana Smith. 24 :160, 26 :230. W. Australia.
G. macdonnelli Braz. 24 :162. N. E. Australia.
G. tatei Pils. 24 :165. Central Australia.
G. (Australbinula) hedleyi Pils. 24:166. Queensland.
G. (A.) mussoni Pils. 24:167. Queensland.
G. (A.) laripinta Tate. 24:168. Central Australia.
G. 1. deserti Pils. 24 :170. Central Australia.
G. (A.) wallabyensis Sm. 24:171. W.Australia.

Gyliotrachela australis Ohdner. 26 :232. Queensland.

Pupilla australis Angas. 26 :218. Australia, Tasmania.
P. ficulnea Tate. 26 :221. Central Australia.

Pupoides adelaidae A. & A. 26 :140. N. S. W. ; S. Australia.
P. pacificus Pfr. 26 :141.

f . sinistralis Pils. 26 :144.
P. contrarius Smith. 26 :144. W. Australia.
P. c. eremicola Tate. 26 :145. Central Australia.
P. beltianus Tate. 26 :145. Central Australia.
P. myoporinae Tate. 26 :146. S. Australia.

DISTRIBUTION OP PUPILLIDAE. 159

P. ischnus Tate. 26 :146. Central Australia.

P. (Glyptopupoides) hedleyi Pils. 27:252. Queensland.

Cylindrovertilla kingi Cox. 26 :44. Queensland.
C. hedleyi Pils. 26 :46. Queensland.
C. fabreana Crosse. 26 :47. Queensland.

Pupisoma orcula Bens. 26 :31, 34. Queensland.
circumlitum Hedley.

' ' Pupa ' ' scotti Braz. 26 :222. Fitzroy I.

Polynesia and Micronesia.

The Gastroeoptas of this region are apparently all due
to importation within the period of human occupation.
Probably Pupoidopsis, which lives near the sea at low ele-
vations, was also carried about by the Polynesians. The
Nesopupae are endemic.

Gastrocopta servilis Gld. Introduced.

lyonsiana Anc. 24:141.
G. (Sinalbinula) pediculus Shuttl. 24:145.

Nesopupa tantilla Gld. 25 :324. Society Is.

N. pleurophora Shuttl. 25 :326. Tahiti, Marquesas ?

N. armata Pse. 25 :327, 28 :112. Society Is. ; Rarotonga.

N. paivae Crse. 25 :328. Gambler Is. ; Taumotus.

N. dentifera Pse. 25 :329. Aitutaki, Cook Group.

N. godeffroyi Bttg. 25 :330. Samoa.

N. tongana Bttg. 25 :331. Tonga Is.

N. vitiana Bttg. 25 :332. Fiji Is.

N. quadrasi Mlldff. 25 :335. Guam.

N. eapensis Bttg. 25 : 335. Yap, Carolines.

N. ponapica Mlldff. 25 :336. Ponape.

Pronesopupa senex Iredale. 26 :1. Kermadec Is.
P. simplaria Pse. 26 :2. Marquesas.

Pupoidopsis hawaiensis P. & C. 26:107. Tuamotus;
Christmas I.

Hawaiian Islands.
As in Polynesia, the species of Gastrocopta are doubtless
due to importation, first {G. pediculus) by the Polynesians,
who probably brought Pupisoma also, and later by the Euro-
peans (6r. servilis). The genus Lyropupa and several sub-
genera of Nesopiipa and Pronesopupa are endemic. All of

160 DISTRIBUTION OP PUPILLIDAE.

the species of these groups are special to these islands. It
is not likely that the Hawaiian Pronesopupae are directly-
related to the type of that genus ; they are more likely to be
a parallel evolution from Nesopupa. For subgeneric classi-
fication of Nesopupa and Pronesopupa see 25:275.

Species referred to Columella should be dissected, as that
reference appears improbable, though the shells are certainly
very similar.

Gastrocopta servilis Gld. Introduced.

lyonsiana Anc. 24 :141. Oahu.
G. (Sinalbinula) pediculusnacca Gld. 24:149. Introduced.

Pupoidopsis hawaiensis Pils. & Cooke. 26:107. Kauai;
Oahu ; Molokai ; W. Maui.

Lyropupa lyrata Gld. 25 :233. Oahu.

L. 1. uncifera P. & C. 25 :236. Oahu.

L. 1. fossilis C. & P. 25 :237. Oahu.

L. microthauma Anc. 25 :238, Oahu.

L. rhabdota C. & P. 25 :239. Molokai.

L. r. pleuris P. & C. 25 :240. Molokai.

L. r. lanaiensis Cooke. 25 :241. Lanai.

L. r. baldwiniaua Cooke. 25 :241. W. Maui.

L. thaanumi C. & P. 25 :242. E. Maui.

L. prisca Anc. 25 :243. Hawaii.

L. striatula Pse. 25 :246, 27 :223. Hawaii.

L. clathratula Anc. 25 :245. Hawaii.

L, truncata Cooke. 25 :247. Hawaii.

L. spaldingi P. & C. 25 :248. Oahu.

L. mirabilis Anc. 25 :249. Oahu.

L. antiqua C. & P. 25 :250. Oahu.

L. hawaiiensis Anc. 25 :251. Hawaii.

L. spama C. & P. 25 :252. Molokai.

L. s. sinulifera P. & C. 25 :253. Molokai.

L, anceyana C. & P. 25 :253. Hawaii.

L. scabra P. & C. 25 :253. Maui.

L. costata Pease. 25 :272 ; 27 :223. Hawaii, Kahoolawe.

kahoolawensis P. &. C. 25 :256.
L. e. puukolekolensis P. & C. 25 :258. Molokai.
L. perlonga Pse. 25 :258. Oahu.
L. p. eylindrata P. & C. 25 :261. Oahu.
L. p. interrupta P. & C. 25 :261. Oahu.
L. p. filocostata C. & P. 25 :262. Kauai.
L. micra C. & P. 25 :263. Oahu.

STROBILOPSID^, PUPILLID^

PLATE 16

PUPILLID^

PLATE 17

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PUPILLID^

PLATE 11

PUPILLIDiE

PLATE 19

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PLATE 21

PUPILLID^E

PLATE 22

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PUPILLID^

PLATE 23

DISTRIBUTION OF PUPILLIDAE. 161

L. m. percostata P. & C. 25 :264. Oahu.

L. m. maunaloae P. & C. 25 :264. Oahu, Molokai.

L. ovatula C. & P. 25 :265. Oahu.

L. 0. kona P. & C. 25 :266. Hawaii.

L. o. moomomiensis P. & C. 27 :225. Molokai.

L. plagioptyx P. & C. 25 :267. Oahu.

L. cyrta C. & P. 25 :268. Hawaii.

L. cubana Dall. 25 :268. Kauai ?

L. thaumasia C. & P. 25 :270. Kauai.

Nesopupa plicifera Ane. 25 :280. Oahu.

N. waianaensis C. & P. 25 :281. Oahu.

N. litoralis C. & P. 25 :283. Oahu.

N. dispersa C. & P. 25 :284. Oahu to Hawaii.

N. baldwini Anc. 25 :287. Molokai, Maui, Hawaii.

N. b. subcostata P. & C. 25 :288. Molokai.

N. b. lanaiensis P. & C. 25 :289. Lanai.

N. limatula C. & P. 25 :290. Maui.

N. dubitabilis C. & P. 25 :291. Molokai.

N. d. kaalaensis C. & P. 25 :292. Oahu.

N. anceyana C. & P. 25 :293. Hawaii.

N. subcentralis C. & P. 25 :294. Hawaii.

N. bishopi C. & P. 25 :296. E. Maui.

N. forbesi C. & P. 25:297. Hawaii.

N. infrequens C. & P. 25 :298. Kauai.

N. wesleyana Ane. 25 :299. Oahu to Hawaii.

N. w. gouveiae C. & P. 25 :301. Hawaii.

N. w. triphera C. & P. 25 :301. Kauai, Oahu, Molokai.

N. w. rhadina C. & P. 25 :301. Oahu, Molokai, Lanai.

N.w. kamaloensis P. & C. 25:303. Molokai.

N. thaanumi Anc. 25 :304. Oahu, Lanai, Maui, Hawaii.

N. newcombi Pf r. 25 :307. Kauai to Hawaii.

N. n. angusta P. & C. 25 :315. Kauai.

N. n. interrupta C. & P. 25 :315. Hawaii, Oahu.

N. n. disjuncta C. & P. 25 :317. Oahu.

N. n. gnampta C. & P. 25 :317. Oahu, Kauai.

N. oahuensis C. & P. 25 :317. Oahu.

N. kauaiensis Anc. 25 :318. Kauai.

N. singularis C. & P. 25 :320. Oahu, Maui.

N. alloia C. & P. 25 :321. Kauai.

Pronesopupa acanthinula Anc. 26 :5. Oahu to Hawaii.

P. hystricella C. & P. 26 :7. Hawaii to Kauai.

P. boettgeri C. & P. 26 :8. Oahu.

P. b. spinigera C. & P. 26 :10. Kauai to Hawaii.

162

DISTRIBUTION OF PUPILLIDAE.

P. admodesta Migh, 26 :11. Oahu to Hawaii.
P. f rondicola C. & P. 26 :13. East Maui.
P. f . eorticieola C. & P. 26 :14. E. Maui.
P. molokaiensis C. & P. 26 :15. Molokai.
P. ineerta C. & P. 26-16. Kauai.
P. sericata C. & P. 26 :17. Hawaii.
P. lymaniana C. & P. 26 :18. Hawaii.
P. orycta C. & P. 26 :18. Hawaii.

Pupisoma orcula Bens. 27 :228 ; 28 :116. Introduced Oahu ;
also Makatea, 28 :116.

Columella sharpi Pils. & Cooke. 27 :247. Hawaii.

C. olaaensis P. & C. 27 :248. Hawaii.

C. alexanderi C. & P. 27 :248. West Maui.

NEiVRCTIC AND NEOTROPICAL REGIONS.

Though there are a good many species, the Americas are
rather poor in genera of Pupillidae, and few of them are
endemic. In the following list the names of subgenera are
italicized.

Nearctic Region

Neotropical Region

Temperate N. Am. Mexico, C. Am. West Indies

Chaenaxis

Gastrooopta s. sir. Gastrooopta s. s<r. Gastrooopta s. sfr.
Immersidens Immersidens

Alb inula

Vertigopsis

Privatula
Pupilla s. str.

Striopupilla
Pupoides s. str.

Ischnopupoides
Vertigo s. str.

Vertillopsis

Angustula
Columella
Sterkia s. str.^

MetasterJcia

Pupisoma i
Zoogenetes

Albinula

Pupilla (?)

Pupoides s. str.

Ischnopupoides
Vertigo s. str.

Angustula
Columella

Pupoides s. str.

Vertigo s. str.

Angustula

MetasterJcia
Bothriopupa
Nesopupa (Cocosl.)
Pupisoma Pupisoma

Metasterkia
Bothriopupa

South America
Gibbulina

Gastrooopta s. str..
Immersidens
Geminidens

Pupoides s. str.
Ischnopupoides

MetasterJcia
Bothriopupa

Phipisoma

1 These genera occur only in the extreme southern edge of the Nearctie
Realm and belong properly to the Neotropical.

DISTRIBUTION OF PUPILLIDAE. 163

Both genera and species are most numerous in the North
Temperate Zone; going south the northern genera gradually
drop out, until in Argentina only Gastrocopta remains ; how-
ever, the South American Pupillidae are quite imperfectly
known. In the tropics several genera are added, Bothriopupa,
Pupisoma and Sterkia, the last two barely entering the lower
border of the Nearctic Realm. The Cocos Island species re-
ferred to Nesopupa seems to be a stray from Polynesia, but it
needs anatomic comparison with Bothriopupa.

The distribution shown in the table above apparently indi-
cates successive invasions from Asia, which spread southward
to distances determined by the date of introduction and the
adaptability of the stocks. The gastrocopts and early verti-
gines apparently came in not later than Oligocene. The
genera which presumably came later, such as Pupilla s. str.,
Vertigo s. str., Columella and Zoogenetes, differ but little
from the old world forms or are specifically identical, and
have made the least progress southward.

The only endemic American genera are GihhuUna, ChaenaxiSy
Sterkia and Bothriopupa. Endemic subgenera of Gastrocopta
are: Immersidens, Vertigopsis, Privatula and Geminidens.
Pupilla has the endemic subgenus Striopupilla ; Pupoides the
subgenus Ischnopupoides, and Vertigo the subgenera Vertil-
lopsis and Angustula. Of the endemic genera, Chaenaxis and
probably Gihhulina are derivatives of the Gastrocopta s. str.
stock. I suspect that Immerside^is came from the same stock,
but pending anatomic evidence or other significant data,
its place in the series remains uncertain. The folicolous
Bothriopupa has the appearance of an Americanized Neso-
pupa, and from Dr. H. B. Baker's observations, agrees with
that group anatomically. The terrestrial Sterkia may be
presumed to be derived from some ancestral N esopupa-\\kQ
group which reached America from Asia, probably in Paleo-
cene times.

Gastrocopta s. str., Pupoides and Pupisoma are common to
the tropics of both hemispheres. Pupoides appeared in a
practically typical species (vol. 27, p. 248) in the Upper
Oligocene (Tampa silex bed) of Florida (at that time insular.

164 DISTRIBUTION OF PUPILLIDAE.

with an Antillean fauna). Microcerion (vol. 26, p. 151) from
the same formation, is, in my opinion, a specialized tangent
from Microstele, with which it agrees in all essential char-
acters. The question arises, whether Gastrocopta s. str.,
Pupoides and Microcerion are traceable to Africa, over a
translantic bridge which is believed to have existed in the
Mesozoic and possibly later,^ or whether they formed an
early migration by the northern route.

United States northward; Lower California.

Chaenaxis tuba P. & F. 24:2. Arizona.
C. intuscostata Clapp. 24:3. Arizona.
C. i. brevicostata Pils. 24 :4. Arizona.

Gastrocopta rupicola Say. 24 :58. Southeastern U. S.

G. r. matacumbensis Pils. 24 :6. Florida Keys,

G. procera Gld. 24 :62. Eastern.

G. p. sterkiana Pils. 25 :357 ; 27 :207. S. Dakota to Texas.

duplicata Sterki. 24 :65.
G. p. mcclungi H. & J. 24 :66. S. Dakota south.
G. cristata Pils. & Van. 24 :68. Texas to Arizona.
G. riograndensis Sterki. 24 :69. Texas.
G. pellucida Pf r. 24 :75. Southern States.
G. p. hordeacella Pils. 24 :78 ; 28 :117. Southern States.
G. p. parvidens Sterki. 24 :80, 27 :207. Arizona.
G. quadridens Pils. 24 :57. Arizona, N. M.
G. (Albinula) armifera Say. 24:15. Eastern States.
G. (A.) a. similis Sterki. 24:17.
G. (A.) a. affinis Sterki. 24:18.
G. (A.) a. abbreviata Sterki. 24:18.
G. (A.) a. ruidosensis Ckll. 24:19. New Mexico.
G. (A.) a. clappi Sterki. 24:20.

G. (A.) contracta Say. 24:22. Eastern U. S. and Mexico.
G. (A.) c. peninsularis Pils. 24:24. Florida.
G. (A.) c. climeana Van. 24:24. Southern States.
G. (A.) holzingeri Sterki. 24:25. Eastern States.
G. (A.) h. agna P. & V. 24 :26. Kansas.
G. (Vertigopsis) pentodon Say. 24:28. Eastern States.
G. (V.) p. gracilis Sterki. 24:31.
G. (V.) tappaniana C. B. Ad. 24:33. Eastern States.
G. (V.) carnegiei Sterki. 24:35. Ohio.

1 CA Schuchert, "Gondwana Land Bridges", Bull. Geol. Soc. of
America, vol. 43, p. 875. 1932.

DISTRIBUTION OP PUPILLIDAE. 165

G. ( V.) pilsbryana Sterki. 24 -M, 28 :64. N. M. to Utah.

sto7ieri C. & J. 28 :64.
G. (V.) p. amissidens Pils. 28:118. N. Arizona.
G. (Immersidens) perversa Sterki. 24:40. Arizona.
G. (I.)p. sanaP. &F. 24:35.
G. (I.) p. minor Sterki. 24:44.
G. (I.) ashmuni Sterki. 24:41. Arizona.
G. (I.) a. imperfecta P. & F. 28 :117.
G. (I.) cochisensis P. & F. 24:45. Arizona.
G. (I.) oligobasodon P. & F. 24:46. Arizona, New Mexico.
G. (I.) dalliana Sterki. 24:49. Arizona.
G. (I.) d. media Pils. 24:50. Arizona.
G. (I.) bilamellata Sterki & Clapp. 24:51. Arizona.
G. (I.) rixfordi Hanna. 27 :207. Montserrate I., Gulf of Cal.
G. (Privatula) corticaria Say. 24:52. Eastern States.

Pupilla muscorum L. 26 :156. Northern States.
P. blandi Mse. 26 :159. Mountain States.
P. b. charlestonensis Pils. 26 :163. Nevada.
P. sonorana Sterki. 26 :163. Arizona.
P. hebes Anc. 26 :164. Mountain States.
P. h. nefas P. & F. 26 :166. Arizona.
P. syngenes Pils. 26 :167. Mountain States.
P. s. dextroversa P. & V. 26 :169.
P. s. avus P. & F. 26 :171. Arizona.

P. (Striopupilla) sterkiana Pils. 26:156; 28:82. L. Cal.
P. (S.) goniodon Pils. 28:82. Guadalupe I., L. Cal.
P. (S.) guadalupensis Pils. 28:83. Guadalupe I., L. Cal.
Pupoides marginatus Say. 26 :111, 27 :249, 28 :80.
P. modicus Gld. 26 :115. S. Florida.
P. catalinensis Hanna. 27 :249. L. California.
P. (Ischnopupoides) hordaceusGabb. Arizona, New Mexico.
eupleura C. & B. 28 :138. Utah.

Vertigo rugosula Sterki. 25:77. Southeastern States.

V. r. oralis Sterki. 25 :78.

V. alabamensis Clapp. 25 :79. Alabama.

V. a. conecuhensis Clapp, 25 :80.

V. morsei Sterki. 25 :81. Northeastern States.

V. ovata Say. 25 :82, 372. Eastern and southern States.

V. 0. diaboli Pils. 25 :88. Texas.

V. o. mariposa Pils. 25 :88. California.

V. 0. santodomingensis Pils. 28 :98.

V. berryi Pils. 25 :89. California.

V. binneyana Sterki. 25 :90. Rocky Mts.

V. ventricosa Morse. 25 :94, 372. Northern States.

166 DISTRIBUTION OF PUPILLIDAE.

V. elatior Sterki. 25 :95, 28 :93. Northern States.

V. pygmaea Drap. 25 :96. Northern States.

V. gouldii Binn. 25 :98. Northern U. S. and Canada.

V. g. hubriehti Pils. 28 :99. Missouri loess.

V. g. paradoxa Sterki. 25 :99, 28 :95. Northern.

V. g. cristata Sterki. 25 :100. Canada.

V. g. eoloradensis Ckll. 25 :115. Colorado, Utah, Arizona.

V. g. basidens P. & V. 25 :117. Rocky Mts.

V. g. arizonensis P. & V. 25 :117. Arizona, New Mexico.

V. g. inserta Pils. 25 :118. Arizona.

V. nylanderi Sterki. 25 :100. Maine.

V. bollesiana Morse. 25 :101. Northeast.

V. hebardi Van. 25 :103, 372. Plorida.

V. perryi Sterki. 25 :103. Mass. and Rhode Island.

V. parvula Sterki. 25 :105. Ohio.

V. tridentata Wolf. 25 :106. Northern States.

V. Columbiana Sterki. 25 :108, 28 :97. Northwestern States.

V. andrusiana Pils. 25 :109. Northwestern States.

V. a sanbernardinensis Pils. 25 :111. Southern California.

V. idahoensis Pils. 28 :100. Idaho.

V. sterkii Pils. 25 :112. California.

V. hannai Pils. 25 :114. Kansas, loess.

V. wheeleri Pils. 28 :96. Alabama.

V. concinnula Ckll. 25 :119. Rocky Mts.

V. arthuri v. Marts. 25 :121. North Dakota.

V. modesta Say. 25 :123, 372. N. States to Alaska.

V. m. ultima Pils. 25 :128. Alaska.

V. m. parietalis Anc. 25 :128. Rocky Mts.

V. m. corpulenta Morse. 25 :130. Rocky Mts.

V. m. insculpta Pils. 25 :131. Arizona, New Mexico.

V. m. castanea Sterki. 25 :132. California.

V. m. microphasma Berry. 25 :376. S. California.

V. m. sculptilis Pils. 28 :100. Montana, Oregon.

V. occidentalis Sterki. 25 :134. S. California.

V. allyniana Berry. 25 :376. S. California.

V. hoppii Moeller. 25 :135, 27 :216. Greenland.

V. dalliana Sterki. 25 :137. California.

V. californica Rowell. 25 :139. California.

V. c. trinotata Sterki. 25 :140. California.

V. c. diegoensis Sterki. 25 :141. S. California.

V. c. Cyclops Sterki. 25 :141. California.

V. c. longa Pils. 25 :377. California.

elongata Sterki. 25 :142.
V. c. catalinaria Sterki. 25 :142, 28 :97. Catalina I. south.
V. c. guadalupensis Pils. 28 :97. Guadalupe I., L. Cal.

DISTRIBUTION OF PUPILLIDAE. 167

V. e. cupressicola Sterki. 25 :143. Monterey, Cal.

V. rowelli Newc. 25 :143, 378. Cal., Oregon.

V. degeneris Pils. 28 :98. Guadalupe Is., L. Cal.

V. (Vertillopsis) oscariana Sterki. 25:144, 378. S. States.

V. (V.) hinkleyi Pils. 26:234. Arizona.

V. (Angustula) milium Gld. 25:146, 378. Eastern States.

Columella edentula Drp. 27 :242. Northern States.

C. alticola Ingersoll. 27 :243. Rocky Mts.

C. hasta Hanna. 27 :245. Pleistocene, Kansas.

Sterkia rhoadsi Pils. 26 :52. Florida.
S. Clementina Sterki. 26 :54. S. California.
S. hemphilli Sterki. 26 :55. S. California.
S. calamitosa Pils. 26 :57. L. California.
S. c. martiniana Pils. 28 :110. L. California.

Bothriopupa variolosa Gld. 24 :230. S. Florida.

Pupisoma dioscoricola C. B. Ad. 26:37, 235. S. Florida,

Texas.
P. minus Pils. 26 :40. Florida.
P. macneilli Clapp. 26 :41. S. Alabama.

Zoogenetes harpa Say. 27 :196. Northeastern and Canada.
Fupillidae of Mexico and Central America.

Gastrocopta servilis Gld, 24 :70. Mexico to Panama.

G. pellueida Pf r. 24 :75. Eastern Mexico.

G. (Immersidens) prototypus Pils. 24:47. Mexico.

G. (I.) p. basidentata Pils. 24:48. Guatemala.

G. (Albinula) contracta Say. 24:22. Eastern Mexico.

Pup ilia (?) oerstedi Moerch. 28:82. Nicaragua.

Pupoides marginatus Say. 26 :111. N. E. Mexico.

P. (Ischnopupoides) chordatus Pfr. 26:119. W.Mexico.

Sterkia eyriesii Drt. 26 :51. Guatemala.

S. bakeri Pils. 26 :236. Southern Vera Cruz.

Vertigo ovata Say. 25 :82. E. Mexico.

V. (Angustula) milium Gld. 25:146. E. Mexico.

Columella polvonensis Pils. 27 :246. Nicaragua.

Nesopupa cocosensis Dall. 25 :323. Cocos I., C. R.

Bothriopupa breviconus Pils. 24 :230. Guatemala.
B. leucodon Morel. 24 :232. Guatemala.

Pupisoma dioscoricola C. B. Ad. 26 :37. Yucatan.
P. minus Pils. 26 :40. Guatemala.

168 DISTRIBUTION OF PUPILLIDAE.

P. comicolense H. B. Bkr. 28 :115. State of Puebla.
P. mediamericanum Pils. 26 :42. E. Mexico, Guatemala.
P. miehoacanense Pils. 26 :40. Michoacan, Mex.
P. bailyi Pils. 28 :116. Cuernavaca, Mexico.

West Indies and Bermuda.

Gastrocopta rupicola Say. 24 :57. Bahamas, Bermuda.

G. r. duplex Pils. 24 :60. Bermuda.

G. r. marginalba Pf r. 24 :60. Cuba, etc.

G. servilis Gld. 24 :70. Generally distributed.

G. s. riisei Pf r. 24 :74.

G. pellucida Pf r. 24 :75. Generally distributed.

G. p. biminiensis Pils. 24 :78. Bahamas.

G. p. delicata Pils. 24 :78. Bahamas.

G. barbadensis Pf r. 24 :83. Generally distributed.

G. b. crassilabris Pfr. 24 :85. Bahamas, Bermuda.

G. b. hojeda Pils. 27 :206. Curacao.

G. polyptyx Pils. 24 :89. St. Thomas to Bermuda.

G. curacoana Pils. 27 :202. Curacao.

longurio Crosse. 24 :82.
G. octonaria Pils. 27 :204. Curacao.

Pupoides marginata Say. 26 :111.

P. m. nitidula Pfr. 26 :113 ; 27 :250.

P. coenopicta Hutt. 26 :123. Cuba, Porto Rico.

Vertigo marki Gul. 25 :107. Bermuda.

"V. numellata Gul. 25 :91. Bermuda.

V. ovata Say. 25 :82.

V. torrei Aguayo & Jaume, 28 :98. Cuba.

V. gouldii Binn. 25 :98.

V. (Angustula) milium Gld. 25:146; 28:101.

V. (A.) bermudensis Pils. 25:149. Bermuda.

Sterkia antillensis Pils. 26 :53.

Bothriopupa tenuidens C. B. Ad. 24 :227.

Pupisoma dioscoricola C. B. A. 26 :36.
P. minus Pils. 26 :40.

' ' Pupa ' ' lata C. B. Ad. 28 :135. Jamaica.

South American Pupillidae.

Gibbulina infundibuliformis Orb. 24:6. Bolivia.

Gastrocopta barbadensis solitaria Sm. 24:88. Fernando
Noronha.

DISTRIBUTION OF PUPILLIDAE. 169

G. servilis oblonga Pf r. 24 :90. Brazil.
G. miliola Orb. 24 :92. Brazil.
G. microdonta Doer. 24 :92. Argentina.
G. clessini Doer. 24 :93. Argentina.
G. nodosaria Orb. 24 :94. Bolivia.
G. wolfi Mill. 24 :94, 27 :208 ; 28 :117. Ecuador.
G. munita Reib. 24 :96, 357 ; 28 :64. Galapagos.
G. duneana Pils. 28 :65. Duncan I., Galapagos.
G. clausa Reib. 24 :99, 357 ; 28 :69. Galapagos.
G. (Immersidens) dicrodonta Doer, 24:100. Argentina.
G. (I.) iheringi Suter. 24:101. Brazil.
G. ( I. ) pazi Hid. 24 :102 ; 28 :64. Peru.
G. (I.) colombiana Pils. 26:228. Colombia.
G. (Geminidens) geminidens Pils. 24:228; 27:202.
Venezuela.

Pupoides simoni Jouss. 26 :114, Venezuela.

P. (Ischnopupoides) paredesii Orb. 26:120. Peru, Bolivia.

Sterkia eyriesii Drt. 26 :51. Guyana to Guatemala.

Bothriopupa tenuidens C. B. Ad. 24 :229. Venezuela.
B. conoidea Newc. 24 :231. Venezuela.
B. peruviana Pils. 28 :114. Peru.

Pupisoma dioscoricola C. B. Ad. 26:36; 28:114. Brazil,

Galapagos.
P. galapagorum Pils. 28 :114. Galapagos.

"Pupa" plusiodonta Holmb. 25:67. Argentina ( ?).

Appendix to Pupillidae.

Gastrocopta jeanneli Germain.

Shell sinistral, a little shortly pupoid, umbilicate. Spire
of 5 convex whorls, regularly and quite rapidly increasing, the
last large, very convexly ventricose; suture rather deep.
Summit obtuse, smooth. Aperture subvertical, semioval, with
separated, slightly converging margins, united by a very thin
callus, with a small, oblique, lamellar parietal fold nearer to
the upper insertion, and a small palatal denticle, hardly
noticeable, in the middle of the outer margin. Peristome
thick, especially in the middle part, white ; columellar margin
obliquely subrectilinear, dilated, yellowish white. The shell
is yellowish amber, glossy, subtransparent, sculptured with
longitudinal, oblique, subundulating striae which are unequal,

170 DISTRIBUTION OF PUPILLIDAE.

quite fine and irregularly spaced, readily visible to the
umbilicus, less crowded on the upper whorls than on the last ;
the embryonic whorls smooth. Length 2.5 mm., diam. 1.6 mm.,
height aperture 1.1 mm., width 1 mm. (Germain).

Kilima Njaro, in upper margin of the forest at Bismarck
Hill, between 2700 and 2800 meters (Ch. Alluaud and R.
Jeannel).

Gastrocopta jeanneli Germain, Bull. Mus. Nat. d'Hist. Nat.
(2), vol. 6, June, 1934, p. 381.

Lauria alluaudi Germain.

Shell subconic, formed of 6J/2 quite convex whorls, increas-
ing slowly, the last moderate. Suture somewhat deep, mar-
ginate. Aperture pyriform oval, quite angular above, the
margins converging very little and quite remote, with a strong
projecting lamellar parietal tooth and a very small supra-
parietal nodosity. Peristome slightly thick (the thickening
more accentuated in the middle of the outer margin) ; colu-
mellar margin dilated. The shell is yellowish chestnut;
sculptured with longitudinal oblique unequal striae, quite
strong but not lamellar. Length 3 to 3.25 mm., diam. max,
1.5 to 1.7 mm.; height aperture 1.3 to 1.5 mm. (Germain) .

Mt. Kinangop, alpine prairies at 3100 meters (Ch. Alluaud
and R. Jeannel).

Lauria alluaudi Germain, Bull. Mus. Nat. d'Hist. Nat. (2),
vol. 6, p. 382, June, 1934.

Lauria karisimbiensis, new species. PI. 25, figs. 2, 3.

The shell is umbilicate, ovate, tapering from the last whorl
to the obtuse apex, the lateral outlines convex ; cinnamon-buff,
subtranslucent, rather glossy, marked with weak, unequal lines
of growth. The whorls are rather weakly convex, the last
rounded at base, curving into the moderately large umbilicus.
The peristome is whitish, the outer margin becoming well ex-
panded in its lower part, basal margin built forward and
subreflected. Parietal margin bearing a very small lamella
situated on the right side of the middle, and extending inward
a short distance (about 0.4 mm.), its inner half being very
low. Length 3.3 mm., diam. 2 mm. ; 5^/2 whorls.

DISTRIBUTION OP PUPILLIDAE. 171

Belgian Congo: Camp Lukumi, Mt. Karisimbi (north of
Lake Kivn), at 11,370 ft., in moss in the subalpine heather
zone. Type 77267 M. C. Z., coll. by Dr. Jos. Bequaert.

By the wide removal of the angular lamella towards the
middle of the parietal margin, from its usual position near
the posterior angle, this species resembled L. horhonensis Pils.
and L. desiderata Preston (vol. 27, p. 62). It appears to be
closely related to the latter, which occurred on Mt. Kenya at
9,000 to 10,000 ft., but it differs by the very small size of the
angular lamella, which is more reduced than in any other
Lauria, and by the less oblique columella margin. I have not
been able to compare a specimen of L. desiderata, but its
habitat is remote from that of our species. The relatively
large umbilicus of L. karisimliensis is one of its notable
features.

L. bequaerti Pils (this vol., p. 138) is very different, being
larger with a strongly developed angular lamella connected
with the outer lip. L. alluaudi according to the description
•differs by possessing a strong tooth on the parietal wall, and
it has one turn more in a shell of the same or smaller size.

Truncatellina flavogilva Germain.

Shell subcylindric, the spire formed of 7 quite convex whorls
•of regular increase, the penult very convex, developed in
width, the last moderate, suture deep. Summit obtuse, flat-
tened. Almost no umbilicus at the bottom of a funnel-shaped
cavity. Aperture semioval, but little oblique, not toothed,
the separated margins joined by an inconspicuous callus.
Peristome white, thick, slightly reflected, the columellar
margin obliquely arched, white. The shell is a little grayish
yellowish. Embryonic whorls very finely striate, the others
-sculptured with oblique, subundulating and very close longi-
tudinal riblets. Length 1.5 mm. (Germain) .

Massif of Kilima Njaro: Neu Moschi, a post on the Rau
river at the foot of the south slope of the mountain, at 800
meters (Ch. Alluaud and R. Jeannel).

Truncatellina flavogilva Germain, Bull. Mus. Nat. d'Hist.
Nat. (2), vol. 6, p. 381, June, 1934.

172 DISTRIBUTION OF PUPILLIDAE.

Columella ninagongonis, new species. PI. 25, fig. 1.

The thin shell is deeply rimate, cylindric in the lower half,
slowly tapering in the upper, to the very obtuse apex, the
lateral outlines being convex; light isabella color, fading to
pale gray on the first two whorls. The whorls are short and
strongly convex, the suture rather deeply impressed. Sculp-
ture of weak lines of growth, the first two whorls microscopi-
cally punctate. The aperture is short, a little longer than
wide. Peristome whitish, the outer margin rather strongly
curved and thin posteriorly, anteriorly becoming well ex-
panded, together with the basal and columellar margins.
Length 1.9 mm., diam. 0.8 mm., length aperture 0.5 mm. j
7 whorls.

Belgian Congo: Mt. Ninagongo (north of Lake Kivu), in
moss of mountain forest at 9000 ft. Type 77268 M. C. Z.,
coll. by Dr. J. Bequaert, Harvard African Exped. 1926-7;
paratype A. N. S. P.

The only nearly related species known is C. pygmaeorum
Pils. & Ckll. (see p. 109), which is sinistral, a little larger and
with the outer lip less arcuate above.

The following names were listed in Servain, Oeuvres Scien-
tifiques de M. J. R. Bourguignat, 1891 (not seen by H. P.).
The list has been reproduced by M. Connolly, 1934, Proc.
Malac. Soc. London, vol. 21, p. 72. All appear to be nude
names.

Sphyradium adamii Bgt. Italy.
Orcula desertorum Bgt. Syria.

hellenica Bgt. Greece.

helvetica Bgt. Switzerland.

mabilleana Bgt. Greece.

pyrgula Bgt. Syria.

rapula Bgt. Syria.

scytoidea Bgt. Anatolia.

Orcula helvetica Bgt. is listed by Connolly as described by
Westerlund, Fauna Europaea, p. 198. The form described
in that place is [Pupa alpestris] "var. helvetica mh.", not the
Orcula of Bourguignat. Westerlund did not ascribe it to

VALLONIIDAE. 173

Bourguignat, but expressly claimed it himself. It is a Vertigo
(vol. 25, p. 201).

Pupa orcutti Pilsbry was mentioned by C. K. Orcutt (West
American Scientist VII, Oct. 1891, p. 270) as from near San
Quentin Bay, Lower California, on lichens. It is a nude name.
I have no record or recollection of this species, but it was
probably one of those described later from the same region.

Family VALLONIIDAE.

Valloniidae Pilsbry, 1900, Proc. A. N. S., Phila., p. 564.

Acanthinulinae Pilsbry, 1918, Man. Conch., vol. 24, p. x;
27, p. 186.

AcanthinuUdae Steenberg, 1918, Vidensk. Medd. Dansk.
Naturh. Foren., vol. 60, p. 14.

Valloniinae Watson, 1920, Proc. Malac. Soc. Lond., vol. 14,
p. 6.

Minute orthurethrous snails with perforate or umbilicate
shells of few whorls, from discoidal to ovate-conic in form,
often with spaced cuticular ribs, and without internal laminae ;
the peristome either expanded, thickened or simple ; toothless
(except in Spelaeodiscus) .

The oviparous or viviparous animal has a long penial ap-
pendix; the retractor muscle forked (except in Planogyra).
The prostate gland is short and posterior. As in many other
unrelated genera of minute orthurethous snails, the terminal
male organs are often wanting. The arcuate jaw is thin, with
weak, flat vertical folds. Central tooth somewhat or decidedly
narrower than the laterals, tricuspid or with ectocones much
reduced. Lateral teeth bicuspid ; marginal teeth wide, pectin-
ate, with several or numerous narrow cusps.

This group is understood in the limits of Steenberg and
Watson except that the genera Pyramidula and Pleurodiscus
are excluded.

The character of being toothless has not, I think, been suffi-
ciently appreciated by those who would unite Valloniidae with
the Pupillidae or the Strobilopsidae. Two leading genera,
Vallonia and Acanthinula, are known in many species from

174 VALLONIIDAE.

the early or middle Paleocene to the present time, without
showing traces of teeth or internal laminae. The only toothed
member of the family is Spelaeodiscus, and here the teeth are
not at all of Pupillid type, being more like the apertural teeth
which have arisen in various stocks of helices.

VALLONIA Risso.

Vallonia Risso, 1826, Hist. Nat. Eur. Merid. vol. 4, p. 101,
sole species V. rosalia Risso = V. costata Miill,

Amplexis Brown, 1827, Illust. Conch. Gt. Brit, and Ireland,
expl. pi, 41, for A. paludosus (= V. pulchella) and A. crenellus
(== V. costata) ; A. paludosus here selected as type.

Amplexus Brown, 1844, Illust. Conch. G. B. and Irel. (2),
p. 45, for Helix pulchella and H. crenella. Not of Sowerby,
1815.

Zurama Leach, in Turton, 1831, Man. Land and Fresh-
water Shells Brit. Is. p. 64, in synon. under Helix pulchella.
Quoted as Zuramia Leach by Kennard & Woodward, 1926,
Syn. Brit. Non-mar, Moll. p. 135, in synon.

Circinaria Beck, 1837, p. 23, in part, — Herrmannsen, Mar.,
1847, Ind. Gen, Malac. vol, 1, p, 237, Helix pulchella selected
as type.

Glaphyra Albers, 1850, Die Hel. p. 87, for Helix tuckeri,
pulchella and costata ; H. pulchella here selected as type.

Lucena ''Hartmann", Gray, 1840, in Turton 's Manual, p.
142. — MoQUiN-TiVNDON, 1855, Moll. France, vol. 2, p. 140, for
H. pulchella. Not Lucena Hartmann, 1821, Neue Alpina I,
208, 245.

Valtonia Kisse, Ancey, 1881, Le Naturaliste, vol. 1, p. 215,
for H. pulchella (typographical errors),

Lurama Leach, Honigmann, 1910, Abh, u, Ber, Mus. fiir
Natur- u. Heimatsk. u. Naturw. Ver. Magdeburg, vol. 2, p. 43
fori, pulchella (typog. err.).

Vol. 8, pp. 247-261. For notes on anatomy see p. 195, pi.
29, fig. 1, y, pulchella (Miill.).

ACANTHINULA Beck.

Vol. 27, p. 188. See also p. 196, where some account of the
anatomy is given. PL 29, fig. 6, genitalia of A. aculeata, after
Steenberg.

VALLONUDAE. 175

0 7 8 12 i« ir

•r

rig. 9. A, Zoogenetes harpa. B, C, Spelaeodiscus triarius.

ZOOGENETES Morse.

Vol. 27, p. 195. Steenberg (1925, pi. 29) has figured the
genitalia of an individual without male terminal organs. In
a specimen from High Pines, Duxbury, Massachusetts, taken
in September, 1911, I found the penis fully developed and the-
uterus containing embryos (fig. 9a.).

The kidney has the pouch-like form usual in Pupillidae,.
the walls of its large cavity having about ten unequal longi-
tudinal ridges. Orthureter long, direct, opening in the lung
some distance from the collar. It is heavily pigmented to-
ward the anterior end, which appears externally as an oblong
black spot.

The penis is rather thick and bears a long appendix en-
larged distally, narrowed in the middle. Epiphallus rather
thick, without a flagellum. The bifid retractor muscle is in-

176 VALLONIIDAE.

serted on epiphallus and appendix. The prostate gland is
small and posterior. The spermoviduct is strongly con-
voluted. Albumen gland multilobular. Uterus much dis-
tended by the contained embryos, its neck much shorter than
the vaginal part of the oviduct. Spermatheca is oblong, but
little larger than its duct, which is of medium length, with-
out diverticulum.

Zoogeiietes differs from Acanthinula chiefly by the absence
of appendages on the epipliallus, the shorter spermathecal duct
and the viviparous reproduction. It is more like Vallonia in
the genitalia.

SPERMODEA Westerl.

Vol. 27, p. 186. Specimens possessing the penis have not
yet been dissected. Without it, the position of the genus can
not be closely estimated.

PLANOGYRA Morse.

Man. Conch, vol. 9, p. 45. Dr. H. Burrington Baker has
shown that this genus is related to Acanthinula. See page
197, pi. 26, figs. 1-3.

Genus SPELAEODISCUS Brusina.

Vol. 27, p. 180. This genus was associated with Pagodulina
on account of some slight similarities in the shells. The ana-
tomical work and figures of Soos (1917, p. 65, figs. 39-42),
partly reproduced here, p. 175, figs. 9 h, c, show a long ap-
pendix, contracted in the middle, as in various Valloniidae.
There is a thick "flagellum", and the penial retractor is
forked. The spermathecal duct is short, as in Vallonia and
Zoogenetes. The prostate gland, composed of very long
tubules, shows unusually strong development.

Steenberg (1925, pp. 184, 202) suggests a subfamily
Spelaeodiscinae, which he includes with some doubt in the
Valloniidae. Watson (1920) judging by the figures of Soos,
considered the genus allied to Vallonia and Acanthinula.
This position appears to me to be logical. The resemblance
of Spelaeodiscus to StroMlops mentioned by H. B. Baker seems

PLEURODISCIDAE. 177

to me a secondary and superficial one, as the former does not
share the essential character of Strohilops, which is the pos-
session of a system of internal laminae from a very early stage
to the adult.

Pagodulina (Vol. 27, p. 166) has not been dissected by any
recent investigator. The figure of Moquin-Tandon, as far as
it goes, agrees with Orcula in form of penis and appendix, as
pointed out by Steenberg (1925).

Family PLEURODISCIDAE.

Pleurodiscidae Wenz, 1923, Fossilium Catalogus, Pars 21,
p. 1069.

Patulastridae (Pleurodiscidae) Steenberg, 1925, Vidensk.
Medd. Dansk. Naturh. Foren. i Kobenh. vol. 185, 202.

The shell is umbilicate, discoidal or turbinate, closely cos-
tulate or striate, the peristome simple and sharp.

The viviparous animal has simple male end organs, the
passage of penis into epiphallus marked by an abrupt con-
traction, the simple penial retractor inserted on epiphallus;
no appendix or flagellum. The spermathecal duct is of
medium length (Pleurodiscus) or short {Pyramidula) , and
not forked. Jaw and teeth substantially as in Valloniidae.

Two Palaearctic genera grouped together in this family are
not closely related, but they agree in the simplicity of the male
organs, and are not strikingly unlike in the shells. To include
them in the Valloniidae would destroy the reasonably homo-
geneous character of that group.

Genera of Pupillidae without penial accessories have been
regarded as secondarily simplified end-products of evolution,
on account of their close relationship by the shell and other
qualities with genitally complicated genera (this vol., Intro-
duction), as in the case of Vertigo and Nesopupa. I can not
see any stem in the Valloniidae which will serve as a plausible
support for the attachment of Pleurodiscus and Pyramidula
thereto as simplified branches.

Dr. H. B. Baker suggests that the Pyramidulinae and
Pleurodiscinae may represent primitive conditions, before the
evolution of complex genitalia (this vol., p. 193). In that

178 PLEURODISCIDAE.

case their divergence from the common stem of the other
groups must have been prior to the branching out of the
Achatinellidae, Tornatellinidae, Pupillidae, Valloniidae etc.,
all of which had acquired similar penial complications before
the parting of their several ways.

On the whole, the inclusion of Pleurodiscidae in Valloniidae
appears to be without satisfactory basis. That classification
was apparently prompted by a desire to restrict the number of
families. I see no advantage in claiming a relationship which
involves so many difficulties.

1, Shell of moderate size, discoidal; penis very short, with
apical papilla through which the long, internally ridged
epiphallus enters; prostate gland long, about equal to the
uterus; spermathecal duct moderately long (Pleurodiscinae)

Genus Pleurodiscus Wenz.

2. Shell minute, depressed conic or turbinate ; penis abruptly
terminated ; prostate gland short, posterior ; spermathecal duct
short. (Pyramidulinae) Genus Pyramidula Fitz.

Genus PLEURODISCUS Wenz.

Pleurodiscus Wenz, 1919, Nachrbl. Deutsch. Malak. Ges.
vol. 51, p. 778, type Goniodiscus (Pleurodiscus) halmei (Pot.
et Mich.).

Helix, Patula and Patulastra of some authors; not Patu-
lastra Pfeiffer as restricted to ^'Patula" pygmaea Drap. by
Kobelt, 1880, 111. Conchylienbuch p. 231.

For anatomy see Hugh Watson, Proc. Malac. Soc. Lond.
vol. 14, p. 6.

The shell is Discus-like, openly umbilicate, depressed, of 5y^
to 6 whorls, the apical 2i/2 smooth, the rest costulate. Aper-
ture lunate; peristome sharp, somewhat dilated at the
columella.

The jaw is thin, closely striate, the edges of the folds slightly
free (Man. Conch. IX, pi. 15, f. 2). Radula of P. lalmei with
17-9-1-9-17 teeth. Central tricuspid, smaller than the laterals,
which are bicuspid. Marginals wide, pectinate.

Reproduction ovo-viviparous. Penis yevj short, the epi-
phallus entering through a short blunt papilla. Epiphallus

PLEURODISCIDAE. 179

long, the penial retractor inserted on it. No penial appendix,
but the epiphallus has internal complications in the form of
spiral ridges. Spermatheeal duct long. Prostate gland about
as long as the uterus, composed of many narrow, separate
tubules (PI. 29, figs. 3-5, after Watson).

This very distinct genus comprises a few species distributed
around the eastern basin of the Mediterranean. They are
closely similar, and possibly not specifically distinct, but only
the Sicilian P. halmei has been dissected. Wenz (Fossilium
Catalogus part 21, pp. 1069-1071) recognized three or four
middle European Tertiary species as follows :

Pleurodiscus falcifertjs (Bttg.), Helix (Patula) falcifera
Boettger, 1870. Upper Miocene, Aquitanian, near Ulm;
Burdigalian, Tuchorschitz. Said to be the same as P.
orbicularis.

P. FRici (Klika), Patula (Anguispira) frici Klika, 1891.
Upper Oligocene, Warzen, Czechoslovakia.

P. MAMiLLATUS (Andreac), Pyramidula mamillata Andreae,
1904, Upper Miocene, Oppeln.

P. ORBICULARIS (Klein), Helix orbicularis Klein, 1846.
Upper Miocene, near Ulm.

Pleurodiscus balmei (Pot. & Mich.). Plate 25, fig. 15.

The shell is depressed with convex spire, umbilicate, the
umbilicus contained about 4^ times in the diameter, some-
what well-like, contracting but slowly within. Whorls convex,
slowly increasing, the last rounded; suture deeply impressed.
Buff, the base pale or whitish. The apical 2y^ whorls are
nearly smooth and glossy; following whorls dull, rib-striate,
the riblets narrow, 3 to 4 in one mm. on the last whorl, a few
fine striae in the intervals. Aperture but slightly oblique.
Peristome thin, blunt, the basal margin straightened or
slightly curving forward, columellar margin dilated.

Height 5.5 mm., diam. 9.5 mm. ; 5% whorls.

Height 6.3 mm., diam. 11 mm. ; 6 whorls.

Sicily: Monte Pellegrino, near Palermo, type loc. ; Malta
(Caruana) ; Sardinia near Laconi; Italy at Reggio in
Calabria ; Algeria.

180 PLEURODISCIDAE.

Helix halmei Potiez et Michaud, 1838, Galerie Moll, de
Douai, I, p. 120. Paulucci, 1879, Journ. de Conch, vol. 27,
p. 7. Westerlund, 1889, Fauna Pal. Reg. Binnenconch,
Helix p. 13. — Helix flavida Zgl. Rossmaessler, 1839, Iconogr.
vol. 2, p. 13, f. 610.— Pfeiffer, Mon. Ilel. Viv. vol. 1, p. 144.—
Helix flavescens Parreyss in coll., Rossmaessler, 1839, I. c,
as syn. of H. flavida. — Helix teresae Benoit, 1845, Ricerche
Malacologiche, p. 9, pi. 2, f. 5 a, h. (Palermo, Cephalu,
Siracusa etc.).

Rossmaessler 's good figures were copied by Tryon in M. C.
vol. 3, pi. 6, figs. 48-50. Potiez and Michaud 's description
and dimensions (7-8x12-14 mm.) were so ambiguous that
though often discussed, no one recognized that the well-known
H. flavida was intended until Mme. Paulucci examined the
types in 1879. It was a pity to displace the well-defined
name flavida.

Pleurodiscus erdeli (Roth). PL 25, fig. 16.

Shell widely umbilicate, depressed, elegantly costulate,
pellucid, buff-corneous, paler beneath. Whorls rounded.
Aperture transversely lunate-ovate ; peristome straight, simple,
flexuous. Alt. 21/^, diam. 3% lines; 5 whorls. Differs from
H. ruderata Stud, by the size, the narrower umbilicus and
more depressed aperture (Roth).

Rhodes, type loc. Roth; Syria at Beirut; Palestine; Asia
Minor; Constantinople.

Helix erdelii Roth, 1839, Moll. Species in Itinere per
Orientem facto, etc., p. 16, pi. 1, f. 4, 5, 20. — Pfeiffer,
Monogr. I, p. 105.

I have not seen specimens from Rhodes. A Beirut example
figured measures height 6.1 mm., diam. 10.7 mm. ; barely 6
whorls. It is cinnamon-buff, the umbilicus noticeably wider
than in P. halmei, more perspective. In the aperture a white
callus lining is seen, stronger than in P. halmei, of which some
authors consider erdeli a variety. It was named for Dr. M.
Erdl, Roth's travelling companion in the Orient.

It has been recorded from Malta by Issel (1868, Bull. Malac.
Ital. I, p. 5, the specimens identified by Bourguignat).

PLEURODISCIDAE. 181

Maltese shells I have seen do not seem to differ from Sicilian
ialmei.

Pleurodiscus sudensis (Pfr.). PI. 25, fig. 14.

Shell umbilicate, depressed, regularly and closely costulate,
thin, diaphanous, reddish-corneous; spire slightly elevated;
whorls 6, convex, the last not descending; umbilicus rather
large, perspective; aperture rotund-lunate; peristome simple,
acute, columellar margin slightly dilated, shortly spreading.
Alt. 7, diam. 13x11.5 mm. (Pfr.).

Crete : Suda, type loc, ; Candia ; Akrotini ; all in the north-
west part of the island.

Helix sudensis Pfeiffer, 1846, Symbolae Hist. Hel. Ill, p.
103. — RossMAESSLER, Icouogr. L. u. Siisswasser Moll. Eur. Ill,
p. 85, f. 901.

The color is *' pinkish-buff", a little browner than P. balmei.
The umbilicus is contained 4j^ times in the diameter. Apical
whorls as in P. halmei. The shell is larger than talmei, with
about half a whorl less. The last whorl is relatively a little
wider and less depressed, the basal margin of the peristome a
little more deeply arcuate. The riblets are slightly closer;
finally, the white callous lining of the last whorl occupies
more space than in halmei, about one-sixth of the base. A
specimen from Akrotini figured measures : height 8 mm., diam.
13.5 mm. ; 5}^ whorls.

P. SUDENSIS CYPRiA (Kob.). PI. 25, figs. 17.

"Differs from the type by the slightly larger shell, more
widely umbilicate, more depressed, lightly flattened below,
aperture ovate, the lower margin of the lip straightened.
Diam. 15 mm., min. 14 mm., alt. 8 mm." {Kobelt).

Cyprus, in several localities (H. Rolle).

Patula sudensis var. cypria Kobelt, 1897, Rossmaessler's
Iconogr., n. F., Erster Supplement-Band, p. 56, pi. 22, f. 5-7.

Kobelt further states that this agrees most closely with the
Cretan P. sudensis, especially in the size and sharp ribbing,
the wide umbilicus and the widening of the last two whorls.

182 PLEURODISCIDAE.

It is flatter, the lower side especially flattened, the umbilicus
is wider, the basal margin nearly straight ; the whorls increase
more rapidly than in the type, etc. I have not seen this form,
which from the above account seems to me except in size to
have more the characters of erdeli than of sudensis.

Genus PYRAMIDULA Fitz.

Pyramidula Fitzinger, 1833, Syst. Verzeich. Oesterreich
Weichthiere p. 95, for Helix rupestris Drap.

For anatomy see : Soos, 1917, Ann. Mus. Nat. Hungarici vol.
16, pp. 114, 160, f. 88, 89.— Hesse, 1918, Nachrbl. D. Malac.
Ges. vol. 50, p. 110.— Watson, 1920, Proc. Malac. Soc. Lond.
vol. 14, pp. 6-30, f. 4a, pi. 2, f. 1, 3, 4.

Shell small (diam. between 2 and 4 mm.), turbinate or de-
pressed with conic spire, of 4 to 5 strongly convex tubular
whorls and moderate or wide umbilicus; aperture toothless;
peristome simple, the columellar margin somewhat expanded.

Jaw very thin, delicately striate. Radula with centrals
about as wide as the laterals, tricuspid with very small
ectocones (Watson), or unicuspid (Soos, Wiegmann) ; laterals
bicuspid; marginals wide, pectinate, with numerous narrow
cusps, no entocones.

Reproduction ovo-viviparous ; penis enlarged distally (see
below), the penial retractor inserted on the vas deferens
(epiphallus ?) ; prostate gland of few tubules, posterior;
spermathecal duct rather short; neck of uterus and vagina
about equal in length (PL 29, fig. 2, P. rupestris, after Watson) .

This genus inhabits the southern part of the Palaearctic
Region from Spain to Japan. Only a few species are recog-
nized. Possibly a few others included by Pfeiffer (Nomencl.
Hel. Viv.) in Patulastra may turn out to be Pyramidulae,
but most of the minute ones seem to belong to Punctum. The
type of Patulastra is Helix pygmaea Drap., designated by
Kobelt, 1880.

The several accounts of the anatomy of P. rupestris differ
somewhat, so that Watson intimates that different species are
involved. Fritz Wiegmann, quoted by Hesse, states that the
penis was undeveloped in his specimens, its Anlage having a

PLEUKODISCIDAB. 183

length of 0.4 mm., consisting of a somewhat curved, club-
shaped, blunt body, which receives the vas deferens at about
the distal third, the penial retractor inserted on the vas
deferens. Soos described the penis as remarkably small, fusi-
form, distally recurved upon itself, the two parts lying close
together; tapering suddenly into the very long, thread-like
vas deferens ; the weak retractor inserted on the thickest part
of the penis. Watson, whose figure is copied in PI. 29, fig. 2,
says "penial appendix much reduced, being represented by a
mere knob, without muscular attachment". It appears to me
very doubtful whether the distal enlargement of the penis is
correctly considered to be homologous with the appendix of
Vallonia, etc.; Soos' figure does not show it, though he men-
tions that the transition from penis to vas deferens is abrupt.
The male organs seem to be very similar to those of some
Vertigininae, such as Columella as figured by Steenberg ( 1925,
p. 100).

According to Moquin-Tandon the uterus contains 3 to 7
embryos; Watson's figure shows 2 or 3. Both Wiegmann and
Soos report not more than one in the specimens they opened.

Wenz (1923, Fossilium Catalogus, p. 1062) recognizes two
Tertiary species:

P. ANTONiNi (Michaud, 1862), Pliocene, Hauterive (Drome).

P. SUBTERES (Clessin, 1877), Upper Mioc, Undorf bei
Eegensberg.

Helix (Patula) recurrecta Oppenheim, 1890, from the
Eocene of Monte Altissimo, Prov. Vicenza, perhaps belongs
here ; see also p. 12.

Pyramidula rupestris (Drap.). PL 25, figs. 9, 9a, 9&.

The low-conic shell is openly umbilicate (the umbilicus con-
tained about five times in the diameter), hazel or russet,
glossy, finely, irregularly striate, the initial 1^^ whorls micro-
scopically granulose; composed of 4 strongly convex whorls,
which are flattened below the deeply impressed suture. The
aperture is basal, oval, the thin peristome expanded a little
at the columella. Parietal callus transparent, very short.
Height 1.8 mm., diam. 2.3 mm.

184 PLEURODISCIDAE.

Middle and southern Europe, type localities Castelnau and
Arbois, in southern France. Algeria; Tunis j Syria. Gener-
ally on limestone rocks.

Helix rupestris Dr^vparnaud, 1801, Tabl. Moll. Fr. p. 71;
Hist. Nat. Moll. terr. fluv. France, p. 82, pi. 7, f. 7-9.—
Pfeipfer, Monogr. IV, p. 71. — Westerlund, 1889, Fauna,
Helix, p. 13. — Helix rupestris mut. rupicola Stabile, 1859,
Prospetto Moll. terr. fluv. Lugano, pp. 21, 51. — Helix rupestris
Drap., BouRGUiGNAT, 1864, Malac. Algerie I, p. 184, pi. 16, f.
24-27; with Var. conoidea on expl. pi. 16, f. 31-33; Helix
scotaea Bgt. in coll. stated to be the same, p. 186. — Helix
rupestris var. meridionalis Issel, 1870, Bull. Soc. Malac. Ital.
Ill, p. 115. — Helix (Patula) jaenensis Clessin, 1882, Malak.
Bl. n. F., V, p. 187, pi. 4, f. 3. — Helix rupestris var. pinii
Adami, 1886, Bull. Soc. Veneto-Trentina Sci. Nat. Ill, p. 172.
— Patula rupestris Drap. var. dalmatina Clessin, 1887, Malac.
Bl. n. F., IX, p. 51. — Helix umhilicata Montagu, 1803, Test.
Brit. p. 434, pi. 13, f . 2. — Pfeipfer, Monogr. I, p. 86 ; Conchyl.
Cab. pi. 89, f. 32-35. — Turho myrmecidis Scacchi, 1833,
Osservazione Zoologiche I, p. 11 (not seen). — Helix aliena
Zgl., Pfeipfer, 1841, Symbolae ad Hist. Hel. I, p. 39. — Helix
spirula Villa, 1841, Dispositio Syst. Conch. Coll. Villa, p. 56
(Prov. Comensi). — Delomphalus rupestris saxatilis Hart-
MANN, 1842, Gasterop. Schweiz, p. 122. — Helix rupestris forma
suhglohosa "Bgt." Westerlund, 1889, Fauna Palaarct. Reg.
Binnenconch., Helix p. 14 (was not named by Bourguignat.
Westerlund evidently refers to Bourguignat 's "var. C",
which that author considered to be typical rupestris of south-
ern Europe, giving the distribution as from Spain to Beirut,
Syria). — Pyramidula rupestris Drap., Kennard & Woodward,
1926, Syn. Brit. Non-mar. Moll., p. 103 (synonymy).

This common and widely spread species varies in size, de-
gree of elevation, width of the umbilicus and in sculpture.
I have made no study of its variations and races, but note
the following, the characters given being taken from the sev-
eral authors.

Var. saxatilis Hartm. Differs from true rupestris by the
larger size, flatter shape and wider umbilicus. 2x3 mm.
Wiirttemberg near Waldhausen, cliffs of Ehningen, etc. A
specimen from Lausanne which I refer to this race measures,
1.7 X 2.7 mm., umbilicus one-third the diameter. PI. 25,
figs. 10, 10a.

PLEURODISCIDAE. 185

Var. rupicola Stabile. Shell more conic, umbilicus nar-
rower. Massagno and Mont S. Salvatore, Lugano. A speci-
men from Pisa measures 2.3 x 2.5 mm., umbilicus 0.5 mm.
PL 25, fig. 8.

Var. scotaea Bgt. PI. 25, figs. 6, 7, after Bgt. Conoid-
globose, the umbilicus small. 3 x 3.5 mm. Constantine, Bou-
Mecid ; Bougie ; the only variety which has been found in
Algeria (== var. conoidea Bgt. not H. conoidea Sowb.).

Var. meridionalis Issel. Spire more elevated, umbilicus
very narrow. 2.5 x 2.5 mm. ; 5 whorls separated by deep
suture. Monte Sant' Angelo, Ferentillo, Umbria.

Var. jaenensis Cless. Conoid-turbinate, deeply and widely
umbilicate, higher than wide. Alt. 2.5 mm., diam. 2 mm.,
5-53^ rounded whorls. Jaen, Spain.

Var. dalmatina Cless. More highly coiled and more globose.
Dalmatia : cliffs over the source of the Jedrobach and in the
Cettina valley near Almissa.

Var. piyiii Adami. Shell larger, pyramidal, strongly striate,
the suture extremely deep, whorls more rounded. Diam. 3-4
mm., alt. 2.3 - 3.3 mm. Mt. Baitone, Bergamasque Alps, on
schistose and granitic rocks.

Helix rupestris var. trochoides (Ferussac, Tabl. Syst. p. 40
(44), not defined; said to be from Castelnau, one of Drapar-
naud's localities for typical rupestris) Paulucci 1879, Fauna
Malac. Calabria, p. 64; also 1881, Bull. Soc. Malac. Ital. VII,
p. 85, referring also to Ferussac, Hist. pi. 80, fig. 3, "la variete
conoide. ' ' That figure represents a specimen somewhat higher
than wide. Mme. Paulucci placed var. meridionalis Issel in
the synonymy of trocJioides Fer. ; but the name cannot fairly
be ascribed to Ferussac, since it was not connected with his
later figure until Mme. Paulucci made that inference in 1879.

Helix rupestris var. trochoides Kreglinger, 1870, Syst. Verz.
Deutsch. Binnen-moll. p. 55, defined by the single word
"elevata^', may be the same. No definite locality given.

Probably the above names rupicola, meridionalis, trochoides
and dalmatina were applied to forms scarcely if at all dis-
tinguishable. My pi. 25, fig. 8, from Pisa, represents this

II MK^

186 PLEUBODISCIDAE.

high type of rupestris. "Whether the high-conic Hispanic
jaenensis is directly related to these forms of the Alpic center
is doubtful.

Anatomical considerations suggest that rupestris of authors
includes more than one species, but those who have published
dissections of these snails have not given the shell characters
of material used. Correlation of the anatomic data with
conchological racial characters given above is therefore im-
possible at present. The variations and races of rupestris
stand much in need of revision by some European investigator,
as only short-range local views have been published.

Pyramidula rupestris chorismenostoma (Blanc & Westerl.).
PI. 25, fig. 11.

Shell turbinate, delicately transversely striate, rufous-
brown, whitish at the aperture, widely and deeply umbilicate ;
whorls 4y^, the upper subcylindric, flattened at the suture,
the penult subangulate in the middle, the last whorl depressed
subcylindric, wholly or in part free in the manner of Ver-
Tiietus. Aperture subcircular [or oval], peristome simple,
acute, straight. Diam. 2.5 mm., alt. 2.25 mm. {Westerl.).

Greece : Syra, around the village of St. Georges, on the inner
faces of piles of stones with Lauria umhilicata (Roth) ; Maco-
lessos, Beotia.

[Helix rupestris] subspecies chorismenostoma Blanc &
Westerlund, 1879, Aperqu Fauna Malac. de la Grece, p. 32.

A Beotian specimen is figured.
Pyramidula rupestris przevalskii Lindh.

Differs from the type by the larger shell with less convex
base, the umbilicus narrower, nearly cylindric, not dilated be-
low; spire much higher for the width of the shell. Shell
narrowly umbilicate, turbinate, depressed, thin, striatulate,
deep reddish corneous. Spire conoid, the apex rather obtuse ;
suture impressed; umbilicus narrow, penetrating, nearly
cylindric, scarcely dilated at the base. Whorls 414 to 41^,
rounded, slowly and regularly increasing, the last distinctly

PLEURODISCIDAE. 187

subangular, scarcely descending in front. Aperture rounded-
lunar, oblique ; peristome simple, straight, the margins distant,
columella a little reflected. Alt. 2.8-3 mm., diam. 3-3.25
mm. (Lindholm).

Eastern Turkestan: Keria Mts. at 10,400 ft. (N. M.
Przevalski).

Pyramidula (Pyramidula) rupestris Drap. subsp, prze-
valskii Lindholm, Annuaire du Mus. Zool. de I'Acad. Sci. de
la Russie, vol. 23, p. 256.

Lindholm further notes that var. conoidea Bgt. reaches
similar size but has the bodywhorl quite rounded and the
umbilicus more open towards the base. The north Indian
P. humilis has a much more depressed shell and a larger
umbilicus.

Pyramidula hierosolymitana (Bgt.). PI. 25, figs. 12, 13.

Shell very small, narrowly perforate, globose-conoidal, very
sharply striatulate, brownish-corneous; 4 convex whorls, in-
creasing regularly; suture deeply impressed; last whorl
rounded; the small perforation narrow. Aperture obliquely
rounded; peristome simple and acute, the columellar margin
reflected subdeflexed in the umbilical perforation; margins
approaching, slightly separated by the last whorl, Alt. 2i/4
mm., diam. 2i/2 mm. (Bgt.).

Palestine : Valley of Jehosaphat, under the sun-baked stones
of tombs, rather abundant (Saulcy) ; valley of Hinnon (Roth).

Helix hierosolymitana Bourguignat, Testacea Novissima
quae Saulcy in itinere per Orientem coll., p. 13; Cat. Moll.
•Saulcy, 1853, p. 22, pi. 1, f. 32-35.— Kobelt, 1877, Ross-
maessler's Iconogr. vol. 5, p. 93, f. 1416.

It differs from P. rupestris by the much narrower umbilicus.
Bourguignat 's figures are copied.

Pyramidula conica Pils. & Hir. PL 23, figs. 3-3c.

Shell umbilicate, conic, rather thin, amber-brown with a
slightly olivaceous tint, slightly translucent. First 1^ whorls
smooth, with rather shallow suture ; the rest closely, finely and

188 PLEURODISCID.VE.

rather irregularly striate. Whorls 4i/^, very convex, separated
by a very deep suture, the last rounded at the periphery and
beneath. Aperture oblique, rounded, about one-fourth of the
circumference excised at the parietal wall by the preceding
whorl; peristome simple and thin, the columellar margin a
little dilated. Umbilicus perspective, contained 3I/2 times in
the diameter. Alt. 1.3, diam. 2.8 mm.

Japan: Suimura, Awa, Shikoku. Types no. 83884, A. N.
S. P.

Pyramidula conica Pilsbry & Hirx\se, 1902, Nautilus XVI,
p. 77.

This species is excessively similar to the common European
P. rupestris, and in separating it specifically I was somewhat
influenced by the enormous distance of its habitat from the
range of the European species. Lately, however, a form of
rupestris has been traced into high Asia, P. r. przevalskii
Lindh., in Eastern Turkestan, about Ion. 81 E. This is still
a long distance from Shikoku, in E. Ion. 134. It is probably
best to treat the insular Japanese form as distinct from the
European, pending a comparison of the anatomy. It differs
by the more tubular, less depressed last whorl and rounder
aperture.

In this connection it may be well to notice some Chinese
snails described as Pyramidula. Pyramidula (Patula) peipi-
nensis Ping & Yen, 1932 (Bull. Fan Mem. Inst. Biol. Ill, No.
2, p. 25) is either a Hippeutis or a small northern form of the
common "Planorhis" umhilicalis Bens., which has a shell like
Hippeutis, but may be a toothless Segmentina. Pyramidula
(Patulastra) grahaui and P. (P.) gracilis Ping & Yen, 1933
(same Bulletin, vol. IV, No. 6, pp. 263-6), are Zonitidae.

Pyramidula humilis (Hutt. & Bens.) . PI. 23, figs. 1, la, lb, 2.

Shell small, convexly depressed, corneous, broadly and
deeply umbilicate; whorls 5, rounded, the last subangulate,
the penult slightly interrupts the circular aperture ; peristome
acute. Diam. 0.125 inch. {Benson).

PLEURODISCIDAE. 189

India: Simla (type loc, Hutton) and Chur (Stoliczka) ;
Landour (Benson) ; Murree (Stol.) ; Tandiana (Theobald).

Helix humilis Hutton, Hutton & Benson, 1838, J. A. S.
Bengal, vol. 7, p. 217.— Pfr., 1847, Monogr. I, p. 106 ; Conchyl.
Cab. Helix III, p. 332, pi. 129, f. 28.— Hanley & Theobald,
1874, Conch. Indiea pi. 61, f. 4-6. — Nevill, 1878, Second
Yarkand Mission, Moll., p. 18. — Pyramidula humilis Pilsbry,
1903, Man. Conch. IX, p. 44.— Gude, 1914, Fauna British
India, Moll. II, p. 43.

The embryonic shell of 1^2 whorls projects somewhat; it
is glossy, with the faintest traces of granulation, the first
whorl being wide, the suture only lightly impressed. Sub-
sequent whorls are dull, brown, with irregular, somewhat
thread-like striae and a very deeply impressed suture. The
last whorl has the periphery above the middle ; it is distinctly
though bluntly angular in immature shells (fig. 2), but the
angulation is almost lost on the last whorl of adult shells.
The umbilicus is widely open, occupying one-third or more
of the diameter of the shell. A Landour specimen received
from Benson measures : height 1.4 mm., diam. 2.3 mm. ; 4%
whorls.

Two species of Punctum are noticed below because they
had been figured on Plate 23 under the impression that they
were Pyramidulae. Probably all Punciuyn show spiral lines.
The species appear to be distributed throughout much of the
Holarctic realm and into the tropics in Mexico. The South
African " Trachycystis" hottentota M. & P. has the appear-
ance of Punctum. In the Atlantic Islands the genus is repre-
sented by P. pusillum (Lowe) and P. placidum (Shuttl.).
The European species have been described by Westerlund,
1889, but his list contains various heterogeneous elements.

Punctum orphana (Heude). PI 23, figs. 5-5c.

Shell umbilicate, orbicular, depressed-conoid, rufous, plicatu-
late-striate, the spire depressed; whorls 4, slowly increasing,
somewhat convex, joined by a rather deep suture, the last
cylindric, at the periphery indistinctly subangular, not
descending; aperture subcircular, nearly straight, peristome

190 PLEURODISCIDAE.

simple, the columellar margin dilated a little ; umbilicus wide,
perspective. Alt, 1. 5 mm., diam. maj. 2.5 mm., min. 2.25 mm.
(Eeude).

China: Shanghai, under fallen bamboo leaves, abundant
(Heude) ; Soo-chow (Teng-Chien Yen).

Helix orphana Heude, 1882, Mem. Hist. Nat. Emp. Chinois:
Notes Moll. terr. Vallee Fleuve Bleu, p. 21, pi. 13, f. 18.

The animal is black. It is found in many bamboo thickets.
Collected in the orphanage of the mission at Shanghai.

My figures are from a specimen received from Pere Heude.
The dentition has not been examined, but from the presence
of microscopic spiral striae I think that it is a species of
Punctum. The shell figured measures, height 1.65 mm., diam.
2.6 mm. ; 4 whorls.

Punctum micra (Pils.). PI. 23, figs. 4, 4a, 4&.

The spiral striation, not represented in the figures, is as
described for P. orphana. The specimen figured measures:
height 1.4 mm., diam. 2.1 mm.

Korea: Bay of Ulsan (Kuroda).

Pyramidula micra Pilsbry, Proc. A. N. S. Phila. 1926, p.
469, pi. 35, f. 1, la, 1&.

pupillidae and related groups. 191

Review of the anatomy op Pupillidae and related groups.

By H. Burrington Baker.

The systematic inferences from these few notes on anatomy
are aired with many misgivings ; any deductions based on the
animal alone would be weak and this would be especially true
of a group in which the shell characters, so far as my slight
knowledge goes, appear manifest while those of the soft parts
are difficult to study and still more so to evaluate. Personally,
I am more inclined towards "Watson's (1920) broad limits of
the Pupillidae than towards the other extreme proposed by
Steenberg in his exquisitely careful ''fitudes" (1925) ; to me,
the total range of anatomical differentiation seems much less
than in most families of pulmonates. Incidentally, the male
genitalia of these Orthurethra go through strikingly parallel
modifications to those in the sigmurethrous Sagdidae (which
I happened to study simultaneously, like Steenberg the Clausi-
liidae) ; in these, the appendicular arm functions as the apex
of the penis. Finally, one must not forget that the soft parts
of a large majority of the species and even of the genera and
subgenera are still unknown. A tentative outline of the rela-
tionships between the subfamilies may be expressed in the
form of a key, with many derivative conditions omitted, as
follows :

A. Valloniinae ( ?) ; shell more heliciform, without teeth
[unless Strohilops (p. 2) or Spelaeodiscus (vol. 27, p. 180)
be included] ; radula with fundamentally bicuspid laterals
and marginals but last with ectoconal accessories; penis
(when present) well differentiated.

1. Penis bifid and sharply separated from its epiphallus
and appendix ; prostate and spermatheca short.
a. Shell dominantly calcareous and with more or less

thickened and expanded peristome Valloniinae.

6. Shell dominantly epidermal, often with membranous

excrescences; peristome usually thin

Acanthinulinae.

192 PUPILLIDAE AND RELATED GROUPS.

2. Penis without appendix ; shell endodontoid (more like b) .

c. Appendicular penial arm vestigial; prostate and
spermatheca short Pyramidulinae.

d. Penis short and simple; epiphallus strong with
prominent papilla ; prostate and spermatheca longer

Pleurodiscinae.

B. PuPiLLiNAE ( ? ) ; shell more pupif orm, usually with pupil-
loid teeth (vol. 24, p. vii) ; radula as in A; penis usually
strong ; inferior tentacles usually present.

3. Oviducal cul-de-sac absent; prostate very short; penis
and retractor bifid, although epiphallar arm weakly
demarcated ; shell with few or extravagantly developed
teeth Pupillinae.

4. Oviducal cul-de-sac short; prostate long; penial retrac-
tor (at least) undivided; penis with modified apex
prolonged beyond epiphallar entrance ; shell with angu-
lar lamella (when present) short Orculinae.

5. Oviducal cul-de-sac long; prostate fairly long; penis
and retractor undivided, with epiphallar entrance near
or at apex, which is not modified; shell usually with
longer angular lamella Oastrocoptinae.

C. Vertigininae ( ? ) ; shell as in B, but usually minute with
prominent epidermis; radula usually with fundamentally
tricuspid teeth and with interstitials prevalent through-
out; penis usually weak; prostate very short; inferior
tentacles (always ?) obsolete.

6. Penis with appendix and with bifid retractor

Nesopupinae.

7. Penis without appendix and with undivided retractor

Vertigininae.
In the preceding treatment, complexity of male genitalia
and lack of shell armature have been considered convergent
characters, which are combined in the Valloniinae (sens, lat.),
specifically in the groups Valloniinae (s. s.) and Acanthi-
nulinae. From this standpoint, simplification of the genitalia
is secondary and is usually accompanied by development of

PUPILLIDAE AND RELATED GROUPS.

193

shell armature. [The Pyramidulinae and the Pleurodiseinae
are exceptions to this general rule and may represent the
most primitive conditions, before the evolution of complex
genitalia.]

From the Valloniinae, a line (or progressive tendency),
usually with more calcareous shells, terminates in the Gastro-
coptinae, which have simplified male genitalia and usually
develop complex shell armature, but which retain, except in
Modicella, a radula similar to that in Vallonia. On the other
hand, from the Acanthinulinae, a divergent line, usually with
more epidermal shells, terminates in the Vertigininae, which
also have simplified male genitalia and usually develop shell
teeth, but which tend to produce interstitial cusps (and
entocones) throughout the radula. This idea may be repre-
sented graphically as in fig. 10.

PUPXILIHAC

ViLLLORIIIIAI VfRTISISISAt

Fig. 10.

An alternative hypothesis would be to consider complexity
of shell armature and simplicity of genitalia as the convergent
characters. This would bring the Vertigininae and the Gastro-
coptinae together but would separate the Valloniinae (s. s.)
from the rest of the Valloniinae (sens, lat.) as the terminals
of the two divergent lines.

Strobilopsidae.

The only reason for the retention of this group as a family
distinct from the Valloniinae lies in its shell characters, es-

194 PUPILLIDAE AND RELATED GROUPS,

pecially its internal teeth. In shell form and in peristome,
the sg. Discostrohilops approaches Vallonia; the principal
anatomical distinction of StroMlops is the apical prolongation
of its penis which somewhat resembles either of the two
"diverticula" of Acanthinula. In addition, its radular cen-
tral is about as wide as its first lateral.

Strobilops aenea Pilsbry. Plate 26, figs. 5 to 8.

The animals studied were collected during May, 1928, in
Springfield Township, near Philadelphia, Pa. ; in most of the
individuals of this lot and in all those from Necaxa, Mexico,
the vas deferens ends near the base of the spermatheca and
the other male organs are lacking. Embryos have not been
observed.

Mantle edge (fig. 8) broad; lappets weak; glands encroach-
ing on lung. Lung more than 3 times its base or 4 times
length of kidney. Principal vein very close to kidney ; weak
venation present along columellar side and anteriad. Kidney
(broader, longitudinally trabeculate region) about as long as
its base or length of pericardium ; orthureter about 3 times
as long, similarly thick- walled but with weak transverse trabe-
culae ; recurved groove short.

Ovotestis (fig. 7) imbedded in basal end of apical liver lobe;
consisting of two lobes, each divided into few, short-clavate
lobules; duct thickened below middle in animals with male
organs; carrefour clavate with ovoid base; talon not devel-
oped. Spermatheca clavate, short-stalked and imbedded in
side of oviduct. Vagina very long. Prostate short with
clavate alveoli. Epiphallus (fig. 5) developed in vas deferens,
some distance above entrance into penial side through a low
papilla. Penis bifurcate; appendicular arm clavate and
shorter than epiphallar, which extends beyond entrance of vas
deferens; appendix elongate fusiform and thin- walled, enter-
ing apex of appendicular penial arm through a thickened
ring. Penial retractor arising from diaphragm, bifurcating
below its middle and inserting near apex of appendicular arm
and alongside entrance of vas deferens on epiphallar branch.
Atrium short, opening near right inferior tentacle.

PUPILLIDAE AND RELATED GROUPS. 195

Columellar retractor (left) gives off almost immediately
the shortly united left free and buccal muscles, soon after
the right free retractor and then continues to form the strong
tail fan. Eight free gives off right ocular, which passes
through penioviducal angle, at about level of salivary glands,
and divides terminally into fan of short muscles, of which first
is inferior tentacular, second goes to vagina and penioviducal
angle and remainder to sides of foot.

Cerebral commissure about % as long as width of each
cerebral ganglion ; pleural connective medium in length ; pedal
much longer. Salivary glands long lanceolate, about length
of buccal mass but more slender; right one % to ^ longer
than left and completely separate from it; ducts short.
Oesophagus long and slender (probably knotted when con-
tracted). Stomach without special gland, but alongside in-
ferior lobe of liver which is narrow and terminally bilobed
like in Vallonia (Steenberg, 1917) although intestinal S-loops
are mainly imbedded in albumen gland. Radular formula
(fig. 6):14 — 1— (6 + 8), with 83 rows (T).

Strobilops texasiana Pils. & Ferr. Plate 26, fig. 4.

Cf. S. lahyrinthica Hanna (1922, Naut. 25: 91) and this
vol., p. 13.

The animal described was collected June 18, 1929, along the
Guadalupe River, north of New Braunfels ; the other 23 indi-
viduals in the lot lacked terminal male genitalia. All indi-
viduals in several lots of the related S. lahyrinthica, from
several localities, are similarly emasculate. Only salient dif-
ferences from the structure of S. aenea will be noted.

Carrefour (fig. 4) enlarged tranversely at apex. Vagina
relatively shorter. Epiphallus nearer penis. Penial appen-
dix long clavate (base much contracted). Penial retractor
bifurcating near origin from diaphragm ; insertion on appen-
dicular penial arm farther from apex.

Valloniinae.

Details of the anatomy of Vallonia (oviparous) are given by
Steenberg (1917), whose figure of the genitalia in V. costata

196 PUPELLIDAE AJSTD RELATED GROUPS.

(Miill.) is reproduced (pi. 29, fig. 1), and by Watson (1920).
Neither describes the internal structure of its penis, but its
appendix (ap') appears shorter than in StroMlops (less so in
Watson's fig.) and one branch of its penial retractor (re)
inserts on the epiphallus (ep). Its radular central is little
over half as wide as its 1st lateral.

Anatomical notes on Spelaeodiscus triarius (Rossm.) are
given by Hesse (191-5) and by Soos (1917). If I interpret
the latter 's figures correctly, its oviduct, spermatheca, vagina,
prostate, apical caecum of the epiphallar penial arm and
penial appendix must be extremely like those of Strohilops;
about the only salient differences appear to be the much larger
appendicular arm of its penis, the insertion of one branch of
its penial retractor below the middle of this appendicular
arm, and its radular central, which is about % as wide as the
1st lateral and appears to have vestigial ectoeones. The two
genera are also similar in shell form and texture.

On the basis of anatomy alone, I would include in this
group Vallonia, StroMlops and probably Spelaeodiscus.

ACANTHINULINAE.

This group develops a weak shell with prominent epidermis,
which often develops riblets or projections. It may include
Acanthinula, Planogyra, Zoogenetes and Spermodea, although
the male genitalia of the last two have not yet been published.

Recent anatomical descriptions of Acantlmiula aculeata
(Miill.) are those of Hesse (1915), Boycott (1917) and Steen-
berg (1917) ; Steenberg's figure of the genitalia (oviparous)
is reproduced (pi. 4, fig. 6). It has a spermatheca (be) of
the short type and a short vagina (va). Its penial appendix
is short, has a terminal sac (ap') and enters a basal swelling
(ap) which looks like the apex of the appendicular penial
arm in Planogyra. The epiphallar arm (pe') of the penis is
bifid apically, with the epiphallar (ep) entrance between the
two "diverticula " (di. di'), and with an insertion of the
penial retractor (re) on one of them.

The female genitalia of Zoogenetes (ovoviviparous) are de-
scribed by Steenberg (1925) and those of Spermodea (ovipar-

PUPILLIDAE AJSTD RELATED GROUPS. 197

ous) by Steenberg (1917), Boycott (1917) and Watson (1920) ;
Steenberg's figure of the jaw of Z. harpa (Say) is reproduced
(pi. 31, fig. 7).

Because of its well-developed appendix, Planogyra is added
to this group, although its shell form is nearer that of Pyra-
midula, which it also approaches in its undivided penial
retractor.

Planogyra asteriscus (Morse). Plate 26, figs. 1 to 3.

P. asteriscus Mse. (1864, Journ. Portland Soc. Nat. Hist.
1: 24), head, jaw and radula (Maine) ; H. B. B. (1928, Naut.
41: 122), preliminary account.

All my adults, from Cheboygan County, Michigan, have
male organs. A very few shells are practically ribless, like in
^'Pyramidula rupestris" nylanderi Morse (1920) from Maine,
which appears only a form of P. asteriscus.

Animal almost white, with short and stout, darkly pig-
mented ommatophores ; inferior tentacles short, lighter in
color. Foot holopod but with serrate line formed by upper
edges of lowest row of tesselloid bosses quite prominent ; short
and broad ; sole lanceolate with rounded tip, displaying coarse
(3 to its length) pedal waves in locomotion. Mantle collar
(fig. 3) thick and swollen, protruding beyond peristome, white
and very noticeable; pneumostome, guarded by thickenings
but without distinct shell-lappets, with anus near inner end.
Lung about 2i/2 times base and 1% times length of kidney;
principal pulmonary vein near orthureter ; rectal surface with
inconspicuous venules. Kidney about li/^ times base and
twice length of pericardium ; trabeculae hea\y, becoming
transverse anteriad. Orthureter slightly over % length of
kidney and much narrower although poorly demarcated ; walls
with weak transverse trabeculae; opening near anterior end;
recurved groove apparently open, also transversely trabecu-
late, extending backwards shortly beyond apex of kidney.

Ovotestis (fig. 1) imbedded about % whorls from liver apex,
consisting of two lobes with few, obovoid lobules; duct long,
greatly swollen, convolute in lower half ; earref our hidden by
prostate ; talon scarcely represented. No embryos observed in

198 PUPILLIDAE AND RELATED GROUPS.

uterus. Walls of free oviduct with longitudinal plicae.
Spermatheca ovoid (often more swollen than in figure), im-
bedded near base of uterine enlargement ; stalk medium, colu-
mellar in position. Vagina long with structure of free ovi-
duct. Prostate short with few digitate alveoli. Epiphallus
(fig. 2) swollen with thick pebbled wall; penial papilla
scarcely developed. Penis slender; epiphallar arm short;
appendicular arm almost filled by a large papilla, through
which appendix opens; appendix of medium length, basally
swollen with heavy, transversely trabeculate walls, apically
enlarged into a thin-walled sac containing vacuolate mucus.
Penial retractor arising from diaphragm and inserting around
base of epiphallus (except for heavy muscular sheath, appen-
dicular arm is without retractor, although apex of appendix
seems to be loosely bound to diaphragm. Atrium quite short,
opening very near base of right inferior tentacle.

Free retractor system much as in Strolilops but origin of
right free muscle and separation of left free from buccal mid-
way between columellar origin and root of tail. Left free
muscle branch to atrium. Digestive system also similar
although salivary ducts about as long as glands.

P. CL.vppi (Pilsbry).

The anatomy of animals from near Seattle, Washington,
collected Aug. 18, 1929, is very similar to that in P. asteriscus.

Pyramidulinae.

The anatomy of the ovoviviparous Pyramidula rupestris
(Drap.) has been described by Soos (1917), by Hesse (1918)
and by Watson (1920) ; a figure of the genitalia (pi. 29, fig.
2) has been copied from the last. The spermatheca is not
much longer than the vagina. The penis appears much sim-
plified although the small diverticulum may represent a ves-
tigial appendicular arm.

Pleurodiscinae.

Watson's (1920) figures of the genitalia (pi. 29, figs. 3-5)
of Pleurodiscus halmei (P. & M.) have been copied. Its vagina

PUPILLIDAE AND RELATED GROUPS. 199

is quite short. Its penis proper appears to be small although
heavy, with internal pilasters. Its epiphallus is exceptionally-
well developed with a spirally folded basal part and a longer
apical region with longitudinal folds ; it opens through a prom-
inent penial papUla. Its simple penial retractor inserts on
the side of the basal region of the epiphallus. It is ovovivi-
parous like Pyramidula.

PUPILLINAE.

The anatomical differences between this subfamily and the
Valloniinae seem much less striking than those of the shell.
Pupilla muscorum (L.) Mgranata {B,ossm.) , alpicola (Charp.),
cupa (Jan.) and triplicata (Stud.) have been dissected by
Steenberg (1925) and his figures of the genitalia (pi. 30, fig.
1) and the jaw (fig. 8) of P. muscorum (L) have been copied.
The uterus (ut) is distended with embryos. The spermatheca
has a variously shaped sac (rs) and develops a cyclindrical
diverticulum (di) which is longer in some of the other species.
The penis is bifid; its epiphallar arm (epj is considerably
swollen and extends a short distance beyond the entrance of
the epiphallus (ep) ; and its appendicular arm (ap) is swollen
around the base of its appendix (apa). The bifid penial re-
tractor (rp) inserts on the epiphallus and near the apex of
the appendicular penial arm.

The male genitalia of Lauria cylindracea (DaCosta) as
figured by Steenberg (1925) are very similar, although the
apical caecum of the epiphallar penial arm is better developed
(approaching Orcula). On the other hand, its spermatheca
is much longer and lacks a diverticulum; in addition, the
embryos are said to develop only in the free oviduct. How-
ever, I would neither create for it a distinct subfamily nor
put in a different family from Orcula on these grounds.

The anatomy of Agardhia ferrari (Porro) has been de-
scribed in detail by Steenberg (1929) and is similar to that
of Pupilla. The principal differences are the oviparity of
Agardhia, its longer spermathecal diverticulum, its shorter
vagina, its larger epiphallus which enters at the apex of the
epiphallar penial arm, the very short appendicular and epi-

200 PUPILLIDAE AJSTD RELATED GROUPS.

phallar branches (and trunk) of its penis, its long appendix
(like Lauria) and the peculiar insertions of its penial retrac-
tor at the bases of the two penial arms.

Anatomically, Pupoides differs from Pupilla in the inser-
tion of its penial retractor at the base (instead of on the side)
of the epiphallus, in the simpler epiphallar arm of its penis
(without apical caecum), by the lack of a spermathecal diver-
ticulum and in the narrower radular central. Its genitalia
appear the simplest in the group, but its reduced penial arms
approach those of Agardhia while its spermathecal stalk,
although much shorter, is unbranched like that of Lauria.

Pupoides margin atus nitidulus (Pfr.). Plate 27, figs. 1
and 2.

Cf. P. marginatus Pils. (1921, vol. 26: 108).

Among many specimens from the Dutch Leeward Islands,
collected during the summer of 1922, one individual from
Aruba was found to have mature male organs.

Ovotestis (fig. 1) with three lobes of short-clavate alveoli;
duct enlarged and convoluted in lower half; carrefour elong-
ate without distinct talon. Oviduct swollen in apical half;
no embryos observed. Spermatheca short, without diverti-
culum (although a muscle attaches across its stalk below sac).
Epiphallus sessile, considerably swollen, internally with long-
itudinal folds. Penis bifid but epiphallar arm demarcated
from epiphallus only by narrower caliber and lack of folds;
appendix short, thin-walled, entering penial apex through a
thickened ring (like in Strohilops) . Penial retractor arising
from diaphragm, bifurcating a little below its middle to in-
sert around apex of epiphallar arm and near middle of appen-
dicular arm of penis. Right ocular retractor in penioviducal
angle. Radular formula (fig. 2) : 16 — 1— (8 + 8), with 82
rows (T) ; central much narrower than inner laterals.

The anatomy of Pupoidopsis hawaiensis Pils. & Cooke,
which has been described and figured by C. Montague Cooke,
Jr., and Marie C. Neal (1928), appears to be very similar to
that of Pupoides but the vagina and atrium are shorter and

PUPILLIDAE AND RELATED GROUPS. 201

the penial retractor seems undivided and attached only to
the epiphallus. The radula has a similarly narrow central
but fewer laterals and marginals.

Orculinae.

In this group, two species of Orcula s. s. (oviparous) have
been dissected by Soos (1917) and one by Steenberg (1925),
whose figures of the genitalia (pi. 30, figs. 2, 2a) and jaw
(fig. 7) of 0. dolium (Drap.) have been reproduced. Besides
the characters mentioned in the key, the spermatheca is very
long and stout with its sac (rs) imbedded in the albumen
gland (ga). The major portion of the unforked penis forms
an apical prolongation (ap), with internal folds, beyond the
entrance of the epiphallus (pi to P4) ; this appears to be a
specialized homolog of the apical caecum of the epiphallar
penial arm in Pupilla, etc.). The true appendix is evidently
absent but is present in the following.

In 0. scyphus hatumensis (Ret.) and 0. orientalis (Pfr.)
[Orculella Steenberg], of which the anatomy (oviparous) has
been described by Hesse (1924), the spermatheca is much
shorter, as if the swollen stalk (pdj in 0. dolium had become
the functional sac, and the penis has an appendix although an
appendicular arm appears to be absent {batumensis) or very
short (orientalis). Also, the swollen prolongation of the
penis (or its epiphallar arm) beyond the epiphallar entrance
is long (orientalis, like in dolium) or short Cbatumensis) .
These species cut down the gap in anatomical characters be-
tween Orcula and Lauria or Pupilla considerably and one
wonders what an examination of the residual species may
show.

Moquin-Tandon 's (1855) figure of the male genitalia of
Pagodulina pagodula (Desm.) does resemble the structures in
Orcula, as claimed by Steenberg, but new researches are
necessary.

Gastrocoptinae.

In this group, Modicella avenacea (Brug.) [+ Chondrina
a.] and Abida frumentum (Drap.) have been dissected by
Soos (1917) ; besides these, Steenberg (1925) has studied M.

202

PUPILLIDAE AND RELATED GROUPS.

similis (Brug.), -^. partioti (Moq.), A. secale (Drap.) and
Sandahlia cylindrica (Mich.), From the latter have been
taken figures of the genitalia (pi. 30, fig. 3) and radula (text-
fig. 11) of M. avenacea, of the jaw (pi. 31, fig. 2) of M.
similis, of the genitalia (pi. 31, fig. 1) and jaw (fig. 3) of
Ahida secale and of the genitalia (pi. 30, fig. 5) and carrefour
(fig. 6) of Sandahlia.

OTTVm.,os

n

23

1 a

Fig. 11. Modicella avenacea, after Steenberg.

In all these, the spermatheca (rs) is of the short type (im-
bedded in the uterus) but long-stalked (pd) ; the uterus (ov)
is highly convoluted; the prostate (pr) is about as long as
the uterus; the oviducal cul-de-sac is long and cylindrical;
the penis (pe) is simple (although with a free apical caecum
in some species) and receives the penial retractor (rp) near
its middle, which is often connected by fibers to the epiphallus
(ep) ; and the right free retractor sends branches to the base
of the penis as well as to the atrium. The unicuspid central
and inner laterals of the radula in Modicella distinguish it
markedly from the other two, which are mainly distinguished
by their shell-characters.

PUPILLIDAE AND RELATED GROUPS. 203

Gastrocopta has a relatively smaller although well formed
penis and a shorter spermatheca than the other three genera ;
also, the attachment of its penial retractor is more nearly-
apical and its tricuspid radular central is narrower than in
Ahida or Sandahlia. Their inclusion in the same subfamily
is anatomically supported. So far as known, all the genera
appear to be oviparous.

Because Chondrina Reichenbach (1828) was proposed as a
substitute for Chondrus Cuvier (1817), the two names are
irrevocably inseparable. If Reichenbach 's (1836) mention of
Helix avenacea as the only species be considered either as a
type designation or as a definite "restriction" (Cf. Opinion
81), his action applies to Chondrus as well, which would be-
come the generic name of avenacea. In any case, the term
Chondrininae is untenable.

Oastrocopta (Albinula) armifera (Say). Plate 27, fig. 5

to 7.

The description is founded on animals from Williams
Canyon, near Manitou, Colorado, collected Sept. 19, 1931, but
specimens from near Spearfish, South Dakota, collected a
week earlier, and from near Philadelphia, collected in the
spring, have the same structure. Male organs, although small,
seem to be always developed. No embryos have been observed.

Ovotestis (fig. 5) with three lobes of clavate alveoli; carre-
four elongate and enlarged at both ends. Oviduct with apical
half (uterus) swollen and apically convoluted; basal half
slender; cul-de-sac slender and almost as long as uterus.
Spermathecal sac imbedded near middle of uterus. Epi-
phallus (fig. 6) swollen and internally papillate in first
half; slender and simpler towards entrance into penis which
is without definite papilla. Penis small and slender; swollen
apically, where it has thick walls and is prolonged some dis-
tance beyond epiphallar entrance. Penial retractor arising
from diaphragm and inserting slightly below epiphallus.
Atrium short. Right inferior as well as ommatophoral re-
tractors passing through penioviducal and attached to both
sides and to vas deferens by slips of muscle; right free re-

204 PUPILLIDAE AND RELATED GROUPS.

tractor with large branch to atrium. Radular formula (fig.
7) : 18 — 1— (8 + 10), with 82 rows (T) ; central about half
width of 1st lateral, with short reflection and 3 small cusps;
marginals with prominent mesocone and with few ectocones.

Gastrocopta (s. s.) octonaria Pilsbry. Plate 27, figs. 3
and 4,

The animals are from Bonaire, Dutch West Indies, collected
August, 1922 ; they may be stout examples of G. curacoana
Pilsbry (teeth of shell destroyed to see animals and vice
versa). Only salient differences from G. armifera are noted.

Free oviduct (fig. 3) stouter; vagina relatively longer.
Epiphallar entrance nearer apex of penis. Radular formula
(fig. 4): 12 — 1 — (4-|-8); marginals usually broader and
often with more ectocones.

Nesopupinae.

This group is evidently most closely related to the Vertig-
ininae but the gap between it and the Acanthinulinae is almost
bridged (except in radular characters) by the genera Pupi-
soma and Pronesopupa. Besides these, it includes Lyropupa,
Nesopupa and Bothriopupa. Ovoviviparity appears common
and may be universal.

Nesopupa (Indopupa) moreleti (Brown) has been studied
by Steenberg (1925) whose figure of the jaw is copied (pi. 31,
fig. 6), and two species of Pupisoma by myself (1927) ; in
plate 28, figures of the jaw (fig. 3) of P. mediamericanum
Pils. and of the pallial complex (fig. 5), the genitalia (fig. 4),
penis with accessories (fig. 7) and radula (fig. 6) of P. comi-
colense H. B. B. have been reproduced. In Pupisoma, the
orthureter has a recurved duct. The uterus, filled with
embryos, is very long but the free oviduct, spermatheca, vagina
and prostate are short. The epiphallus enters through a
small papilla into the bifid penis, with its appendicular arm
poorly demarcated from the short appendix. The branched
retractor inserts on the apices of both penial arms. The jaw
consists of narrow, overlapping plates. All radular teeth are
fundamentally tricuspid but develop many accessory cusplets.
I am now dubious about the presence of inferior tentacles in

PUPILLIDAE AND RELATED GROUPS. 205

Pupisoma; preserved specimens of Lyropupa also appear to
have inverted ones, each with a definite retractor, although
their absence has been observed in living animals.

Pronesopupa (Edentulopupa) admodesta (Mighels). Plate
28 figs. 1 and 2.

Two specimens from Nuiianu, Oahu, collected by Dr. C. M.
Cooke, has been so long in alcohol and were so full of embryos
that only the penis is figured.

Pallial complex very like that of Pupisoma comicolense; re-
curved ureteric duct closed over for at least part of its length.
Female genitalia similar to those of Lyropupa perlonga, with
large embryos even in vagina. Male organs (fig. 1) excep-
tionally large. Penis bifid ; epiphallar arm demarcated from
epiphallus by a swollen constriction; base of appendix with
thickened walls and entering through a small papilla. Penial
retractor arising from diaphragm and inserting near bases of
both appendix and epiphallus. Radular formula (fig. 2) :
11 — 1 — (5 + 6), with 106 rows; central tricuspid with an
accessory on each ectocone; inner laterals tricuspid with ec-
toconal accessories ; marginals usually fundamentally tricuspid
but with many interstitials.

The penial appendix in Pronesopupa is better demarcated
than in Pupisoma; the differentiation of the male genitalia in
both genera is in marked contrast to the weak development in
Lyropupa, Nesopupa and Bothriopupa.

Pronesopupa (s. s.) acanthinula (Ancey).

Specimens from Nuuanu, Oahu (Cooke) have the same
radular formula and very similar teeth to P. admodesta.

Lyropupa (Mirapupa) perlonga (Pease). Plate 28, figs.
10, 11.

A specimen from Luakaha, Nuuanu Valley, Oahu, collected
Aug. 3, 1933 by Dr. Pilsbry, has male genitalia.

Ovotestis is in newcomhi; carrefour (fig. 10) ellipsoid.
Uterus of non-gravid example with an apical, spongy swelling,
continued by a longer one that appears glassy. Spermatheca
of short type with sac imbedded in uterus ; vagina quite long.

206 PUPILLIDAE AND RELATED GROUPS.

Prostate short. Male organs small and poorly differentiated ,•:
appendix weakly marked off from penial apex; epiphallus
apparently entering directly into side of penis. Penial re-
tractor bifid; arising from diaphragm and inserting on
appendix and epiphallus. Atrium very short. Radular
formula (fig. 11): 10 — 1 — (4 + 6); central half again as
broad as 1st lateral, tricuspid but ectocones with one or two
accessories; laterals and marginals fundamentally tricuspid
but with an increasing number of accessories or interstitials.
The remarkably broad central of Lyropupa appears to be
its most distinct anatomical character. Its genitalia seem very
similar to those of Nesopupa as figured by Steenberg (1925).

Lyropupa (s. s.) microtrauma Ancey.

An animal from Nuuanu, Oahu (Cooke; ANSP. 117154)
has the same radular formula and very similar teeth.

Nesopupa (Limbatipupa) newcombi (Pfr.). Plate 28, figs.
12, 13.

An animal from Kalihi, Oahu (Cooke: ANSP. 117154)
has similar (ovoviviparous) anatomy to that in L. perlonga
and only differences will be noted.

Ovotestis (fig. 13) not lobed, with short-clavate alveoli;
carrefour more elongate. Spermatheca longer with sac near
apex of uterus. Penial retractor inserting around appendix
on apex of its penial arm. Radular formula (fig. 12) : 9 — 1
— (4 -j- 6) ; marginals often with fewer cusps. Jaw (f. multi-
dentata) with 11 plaits; similar to that of N. moreleti (pi. 31,.
fig. 6) in structure.

This species has a broad central like Lyropupa. Similar
radulae were found in dried-in animals of f. multidentata C.
& P. from Glen Ada, Nuuanu, Oahu (ANSP. 44717; same
formula, with 97 rows) and of N. kauaiensis Ancey from Kipu^
makai of Gap, Kauai (ANSP. 44743; one more marginal).

Nesopupa (Nesodagys) w^esleyana Ancey. Plate 27, fig. 8.

An animal from Kalihi, Oahu (Cooke; ANSP. 117155) has
the radular formula (fig.): 10 — 1 — (4 -|- 6) ; central and

PUPILLIDAE AND BELATED GROUPS. 207

first lateral subequal ; other laterals smaller ; marginals broad
with numerous interstitials. This is very similar to the radula
of N. (Indopupa) moreleti (Brown) as figured by Steenberg
(1925) except for his abnormal inner laterals. The central
in these two and the next species is very different from that
in Lyropupa and Limhatipupa.

Nesopupa (Nesopupilla) plicifera Ancey.

Four ovoviviparous animals from Nuuanu, Oahu (Cooke:
ASP. 117149) lack male genitalia; one gives the radular for-
mula : 12 — 1 — (5 + 7) ; teeth as in preceding species.

Nesopupa (Cocopupa) cocosensis (Dall). Plates 28, figs.
8, 9.

The anatomy of specimens from Wafer Bay, Cocos Island,
collected by Dr. Pilsbry on the Gifford Pinchot Expedition, is
very similar to that of N. newcomhi and only differences will
be noted.

Vagina (fig. 8) shorter and stouter. Penial retractor
scarcely branched although with both insertions. Radular
formula (fig. 9) : 16 — 1 — (9 + 7) with 106 rows; central
and lateral cusps shorter and more aculeate.

The radula of this species looks more like that of Bothrio-
pupa, both in relative breadth of the lateral field and in cusp-
shape, than like that of the Hawaiian Nesopupae. Incident-
ally, Ochrodermella, from the same locality, belongs in the
Ferussaciinae (Achatinidae) and has anatomy similar to that
of NeosuhuUna from the Dutch West Indies.

BoTHRiOPUPA tenuidens (C. B. Adams). Plate 27, figs, 9, 10.

A specimen from foothills behind the reservoir for Port
Antonio, Jamaica, collected Aug. 21, 1933, has very small male
genitalia (fig. 10). The anatomy (ovoviviparous) is very
similar to that in Nesopupa and the generic status of the latter
must depend on its shell characters.

Radular formula (fig. 9): 13 — 1 — (7 + 6); central and
first lateral subequal with slender mesocones and 2-4 ectoconal
accessories.

208

PUPILLIDAE AND RELATED GROUPS.

Vertigininae.

In this group, the anatomy of Columella edentula (Drap.)
was studied by Hanna (1912) and Vertigo pusilla Miill., anti-
vertigo (Drap.), substriata (Jeffr.), pygmaea (Drap.), moul-
insiana (Dup.) and alpestris Aid., Truncatellina rivierana
(Bens.) and C. edentula were dissected by Watson (1923).
Steenberg (1925) gave more careful accounts of all these
except V. alpestris and added V. (Vertilla) angustior Jeffr.,
T. cylindrica (Fer.) and T. costulata (Nilss.) ; his figures of
the genitalia of V. pusilla (pi. 31, figs. 4, 5), T. cylindrica
(fig. 9) and C. edentula (fig. 8) are reproduced, also that of
the radula of C. edentula (text, fig. 12, 2). Male organs are
commonly absent. Oviparity prevails.

^o

<f^^^

o

I i^^

Columella edentula Drap.

XI 500

\/\maIIID(

'C=:^

Truncatellina monodon.

Fig. 12. — 2, Columella edentula, after Steenberg. 3. Truneatellina
monodon, after Odhner.

PUPILLIDAE AND RELATED GROUPS. 209

The uterus (ov) is simple in Vertigo but more or less folded
in the other two genera. The free oviduct (ol) is long in
Vertigo, short or absent in the others. The spermatheea (rs)
is of the short type, imbedded in the uterus, but has much the
longest stalk in Vertigo. The prostate (pr) appears almost
vestigial. The epiphallus (ep) is poorly developed but re-
ceives the penial retractor (rp) in Columella (penial entrance
subapical) and in Vertigo (entrance apical) but is obsolete in
Truncatellina, in which the retractor inserts on the penial
apex. The radular teeth of Vertigo are fundamentally tri-
cuspid and develop numerous interstitial cusplets, but the
laterals of Columella and Truncatellina have only two large
cusps and interstitials appear to be lacking on the central (in
Truncatellina on the inner laterals as well ; text fig. 12, 3, T.
monodon Held, after Odhner, 1924). While Vertigo does
seem to form a group apart from the others, the differences
seem of no more than generic importance.

Bibliography op Papers Cited.

Baker, H. Burrington: 1927, Proc. Acad. Nat. Sci. Philadel-
phia 79 : 223.
Boycott, A. E.: 1917, Proc. Malac. Soc. London 12: 221.

1917, Jour, of Conch. 15 : 175.

Cooke, C. Montague, Jr., and Marie C. Neal. 1928, B. P.

Bishop Museum Bull. 47 : 28.
Hanna, G. Dallas : 1912, Proc. U. S. Nat. Mus. 41 : 371.

1922, Naut. 35 : 91.
Hesse, P. : 1915, Nachr. Deutsch. Mai. Ges. 47 : 55.

1918, Nachr. Deutsch. Mai. Ges. 50: 110.

1924, Arch. MoUusk. 56 : 1.

Moquin-Tandon, A. : 1855, Hist. nat. Moll. terr. fluv. France.
Odhner, Hj. : 1924, Arkiv for Zoologi 16, No. 29 : 3.
Soos, L. : 1917, Ann. Mus. Nat. Hungarici 15 : 1.
Steenberg, C. M. : 1917, Vidensk. medd. f ra Dansk. naturh.
Foren. 69 : 1.

1925, Vidensk. Medd. fra Dansk. naturh. Foren. 80:1.
1929, Vidensk. Medd. fra Dansk. naturh. Foren. 88 : 15.

Watson, Hugh: 1920, Proc. Malac. Soc. London 14: 1.
3923, Proc. Malac. Soc. London 15: 270.

210 EXPLANATION OP PLATES, VOL. 28.

EXPLANATION OF PLATES.

Plate 1.

figures page

I, 2. Strohilops lahyrinthica {^ay) . Philadelphia 20
3, 4. Strobilops lahyrinthica Say). Type speci-
men 20

5. Strobilops lahyrinthica ( Say ) . Kittery,York

Co., Maine. 114775 20

6. Strobilops labyrinthica (Say). Philadel-

phia. 27281 20

7. Strobilops labyrinthica (^Siy) . Ann Arbor,

Mich. 122 20

8,9. Strobilops labyrinthica {Say) . Silver Lake,

Kansas. 64500 20

10. Strobilops labyrinthica {Say) . Silver Lake,

Kansas. 58319 20

II. Strobilops labyrinthica {Say.) Dallas Lake,

Wright Co., Minnesota. 67353 20

Plate 2.

1-4. Strobilops texasiana floridana Pils, Type

and paratypes, Miami, Fla. 77044 26

5-10. Strobilops texasiana P. & F. Type (figs. 7)

and paratypes; near New Braunfels,

Texas. 64498 24

Figures 2, 6, 11, X 15 ; the others X 13.

Plate 3.

1-5. Strobilops affinis Pils. Upper Red Hook,
Dutchess Co., N. Y. 74414. Figs. 1, 2
the type 27

6-11. Strobilops aenea Pils. Cazenovia, N. Y.,

90785. Figs. 7, 8 the type 29

12. Strobilops aenea Pils. Brandywine Valley,

Delaware. 85244 29

1-5. Strobilops affinis Pils. Figs. 1, 2, the type ;

fig. 5, base of young specimen 27

6-11. Strobilops aenea Pils. Cazenovia, N. Y.,
90785. Figs. 6, 7, the type; fig. 8, 9,
young specimens ; fig. 10, edge of parietal
lamella 29

12. S. aenea. Brandywine Valley, Delaware.

85244 29

explanation of plates, vol. 28. 211

Plate 4.
figures page

1. Strohilops salvini Tristr. After von Mar-

tens 37

2. S. aenea Pils. Parietal lamellae, near their

inner ends 29

3. 4. S. lahyrinthica form virgo Pils. Type 23

5. S. veracruzensis crossei Pils. After Strebel 36

6. S. veracruzensis Pils. After Strebel 35

7. 8. S. lahyrinthica Say. Genitalia and kidney,

after Hanna 1

9, 10, 11. 8. lahyrinthica Say. Living animal and

teeth. Pilsbry del 1

Plate 5.

1-4. Strohilops streheliFfr. Mirador. 11546.. 33

5. S. strebeli guatemalensis Hinck. Young.

9212a 34

6-8. S. streheli guatemalensis Hinck. Type 9272 34
9-13. S. aenea mexicana Pils. El Diente, near
Monterey, Mexico. 77194. Figs. 10, 11,
the type ; fig. 12, young example 32

Plate 6.

1-3. Strohilops helleri Dall. Type. 108515

USNM 41

4. 6. S. morsei Dall. Type. 24013 USNM .... 39

5. S. morsei Dall. Venezuela. 22923 ANSP. 39
7-11. 8. hrasiliana F. Bkr. Figs. 7, 8 the type.

Fig. 11, edge of the parietal lamella 40

Plate 7.

1-3. Strohilops huhhardi Brn. Calhoun Co.,

Texas. Type. 124 47

4. Strohilops huhhardi. Valles, San Luis

Potosi. 107580 50

5. Strohilops huhhardi. Tampico. 96558 ... 50

6. Strohilops huhhardi. El Abra, Vifiales,

Cuba 49

7-9. Strohilops huhhardi. Bellevue, Jamaica.

11523 48

10-12. Strohilops huhhardi. Admiral's Cave, Ber-
muda, 131597 49

212 explanation op plates, vol. 28.

Plate 8.

FIGURES PAGE

1-7. Strolilops huhhardi Brn. Miami, Florida.

77087 and 82829. (Figs. 1, 2, 3, 4 are
the form named stcvensoni. Fig. 5, diam.
1.2 mm., and fig. 6, immature specimens). 48

8, 9. Strohilops huhbardi. Lake Helen, Florida.

65371 48

10-13. Strohilops iiniplicata Sdbgr. Budenheim.
4153. Fig. 12, spire removed, showing
lamellae by transparence through base of
penult whorl 9, 46

Plate 9.

1, 2, 3. Strohilops veracruzensis Pils. Type and

paratype 35

4, 4a. Strohilops v. crossei Pils. After Fischer &

Crosse 36

5, 6, 7. Strohilops aenea spiralis Pils. Type, Maga-

zine Mt., Ark. 91331 31

8, 9. Strohilops aenea f . micromphala Pils. Char-

lieville, La. 98626 30

10, 11. Strohilops v. parietalis Pils. Ardsley, Pa.

105695 24

Plate 10.

1, 2, 3, Strohilops diodontina Hde. After Heude 43

4-7. Strohilops hirasei Pils. Type and paratype.

95251 44

8-10. Strohilops coreana Pils. Type. 99967 ... 43

11-14. Strohilops nipponica Pils. Type 45

Plate 11.

1-4. Strohilops hoholensis Gude. 62067 54

4a. Strohilops hoholensis Gude. After Gude.. 54

5a-5c Strohilops trochospira MUdff. After Moell-

endortf 52

6, a-e. Strohilops trochospira Mlldff. After Gude 52

7, Strohilops hoholensis Gude. Parietal lam-

ella 55

8-10. Strohilops hoholensis Gude. 78457, 32117 54

11-14. Strohilops quadrasi Mlldff. 96492-3 ; at 13

the inner ends of the parietals 55

explanation of plates, vol. 28. 213

Plate 12.
figures page

1-3. Dimorphoptychia arnouldi (Mich.). After

Sandberger 11

4-6. Kanahohelix kanabensis (White). After

White 11

7-8. Pseudostrohilus riethmulleri Oppenh. After

Oppenheim 9

9. Strohilops hubhardi A. D. Brn. Lake Helen.

65371 50

10. Ohhi7iula anthracophila Stache. After

Stache 10

11-13. Strohilops romani Wenz. Hauterive. 2800 8

14-16. Strohilops polyptychia Mlldff. 114168 .... 52

Plate 13.

1-7. Strohilops wenziana Pils. Type and para-
types 60

8-11. Strohilops piratica Pils, Type and para-
types 58

Plate 14.

1, la, 2, 3. Fauxulus hurnupianus Pils. Type and

paratypes, 144265, 144264 73

4, 4a. Fauxulus falconianus Pils. Type. 145477. 75

5, 6. Gastrocopta thomasseti Pils. Type. 145474 71

Plate 15.

1. Vertigo wheeleri Pils. Type 96

2. Vertigo loessensis Bkr. {=V. elatior St.).

Paratype, 145107 93

3. Gastrocopta pilshryana Sterki (Holotype of

Pupilla stoneri Cham. & Jones) 64

4. Sterkia c. martiniana Pils. Type 110

5. Vertigo gouldii paradoxa St. Newfoundland.

150044 95

6. Pupisoma comicolense H. B. B. Type .... 115

7. Vertigo californica guadalupensis Pils.

Type 97

8. Vertigo californica catalinaria Pils. Type 97

9. Vertigo degeneris Pils. Type. Guadalupe

Island. 141405 98

10. 11. Pupilla guadalupensis Pils. Type and

paratype. 141452 83

12. Pupilla goniodon Pils. Type. 141410 82

FIGURES

1,

la.

2,

2a.

3,

3a, 4,

5.

6.

7,

8, 9.

1,

2.

3,

4.

214 explanation of plates, vol. 28.

Plate 16.

PAGE

Strohilops helleri (Ball). Narborough

Island 62

Strohilops helleri (Dall). James Island .. 62

Strohilops hannai Pils., type and paratype 59

Pupisoma mcneilli Clapp. Cotype, 128335 115

Bothriopupa peruviana Pils. Type 114

Pupisoma galapaganum Pils., Type, 152716,

and paratype, 152314 114

Plate 17.

Gastrocopta duncana Pils. Type, 152689 65
Gastrocopta munita f . hr^mnea Pils. Kicker

Rock. 152701 69

5, 6. Gastrocopta munita f. hrunnea Pils. (Para-
type of P. reihischi Dall, 1928, not 1919).
Tagus Cove. 5613 68

7. 8, 9. Gastrocopta munita (Reib.). Albemarle,

152702 67

10, 11, 12. Gastrocopta clausa (Reib.). Albemarle.

152715 70

13, 14, 15. Gastrocopta clausa ( Reib. ). Chatham.

152713 70

16, 17. Gastrocopta clausa (Reib). Duncan I.,

151479 70

18. Gastrocopta clausa (Reib.). Charles I.

152703 69

19. Gastrocopta clausa (Reib.). Duncan I.,

151653 69

Plate 18.

1. Gastrocopta pazi (Hid.) . After Moros ... . 64

2. Gastrocopta dahli Thiele. After Thiele . . 72

3. Gastrocopta semiclausa Thiele. After Thiele 73

4. Granopupa marmonchana Pallary. After

Pallary 77

5. 6, 7. Ahida variahilis villarensis Pier. After

Piersanti 76

8. Pupoides karachiensis Peile. After Peile . . 80

9. Lauria wouramhoulchiensis Con. After

Connolly 86

10. Lauria leprevieri Pallary. After Pallary . 86

11. 12. "Terftfifo ^sc/joA'/tTi^arctica Wallenb. After

G. Mermod 103

EXPLANATION OP PLATES, VOL. 28. 215

FIGURES PAGE

13, 15, 16. Vertigo genesii Gredler. Ireland. After

Phillips 104

14. Vertigo genesii Gredl., Topotype, Salten.

After Phillips 104

17. Vertigo genesii geyeri Lindh. After Lind-

holin 104

Plate 19.

1, 2. Orcula kaznakovi Rosen. After Rosen .... 91

3, 4. Orcula gularis pseudodolium Wagn. After

Wagner 90

5. Orciila dolium pseudogularis Wagn. After

Wagner 88

6. Orcula dolium tatrica Wagn. After Wagner 89

7. 8. Orcula tingitana Pallary. After Pallary . . 91

9. Orcula tolmi7iensis Wagn. After Wagner 90

10, 10a, 11. Orcula orientalis antiochensis Pils. Type

and paratype 93

12, 13, 13a. Orcula o. moussoni Reinh. Aleppo 93

Plate 20.

1. Ahida polyodon montserratica f. monsjovica

Bof . After Aguilar-amat 122

2. Ahida catalonica valida Bof. After Aguilar-

amat 122

3. Ahida p. m. f, vidali Bof. After Aguilar-

amat 122

4. Chrondrina clienta caucasica Ehrm. x 7.5.

After Ehrmann 79

5-7. Orcula fuchsi Zimm. After Zimmermann 89

8. 8a. Ahida hrauniopsis Bof. After Aguilar-

amat 122

9. Solatopupa similis monterosatoi Bof. After

Aguilar-amat 123

10. Chondrina higorricnsis tenuimarginata f.

flaccida Bof. After Aguilar-amat 123

11. Chrondrina higorriensis t. f. macrochilus

Bof. After Aguilar-amat 123

12. Chondrina avenacea suhcornea Bof. After

Aguilar-amat 122

13. Chondrina dertosensis farta Bof. After

Aguilar-amat 123

216 explanation of plates, vol. 28.

Plate 21.
figures page

1-3. Truncatellina sundleri Odhner. After

Odhner 106

4. Fauxulus harnardi Conn. After Connolly 121

5. Nesopupa novopommerana Rensch. After

Rensch 112

8. Nesopupa rodriguezensis Conn. After

Connolly 110

9. Nesopupa vengoensis Connolly. After

Connolly Ill

12. Orcula tomlini Conn. After Connolly .... 133

13. Bifidaria hannertonensis {= G. pediculus).

After Gabriel 118

14. Fauxulus fryanus (Bens.). After Connolly 121

15. ''Pupa'' lata C.B. Ad. Type 135

16. 17. Vertigo milium Gld. Santo Domingo 101

Plate 22.

1. Columella pygmaeorum P. & C. Type .... 109

2. Oastrocopta niole Fulton. Paratype 119

3-6. Oastrocopta euryomphala Pils. Type and

paratype. 160431 120

7. Oastrocopta p. Jiordeacella Pils. California.

161037 117

8. Vertigo modesta Say, var., Mona, St. Louis

Co., Missouri. No. 160361

7-11. Vertigo idahoensis Pils. Type. 158670 . . 100

12-14. Vertigo gouldi huhrichti Pils. Type and

paratype. 160362 99

Plate 23.

1, la, 1&, 2. Pyramidula humilis Hutt. & Bs. Landour.

1336 188

3-3c. Pyramidula conica Pils. Type. 83884 187

4, 4a, 46. Punctum micra (Pils.). Type. 79557. .. 190
5-5c. Punctum orphana (Heude). 49501 189

Plate 24.

1. Vertigo torrei Aguayo & Jaume. After

Aguayo 98

2-4. Vertigo modesta f. sciilptilis Pils. Type

and paratypes. 162884 100

5, 6. Oastrocopta pilshryana amissidens Pils.

Type and paratypes. 161437 118

EXPLANATION OF PLATES, VOL. 28. 217

FIGURES PAGE

7, 8, 9. Vertigo teilhardi Ping & Yen. After Ping

and Yen 101

10. Pupilla pseudocylindrica Ping & Yen, After

Ping and Yen 136

11. Pupilla alpicola Ch. = ciipa Jan. After

Mermod 135

12-14. Lauria hequaerti Pils. Cotypes 138

15. Pupisoma bailyi Pils. Type. 156099 .... 116

Plate 25.

1. Columella ninagongonis Pils. Type 172

2, 3. Lauria karisiinhiensis Pils. Type 170

4, 5. Pyramidula rupestris jaenensis Cless. After

Clessin 185

6, 7. Pyramidula rupestris conoidea Bgt. (== var.

seotaea Bgt. ) . After Bourguignat 185

8, Pyramidula rupestris rupicola Stabile. Pisa.

47692 185

9, 9a, 96. Pyramidula rupestris Drap., typical. Grasse.

47685 183

10, 10a. Pyramidula rupestris saxatilis Hartm.

Lausanne. 47684 184

11, Pyramidula rupestris chorismenostoma Bl.

& Wstl. Beotia. 47696 186

12, 13. Pyramidula hierosolymitana Bgt. After

Bourguignat 187

14. Pleurodiscus sudensis Pfr. Akrotiri, Crete.

153775 181

15. Pleurodiscus halmei P. & M. Palermo.

152807 179

16. Pleurodiscus erdeli Roth. Beirut. 152808 180

17. Pleurodiscus s. cypria Kob. After Kobelt. 181

Plate 26.

In this and next two plates, scales are collected at bottom
of page and represent .02mm. for figs, of radular teeth, .1 mm.
for radular transverse rows (T) and jaws, and 1 mm. for all
others.

A, penial appendix. C, carrefour. D, vas deferens. E, epiphallus.
G, albumen gland. H, hermaphroditic duct. L, radular laterals. M,
radular marginals. O, ovotestis. P, penial trunk. PA, appendicular
arm. PE, epiphallar arm. PR, penial retractor. PT, prostate. R,
radular central. S, spermatheca. T, outline of radular transverse row
(right half). U, oviduct (uterus -f- free oviduct. UC, oviducal cul-de-sac.
V, vagina. X, atrium.

218 EXPLANATION OF PLATES, VOL. 28.

FIGURES PAGE

1-3. Planogyra asteriscus Mse. Douglas Lake,
Michigan. 1, genitalia. 2, penis and ac-
cessories from glycerin jelly mount. 3,
internal view of pallial complex 197

4. Sirohilops texasiana Pils. & Ferr. New
Braunfels, Texas. Genitalia (ovotestis
omitted) 195

5-8. StroMlops aenea Pils. Philadelphia, Penna.

5, penial apex from glycerin jelly mount.

6, radula. 7, genitalia. 8, internal view

of pallial complex 194

Drawn by H. B. B.

Plate 27.
Scales and labels as in plate 26.

1, 2. Pupoides marginatus nitidulus Pfr. Aruba,
Dutch West Indies. 1, genitalia. 2,
radula 200

3, 4. Gastrocopta octonaria Pils. Bonaire, Dutch
West Indies. 3, terminal genitalia. 4,
radula 204

5-7. Gastrocopta armifera Say. Williams

Canyon, Colorado. 5, genitalia. 6, penis
and accessories from glycerin jelly mount.

7, radula 203

8. Nesopupa wesleyana Anc. Kalihi, Oahu.

Radula 206

9, 10. Bothriopupa tenuidens C. B. A. Jamaica,

British West Indies. 9, radula. 10, ter-
minal genitalia 207

Drawn by H. B. B.

Plate 28.
Scales and labels as in plate 26.

1, 2. Pronesopupa admodesta Migh. Nuuanu,
Oahu. 1, terminal genitalia from glycerin
jelly mount. 2, radula 205

3. Pupisoma mediamericanum Pils. Necaxa,

Mexico. Jaw 204

4-7. Pupisoma comicolense H. B. B. Necaxa,

Mexico. 4, genitalia. 5, internal view of
pallial complex. 6, radula. 7, penis and
accessories from glycerin jelly mount . . . 204

EXPLANATION OF PLATES, VOL, 28. 219

FIGURES PAGE

8, 9. Nesopupa cocosensis Dall. Cocos Island. 8,

terminal genitalia. 9, radula 207

10, 11. Lyropupa perlonga Pse. Nuuanu, Oahu. 10,

genitalia (ovotestis omitted). 11, radula 205

12, 13. Nesopupa newcomhi Pfr. Kalihi, Oahu.

12, radula. 13, genitalia 206

Drawn by H. B. B.

Plate 29.

ap, appendicular arm of penis (penial caecum in pi. 5, fig. 8). ap' or
apg, penial appendix, be, spermathecal sac. eg, cl, atrium, dh, herma-
phroditic duct, ep, epiphallua. ep^, epiphallar arm of penis, ga, albu-
men gland, gh, ovotestis. ol, free oviduct, ov, uterus, pd, spermathecal
stalk, pf, carrefour. pr, prostate, re, rp, penial retractor, rs, sperma-
thecal sac. ut, uterus, va, vagina, vd, vas deferens.

FIGURES PAGE

1. Vallonia costata Miill. Genitalia after Steen-

berg (1917) 195

2. Pyramidula rupestris Drap, Genitalia after

Watson (1920) 198

3-5. Pleurodiscus halmei P. & M. After Watson
(1920). 3, genitalia. 4, penis and ac-
cessories. 5, spermatozoon 198

6. Acanthinula aculeata Miill. Genitalia after

Steenberg (1917) 196

Plate 30.

All figures after Steenberg (1925). Labels
as in plate 29.

1. Pupilla nniscorum L. Genitalia 199

2. 2a. Orcula dolium Drap. 2, genitalia. 2a, base

of spermoviduct with cul-de-sac (C SO) .. 201

3. Modicella avenacea Brug. Genitalia. 4,

stomach 202

5, 6. Sandahlia cylindrica Mich. 5, genitalia.

6, carrefour 202

7. Orcula dolium Drap. Jaw 201

8. Pupilla muscorum L. Jaw 199

Plate 31.

All figures after Steenberg (1925). Labels
as in plate 29.
1. Ahida secale Drap. Genitalia 202

220 EXPLANATION OF PLATES, VOL. 28.

FIGURES PAGE

2. Modicella similis Briig. Jaw 201

3. Ahida secale Drap. Jaw 202

4. 5. Vertigo pusilla Miill. 4, genitalia. 5,

spermatheca 208

6. Nesopupa moreleti Brown. Jaw 204

7. Zoogenetes harpa Say. Jaw 196

8. Columella edentula Drap. Genitalia 208

9. Truncatellina cylindrica Fer. Genitalia . . 208

INDEX

Abida Leach, 122, 202
Aoanthinula Beck, 12, 174
Acanthinulinae, 196
aculeata Miill. (Acanthinula), 116
adamii Bgt. (Sphyradium), 172
aenea Pils. (Strobilops), 29, 194
affinis Pils. (Strobilops), 27
Agardhia Westerl., 199
aigenensis Tauch, (Helix), 10
aigenensis Taueh. (Pseudostro-

bilus, 10
albina Bttg. (Pupa), 86, 139
alluaudi Germ, (Lauria), 170
alpicola Charp. (Pupa), 135
amblya Bgt. (Orcula), 92
Amplexis Brown, 174
Amplexus Brown, 174
amissidens Pils. (Gastrocopta), 118
anthracophila Stache (Helix), 10
anthracophila Staehe (Obbinula), 10
Anthracopupa Whitf., 117
antiUarum Pils. (Vertigo), 101
antiochensis Pils. (Orcula), 92
antonini Mich. (Helix), 183
arctica Wall. (Vertigo), 103
armata Pse. (Nesopupa), 112
armifera Say (Gastrocopta), 203
arnouldi Mich. (Dimorphoptychia),

11
arnouldi Mich. (Helix), 11
asteriscus Mse. (Planogyra), 197
austriaca Zim, (Orcula), 132
avenacea Brug. (Chrondrina), 78

B

bailyi Pils. (Pupisoma), 116
balmei P. & M. (Helix), 180
balmei P. & M. (Patulastra), 180
balmei P. & M. (Pleurodiscus), 179
bannertonensis Gabr. (Bifidaria),
118

barnardi Conn. (Fauxulus), 121
bequaerti Pils. (Lauria), 138
bilamellatus Cless. (Strobilus), 5
boettgeri Andr. (Strobilops), 5
boettgeri Andr. (Strobilus), 5
boholensis Gude (Plectopylis), 54
boholensis Gude (Strobilops), 54
Bothriopupa Pils., 207
bourguignati(Pagodina), Cout., 108
brasiliana F. Bkr, (Strobilops), 40
brauniopsis Bof. (Abida), 122
britannica Cox(Anthracopupa), 117
brunescens Wenz. (Strobilops), 55
brunnea Pils. (Gastrocopta), 69
burnupianua Pils. (Fauxulus), 73

eatalinaria St. (Vertigo), 97
caucasica Ehrm. (Chrondrina), 79
Cavipupa Pils., 120
cereana Muhlf. (Pupa), 78
Chondrina Rch., 122, 203
chorismenostoma Bl. (Helix), 186
chorismenostoma Bl. (Pyramidula),

186
Circinaria Beck, 174
clappi Pils. (Planogyra), 198
clausa Reib. (Gastrocopta), 69
clausilioides Boub. (Abida), 77
clavella Eeinh. (Truncatellina), 108
clienta West. (<Uhondrina), 79
Coelostrobilops Pils., 57, 60
coloradensis CkU. (Vertigo), 95
Columbiana St. (Vertigo), 97
Columella West., 208
comieolense H. B. B. (Pupisoma),

115
conica P. & H. (Pyramidula), 187
conica Rm. (Orcula), 132
concinnula Wheeler (Vertigo), 96
conoidea Bgt. (Helix), 184
coreana Pils. (Strobilops), 43

(221)

222

INDEX, VOL. XXVIII.

corrugata Prest. (Nesopupa), 110
cossmanni Wenz (Strobilops), 7
cofitata Cless, (Strobilops), 5
costata Cless. (Strobiius), 5
costata K. Mill. (Strobiius), 7
costigerella Lindh. (Truncatellina),

105
critica Zel. (Pupa), 92
crossei Pils. (Strobilops), 36
cupa Jan. (Pupilla), 85, 135
curdoofensis Cless. (Strobiius), 5
cylindrata Bttg. (Pupa), 85
cylindrica Colb. (Vertigo), 100
eylindrica Fer. (Truncatellina), 105
eypria Kob. (Patula), 181

D

dahli Thiele (Gastrocopta), 72
dalmatina Cless. (Patula), 184
degeneris Pils. (Vertigo), 98
depressa "Wenz (Strobilops), 9
desertorum Bgt. (Oreula), 172
desmarestina Brong. (Helix), 6
desmarestina Brong. (Strobilops), 6
desmoulinsiana Jeffr. (Pupa), 103
diecki Gredl. (Pupilla), 85
diezi Cless. (Strobiius), 9
Dimorphoptychia Sandb., 11
diodontina Hde. (Helix), 43
diodontina Hde. (Strobilops), 43
ddoscoricola C. B. A. (Pupisoma),

114
diptyx Bttg. (Helix), 6
diptyx Bttg. (Strobilops), 6
diptyx Klika (Strobiius), 6
Diseostrobilops Pils., 18, 46, 56
doliolum Brug. (Oreula), 133
dolium Drp. (Oreula), 124
dollfusi Collet (Pitys), 6
dollfusi Collet (Strobilops), 6
duncana Pils. (Gastrocopta), 65
duvali Mich. (Helix), 6
duvali Mich. (Strobilops), 6

E
ectina Bgt. (Oreula), 9i2
edita Zim. (Oreula), 125
edentula Drap. (Columella), 109
elasmodonta Reuss. (Helix), 6
elasmodonta Reuss. (Strobilops), 6'
elatior Stki (Vertigo), 93

elongata Stoss. (Torquilla), 78
Enteroplax Gude, 18, 50, 56
Eostrobilops Pils., 18, 42, 56
erdeli Both (Helix), 180
erdeli Roth (Pleurodiscus), 180
eumicra Bgt. (Vertigo), 85, 103
eupleura Ch. & B. (Pupoides), 138
euryomphala Pila. (Gastrocopta),
120

F
faleifera Bttg. (Helix), 179
falciferus Bttg. (Pleurodiscus), 179
falconianus Pils. (Fauxulus), 75
farta Bof. (Chondrina), 123
Fauxulus Schauf., 73, 121
fischeri Wenz (Strobilops), 6
flaccida Bof. (Chondrina), 123
flavescens Parr. (Helix), 180
flavida Zgl. (Helix), 180
flavogilva Germ. (Truncatellina),

171
floridana Pils. (Strobilops), 26
frici Klika (Patula), 179
fryanus Bs. (Fauxulus), 121
fuehsi Zim. (Oreula), 89, 132

G

galapagorum Pils. (Pupisoma), 114
Gastrocopta Woll., 63, 117, 201
Gastrocoptinae Pils., 201
gedanensis Klebs (Strobilops), 7
gedanensis Klebs (Strobiius), 7
Geminidens Pils., 64
geminidens Pils. (Gastrocopta), 64
genesii Gredl. (Vertigo), 104
geyeri Lindh. (Vertigo), 104
gigantea Schr. (Abida), 77
gigantea Schr. (Pupa), 77
Glaphyra Albers., 174
godetiana Piag. (Vertigo), 102
goniodon Pils. (Pupilla), 82
grabaui Ping (Pupa), 137
grabaui Ping (Pupilla), 137
grabaui P. & Y. (Pyramidula), 188
gracilior Zim. (Oreula), 128
gracilis P. & Y. (Pyramidula), 188
graeeus Pils. (Oreula), 92
Granopupa Bttg., 77
granum Drap. (Granopupa), 78
guadalupensis Pils. (Pupilla), 83

INDEX, VOL. XXVni.

22S

guadalupensis Pils. (Vertigo), 97
guatemalensis Hink. (Strobilops),

34, 60
gularis Bm. (Orcula), 129
Gyliotrachela Tomlin, 73
Gyraulus Ag., 117

H
halleriana Jeffr, (Pupa), 135
hannai Pils. (Strobilops), 59
headonensis Edw. (Helix), 7
headonensis Edw. (Strobilops), 7
helleniea Bgt. (Orcula), 172
helleri Dall (Endodonta), 42, 62
helleri Dall (Strobilops), 41
helvetica Bgt. (Orcula), 172
helvetica West. (Pupa), 172
helvetica West. (Vertigo), 172
hierosolymitana Bgt. (Helix), 187
hierosolymitana Bgt. (Pyramidula),

187
Hippeutis, 188
hirasei Pils. (Strobilops), 44
hoggarensis Pall. (Leucoohiloides),

81
hoggarensis Pall. (Pupoides), 81
hopeiensis Ping (Pupa), 137
hopeiensis Ping (Pupilla), 137
hordaceus Gabb (Pupoides), 138
hordeacella Pils. (Gastrocopta), 117
hubbardi A. D. Br. (Helix), 48
hubbardi A. D. Br. (Strobilops), 47
hubrichti Pils. (Vertigo), 99
humilis Hutt. (Helix), 188
humilis Hutt. (Helix), 188

idahoensis Pils. (Vertigo), 100
imperfecta P. & F, (Gastrocopta),

117
inermis Eussell (PupiUa?), 137
infima Zim. (Orcula), 125
infrapontica Wenz (Gastrocopta),

e3

jaenensis Cless. (Helix), 184
jeanneli Germ. (Gastrocopta), 169
joossi Gottsch. (Strobilops), 7
joossi Gottsch. (Strobilus), 7

kanabensis White (Helix), 11
kanabensis White (Kanabohelix),ll
Kanabohelix Pils., 11
karachiensis Peile (Pupoides), 80
karisimbiensis Pils. (Lauria), 170
kaznakovi Rosen (Orcula), 91
kaznakovi Kosen (Pupa), 91

labassetieri Morg. (Helix), 7
labassetieri Morg. (Strobilops), 7
labyrinthica Say (Helix), 20
labyrinthica Say (Strobilops), 20
labyrinthieula Mich. (Helix), 7
labyrinthicula Mich. (Strobilops), 7
laevigata Kok. (Vertigo), 103
lamarckii Aud. (Pupa), 78
lata C. B. Ad. (Pupa), 135
Lauria Gray, 138, 199
leiodus Hutt. (Strobila), 12
laprevieri Pall. (Pupilla), 86
Leucochilus Bttg., 63
lilljeborgi West. (Vertigo), 103
Ijubetenensis Stur. (Orcula), 92
loessensis Bkr. (Vertigo), 93
Lucena Moq., 174
Lurama Honigm., 174
Lyropupa Pils., 205

M

mabilleana Bgt. (Orcula), 172
macrochilus Bof. (Chondrina), 123.
macrodonta Hesse (Agardhia), 88
mamillata Andr. (Pyramidula), 179
mamillatus Andr. (Pleurodiscus),,

179
marginatus Say (Pupoides), 80
marmouchana PaU. (Granopupa),

77
marmouchana Pall. (Pupa), 78
martiniana PUs. (Sterkia), 110
menardi Brong. (Helix), 7
meridionalis Iss. (Helix;, 184
mexicana Pils. (Strobilops), 32
micra P. & Y. (Pupa), 108
micra P. & Y. (Truncatellina), 10-8
micra Pils (Punctum), 190
micra Pils. (Pyramidula), 190

224

INDEX, VOL. XXVni.

micromphala Pils. (Strobilops), 30
Microstele, 137
milium Gld. (Vertigo), 101
minor Gall. (Orcula), 133
menardi Brong. (Strobilops), 7
Modicella Ads., 201, 203
monile Sandb. (Helix), 8
monilia Dh. (Helix), 8
monilia Dh. (Strobilops), 8
monsjovica Bof. (Abida), 122
monterosatoi Bof. (Solatopupa),

123
morsei Dall (Strobila), 40
morsei Dall (Strobilops), 39
moussoni Reinh, (Orcula), 93
moulinsiana Dup. (Vertigo), 103
munita Reib. (Gastroeopta), 66

N
neglecta Poey (Vertigo), 99
Nesopupa Pils., 204, 206
Nesopupinae, 204
Nesostrobilops Pils., 57, 62
ninagongonis Pils. (Columella), 172
niobe Fult. (Gastroeopta), 119
nipponica Pils. (Strobilops), 45
nitida Mouss. (Pupa), 93
nitidulus Pfr. (Pupoides), 200
novopommerana E«h. (Neaopupa),

112
nucleata Turt. (Helix), 116
nylanderi Mse. (Pyramidula), 197

O

Obbinula Stacbe, 10
octonaria Pils. (Gastroeopta), 204
octoradiata Sars (Strobila), 12
odontostoma West. (Pupa), 105
oerstedi Mch. (Pupa), 82
oerstedi Mch. (Pupilla), 82
orbicularis Kl. (Helix), 179
orbicularis Kl. (Pleurodiscus), 179
orcula Bs. (Pupisoma), 116
Orcula Held., 88, 123, 201
Orculella Steenb., 92
Orculinae, 201

orcutti "Pils." (Pupa), 173
oreina Zim. (Orcula), 125
orientalis Pfr. (Orcula), 92
orphana Hde. (ITelix), 189
orphana Hde. (Punctum), 189
orthostoma Pier. (Pupa), 76

pachygaster Jens. (Vertigo), 103
Pagodinella Hesse, 138
Pagodulina Cless., 138, 201
paradoxa Stki. (Vertigo), 95
parietalis Pils. (Strobilops), 24
Patulastra Pfr., 182
Patulastridae Steenb., 177
patuliformis Sacco (Helix), 8
patuliformis Sacco (Strobilops), 8
pazi Hid. (Gastroeopta), 64
pediculus Shutt. (Gastroeopta), 118
peipingensis P. & Y. (Pyramidula),

188
peruviana Pils. (Bothriopupa), 114
Petrarca Pils., 88
pilsbryana Sterki (Gastroeopta), 64
pingi Pils. (Microstele), 137
pinii Ad. (Helix), 184
piratica Pils. (Strobilops), 58
plagiostoma Pier. (Pupa), 76
plana Cless. (Strobilops), 9
Planogyra Mse., 176, 197
Planorbis, 188
planus Cless. (Strobilus), 9
neurodiscidae Wenz., 177
Pleurodiscus Wenz., 178, 198
polyptychia MUdflf. (Plectopylis),

52
polyptychia MUdff. (Strobilops), 52
Pronesopupa Ired., 205
przevalskii Lindh. (Pyramidula),

186
pseud ocylindrica P. & Y. (Pupilla),

136
pseudocylindrica P. & Y. (Trunca-

tellina), 137
pseudodolium Wagn, (Orcula), 90,

128
pseudogularis Wagn. (Orcula), 88,

128
pseudolabyrinthica Cossm., 7
pseudolabyrinthica Sandb. (Helix),

8
pseudolabryrinthica Sandb. (Stro-
bilops), 8
Pseudostrobilus Oppenh., 9
Ptyehospira Slavik, 117
Punctum Mse., 189
Pupilla Leach, 81, 199
Pupillinae, 192, 199

INDEX, VOL. XXVIII.

225

Pupisoma Stol., 114
Pupoides Pfr., 80, 137, 200
Pupoidopsis P. & C, 200
pupula I>h. (Pupilla), 86
Pyramidula Fitz., 18i2, 198
pyrenaearia Mich. (Abida), 77
pyrgula Bgt. (Orcula), 172
pygmaeorum P. & C. (Columella),
109

Q

quadrasi MUdff. (Plectopylis), 55
quadrasi Mlldff. (Strobilops), 55

E
rapula Bgt. (Orcula), 172
recurrecta Oppenh. (Helix), 12, 183
reibischi Dall (Pupilla), 67, 117
restituta West. (Orcula), 129
riethmiilleri Tauch (Helix), 10
riethmiilleri Tauch (Pseudostro-

bilus), 10
rodriguezensis Conn. (Nesopupa),

110
romani Wenz. (Strobilops), 8
rupestiis Drap. (Helix), 184
rupestris Drap. (Pyramidula), 183
rupicola Stab. (Helix), 184

S
ealvini Tristr. (Helix), 38
salvini Tristr. (Strobilops), 37
sandbergeri Lomm. (Strobilops), 8
eandbergeri Lomm. (Strobilus), 8
saxatiUs Hartm. (Helix), 184
saxicola M.-T. (Abida), 77
scotaea Bgt. (Helix), 184
sculptilis Pils. (Vertigo), 100
Scyphus Monts., 88
scytodea Bgt. (Orcula), 172
semielausa Thiele (Gastrocopta), 73
eenegalensis Malz. (Gastrocopta),72
Senilauria Pils., 88
sesqueplicata Bttg. ( Strobilops) , 9
eesqueplicata Bttg. (Strobilus), 9
senaaricus Pall.(Leucochiloides), 81
signata Mouss. (Pupilla), 85
sillarensis Pier, (Abida), 76
sillarensis Pier. (Pupa), 76
Spelaeodiscus Brus., 176
spiralis Pils. (Strobilops), 31
spoliata Rm. (Orcula), 132

sterkiana Pils. (Pupilla), 82
Sterkia Pils., 109
sterri Voith (Pupilla), 136
stevensoni Pils. (Strobilops), 49
stoneri Cham.b. & Jones (Pupilla),

64
strebeli Pfr. (Helix), 33
strebeli Pfr. (Strobilops), 33
strebeli Pils. (Strobilops), 30
strobeli Gredl. (Truncatellina), 108
Strobila Morse, 12
St-obilidae, 1
StrobUopsidae, 1, 193
Strobilops Pils., 4, 12
Strobilus Sandb., 12
subconica Ping (Pupa), 137
subconiformis "Bttg." (Pupilla),

85
Bubconoidea Jooss. (Strobilops), 8
subconoideus Jooss. (Strobilus), 8
subcornea Bof. (Chondrina), 122
subglobosa Bgt., West. (Helix), 184
sublabyrinthica Edw. (Helix), 12
suboviformis Bttg. (Pupilla), 85
substriata Jeffr. (Vertigo), 103
subteres Cless. (Pyramidula), 183
sudensis Pfr. (Helix), 181
sudensis Pfr. (Pleurodiscus), 181
sundleri Odh. (Truncatellina), 106
suprema Wenz. (Strobilops), 6

T

tatrica Wagn, (Orcula), 89
tauricola Lindh. (Truncatellina),

106
teilhardi P. & Y. (Vertigo), 101
teresae Ben. (Helix), 180
texasiana P. & F. (Strobilops), 24,

195
Themapupa Iredale, 80
thieuxi Loc. (Solatopupa), 123
thomasseti Pils. (Gastrocopta), 71
tiarula Sandb. (Strobilops), 9
tiarula Sandb. (Strobilus), 9
tingitana Pall. (Orcula), 91
tolminensis Wagn. (Orcula), 90, 13Q
tomlini Conn. (Orcula), 133
torrei Aguayo & Jme. (Vertigo), 98
transaequatorialis Rch. (Neso-
pupa), 113
triplicata Stud. (Pupilla), 85

226

INDEX, VOL. XXVin.

trochoides Kreg. (Helix), 184
trochoides Paul. (Helix), 184
trochospira Mlldifif. (PleetopyUs), 53
trochospira MUdflf. (Strobilops), 52
Truncatellina Lowe, 105, 208
turcica Let. (Orcula), 92
turcmenia Bttg. (Pupilla), 85

U
umbilicata Mont. (Helix), 184
undorfensis Cless. (Strobilus), 5
uniplieata A. Brn. (Helix), 9
uniplicata A. Brn. (Strobilops), 9

V

valida Bof. (Abida), 122
Vallonia Eisso, 174, 195
Valloniidae, 173
Valtonia Anc, 174
vendryesiana Gloyne (Helix), 49
vendryesiana Gloyne (Strobilops),
48 '

vengoensis Conn. (Nesopupa), 111
veracruzensis Pils. (Strobilops), 35
Vertigininae, 208
Vertiginidae Stimps., 93
Vertigo Mull., 93, 208
vidali Bof. (Abida), 122
Virgo Pils. (Strobilops), 2^

W
wenziana Pils. (Strobilops), 60
wheeleri Pils. (Vertigo), 96
wolfii Mill. (Gastrocopta), 117
wouramboulehiensis Conn. (Lauria),
86

zonata Bttg. (Lauria), 86
zonata Bttg. (Pupa), 139
zonifera Pils. (Lauria), 86, 139
Zoogenetes Mse., 175
zschokkei Biitt. (Vertigo), 103
Zurama Leach., 174

PUPILLIDAE

PLATE 24

(-

13 14-

PUPILLIDAE. ETC.

PLATE 25

VALLONIIDAE, STROBILOPSIDAE

PLATE 26

1-3, Planogyra asteriscus. 4, Strobilops texasiana.

5-8, Strobilops aenea.

PUPILLIDAE

PLATE 27

1, 2, PuPOiDES M. NiTiDULUS. 3. 4, Gastrocopta OCTOXARIA.

5-7, G. armifera. 8, Xesopupa wesleyaxa.

9, 10, Bothriopupa tenuidens.

PUPILLIDAE

PLATE 28

1, 2, Pronesopupa admodesta. 3, Pupisoma mediamericanum.

4-7. Pupisoma comicolense. 8, 9. Xesopupa cocosensis.

10, 11, Lyropupa perloxga. 12, 13, Nesopupa xewcombi.

VALLONIIDAE, ETC.

PLATE 29

1, VaLLONIA PtJLCHELLA.

3-5, Pleurodiscus balmei.

2, Pyramidula rupestris.
6, Acaxthixula aculeata.

PUPILLIDAE

PLATE 30

1, 8, PUPILLA MUSrCORUM.

3, 4, Choxdrixa avexacea.

2, 7, Orcula dolium.

5, 6, Sandahlia cylindrica.

PUPILLIDAE

PLATE 31

1, 3, Abida sec.'U.e. 2, Chondrina similis.

4, 5, Vertigo pusilla. 6, Nesopupa moreleti.

7, Zoor.EXETES HARPA. 8, COLUMELLA EDEXTULA.

9, Truxcatellina cylixdrica.

SECOND SERIES: PULMONATA.

MANUAL

CONCHOLOGY

Structural and Systematic

WITH ILLUSTRATIONS OF THE SPECIES

FOUNDED liY

GEORGE W. TRYON, Jr.

OONTINUKD BY

HENRY A. PILSBRY, Sc.D..

Speciai. Curator Department op Mollusca, Academy of Natural
Sciences of Philadelphia.

109 Piain.

Part

PHILADELPHIA :

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT

ACADEMY OF NATURAL SCIENCES

OF PHILADELPHIA.

NOV 1 1 1927

ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA

MANUAL

OF

CONCHOLOGY

STRUCTURAL and SYSTEMATIC

SECOND SERIES: PULMONATA.

BY

HENRY A. PILSBRY

Part

1 1 u im.

PHILADELPHIA :

PUBLISHED BY THE DEPARTMENT OF MOLLUSCA

ACADEMY OF NATURAL SCIENCES

OF PHILADELPHIA

APR - 2 193t

■

JV^

T

7

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ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA

MANUAL

OF

CONCHOLOGY

STRUCTURAL and SYSTEMATIC

SECOND SERIES: PULMONATA. --1^ "^"k^

BY

HENRY A. PILSBRY

P„. LI' P^n-

PHILADELPHIA:

PUBLISHED BY THE DEPARTMENT OF MOLLUSCA

ACADEMY OF NATURAL SCIENCES

OF PHILADELPHIA

JUN 14 1934

'7/

SECOND SERIES: PULMONATA.

MANUAL

CONCHOLOGY

Structural and Systematic

WITH ILLUSTRATIONS OF THE SPECIES

FOITNDED ItY

GEORGE W. TRYON, jR.

CONTINOKD BY

HENRY A. PILSBRY, Sc.D.,

Special Curator Department of Mollusca, Academy of Natural
Sciences of Philadelphia.

pofft 12 flail).

PHILADELPHIA :

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT

ACADEMY OF NATURAL SCIENCES

OF PHILADELPHIA.

NOV 7 1935

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