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


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SCliNCB 



SECOND SERIES: PULMONATA 



MANUAL 



OF 



CONCHOLOGY 

VOL. XXV 

PUPILLID^: 

(GASTROCOPTUSLE, VERTIGININ^E) 



BY 

HENRY A. PILSBRY, Sc.D. 

AND 

C. MONTAGUE COOKE, Ph.D. 

Collaborating on Hawaiian forms 



PHILADELPHIA : 
PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA 
I9I8-I920 




EAftftt 

scisNd 
LIBRARY 



PUBLICATION COMMITTEE i 

HENRY SKINNER, M.D., Sc.D. WITHER STONE, A.M., ScJX 
HENRY A. PILSBRY, Sc.D. WILLIAM J. Fox 

EDWARD J. NOLAN, M.D. 



EDITOR : HENRY A. PILSBRY. 
TREASURER : S. RAYMOND ROBERTS, 



CONTENTS 

PAGE 

Introduction 

Terminology of the teeth of Pupillidse vii 

Descriptions of genera and species 
Subfamily Gastrocoptinse. 

Genus CHONDRINA Reichenbach 1 

Subfamily Vertigininae 68 

Genus VERTIGO Miiller 69 

American species 74, 372 

Species of Japan and Eastern Asia 150 

Palaearctic species, Europe, Asia, Northern 

Africa 159, 378 

Fossil Vertigininae 214, 379 

Genus STAURODON Lowe 224 

Genus LYROPUPA Pilsbry (Hawaiian) 226 

Genus PTYCHALAEA Boettger 273 

Genus NESOPUPA Pilsbry 274 

Systematic classification 275 

Hawaiian species 278 

Cocos Island species 322 

Polynesian, Micronesian and Melanesian 

species 324 

East Indian species ^. . . . 336 

Species of India and Ceylon 345 

Species of the Mascarene Islands 349 

African species 356 

St. Helena species 363 

Genus CAMPOL^MUS Pilsbry 364 

Genus COSTIGO Boettger 366 

Appendix : additions and corrections 370 

Explanation of Plates 381 

Dates of publication, parts 97-100 392 

Index 393 

(iii) 

696292 



PREFACE 

The present volume is chiefly concerned with the subfamily 
Vertiginina, to be completed in the next and final volume on 
Pupillidae. 

In treating of these snails the author has had the advan- 
tage of assistance from Dr. V. Sterki with the American Ver- 
tigines. The portions relating to Hawaiian species are in 
large part due to the collaboration of Dr. C. Montague Cooke, 
though all of the species have been studied by both authors. 

The work is based on material contained in the following 
collections, chiefly in the first two : 

The Academy of Natural Sciences of Philadelphia. 

The Bishop Museum, Honolulu. 

The Museum of Comparative Zoology, Cambridge. 

The United States National Museum. 

The Carnegie Museum, Pittsburgh (Sterki collection). 

The Indian Museum, Calcutta. 

The Bryant Walker collection, Detroit. 

In addition to the naturalists mentioned in Vol. XXIV, 
page vi, the author desires to express his thanks for mate- 
rial to 

Dr. S. Stillman Berry of Eedlands, California; 

Mr. William H. Clapp, Cambridge, Massachusetts; 

Major M. Connolly, London. 

H. A. P. 

PHILADELPHIA, NOVEMBER, 1919. 



INTRODUCTION 



For convenient reference a diagram showing the termin- 
ology of the teeth is repeated here. 



Lamellae 



Sapracolumellar ~ 

COLUMBLLAR 




'""'jir Suprapalatal 

UPPER PALATAL 
-* Inter palatal 
^ LOWER PALATAL 



I Iniracolumellar -....... 

V *\ ft ^7^ i 

~~z& Infrapalatal 

**^, BASAL 
FIG. 1. TERMINOLOGY OF LAMELLAE AND FOLDS. 

The names of primary teeth are in small capitals. 

The bay formed by the upper arc of the outer lip, and 
usually limited by the upper palatal and angular teeth, is 
called the SINULUS. This term has also been used extensively 
in Clausilia, Ennea, etc., in the same sense. 

In Vertigo, the point of the outer lip, running forward or 
bent inward, has been termed the AURICLE by Dr. Sterki. 

The ridge encircling the whorl behind and parallel to the 
free margins of the peristome is known as the CREST (collar, 
calliis cervicalis, ~bourrelet exterieur, Nackenwulst) . 

(vii) 



Viii INTRODUCTION. 

The use of formulas for expressing succinctly such struc- 
tures as the teeth of bivalves or gastropods would be a great 
convenience if it were not that nearly every investigator in- 
vents a new system for himself. Without uniform currency, 
terminology loses its value as an intellectual tool. For this 
reason, and because such formulas appeal to some but not to 
all naturalists, little use has been made of them in this work. 

In the Introduction to Vol. XXIV, p. viii, the tooth-formula 
used by Westerlund was noticed, and a more elaborate one 
was proposed. As the latter has never been used practically, 
it need not be considered further, being at best rather clumsy. 

Westerlund 's formula gives merely the number of teeth on 
each margin of the aperture, parietal, columellar, and palatal 
including basal, thus: 2-1-3. This is easily understood and 
often convenient for succinct statements in keys or tables. 

More elaborate formulas for brief statement of the aper- 
tural denticulation of Vertigo (but applicable also to other 
Pupillida3) have been published by Dr. V. Sterki (Proc. U. S. 
Nat. Mus., xi, 1889, pi. 42, fig. 5) and by Mr. C. M. Steenberg 
(Danmarks Fauna, Landsnegle, 1911, p. 156, fig. 127). 
Though these formulas have been used but little outside of 
the publications mentioned, the lists of symbols are given 
below as a matter of record. 

Both the Sterki and the Steenberg formulas express the 
structures fully. The chief objection to them is that they are 
rather difficult for one not a specialist on Pupillidae to inter- 
pret. Steenberg 's symbols do not distinguish between pri- 
mary and secondary teeth, and are based upon the Danish 
language ; Sterki does not recognize the lamella angidaris as 
a primary tooth. In writing his formulae, Dr. Sterki uses dots 
in place of figures for the secondary teeth. 

It appears to me that an ''international" formula uniting 
the best features of both could be constructed by designating 
the six primary teeth by the initials of their Latin names, 
and the secondary denticles by dots; the parietal, columellar 
and baso-palatal parts of the formula to be separated by 
hyphens. In all formulae the enumeration proceeds anti- 
clockwise. 



INTRODUCTION. IX 

Terminology of "International" Sterki Stcenberg 

this work. formula. formula. formula. 

Parallel and Twin lamellae la. V 

Angular lamella A 1 V 2 

Parietal lamella P A V 3 

Inf raparietal lamella I . 2 V 4 

Supracolumellar lamella . A t 

Columellar lamella C B A 2 

Subcolumellar lamella . A 3 

Basal fold B C B 

Infrapalatal folds 3 G 6 - 8 

Lower palatal fold Pi D G 5 

Interpalatal fold 4 G 4 

Upper palatal fold Ps E G 3 

Suprapalatal fold . 5 G 2 

Suturalfold 6 G t 

In the second column the letters i and s are added to the 
initial P to indicate inferior and superior palatals. 

As an example, the teeth of an 8-toothed Vertigo antwertigo 
(angular, parietal, inf raparietal, columellar, basal, infra- 
palatal, lower and upper palatals) would be represented thus: 

Westerlund formula: 3 1 4 

Sterki formula: .A.BC.DE 

Steenberg formula : V 2, 3, 4 A B Gs, 5, e 

"International" formula: AP. C B. Pi Ps 



v/. 
SECOND SERIES: PULMONATA. 

MANUAL 

OF 

CONCHOLOGY 

STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED T5Y 



GEORGE W./TRYON, JR. 



HENRY A. PILSBRY, Sc.D., 

SPECIAL CURATOR DEPARTMENT OF MOLLUSCA, ACADEMY OF NATURAL 
SCIENCES OF PHILADELPHIA. 



Part 



PHILADELPHIA : 

PUBLISHED BY THE CONCHOLOG1CAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES 

OF PHILADELPHIA. 



MANUAL OF CONCHOLOGY 



Subfamily GASTROCOPTIN^ (continued). 

Genus CHONDEINA Reichenbach. 

Pupa, in part, DRAPARNAUD 1805, and of most subsequent 
authors ; not of Bolten, 1798. Chondrus CUVIER, 1817, in part. 
Cochlodonta FERUSSAC, 1821, in part. 

Torquilla STUDER, Kurzes Verzeich., etc., 1820, p. 89. VON 
MARTENS, Die Heliceen, 1860, p. 287, type Pupa avenacea 
Brug. Not Torquilla Brisson, 1760 (Aves). 

Chondrina REICHENBACH, Allgemeine Taschenbibliothek der 
Naturwissenschaften. 5ter Theil. Zoologie oder Natur- 
geschichte Thierreichs, i, 1828, p. 93, as substitute for Chon- 
drus Cuv. ; ii, 1836, p. 152, for H. avenacea. 

Modicella H. and A. ADAMS, Genera Rec. Moll., ii, 1855, p. 
169. BOETTGER, Jahrb. Nassau. Ver. Naturk., xlii, 1889, p. 
249. WESTERLUND, Fauna Europaea Moll. Extramar. Prodr., 
ii, 1878, p. 178 ; Synopsis, 1897, p. 94. Cf. also CAZIOT, Bull. 
Soc. Zool. France, vol. 33, 1908, p. 195. 

Alloglossa LINDSTROEM, Om Gotlands Nutida Mollusker, 
1868, p. 18; monotype A. a-venacea (Brug.). 

The shell is rimate, cylindric-conic or conic, brown (except 
the group of C. similis, which have the shell opaque-whitish), 
of 4% to 10 whorls. Aperture with teeth arranged as in 
Abida, but often small, reduced in number or wanting; palatal 
plicae not entering deeply and not enlarged to form an internal 
barrier. Peristome somewhat expanded. 

Radula having the central and lateral teeth narrow, uni- 
cuspid, the cusp long and curved in profile; marginals with 
several cusps. 

Type C. avenacea (Brug.). Distribution, southern Europe 
from Portugal to the Caucasus, north to Gotland; North 
Africa. 



2 CHONDRINA. 

The radular teeth of Chandrina differ a good deal from 
those of other Pupillid<e, yet the peculiarity is probably not 
indicative of great systematic separation. Similar simplifica- 
tion of the teeth is seen in Allognathus, which in other char- 
acters differs but little from the rest of the Pentataeniate Heli- 
ces ; also in some tree snails. 

C. avenacea is probably as near as any recent species to the 
ancestral forms of the genus, its aperture having the full 
tooth-formula of the Abida group of genera. The other sec- 
tions of the genus (Modicella and Solatopupa) appear to be 
regional specializations from ancestors belonging to the aven- 
acea group (Chondrina s. sir.). The specialization usually 
was in the direction of tooth-reduction; but in the case of 
Solatopupa (C. similis and its allies) there has been adapta- 
tion to direct, brilliant sunlight, the shell becoming conspic- 
uously calcareous. 

Under Abida (Vol. XXIV, p. 263) the author has men- 
tioned his diffidence in offering a compilation in place of the 
monograph by an expert which these genera stand in need of. 
It seems best not to leave the account of Pupillidse incom- 
plete, and it is not practicable at present to have the work 
done abroad. 

Radula. The dentition of this genus was first investigated 
by G. Lindstrom (Om Gotlands nutida mollusker, 1868, p. 18), 
who described and figured the teeth of Pu^pa avenacea. On 
account of the narrow, simple cusps he formed a new genus, 
Alloglossa, for this species. It is a pity that a prior name has 
to be used, as Lindstrom was the first to define the group 
scientifically. 

Schako (in Westerlund, Fauna Europsea, etc., 1878, p. 167) 
stated that he had found dentition like that described by 
Lindstrom for avenacea in the species 

avenacea Brug. goniostoma Rm. 

quinquedentata Born. ventilatoris Parr. 

megachettos Jan. mMfeldii Kiist. 

Finally, the Rev. Prof. H. M. Gwatkin (Proc. Malac. Soc. 
London, iii, 1897, p. 227) gave the results of his examination 



CHONDRINA. 6 

of Pupillid dentition with lists of the species described. The 
following are stated to have teeth of the megacheUos-avenacea 
type: 

algesirae Kob. megacheilos Jan. 

amicta Parr. quinquedentata Born. 

avenacea Brug. scalaris Ben. 

bergomensis Charp. secale Drap. 

bigorrensis Charp. similis Brug. 
calpica West. 

The inclusion of secale must be due to an error of some sort, 
as I have found that species to have broad lateral teeth, all of 
them bearing ectocones. 

Distribution. 

Chandrina, like Abida, is generally found upon rocks, walls 
and cliffs, preferably on limestone, and therefore is chiefly de- 
veloped in broken country or mountains, where such stations 
abound, as are most of the Clausilias. Favorable stations are 
limited and discontinuous, leading to the segregation of colo- 
nies of snails having the habits mentioned. 

In distribution, CKondrina is much like Abida, the Alps and 
the Pyrenees being evolution centers, with one plastic and 
adaptable species, C. avenacea, widely spread over central 
Europe. 

About the Alpic center there is the typical or avenacea 
group of Chondrina, and the section Solatopupa, the similis 
group. 

In the Pyrenean center and through Spain and Portugal 
there is the bigorriensis group of Chondrma, and the weakly 
differentiated section Modicella; both probably derivatives of 
avenaceous ancestors. Species and races are very numerous, 
though the structural differentiation has been less, perhaps, 
than in the Alpic center. The number of species diminishes 
rapidly westward from the Pyrenees, very few being found in 
Portugal and northern Morocco. 

It appears reasonable to suppose that an avenacea-like stock, 
widely spread in southern Europe in late Tertiary time. 



4 CHONDRINA. 

was subjected locally to a great variety of conditions conse- 
quent upon the elevation of the Alps and Pyrenees, with pro- 
gressive interruption of limestone terranes by folding, fault- 
ing and erosion. To these isolating factors, probably, the pro- 
lific speciation is chiefly due. 

C. tingitana in Morocco is very closely related to forms of 
the adjacent part of Spain. The group of C. goniostoma, pen- 
chinatiana, letourneuxi and lallemantiana, reported from Al- 
geria by Bourguignat and Letourneux, have an aspect so 
Pyrenean that the possibility of importation occurs to one. 
They appear out of place in Algeria. C. avenacea reported by 
Letourneux also seems to be a stranger in Algeria, and Caziot 
directly denies its occurrence there (Ann. Soc. Linn. Lyon, 
Annee 1908, p. 160, footnote 1). 

Ch&ndrina is not certainly known anterior to the Pleisto- 
cene. Several forms possibly referable to the genus are enu- 
merated below, but by being toothless they resemble a few 
aberrant terminal products of evolution in the recent fauna 
rather than the more normal toothed forms which must have 
been ancestral. There is little reason for referring any of 
them to Chondrina, yet no other generic reference is obvious. 
The absence of definite traces of this genus in known Tertiary 
deposits leaves the question of its origin highly problematic. 

CHONDRINA(??) TROCHULUS (Sandb.). Pupa (Modicella) 
trochulus SANDBERGER, Vorwelt, 1875, p. 601, pi. 29, f. 25. 
CLESSIN, Malak. Bl. n. F., vii, p. 86. 

Upper Miocene : Morsingen, Undorf . Similar to Granopiipa 
rupestris Ph. ; toothless ; but in view of the frequency of tooth 
degeneration in Pupillida this character is almost worthless 
as a test of relationship. 

PUPA MULTICOSTULATA Gutzwiller. Abhandl. Schweizer- 
ischen palaont. Ges., vol. 32, 1905 (1906), p. 28, pi. 4, f. 30. 
Eocene, Lausen near Basel. Aperture unknown, and position 
doubtful ; externally resembles Chondrina. 

CHONDRINA(?) TOURNOUERI (Ancey). Pupa lusitanica 
TOURNOUER, Journ. de Conchyl., vol. 27, 1879, p. 176, pi. 6, f. 
3 ; not of Eossmaessler. Pupa tournoueri ANCEY, Le Natural- 
iste, i, p. 414, n. 11. for lusitanica Tour. Pupa tournoueri 



CHONDRINA. O 

COTTER, Communicaqoes da Direc^ao dos Services Geologicos 
de Portugal, iv, 1900-1901, p. 144, plate, f. A, 24, 25, n. n. for 
lusitanica Tour. Lower Miocene ( ?) near Lisbon, Portugal. 

Nomenclature. 

The species now referred to Chondrina have been known 
almost universally as Pupa or Torquilla. Both of these generic 
names were originally used for other groups of animals, and 
hence must be dropped in this connection. Chondrina was not 
discriminated from Abida until 1868, and in fact up to the 
present time few authors make the distinction, so that the 
notes on nomenclature given under Abida (Vol. XXIV, p. 
267) are to be consulted in connection with the following: 

The genus Chondrus Cuvier, 1817, comprised species of 
three modern genera, arranged in two divisions, as follows : 

1. Bulimus zebra Oliv. 2. Pupa f rumen turn Drap. 
Pupa tridens Drap. Bulimus similis Brug. 
Pupa variabilis Drap. Pupa polyodon Drap. 

2. Bulimus avenaceus Brug. Pupa quadridens Drap. 
Pupa secale Drap. 

In 1821 Gray mentioned Pupa cinerea as an example of the 
genus, and Hartmann, in the same year, used Chondrus for 
species now referred to Abida, Chondrina, etc.; but no type 
was expressly named until Gray, in 1847, selected Bulimus 
zebra. Chondrus, however, was not valid, having been used 
previously by Lamouroux for another group. 

Pfeiffer says of Chondrus Hartmann, 1821, "genus restric- 
tum ex typo P. frumenti," but this is not the case. In his 
article, System der Erd- und Plussschnecken der Schweiz, mit 
vergleichen der Aufzahlung aller auch in den benachbarten 
Landern, Deutschland, Frankreich und Italien sich vorfind- 
enden Arten (in Neue Alpina, Bd. i, pp. 218-220) Hartmann 
used Chondrus in somewhat wider than Cuvierian limits, in- 
cluding species of the modern genera Chondrina, Abida, Chon- 
drula, Vertigo, Pupilla, Truncatellina and Lauria. 

Recognizing the preoccupation of Chondrus, Reichenbach, 
1828, proposed to substitute Chondrina, without mentioning 
any species. He considered it a subgenus or section of Helix 



6 CHONDRINA. 

In 1836 he defined Chondrina briefly, mentioning one species, 
H. avenacea, as an example of his subgenus. This becomes, 
necessarily, the type of Chondrma. 

Cochlodonta Ferussac, Tableau Syst, 1821, pp. 28, 58, cov- 
ered lengthened, toothed snails of many genera, including 
various Pupillidae. As no type appears to have been selected, 
Helix uva Fer. (Cerion uva L.) has been named as the geno- 
type (Man. Conch., XXIV, p. 268). 

Modicella was also a composite group, proposed for tooth- 
less Pupae, with the following species, the modern genera 
being added in parentheses : 

conoidea Nc. (Bothriopupa) pacifica Pfr. (Pupoides) 

fallax Say (Ena, Pupoides) pallida Ph. (Chondrina) 

farinesii Desm. (Chondritia) putilla Shuttl. (Pupoides) 

freyeri Schm. (Zospeum) rupestris Phil. (Granopupa) 
modica Gld. (Pupoides) 

No type has been selected, but Westerlund, Boettger and 
some other authors have used Modicella for the groups of 
avenacea, rupestris and their allies. Ct. Caziot has contended 
that the name should be dropped. Pupa farinesii is selected 
here as the type species. 

Chondrina is here limited to species known or believed to 
have similar radulae. It contains many of the species grouped 
in the genus Modicella by Westerlund (1897), but it differs 
by the exclusion of the Rupestrellae ( Westerlund 's groups 5 
to 8) and by the inclusion of the similis group, which Wester- 
lund leaves with the Abidse. 

Westerlund has classified the species in eight groups desig- 
nated by numbers. Commandant Caziot has offered a re- 
arrangement into five groups (Ann. Soc. Linn, de Lyon, vol. 
53, 1907, pp. 196-7), uniting Westerlund 's 3d and 4th groups, 
and making two groups out of Westerlund 's 5th to 8th, with 
also numerous transpositions of species. Most of these altera- 
tions appear to be improvements. In the following year Caziot 
and Margier outlined eleven groups to contain the Chon- 
drinas and Rupestrellas, mentioning a few species under each 
group. Connecting links are so numerous that any linear 



CHONDRIN.E OF THE ALPIC CENTER. 7 

arrangement has its flaws, and the number of subordinate 
groups into which the three main series or sections may be 
split is here, as everywhere, purely a matter of convenience. 
The presence of several series of independent origin, in which 
the teeth have been reduced or lost, adds to the complexity of 
specific classification. 

Sections of Chondrina. 

a. 1 Shell brown, of different shades. 

6. 1 Lamellae and plicae well developed, 6 or more. Sec- 
tion Chondrina, p. 7. 

c 1 . Alpic; series of C. avenacea, no. 1 to 8. 

c 2 . Pyrenean ; series of C. bigorriensis, no. 9 to 28. 

6. 2 Teeth small or wanting; not more than 2 plicae in 
the outer-basal lip. 

c 1 . Pyrenees and Spain. Section Modicella, p. 44. 
c 2 . Alpes Maritimes and Liguria, species no. 3, 4. 
a 2 . Shell opaque, whitish or very pale. Southern France and 
Italy. Section Solatopupa, p. 57. 

It should be noted that some species of the avenacea group 
and of Solatopupa have degenerate teeth, as in the section 
Modicella. See species no. 3, 4, 42, 43. 

Section CHONDRINA s. str. 

Series of C. avenacea. 

1. CHONDRINA MEGACHEILOS (Cr. et Jan). PL 1, figs. 1, 2, 3. 
The shell is more or less openly umbilicate, nearly smooth 
or somewhat wrinkled, chestnut-brown to chocolate, paler 
above, shining. Spire slowly tapering, the penult whorl nearly 
as wide as the last ; latter part of last whorl compressed into 
a narrow ridge below, flattened laterally, and often weakly 
impressed over the upper and lower palatal plicae. Aperture 
oval, armed with 9 teeth : angular lamella long, continuous, or 
almost joining the short, higher, spiral lamella; parietal 
lamella short. Columellar lamella is strong and rather long, 
not quite reaching the peristome; subcolumellar smaller, 
shorter. Five palatals, the sutural and suprapalatal plicae 



8 CHONDRIN.E OF THE ALPIC CENTER. 

very small, upper and lower palatals long, the former emerg- 
ing to the peristome, basal plica small, rather deep. Peri- 
stome is expanded and strongly thickened within, white. 

Length 7, diam. 3 mm.; 7% whorls (Como). 

Length 8, diam. 3.2 mm.; 8 whorls (Como). 

Length 10.3, diam. 4 mm.; 9 whorls (Milan). 

Italy: Lombardy and Piedmont; also in adjacent parts of 
the Tyrol and Switzerland, and rare west to Grasse (Alpes- 
Maritimes) ; valley of the Verdon, Basses- Alpes, abundant 
and of good size (Margier). 

Chondrus megacheilos DE CRISTOFORIS et JAN, Catalogus 
rerum nat. in Museo extantium Joseph! de Cristof ori et Georgii 
Jan, sect, ii, pars 1, fasc. 1, p. 5, Mantissa, etc., 1832, p. 3. 
Pupa megacheilos Jan, ROSSM., pt. 5, 1837, p. 13, pi. 23, f . 318 ; 
iii, pt. 17, p. 106, pi. 85, f. 938. KUESTER, Conchyl. Cab., p. 
46, pi. 6, f. 6-8. DBS MOULINS, Actes Soc. Linn. Bord., vii, 
1835, p. 162, pi. 2, f. Al, A2. MOQ.-TAND., in part, Moll. Fr., 
p. 354, pi. 25, f . 23-32 ; with var. rufula. PFR., Monogr., ii, 
346; iii, 547; iv, 673; vi, 315; viii, 384 (see for older refer- 
ences). WESTERLUND, Malak. Bl., xxii, 1874, p. 62 (new 
diagnosis, and notes on transitions to avenacea) . SCHROEDER, 
Nachrbl. d. m. Ges., vol. 45, 1913, p. 171 (varieties, etc.). 
E. MARGIER, Feuille Jeunes Naturalistes, 1904, no. 399, p. 65. 
CAZIOT, Ann. Soc. Linn. Lyon, 1908, p. 158 (distribution, 
etc.). Torquilla tricolor ANT. et J. B. VILLA, Dispositio Syst. 
Conch., etc., 1841, p. 57. P. albilabris ZIEGLER MS., according 
to VILLA, Disp. Syst., 1841, p. 24. Pupa megacheila CAZIOT, 
Etude Moll. terr. et fluv. Monaco, etc., 1910, p. 316, with var. 
minor, p. 317, pi. 9, f. 19, from north of Grasse, on the short- 
cut leading to Saint-Vallier. Pupa megacheylos DUPUY, Hist. 
Nat. Moll. France, p. 394. 

In its typical form this species differs conspicuously from 
avenacea by its strongly thickened white lip, the columellar 
border more straightened, and by the development of 9 teeth, 
the sutural fold never being found in avenacea; also by the 
strongly compressed base and the more open umbilicus; yet 
there are transitional forms in the Tyrol which appear to 
connect the two stocks. These have been left as "varieties" 
under avenacea in this work. 



CHONDRIN^E OF THE ALPIC CENTER. 

Doubtless all records from the Pyrenees, from, those of 
Boubee and Des Moulins on, pertain to other species. It is 
not found in southern France beyond the dep. Alpes-Mari- 
times and Alpes-Basses, and even there the colonies are ap- 
parently few and small. According to Adami, the large, 
typical forms are at elevations above 2000 meters, smaller 
forms being found at lower levels. 

Commandant Caziot regards the big&rriensis group of the 
Pyrenees as modifications of a megacheilos stock which radi- 
ated from the Alpic center, but I am disposed to look upon the 
bigorriensis group and megacheilos as parallel mountain modi- 
fications of an early avenacea-like stock, and not otherwise 
directly related. 

Jan's original description is as follows: Chandrus mega- 
cheilos. (7/i. with the shell dextral, conic-cylindric, the sum- 
mit a little obtuse (alt. 4%, width 1% lines). Aperture 7- 
plicate (alt. 1^, width 1 line) peristome wide, white. North- 
ern Italy. 

Torquilla tricolor Villa was said to differ from megacheilos 
by the greater size, thicker shell, more reflected peristome and 
more numerous apertural folds (8) ; length 11-13, diam. 5 mm. 
In humid, limestone mountains of Valsassina, province of 
Como. This name may be used for the large form of mega- 
cheilos if one is needed. 

Form minor West. Length 5%-6, diam. 2 mm. Different 
from avenacea by the compressed, keeled base of the last whorl, 
reflected lip, the mouth angular below, the straightened colu- 
mellar lip, etc. (Fauna, iii, 1887, p. 96). No locality was 
given by Westerlund, but Schroder has reported it from near 
Gargnano, Salo, etc., on Lake Garda (Nbl., 1913, p. 174). 
Caziot also figured a "var. minor" from north of Grasse, 
length about 5.7 mm. 

Var. toscolana Schroder. Shell similar to the type except 
that it is stronger ; whorls 8-9 ; throat set with many folds and 
tooth-like tubercles (up to 16 !) . Length ll-14i/ 2 , breadth 4-5 
mm. On cliffs in Toskolano valley, in the mountains west of 
Lake Garda (Modicella megacheilos Jan, var. toscolana Schro- 
der, Nachrbl. d. m. Ges., vol. 45, 1913, p. 174). 



10 CHONDRIN^E OF THE ALPIC CENTER. 

Var. caziotana n. n. A variety rather of megacheila than of 
avenacea; in the latter species the palatal folds are opposite 
the columellars, and do not reach the peristome, while in 
megacheilas the stronger palatal folds alternate with the colu- 
mellars, and one at least, or several, reach the peristome, and 
this is the case with our variety, which is therefore closely re- 
lated to the var. maritima. It forms a passage to Pupa mega- 
cheila, but has not an angular aperture. Alpes-Maritimes : 
Gorges of Cians, north of Beuil, at 1500-1600 meters eleva- 
tion, and on the limestones of the plateau de Caussols, in the 
ravine near the road from Grasse to Thorenc, at the place 
called le Planesteli, at about 1000 meters (Pupa megacheila 
var. Idbiosa Caziot, Etude moll. terr. et fluv. Monaco, 1910, p. 
317, pi. 8, fig. 9). It may perhaps be equivalent to Pupa ave- 
nacea var. maritima Loc., according to Caziot, t. c., p. 316, 
footnote. The names labiosa and maritima are preoccupied in 
Pupa. Caziot 's figures are too small to show any distinctive 
characters. His description is given above. 

Var. galloprovincialis Margier. Hauteville (Ain). A form 
said to resemble that of the Tyrol. I have not found a descrip- 
tion of it, and suspect that it is a MS. name. Mentioned by 
Caziot, Ann. Soc. Linn. Lyon for 1908, p. 159. 

2. CHONDRINA AVENACEA (Brug.). PL 1, figs. 4, 5, 6. 

The shell is perforate, rimate, cylindric-conic, thin, chest- 
nut-brown or carob-brown; weakly, rather finely, irregularly 
striate. Whorls somewhat convex, the last well rounded at 
base, slowly ascending forward. Aperture with 7 teeth. An- 
gular lamella low, rather long, either weakly joining the spiral 
lamella or shorter and discontinuous with the spiral ; the spiral 
lamella is often rudimentary or absent. Parietal lamella 
short. Columellar lamella short, horizontal, a smaller, shorter 
subcolumellar below it. Upper and lower palatal and basal 
plicas rather short, the upper reaching the peristome, the 
others a little immersed; frequently a minute suprapalatal 
plica is developed. Peristome thin, whitish, a little expanded ; 
parietal callous very thin, transparent. 

Length 6.8, diam. 2.8 mm. ; 7% whorls. 



CHONDRIN^E OF THE ALPIC CENTER. 11 

Length 5.8, diam. 2.2 mm. ; 7 whorls. 

Europe, from the French Pyrenees to Greece, north to Oe- 
land and Gotland, but not in the British Isles. Western Cau- 
casus. Type loc., near Paris. 

Bulimus avenaceus BRUG., Encycl. Meth., i, 1783, p. 355 (en- 
virons de Paris) . Pupa avenacea Brug., PFR., Monogr., ii, 
347; iii, 548; iv, 674; vi, 316; viii, 385 (see for older refer- 
ences). CAZIOT, Ann. Soc. Linn. Lyon for 1908, pp. 160-169 
(distribution, synonymy, etc.). WESTERLUND, Fauna, iii, 1887 ? 
p. 97, with forms eupora, paucidens, and p. 98, var. subhor- 
deum, lepta, transiens West., etc. LOENS, NachrbL, xxiv, 
1892, p. 172 (varieties about Lake Lugano). Pupa (Modi- 
cello] avenacea Brug., BOETTGER, Jahrb. d. m. Ges., viii, 1881, 
p. 226, and x, 1883, p. 179 (Caucasus and Transcaspian Terri- 
tory). Alloglossa avenacea Brug., LINDSTROM, Om Gotlands 
nutida Mollusker, 1868, p. 18, pi. 1, f. 11, 13 (jaw and radula). 
STEENBERG, Danmarks Fauna: Landsnegle, 1911, p. 178, f. 
147-149 (good). Torquilla avenacea Brug., CLESSIN, Mollus- 
kenfauna Oesterreich-Ungarns u. der Schweiz, 1887, p. 227, f. 
129. Modicella avenacea WEST., Synopsis, 1897, p. 97. 
STEENBERG, NachrbL d. Malak. Ges., vol. 45, 1913, p. 169 (oc- 
currence in Denmark doubtful) ; Danmarks Fauna, Land- 
snegle, 1911, p. 178, f. 147-149 (teeth, genitalia and shell). 
Pupa avena DRAPARNAUD, Hist. Moll. France, 1805, p. 64, pi. 
3, f . 47, 48. ROSSMAESSLER, Iconogr., i, p. 82, pi. 2, f . 36 ; pt. 
5, p. 13, f . 319. KUESTER, C. Cab., p. 48, pi. 6, f . 12-14. Pupa 
hassiaca PFR., Symbolae ad Hist. Hel., i, 1841, p. 45 ; Monogr., 
ii, p. 334 (Habichtswald, Cassel, Hessen). KUESTER, Con- 
chyl. Cab., p. 85, pi. 12, f. 10, 11. VON MARTENS, NachrbL d. 
m. Ges., x, 1878, p. 89 (type specimen is an avenacea, injured 
in the penult whorl). CAZIOT, Bull. Soc. Zool. France, xxxiv, 
1909, p. 221. Jaminea septemdentata Risso, Hist. Nat. Eur. 
Merid., iv, 1826, p. 91. Pupa lucana BRIGANTI, Su due nuove 
specie di testacei spettanti al genere Pupa, 1832, in Atti 1st. 
Incoraggiamente, Napoli, v, 1834, pp. 221-238 (not seen) ; 
placed as a synonym of Pupa avena by Scacchi, Catalogus 
Conchyliorum Reg. Neapolitaiii, 1836, p. 16. 

This common snail is more widely distributed than any other 



12 CHONDRIN^E OF THE ALPIC CENTER. 

species of this or related genera except Granopupa granum. 
No satisfactory record for Spain has come to my notice, but 
Caziot states that, according to Margier, it has been found in 
the neighborhood of Gerona, Catalonia. Eastward the locality 
Caucasus is a long way from any other record, but the species 
probably extends through the southern provinces of Russia. 
Sicily and Greece are the southern extremes of its well-known 
range, but there is a single Algerian record, limestone rocks at 
the foot of the great massif of Thabbourt Bouzgueur, Kabylia 
(Letourneux, La Kabylie, i, 1872, p. 227). 

The thin lip, dark color, rounded base and 7 or 8 teeth are 
the chief characters ; but there are forms more or less connect- 
ing with C. megacheilos. 

Westerlund has compiled a formidable list of varieties; to 
it considerable additions are here made. It is an undigested 
mass of forms of different grades which nobody has yet studied 
in a broad, comparative manner. Some of them probably 
pertain to C. megacheilos. 

Form eupora West. Shell typical; aperture with 2 or 3 
parietal and 2 columellar lamellae, 4 palatal plicae, the upper 
one punctiform, the fourth short (West., 1887). Locality not 
given. Caziot states that most specimens of middle France 
are of this form. 

Form cerealis Ziegler MS. Shell larger, length 8, diam. 
21/2-3 mm. (Westerlund). 

Form paucidens West. Shell typical, but with 2 palatal 
plicae (West., 1887). Locality not given. 

Var. subcereana West. Aperture with 2 parietal, 2 colu- 
mellar, 4 palatal plicae, the angular lamella very large, pala- 
tals subequal, remote ; length iy 2 -8, diam. 2% mm. Sweden : 
Gotland, Oeland; Denmark; also Tirol, Tauria (Alloglossa 
avenacea var. subcereana Westerlund, Fauna moll. terr. et 
fluv. Suecica, etc., 1873, p. 280 ; Modicella a. s., West., Synop- 
sis, 1897, p. 97). 

Var. ferruginea West. Shell perforate, narrowly ovate - 
conic, rufous-ferruginous ; very delicately, more distinctly stri- 
ate at the suture, with few, distinct, whitish delicate riblets; 
whorls convex, the last a little convex at base ; aperture with 



CHONDRIN^E OF THE ALPIC CENTER. 13 

3 parietal, 2 columellar, 4 palatal plicae, the outer parietal fold 
very small, very deeply placed below the angular; the upper 
palatal minute, deeply placed, the median plicae longest, 
higher within. Length 4-5, diam. l%-2 mm. Agen, France. 
(Pupa ferruginea West., Fauna Europaea Moll, extramar. 
Prodr., ii, 1878, p. 180; Fauna, iii, 1887, p. 98; Synopsis, p. 
97, as var. of avenacea) . According to Margier, it was based 
upon decolored examples of avenacea. 

Var. subhordeum West. (pi. 1, fig. below f. 8). Shell 
smaller, with the same number of whorls, nearly smooth; 2 
folds on parietal wall, columella and palate ; length 5, diam. 2 
mm. ("P. hordeum of many authors, P. avena v. minor Kiis- 
ter, Mon., pi. 6, f. 15-16," Westerlund, Fauna, iii, 1887, p. 98). 
Kuster's figure is copied. He gives the localities, Bavaria, 
Wiirtemberg, Switzerland, Carinthia and Carniola as far as 
Triest. 

Var. cocchi Ben. Benoit states that in Sicily Pupa avenacea 
is common in woods and in the high mountains of Madonie, of 
Fiumedinisi ; found also at Marineo and Busambra. The Sicil- 
ian specimens are usually a little smaller than those of the 
rest of Europe (Nuovo Catalogo, 1881, p. 93). This form is 
evidently that which Benoit tentatively named Papa [sic] 
cocchi in 1845, in this manner. 

PAPA AVENA? Drap. Shell subconic, umbilicate, chestnut; 
whorls 7, obliquely striate ; aperture subrotund ; 7-toothed, lip 
emarginate ; apex rather obtuse. 

The shell which is found in Sicily is different, I believe, 
from that which lives in France ; yet it is closely related, for 
the Sicilian form differs only by being less conic, by the num- 
ber of whorls, by the shape of the aperture and by the ar- 
rangement of the folds in the same. According to Baron 
Mandralisca it is found abundantly at Madonie, on limestone 
rocks. Size 2% lines. PAPA COCCHI? Mihi. (L. Benoit, Ri- 
cerche Malacologiche, 1845, p. 11, pi. 2, f. 7). 

De Gregorio has named the same form Pupa avenacea Brug. 
var. sicula, II Nat. Siciliano, xiv, 1895, p. 203; referring to 
Benoit 's figures, II. Sist., pi. 5, f. 33, and his Nuovo Cat., and 
stating that the examples differ from the typical form by the 
smaller size and by having the lip of the aperture simple. 



14 CHONDRESLE OF THE ALPIC CENTER. 

Var. minima Loc. Shell a third smaller than the type, of 
a little more thick-set form, somewhat ventricose, the striae 
more strongly marked. Not common ; le Mont-d 'Or lyonnais 
(Locard, Variations Malac., i, 1880, p. 253). 

Var. aureaccnsis Loc. Shell much lengthened, slim, little 
tapering ; 10 nearly flat whorls, the last a little larger ; suture 
very large, as though channelled ; aperture small, rather nar- 
rowly oval, retracted at base ; 2 separated superior folds, the 
second immersed; 2 deeply placed columellars ; 3 immersed 
palatals, the first barely reaching the peristome; peristome 
thin, sharp, straight; shell vinous-brown, with little gloss, 
ornamented with very strong and irregular ridges. Length 
9-10, diam. 2 mm. Saint-Didier-au-Mont-d 'Or (Rhone) ; Cau- 
terets (Hautes-Pyrenees). (Pupa aureacensis Locard, Ann. 
Soc. d'Agric. Lyon (7), iii, 1906, p. 186.) 

The two widely separated localities, in diverse faunas, raise 
a suspicion that it may have been based on abnormal shells. 

Var. aveniculum 'Hartm.,' Clessin. Shell small (smaller 
than that of hordeum), of circular form. Length 5-6 mm. 
Switzerland, environs of Chur, Malans, right bank of the 
Landquart, Zizers, in the Schlundtobel on the Greta v. Schu- 
ders (Hartm. in sched. Am. Stein Graubiinten, p. 61. Tor- 
quilla avenacea var. aveniculum Clessin, Moll.-fauna Oester- 
reich-Ungarn, 1887, p. 229). 

Var. duplicata Kiister (pi. 1, fig. below fig. 7) differs from 
avenacea by having four palatal folds, the upper one deeper 
and shorter, the other three not reaching to the peristome ; the 
upper fold at the insertion of the right margin of the peri- 
stome is weak, not penetrating deeply; nearly behind it the 
second fold of the parietal wall is doubled, the right part 
hardly half as large as the left. Length 2, diam. 1 line. Tou- 
lon, southern France (Pupa duplicata Kiister, Conchyl. Cab., 
Pupa, p. 109, pi. 14, f. 37-39). The separation of angular and 
spiral lamellae is by no means rare among French specimens of 
avenacea, but the present form is also distinguished by having 
a suprapalatal plica and by small size. 

Var. lepta West. Fusiform-turrited, slenderly elongated, 
acuminate, chestnut-brown or corneous-brown, nearly smooth ; 



CHONDRINyE OF THE ALPIC CENTER. 15 

whorls 8, convex. Aperture oblong, outer lip more strongly 
arcuate, thin ; 7 lamellae and folds : 2 parietal, 2 columellar, 3 
palatal, these of nearly equal length. Length 5, diam. 2 mm. 
Adelsberg. (Pupa avenacea var. lepta Westerlund, Fauna, iii, 
1887, p. 98.) 

Var. clienta West. (pi. 1, figs. 7, 8). Slenderly fusiform, 
regularly, densely striatulate, aperture 8-toothed: 4 palatals, 
the two median pliciform, the upper produced to the margin, 
the one above it punctiform ; peristome with the outer margin 
regularly arcuate (not angular above) ; length 7, diam. 2 mm. 
Choc, in the Tatra Mts., Galicia. (Pupa avenacea var. clienta 
Westerlund, Jahrb. d. m. Ges., x, 1883, p. 60.) 

Two specimens figured were received from Bielz. 

Var. melanostoma ' Paul. ' West. Very dark chestnut-colored 
suffused with blue, mostly shortly ovate with very ventricose, 
rarely with conically elongated, spire; irregularly striate; 
whorls 6%-7, convex; peristome very narrowly expanded, 
thin; lamellae and plicae typical. Length 5-6, diam. 
mm. Opcina and Nabrecina near Triest ; a var. is 4% 
mm. (Pupa avenacea var. melanostoma Paul, in sc., Wester- 
lund, Fauna, iii, 1887, p. 98). 

Var. apuana Issel. PI. 1, fig. 12. This variety has some 
analogy with P. hordeum, but differs by the shape of the aper- 
ture. The shell is more slender than that of the type, of a 
brown color, and has the right margin of the aperture more 
regularly arcuate. Punta del Sagro in the Apuan Alps (Pupa 
avenacea var. apuana Issel, Dei Molluschi race, nella Prov. di 
Pisa, in Mem. Soc. Ital. di Sci. Naturali, ii, no. 1, 1866, p. 21. 
Del Prete, Bull. Soc. Malac. Ital., v, p. 81, pi. 1, f. 10-12). 

According to Del Prete, it is found in abundance at Carrara 
above Torano, at Tambura, and rare at Matanna. One of his 
figures is copied, pi. 1, fig. 12. His natural-size figure is 6.3 
mm. long. The name appeared as Alloglossa avenacea var. 
apuna Issel, in Westerlund, Malak. Bl., vol. 22, p. 130. 

Form elatior Paulucci. Larger, both in length and width, 
but differing chiefly by being much more slender. Avellana in 
Umbria and at Caramanico (Pupa avenacea form elatior Paul., 
Bull. Soc. Malac. Ital., vii, 1881, p. 119). 



16 CHONDRIN^E OF THE ALPIC CENTER. 

Var. lessinica Adami. Elongate, brown, the aperture irreg- 
ularly subangulate, toothless; whorls 8-10; length 8-11, diam. 
2y 2 mm. Breonio, Prov. of Verona (Pupa avenacea var. lessi- 
nica Adami, Bull. Soc. Malac. Ital., xi, 1885, p. 222). 

" Found sporadically in many localities.' 7 From this re- 
mark it appears to be an occasional mutation rather than a 
race. 

Var. arcadica Reinh. Two palatal plicae, one parietal in the 
middle, the other vanishing, upper columellar fold strong, the 
lower deep, weak and vanishing. Otherwise strongly devel- 
oped, resembling normal avenacea in color, striation and size, 
6.5 to 7.5 mm. Mountain region of Kyllene, Arcadia, Greece. 
(Torquilla avenacea var. arcadica Reinhardt, Sitzungsber. Ges. 
naturforsch. Freunde Berlin, 1881, p. 137). 

Form tirolica West. One parietal, 2 columellar, 4 palatal 
folds. Length 5%, diam. 2y 2 mm. Vallarsa, Tyrol (Modicella 
avenacea var. arcadica form tirolica Westerlund, Synopsis, 
1897, p. 97). Why considered a form of arcadica is not ap- 
parent. 

Var. abimdans West. Shell subturrite-tapering, acuminate, 
rufescent, obsoletely, irregularly striate; whorls 8, convex, 
especially the upper ones ; suture impressed ; aperture oblong, 
rounded at base, the margins subparallel, outer one curved 
above, columellar margin long, straightened, vertical ; with 8 
teeth ; upper palatal internal, very short. Length 8, width 3% 
mm. (Pupa [Modicella] avenacea var. abundans Westerlund, 
Verh. k.-k. zool.-bot. Ges. Wien, 1902, p. 406. Pupa [Tor- 
quilla] avenacea Brug. var. abundans Westerlund, Nachrbl. d. 
m. Ges., xxvi, 1894, p. 172). Greece: Kylteni in the Morea 
(Th. Kruper). 

(Races having some characters of megacheilos, from the 
Tyrol, Venetia. and Alpes-Maritimes.) As I have seen only 
riviana, I can form no opinion upon the value or synonymy of 
these races. 

Var. avenoides West, (transiens West.). Large, strong- 
shelled, cylindric-fusiform, very indistinctly striate, glossy, 
reddish-brown with bluish suffusion, the spire produced; 
whorls 8-8%, very .convex ; aperture oblong, with nearly equal, 



CHONDBIN^E OF THE ALPIC CENTER. 17 

weakly arcuate margins; lamellae and plicae 9: 3 parietal, 2 
columellar, 4 palatal, none of them reaching the peristome, 
the upper one very small; margin whitish, somewhat ex- 
panded below and on the left. Length 7%-9%, diam. 2%-3% 
mm. Lake Garda, Tirol. (P. avenacea transiens ad mega- 
cheilos Strobel. P. megacheilos var. avenoides and var. bigor- 
riensis Westerlund, Fauna Europaea Moll, extramar. Prodr., 
ii, 1878, p. 168. P. avenacea var. transiens West., Fauna, iii, 
1887, p. 98. Var. transiens W., Caziot, Ann. Soc. Linn. Lyon, 
1908, p. 165.) Differs from megacheilos by having the last 
whorl rounded at the base, the base of the aperture regularly 
rounded, the lip not reflected and the more deeply placed 
palatal plicae (West.). This appears to be a well-marked sub- 
species, which should evidently bear the earliest name, C. ave- 
nacea avenoides West. See also Gredler, Nachrbl., 1878, pp. 
20, 21. 

Var. megacheUoides Clessin. Shell moderately umbilicate, 
ovate-conic, usually of lighter color; peristome somewhat 
lipped; aperture with 3 nearly equal palatal plicae standing 
directly opposite the columellar folds and the inner fold of the 
parietal wall, and one small upper palatal plica far within. 
Neck obtuse on the margins, usually without furrows. South- 
ern Tirol, Doss Trento and S. Bernardino near Trient, Segno 
in the Nonsberge, etc. (Pupa avena Drap. Gredler, Conch, aus 
den Nordosten Tirols, 1860, p. 104. Torquilla avenacea var. 
megacheUoides Clessin, Molluskenfauna Oesterreich-Ungarns 
u. der Schweiz, 1887, p. 229). Probably a synonym of var. 
avenoides West. 

Var. transiens Clessin. Shell as in the preceding variety, 
but more slender, regularly striate, darker colored ; peristome 
less lipped. South Tirol, Bozen on porphyritic rocks, on the 
Schloss Rungelstein, near Dolsach in the Drauthal, on primi- 
tive rocks (the first fold rudimentary). (Pupa avena Gred- 
ler, Tirol, p. 104. Torquilla avenacea var. transiens Cless., 
Moll.-fauna Oesterreich-Ungarns, 1887, p. 229.) This is not 
transiens West., which probably has priority. 

Var. gredleriana Clessin. Shell somewhat smaller; neck 
compressed, keel-like, the upper palatal fold wanting or rudi- 



18 CHONDRIN^ OF THE ALPIC CENTER. 

mentary and almost concrescent with the second fold. Aper- 
ture not rarely narrowed angularly at the base. South Tirol, 
Salurn on the Litschbach, on limestone (Pupa avena Gredler, 
Tirol, p. 104. TorquUla avenacea var. gredleriana Clessin r 
Moll.-fauna Oesterreich-Ungarn, p. 229). 

Var. riviana Schroder. PL 1, figs. 9, 10. Shell narrowly 
umbilicate, fusiform-conic, finely striate or smooth, corneous- 
brown to dark brown, often with a bluish wash; 7-8 weakly 
convex whorls; neck somewhat flattened, compressed into a 
weak keel beneath; aperture large, ovate, obliquely truncate 
above; throat with 8 folds: 2 each on the parietal wall (the 
one next to the outer lip sometimes divided) and on the colu- 
mella, 4 on the palate, the upper small, the next long and 
reaching the margin, the last somewhat shorter; peristome 
sharp, white, weakly lipped, somewhat reflected at the colu- 
mellar margin. Alt. 7-10 1 /2, breadth 3-4 mm. (Schroeder). 

Austrian Tyrol : cliffs on the banks of Lake Garda, at Tor- 
bole and Kiva, especially on the nummulitic limestone of Monte 
Brione, usually in company with Chondrula quadridens and 
TorquUla frumentum (Schroder). 

Modicella riviana SCHROEDER, Nachrbl. d. Malak. Ges., vol. 
45, 1913, p. 173. 

It differs from Modicella avenacea by its size, the wide 
mouth, the number and strength of the folds (avenacea 7, 
riviana 8) ; from M. megacheilos Jan by the more fusiform 
shape, the sharp peristome which lacks a flattened lip, and the 
number of teeth (megacheilos 9, riviana 8). The varieties in- 
troduced by Clessin, megacheiloides, transient and gredleriana, 
from other districts of the southern Tyrol, have nothing to do 
with it (Schroeder). 

Cf. also Westerlund's notes on the same form. Malak. Bl., 
xxii, p. 63, and Gredler, Nachrbl., 1885, p. 37, paragraph 3. 
Specimens collected at Riva by J. L. and A. L. Baily (pi. 1, 
figs. 9, 10) show considerable variation in shape and size. 
Whether it is to be ranked as a species, or as a subspecies of 
avenacea or megacheilos is a question to be decided when the 
intermediate forms abounding in the Tyrol shall be revised 
with adequate material. 



CHONDRIN.E OF THE ALPIC CENTER. 19 

Var. plicatula Adami. Shell subfusiform, brown, the aper- 
ture subangular below, brown within ; 5 subequal palatal plicae ; 
columellar margin plicatulate. Whorls 8-9. Length 8-9, 
diam. 2^-3 mm. Heights of the limestone mountains in the 
Illasy valley (Mt. Malera, Zevola and Campobruno), Province 
of Verona. Near to megachettos Jan. (Pupa avenacea var. 
plicatula Adami, Bull. Soc. Malac. Ital., xi, 1885, p. 221.) 

Var. maritima Loe. Subcylindric-elongate, slowly tapering, 
8-9 quite convex whorls, the last a little larger ; suture deep ; 
aperture quite large, oval-rounded, a little angular at the 
base; 2 superior folds, the second approaching and much im- 
mersed ; 2 subequal and deeply placed columellar folds ; 3 pala- 
tals, of which only one or two reach the peristome ; peristome 
sharp, expanded, white-rose or red ; shell vinous-brown, glossy, 
ornamented with striae, occasionally obsolete. Length 10, 
diam. 3 mm. Saint-Martin-de-Lentosque (Alpes-Maritimes). 
(Pupa maritima Locard, Ann. Soc. d'Agric. Lyon (7), iii, 
1906, p. 186. Cf. CAZIOT, Etude Moll. Monaco, 1910, p. 316.) 
Not Pupa maritima Pfr. 

This form may perhaps be referable to megacheUos. The 
name was preoccupied. 

3. CHONDRINA OLIGODONTA (Del Prete). PL 1, figs. 13, 14. 

Shell cylindric-conic, ashy-brown ; aperture having a single 
columellar lamella, the parietal and palatal folds rudimentary, 
often wanting ; right lip more arcuate. 

This variety, which in external appearance differs little 
from the type [avenacea] and even less from var. apuana 
Issel, is well distinguished by the characters of the aperture, 
which has a single superior columellar tooth (and even this 
smaller than in the type), whilst the other superior and 
palatal teeth can scarcely be seen, or are wholly wanting. The 
exterior wall is characterized by wanting the three little 
white, parallel lines of the type [of avenacea], which corres- 
pond to the internal palatal teeth (Del Prete). 

Italy : Apuan Alps, Liguria, only at considerable elevations. 

Pupa avenacea Brug., var. oligodonta Del Prete, Bull. Soc. 
Malac. Ital., v, 1879, p. 80, pi. 1, f. 13-15. 



20 CHONDRIN^E OF THE ALPIC CENTER. 

My knowledge of this form is solely from the original ac- 
count. Probably the shell which has been identified by Sette- 
passi as Pupa farinesii, from the same region, is a form of 
oligodonta more advanced in tooth-reduction. I do not think 
it is at all likely to be the Pyrenean species. Part of Sette- 
passi's note follows. Cf. C. a. lessinica, p. 16. 

In the course of my researches in the Apuan Alps (a part of 
the Apennines penetrating Tuscany from the Magra river, 
Spezia, in Liguria, to the Serchio river, Lucca, in Tuscany) I 
found Pupa farinesii almost everywhere. It is the real /an- 
nesii, without teeth, mingled with P. avenacea Brug., but 
always at high elevations. Its presence has already been an- 
nounced by Dr. Raymond del Prete, who has had it in his col- 
lection for several years. It is found together with another 
form which has one tooth, the variety described by Moquin- 
Tandon, 1855, p. 359, as var. denticus [sic], and with another 
having the same tooth and two columellar callosities above, 
var. oligodonta Del Prete (F. Settepassi, in Feuille Jeunes 
Naturalistes, no. 520, 1914, p. 75). 

4. CHONDRINA CIANENSIS (Caziot). PL 1, fig. 15. 

Shell dextral, subfusiform, a little swollen, tapering above ; 
8 convex whorls, slowly and regularly increasing, the last a 
little less convex, not ascending to the aperture, having a 
height of 3 mm. in a total height of 11. Suture oblique, well 
impressed throughout; summit obtuse. Umbilicus narrow, 
half encircled by a not very pronounced cervical gibbosity. 
Aperture ample, nearly vertical (the axis a little inclined to 
the right), oval, a little elongate, the upper border not an- 
gular, lower margin regularly oval, columellar margin straight, 
the margins converging and joined by a thin callous. In some 
specimens there is a very thin angular tooth, a reduced pari- 
etal, and a slightly more distinct columellar, but these teeth 
do not exist in all individuals. There are no traces of palatal 
teeth. Peristome thin, sharp, reflected chiefly in the lower 
part and the upper part of the columella. The shell is ochra- 
ceous, uniform, nearly of the same color as the rocks it lives 
upon ; the first whorls are darker ; ornamented with irregular, 
unequal, very oblique striae, more or less crowded, the upper 
whorls not striate. Alt. 9, diam. 2% mm. (Caziot). 



CHONDRIN.E OF THE ALPIC CENTER. 21 

Alpes-Maritimes : gorges of the Cians, at about 1200 meters 
elevation, nearer to Beuil than Rigaut, living on Permian 
rocks in very numerous colonies, on the right bank of the 
torrent. 

Pupa cianensis CAZIOT, Etude Moll. Monaco et Dep. Alpes- 
Maritimes, 1910, p. 318, pi. 9, f. 16. 

P. cianensis differs from pallida by the coloration, the slower 
increase of the whorls, the deeper suture, stronger, coarser 
and more irregular striation ; the more lengthened last whorl, 
which is less convex, and the longer aperture. Pupa cianensis 
does not belong to the group of pallida, but to that of mega 
cheilos and avenacea. In the Alpic center it takes the role of 
farinesii in the Hispanic (Caziot). 

It lives in a much colder zone than pallida, in gorges shaded 
from the rays of the sun except for short periods. Just what 
relation it holds to C. oligodonta of the Apuan Alps has not 
been touched upon by Ct. Caziot. 

5. CHONDRINA CEREANA ('Mtzhlf.' Kiister). PL 1, fig. 11. 

Shell subperforate, cylindric-subfusiform, lightly wrinkle- 
striate, glossy, purplish-brown ; spire turrited, rather acute ; 8 
slightly convex whorls, the last somewhat compressed at the 
base. Aperture oblong, brown, 8-plicate within : angular and 
parietal folds deeply placed, of moderate size ; 2 columellars, 
the lower dentiform ; 4 palatal plicae, the upper one minute or 
obsolete, the rest subequal, nearly reaching the margin ; peri- 
stome simple, narrowly expanded, the margins converging. 
Length 7, diam 2| mm.; aperture 2 mm. long, If wide (Pfr.). 

Southern Germany and France (Kuester) ; Carinthia, south- 
ern France, northern Spain (Westerlund). 

Pupa cereana Miihlfeldt in coll., KUESTER, Conchyl. Cab., 
1845, p. 47, pi. 6, f. 9-11. PFR., Monogr., ii, 348. WESTER- 
LUND, Fauna, iii, 1887, p. 99. Modicella cereana Ch., Kiist., 
WESTERL., Synopsis, p. 98. 

With the appearance of P. avenacea, it agrees more with P. 
megacheilos in structure of the plicae (Pfr.). The distribu- 
tion given by Westerlund does not appear plausible. 



22 CHONDRIN^E OF THE ALPIC CENTER. 

6. CHONDRINA CIRCUMPLICATA ('Mouss.' West.). PL 4, figs. 
12, 13. 

Shell shortly conic-turrited, dark purple to chocolate-brown, 
nearly smooth ; whorls 8-9, convex, the last divided above the 
middle by a spiral furrow. Aperture with 7 or 8 lamellae and 
plicae: 2 parietal lamellae, the angular lamella very long; 2 
columellar lamellae, the upper one prolonged to the margin; 
3 or 4 palatal plicae, the margin moreover closely set with 
small white folds. Length 6%, diam. 2y 2 mm. (West.). 

Italy : Val Membrano above Bergamo, Lombardy. 

Pupa circumplicata Mss., WESTERLUND, Malak. BL, xxii, 
1875, p. 131 ; Fauna Europaea, 1878, p. 168 ; Fauna, iii, 1887, 
p. 99. PFR., Monogr., viii, p. 385. Modicella c., WEST., 
Synopsis, 1897, p. 98. Pupa multidentata STROBEL, Essai 
d'une Distrib., 1857, not J. multidentata Risso, 1826. 

The lamellae and plicae are all stronger and longer than in 
avenacea. The spiral lamella is either continuous with the 
angular or separated from it. There is a deeply immersed 
suprapalatal plica. The columellar lamella sometimes emerges 
weakly to the lip, sometimes not, but there is a slender lamella 
in the angle of columella and parietal wall. The specimens 
figured are from the Marchesa Paulucci. 

7. CHONDRINA BERGOMENSIS ('Charp.' Kiister). PL 4, fig. 10. 

Shell short, deeply rimate, ovate-conic, lusterless, translu- 
cent, closely, obliquely rib-striate, pale corneous-yellow, the 
spire rather long, turrited, obtuse at apex ; 7-7% whorls are 
weakly convex, with simple, strongly impressed suture, the 
last somewhat ascending in front, slightly compressed below; 
the neck ribbed, impressed above the middle anteriorly. Aper- 
ture large, nearly ovate, 7-folded, 3 folds stand in the palate, 
2 on the parietal wall, and 2 on the columella. The palatal 
folds are long and show outside as white lines; peristome ex- 
panded, scarcely thickened, whitish, the margins strongly con- 
verging, joined by a thin, glossy callous. Length 2^-2%, 
diam. % lines (Kuester). 

Italy: around Bergamo, especially in the villages Valbona, 



CHONDRIN^E OF THE ALPIC CENTER. 23 

Rosciano and Ponteranica, on walls and limestone cliffs 
(Charpentier) ; Bergamasker Alps as far as Lake Garda 
(Kobelt). 

Pupa bergomemis Charpentier MS., KUESTER, Conchyl. 
Cab., p. 114, pi. 15, f. 14-16. PFR., Monogr., iii, 548. WES- 
TERLUND, Fauna, 1887, p. 99. KOBELT, Iconogr. n. F., ix, 
1902, p. 70, f. 1696. 

It is distinguished from all its relatives (megachettos, avena 
and secale) by its more slender shape, grayish horn-color, 
smaller size, and entire lack of gloss ; but chiefly also by the 
rib-striae, and by the very regularly and gradually increasing 
whorls (Charp. in Kuester). 

The angular lamella is usually separated from the spiral, 
but sometimes they are weakly connected. There is no supra- 
palatal plica. The upper palatal reaches the peristome, the 
lower scarcely, and the basal is decidedly shorter. The base is 
rounded, though the last whorl is distinctly compressed later- 
ally. Lip is thin and somewhat expanded throughout. Stria- 
tion is fine, close and regular. Length 6, diam. to edge of lip 
2.4 mm. 

Var. lepida West. Shell longer, the last whorl wider and 
higher, the outer margin of aperture more strongly angular 
above, columellar margin strongly converging at the insertion ; 
length 6%, diam. 2% mm. Lago di Garda. (Westerlund, 
Fauna Europaea, etc., 1878, p. 173.) 

8. CHONDRINA SPELTA (Beck). Vol. 24, pi. 48, figs. 6, 9. 

Shell more conic than the typical form [of avenacea] , um- 
bilical crevice larger, whorls more convex, the suture therefore 
deeper, the last whorl almost globular ; neck not compressed at 
base ; lower columellar fold usually rather degenerate ; always 
two folds in the palate; aperture somewhat more rounded; 
length 2, diam. 1 line, 7 whorls; in all other characters like 
the type. Dalmatia (Rossm., P. avena var. minor) . 

Shell with a narrow but penetrating umbilicus, ovate-conic, 
obtuse, from reddish-brown to brownish purple-red, frequently 
one of the whorls pale red, from the light-colored body show- 
ing through. The surface is closely rib-striate, almost luster- 



24 CHONDRIN.E OF THE ALPIC CENTER. 

less, the walls rather thick and scarcely translucent. The 
spire, which terminates in an obtuse apex, consists of 7 rather 
convex whorls separated by an impressed suture, and becom- 
ing rather high below. The last whorl is the broadest, nar- 
rowed below without a carina. Aperture almost round, red- 
dish-yellow, rather large, with 6 rather strong teeth, of which 
2 are in the palate, 2 on the parietal wall and 2 on the colu- 
mella. The peristome is only slightly thickened, somewhat re- 
flected towards the umbilicus only, the insertions approaching 
somewhat, the outer one joined with the upper fold of the 
parietal wall. Length 2%, diam. 1% lines (Kuester, P. miihl- 
feldtii) . 

Dalmatia: Almissa, Bagusa, and on the islands Lesina, 
Brazza and Bua, on rocks and under stones (Kiister). Servia 
(Mollendorff). Montenegro at Cettinje (Margier). 

Pupa avena var. minor Menke, (syn. : Torquilla hordeum), 
EOSSMAESSLER, Iconogr. 1, pt. v, vi, 1837, p. 13 ; Pupa hordeum 
on pi. 23, fig. 32Q.TorquiOa spelta BECK, Index Moll., 1837, 
p. 86 (based upon Rossmaessler's fig. 320). Pup [a] nana 
POTIEZ et MICHAUD, Galcrie des Moll., Mus. de Douai, i, 1838, 
p. 169, pi. 17, f. 3, 4 (La Dalmatie). Pupa miihlfeldtn KUES- 
TER, Conchyl. Cab., p. 28, pi. 4, f. 1-3, with var. major, p. 29, 
f. 4, 5, and var. minor (Pupa obscura Miihlf. coll.), p. 29, f. 6, 
7 ; Bericht nat. Ges. Bamberg, ix, 1870, p. 97. PPR., Monogr., 
ii, 332. WESTERLUND, Fauna, iii, 1887, p; 104. CAZIOT, Bull. 
Soc. Zool. France, vol. 35, 1910, p. 149. MARGIER, Feuille 
Jeunes Nat., vol. 40, 1910, p. 95. Pupa quinqueplicata Miihl- 
feldt MS., ANTON, Verzeichniss Conch., 1839, p. 47, no. 1713 
(nude name; not P. quinqueplicata P. & M., 1838). Allo- 
glossa muhlfeldti K., MOELLENDORPP, Malak. BL, xxi, 1873, p. 
134 (Servia). 

A markedly conic, chestnut-brown shell, of stouter figure 
than shells of the phUippii group, and differing from avenacea 
and its immediate allies by the more conic shape and by hav- 
ing only two palatal folds, by the shorter angular lamella, etc. 
It is a rather isolated species. 

Nomenclature. This species, usually known as Pupa muhl- 
feldti or miihlfeldi K., has not been correctly designated dur- 



CHONDRIN/E OP THE ALPIC CENTER. 25 

ing the last 80 years. Pupa avena var. maior and var. minor 
Menke, Synops. Meth., 1828, p. 18, were not defined or local- 
ized, and in using var. minor for the present form, Ross- 
maessler admitted that he was in the wrong, as there is no 
evidence that Menke ever saw the Eossmaesslerian form, and 
apparently intended merely to designate small examples of 
avenacea. T. hordeum Studer, mentioned in this connection, 
is an undetermined Swiss species, certainly not the Dalmatian 
form. After Rossmaessler, Beck was the first to name the 
shell, his Torquitta spelta being based upon Rossmaessler 's 
figures, one of which is copied in our pi. 48, fig. 6. This an- 
ticipated the subsequent names nana, miihlfeldtii, obscura and 
quinqueplicata. 

The typical form of spelta is small, length 2 lines according 
to Rossmaessler 's text, 5 mm. in the natural-size figure on his 
plate. There are but two palatal and no basal plicae, and the 
angular lamella is distinct but not long, entering but little be- 
yond the anterior end of the parietal lamella. This form is 
also represented by pi. 48, fig. 9, length 5, diam. 2.5 mm., 7% 
whorls. Another of the same lot is 5.4 mm. long. 

Kiister (1870) notes that this species "is found almost 
everywhere in middle Dalmatia, from the interior (Verlika) 
to close to the coast, on cliffs, common in places, as on the fine 
cliffs near Almissa ; also on the islands, where I obtained it on 
Lissa and Lesina. It appears to remain active also in summer. 

"Two forms of this species occur, sometimes in the same 
places, sometimes segregated, the one double the size of the 
other, 6-6y 2 mm. high, 3 mm. wide; the smaller one only 4 
mm. high, 2 broad. The larger form is usually darker even 
deep cherry-brown, with 6 to 8 folds in the aperture, there 
being a little plica above the two normal palatal plicae, and a 
fourth, rarely developed, in the base. The small form is 
brownish corneous-red, normal examples with 6 folds, 2 in the 
palate, 2 on the columella and 2 on the parietal wall are often 
present; but not rarely one disappears from each of the last 
two pairs, or only the lower columellar fold is lacking. 

"At Ragusa I found both forms, but not together; on Le- 
sina an intermediate form; the largest are at Spalato; fur- 



26 CHONDRIN^ OF THE ALPIC CENTER. 

ther inland there is only the small form at Makarska. On 
Lissa, rather large, yet not reaching the size of the Spalato 
shells." 

These notes give the proper perspective for viewing the 
named varieties; Kuster's large form being ventUatoris, his 
small one typical spelta and var. obscura. 

Var. venttiatoris ('Parr.' West). Vol. 24, pi. 48, fig. 11. 
Larger, more openly umbilicate, irregularly rib-striate, from 
dark chestnut-brown to deep cherry-brown ; sometimes having 
a third palatal fold at the base, and a fourth little one above 
near the suture. Length 6%-7, diam. hardly 3 mm. (West.). 

Dalmatia: Spalato. 

Pupa ventilatoris Parreyss, MARTENS, Die Heliceen, 1860, p. 
288 (name only). WESTERLUND, Malak. Bl., xxii, 1875, p. 131 
(description). PFR., Monogr., viii, p. 264. CAZIOT, Bull. Soc. 
Zool. France, vol. 35, 1910, p. 151. Pupa muhlfeldi [sic] var. 
ventilatoris Parr., WESTERLUND, Fauna, iii, 1887, p. 104. 
Pupa muhlfeldtii var. major KUESTER, Conchyl. Cab., p. 29, 
pi. 4, f. 5, teste West., Nachrbl., vii, 1875, p. 73. 

Kuester recognized this race, but used a preoccupied name. 
His description follows : 

Var. major Kiister. Vol. 24, pi. 48, fig. 11. Shell in general 
like the typical form, but the color is generally darker, the 
whorls flattened in the middle, aperture with 6 or 7 folds, the 
seventh standing basally on the palate, where the peristome 
curves into the columella. Length 3, diam. 1% lines. Under 
stones near Spalato, Dalmatia, above Salona towards the fort- 
ress Clissa, in company with ClausUia crenulata, not uncom- 
mon (K.). 

Var. obscura Mtihlf., Kiist. Vol. 24, pi. 48, fig. 4. Shell 
rather thin-walled, long-conic, the whorls flatly convex, with 
the suture but little impressed, aperture 6-plicate, the outer 
[angular] fold of the parietal wall sometimes almost imper- 
ceptible. Length scarcely 2 lines, diam. 1 line. Within the 
Spalatine district, on cliffs (K.). This is var. minor Kiister. 

The figures of this and the preceding variety are from 
Kiister. A specimen having the angular lamella extremely 
low, but with convex whorls, is drawn in vol. 24, pi. 48, fig. 



CHONDRIN^E OF THE PYRENEAN CENTER. 27 

10; length 4.7, diam. 2.5 mm., 6y 2 whorls. Another of the 
same lot is 5.3 mm. long, with 7 whorls. 

Series of C. bigorriemis. 

The forms of this eminently Pyrenean and Spanish series 
have often been referred to C. megacheilos of northern Italy, 
on account of their strong teeth and lip ; yet the relationship 
appears to be an indirect one, through a common ancestry in 
the avenacea stock. The series, while natural enough, is here 
segregated from the avenacea series rather as a matter of con- 
venience in treating the numerous forms than because of any 
important structural features. However, the lip is usually 
thickened within. 

The various varieties of P. megacheilos described by Des 
Moulins and Moquin-Tandon belong to the present group of 
species and races. P. megacheilos var. rufula Moq., Moll. Fr., 
p. 354, with the "shell reddish, more or less pale, Saint Sau- 
veur, ' ' has not been recognized by subsequent authors, though 
doubtless known under some other name. 

They may naturally be considered under these geographic 
groups : 

a. Species of the Pyrenees, Catalonia and adjacent prov- 
inces of Spain, no. 9 to 20. 

b. Species of central Spain, no. 21. 

c. Species of northern Spain, no. 22, 23. 

d. Species of Portugal, Cadiz and Morocco, no. 24 to 26. 

e. Species of Algeria, no. 27, 28. 

The group (a) of the Pyrenees and Catalonia is intricate, 
containing many forms which have not been figured or de- 
scribed in comparative terms, and which are scarcely known 
outside of the collections of their describers. A table giving 
some of the main characters is given as an aid to identification 
of species. The Algerian species also are known by the orig- 
inal account only. The few forms recorded up to this time 
from northern and western Spain, Portugal and Morocco are 
better understood. 



28 



CHONDRIN^E OF THE PYRENEAN CENTER. 






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CHONDRIN^E OF THE PYRENEAN CENTER. 29 

(a. Species of the Pyrenees, Catalonia and Aragon, to 
Valencia.) 

9. CHONDRINA BIGORRIENSIS ('Charp.' Des M.). PL 2, figs. 1, 

2, 3, 7. 

The shell is fusiform-conic, perforate and rimate, hessian- 
brown, fading to cinnamon-brown at the summit ; but slightly 
glossy ; finely and regularly rib-striate (but varying in degree 
and distinctness). Whorls moderately convex, the last a little 
flattened over the palatal region, which is marked with three 
white lines ; the base compressed into a rounded keel ; whitish- 
rufous towards the lip. Aperture oblong, cinnamon within, 
having 8 teeth: angular lamella long, continuous with the 
spiral; parietal long and low; columellar larger and longer 
than the subcolumellar ; 3 equal, rather long palatals, and a 
very minute or subobsolete and inconspicuous suprapalatal. 
Peristome white, but slightly expanded, a rather thick cinna- 
mon callous within the throat, but not thickened at the edge. 

Length 9.2, diam. incl. perist. 2.8 mm. ; Sy 2 whorls. 

Length 7, diam. incl. perist. 2.4 mm. ; iy 2 whorls. 

Central Pyrenees: type loc. Bagneres de Bigorre (Hautes- 
Pyrenees). 

[Pupa megacheilos] var. d, pusilla. (Pupa bigorriensis 
Charp.), DES MOULINS, Actes Soc. Linn, de Bordeaux, vii, 
1835, p. 163, pi. 2, f. D!, o2. ? P. farinesii var. bigoriensis 
Charp. subvar. ventricosa DUPUY, Bull. Soc. d'Hist. Nat. Tou- 
louse, xiii, 1879, p. 55 (La Preste; no description). Pupa 
bigorriensis GOURDON, Moll. Mts. de Luchon et de la Barousse, 
Bull. Soc. d'Hist. Nat. Toulouse, xv, 1881, p. 92 (dist. in 
Hautes-Pyrenees and Haute-Garonne). FAGOT, same Bull., 
xvi, 1882, p. 73 (Pic du Gar). Pupa megacheilos var. gracilis 
ROSSM., Iconogr., ii, pt. 11, 1842, p. 10, pi. 53, f. 729 (Mont- 
pellier and Barrege). 

Mr. Ancey has reported (as Pupa bigorriensis Charp. var. 
sinistrorsa) finding a single sinistral specimen among normal 
specimens from Cazaril, Bigorre (Le Naturaliste, i, 1881, p. 
403). 

Des Moulins' original description follows: 



30 CHONDRIN^] OF THE PYRENEAN CENTER. 

Var. d. (pusilla). It is this shell which received from M. 
de Charpentier the name of Pupa bigorriensis. It is smaller 
than the preceding variety [teMMmargwata] ; its peristome is 
less widened; and the aperture, of which the basal angle is 
almost effaced, is less lengthened. The effacement of this in- 
ferior angle loses all really characteristic importance by its 
inosculation with the type [that is, the typical megacheilos, in 
which the aperture is somewhat angular at the base] equally 
in the four varieties. It only forms an individual variation 
which dominates the type at Bagneres de Bigorre, whilst it is 
the type which dominates at la Preste (Pyr.-Oriental.). 
Length 7-8, diam. 2% mm. Central Pyrenees. 

Des Moulins' figure, copied in pi. 2, fig. 3, shows a fourth 
fold in the base. The specimens from Bigorre (received from 
Locard, and from other sources), do not show this fold. 

Perhaps the name pusilla should be used for this species, 
since it was that adopted by Des Moulins ; yet as pusilla and 
bigorriensis were published simultaneously ( as above ) , it may 
not be necessary to reject the selection of the second by nearly 
all subsequent authors. Rossmaessler states that he received 
pusilla from the Pyrenees from Parreyss in 1845 under the 
name P. consobrina (Iconogr., iii, 1859, p. 108). 

Pupa moquiniana is ranked as a variety of bigorriensis by 
Westerlund. There does not seem to be much difference. 

Var. moquiniana Kiister. PL 2, figs. 8, 9. The shell is elon- 
gate, rather obtuse, conic-cylindric, rimate, finely rib-striate, 
the striae rather oblique, having a silky luster, dark reddish- 
brown, the apex rust-yellowish. The 7-8 whorls are flatly con- 
vex, contracted towards the base, joined by a simple, slightly 
impressed suture, and increasing in height very gradually. 
Neck flat, compressed at the base, rounded, sometimes quite 
keel-like, whitish; also the rest of the neck is light flesh- 
colored, with 4 unequal whitish streaks from the folds show- 
ing through. Aperture wide, rust-reddish, vertical, higher 
than wide, with 9 unequal folds in the contracted throat : two 
on the parietal wall, the inner one far within, small ; 2 on the 
columella, the upper one much larger, sometimes emerging 
far; 5 in the palate, the upper one small, far within, like the 
rest in this place, elevated tooth-like; the 2d, 3d, and often 



CHONDRIN.E OF THE PYRENEAN CENTER. 31 

the 4th emerging to the peristome, the 5th small, obtuse, almost 
always in the obtuse angle formed by columella and lip, often 
even standing on the columella, frequently turned towards 
the right. Peristome sharp, broadly expanded, white-lipped; 
columella nearly straight ; terminations approaching. Length 
31/^-4, diam. li/4 lines (Kuester). 

France: Mt. Beudar near Pau (Basses-Pyrenees). 

Pupa moquiniana KUESTER, Conchyl. Cab., Pupa, p. 52, pi. 
7, f. 4, 5. PFR., Monogr., ii, 347. 

10. CHONDRINA TENUIMARGINATA (Des Moulins). PL 2, figs. 
4, 5. 

"Very brown; less solid and less lengthened than the pre- 
ceding [elongatissima] , smaller and more striate than var. a 
[Italian megacheilos] , with the same variations as to the in- 
ferior angle of the aperture, but having the margin of the 
peristome thinner, sharper, and reflected less flatly; 7-8 
whorls. Length 8-9, diam. 3 mm. Pyrenees-Orientales. It 
was this which M. Michaud had intended to publish under the 
name P. farinesi, and M. Farines under that of P. pyrenaica. 
It offers a subvariety of a very light corneous color, transpar- 
ent, fragile, very much striate, from Barcelona (Des Moulins). 

Pyrenees-Orientales: la Preste (Farines, Fagot); also re- 
ported from Luchon (Haute-Garonne) by Moquin-Tandon. 

Pupa frumentum BOUBEE, Bull. Hist. Nat. France, 3e. sect., 
moll, et zooph., 1833, pp. 10, 11 ; edit, of 1834, p. 30 ; not of 
Drap. Pupa megacheilos var. c (tenuimarginata) DES MOU- 
LINS, Actes Soc. Linn. Bord., vii, 1835, p. 163, pi. 2, f. cl-4. 
Pupa farinesi Mich. MS., and P. pyrenaica Farines MS., ace. 
to Des Moulins, in synonymy of tenuimarginata, 1835. Pupa 
~badia MOQUIN-TANDON, as synonym of tenuimarginata, Hist. 
Moll. France, ii, 1855, p. 354. Pupa leptocheilos FAGOT, Bull. 
Soc. d'Hist. Nat. Toulouse, 1879, p. 241. Pupa leptochila 
Fagot, LOCARD, Ann. Soc. Agric. Lyon (7), iii, 1896, p. 188. 

The arrangement of teeth is practically the same as in bigor- 
riensis, but it is a less cylindric, lighter colored shell, with the 
aperture more angular below, at least typically. There is 
sometimes a small sutural fold. The angular lamella is con- 



32 CHONDRIN^E OF THE PYRENEAN CENTER. 

tinuous with the spiral. Des Moulins' figure is copied in pi. 2, 
fig. 5. The name was altered by Fagot on the ground that 
* ' sesquipedalia verba excludenda sunt" as specific names a 
salutary but obsolete Linnean regulation. 

Var. elongatissima (Des Moulins). PL 2, fig. 6. Remark- 
able for its elongation, the fusiform appearance, the narrow- 
ness of the umbilicus (consequent upon the elongation), and 
by the smallness of the whorls of the point, characters which 
make it resemble a Clausilia. The shell, more transparent 
than that of Italy [megacheilos], is more solid than in the 
following varieties [tenuimarginata, bigorriensis] . The peri- 
stome thickened and reflected almost as flatly as in that of 
Italy, presents the same variations as to the inferior angle of 
the aperture. The size of aperture is equal to that of var. a 
[megacheilos], which results in a different proportion which 
is very characteristic of the variety ; 9-10 whorls. Length 12- 
13 mm., diam. 3^-4 mm. Pyrenees. The celebrated ento- 
mologist and lichenologist M. Leon Dufour gave four speci- 
mens to M. de Grateloup, biit the precise locality is unknown 
to me (Pupa megacheilos var. b (elongatissima), Des Moulins, 
Actes Soc. Linn. Bord., vii, 1835, p. 162, pi. 2, f. si, s2). 

Moquin-Tandon has recorded this variety from back of 
Gavarnie on the Spanish side of the boundary; perhaps he 
intended the following form. 

11. CHONDRINA GIGANTEA ('Moq.' Bossm.). PL 3, figs. 7, 8. 

Its color is light corneous-brown ; while very finely striate, 
it is much more decidedly so than P. megacheilos Jan. The 
umbilical orifice, narrow but very distinctly developed in 
megacheilos, is only weakly indicated ; the back of the neck is 
less strongly compressed, and the aperture therefore less an- 
gular below. It is especially remarkable that the two middle 
palatal folds become weaker inwards, as usual, while in mega- 
cheilos they always begin there with a tubercle. The lip of 
the less expanded peristome is weaker and not so broad and 
flat as in the other species (Rossm.). 

Cirque de Gavarnie, on the Spanish border, beyond the 
Pont de Neige (Partiot). 



CHONDRIN^ OF THE PYRENEAN CENTER. 33 

Pupa badia var. gigantea Moquin-Tandon MS., ROSSMAES- 
LER, Iconogr., iii, 1859, p. 106, pi. 85, f. 937 (as var. ? of mega- 

cheilos) . 

12. CHONDRINA GONIOSTOMA (Kiister). PL 2, figs. 13, 14. 

Shell somewhat cylindric-conic, rather thin, translucent, 
narrowly but deeply umbilieate, very finely rib-striate, glossy, 
especially on the penult whorl, reddish corneous-brown, the 
summit yellowish ; the spire is rather drawn out, the 7-8 whorls 
flatly convex, appearing separated by a deep suture, slowly 
increasing ; neck flat, yellowish behind the lip, with three whit- 
ish lines; base compressed, sharpened keel-like. Aperture 
rather broad, rounded-triangular, with a sharp angle below at 
the passage of the lip into the columella; at the base of the 
latter there is sometimes a tooth-like projection. Four plicae 
in the palate, the first small, the second very large, emerging 
to the peristome and forming a blunt tubercle there; colu- 
mellar lip ascending in a curve, much shorter than the peri- 
stome. Length 4, breadth 1% lines (Kuester). 

France: in the eastern Pyrenees (Kiister). 

Pupa goniostoma KUESTER, Conchyl. Cab., p. 53, pi. 7, f . 1-3. 
PFR., Monogr., ii, 345 ; iii, 547 ; iv, 673 ; vi, 314. BOURGUIG- 
NAT, Moll, de San- Julia de Loria, 1863, p. 22; Malac. de PA1- 
gerie, ii, 85, pi. 5, f. 35-38. 

A species more closely related to P. megacheilos than the 
preceding [P. moquiniana], but readily distinguished by the 
peculiar shape of the aperture as well as by the lack of a 
white lip. The finer striation, stronger gloss, smaller number 
of folds and more purely conic form separate goniostoma from 
moquinianci, in which there is also a tendency towards the 
formation of an apertural angle. From cereana the present 
species is distinguished by the greater size and especially by 
the union of the second palatal fold with the peristome ; from 
avena by the greater number of palatal folds (Kuester}. 

This appears to be a variety of tenuimarginata Des Moulins. 

Form aucta West. With 6 palatal folds, the first very short, 
sutural, 2d short, 3d to 5th marginal, 6th below, near the 
columella (Fauna Pal. Reg. Binnenconch., iii, 1887, p. 96). 



34 CHONDRIN/E OF THE PYRENEAN CENTER. 

Var. juliensis Bgt. PL 2, figs. 15, 16. Shell resembling the 
type, but having two folds (instead of one) at the insertion of 
the right margin. San Julia de Loria, Andorra. The typical 
form is quite rare, but the var. julieiisis is much more abun- 
dant (Pupa, goniostoma var. juliensis Bourguignat, Moll, de 
San-Julia de Loria, 1863, p. 24). 

13. CHONDRINA ANGULATA (Fagot). 

Shell perforate-rimate, conic-tapering, the spire fusiform, 
densely, subregularly striate, corneous-buff; whorls 12, the 
upper convex, lower flattened, the last with a strong white 
crest at base. Aperture oblique, oval-compressed, angular 
below; 2 parietal, 2 columellar and 5 or 6 palatal folds, the 
angular long, 3d palatal emerging, 4th and 5th punctif orm ; 
outer margin strongly thickened, the columella porcellanous. 
Length 11, diam. 2 mm. (Westerlund). 

Spain. 

Pupa angulata FAGOT, in Cronica Cientifica, 1888, not seen. 
Modicella angulata Fag., WESTERLUND, Synopsis, p. 95. 

Appears to differ from others of the group by the large 
number of whorls. 

14. CHONDRINA ARIGONIS (Rossm.). PL 2, figs. 10, 11, 12, 17, 
18. 

Shell perforate-rimate, ovate-turrited, the spire acutely 
tapering ; violaceous-brown, substriate ; whorls 7-8, convex, the 
last finally ascending, the neck a little swollen, the base 
rotundly compressed a little, buff, having two short white 
lines; suture deeply impressed. Aperture rounded semi- 
ovate, 6-folded : two folds on the parietal wall, one oblique, at 
the insertion of the lip, the other elevated, immersed ; two on 
the columella ; two rather deeply placed plicae of medium size 
in the somewhat calloused palate ; peristome somewhat spread- 
ing, with a very fully developed white or brownish lip. 
Length 6%-8, diam. 3% mm. (Rossm.). 

Spain: Pego, Prov. Valencia (Jose Arigo). 

Pupa arigonis ROSSMAESSLER, Iconogr., iii, p. 105, pi. 85, f. 
936. PPR., Monogr., vi, 316. Pupa arigoi Rssm., WESTER- 
LUND, Fauna, iii, p. 101. 



CHONDRINA OF THE PYRENEAN CENTER. 35 

By the form and the much developed lip it groups with the 
smaller forms of megacheUos, but differs by the base of the 
neck not compressed, and by having only 2 palatal plicae, as 
well as by the upper anterior fold [angular lamella] which 
never penetrates inward to the inner end of the parietal 
lamella, but always terminates at its forward end (Rossm.). 

The outer lip expands very little or not at all, but the whole 
peristome is widened by the conspicuous thickening of the 
inner edge. There is also a wide buff band behind the lip. 
Typically the teeth are strongly developed, but in one lot 
labeled Catalonia they are much reduced, low and weak, 
length 7.9, diam. 2.9 mm., 8 whorls. However, this lot is per- 
haps not really referable to arigonis. 

In another set from the type locality, the shell is more 
conic, and there are very minute, deeply immersed supra- 
palatal and basal plicae (figs. 17, 18, length 6, diam. 2.7 mm.). 
The surface, in all specimens seen, is nearly smooth or has 
very low, coarse wrinkles. 

Other described forms of the series follow; none of them 
have been figured, and their relations with the better known 
species remain to be indicated. 

15. CHONDRINA BAREGIENSIS ('Bgt.' Loc.). Cylindric, a 
little ventricose, 8 slightly convex whorls, the suture well 
marked; aperture subtriangular, a little contracted, angular 
at the base ; 2 small superior folds, the lower well immersed ; 
1 columellar at the superior angle; 4 narrow palatals, the 
upper obsolete, not reaching to the peristome ; peristome thin, 
very little reflected, sharp ; shell a little shining, dark red, 
ornamented with coarse striae. Length 9, diam. 3 mm. En- 
virons of Bareges (Hautes-Pyrenees). (Pupa baregiensis 
Bourguignat in coll., Locard, Ann. Soc. d'Agric. Lyon. (7), 
iii, 1896, p. 187.) 

It appears to be distinguished by the single columellar 
lamella. 

16. CHONDRINA CENTRALIS (Fagot). Quite small, conic, 
short and squat, rapidly tapering; 7 convex whorls with im- 
pressed suture. Aperture rounded basally, with parallel mar- 
gins ; 2 superior folds, one moderate, the other small ; 2 colu- 
mellars, the lower one more slender; 2 palatals, the lower 
obsolescent, not reaching the peristome. Peristome acute, 



36 CHONDRIN^E OF THE PYBENEAN CENTER. 

little reflected. Shell wine-brown, ornamented with quite 
spaced lamellar striae. Length 7, diam. 3 mm. Plateau de 
Lourdes, vallee du Gave d'Ossoii (Hautes-Pyrenees). (Pupa 
centralis, Fagot, Hist, malacol. des Pyrenees Francaises et 
Espagnoles, Bull. Soc. Ramond, 1891, xxvi, not seen; desc. 
from Locard, Ann. Soc. d'Agric. Lyon, (7), iii, 1896, p. 188.) 

17. CHONDRINA DERTOSENSIS (Bofill). Shell perforate-sub- 
rimate, ovate-subeonoid, obtusely tapering at the apex, cor- 
neous-reddish, a little glossy, obliquely striatulate ; 7-8 convex 
whorls, regularly increasing, separated by an impressed 
suture, smooth at the apex ; the last whorl a little swollen, a 
little compressed at the rimation, slightly ascending to the 
aperture. Aperture subvertical, semiovate-rounded, plicate as 
follows: An angular fold at the insertion, another rather 
prominent, immersed parietal in the middle ; 2 on the colu- 
mellar lip, the lower one smaller; 2 or 3 palatals (in adults), 
the upper one stronger, reaching the lip, the others shorter, a 
little immersed; peristome unexpanded, acute, with a tawny 
lip within, the margins strongly approaching, the outer curved 
above, columellar slightly reflected. Length 6, diam. 2, aper- 
ture 2 x 1 l / 3 mm. (Pupa dertosensis Bofill, Bull. Soc. Malac. 
France, iii, 1886, p. 162). 

Spain: rocks at the peak of the Serra de Cardo, Tortosa 
Catalonia) . 

This form has been placed by Westerlund in the avenacea 
group, by Caziot in the farinesii group near pulchella. 

18. CHONDRINA CRASSATA ('Bofill', Fagot). Shell nearly 
covered rimate, almost cylindrical, rather rapidly tapering at 
the summit, corneous- violaceous or rufescent, nearly smooth. 
Whorls 9, rapidly regular, the last compressed basally, broadly 
crested, descending at the aperture. Aperture produced to 
the left, ovate, with 2 parietal, 2 columellar and 4 palatal 
folds. Columellar lamellae nearly equal ; first palatal puncti- 
form, the rest subequal, not emerging; peristome expanded 
throughout, strongly thickened, porcellanous, the columellar 
margin oblique, straight, outer margin arcuate. Length 10, 
diam. 2y 2 mm. Spain. (Pupa crassata Bofill, in Fagot, Cro- 
nica Cientifica, 1888, not seen ; desc. from Westerlund, Synop- 
sis, 1897, p. 96.) 

19. CHONDRINA SCHISTA (West.). Shell deeply perforate, 
cylindric-conic, cherry-brown, rarely reddish-corneous, smooth 
(sometimes, under strong magnification, very irregularly stri- 



CHONDRIN.E OP THE PYRENEAN CENTER. 37 

ate), with oblique apex. Whorls 7-8, somewhat convex, the 
upper very rapidly increasing, the two before the last nearly 
equal, the last anteriorly obtusely angular above ; flattened in 
the middle, strongly blunt-keeled at the base. Aperture oval, 
with 2 parietal lamellae (the angular lamella long, interrupted 
deep within) ; 2 columellar lamellae, strong and horizontal; 4 
palatal plicae (the first very short, deep within, the rest very 
long, interrupted, the second nearly reaching the margin. 
Peristome narrowly expanded, brownish yellow ; margins sep- 
arated, the outer lip more arcuate above. Length 5-G 1 /^, diam. 
2-21/2 nim. Pyrenees. (Pupa schist a Westerlund, Fauna Pal. 
Reg. Binnenconch., iii, 1887, p. 112.) 

20. CHONDRINA HOSPITII (Fagot). Shell cylindric-conic, 
corneous-buff, a little rufous, not glossy, striate; whorls 8, 
convex, regular, the last a little larger, swollen in the middle, 
compressed below, much ascending. Aperture lunate-oval, 
large, with 2 parietal, 2 columellar and 3 or 4 palatal folds. 
Angular lamella thin, entering ; parietal deep ; columellar 
strong, immersed; 2 upper palatals subequal, the first some- 
what immersed ; sometimes with a punctiform fourth palatal 
fold. Peristome expanded throughout, white, thickened, the 
external margin much incurved, columella arcuate. Length 
7-8, diam. 2 mm. Spain : Venasque. (Pupa hospitii Fagot, 
Cronica Cientifica, 1888, not seen; desc. from Westerlund, 
Synopsis, 1897, p. 95.) 

PUPA ORTIGOSANA Fagot, of Ortigos (Logrona) is an unpub- 
lished species mentioned by Caziot, Ann. Soc. Linn, de Lyon, 
liii, 1907, p. 196, who placed it between hospitii Fag. and lusi- 
tanica Rm. 

(b. Species of Central Spam,} 
21. CHONDRINA GRATIOSA (West.). 

Shell rimate, oblong-turrited, cylindric, brown, delicately 
rib-striate; whorls 7, convex. Aperture oblong-ovate, with 1 
parietal, 2 columellar, 4 palatal folds (no angular; parietal 
deeply placed, high, compressed and short ; columellars small, 
immersed, white; 1st to 3d palatals remote, outwardly thick, 
inwardly tapering, long and slender, the 4th basal, smallest, 
punctiform, more deeply immersed). Peristome unexpanded, 
thin, the margins distant, very lightly curved, the outer 
shortly arcuate above. Length 4 mm. ( West.). 



38 CHONDRIN.E OF THE PYRENEAN CENTER. 

Central Spain, in drift of the Jarama river (S. Calderon). 
P.[upa] (Modicella) gratiosa WESTERLUND, Ann. Mus. Zool. 
Acad. Imp. Sci. St. Petersb., iii, 1898, p. 167. 
It seems to be near P. sardoa Cantr. (West.). 

(c. Species of Northern Spain.) 

Besides the following forms, C. gorbeana, no. 40, belongs to 
this district. 

22. CHONDRINA VASCONIC A (Kobelt). 

Shell fusiform-turrited, the spire slender, rather obtuse at 
the apex, rufescent-corneous, finely rib-striate; 8 convex 
whorls, separated by a deep suture, slowly increasing, the last 
a little longer than the penult, compressed at base, distinctly 
ascending in front, compressed behind the aperture, and 
lightly scarred over the lamellae. Aperture small, truncate- 
ovate, with the peristome thickened, margins joined by a very 
thin callous, contracted by 6 lamellae : two in the parietal wall, 
one larger, reaching the callous at the insertion of the outer 
margin, the other deep, entering, small ; a third at the begin- 
ning of the basal margin ; and three parallel folds in the outer 
margin, shining through to the outside, of which the lower is 
scarcely visible in the aperture. Length 6, diam. 2, alt. aper- 
ture scarcely 1.5 mm. (Kobelt). 

Spain: Orduna, Biscaya (Kob.). 

Pupa vascomca KOBELT, Jahrb. d. Malak. Ges., ix, 1882, 
p. 72. 

23. CHONDRINA KOBELTI ('Hid/ Westerlund). 

Shell oblong with rather conic spire, irregularly striate, 
chestnut-brown, whitish near the aperture. Whorls 7, very 
convex, the last rounded beneath, flattened anteriorly, slowly 
ascending above; suture impressed. Aperture triangular- 
oblong, with 6 or 7 teeth : 2 parietal lamellae, the angular long, 
1 columellar lamella, situated high, horizontal in front, not 
running forward, curving upward rapidly within; 4 palatal 
folds, the first very small, within, the other three nearly equal, 
separated, elevated within, very shortly entering. Peristome 



CHONDRIN^] OF THE PYRENEAN CENTER. 39 

simple, the margins separated; outer margin thin, arcuate 
above, then straightened; columellar margin nearly vertical, 
somewhat thickened, dilated; columella somewhat calloused. 
Length 5y 2 , diam. 2 mm. (West.). 

Spain : Oviedo, Asturias. 

Pupa kobelti Hid., WESTERLUND, Fauna, Pal. Reg. Binnen- 
conch., iii, 1887, p. 102. Modicella kobelti WEST., Synopsis, 
p. 95. 

23a. C. kobelti ordunensis n. subsp. PL 3, figs. 2, 3, 4. 

The shell is shortly rimate, cylindric-fusiform, rather solid, 
carob-brown, with an irregular cinnamon band behind the 
peristome and on the base; penult and last whorl flattened, 
tapering downwards, furrowed above the rounded, projecting 
base. Spire stout, convexly conic, not paler. Whorls con- 
vex. Sculpture of fine oblique striae. The aperture is much 
longer than wide, subtriangular, being narrowed below, 
rounded basally, tawny within, armed with 7 (or 8) white 
lamella? and plica? : angular lamella low, rather long, extend- 
ing a little beyond the outer end of the parietal, which is 
stronger ; columellar lamella horizontal, strong, not emerging, 
curving upward deep within; the subcolumellar lamella ob- 
tuse, tuberculiform, tinted, not visible in a direct front view. 
Three subequal plica? are evenly spaced, the upper palatal 
reaching to the peristome, the others shorter; they show as 
white or whitish streaks outside (and there is sometimes a 
minute, immersed, suprapalatal fold). The columellar lip is 
dilated, peristome otherwise unexpanded, with a thin, blunt 
edge ; but it is rather strongly thickened a little distance within 
the palate. 

Length 5.5, diam. 2.1 mm.; 6y 2 whorls (fig. 3). 

Length 5, diam. 2.1 mm. 

Spain: Pena de Ordufia; type no. 22791 A. N. S. P. 

This form differs from the unfigured C. kobelti West., from 
further west in northern Spain, by having a well-developed 
subcolumellar lamella, which, however, is immersed and 
scarcely visible in a front view. I have not seen C. kobelti. 



40 CHONDRIN^E OP THE PYRENEAN CENTER. 

(d. Species of Portugal, the Province of Cadiz, and Morocco.) 
24. CHONDRINA LUSITANICA ('Rossm.' Pfr.). PI. 3, figs. 5, 6. 

The shell is subperforate-rimate, cylindric-conic, very ob- 
liquely, irregularly ribbed, brown ; spire tapering, obtuse ; 
whorls 8, convex, the last ascending at the end, the neck 
sharply costulate, buff; base somewhat compressed obliquely; 
with 4 white lines. Aperture suboval, liver-colored, 8-plicate : 
2 rather long lamellae, near together, on the parietal wall, one 
extending to the outer margin where it connects with the in- 
sertion, the other more immersed; 2 moderate folds on the 
columella ; 4 in the palate, showing outside white through the 
tawny throat, the upper one remote and small, the second 
curved, reaching the margin, the third and fourth shorter and 
a little further in. Peristome spreading, somewhat thickened, 
the outer margin angularly curved above. Length 7, diam. 3 
mm. (Rossm.). 

Portugal: Serra d'Arzabida [Arrabida] (type loc.) ; many 
places in provinces Douro, Beira and Estremandura (Nobre). 

Pupa lusitanica Rossm. in litt., PFR., Monogr., ii, 1848, p. 
347. ROSSMAESSLER, Iconogr., iii, 1859, p. 105, pi. 85, f. 935. 
LOCARD, Conchyl. Portugaise, Arch. Mus. d'Hist. Nat. Lyon, 
vii, 1899, p. 147. Pupa avenacea var. luzitanica Rossm., 
NOBRE, Annaes Sci. da Acad. Polytech. do Porto, iii, 1908, p. 
50, pi. 1,13, 4. 

This form is distinguished from avenacea chiefly by the 
strong rib-striation and the internally thickened lip, produc- 
ing a light tawny area behind the lip. The teeth are strongly 
developed, the angular entering as far as the parietal, no divi- 
sion into angular and spiral lamellae being evident. The 
minute denticle in the base is sometimes nearly obsolete, and 
the suprapalatal is minute. 

Some authors have considered lusitanica a variety of ave- 
nacea, but it is surely quite distinct by the callous thickening 
in the throat and the strong sculpture. It belongs to a group 
of forms chiefly developed in the Pyrenees, and not to the 
immediate group of avenacea. Whether the true avenacea 
occurs in Portugal is highly doubtful; its western limit has 



CHONDRINA OP THE PYRENEAN CENTER. 41 

not been determined, and even reliable Spanish records are 
lacking. M. Morelet at one time cited lusitanica as P. secale, 
a species not found in Portugal. The best recent account of 
lusitanica is that of Nobre. 

25. CHONDRINA CALPICA (West,). PI. 3, figs. 10, 11. 

Shell perforate, ovate-turrited, horn-yellow, often densely 
suffused with bluish, ribbed; 7% whorls, the upper convex, 
the last depressed on the neck, somewhat compressed at the 
base, ascending above. Aperture oblong-semiovate, with 6 
strong white, immersed lamellaB and plicae : 1 parietal lamella, 
2 columellar, the lower one very small; palatal folds 3, the 
upper very long ; peristome thin, simple, acute ; outer margin 
strongly arched above. Length 7, diam. 3 mm. ( West.). 

Gibraltar: along the way from the city to the fortress (J. 
Lindahl). 

Pupa calpica WESTERLUND, Nachrbl. d. Mai. Ges., 1872, p. 
27 ; Malak. Bl., xxii, 1874, p. 58, pi. 2, f . 6-8 ; Fauna, iii, 1887, 
p. 103. PFR., Monogr., viii, 386. KOBELT, Iconogr. n. F., ix, 
1902, p. 70, f. 1694. 

The angular lamella is rudimentary or obsolete. In fresh 
shells (Westerlund's had been preserved in spirits) the color 
is. carob-brown or chocolate, fading more or less upward, but 
sometimes the whole tint is lighter, nearer walnut-brown. 
Sometimes there is a small suprapalatal plica, as in fig. 11. 
Length 6.5, diam. 2.8 mm. It is related to tingitcwia. 

26. CHONDRINA TINGITANA (Kobelt). PL 3, fig. 9. 

Shell rimate-perforate, fusiform-turrited, the spire slender; 
corneous; sculptured with oblique, rather widely spaced, ar- 
cuate ribs; 7 slowly increasing whorls, separated by a deep 
suture, the last nearly twice as long as the penult, the base 
compressed in an obtuse crest. Aperture ovate, somewhat ob- 
lique, with the margins of peristome converging, joined by a 
thin callous, subtuberculiferous near the insertion of the outer 
lip ; contracted within by 6 lamellae : one compressed parietal ; 
two, the upper larger, in the basal [columellar] margin; 3 
parallel in the outer margin. Alt. 7.5; a variety is smaller 
and more swollen (Kobelt). 



42 CHONDRIN^E OF THE PYRENEAN CENTER. 

Morocco : mountains of Beni Hosemar, opposite Tetuan, on 
limestone cliffs (Kobelt). 

Pupa tingitana KOBELT, Jahrb. d. malak. Ges., ix, 1882, 
p. 71. 

The specimen figured, labeled Tetuan, determined by Cles- 
sin, is not exactly typical of the species or the variety. The 
peristome is quite thin; angular lamella very low, irregular, 
composed of low tubercles weakly connected, and sometimes 
penetrating as far as the parietal lamella, but usually shorter 
or interrupted. Parietal lamella high. Upper and lower 
palatal plicas are rather long, a minute suprapalatal above 
them. Length 7, diam. 2.9 mm. 

Chondrina tingitana algesirae (Kob.). Differs from the 
preceding, which is close, by having the outer margin only 
armed with two lamellae, and also the tubercle of the parietal 
callous far more distinct. Found by me in multitudes on a 
limestone ridge near Algeciras ; seems constantly distinct from 
the preceding by the lack of the third lamella on the palate, 
yet is apparently only a local variety (Kobelt). 

(e. Species of Algeria.) 
27. CHONDRINA LETOURNEUXI (Bgt.). PL 3, figs. 14. 

Shell rimate, obese-oblong, ventricose, rather solid, corneous, 
sharply and strongly costulate. Spire short, tapering, the 
apex obtuse, smooth. Whorls 8, slightly convex, slowly in- 
creasing, parted by a moderately impressed suture, the last 
whorl tapering, compressed-carinate at base, deeply furrowed 
in the middle, and strongly ascending at the insertion of the 
outer lip. Aperture semioval, 7-folded ; a lamelliform fold at 
the outer lip, a stronger, more deeply placed parietal fold, 
two obliquely descending, emerging columellar folds, of which 
the lower is stronger, and three lamelliform, emerging palatal 
plicae, of which the upper is very strong, converges towards 
the parietal fold, which it almost touches. Peristome thick- 
ened, white, expanded, the outer margin arcuately angular 
anteriorly; margins joined by a thick callous cut into seven 
little teeth, one in the upper angle between the outer lip and 
the large lamelliform fold, and six others between the large 



CHONDRIN.E OP THE PYRENEAN CENTER. 43 



fold and the columellar margin. Length 5%, diam. 2 l / 2 mm - 
(Bgt.). 

Algeria : debris of the ravine of Chabet-Beinan, near Cape 
Caxine, 14 kilometers west of Algiers (Letourneux, Poupil- 
lier). 

Pupa letourneuxi BOURGUIGNAT, Malacologie de 1'Algerie, 
ii, 1864, p. 78, pi. 5, f. 19-22. 

The exact relations of this species and the next are in doubt, 
as we do not know whether there is an internal palatal plica 
above the upper emerging one, and the presence of a spiral 
lamella is uncertain. Their presence in Algeria is rather 
anomalous, as they appear to be of Pyrenean type. I have 
elsewhere (p. 4) referred to the Algerian records for C. 
goniostoma and other Pyrenean species. 

28. CHONDRINA LALLEMANTIANA (Bgt.). PL 3, fig. 13. 

Shell rimate, acuminate-elongate, ventricose at base, solid, 
somewhat glossy, subpellucid, under the lens regularly and 
obsoletely subcostulate, pale corneous or whitish-ashen. Spire 
acuminate, the apex smooth, rather acute. Whorls 9, a little 
convex, slowly increasing, separated by an impressed suture, 
the last whorl tapering, compressed at the base (as though 
gibbous) and carinate (the keel acute, strong, nearly disap- 
pearing at the peristome), at the periphery thicker, whitish, 
and strongly ascending at the outer lip. Aperture semiovate, 
slightly contracted below, having 7 folds: a lamelliform fold 
at the outer lip ; parietal fold minute, lamelliform ; 2 strong 
columellar folds; and 3 emerging, lamelliform palatal plicae, 
of which the upper is stronger. Peristome white-thickened, 
expanded, strongly reflected at base; outer margin arcuate 
anteriorly; the margins joined by a whitish callous orna- 
mented with 7 or 8 very long little plications. Length 7, diam. 
3mm. (Bgt). 

Algeria: debris of the ravine Chabet-Beinan, near Cape 
Caxine, 14 kilometers west of Algiers (Letourneux, Poupil- 
lier). 

Pupa lallemantiana BOURGUIGNAT, Malacologie de TAlgerie, 
ii, 1864, p. 80, pi. 5, f. 23-27. 



44 CHONDRINJE OP THE PYRENEAN CENTER. 

Differs from letoumeu&i, according to Bourguignat, by the 
conoid form, more glossy shell of a lighter tint, by the more 
separated, blunter, much less strong costulations, by the ab- 
sense of a median furrow on the last whorl, as in letourneuxi; 
by the outer margin not angular and not projecting forward 
by the less projectng columellar folds, not obliquely entering ; 
by the upper palatal plica not converging to the parietal fold, 
and by the plications of the parietal callous, which have a 
lamelliform, not a punctiform appearance as in letourneuxi. 

Section MODICELLA Ads. 

Series of C. farinesii. 

These forms of the Pyrenees and southeastern Spain differ 
from avenacea by the weakness or loss of teeth; of palatal 
folds there are not more than two, often one or none. Parietal 
and columellar margins each bear two, one or no lamellae. 
When present, the lamellae and folds are small and short. 

As is usual in series showing degeneration of the teeth, the 
individual lamellae and plicae become highly variable, and their 
value as specific characters is correspondingly diminished. 

There has been no rational revision of the many local forms, 
races or species. The work can be done only by a naturalist 
able to get a practical knowledge of them in the field and by 
study of the types or topotypes. 

As the data are not at hand for construction of a key, some 
of the more obvious characters have been tabulated below as 
an aid in determination. In the second column of the table, 
a indicates the presence of an angular lamella. 



CHONDRIN^E OF THE PYRENEAN CENTER. 



45 





Teeth 


00 

*z 



BE 


1 


a 

CS 

5 


Sculpture 


k 


a 


2 















h i 

2 
1 

2 

2 

2 
2 
3 
2 
2 
2 



farinesii. . . . -V 
farinesii var. . < 
speluncae ... 
tarraconensis . < 



a 
a 

a 
2 
2 


a 
a 
a 
a 
2 
2 



2 
2 
2 
2 
2 
2 
2 
2 
2 






1 



I 
1 
i 

i 


2 
2 
1 
1 
1 
1 
1 
1 
2 
2 
1 
2 
2 
2 
2 



7-7* 

9-10 

8 

7? 
6* 

io-ii 

8 
8 

9-10 
8 
8 

7 


5-6.6 

8 
6 

6 
4 
6.5 

8 
7-8 

7-8 
7 
0| 

7 


2-2.6 

3 

2 

2 

If 

1.5 
3| 
3-3J 

y 

2 
2| 

2f 


striate. 

fine crowded wrinkles, 
irregularly striatulate. 

" undulately striatulate. 
smooth, 
finely costulate. 
wrinkle-striate. 
very finely striolate. 
wrinkle-striate. 
hair-striate. 
distinctly, irregularly striate. 
striate. 

strongly striate. 
very finely, closely striate. 

irregularly hair striate. 
ribbed, 
inconspicuous striae, 
costulate-striate. 
striatulate. 
fine, uneven striae. 


microdon. . . . 
boettgeriana. . 
jumillensis . . . 
ignota 
guiraonis . . 
saltus . ... 


obliterata . . . 
massotiana . . . 
m. penchinatiana 
m. angustata . . 
m. confusa . . . 
m. sexplicata . . 
aragonica. . . . 
domicella. . . . 
pulchella .... 
p. manotiana . . 
ilendensis. . . . 
gorbeana .... 


10 
8 
8 

7* 
8 
7* 


6-9 
7 
4.5 
6 
6 
7 


2-2J 
2.5 
1.5 
2 

2 

2.8 



29. CHONDRINA FABINESI (Des Moulins). PI. 5, figs. 1, 2. 

Shell cylindric-conic, more or less lengthened, striate, brown ; 
whorls 6-7, a little convex, the suture deep. Aperture sub- 
oval, toothless; peristome whitish, simple, not reflected; um- 
bilicus patulous virguleeform. Length 6-6%, diam. 2 mm. A 
variety is shortened, more conic, length 5, diam. 2 mm. 
(DesM.). 

The shell is thin, chestnut-brown or carob-brown, finely and 
irregularly striate, the striation sometimes almost effaced, 
sometimes distinct. The whorls are strongly convex, the last 
somewhat compressed laterally, rounded basally. The peri- 
stome is thin, whitish, the columellar margin dilated, outer lip 
unexpanded, strongly arcuate above. Typically there is no 
trace of lamellae or plicae. 



46 CHONDRIN.E OF THE PYRENEAN CENTER. 

Length 6.6, diam. 2.6 mm.; 7% whorls (La Preste). 

Length 6, diam. 2.3 mm. ; 7 whorls. 

Length 5 mm.; 7 whorls (Perpignan). 

Pyrenees of southern France and eastern Spain, generally 
distributed; type loc. La Preste (Pyrenees-Orientales), on 
limestone back of the thermal baths (J. N. Farines) ; north- 
eastward to dep. Lozere. 

Pupa farinesii DES MOULINS, Actes Soc. Linn. Bordeaux, 
vii, 1835, p. 156, pi. 2, f. El-3. PFR., Monogr., ii, p. 308. 
ROSSM., Iconogr., ii, pt. 10, p. 25, f. 639. KUESTER, Conchyl. 
Cab., p. 51, pi. 6, f . 21, 22. MOQUIN-TANDON, Moll. Fr., ii, 
1855, p. 359, pi. 26, f. 5-10, with var. dentiens, I. c., f. 11. 
BOURGUIGNAT, Mollusques de San-Julia de Loria, 1863, p. 15, 
pi. 2, f. 1-3, with var. dentiens, p. 17, pi. 2, f. 5, var. obesa,, p. 
17, pi. 2, f. 4, and var. subcarinata, p. 17, pi. 2, f . 6 (in Rev. 
et Mag. de Zool., xv, 1863, p. 58). CAZIOT, Ann. Soc. Linn. 
Lyon, liii, 1907, p. 194 (distribution). MARGIER, Feuille 
Jeunes Naturalistes, no. 516, 1913, p. 199 (distribution). 
Probably not P. farinesii of SETTEPASSI, Feuille J. Nat., no. 
520, 1914, p. 75. 

A well-differentiated species, by the lack or very small size 
of teeth, recalling the Sicilian G. rupestris and the Algerian 
G. dupoteti, yet not closely related to either. 

The distribution of farinesii has been worked out by Mar- 
gier and Caziot. It is generally distributed in the dep. Pyre- 
nees-Orientales, and occurs also in the Spanish Provinces of 
Gerona and Barcelona. There is a record from Portugal, no 
doubt erroneous. In France it extends northward from the 
dep. Pyrenees-Orientales through Aude, Tarn, and Averyron 
to Lozere, that is, along the Cevennes uplift (using this term 
in a wide sense) . There is even a single record from Langogne 
(dep. Allier), but I regard this as dubious, since the Abbe 
Dumas, who seems to have collected assiduously, did not find 
it (Rev. Sci. du Borbonnais, 1901). It is very local in these 
eastern departments, special localities being on limestone 
rocks of the gorge of Malefosse, of the Tarn, as far up as Bla- 
joux. M. Locard found specimens in flood debris of the 
Rhone, doubtless washed in from a western affluent. There is 



CHONDRIN.E OF THE PYRENEAN CENTER. 47 

no satisfactory evidence that farinesii occurs anywhere east of 
the Rhone. It has been reported from the Apuan Alps of 
Tuscany, but while I have not seen specimens, I am disposed 
to think them terminal members of a parallel and independent 
evolution-series from the avenacea stock. See under oligo- 
donta. 

Of the several varieties described, the first three appear to 
occur with the typical form, and seem to be merely mutations. 

Var. obesa Bgt. Shell shorter, more obese than the type. 
San-Julia de Loria, Andorra. 

Var. subcarinata Bgt. Shell a little more lengthened than 
the type, with an obsolete carina at the base, which makes the 
aperture larger. San-Julia de Loria. 

Var. dentiens Moq.-Tand. PL 5, fig. 4. The angular lamella 
present as a whitish deposit, diffuse or in form of a little tooth, 
and a low prominence may usually be seen on the columella. 
La Preste. 

Var. biplicata Bgt. Shell of a little smaller size, having 
only one tooth on the columella instead of two as in the type 
[of jumillensis] . Eocks of San- Julia de Loria ; also valleys of 
Ussat and Vic-Dessos (Ariege). (Pupa jumillensis var. bipli- 
cata Bourguignat, Moll, de San-Julia de Loria, 1863, p. 18, pi. 
2, f. 10-12.) It differs from var. dentiens by having a low 
columellar lamella as well as an angular. See p. 50. 

Var. ascendens West. Shell widely perforate; whorls 7%, 
the ascent of the last high and long, rounded at base. Aper- 
ture with the base narrower, two-folded; outer margin angu- 
larly curved above. Length 6, diam. 3 mm. St. Beat, Pyre- 
nees. (Pupa jumillensis var. ascendens Westerlund, Fauna 
Europasa Moll. Extramar. Prodr., fasc. ii, 1878, p. 179.) 

PUPA SPELUNCAE ('Bgt.,' Loc.). Cylindric-elongate, feebly 
tapering; 9-10 slightly convex whorls, the last a little larger, 
suture moderate ; rimation quite strongly developed ; aperture 
subrectangular, a little excentric ; peristome thin, sharp, with- 
out external crest; shell quite solid, slightly glossy, deep 
brown, ornamented with fine, crowded wrinkles. Length 8, 
diam. 3 mm. Entrance of the grotto of Eaux-Chaudes (Basses- 
Pyrenees). (Pupa speluncce Bourguignat in coll., Locard, 
Ann. Soc. d'Agric. Lyon, (7), iii, 1896, p. 185.) 



48 CHONDRINA OF THE PYRENEAN CENTER. 

Probably a large form of farinesii. 

30. CHONDRINA TARRACONENSIS (Fagot). Shell rather openly 
perforate, regularly conic from the base, corneous-buff, irreg- 
ularly striatulate ; whorls 8, somewhat convex, slowly increas- 
ing, the last larger, ascending, rounded at the base. Aperture 
vertical, ovate-lunate, with 1 to 3 teeth: angular smaller, 
parietal more or less lengthened, or punctiform or wanting; 
columella thick, toothed; peristome unexpanded, margins 
nearly parallel. Length 6, diam. 2 mm. Spain. (Pupa tarra- 
conensis Fagot, Catal. Moll. Essera (not seen) ; Westerlund, 
Synopsis, 1897, p. 100.) Cfr. M. jumttl. v. ascendens W. 
( Westerlund) . 

31. CHONDRINA LONGINI (Fagot). Shell rimate-perf orate, 
the perforation nearly covered by the columellar margin, 
cylindric-conic, opaque, glossy, corneous-violaceous, at the 
summit yellowish; irregularly and obliquely undulately stri- 
atulate; spire tapering from base to summit, especially so in 
the upper part, the apex corneous, smooth, mamillate, not 
prominent, as though truncate; whorls 7%, a little convex, 
separated by a well-impressed suture, regularly increasing, 
the last larger, not compressed below but rounded, ascending 
a little to the aperture. Aperture slightly oblique, ovate- 
rounded, 3-plicate as follows: 1 tooth-like, porcellanous an- 
gular ; 1 immersed, lamellif orm, compressed, tapering parietal, 
also porcellanous ; 1 straight columellar, more or less incurved 
at the summit ; peristome thickened at the aperture, the mar- 
gins little approaching, the columellar margin lightly in- 
curved, reflected at the umbilicus, outer margin simple, regu- 
larly curved. Alt. 6, diam. 2 mm. ; alt. aperture l 1 ^, width 1 
mm. (Pupa longini FAGOT, in Navas, Boletin Soc. Aragonesa 
de Cienc. Nat., v, 1906, p. 201). 

Spain : pena Foradata above the Salient, Aragon, at an ele- 
vation of over 2000 meters (R. P. Longin Navas). 

It is perhaps approached only by our Pupa saltus of the 
Essera valley, and ignota of the valley of Panticosa ; but it is 
distinguished from the first by the less numerous whorls (8 
instead of 10), the last whorl rounded at the base and not 
compressed, the aperture more oval-rounded, by its columellar 
fold more immersed and curved at the top, etc.; from the 
second it differs by the greater size, more swollen whorls, the 
feebler teeth, etc. (Fagot). 

This is apparently the Pupa navasi Fagot, ined., mentioned 
by Caziot, Ann. Soc. Linn, de Lyon, liii, for 1906 (1907), p. 
197, from the same place. 



CHONDRIN^E OF THE PYRENEAN CENTER. 49 

32. CHONDRINA MICRODON (West.). Shell rimate-perf orate, 
ovate-conic, with the spire 2~y 2 times longer than the aperture, 
brown, smooth, the apex obtuse. 6% strongly, beautifully 
convex whorls, regularly increasing, the last rounded below, 
ascending in front; suture deep, narrow. Aperture strongly 
receding below, obliquely oblong, with one conic tooth deep 
within and high up on the columella; peristome simple, nearly 
unexpanded, only the columellar margin being dilated above ; 
margins very little and almost equally arcuate, except that the 
outer lip is strongly curved and protracted towards the colu- 
mella above. Length 4, diam. 1% mm. (Pupa microdon WES- 
TERLUND, Fauna, Pal. Reg. Binnenconch., iii, 1887, p. 104. 
Modicella m., WEST., Synopsis, 1897, p. 100). 

Spain : Montserrat, prov. Barcelona. 

33. CHONDRINA BOETTGERIANA (Clessin). PL 5, fig. 3. 

Shell rimate, subfusiform, thin, brown, pellucid, finely cos- 
tulate, the ribs narrow, regular; 10-11 convex whorls, slowly 
and regularly increasing, separated by a moderately impressed 
suture, the last somewhat compressed at base, about equalling 
one-fifth the length. Aperture nearly quadrangular, toothless. 
Peristome acute, a little reflected, the margins connected by a 
scarcely perceptible parietal callous, the outer margin lightly 
impressed. Length 6.5, diam. 1.5 mm. (Clessin). 

Spain: Jaen. 

Pupa boettgeriana CLESSIN, Malak. Blatter (n. F.), v, 1882, 
p. 188, pi. 4, f. 4. 

Described from one specimen. From farther southwest 
than other toothless species. 

34. CHONDRINA JUMILLENSIS ('Guirao,' Pfr.). PI. 5, figs. 6, 7. 
Shell perforate, ovate-turrited, rather solid, obliquely 

wrinkle-striate, slightly glossy, purplish-brown. Spire con- 
vex, slowly tapering, the apex rather acute; suture deep, 
simple. Whorls 8, convex, the last scarcely forming one-third 
of the length, slightly compressed around the punctiform per- 
foration. Columella longitudinally folded within and trans- 
versely two-toothed. Aperture slightly oblique, truncate- 
oblong ; parietal wall having a little lamella at the right angle ; 
peristome simple, slightly expanded, the right margin strongly 
curved above, toothless, columellar margin widely dilated. 
Length 8, diam. 3% mm., aperture 3 mm. long (Pfr.). 



50 CHONDRIN^E OP THE PYRENEAN CENTER. 

Southeastern Spain: Jumilla and Orihuela (Bourgeau). 

Pupa jumillensis Guirao, PFR., Monogr., iii, 1853, p. 540 
(Pupa bourgeaui Shuttlew. Mss. cited as a synonym). BOUR- 
GUIGNAT, Moll, de San-Julia de Loria, 1863, p. 17, pi. 2, f. 7-9. 

The original description of jumillensis is translated above. 
The figures are from Bourguignat, who did not give the source 
of the specimens, remarking that it is " found in a great num- 
ber of Spanish localities. ' ' 

Nomenclature. Rossmaessler, in 1859, described as the true 
Pupa jumillensis of Guirao a form much smaller than Pfeif- 
fer's and having but one columellar lamella instead of two. 
He was at a loss as to the source of Pfeiffer's specimens, which 
were supposed to have been furnished by himself. The matter 
is discussed at length in Iconographie, iii, p. 110. Bourguig- 
nat, in his Mollusques de San-Julia de Loria, cited above, re- 
described jumillensis Pfr., and synonymized jumillensis Em. 
with his own P. jumiUensis var. biplicata, from the Pyrenees 
a locality far from Rossmaessler 's. Westerlund ignores Pfeif- 
fer's jumillensis of 1853, accepting Rossmaessler 's species of 
1859, a proceeding clearly inadmissible. The untangling of 
this snarl must be left to European students who must elu- 
cidate the zoological relations of the forms as well as their 
nomenclature. 

Various forms which have been described as varieties of 
jumillensis appear referable rather to farinesii. A variety 
with three folds, 1 angular, 1 immersed parietal and 1 colu- 
mellar, is noted by Bofill in the Province of Barcelona (Bull. 
Soc. Malac. Fr., iii, 1886, p. 154). Possibly some form of fari- 
nesii is intended. Cf. P. tarraconensis Fagot. 

Pupa ignota appears to have been proposed as a substitute 
for P. jumillensis Pfr. not Rossm., and if so it becomes an 
exact synonym of jumillensis as here understood. Since I 
have not seen the original publication, Locard's description of 
ignota is given here. 

Pupa ignota Fagot. Cylindric-turriculate, a little elongate, 
spire high ; 8 convex whorls, the suture well marked ; aperture 
oblong-rounded, triplicate: 1 superior fold, lodged at the 
suture; 2 columellar folds; peristome a little everted; shell a 



CHONDRIN^E OF THE PYRENEAN CENTER. 51 

little glossy, violaceous-brown, very finely striolate. Length 
7-8, diam. 3-3% mm. Haute-Garonne, Hautes et Basses- 
Pyrenees, etc. (Pupa ignota Fagot, Cat. Moll. Essera, in Cro- 
nica Cientifica, 1888, p. 23; Locard, Ann. Soc. d'Agric. Lyon, 

(7), 1896, p. 185). 

34a. Chondrina jumillensis guiraonis n. n. Vol. XXIV, pi. 
47, fig. 9. 

The color is between russet and chestnut-brown, somewhat 
glossy, wrinkle-striate. The aperture is shorter and more ob- 
lique than that of farinesn. Angular lamella small and slen- 
der ; columellar lamella low, obtuse, so deeply immersed that it 
does not show in a direct face view. Length 4.7, diam. 2.2 mm. 

Rossmaessler's description of jumillensis follows: "Shell 
most minutely perforate, ovate-turrited, brown, obliquely stri- 
ate, the apex somewhat obtuse; whorls 6, convex, the last 
slightly compressed at base, a little ascending; aperture 
rounded semi-ovate, having 2 folds: one very slender fold at 
the insertion of the outer lip, one deep within on the straight- 
ened columella ; peristome a little expanded, thin, simple, the 
outer margin strongly curved, columellar margin rather 
straightened. Alt. 5, diam. 2% mm." (Rossm.). 

Rocks of Monte San Miguel near Orihuela, prov. Alcante, 
and of Monte de Santa Ana near Jumilla, prov. Murcia, Spain 
(A. Guirao). 

Pupa jumillensis Guirao, ROSSM., Iconogr., iii, p. 110, pi. 85, 
f. 943. Cf. Pupa jumillensis var. biplicata BOURG., Moll, de 
San-Julia de Loria, 1863, p. 18, pi. 2, f. 10-12. 

This shell is shorter and rougher than C. farinesii dentiens, 
with a shorter aperture. The type specimen figured is from 
Jumilla, no. 22790 A. N. S. P. 

Bourguignat's var. biplicata was said by him to equal jumil- 
lensis Rm., which I doubt. It is from Andorra in the Pyre- 
nees, and is probably a form of farinesn. 

35. CHONDRINA SALTUS (Fagot). Shell nearly covered 
rimate-perf orate, corneous-buff, reddish, hair-striate ; spire 
regularly tapering from the base, subulate ; whorls 9-10, the 
last ascending; aperture oval, with single folds on parietal 
wall, columella and palate; angular tooth-like callous, colu- 



52 CHONDRIN^E OP THE PYEENEAN CENTER. 

mellar immersed, palatal small, immersed ; outer margin thick- 
ened, arcuate, columella straightened, reflected above. Length 
7-8, diam. 2 mm. Spain. (Pupa saltus Fagot, Catal. Moll. 
Essera, Cronica Cientifica, 1888, not seen; descript. from 
Westerlund, Synopsis, 1897, p. 101.) 

36. CHONDRINA OBLITERATA (Charp.). PI. 5, figs. 9, 10. 

The shell is long, cylindric-conic, thin, slightly glossy, 
purple-brown, distinctly but irregularly striate, composed of 
nearly 8 whorls, rather rapidly increasing in height, weakly 
convex, joined by a whitish, thread-like, somewhat impressed 
suture; the apex bluntly rounded, the last whorl contracted 
below, with flat neck. The aperture is semiovate; on the 
straight columella an indistinct fold stands; two of them in 
the palate, the upper one indistinct, punctiform, the lower 
somewhat longer ; a fourth indistinct one on the parietal wall 
near the insertion of the peristome. The columellar reflection 
is wide and almost half covers the large, deep umbilical crev- 
ice. Length 3% lines, width 1 line (Kuester). 

Portugal (Holl). 

Pupa obliterata CHARPENTIER, in Kuester, Conchyl. Cab., p. 
112, pi. 15, f . 7-9. PPR., Monogr., iii, 544. 

This species is known only by the description of Charpen- 
tier, and the figure and description of his type by Kiister. It 
has not been found by recent collectors in Portugal, and that 
locality must be considered dubious. It has some resemblance 
to forms of eastern Spain. Cf. also M. avenacea apuana. 

37. CHONDRINA MASSOTIANA (Bgt.). PL 5, fig. 5. 

Shell oval-turriculate, deeply rimate, corneous-chestnut, 
with a very fugacious blue-white bloom when fresh, obliquely 
striate. Spire lengthened, the summit smooth, glossy and quite 
obtuse; 8 whorls, regularly increasing; suture deep, making 
the whorls very convex and a little flattened towards the 
suture ; last whorl slightly keeled around the umbilical fissure, 
and slightly ascending in front. Aperture slightly oblique, 
rounded-oblong, with four little teeth, as follows: one at the 
insertion of the outer lip, the second, deeply immersed, in the 
middle of the convexity of the penult whorl, the third in the 



CHONDRIN^ OP THE PYBENEAN CENTER. 53 

upper part of the columellar lip ; finally a fourth, lamelliform, 
in the middle of the outer lip. These teeth exist only in quite 
adult individuals. Peristome a little everted. Columellar 
margin rectilinear, reflected and dilated, the outer lip strongly 
arched, the margins converging and strongly approaching. 
Length 6y 2 , diam. 214 mm. (Bgt.). 

Pyrenees: damp rocks along the pathway, San-Julia de 
Loria, Andorra. 

Pupa massotiana BOURGUIGNAT, Moll, de San-Julia de Loria, 
1863, p. 18, pi. 2, f. 13, 14 (in Rev. et Mag. de Zool., 1863). 
Pupa penchinatiana BGT., t. c., p. 20, pi. 2, f. 15, 16. 

Closely related to jumttlensis and farinesii, from which it 
differs by having a palatal fold and a small parietal lamella. 

Var. penchinatiana Bgt. PL 5, fig. 11. Umbilical crevice 
more open, and there are two palatal folds, not reaching the 
peristome, while the single one of massotiana is on the peri- 
stome and extends a little way inward. Pathway at San- Julia 
de Loria. 

Pupa penchinatiana has been recorded from the mountains 
of Kabylia in Algeria (Hanoteau et Letouriieux, La Kabylie, 
1872, p. 227). Such a distribution appears anomalous, yet, as 
in the case of Abida brauni, one is perhaps not justified in ex- 
pressing more than surprise. 

Var. sexplicata Bofill. With 2 columellar folds. Catalonia, 
at the Escalas de Sopeira; limestone rocks at Pont de Mon- 
tanyana, and on the margins of the Noguera Ribagorzana 
(Pupa penchinatiana var. sexpUcata Bofill, Bull. Soc. Malac. 
France, iii, 1886, p. 160). 

Var. angustata West. Shell broadly perforate, conic, 
strongly striate, with 7 whorls ; margins of the peristome ob- 
liquely produced downward, approaching one another; the 
aperture therefore narrowed below, outer lip strongly curved 
above. Two lamella on parietal wall, one on columella, two 
folds in palate. Length 7, diam. 2% mm. Alcazar, Spain. 
(Pupa massotiana var. angustata West., Fauna, iii, 1887, p. 
102.) 

Var. confusa West. Shell ovate-turrited, very finely, very 
obliquely and closely, regularly striate, dark cherry-brown; 



54 CHONDRIN^E OF THE PYRENEAN CENTER. 

the last whorl rounded below. Aperture with 4 white teeth: 

1 small angular lamella, 1 sunken, short, strong, parietal, and 

2 horizontal columellar teeth, deep within, high on the colu- 
mella, the second one very small. No palatal folds. Catalonia. 
Coll. Ponsonby, sent by Bofill as P. dertosensis Bof. (P. masso- 
tiana, var. confusa West., Fauna, iii, 1887, p. 102). 

Var. aragonica Fagot. Shell rimate, cylindric-conic, elon- 
gate, buff-reddish, very delicately, irregularly hair-striate ; 
whorls 10, somewhat convex, rapidly and regularly increasing, 
the last ascending, compressed at the base, but not carinate. 
Aperture oblique, oval, with 2 parietal, 1 columellar, 2 palatal 
folds. Angular lamella tooth-like ; columellar small, oblique ; 
palatals rather short, remote ; peristome expanded throughout, 
white, the columellar margin nearly straight, outer margin 
longer, arcuate. Length 6-9, diam. 2-2% mm. Spain. (Pupa 
aragomca Fagot, Catal. Moll. Essera, 1888, p. 17 ; descript. 
from Westerlund, Synopsis, p. 101.) 

38. CHONDRINA DOMICELLA (Westerlund). Shell rimate, 
ovate-conic, obliquely ribbed, rufous-brown. Whorls 8, con- 
vex, regularly increasing, last slightly compressed at base; 
suture deep. Aperture oval, rounded beneath, with 6 lamellae 
and plicae : 1 long angular lamella, 1 much shorter, immersed 
parietal fold, 2 horizontal columellar lamellae, situated high 
and immersed, the upper much the stronger, opposite the 
palatal laminae ; 3 palatal laminae, not marginal, the 1st rather 
immersed, very short, 2d and 3d prolonged inward, especially 
the 2d. Peristome but slightly expanded, the outer margin 
obtusely, angularly curved above. Length 7, diam. 2.5 mm. 
(West.). 

France: Lourdes (Fagot). 

Pupa (Torquilla) domicella West. Verh. zool.-bot. Ges. 
Wien, xlii, 1892, p. 36. Lourdes, France. 

"This species approaches nearest to P. massotiana" (W.). 

39. CHONDRINA PULCHELLA (Bofill). 

Shell perforate-subrimate, conoidal, rather glossy, of a deep 
reddish color, obliquely ornamented with irregular, inconspic- 
uous striae; regularly tapering from the convexity of the last 



CHONDRIN^E OF THE PYRENEAN CENTER. 55 

whorl to the summit; 8 very convex whorls of slow, regular 
increase, separated by a deep suture, the first whorls smooth, 
summit quite obtuse; last whorl a little ascending above 
towards the aperture, ornamented with 2 transverse lines v 
not very long and not reaching the peristome, corresponding 
to palatal folds. Aperture nearly vertical, suboval-truncate, 
with 6 lamellif orm denticles : one angular, white, approaching 
the outer lip at the beginning but diverging inwardly; one 
apertural [parietal], white, immersed, quite near the end of 
the angular lamella; two whitish columellars; two yellowish 
palatals, immersed, feebly prolonged inwardly. Peristome 
simple, narrow, curved towards the apertural angle, slightly 
everted at the base and the columella, the margins approach- 
ing, united by a weak callous. Length 4%, diam. 1%, aper- 
ture 1 x % mm. (Bofitt). 

Spain : arid places of Montsech, Catalonia, on rocks of the 
road of Pont de Mont any ana, at "Ca'n Quinquilla." 

Pupa pulchella BOFILL, Bull. Soc. Malac. France, iii, 1886, 
p. 161. 

By its teeth this Pupa has some resemblance to our var. 
sexplicata of P. penchinatiana (Bourg.), but it is readily dis- 
tinguished by the conoidal form, by the regular taper of the 
spire from the convexity of the last whorl to the summit, by 
the very convex whorls, smaller size, slimmer form, and the 
reddish color (Bofill). 

Possibly related to granum Drap. 

Var. manotiana 'Bgt./ West. Shell umbilicate, oblong- 
conic, costulate-striate, brown; whorls T^, regularly increas- 
ing, convex, separated by a rather deep suture, the last some- 
what ascending in front. Aperture oval, an angular fold at 
the insertion of the outer margin, and a parietal fold in the 
middle, a columellar fold, and two palatal folds, all deeply 
placed, white, short and high, a denticle on the columella 
below ; peristome unexpanded, thin, the outer margin strongly 
arcuate above and approaching the columellar margin. Length 
6, diam. 2 mm. Spain. (Pupa [TorquUla] pulchella Bof. 
var. manotiana Bgt. in sc., Westerlund, Nachrbl. d. m. Ges., 
1894, p. 172.) 



56 CHONDRIN^E OF THE PYRENEAN CENTER. 

Var. ilendensis (Fagot). Shell rimate-perf orate, conic, stri- 
atulate, corneous-buff; spire lengthened-turrited ; whorls 8, 
regular, the last slightly larger, a little compressed at base. 
Aperture oblique, ovate, with 2 parietal, 2 columellar and 2 
palatal folds. Angular lamelliform, the parietal regular, ar- 
cuate ; upper columellar tooth-like, the lower remote ; palatals 
equal, not emerging, exactly opposite the columellar. Peri- 
stome unexpanded, reflected at the umbilicus, the columellar 
margin almost straight. Length 6, diam. 2 mm. Spain. 
(Pupa ilende-nsis Fagot, Catal. Moll. Essera, 1888 ; descript. 
from Westerlund, Synopsis, p. 102.) 

40. CHONDRINA GORBEANA n. sp. PL 3, fig. 12. 

The shell is deeply, rather shortly rimate, long-conic, slowly 
tapering from the last whorl to the obtuse summit, glossy, 
carob-brown, the apex paler, last whorl with a tawny or white 
band behind the lip. Sculpture of fine but irregularly devel- 
oped oblique striae. Whorls moderately convex, the last ascend- 
ing in front, slightly compressed, the base quite narrowly 
rounded. The suture is impressed, bordered by a pale gray 
line. The aperture is truncate-oval, brown within. Angular 
lamella very low; parietal lamella represented by a barely 
perceptible callous ; columellar lamella small, far within, high 
on the pillar. No palatal folds. The peristome is dilated 
above the umbilical fissure, the outer lip not in the least ex- 
panded ; within its edge there is a rather strong, white callous 
throughout. Parietal callous thin and transparent. Length 7, 
diam. to lip-edge 2.8 mm. ; iy 2 whorls. 

Spain : Pena de Gorbea (southern border of prov. Vizcaya). 

This species differs from avenacea by the strong white cal- 
lous within the peristome, as well as by the obsolescence of 
the lamellae and absence of plicae. In one specimen the an- 
gular lamella is short and tubercular but stronger than in the 
type. The species appears to be a degenerate member of the 
bigoriensis group. The locality is not far east of Orduna. 



CHONDRINA, SECTION SOLATOPUPA. 57 

Section Solatopupa Pilsbry. 

Solatopupa PILS., Man. Conch., XXIV, p. 234, type Pupa 
similis (Nov. 9, 1917). 

The shell is calcareous though thin, opaque, whitish with 
more or less cinereous, violaceous or bluish tint, and often 
marbled with brown ; having angular, parietal and columellar 
lamella*, upper and lower palatal and basal folds, or lacking 
teeth. Type C. similis (Brug.). 

Distribution : southern France, northern and western Italy 
south to Tuscany; a group of the olive zone, living on lime- 
stone rocks exposed to the sun. 

a 1 . Palatal folds well developed ; 5 to 7 teeth. 

C. similis, no. 41. 
or. No palatal folds. 

6 1 . Shell conic, thin and fragile, toothless. 

C. psarolena, no. 43. 

6 2 . Shell cylindric, opaque, teeth 3 to 0. 

C. pallida, no. 42. 

41. CHONDRINA SIMILIS (Brug.). PL 4, figs. 1 to 4. 

The shell is shortly rimate, cylindric-fusiform, opaque, pale 
cinereous or nearly white, generally more or less clouded and 
obliquely flamed with cinereous or vinaceous gray, the upper 
whorls of a chamois tint. More or less distinctly striate. 
Upper whorls strongly, the later ones weakly convex, the base 
rounded. Aperture is shortly oval, brown within, having 5 
whitish teeth : angular lamella short and low, joining the lip. 
Parietal lamella rather strong. Columellar lamella obliquely 
ascending within, not emerging, a very low prominence (or a 
small, blunt lamella) below it. Upper and lower palatal folds 
subequal, not approaching the peristome. Peristome white, 
somewhat expanded, scarcely thickened. 

Length 10.5, diam. above aperture 3.2 mm. ; Sy 2 whorls 
(Marseilles). 

Length 14, diam. above aperture 3.6 mm. ; 10 whorls 
(Grasse). 

Length 13, diam. above aperture 4 mm.; 9y 2 whorls (Nice). 



58 CHONDRINA, SECTION SOLATOPUPA. 

Length 8.6, diam. above aperture 2.9 mm. ; 7% whorls 
(Cette). 

Southern France: northern and western Italy south to 
Tuscany; Corsica, 

Bulimus similis BRUGUIERE, Encyclop. Meth., i, 1792, p. 355. 
Pupa simUis Brug., DUPUY, Hist. Moll. France, p. 401, pi. 
20, f. 6. WESTERLUND, Fauna, iii, 1887, p. 94; with (p. 95) 
forms major, minor, variegella, pachygastra, laevigata, dis- 
similis; var. porcellata, in Supplement, 1890, and Synopsis, 
1897, p. 79. MARGIER, Feuille Jeunes Nat., 1901, no. 365, p. 
139 (distribution). CAZIOT, Ann. Soc. Linn, de Lyon, vol. 
50, 1904, pp. 147-154; var. fasciata, and var. guidoni, p. 151 
(synonymy, varieties, distribution) ; Etude sur les Moll. terr. 
et fluv. de la Principaute de Monaco et du Dep. Alpes-Mari- 
times, 1910, p. 309. MERLE, La Nature, 1912, no. 2055, p. 
305, f. 2 (radiograph). Pupa cinerea DRAP., Tableau, p. 61; 
Hist, Moll. Fr., p. 65, pi. 3, f. 53, 54. Ross., Iconogr., pt. 5, 
p. 19, f. 336. KUESTER, Conchyl. Cab., p. 36, pi. 5, f. 4-8 
SCHMIDT, Abh. Nat. Ver. Sachsen u. Thiiringen in Halle, i, p. 
42, pi. 10, f. 82 (anat.). PAGET, Ann. and Mag. N. H., (2), 
xiii, 1854, p. 455 (tooth variation and banding at Nice; in- 
cludes several species). Pupa cinerea Drap., var. pachygaster 
SHUTTLEWORTH, Mittheil. Naturforsch. Ges. in Bern, no. 3, 
June, 1843, p. 18, no description. Pupa quinquedentata Born, 
Pfeiffer, Monogr., ii, 345, with var. minor = Pupa variegella 
Ziegl. in coll.; iii, 547; iv, 673; vi, 314; viii, 384. ROSSM., 
Iconogr., iii, pt. 17, p. Ill, pi. 85, f. 945. MOQUIN-TANDON, 
Hist. Moll. Fr., ii, 1855, p. 352, pi. 25, f. 15-22; with varr. 
major, variegella, minor, pachygaster. BOURGUIGNAT, Malac. 
Chateau d'lf, p. 24, pi. 1, f. 14, 15, with var. turriculata, p. 
25, pi. 1, f. 16. G. NEVILL, P. Z. S., 1880, p. 125 (all along 
the Genoese Riviera, at Menton from the sea to 4000 ft.) ; also 
varr. pr 'cehist orica and speluncarum. Probably not Turbo 
quinquedentatus Born, Test. Mus. Caes. Vindob., 1780, p. 359, 
pi. 13, f. 9. Pupa quinquedentata Born (cinerea Drap.) 
WIEGMANN, Nachrbl. d. Malac. Ges., 1901, p. 12 (anatomy). 
Jaminia quinquelamellata Risso, Hist. Nat. Eur. Merid., iv, 
1826, p. 91, teste Bourguignat. Pupa quinqueplicata POTIEZ 
et MICHAUD, Galerie, etc., i, 1838, p. 163. 



CHONDRINA, SECTION SOLATOPUPA. 59 

This very abundant species varies widely in size, degree of 
variegation and in sculpture, the greatest diversity being 
among lots from the Alpes-Maritimes. Bruguiere's type was 
about 10 mm. long, from the south of France. 

In several lots from Italy there is a short basal fold which 
is often so far immersed that it is not visible in a direct face 
view (pi. 4, fig. 2, Florence). It is not constantly present 
in any lot examined. Paget appears to have noticed this fold 
in specimens from the Riviera, but no other author has men- 
tioned it. 

The subcolumellar lamella varies from distinct to so deeply 
immersed as to be nearly invisible, in different individuals of 
most lots examined. 

In the Alpes-Maritimes it lives in populous colonies, accord- 
ing to Caziot, in well-exposed, sunny places, at Saint-Jeannet, 
Eze, Saint Vallier, etc., up to 1000 m., on calcareous rocks. It 
avoids alluvial plains and is very rarely found on igneous 
rocks. A length of 16 mm. is not common, but specimens that 
long have been found on the rocks of Eze, etc., the diam. 4 mm. 

The distribution of C. similis has been worked out by Mar- 
gier and Caziot. In northern Italy it is rare in Piedmont, 
but extends through Lombardy and as far as Verona in Vene 
tia, according to specimens in this collection. Southward it 
appears to be confined west of the Apennines, being very com- 
mon in Liguria, also in Tuscany (Leghorn, Florence, etc.) 
south to a point 17 kilom. southeast of Rome. In France it is 
found in Corsica, in the departments of Alpes-Maritimes, 
Alpes Basses, and westward, north to Drome and Estaing 
(Aveyron), west to Amelie-les-Bains, in the Pyrenees. It has 
also been recorded from the province of Gerona, eastern 
Spain, from Fonteta and Figuera on the Ampurdan, its south- 
ern known limits. The details of distribution are given by 
Margier, Feuille Jeunes Nat., 1901, pp. 139-141. 

A scalariform abnormal form resembling Clausilia scalaris 
Pfr. has been noted by Caziot (1904, p. 150). 

Since 1848 this species has often been called P. quinqueden- 
tata (Born), but the identity of Bern's species is very uncer- 
tain. The figure looks more like a Clausilia, and may possibly 



60 CHONDRINA, SECTION SOLATOPUPA. 

be one of the long and slender varieties of Abida variabilis; 

the description is inconclusive, and the locality unknown. 
The following described forms are synonyms : 
Form dissimilis West. Quite cylindric, with the spire 

shortly, conically tapering only near the summit, all whorls 

equally and very slowly increasing, somewhat angular below ; 

no angular lamella; parietal lamella very high, very oblique, 

long. Length 12-14, diam. scarcely 3 mm. ; aperture 2% mm. 

See under P. olivetorum. 

Pupa olivetorum Loc. Cylindric, very narrowly elongate, 
the spire feebly acuminate ; 10-12 quite convex whorls, the last 
angular towards the rimation ; suture well marked ; aperture 
small, oboval ; peristome not very thick ; 2 superior folds, one 
very small at the suture, the other long and deeply placed; 
columella simply folded; two deep palatal folds; shell some- 
what thin, ashy white, marbled, finely striolate. Length 15- 
16, diam. 3 mm. Le Midi, Alpes-Maritimes. Gard, Herault, 
etc. (Locard, Ann. Soc. Agricult. Lyon, (7), iii, 1896, p. 183). 

Identical with dissimilis West, and not a species or even a 
valid variety, according to Caziot (1904), but simply a slen- 
der mutation occurring individually in normal colonies in 
many places, never in pure colonies. The tendency to produce 
such cylindric individuals is common in many species of the 
genus. 

Pupa plagionixa 'Bgt.,' Loc. Quite small, almost regularly 
conic, rather squat. 8-10 nearly flat whorls, the last well 
keeled at the base ; suture not very deep ; aperture subrec- 
tangular, with the same apertural teeth as similis; shell solid, 
ashy-white, marbled with blue, finely wrinkled. Length 9-11, 
diam. 3% mm. Alpes-Maritimes, Var, Bouches du Rhone, 
Isere, etc. (Locard, Ann. Soc. d'Agric. Lyon, (7), iii, 1896, 
p. 184). According to Caziot (1904) this is certainly an ab- 
normal shell. 

Pupa variegella Ziegl. This form was mentioned as Chon- 
drus variegellus Ziegl., but not defined by Cristofori and Jan 
(Catalogus, etc., sect, ii, 1832, p. 5, no. 18) ; they give the 
locality southern France. The next notice and first definition 
is by Beck, who has a var. c. varia, with the synonym Pupa 
variegella Zigl., under Torquilla cinerea (Index Moll., 1837, 
p. 87). He refers to Rossmaessler's fig. 336 as illustrating his 
variety. This figure appears to be a strongly variegated speci- 
men of the typical form, about 13 mm. long. Pfeiffer, in 



CHONDRINA, SECTION SOLATOPUPA. 61 

1848, recognized a var. minor, of which he cites P. variegella 
Ziegl. as a synonym, with the description, "Whorls 8, length 
9y 2 , diam. 3 mm." No locality is given. Moquin-Tandon, 
1855, recognized var. variegella for a "shell larger [than 
typical similis], very distinctly marbled," with the locality 
Grasse. This interpretation has been accepted by Ct. Caziot 
(1904). If the name variegella is used at all, it should prob- 
ably be restricted to the shell figured by Eossmaessler, but a 
better course would be to discard it as superfluous. 

The following varieties are given for what they are worth 
not much in most cases. 

Var. major Moq. Shell larger, of the same color. Sainte- 
Lucie near Narbonne; Draguignan (Moq.-Tand., L c., pi. 25, 
f. 22). The figure measures 16.5 mm. long, 4.5 diam. 

Var. minor Moq. Shell smaller. Caziot states that this 
form is common at the chateau of Nice and other places in the 
Alpes-Maritimes. Length 9-10, diam. 3 mm. ; on the Var 
river, 8x2% mm. 

Var. pachygaster 'Shuttl.,' Moq. Shell of the same size as 
the typical form [9-13x3-4 mm.] or smaller, more ventricose. 
Corsica: Ajaccio, Saint-Florent (Moq.-Tand.). The name is 
preoccupied. 

Form laevigata West. Irregularly, finely striate or almost 
smooth. 

Var. porcellata West. Shell subfusiform, rather swollen in 
the middle, strongly tapering above, very acute, regularly and 
closely rib-striate; length 11, diam. in the middle 3% mm. 
Spezzia, Italy. 

Var. fasciata Caziot. Each whorl ornamented with a brown 
median band, as in Cochlicella. Aqueduct de Carpentras 
(Alpes-Maritimes) ; also in Provence. 

Var. guidoni Caziot. Differs from the type by its colora- 
tion, the cask-like form of the whorls of the spire, the more 
pronounced suture and the nearly smooth shell. Pioggiola 
pres le Mont Padro, Corsica, on granite. 

Var. Juliana Issel. Color reddish, more or less intense, 
probably due to the ferruginous material abundant in the 
same locality. Baths of S. Giuliano, Prov. Pisa. (Pupa quin- 
quedentata var. Juliana ISSEL, Memorie Soc. Ital. di Sci. Nat., 
ii, no. 1, 1866, p. 21.) 



62 CHONDRINA, SECTION SOLATOPUPA. 

Var. pr cellist orica G. Nevill. The commonest subfossil shell 
in all the deposits except F ; though varying in countless ways, 
it always preserves a distinct "facies" from that of the pre- 
ceding living form [similis] ; it can invariably be distin- 
guished by the much stronger and more regular striation, not 
so oblique, so flexuous, or so inclined to be subobsolete; the 
aperture is also invariably smaller and more contracted, less 
everted as a rule, with both margins straight and parallel, in- 
stead of more or less rounded; the folds appear scarcely to 
differ at all. Long. 14, diam. 4 mm. Type of the var. from 
deposit B, Menton (NevUl) . 

Var. speluncarum G. Nevill. A more distinct and charac- 
teristic variety than the preceding, which I only found in de- 
posit F, where it was abundant, and in the interior of the cave 
itself (as I have already mentioned). A very short, convexly 
swollen, tumid form, of only 9 instead of 10 whorls, the last 
being proportionally much broader; striation more like that 
of the typical form than of the preceding variety; the short, 
quadrangular, unusually everted aperture, with remarkably 
thickened and reflected peristome, seems to be its most char- 
acteristic feature, the margins being more convexly rounded 
and united by a distinct (instead of subobsolete) callosity. I 
can see no difference in the folds (or teeth). Long. 10%, 
diam. 3% millim. From deposit F (Nevill). Menton. 

Types, Indian Museum, Calcutta. 

Var. isabella Caziot. A variety of lengthened shape, very 
slightly swollen, robust, 11 very slightly convex whorls. It 
differs from the type especially by the dimensions, length 17- 
18, diam. 4%. mm. 

The typical similis varies between length 9-15, diam. S-S 1 /^ 
mm. The var. major West, has, length 16 mm., with a diam- 
eter equal to my variety ; it is thus more obese. The var. isa- 
bell& approaches var. variegella Ziegler which has been found 
at Grasse, and which is more ventricose and more distinctly 
marbled. The variety of the Courmettes has, besides, the 
depth of the aperture of a dirty yellow color. Some specimens 
show a single band, such as is characteristic of my var. uni- 
fasciata. Southern slope of the Courmettes between the tor- 



CHONDRINA, SECTION SOLATOPUPA. 63 

rent du Loup and the village Tourette (Alpes-Maritimes) 
(Caziot). 

Pupa similis Brug. var. isabellcu CAZIOT, Feuille Jeunes 
Naturalistes, no. 513, 1913, p. 148, fig. on p. 147. 

42. CHONDRINA PALLIDA ('Phil.,' Rossm.). PL 4, figs. 5 to 8. 

Shell with a shallow umbilical crevice, ovate-fusiform, 
pointed; lilac-ash-gray, with irregular brownish flecks and 
horn-colored apex; weakly striate, somewhat glossy. Eight 
somewhat convex whorls. Aperture semi-ovate, the throat 
yellowish-brown ; peristome expanded, sharp, simple, the colu- 
mellar margin shorter and straighter than the outer lip. 
Columella somewhat callous, advancing. Length 4, diam. 1^ 
lines, 7 whorls (Rm.). 

Upper Italy (Philippi) ; Riviera from Toulon to Porto 
Venere near Spezia (v. Martens). 

Pupa pallida Phil, in lit., ROSSMAESSLER, Iconogr., ii, pt. 11, 
1842, p. 11, pi. 53, f. 732. KUESTER, C. Cab., p. 86, pi. 12, f. 
12, 13. PFR., Monogr., ii, p. 308. VON MARTENS, Nachrbl. d. 
m. Ges., 1900, p. 70 (between Rapallo and Zoagli, very abun- 
dant on limestone cliffs) . Pupa patula Menke, teste Pfr., 
Monogr., viii, p. 363. 

The type figure is copied, fig. 7. It is not known where 
the very small, toothless form which served as type of this 
species occurs; but it is doubtless merely a local race or 
colony of the prevalent larger form known as amicta Parr. 
It is unfortunate that the species was first described from a 
peripheral mutation. 

Var. amicta Parreyss. Shell deeply rimate, cylindric-tur- 
rited, solid, smooth, bluish-white; spire long, corneous above, 
gradually terminating in a rather obtuse cone ; suture simple. 
Whorls 9, scarcely convex, the last about two-sevenths the 
total length, subtuberculate-compressed at base. Aperture 
nearly vertical, truncate-oblong, fleshy-brown within, with 3 
teeth: the first angular, second standing apart deep on the 
belly of the penult whorl, the third fold-like, deep on the colu- 
mella. Peristome white, a little expanded ; the margins some- 
what unequal, right margin toothless. Length 12, diam. 3 
mm.; apert. 3% mm. long, 3 wide (Pfr.). 



64 CHONDRINA, SECTION SOLATOPUPA. 

Pupa amicta Parreyss MS., PPR., Malak. Bl., i, 1854, p. 67 
Monogr., iv, 668 ; vi, 307. BOURGUIGNAT, Malac. terr. chat, 
d'lf, p. 25, pi. 1, f. 11-13. ISSEL, Bull. Soc. Malac. Ital., vii, 
1881, pp. 208-212; with var. excelsa, p. 210. Pupa pallida 
var. tridentata EOSSMAESSLER, Iconogr., iii, pt. 17, 1858, p. 
Ill, pi. 85, f. 944. P. pallida var. dentiens MARTENS, Die 
Hel., 1860, p. 287. 

Distinguished from typical pallida by the presence of small 
angular, parietal and columellar teeth ; but these are variable 
in development in adult shells of the same lot. In some of the 
examples from Nervi, Liguria, the angular and parietal are 
reduced to slight traces (pi. 4, fig. 5), and there is no colu- 
mellar. Such specimens approach the original pallida rather 
closely, but all I have seen are slightly larger, 9y 2 to 10 mm. 
long. Other shells of the same lot have two distinct teeth and 
a trace of the columellar, or none. Specimens from Spezia 
have the three teeth distinct (pi. 4, fig. 8). This may be 
taken as type locality of amicta, since Pfeiffer gave only Sicily 
(1854) and northern Italy (1859). 

Specimens from Grasse (Terver) have the lip more ex- 
panded than Italian shells seen; two teeth, with rarely a 
small columellar also (pi. 4, fig. 6). The parietal is often not 
visible in a direct face view. They vary in size : 

Length 12, diam. above aperture 3.3 mm. ; 9 whorls. 

Length 9.5, diam. above aperture 3 mm. ; iy 2 whorls. 

Although amicta has been reported from the Chateau d'lf 
(Dep. Bouches-du-Ehone) and the peninsula of Saint-Man - 
drie, near Toulon (Bourguignat, 1860), and I have specimens 
from Grasse (Var), yet it appears to be w r anting in the Dep. 
Alpes Maritimes, where the snail fauna has been studied care- 
fully by Ct. Caziot and many others. It occurs at suitable 
places along the whole coast of Liguria, but the colonies, while 
rather numerous, appear to be of limited extent. 

According to Issel, C. amicta ordinarily lives on limestone 
rocks near sea level, but it has also been found inland and at 
some elevation, at Bavari and Traso in the Bisagno valley, five 
or six kilometers from the coast, at about 100 meters elevation, 
and above Cassagna and Statale, ten to twelve kilometers in- 



PLATE 1 



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SECOND SERIES: PULMONATA. 



MANUAL 



OF 



CONCHOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED BY 

GEORGE W. TRYON, JR. 

OONTINITKD BY 

HENRY A. PILSBRY, Sc.D., 

SPECIAL CURATOR DEPARTMENT OF MOLLUSCA, ACADEMY OF NATURAL 
SCIENCES OF PHILADELPHIA. 



Part 



98. Col red 



PHILADELPHIA : 

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES 

OF PHILADELPHIA. 



CHONDRINA, SECTION SOLATOPUPA. 65 

land at about 450 meters above the sea. Upon these specimens 
he bases the following variety. 

Var. excelsa Issel. The shell more slender, more acuminate, 
smaller, and having the aperture more regularly oval, and 
with the margins more approaching. 

43. CHONDRINA PSAROLENA (Bgt.). PL 4, figs. 9, 11. 

Shell rimate-p erf orate, conic-oblong, fragile, a little pellu- 
cid, obliquely striatulate; corneous, irregularly marked with 
longitudinal streaks of cinereous or whitish-blue. Spire conic, 
the apex acute, corneous, smooth; whorls 7, very convex, 
parted by a deeply impressed suture, the last whorl not one- 
third the whole length. Aperture rounded, the peristome 
simple, acute, not reflected; columella simple; columellar 
margin dilated, a little expanded ; margins strongly converg- 
ing, joined by a thin callous. Length 7-8, diam. 4 mm. ; height 
of aperture 2.5, width 1.75 mm. (Bgt.). 

Alpes-Maritimes, in the gorges of the Saorge (type loc. ? 
Bgt., Caziot) ; also cavern la Giachera in the Nervia valley, 
Liguria, and on the Costa di Drego above a little affluent on 
the left of the Argentina or Tazzia river, at 1000 meters eleva- 
tion (Issel) ; fossil in the pleistocene deposits of Menton 
(Nevill). 

Bulimus cinereus MORTILLET, Coq. fluv. et terr. de Nice, in 
Bull. Soc. Hist. Nat. Savoie, iii, 1851, p. 96. Bulimus cinereus 
DUMONT et MORTILLET, Descript. somm. des esp. nouv., in 
Prospectus de 1'Hist. Moll, terr., etc., Savoie et du Basin du 
Leman, 1852, p. 3. ROSSMAESSLER, Iconogr., iii, pt. 17-18, 
1858, p. 102, pi. 84, f. 929. Not Bulimus cinereus Reeve, 1848. 
Bulimus psarolenus BOURGUIGNAT, Amenites Malac., ii, 1859, 
p. 116, pi. 15, f . 1, 2. Pupa psarolena Bgt., STABILE, Bull. 
Malac. Ital., i, 1868, p. 33 (discussion of synonymy). NEVILL, 
P. Z. S., 1880, p. 124. CAZIOT, fitude Moll. Monaco, et Alpes- 
Marit., 1910, p. 313, pi. 9, f. 23, 24. Pupa mortilleti v. MAR- 
TENS, Die Hel., 1860, p. 287 (n. n. for B. cinereus Mort). 

I have not seen this species, which appears to take the place 
of pallida in the Alpes-Maritimes, but apparently at much 
greater elevations than pallida. It lives on dusty limestone 



66 UNDETERMINED PUP^E. 

rocks according to Caziot. Bourguignat found it on rocks of 
the gorges above the village of Saorgio, near Nice, in great 
abundance. Mortillet's locality was back of Menton. G. 
Nevill found an elongated form of the species in the pleisto- 
cene of Menton. Caziot 's figures also represent more length- 
ened examples. 

Unidentified and Undescribed Species of Abida, Granopupa 
or Chcmdrina. 

Pupa affinis Aradas et Maggiore, Catalogo regionato della 
Conchiglie viventi e fossili di Sicilia, p. 116 (Atti Accad. 
Gioenia, xv, 1839; xx, 1843). Shell conic-cylindric, ashy, 
smooth, solid; aperture 5-toothed; margin white, slightly re- 
flected. Alt. 3%, diam. 1% lines. Coast of Catania. Differs 
from P. variabiUs by the form, cylindric to the upper third, 
and the situation of the teeth, of which one is in the insertion 
of the lip, second in the labrum, two minute and contiguous 
on the columella, and the fifth, laminar, in the labium. I have 
not seen this publication, and take the diagnosis from Philippi, 
Enum. Moll. Sicil., ii, 220, who quotes from A. & M. Ross- 
massler's use of the name P. affinis was probably prior. 

Pupa contorta Calcara. T. cylindraceo-conica, striata, cor- 
neo-cinerascens ; anfr. 8 plano-convexi, ultimus ante finem 
notabiliter contortus; apertura alba, 8-dentata, margine eras- 
siusculo, subreflexo. Long. 3 lin. Palermo, Sicily. (Calcara, 
Effem. scient. per la Sicil., N. 75, p. 101.) 

Torquilla gastrodes Zigl. Beck, Index Moll., 1837, p. 58. 
Dalmatia. 

Jaminia heptodonta Risso. T. glabra, nitida; anfr. 11, line- 
olis obliquis impressis sculpti; apertura septemdentata ; epi- 
dermis fuscescens. Circa Nizza. Long. 0.009 (Risso, Hist. 
Nat. Eur. Merid., iv, 1826, p. 92). 

Torquilla hornbeckii Villa. See vol. xvi, p. 24. 

Pupa libanotica Tristram. Shell cylindric-oblong, corneous 
or whitish, under the lens irregularly striate, the apex very 
obtuse ; whorls 10, a little convex, the suture impressed ; first 
5 whorls very obtuse and rapidly increasing ; 6th whorl about 
equal to the 7th ; the last and penult strongly contracting, the 



UNDETERMINED PUP^. 67 

last carinate at the base and contracted at the aperture. 
Aperture semi-oval, one-toothed, the tooth standing on the 
callous; peristome whitish, reflected, continuous in a callous. 
Length 11, diam. 4% mm. (Tristram). 

Palestine: Ainat, Lebanon (Tristram). 

Pupa libanotica TRISTRAM, Proc. Zool. Soc. Lond., 1865, p. 
538. 

This species has not been figured. It may belong to the 
Enida, but has been considered a Torquilla. 

Pupa plusiodonta Holmberg. Shell subperforate, fusiform- 
ovate, thick, scarcely shining, unequally subplicate and stri- 
ate, brownish fulvous, paler towards the apex ; spire subovate- 
conic, the apex obtuse, suture deep; whorls 7%, gradually 
increasing, convex, the first brownish-tawny, smooth, the rest 
darker, the last very little ascending, one-third the length. 
Aperture oval, contracted by 8 pliciform teeth : two parietal, 
of which one is produced to the suture, the other more deeply 
placed, parallel, further from the right margin; two on the 
columella, of which one further from the base is higher than 
the other ; one basal, is smaller than the rest ; and three mar- 
ginal, parallel and equidistant, the first nearly basal, the 
second going further in, the third deep ; these being visible ex- 
ternally as whitish lines. Peristome acute, expanded, pale, 
whitish externally and within; columellar margin straight, 
the basal regularly curved, right margin moderately curved, 
becoming strongly so near the suture; margins separated, 
joined by a thin callous. Length 9%, diam. 3%, aperture 3^2 
mm. long, 2% wide (Holmberg}. 

Argentina: on the left bank of the Rio Negro near where 
it enters the sea, a single specimen found among fluviatile, 
land and sea shells (Pupa plusiodonta HOLMBERG, Apuntes de 
Historia Natural, i, no. 2, Feb. 1909, p. 27). 

It has not been figured. The description suggests a form 
of Abida or Chondrina. As the unique type was found among 
shells which had been collected some twenty-five years before, 
it appears likely that a stray European shell, perhaps from 
Italy, had got among them in the meantime. 

The following names, without further information, except 



68 VERTIGININAE. 

that they belong to the submenus TorquiUa, are from Schau- 
fuss, Molluscorum Systema et Catalogus, Samml. Fr. Paetel, 
1869: 

Pupa dalmatina Prtsch. Dalm. P. 85. 

Pupa parmjena d'Orb. Cuba. P. 85. 

Second edition of the same: Catalog der Conchylien- 
Sammlung von Fr. Paetel, 1873. 

Pupa baldensis Parr. M. Bald. P. 107. 

Pupa depresstiabris Parr. Gorz. P. 107. 

Pupa formosa Parr. Balkan. P. 108. 

Pupa inconspecta Parr. Serb. P. 108. 

Pupa labiosa Parr. Grasse. P. 108. [==P. braunvi var. 
labiosa Moq. ?] 

Pupa oblonga Parr. Italia. P. 108. 

Pupa prolongata Parr. Banat. P. 108. 

Pupa proximo, Rm. Illyr. P. 108. 

Pupa striatula Parr. Hispan. P. 108. 

Pupa turgida. Zglr. Illyr. P. 109. 

The same, edition of 1883 : 

Pupa anceps Fagot. Europ. P. 158. 

The same, 4th edition, 2d part, 1889 : 
Pupa scalariformis Em. Sicil. P. 304. 

Pupa tricolor Sowerby, Conch. Icon., xx, pi. 20, f . 190. ? = 
tricolor Villa. 

Subfamily VERTIGININAE. 
Pupillid snails without inferior tentacles. 

This group comprises Pupillidae having compact, oval, 
ovate or cylindric shells of quite small or minute size, usually 
iy 2 to 3 mm. long,' from brown to amber or olivaceous color, 
having the typical 6 teeth of the family, or varying to none 
or to a greater number. The axis is perforate, but usually 
closed in the adult stage. Vertigininae are abundant in 
Holarctic, Polynesian and Hawaiian faunas, almost wholly 
wanting in South American and African. 



VERTIGO. Oil 

The subfamily, in its present limits, is a new group. The 
family Vertigimdae of Stimpson (Shells of New England, 
1851, p. 53) comprised only the genus Vertigo including 
V. simplex Gld., but Stimpson also characterized his family 
by the want of inferior tentacles. 

From the literature it may be gathered that some Pupillidae 
certainly belonging to other subfamilies are deficient in ten- 
tacles, and according to Gredler, a reliable observer, V. genesii 
has inferior tentacles. Further careful observations on many 
species are needed. The inferior tentacles are usually very 
short in Pupillidae, and may therefore be overlooked easily. 

A general likeness may be traced between the shells of the 
genera now referred to Vertigininae ; it appears to be a na- 
tural group; though the variation is so great throughout the 
family that definitions sufficiently elastic to cover any of the 
groups larger than genera become too comprehensive to be 
diagnostic. 

The genera fall into two main geographic divisions: 1. 
northern or mainly Holarctic, including Vertigo, Trunca^ 
tellina, Sterkia and their satellite groups, and 2. Polynesian 
and Tropical, with Nesopnpa and the associated groups. 
Places where these divisions overlap are extremely few. 

A list of fossil genera and species of Vertigininae follows 
the account of the genus Vertigo. 

Genus VERTIGO Muller. 

Vertigo 0. F. MULLER, Vermium terrestrium et fluviatilium 
Hist., ii, 1774, p. 124, monotype V. pusilla Mull. 

Isthmia GRAY, London Medical Repository, xv, 1821, p. 239, 
for "Helix Isthmia cylMrica Drap, t. 3, f. 30, 31," => V. 
pygmcea Drap. Cf. BALL. Tr. Wagner Inst. iii, pt. 2, p. 248 ; 
Nautilus xvii, 1904, p. 114, and NEWTON and HARRIS, Proc. 
Malac. Soc. London i, p. 72, footnote 1. GRAY, P. Z. S. 1847, 
p. 176, type Vert, nitida edentida Drap. 

Al&a JEFFREYS, Trans. Linn. Soc. London xvi, 1830, p. 357. 
GRAY, P. Z. S. 1847, p. 176, type A. paliwtris Jeffr. = 
V. antwertigo. PILSBRY, Nautilus xviii, 1905, p. 119. 



70 VERTIGO. 

Staurodon LOWE, Proc. Zool. Soc. 1854, p. 214, type 
P. pygmaa Drap. Not Staurodon Lowe, 1852. 

Dexiogyra STABILE, Moll. terr. Viv. du Piemont, 1864, p. 
104 (in Atti della Soc. di Scienze Nat., Milano, vi), for V. 
moulinsiana, V. pygm&a, V. antivertigo. V. antwertigo here 
designated as type. 

Dexiogira De BETTA, Moll. Prov. Veron., 1870, p. 83. 

Nearctula STERKI, Nautilus vi, 1892, p. 5, type by orig. des., 
V. calif ormca Rowell. 

Haplopupa PILSBRY, Nautilus xi, Feb. 1908, p. 119, mono- 
type V. daUiana. 

Pupa DRAPARNAUD, 1801, and of many subsequent authors. 
Not Pupa Bolten, see Vol. XXIV, p. 267. 

Small, oval, oblong or ovate, compact Pupillids, the summit 
very blunt; usually glossy and some shade of brown. Aper- 
ture having the typical six teeth of Pupillidae, none of them 
concrescent, part or all of them sometimes wanting; angular 
lamella not marginal when present. Outer lip straightened or 
bent inward in the middle. Animal without inferior tentacles. 

Type, V. pusflla Mull. 

The apertural teeth of Vertigo show great modifications in 
number and size. There are often small teeth accessory to 
the six typical ones, and in other forms there has been re- 
duction of teeth, the basal, angular and . upper palatal being 
lost in many species. Other forms have gone further, tooth- 
less species resulting from degeneration of teeth in several 
phyletic lines within the genus in both Europe and America. 
These modifications make a diagnosis of the genus almost im- 
possible. 

These teeth, though among the best specific characters, are 
subject to wider variation than usual. The secondary den- 
ticles are especially likely to vary ; but also the basal fold, the 
angular lamella and often the upper palatal fold are variable, 
being either present or absent in many species. 

In many Vertigos the basal fold is subcolumellar in position, 
rather than on the basal margin at the foot of the eolumella. 
It has therefore often been counted as a second columellar 
tooth in descriptions. 



VERTIGO. 71 

In Europe, several diverse forms have given rise to smistral 
species, but up to this time no sinistral species or specimens 
have been reported from America or Japan. 

The shells of Vertigo are usually clean, but V. dalliana at 
least occurs daubed with slime and dirt, like many 
Gastrocoptinae. 

Key to the subgenera of Vertigo. 

a 1 . Angular lamella curving towards and connected with the 
outer lip ; lamella and plicae well developed. 

Subgenus Ptychal&a, species no. 41. 

a 2 . Angular lamella, when present, standing free of the peris- 
tome, not emerging to the edge. 

6 1 . Columellar lamella short, lamellar or toothlike, sub- 
horizontal or inwardly ascending a little, sometimes 
absent. Subgenus Vertigo. 

6 2 . Columellar lamella subvertical, spirally ascending 
(pi. 5, fig. 18) ; upper palatal fold entering very 
deeply, the lower shorter or wanting. 

Subgenus Vertilla, species nos. 81, 82. 
b 3 . Columellar lamella subvertical, the upper palatal 
fold shorter than the lower or wanting; no angular 
lamella or basal fold in known species. 

Subgenus Vertttlaria, species no. 31. 
& 4 . Columellar lamella curving down at its inner end 
(pi. 13, figs. 2, 4) ; lower palatal fold entering very 
deeply, much longer than the upper. 

Subgenus Angustula, species nos. 32, 33. 

Nomenclature. 

In his original publication of Isthmia, 1821, Gray refers to 
only one species, identical with Vertigo pygnuea. On ac- 
count of his somewhat irregular system of indicating the 
groups of " Helix," Harris & Newton have ruled this paper 
out as not binomial, stating that the name must date from 
Gray, 1840. It may be observed that their ruling has not 
been followed by any other author. All the names of a work 
stand or fall together ; one cannot accept part as binomial and 



72 VERTIGO. 

reject the remainder as polynomial. In the present case, Gray 
was in effect using subgenera, and not what is commonly 
understood as polynomial terms; I have not found Gray's 
publication of Isthmia in 1840, though he quotes that date 
in his paper of 1847, where V. mtida (presumably = Pupa 
edentula Drap.) is mentioned as the type. Isthmia has been 
extensively used in Europe for the genus Truncatellwa Lowe. 

Al&a was proposed by Jeffreys for dextral Vertigines, but 
later, in the British Conchology, he considered "this generic 
addition to be quite useless and untenable." Dall has ex- 
pressed a similar view; and I fully agree with these authori- 
ties, although Boettger, Westerlund and most German authors 
have used Al&a. Dexiogyra was proposed for the same group, 
and is even less defensible. 

The value of Nearctula and Haplopupa remains to be as- 
certained when the affinities of their type species are better 
understood. They do not seem separable from the subgenus 
Vertigo by sufficient characters, as at present known. 

Distribution of Vertigo. 

Vertigo is essentially a northern genus of humid stations 
or regions. The group of Vertigo modest a is circumpolar, 
numerous in races and individuals above the parallel of 60, 
and extending south at increasing elevations, in the Canadian 
and Transition zones, to about lat. 33 in Arizona, there only 
in mountain-top colonies. In Europe it does not go so far 
south, only to the Austrian Tyrol. 

Only 5 species of Vertigo have been reported from the 
American tropical fauna (Mexico and the West Indies), 3 of 
them being identical with northern species, the other 2 doubt- 
fully distinct from the widely spread V. ovata. This identity, 
together with the fact that they appear to be among the rarest 
snails in those regions, surely indicates that Vertigo is a late 
comer in the American tropics, if not, as I suspect, imported 
by man. Species have been found as far south as about 
18 N. Lat. 

In the Palsearctic fauna, Scandinavia and the eastern Alps 
(Tyrol) have the greatest development of the genus. No 



VERTIGO. 73 

species is known from as far south as the 35th parallel, and 
only a few species (in Algeria and Tunis) approach it. 

Where Vertigo occurs in arid regions it inhabits humid 
places, usually at high levels. Thus, in the arid zone of south- 
ern New Mexico and Arizona, Vertigo is often abundant in 
the forests mainly above 8000 ft., where there is abundant 
shade and moisture. It may usually be found where there 
are aspens. 

Part of the species of temperate latitudes of Europe and 
America are identical or very similar. Their distribution 
probably dates from the Pliocene. Doubtless much if not all 
of the evidence of the route has been destroyed by glacial 
action. While the present distribution of these closely allied 
forms suggests emigration around the north Atlantic, a 
Pliocene distribution through northern Asia seems far more 
likely, though northern Pliocene fresh-water deposits which 
might contain traces of this fauna are still unknown. 

This interpretation is in harmony with the facts of present 
distribution relative to the circumpolar modesta group, where 
the data indicate continuity of distribution within post- 
Pliocene time from Alaska west to Lapland, and a southern 
movement during Glacial time. The other Vertigines, less 
tolerant of cold and for the greater part more differentiated, 
no doubt were pressed south earlier, and were exterminated 
in Siberia and the mountain states of western America. The 
survivors are chiefly at the extremes of their former range, in 
Europe and in the eastern half of North America. 

Vertigo pygm&a is to be regarded as a member of that 
Pliocene fauna, still unchanged in the two hemispheres, though 
in both giving rise to various divergent races. V. ovata and 
antivertigo are but slightly differentiated. V. moulinsiana 
and V. ventricosa differ slightly more, and a few other species 
could be similarly paired. In both hemispheres there are also 
far more differentiated forms of Vertigo, mainly southern in 
distribution, indicating a long period of evolution in America 
as well as in Europe. 

Where Vertigo arose may perhaps never be known. The 
presence of typical species in the Oligocene of Central Europe, 



74 AMERICAN SPECIES OF VERTIGO. 

and of highly peculiar species in the Bermudas and Bonins 
outlying, degraded old volcanic masses which have been base- 
levelled and further reduced by subsidence indicate a long 
period of evolution in the northern continents. Its absence 
in the Atlantic archipelagos Azores, Madeira, etc., is per- 
haps due to belated arrival of Vertigo proper in extreme 
western Europe. There are a few Eocene species probably 
belonging to the Vertigininae, which indicate the presence of 
the subfamily in western Europe prior to the Oligocene. 
The single genus Stawrodon in Madeira points in the same 
direction. 

The species of Vertigo are considered in four groups : 

I. American species. 

II. Species of Japan and the eastern border of Asia. 

III. Palaearctic species of Europe, Asia and northern Africa. 

IV. Tertiary species. 

I. AMERICAN SPECIES OP VERTIGO. 

After the early work of Say and A. Binney, the American 
Vertigines were studied by Dr. Gould, and twenty years later 
by Professor E. S. Morse, who was a pioneer in the critical 
study of our minute land shells. His results, published in 
various papers between 1864 and 1868, were incorporated 
by W. G. Binney in his Terrestrial Mollusks vol. V, becoming 
a permanent addition to science. In 1889 Dr. V. Sterki pub- 
lished the first of a long series of papers on Pupillidas which 
have made him the first American authority on the subject. 
We owe to him the discovery of many of the most remark- 
able of our Vertigos, and also valuable critical work on the 
older species, their mutations and races. Many collectors 
were induced by him to search for the small Pupillidse, with 
gratifying results. 

In the preparation of the following monograph the author 
has had frequent occasion to consult Dr. Sterki. It will be 
seen that his published work, letters and MS. notes have been 
quoted freely ; and the monograph is much the better for his 
generous assistance. In every case Dr. Sterki 's contributions 
have been acknowledged in the text. 



AMERICAN SPECIES OF VERTIGO. 75 

Specimens of all of the American species except V. arthuri 
have been examined in the course of this work, including types, 
paratypes or specimens from the describer of all except a few 
of the older species. 

SPECIES OF BERMUDA. The species, while strongly differen- 
tiated, are evidently related to forms of the eastern United 
States. 

V. marki Gul., no. 18 (related to V. trident ata] . 

V. numellata Gul., no. 7 (remotely related to the ovata 
group). 

V. bermudensis Pils., no. 33 (related to V. milmm). 

SPECIES OF THE WEST INDIES. These tropical forms may 
be presumed to be of northern origin, since all of them are 
either identical with species of the United States or at least 
not satisfactorily shown to be different. Possibly all may 
be importations within the period of commerce ; I cannot other- 
wise understand the absence of differentiation in those I have 
been able to examine. The following have been recorded. 

V. ovata Say, no. 4. Cuba, Haiti, Porto Rico. 

V. neglecta Poey. Cuba ( ? = ovata, no. 4) . 

V. hexodon C. B. Ad. Jamaica ( ? = ovata, no. 4) . 

V. gouldii Binn., no. 11. Jamaica, Cuba, St. Croix, Tortola. 

V. milmm Gld., no. 32. Jamaica. 

V. cubana Dall. "Cuba" (== Hawaiian Lyropupa). 

Key to American groups of Vertigo. 

ft 1 . Columellar lamella crescentic, its inner end turning down- 
ward; lower palatal fold entering to the dorsal side and 
turning down at the end. Teeth all well developed. 

Subgenus ANGUSTULA, species nos. 32, 33. 
ft 2 . Columellar lamella subvertical; lower palatal fold short, 
a conspicuous external constriction over it (pi. 8, fig. 10) ; 
3 or rarely 4 small teeth. Southern states. 

Subgenus VERTILLARIA, species no. 31. 

ft 3 . Columellar lamella short, entering horizontally or ascend- 
ing inwardly, or wanting. Subgenus Vertigo. 
6 1 . Lower palatal fold entering to the dorsal side; 6 
well developed teeth; a very high crest. Bermuda. 
Group of V. numellata, species no. 7. 



76 AMERICAN SPECIES OP VERTIGO. 

& 2 . Without the above combination. 

c 1 . 6-9 teeth; crest or auricle generally well de- 
veloped; rather strong, dark colored forms 
(pi. 6). Group of V. ovata, species nos. 1-6. 
c 2 . 2-6 teeth (pi. 7). 

Group of V. pygmcea, species nos. 8-20. 
c 3 . 4-6 teeth ; shell cylindric or oblong, striate. 

Group of V. color adensis, species nos. 21-24. 
c 4 . 0-5 teeth, never a basal fold ; cylindric or ovate 
(pi. 10). 

Group of V. modesta, species nos. 25-28. 

c 5 . 0-4 teeth, no basal fold; no trace of a crest; 

cylindric, ribbed or with traces of ribs (pi. 9). 

Group of V. caUfornica, nos. 29-30. 

The keys to species are often insufficient for the identifica- 
tion of critical or variable forms, but they are offered as better 
than no guide. I have not been able to make differential 
definitions of the groups of V. ovata, V. ventricosa, V. modesta 
and V. caUfornica. 

Group of Vertigo ovata. 

Rather dark-colored, glossy shells usually having 7 to 9 
teeth, never less than 5, the basal fold always present, but 
often subcolumellar in position ; angular lamella present, often 
an infraparietal also. Crest and auricle generally distinctly 
developed. 

Key to species. 

1. Shell distinctly striate, the palatal callus, crest and 
impressions behind the lip weak or wanting. Austro- 
riparian. V. rugosula, no. 1. 
Shell weakly or scarcely striate. 2. 

2. Last half whorl deeply impressed or constricted over the 
palatal folds (pi. 8, fig. 14). Lamellae and lower palatal 
fold long. Southern states. 3. 
Last whorl only moderately or slightly impressed over 
the lower palatal fold (pi. 6, fig. 2). 4. 



AMERICAN SPECIES OF VERTIGO. 77 

3. 1.8 x 1 mm. ; seven teeth. V. alabamensis, no. 2. 
1.5 x 1 mm. ; six teeth. V. a. conecuhensis, no. 2a. 

4. More than 2.6 mm. long, of 6 whorls; spire long, the 
aperture relatively small. V. morsei, no. 3. 
Shell smaller. 5. 

5. Ovate, the spire very convexly conic; length about 2.2, 
diam. 1.3 to 1.4 mm. ; 5 whorls ; usually with 7 to 9 teeth. 

6. 

Oblong-conic, the spire more slender than in V. ovata ; 2.5 
x 1.35 mm., 5y 2 whorls. California. V. berryi, no. 5. 
Oblong-cylindric, 2 x 1.1 mm., 5 whorls. Rocky Moun- 
tains. V. binneya/na, no. 6. 

6. Crest and auricle well developed. 7. 
Crest and auricle very weakly developed. California. 

V. o. mariposa, no. 4&. 

7. Teeth moderately large. V. ovata, no. 4. 
Teeth longer and stronger. Texas. V. o. diaboli, no. 4a. 

1. VERTIGO RUGOSULA Sterki. PL 8, figs. 1, 2, 3. 

"Related to V. ovata and gouldti, in shape more elongated 
than the latter, more cylindrical and somewhat larger. Aper- 
tural parts and lamellae much like those of ovata, but the 
columella is decidedly longer and straighter, and the inferior 
columellar lamella is distinctly placed on it. Of a peculiar 
formation is the surface : of the 5 well rounded whorls, about 
one and a half of the upper are nearly smooth ; the following 
with exception of the last are distinctively and regularly 
striated, the last very finely but distinctly rugose in the sense 
of the lines of growth; near the aperture again striated. 
Color, dark chestnut. Length 1.8 to 2, diam. 1.1 mm." 
(Sterki). 

South Carolina: Sullivan's Island, type loc., W. G. Mazyck. 
Tennessee: near Cowan and near Sherwood, H. H. Smith. 
Mississippi: Pass Christian, B. Walker. Louisiana: De Soto 
Parish, L. S. Frierson; Lovett Landing, Morehouse Parish, 
C. B. Moore. Oklahoma: Limestone Gap, Ferriss & Pilsbry. 
Texas : Lee Co., J. A. Singley ; Navidad River bottom, Jackson 
Co., J. D. Mitchell. 



/8 AMERICAN SPECIES OP VERTIGO. 

Vertigo rugosula STERKI, Proc. A. N. S. Phila. 1890, p. 34 ; 
Nautilus iv, p. 39, pi. 1, f. 3; reprinted by Binney, Fourth 
Supplement T. M. vol. v, in Bull. Mus. Comp. Zool. vol. 22, 
no. 4, 1892, p. 201, figure. WHEELER, Nautilus xxv, p. 124 
(Monte Sano, Madison Co., Ala.). 

Besides the smaller size, darker color, and characteristic 
striation, which is much more regular and stronger than in 
V. ovata, this species differs by the shape of the angular 
lamella, which is rather long, low in front, rising inwardly. 
It emerges about as far as the parietal lamella, which is strong 
and quite long. The infraparietal lamella is a very small 
tubercle, as a rule, sometimes scarcely or not perceptible. The 
inner end of the upper palatal fold turns downward. Further 
differences are given by Dr. Sterki: "The last whorl is rela- 
tively smaller, the aperture is somewhat less broad ; the crest 
and impressions over the palatal folds are less marked or 
wanting ; the palatal callus is thin or wanting ; the basal fold 
is situated higher on the columellar margin, in the position 
of a subcolumellar lamella, while in V. ovata it is basal. ' ' 

In profile view there is a slight prominence of the outer 
lip, at the termination of a shallow depression, but it is far 
less developed than the "auricle" of V. ovata. 

A topotype received from Dr. Sterki (fig. 3) measures, 
length 2, diam. 1.2 mm. ; barely 5 whorls. 

Dr. Sterki gave also the locality Fish Camp, Fresno Co., 
California, H. Hemphill; I have not seen the specimens but 
possibly they are what I am calling V. ovata mariposa, a 
shell which does not have the distinct striation of V. rugosula. 

la. Vertigo rugosula oralis Sterki. PI. 8, figs. 5, 6, 7. 

"It is somewhat smaller [than rugosula] , ovate, the striation 
and rugosity of the surface are less marked, and the inferior 
apertural [infraparietal] lamella is wanting; in turn it has in 
most examples a lamella at the base (between inferior colu- 
mellar and inferior palatal) and the callus in the palatal wall 
is rather strong. The coloration of part of them is somewhat 
lighter. It cannot be confounded with V. ovata Say, its 
relations to the type of rugosula being evident, and in addition, 



AMERICAN SPECIES OP VERTIGO. 79 

ovata has been found with it. Nor can it be referred to 
ventricosa it is larger and stronger, of much darker color, 
its surface is not so smooth and polished, it has 3 or even 4 
lamellae more, and the columella is longer" (Sterki, for var. 
ovulum) . 

Florida: Volusia Co., type loc., G. W. Webster, Pilsbry 
and others. Alabama: Stevenson; Tallapoosa R., about 7 
miles S. E. of Wetumpka. 

[Vertigo rugosula var.] ovulum STERKI, Proc. A. N. S. Phila. 
1890, p. 35. Not Pupa ovulum Pfr., 1841, also a Vertigo. 
Vertigo rugosula ovalis Sterki, PILSBRY & VANATTA, Proc. 
A. N. S. Phila. 1900, p. 608 (substitute for ovulum Sterki). 
V. r. oralis Sterki, PILSBRY, Nautilus xix, 40 (Cape Sable, 
Fla.) WHEELER, Naut. xxvi, p. 15 (region of Huntsville, 
Ala.) VANATTA, Naut. xxvi, 20, 33 (near Marco; Lee Co.; 
Seminole Pt., Monroe Co., Fla.). 

It is generally stouter in figure than V. rugosula, but the 
contour varies widely. Some shells are very globose. 
Length 1.7, diam. 1.1 mm., 4% whorls ) Tallapoosa R. 7 miles 
Length 2, diam. 1.2 mm., 5 whorls J s. E. of Wetumpka, Ala. 

Length 1.6, diam. 1.1 mm. "I __ , . ~ ., 

I Volusia Co., Fla. 
Length 1.9, diam. 1.15 mm. j 

The angular lamella is shorter than in V. rugosula, and 
it is not lower in front. The lip-point is more strongly de- 
veloped in some specimens from Alabama (fig. 5) than in 
most of those from Florida (figs. 6, 7) . There is often a notice- 
able light-colored crest behind the lip. In a specimen from 
"Mt. Taylor," Volusia Co., the suprapalatal and infrapalatal 
folds are strongly developed. 

Though this form was at first described as a variety of V. 
rugosula, Dr. Sterki regards it as nearer to V. ovata, and very 
close to the European V. antivertigo. The name was pub- 
lished in 1900 as V. r. ovalis, but this appears to have been an 
error for oralis, under which name it has appeared in several 
subsequent lists. 

2. VERTIGO ALABAMENSIS Clapp. PI. 8, figs. 9, 13, 14. 

The shell is cylindric-oval, perforate, convex, sutures well 



80 AMERICAN SPECIES OF VERTIGO. 

impressed, the last whorl somewhat tapering below, bulging 
above, and deeply constricted over the palatal folds ; crest well 
marked. Lip well reflected, pale brown, and deeply con- 
stricted opposite the upper palatal, simple above the con- 
striction, and with a strong callus or internal collar running 
down and connecting with the columella. Denticles 7, the 
parietal high, sinuous and deeply entering; angular strong, 
flat; upper palatal deep, very high in front and tapering to 
the rear, lower palatal very deeply seated, inner end back of 
the subcolumellar lamella, both palatals distinctly showing 
from the outside as white lines. Columellar lamella strong 
flat ; subcolumellar bifid, strong ; basal fold distinct, set on the 
callous collar just below the subcolumellar lamella. Length 
1.8, diam. 1.10 mm. ; length of aperture 0.69, width 0.63 mm. 
(Clapp). 5% whorls. 

Alabama : among rotting leaves in a ravine near junction of 
North River with Black Warrior, Tuscaloosa Co., H. H. Smith. 

Vertigo alabamensis CLAPP, Nautilus xxviii, April, 1915, 
p. 137, pi. 6, f. 6, 6a, 6fe. 

This is a peculiar species. The parietal lamella is very 
strongly developed, high and long. The angular lamella is 
far longer than usual in Vertigo, and converges inwardly 
towards the parietal. The columellar lamella descends near 
its outer end, and penetrates deeply. _The "subcolumellar" 
descends obliquely inward, its crest either emarginate or level 
(pi. 8, fig. 9). It is apparently to be considered a basal fold, 
since the palatal callus extends to it. If this interpretation 
is correct, the tooth described as the basal must be an infra- 
palatal fold. It is usually a very low tubercle, and very 
rarely "distinct," as described and figured by Dr. Clapp. 
The palatal folds are very strongly developed, the upper a 
high lamina, the lower rising to a peak deep within. 

The types of this and the following form are in coll. G. H. 
Clapp ; figured from paratypes in coll. A. N. S. P. 

2a. Vertigo alabamensis conecuhensis n. subsp. PL 8, fig. 12. 

At Evergreen, Conecuh Co., Ala., Mr. Smith found a variety 
which differs from the type in being shorter and more globose 



AMERICAN SPECIES OF VERTIGO. 81 

with thii teeth less strongly developed and the basal fold 
absent in all shells examined. Length 1.53, diam. 1 mm. 
Length of aperture 0.58, width .63 mm. (Clapp). 

Vertigo alabametisis co-necuhensis CLAPP, Nautilus xxviii, 
April, 1915, p. 137, pi. 6, f. 7, la. 

The size varies from 1.6 x 1.05 mm. to 1.5 x 1 mm. 

3. VERTIGO MORSEI Sterki. PL 6, figs. 8, 9. 

"Shell large (for the group), cylindrical-turriculate, with 
a rather acute apex, imperforate rimate, with few obsolete 
striae of growth, shining, translucent. Whorls six, rather 
slowly and regularly increasing, the last scarcely higher than 
the penultimate and rather narrower, somewhat sloping to- 
wards the base, slightly ascending at the aperture; suture 
deep. Aperture lateral, scarcely oblique, comparatively small, 
inferior and palatal part well-rounded, the latter with an angu- 
lar impression and slightly protracted in about its middle, 
the upper half more strongly curved, peristome everted ; on 
the palatal wall, at some distance from and parallel with the 
margin [externally] a moderate crest, behind it a deep and 
large impression over the palatal folds, and in front of it 
a groove corresponding with the impression at the auricle; 
inside the crest there is a distinct callus of the same color as 
the shell; apertural lamellae and folds typically nine: three 
on the parietal wall (the same as in V. ovata), the largest 
whitish; two on the columella, the superior strong, vertical 
above, then in an angle turning horizontally, the inferior 
horizontal, lamelliform, thin, high and directed obliquely up- 
ward ; basal small, sometimes double, rarely ; palatals high, 
and rather long, curved and directed upward; suprapalatal 
small, nodule-like. 

"Alt. 2.7, diam. 1.3; apert, alt. 0.9, lat. 0.8 mm." (Sterki). 

New Jersey : White Pond, Warren Co., Pilsbry. Michigan : 
Kent Co., type loc., Dr. De Camp. Ohio: Sandusky; fossil 
in marl, Castalia, Erie Co. Indiana : margins of Lakes James, 
Tippecanoe and Maxinkuckee under dead weeds, wood etc. 
L. E. Daniels. Illinois: Joliet, J. H. Ferriss. 

Vertigo morsei STERKI, Nautilus viii, Dec., 1894, p. 89. 



82 AMERICAN SPECIES OP VERTIGO. 

BLATCHLEY and DANIELS, 27th Ann. Rep. Dep. Geol. and Nat. 
Res. Indiana, 1902, p. 587, 632 (Kosciusko, Marshall and 
Steuben counties, Ind. WALKER, Moll, of Michigan, 1906, 
p. 516, f. 149 (Dewey's Mill, near Grand Rapids, type loc. ; 
Eaton, Bay and Washtenaw counties, Mich.). 

"In one example the inferior parietal tooth is wanting, in 
another the basal only trace-like, and in a third the same is 
double, as frequently found in V. ovata. With the latter 
species, V. morsei has much resemblance, especially in the 
aperture: the configuration and the 'teeth' are the same, but 
the inferior columellar, and the two principal parietals, are 
rather larger, and markedly directed upward. The main dif- 
ference is in the number and relative size of the whorls : while 
in V. ovata they are five, and rapidly increasing, the last pre- 
dominating, in our species there are six, slowly increasing, 
the last, and consequently the aperture, comparatively small. 
This is a radical difference, and gives the shell quite another 
aspect, so that there can be no question about its being dis- 
tinct. But V. morsei is also decidedly larger, V. ovata not, 
or little, exceeding 2 mm. in altitude" (Sterki). 

The species is named in honor of Prof. E. S. Morse. 

Specimens determined by Dr. Sterki measure : 

Length 3, diam. 1.5 mm., 6y 2 whorls. 

Length 2.6, diam. 1.3 mm., 6 whorls. 

The color is from auburn to chestnut-brown. There is a 
distinct crease behind the lip, from the crest to the lip- 
point, and over both palatal plicae there is a wide depression 
externally. Dr. Sterki writes that out of 100 specimens ex- 
amined from the marl of Erie Co., Ohio, 10 have an infra- 
parietal, quite small or vestigial. 

While it stands very close to V. ovata, this species does not 
appear to intergrade, so far as seen. It is the largest Vertigo 
known. 

4. VERTIGO OVATA Say. PL 6, figs. 1 to 4, 7. 

"Shell dextral, subovate, brown; apex obtuse; whorls five, 
glabrous; suture not very deeply impressed, body whorl in- 
dented near and upon the labrum ; aperture semioval ; labrum 



AMERICAN SPECIES OF VERTIGO. 83 

five-toothed, of which three are situate on the transverse por- 
tion of the lip, parallel to each other, equidistant, the superior 
and inferior ones being small, the latter sometimes obsolete, 
the intermediate one lamelliform, prominent, and the two 
others situate on the columella, approximate, extending at 
right angles to the three preceding ones, the superior one 
oblique and smaller; labrum reflected but not flattened, bi- 
dentate, teeth lamelliform, prominent; umbilicus distinct. 
Length less than one-tenth of an inch. Breadth nearly 1/20 
inch" (Say). 

Length 2.2, diam. 1.4 mm. ; 5 whorls (Dutchess Co., N. Y.). 

Length 2.3, diam. 1.4 mm. (Ottawa, Canada). 

Length 2.15-2.35, diam. 1.35 mm. (Oswego, Ore.). 

Labrador and Alaska, south to Alabama, Texas and 
Arizona ; State of Vera Cruz, Mexico and West Indies. Type 
loc. Philadelphia. 

Vertigo ovata SAY, Journ. A. N. S. Phila. ii, 1822, p. 375. 
BINNEY, Man. Amer. Land Shells, 1885, p. 334, f. 362, 363. 
MORSE, Amer. Nat. i, 1868, p. 668, f. 67, 68. TAYLOR, Nautilus 
v, 91 (Vancouver I.). WEBSTER, Naut. v, 119 (Florida). 
STUPAKOFF, Naut. vii, 135 (Allegheny Co., Pa.). PLEAS, 
Naut. vii, 68 (Henry Co., Ind.). SQUYER, Naut. viii, 63 
(Mingusville, Mont.). PRIME, Naut. viii, 70 (Long Island, 
N. Y.). STERKI Proc. U. S. N. M. xi, 1888, p. 375, pi. 42, f. 
5-7; Naut. viii, 89 (Kent Co., Mich.) ; ix, 116 (San Marcial, 
N. M.); xxix, 123 (Geneva, O.). SCHICK, Naut. viii, 137 
(Philadelphia). SARGENT, Naut. ix, 89 (Clearwater, Minn.). 
HANHAM, Naut. x, 101 (Quebec); xiii, 3 (Winnepeg). 
PILS. & CKLL., Naut. xiv, 86 (Las Vegas, N. M.). COCKERELL, 
Naut. x, 41-43 (Mesilla, near Silver City and Eincon, N. M.) 
Chadwick, Naut. xix, 58 (near Milwaukee, Wis.). PILSBRY, 
Naut. xix, 130 (Grant, N. M.) ; xxv, 75 (Monroe Co., Pa.). 
WALKER, Naut. xx, 81 (Monroe canyon, Neb.) ; Moll, of 
Michigan, 1906, p. 516, fig. 148 (entire state). SMITH, Naut. 
xx, 90 (Otsego Co., N. Y.). WHEAT, Naut. xx, 101 (Cayuga 
Lake, N. Y.). HANNA, Naut. xxiii, 95 (Douglas Co., Kansas). 
NYLANDER, Naut. xxii, 143 (Aroostook Co., Me.). HENDER- 
SON, Naut. xxii, 9 (Amarillo, Texas). PILSBRY & 



84 AMERICAN SPECIES OP VERTIGO. 

Naut. xxii, 104 (Albuquerque, N. M.). BERRY, Naut. xxiv, 
63 (Unity, Me.) ; xxix, 125 (Winnecook, Mont. ) .WHEELER, 
Naut. xxv, 124 (Monte Sano, Madison Co., Ala.). GREEGER, 
Naut. xxix, 89 (Payne Co., Okla. ) . OVER, Naut. xxix, 91 
(Deuel and Clay Co., S. Dak.). DANIELS, 27th Ann. Rep. 
Dep. Geol. and Nat. Res. Indiana, 1902, p. 632 (Kosciusko 
and Steuben counties, Dunreith, Arlington, Lawrenceburg 
and Indianapolis, Ind.). J. HENDERSON, Univ. of Colo. 
Studies iv, p. 171, fig. 9 (Twin Lakes and Saguache, West 
Cliff, Trinidad, and V. o. antiquorum CklL, Grape Creek, 
Colo.). JOHNSON, Fauna of New England 13, 1915, p. 215 
(all New England states). DALL, Land and Fresh Water 
Mollusks, Harriman Alaska Exped. xiii, 1905, p. 32 ("Ungava 
Bay, Labrador! Victoria, British Columbia! St. Paul, Kadiak 
I. ! Alaska ; Tigalda Island, Aleutian chain ! Laggan, Alberta, 
Manitoba"). 

Pupa ovata Say, GOULD, Boston Journ. Nat. Hist, iv, 1843, 
p. 350, pi. 16, f. 7, 8. PFR., Monogra. ii, p. 360. v. MARTENS, 
Biol. Centrali Amer., Mollusca, p. 327. Isthmia ovata Say, 
MORSE, Journ. Portland Soc. N. H., i, 1864, p. 38, f. 93; pi. 
10, f. 94. 

Zonites upsom CALKINS, Valley Naturalist ii, Dec., 1880, p. 
53, fig. . Cf. W. G. BINNEY, Suppl. to Terr. Moll. V, Bull. 
M. C. Z. xi, no. 8, Dec., 1883, p. 149, pi. 1, f. L (immature 
stage). 

Pupa ovata forma nov. antiquorum COCKERELL, Zoe ii, 
April, 1891, p. 18. 

In the collection of the Academy V. ovata is present from 
Prince Edwards Island, Quebec and Ontario, through all the 
eastern states south to the Potomac, and west of the Alle- 
ghanies south to Alabama and Galveston, Texas; through all 
of the northern states west to Montana and Colorado ; also in 
the Rio Grande Valley, south to Mesilla, New Mexico; in 
southern Arizona near the Mexican boundary in the Huachuca 
Mts., and westward to Tempe and Jerome. It is, however, 
decidedly local in Arizona and New Mexico, and except in 
the Huachucas, the specimens are from stream debris, per- 
haps always washed from greater elevations than the places 



AMERICAN SPECIES OF VERTIGO. 85 

mentioned. In the Pacific states, there are specimens from 
Seattle, Washington (P. B. Randolph) and Oswego, Clackamas 
Co., northern Oregon (J. A. Allen). I have not seen it from 
Idaho, Utah, Nevada or California, but allied or subspecific 
forms occur in California. Specimens are also wanting in 
this collection from the southern Atlantic states, from Virginia 
to Florida. Antillean specimens are noted below. 

The Alaskan records are all from Dall. It appears to ex- 
tend north to about 58, south to about 18. If the specimens 
from these extremes are reliably determined, V. ovata has 
the greatest range in latitude and climate of any Vertigo or 
other Pupillid snail in the world, so far as I know. 

' ' V. antivertigo Drap. and V. ovata Say have almost exactly 
the same dentition as to number, size, and shape, and places 
of the single lamellae, and also as to other features of the 
shells they are very similar. V. ovata, in general, is a little 
larger, of lighter coloration, the margin is somewhat more ex- 
panded; the aperture appears larger because the columellar 
margin is relatively a trifle shorter; the whorls increase in 
size somewhat more rapidly, and the suture is a trifle deeper. 
Specimens from Massachusetts are not only of the same or 
even a darker shade [than V. antivertigo], but also of the 
same and sometimes of a smaller size, and other distinguish- 
ing features are slightly marked. They resemble each other 
so much that it is difficult to separate them, and if collected 
at the same place no one would regard them as even distinct 
varieties. I hesitate, however, to declare them identical be- 
cause I have not yet made a comparison of the soft parts. 
Doubtless they are parallel forms, and if found on the same 
continent would be regarded as varieties of one species. 
V. antivertigo is remarkably constant throughout Europe so 
far as I know it, while V. ovata is on the contrary quite vari- 
able, owing, possibly, to the greater differences in climate. 
Also a few decided varieties exist and maybe more will be 
found ; the future study of these forms is likely to be of great 
interest" (Sterki). 

I have not seen var. antiquorum CklL, but no differential 
characters of importance appear in the description, which 
follows. 



86 AMERICAN SPECIES OF VERTIGO. 

"Pupa, ovata forma nov. antiquorum. 2 mm. long; lamellae 
7; three on parietal wall, the central one large and slender, 
the others very small: two on columella, these approximately 
of equal size, and both rather slender; two on external wall, 
continued backwards, the upper one large and curved down- 
wards about its middle. Whorls 4%, body-whorl inflated, the 
others diminishing regularly and rather rapidly towards the 
apex, outer lip conspicuously curved inwards opposite the 
upper tooth on external wall. Posttertiary deposit at West 
Cliff, Colorado. This form also occurs living, as Dr. Sterki, 
to whom I sent a specimen says it 'is of a form of which I 
have seen examples from many parts of the country, of the 
same size, shape and formation of lamellae.' " (CHI.). 

Translation of the original description of Pupa ovulum 
follows : 

"Pupa ovulum Pfr. Shell minute, dextral, slightly rimate 
at base, apex obtuse, shining, brown ; whorls 5, a little convex, 
narrow, the last one inflated and impressed in the middle; 
aperture 6-toothed: two equal teeth on the columella, two in 
the right margin and two in the left. Length 1, diam. y 2 lin. 
Vera Cruz (Hegewisch). Very like Vertigo pusilla Mull., 
but dextral, brown, the whorls less convex" (Pupa ovulum 
PFEIFFER, Symbolae ad Hist. Hel. i, 1841, p. 46). 

P. ovulum was placed in the synonymy of Pupa ovata Say 
in the Monographic ii, 1848, p. 361 ; Pfeiffer there referring 
the figures in Kiister (pi. 14, figs. 1, 2) to ovulum, so it may 
be inferred that they were drawn from Vera Cruz specimens ; 
though in the text of Kiister they are called Pupa ovata Say, 
and a different description is given. The figure is so poorly 
drawn that it might represent any form of the ovata group. 

Subsequent collectors do not appear to have found the 
Vera Cruz form. Its identity with V. ovata remains to be 
verified. There are no recent records of V. ovata from Mexico. 

Antillean forms provisionally referred to V. ovata. 

Forms which appear referable to this species are before me 
from Porto Eico, Santo Domingo and Cuba ; and the species 
described as Vertigo neglect a and Pupa hexodon, from Cuba 






AMERICAN SPECIES OP VERTIGO. 87 

and Jamaica, have not been satisfactorily differentiated. Un- 
til sufficient material is brought together for definite con- 
clusions the whole may be referred provisionally to V. ovata. 

A bleached Vertigo from Santo Domingo, collected by W. M. 
Gabb, measures 1.9 x 1.2 mm. There is a suprapalatal tubercle 
but no other accessory teeth. The surface is distinctly, finely 
but not sharply striate, but otherwise agrees with ovata 
(pi. 13, fig. 13). 

Three specimens from Humacao, Porto Rico (pi. 13, fig. 16), 
similar to V. ovata in color and surface, are small, length 1.7, 
diam. 1.15 mm., with 4% whorls. There are weak infra- 
palatal nodules in two of them, all having a suprapalatal. 
None has an infraparietal lamella. Possibly this and the pre- 
ceding forms are separable as subspecies, but the material seen 
is insufficient for a decision. 

The J. B. Henderson collection contains a Cuban specimen 
of V. ovata, received from Poey through Gill, 2.1 x 1.36 mm., 
having a minute suprapalatal but no other accessory teeth. It 
does not differ from many northern examples. 

Pupa hexodon is not known to me by Jamaican specimens. 
In the description, repeated below, there is nothing to dif- 
ferentiate it from six-toothed individuals of V. ovata. 

Pupa hexodon (C. B. Adams). " Shell ovate; brown; 
smooth and shining; spire with convex outlines; whorls 5, 
rather convex, with a well impressed suture; aperture semi- 
oval, with the transverse lip rather oblique, constantly with 
six teeth, of which two are on the transverse lip, the inner one 
being a little larger; two teeth are on the columella, the 
upper one being a little larger, and on the outer lip are two, 
of which the lower one is a little larger ; umbilicus very small. 
This species resembles P. ovata Say. Length .085 inch, 
breadth .055 inch" [about 2.12x1.4 mm.] (Adams). 

Jamaica, C. B. Adams. 

Pupa hexodon C. B. ADAMS, Contributions to Conchology, 
no. 3, p. 37. SHUTTLEWORTH, Diagn. n. Moll., no. 6, p. 145. 

Vertigo neglect a Poey (pi. 13, fig. 11). Shell dextral, 
rimate-perforate, ovate, very delicately striate, thin, pale 
brown; spire conic, obtuse; whorls 4%, convex, the last ex- 



88 AMERICAN SPECIES OP VERTIGO. 

ceeding half the total height of the shell. Aperture sub- 
rotund, coarctate, 4-toothed : two very minute marginal teeth 
on the columella; one palatal, short; one strong, marginal in 
the process of the right margin. Peristome simple, expanded, 
the margins seperated by the lack of a callus; no apertural 
tooth. Length 1%, diam. 1 mm. Aperture small (Poey). 

Cuba: Cardenas, on the sandy shore (R. Arango). 

Vertigo neglecta Ar. mss., POEY, Memorias sobre la Historia 
Natural de la isla de Cuba, ii, 1856, p. 30, pi. 2, f. 17, 18. 
PFR., Monographia v, p. 328. 

This has been surmised to be a V. ovata without the parietal 
teeth; and as the type was taken on the shore, it may very 
likely have been a dead shell in which the parietal callus 
bearing the teeth had scaled off, not an unusual condition 
in drifted shells. 

4a. Vertigo ovata diaboli n. subsp. PL 6, figs. 11, 12. 

All of the lamellae and plicae are larger than in V. ovata, 
entering further; infraparietal lamella developed. Palatal 
plicae, especially the upper, which converges towards the 
lower, very strong and long. The entering point of the 
outer lip is well developed, and the crest behind the lip moder- 
ately strong, but lower than in typical ovata. 

Length 2.2, diam. 1.25 mm. ; 5 whorls (type). 

Length 2.3, diam. 1.3 mm. 

Texas : drift debris of Devil 's River, about 4 miles from the 
Bio Grande, Val Verde Co. ; collected by Ferriss and Pilsbry, 
1903. No. 90437 A. N. S. P. 

The specimens are all "dead" shells, but appear to have 
been lighter-colored than typical V. ovata. The point of the 
outer lip is seen to project well forward, in a profile view. 
The impressions behind the lip are about as in V. ovata. 
Perhaps a distinct species, but as it is known by a single gath- 
ering, it is left under V. ovata for the present. 

4&. Vertigo ovata mariposa n. subsp. PL 6, figs. 5, 6. 

The shell has the usual broadly ovate shape and auburn or 
darker color. The teeth are about as in typical V. ovata. 



AMERICAN SPECIES OP VERTIGO. 89 

Lower palatal is longer and enters more deeply than the upper. 
The basal is subcolumellar in position. The outer lip bends 
inward only very slightly ; the crest and the external im- 
pression over the palatal plicae are but weakly developed. 
Length 2.2, diam. 1.35 mm.; length of aperture 0.85 mm.; 
4% whorls. 

California: Mariposa Co., type no. 11644 A. N. S. P.; 
meadow near Wawona, Sequoia Park, in the same county, H. 
N. Lowe, 1916. 

Easily recognized by the shape of the outer lip. So far as 
known, typical V. ovata does not occur in California. 

Mr. Lowe's specimens are a trifle smaller, 2.1 mm. long, 
and the color is very dark. 

5. VERTIGO BERRYI n. sp. PL 6, figs. 10, 13. 

The shell is oblong-conic, auburn, glossy, slightly irre- 
gularly striate. The spire tapers from the last whorl, the 
lateral outlines being slightly convex; summit is very obtuse, 
of a paler tint. The whorls are rather strongly convex; the 
last having an inconspicuous narrow, low ridge close behind 
the lip expansion, preceded by a broad and deep impression 
over the palatal region, above which it remains strongly con- 
vex. Except for its smaller size, the aperture is about as in 
V. ovata. There is a small angular lamella near the large 
parietal ; columellar lamella rather massive, ascending inward. 
Palatal plicae subequal, rather long, the basal fold smaller; 
suprapalatal quite small. The teeth and palatal callus are 
much lighter than the lip. The peristome is expanded ; outer 
border is slightly curved in, but far less than in V. ovata. 

Length 2.5, diam. to lip edge 1.35 mm. ; length of aperture 
0.9 mm. ; 5% whorls. 

California : Mill Creek Canyon, at 4600 ft., in the San Bern- 
ardino Mountains; collected by Dr. S. S. Berry, July, 1910. 
Type 105166 A. N. S. P. Also Valle Trinidad, Lower Cali- 
fornia, C. R. Orcutt, 1901 ; no. 308964 U. S. N. M. 

This species differs from V. ovata by the relatively narrow, 
almost straightly tapering spire, the much weaker inbending 
of the outer lip, the minute crest outside the peristome, and 



90 AMERICAN SPECIES OF VERTIGO. 

the capacious external excavation over both palatal plicae. 
Vertigo morsel is similar to V. berry i in having a long spire 
relative to the aperture, but the Eastern species is much larger, 
has more convex lateral outlines, a much more developed point 
on the outer lip and a stronger crest behind the lip. 

It was reported as Vertigo ovata (Say) var., in Nautilus 
xxx, p. 38. 

6. VERTIGO BINNEYANA Sterki. PL 11, fig. 12. 

The shell is cylindric-oblong, auburn, somewhat transparent, 
glossy, weakly, irregularly striate. The whorls are moderately 
convex, the last having a low but distinct crest behind the 
lip, a small impression between the crest and the point of the 
lip, and a rather large impression behind the crest, over the 
palatal folds. The aperture is rather small. The teeth are 
whitish; parietal lamella strong but rather short; a quite 
short angular lamella stands even with its outer end. 
Columellar lamella strong, but not long, ascending a little in- 
wardly. The palatal folds are both quite strong, the lower 
entering much further. Basal fold short. The peristome is 
somewhat expanded, the outer lip a little bent inward above 
the middle. There is a moderate palatal callus. Length 
2.1, diam. 1.1, length of aperture 0.75 mm. ; 5 whorls. 

Montana: Helena, type loc., H. Hemphill; Glendive, Dr. 
Sterki; drift debris of Musselshell River, Winnecook, S. S. 
Berry. Manitoba: Winnipeg, Sterki. Nanaimo, Vancouver 
Island, G. W. Taylor. ( ? New Mexico : Albuquerque, Dr. 
Sterki.) 

Vertigo Unneyana STERKI, Proc. A. N. S. Phila. 1890, p. 
33; Nautilus iii, 1890, p. 125 (Helena and Glendive, Mont.; 
Winnipeg, Man., Albuquerque, N. M.) ; iv, p. 39, pi. 1, f. 1. 
BINNEY, Fourth Suppl., Bull. M. C. Zool. xxii, 1892, p. 
198, fig. PILSBRY, Proc. A. N. S. Phila. 1899, p. 315, fig. 2 
(specimen from Winnepeg). RANDOLPH, Naut. ix, 102 
(Seattle, Wash.). SQUYER, Naut. viii, 63 (Mingusville, 
Mont.). BERRY, Naut. xxix, 125 (Winnecook, Mont.). 

It is smaller and more cylindric than any form of V. ovata, 
has a longer lower palatal fold, and a less impressed point in 



AMERICAN SPECIES OP VERTIGO. 91 

the outer lip. V. pygmaa is wider, has a much stronger crest, 
and the angular lamella is small or wanting. 

Dr. Sterki informs me that it has been found in drift debris 
of the Missouri River, Iowa. 

Group of Vertigo numellata. 

The aperture has the typical six teeth well developed, the 
lower palatal fold long, entering to the dorsal side, columellar 
lamella receding, ascending somewhat inwardly. There is a 
very high, rounded crest. 

The single species inhabits Bermuda. 

7. VERTIGO NUMELLATA Gulick. PL 13, figs. 14, 15. 

The shell is shortly rimate, oval, auburn, the surface dis- 
tinctly but weakly striate, glossy. Whorls weakly convex, the 
last having a very high, rounded crest, then deeply contracted 
behind the peristome ; behind the crest it is flattened, and nar- 
rowly furrowed over the palatal folds, one or both of the 
furrows usually extending upon the crest. Aperture is ob- 
structed by 6 teeth: the angular lamella is rather long and 
stands remote from the parietal. Parietal lamella strong and 
entering deeply. The columellar lamella is deeply placed, 
rather massive, and ascends somewhat inwardly. The upper 
palatal fold is strong and rather long. Lower palatal is some- 
what more immersed and longer, entering to a dorsal position. 
The basal fold is low and small, sometimes subobsolete. The 
palatal callus is not conspicuous. The peristome is thin, well 
expanded, prominently projecting forward and bent inward 
above the middle of the outer lip, thinner and retracted above 
the prominence. 

Length 1.8, diam. 1.05 mm. ; 5 whorls (type). 

Length 1.8, diam. 1 mm. ; 5y 2 whorls. 

Length 1.6 mm. ; 4*/2 whorls. 

Bermuda: Paynter's Vale, between Tucker's Town and 
Walsingham Bay, pleistocene and recent; also Knapton Hill, 
fossil; near Bailey Bay in the red clay breccia near Castle 
Harbor. Type no. 85583 A. N. S. P. 

Vertigo numellata GULICK, Proc. A. N. S. Phila. 1904, p. 



92 AMERICAN SPECIES OP VERTIGO. 

413, pi. 36, f. 6. VERRILL, Trans. Conn. Acad. xii, 1907, p. 
170, f. 54c. 

While highly specialized and peculiar, this species appears 
to be somewhat related to V. oralis and especially V. 
aldbamensis. It has also some resemblance but probably no 
direct relationship to V. ovata. It differs from all American 
species by the far more massive crest. There is a long lower 
palatal fold, but it is not turned down at the end as in 
V. milium, and the shape of the columellar lamella shows 
that V. numellata is not an Angustula-. 

This species was described from fossil examples in the lime 
rock, but it is abundant as a recent shell in Paynter's Vale, 
about Church Cave, near Tucker's Town. Except in color, 
there is no difference between recent and fossil examples. 

Group of Vertigo pygmaea. 

It is scarcely possible to define this group, some species 
approaching the group of V. ovata closely, while others are 
hardly distinguishable from the modest a group. As a whole 
the group is equivalent to the European group of V. 
moulinsiana and pygmaea. 

Key to species. 
(Eastern and central species). 

1. Aperture having 3 teeth, parietal, columellar and lower 
palatal ; sometimes a minute upper palatal. 2. 
Aperture having at least 4 well developed teeth. 5. 

2. 1.8 to 2.2 mm. long; upper palatal fold generally present 
but minute. 3. 
1.5 to 1.6 mm. long ; upper palatal fold rarely developed. 4. 

3. Palatal callus slight or wanting ; Maine to Colorado. 

V. tridentata, no. 16. 
Palatal callus strong ; Bermuda. V. marki, no. 17. 

4. Subcylindric, yellowish, the lower palatal fold penetrat- 
ing to the dorsal side ; Ohio, N. C. V. parvula, no. 15. 
Ovate, dark olive buff, fragile, the lamella and folds small 
and short. New England. V. perryi, no. 14. 

5. Lower palatal fold long, very deeply immersed, an im- 



AMERICAN SPECIES OF VERTIGO. 93 

pression on the back over it ; 6 teeth ; 1.7 x 0.95 mm. 
Maine. V. nylanderi, no. 11. 

Lower palatal fold not notably immersed. 6. 

6. Shell sharply but very finely striate, especially the penult 
whorl; no palatal callus; 1.85 to 1.95 x 1 mm.; 5 to 6 
teeth. V. gouldii, no. 10. 
Shell nearly smooth, or but weakly striate. 7. 

7. Crest and palatal callus moderately or strongly de- 
veloped. 8. 
Crest very low when present; palatal callus very weak 
or wanting. 11. 

8. Shell cylindric-oval, 1.8 to 2 mm. long ; no angular lamella 
and the basal fold small or wanting. Maine to Va., 
west to Lake Superior. V. pygm&a, no. 9. 
Ovate, 1.8 to 2.2 mm. long. 9. 

9. No angular lamella or basal fold; upper palatal fold 
small or minute. 10. 
A basal fold, and often an angular also; ovate-conic; 
upper palatal fold well developed. Northern states. 

V. v. elatior, no. 8a 

10. Palatal callus moderate or thin ; Maine to Colo. 

V. trident at a, no. 16. 
Palatal callus heavy ; Bermuda. V. marki, no. 17. 

11. Angular, parietal and columellar lamellae, upper and 
lower palatal folds ; no crest ; 1.25 x 0.84 mm. Florida 
Keys. V. hebardi, no. 13. 
No angular lamella, 4-5 teeth, a basal fold present or 
wanting; Northern States and Canada. 12. 

12. 1.75 x 1.1 mm. V. ventricosa, no. 8. 
1.5 x 0.9 mm. V. bollesiana, no. 12. 

(West Coast species). 

1. With 4 or more well developed teeth. 2. 
With 2 or 3 weak teeth; no trace of crest or palatal 
callus; 2 x 1.23 mm. ; California. V. sterkii, no. 20. 

2. 4 teeth; cylindric-oval; Oregon to Alaska. 

V. columbiana, no. 18. 
5 to 6 teeth. 3. 



94 AMERICAN SPECIES OP VERTIGO. 

3. Ovate-conic, with a strong palatal callus; British 
Columbia. V. v. elatior, no. Sa. 

Cylindric-oblong ; palatal callus very weak or wanting; 
Washington to California. V. andrusiana, no. 19. 

(Eastern species, Atlantic to Rocky Mountains) . 
8. VERTIGO VENTRICOSA (Morse). PI. 7, figs. 1, 2, 3. 

"Shell umbilicate, ovate, conic, smooth, polished; apex ob- 
tuse; suture deep; whorls four, convex. Aperture, semi- 
circular, with five teeth, one prominent on the parietal margin, 
two smaller on the columellar margin, and two prominent 
within, contracting the aperture at the base ; peristome widely 
reflected, the right margin flexuose, within thickened and 
colored. Length .07 inch; breadth .45 inch" [1.75 x 1.1 mm.] 
(Morse). 

Magdalen and Prince Edward Is., Quebec, New England 
and New York, west to Ohio, Michigan and Illinois. 

Isthmia ventricosa MORSE, Ann. Lye. Nat. Hist. N. Y. viii, 
Nov., 1865, p. 207, fig. 1. Vertigo ventricosa Morse, BINNEY, 
Terr. Moll, v, 1878, p. 218. STERKI, 8th Ann. Rep. Ohio 
State Acad. Sci., 1900, p. 32 (Tuscarawas Co., 0.). 
WHITEAVES, Ottawa Naturalist 1905, 171 (Riviere du Loup, 
Quebec). NYLANDER, Nautilus xiii, 103 (Aroostook Co., Me.). 
HANHAM, Naut. x, 101; xi, 111 (Isle d 'Orleans, Quebec). 
WHEAT, Naut. xx, 161 (Cayuga L. } N. Y.). BLANEY, Naut. 
xviii, 46 (Ironbound I., Me.). WALKER, Moll, of Michigan, 
1906, p. 517, f. 150 (Grand Rapids and Beulah, Benzie Co.). 
JOHNSON, Fauna of New England no. 13, 1915, p. 214 (Me., 
N. H., Conn.). V.[ertigo] approximans STERKI, Nautilus 
iii, 1890, p. 136. 

The author has seen typical ventricosa only from Canada, 
New England and New York, but it has been reported by 
Walker and Sterki from states bordering the Great Lakes as 
far west as Illinois. 

V. ventricosa differs from V. ovata by the constantly 
smaller size, absence of an angular lamella, and smaller num- 
ber of teeth. It is of an auburn color, somewhat transparent, 
glossy, with only a trace of striation. Under a high power 



AMERICAN SPECIES OP VERTIGO. 95 

it shows microscopic punctation or granulation. The basal 
fold is usually quite small, though sometimes, as in the speci- 
men from Prince Edward Island figured (fig. 1), it is well 
developed. In many Maine examples it is absent (fig. 3, Buck- 
field, Oxford Co., Me.). The degree of prominence of the in- 
wardly bent point of the lip-edge varies a good deal. The 
crest is quite low. Specimens measure: 

Length 1.7, diam. 1 mm.; 4% whorls (Hebron, Me., fig. 2). 

Length 1.95, diam. 1.2 mm. ; 4% whorls (Buckfield, Me., 
fig. 3). 

A form which Dr. Sterki listed as V. approximans was 
originally stated to be "characterized by the two palatal 
lamellae being close together." It was from Rockford, 111. 
The original specimens are lost, but what appears to be the 
same form has been found in Ohio. " It is rather small, length 
1.3, diam. 0.9 mm., more or less; short, slight, with no callus 
in the palate or a slight one. There are parietal and colu- 
mellar lamellae and two palatal folds, all small, the palatals 
close" (Sterki). 

Dr. 0. Boettger considered ventricosa a synonym of V. 
moulinsiana of Europe (Jahrb. Nassauischen Vereins, 1889, 
p. 307) ; but the resemblance does not appear sufficiently close 
for specific identity. 

8a. Vertigo ventricosa elatior Sterki. PL 7, fig. 6. 

"Larger and more elevated than ventricosa, with a rather 
acute apex; a strong callus in the palate, into which the 
palatal plicae merge, a strong tooth-like lamella in the base" 
(Sterki). 

Length 2.15, diam. 1.2 mm. ; 5 whorls. 

New York, Ohio, Michigan and west to Montana (Sterki) ; 
Aroostook and Hancock counties, Maine ; Darby and White 's 
Springs, west of Ward, Mont.; Field, B. C. and Laggan, 
Alberta ; Oscuro Mts., Socorro Co., N. M. 

Vertigo ventricosa var. elatior STERKI, The Land and 
Fresh Water Mollusca in the vicinity of New Philadelphia; a 
contribution to the Natural History of Tuscarawas Co., Ohio, 
p. 5, 1894. Eighth Ann. Rep. Ohio State Acad. Sci., 1900, p. 



96 AMERICAN SPECIES OF VERTIGO. 

33; Nautilus viii, 107; xxix, 123 (Geneva, Ohio). SARGENT, 
Naut. ix, 89 (Clearwater, Minn.). NYLANDER, Naut. xiii, 103 
(Aroostook Co., Me.). HENDERSON, Naut. xx, 97 (Cazenovia, 
N. Y.). WALKER, Moll. Michigan, 1906, p. 517 (throughout 
the Lower Peninsula) . J. HENDERSON, Univ. of Colo. Studies 
iv, 172 (Lake George, Colo.). Vertigo gouldi lagganensis 
PILSBRY, Proc. A. N. S. Phila, 1899, p. 314, fig. 1 (Laggan, 
Alberta). 

Dr. Sterki reports it from the loess at New Harmony, 
Indiana, and from marl deposits, Castalia, Erie Co., Ohio. 

The shape, more conic than V. ventricosa, the strong palatal 
callus and teeth, the well developed basal fold and the larger 
size give this race individuality. It has an extensive range 
west of that of V. ventricosa, though also occurring in territory 
of the latter in the northeast, as far as northern Maine. 

Compared with V. pygmaa, V. g. elatior is more conic, the 
outer lip has a more distinct point or "auricle," the crest is 
less massive, usually weak, and there is a deeper external 
impression over the lower palatal fold. 

Dr. Sterki notes that there is often a supra-palatal fold 
developed, and occasionally an angular lamella. 

It would not be amiss to rank elatior as a species. 

9. VERTIGO PYGMAEA (Drap.). PL 7, figs. 11, 12. 

The shell is cylindric-oval, auburn or chestnut-brown, glossy, 
having only weak traces of striation, but the surface appears 
densely weakly pitted microscopically. The whorls are moder- 
ately convex, the last having a strong rounded crest a short 
distance behind the peristome, separated from it by a con- 
cavity, and somewhat paler colored than the rest of the shell. 
The parietal lamella is strong but rather short, median. 
Columellar lamella deeply placed, short, ascending inwardly. 
Both palatal folds are strong, the lower one longer, as usual. 
They stand on a strong callus. The basal fold is very small, 
rarely absent. There is often a low suprapalatal fold. The 
peristome is well expanded, somewhat reflected, colored like 
the shell. The outer lip is only slightly incurved. 

Length 2, diam. 1.1 mm. ; 5 whorls. 



AMERICAN SPECIES OP VERTIGO. 97 

Length 1.8, diam. 1 mm. 

Maine : Eockland, Thomaston and Warren, Knox Co., N. W. 
Lermond. Massachusetts, E. W. Roper. New York: Staten 
Island, E. W. Hubbard; Westchester Co., E. G. Vanatta; 
Dutchess Co., W. S. Teator; Madison Co., Henderson, Pilsbry; 
Herkimer Co., A. Bailey. New Jersey: Burlington Co., B. 
Long. Pennsylvania, Philadelphia Co., Montgomery Co., 
Pilsbry; Chester Co., W. D. Hartman. D. C., Sterki. Vir- 
ginia: Alexandria, Sterki. Ohio: Columbus, H. Moores; 
Lake Superior, J. T. Crans. 

Vertigo callosa STERKI, Proc. A. N. S. Phila. 1890, p. 31 
(Columbus, 0., Mass., N. Y.) not of Reuss, 1849. Pupa 
(Nearctula) superioris PILSBRY, Nautilus xii, 1899, p. 103 
(Lake Superior). Vertigo pygmcea Drap., STERKI, Nautilus 
vi, May, 1892, p. 5 ; Nachrbl. d. m. Ges. 1889, p. 114. PILSBRY, 
Proc. A. N. S. Phila. 1900, p. 608. HANHAM, Nautilus xi, 111 
(Quebec). HENDERSON, Naut. xx, p. 97 (Cazenovia, N. Y.). 
JOHNSON, Fauna of New England no. 13, 1915, p. 216 (Me. 
and Mass. ) . 

This species was first recognized in America by Dr. Sterki ; 
it had long been in collections, confused with other species. 
It proves to be somewhat widely spread, having been found in 
many places between Quebec, Maine, Virginia and Ohio. The 
strong, continuous crest behind the well expanded lip and the 
absence of a distinctly defined upper arc or sinulus of the 
outer lip are its more prominent features. V. gouldii is a 
paler, much more sharply striate shell with weaker crest and 
distinct sinulus. V. ventricosa has a weaker and interrupted 
crest. 

In some lots, as that from Cazenovia, Madison Co., N. Y., 
the basal fold is small or rarely absent, only five teeth 
present. In others of the same lot there are both basal and 
suprapalatal folds, the latter weak ; these having seven teeth. 
Dr. Sterki notes that sometimes the basal fold is bifid, as an 
individual variation, and occasionally a small angular lamella 
is present. The crest and the palatal callus vary in strength. 

For other references to V. pygm&a see under Palaearctic 
species, no. 52. 



98 AMERICAN SPECIES OP VERTIGO. 

10. VERTIGO GOULDH (Binney). PI. 7, figs. 4, 5, 8. 

Shell light chestnut, cylindrical-ovate ; whorls between 4 and 
5, ventricose, the last occupying nearly half the length of the 
axis ; apex obtuse. Aperture lateral, composed of two unequal 
curves meeting in the center of the outer lip ; with 5 prominent 
white teeth: one upon the transverse margin, two uppn the 
umbilical margin and two upon the labial margin; lip thick- 
ened, not reflected, umbilicus a little open (A. Binney). 

Length 1.85 to 1.95, diam. 1 mm. ; Brookline, Mass. 

Prince Edward and Magdalen Islands west to Alberta, 
Field, B. C. and Montana, south to the Potomac River, North 
Carolina (Sterki), Tennessee at Cade's Cove, Blount Co. ; near 
Valley Head, Alabama (H. H. Smith), and Pleistocene of 
Kansas ( ?) ; also reported from the West Indies. 

Pupa gouldii BINNEY, Proc. Boston Soc. N. H. i, 1843, p. 
105 ; Terrestrial Moll, ii, p. 332, pi. 51, f . 2. Vertigo gouldi 
W. G. BINNEY, Terr. Moll, v, p. 214. STERKI, Proc. A. N. S. 
Phila. 1890, p. 31 ; 8th Ann. Rep. Ohio State Acad. Sci., 1900, 
p. 32 (Tuscarawas Co., O.) ; Nautilus xxix, 123 (Geneva, 0.) ; 
iii, p. 125 (Helena, Mont.; Ottawa, Ont.). NYLANDER, 
Nautilus viii, 126 (Aroostook Co., Me.). HANHAM, Naut. x, 
191 ; xi, 111 (Quebec). CLAPP, Naut. xiv, 64 (Kennebunkport, 
Me.). HENDERSON, Naut. xx, 97 (Cazenovia, N. Y.). JACK- 
SON, Naut. xxi, 144 (North Haven, Me.). BERRY, Naut. xxiv, 
p. 63 (Unity, Me.). PILSBRY, Naut. xxv, 75 (Monroe Co., 
Pa.). VANATTA, Naut. xxviii, 11 (Sussex Co., N. J.). 
DANIELS, 27th Ann. Rep. Dep. Geol. and Nat. Res. Indiana, 
1902, p. 632 (Henry Co., Connersville, Dunreith, Ind.). 
COCKERELL, Nautilus x, p. 143 (post-Tertiary deposit at West 
Cliff, Colo.). WALKER, Moll, of Michigan, 1906, p. 517, f. 151 
(generally distributed). JOHNSON, Fauna of New England, 
no. 13, 1915, p. 214 (Westbrook, Bethel and Woodland, Me.; 
Cambridge, Roxbury and Westport, Mass.; Tiverton, R. I.; 
Northfield, Conn.). ? HANNA and JOHNSTON, Kansas Univ. 
Sci. Bull, vii, no. 3, Jan. 1913, p. 120, pi. 18, f. 4 (Pleistocene, 
Phillips Co., Ks.). 

Vertigo gouldii paradoxa Sterki, in NYLANDER, Nautilus xiii, 
Jan., 1900, p. 103. 



AMERICAN SPECIES OP VERTIGO. 99 

It is related to V. pygmcca, but the crest behind the lip is 
not so strong, the whorl is more flattened or impressed at and 
behind the point or "auricle" of the lip, giving the latter the 
appearance of being biarcuate, though it is not as conspicu- 
ously so as Binney's description would lead one to suppose. 
The surface is very distinctly striate, especially the penult 
whorl. The basal plica is subcolumellar in position. The 
parietal lamella is strong and rather long. There is never any 
trace of a palatal callus. 

"The angular lamella may be present or absent. The 
inferior columellar (basal) is by no means constant, and in 
some forms is generally wanting. In a few specimens from 
Summit Co., Ohio, there is a very small but distinct infra- 
parietal nodule ; one of these has an angular also, and is thus 
7-toothed. The palatal folds are rather variable in position 
and shape, and there may be a well marked external im- 
pression over them, or none" (Sterki). 

V. gouldn is rather commonly distributed in New England 
and New York, but more local southward, where it appears 
to follow the mountains to Tennessee and northern Alabama. 
The Pleistocene form from Kansas, as figured by Hanna and 
Johnston, differs so much in the position of the basal fold 
that its reference to gouldvi appears doubtful. So far as I 
know, there is no Austroriparian record, so that the West 
Indian occurrence appears anomalous, and requires confirma- 
tion. Dr. Sterki states that it is found in Cuba and Jamaica, 
and single specimens of this species are in coll. A. N. S. P. 
labelled St. Croix (Griffith), and Tortola (R. Swift). While 
these localities appear improbable, the high degree of accuracy 
of the Swift collection labels causes me to mention them. The 
possibility of importation or mixture of specimens must be 
considered. 



Vertigo gmtldii paradoxa Sterki, n. subsp. PL 12, figs. 
6,8. 

' ' Rather small, generally cylindrical, with the surface striae 
well developed, the palatals, usually somewhat long and thin, 
are close together, and the lower palatal is placed markedly 



100 AMERICAN SPECIES OP VERTIGO. 

inward. Inferior columellar (basal) is usually wanting or 
quite small ' ' (Sterki) . 

Length 1.75, diam. 1 mm. (fig. 6). 

Maine: Woodland, Aroostook Co., type loc., Nylander. 
Quebec, Sterki. 

Figured from cotypes, no. 119007 A. N. S. P. It had been 
mentioned before (Nautilus xiii, 103), with the note "Fine 
examples with the two palatal folds continuous and one an- 
gular" terms not diagnostic of the race. It stands midway 
between gouldii and nylanderi. 

10&. Vertigo gouldvi cristata Sterki, n. subsp. PL 12, figs. 
4,5. 

"The shell is rather large, about 2 mm. long (1.8 to 2.1) ; 
form cylindrical to somewhat oblong, barrel-shaped. Surface 
striae rather fine. Some distance behind the outer and basal 
lips there is a rather large, conspicuous crest, which does not 
extend above the middle ; behind it there is a broad flattening 
or impression over the palatals, the base being narrow there, 
then becoming rather broadly rounded towards the aperture. 
A small angular lamella may be either present or wanting. 
There is no basal ('lower columellar 7 ) fold." (Sterki.) 

Length 2.1, diam. 1,2 mm. (fig. 4). 

Canada : Quebec, rather abundant, A. W. Hanham. 

A strongly marked race, having the crest as well developed 
as in many examples of V. pygmcea,, but without a callus in 
the palate, and with the sculpture of gouldii. Figured from 
cotype no. 119008 A. N. S. P. 

11. VERTIGO NYLANDERI Sterki. PL 7, figs. 13, 14, 15. 

Shell rimate, oblong, with a rather acute apex, cinnamon 
colored, pellucid; whorls 4%-5, quite convex, with a deep 
suture; sculptured with somewhat irregular, crowded striae 
(except the embryonic whorl) ; the last occupying about one- 
half of the altitude, gradually narrowed towards the aperture, 
which is small. Peristome slightly everted, margin not thick- 
ened. The outer margin has an indentation barely above its 
middle, forming a well-marked sinulus ; behind it a trace of a 



AMERICAN SPECIES OP VERTIGO. 101 

crest, and behind that a long, deep, furrow-like impression 
over the palatal folds, ascending obliquely from near the base ; 
no callus within ; lamellae and plicae 6 ; parietal lamella long 
and curved ; parallel with it is a thin, lamellif orm angular ; 
columellar lamella ascending inwardly. Palatal folds long, 
the lower palatal deep-seated, emerging only about to the inner 
end of the upper. Basal fold small, subcolumellar in position. 

Length 1.7, diam. 0.95 mm. 

Length 1.6, diam. 0.9 mm. 

Maine : Woodland, Aroostook county, 0. 0. Ny lander. 

Vertigo nylanderi STERKI, Nautilus xxii, Feb., 1909, p. 107. 

By the fine striation it has some resemblance to V. gouldti 
and might be regarded as an extreme form of V. g. paradoxa, 
as Dr. Sterki has suggested to me. The color is more that 
of V. bollesiana. It differs from those, and from all other 
American species, by the deeper immersion of the long lower 
palatal fold, and the deep impression in the back, over the 
palatals. 

It is remarkable that this strongly characterized species has 
been found in only a single locality. Type no. 1075 Sterki 
coll. Description and figures from a cotype, 98331 A. N. S. 

12. VERTIGO BOLLESIANA (Morse). PI. 7, figs. 9, 10. 

Shell minutely perforate, cylindrical ovate, delicately 
striated, subtranslucent ; apex obtuse; suture well defined; 
whorls four, sub-convex ; aperture suborbicular, somewhat flat- 
tened on its outer edge; with five teeth, one prominent and 
rather curved on the parietal margin, two similar in form, the 
low^er one the smaller, on the columellar margin, and two 
slightly elevated lamelliform teeth within and at the base, 
peristome subreflected and thickened. 

Length .065 inch; breadth .035 inch (Morse). 

Maine: throughout the State, type loc. Orono. Massachu- 
setts, New Hampshire, New York. Also reported from 
Indiana and Michigan, south to Norfolk, Virginia (Morse) 
and mountains of Bast Tennessee. 

Isthmia bollesiana MORSE, Ann. Lye. of Nat. Hist, of N. Y., 
viii, 1865, p. 209, figs. 4-6. W. G. BINNEY, Terr. Moll. vol. v, 



102 AMERICAN SPECIES OF VERTIGO. 

1878, p. 215, f. 120; Man. Amer. Land Sh., 1885, p. 191. 
STERKI, Nautilus iv, p. 9, footnote; iii, 125 (Sewanee, Tenn. ; 
St. Croix, W. I.). PILSBRY, Proc. A. N. S. Phila. 1900, 133 
(Cades Cove and Tuskeegee Mt., E. Tenn.). TEATOR, Nau- 
tilus iii, 69 (Dover Plains, Dutchess Co., N. Y.). PLEAS, Naut. 
vii, 68 (Henry Co., Indiana). GARDNER, Naut. viii, 76 (Long 
Island, N. Y.). CLAPP, Naut. xiv, 64 (Mt. Agamentieus, 
Me.). BLANEY, Naut. xviii, 46 (Ironbound I., Me.). 
WALKER, Moll, of Michigan 1906, p. 518, f. 152 (Petoskey, 
Charlevoix, Crystal Lake, Benzie Co. and Huron Mts., Mar- 
quette Co., Mich.). JOHNSON, Fauna of New England, no. 
13, 1915, p. 214 (Bethel, Me., Francestown, N. H.). 

There is an extremely small crest close behind the lip, and 
a rather large oblique depression over the palatal folds. The 
basal fold is subcolumellar in position, and rarely it is absent. 
As Morse says, it is smaller, lighter-colored and more trans- 
parent and delicate than V. gouldii, and it is less distinctly 
striated. The teeth are smaller, especially the palatals. 
Length 1.5, diam. 0.9 mm. ; 4% to 4% whorls. 

According to Morse, it occurs under dead leaves and on 
bark, in hardwood groves. It appears to be rare except in 
Maine. 

The specimens at hand are from New England and New 
York. The western records, Michigan, Ohio, Indiana, and 
those from East Tennessee I have not been able to verify 
personally. Dr. Sterki writes that he has ' * no specimens from 
Michigan and Ohio; some records may have been founded on 
misidentification. It appears to be northeastern." 

Dr. Sterki contributes the following notes: "After again 
looking over a good deal of material, including a number col- 
lected by E. S. Morse, I come to the conclusion that V. ~bol- 
lesiana is specifically distinct from gouldii, though some speci- 
mens of the latter resemble it closely. V. bollesiana is rather 
uniform in size, length 1.3 to 1.6 mm. (1) The striae are 
slighter, finer, than in goiddvi, often subobsolete, sometimes 
subregular, very fine and crowded. (2) the form is oval or 
ovoid, never cylindric. (3) it is generally of lighter color, 
corneous, not reddish or brownish. (4) the configuration of 



AMERICAN SPECIES OF VERTIGO. 103 

the palate externally is different, though gouldii varies in that 
respect. In bollesima there is generally a small, narrow crest 
close behind the lip-margin, and a slight impression at the 
auricle, suggesting a double curve of the outer lip. This is 
less marked in gouldii. 

"The inferior columellar (basal) is often wanting or 
vestigial. In some specimens it is bifid. Some specimens 
have a very small angular. The peristome was described as 
subreflected and thickened, but it is narrowly everted, and 
thus only apparently thickened in a front view. ' ' 

13. VERTIGO HEBARDI Vanatta. PL 8, fig. 4. 

The shell is minute, distinctly perforate, very shortly rim- 
ate, shortly oval, fragile, corneous; first whorl smooth, the 
penult irregularly, finely striate, last whorl with few striae; 
glossy. The whorls are rather strongly convex, the last not 
noticeably flattened or grooved, and without a crest behind the 
lip. The aperture is indistinctly triangular and has 5 teeth : 
angular lamella very low, half as long as the rather short, 
high parietal. The columellar lamella enters deeply and 
horizontally, its crest slanting downward. The palatal folds 
are rather short, subequal, the lower being slightly stouter and 
a trifle further in. The outer lip is scarcely expanded, some- 
what straightened, without a projecting point. Length 1.25, 
diam. 0.84 mm. ; 4 whorls. 

Florida: Long Key, Morgan Hebard. 

Vertigo hebardi VANATTA, Proc. A. N. S. Phila. 1912, p. 
445, fig. 

It is most like V. bollesiana, but the shell is shorter, has a 
distinct angular lamella and no external crest or impression 
behind the lip. Figure and description from the type. 

14. VERTIGO PERRYI Sterki. PI. 7, fig. 7. 

' ' Shell minute, dextrorse, ovate with the apex rather acute, 
rimate ; thin, transparent, of rather dark brown color with a 
slight greenish tinge [dark olive buff]. Whorls 4%, rather 
rapidly increasing, separated by a moderately deep suture, the 
last comparatively large, occupying over one-half of altitude, 



104 AMERICAN SPECIES OP VERTIGO. 

rounded ; with a slight impression over the palatal fold ; aper- 
ture well rounded, truncate, the margins slightly everted, the 
outer margin barely impressed at the auricle which is marked 
by a slight angle projecting over the level of the peristome ; 
no callus in the palate ; lamellae and folds three or four, small, 
very short, of brownish color ; the parietal, columellar and in- 
ferior palatal, and sometimes there is also a superior palatal. 
Surface with very fine, irregular striae, somewhat shining,. 
Alt. 1.5 to 1.6, diam. 1.1 mm.; aperture, alt. 0.6 mm." 
(Sterki). 

Ehode Island : Warwick, J. F. Perry. Massachusetts: Dux- 
bury, W. F. Clapp. 

Vertigo perryi STERKI, Nautilus xix, Sept., 1905, p. 53. 

"The present species resembles the low form of Vertigo 
ventricosa Mse. in the shape and size of the shell, but the 
formation of the aperture and its lamellse and folds is quite 
different, the color is deeper and the surface less shining. 
From the other three described, typically three-toothed, east- 
ern Vertigos: tridentata Wolf, oscaricwia Sterki and parvula 
Sterki, V. perryi is also very different ; in all of the three, the 
parietal lamellae and palatal folds are much larger, longer, and 
of whitish color ; the aperture is higher than wide ; the shells 
are more elevated and of lighter color" (Sterki). 

The two localities now known for this strongly distinct 
species are only about 45 miles apart, and both are close to 
the sea. It is remarkable that a species so well characterized 
could exist undetected for so long, close to centers of concho- 
logical work. Specimens are contained in the collections of 
Dr. Sterki, Mr. Perry, Mr. H. F. Carpenter, the Academy of 
Natural Sciences, The Museum of Comparative Zoology and 
perhaps some others. The example figured measures: length 
1.6, diam. 1.05 mm. In some others the teeth are weaker, and 
rarely there is the trace of an upper palatal fold. Often the 
columellar lamella is hardly visible in a front view. The 
lower palatal fold is often developed when the other teeth are 
scarcely noticeable. The shell is very fragile. At Duxbury, 
where it was collected by Mr. and Mrs. W. F. Clapp, it lives 
on grass in a swamp, in wet weather as much as a foot above 1 
the ground. 



AMERICAN SPECIES OP VERTIGO. 105 

15. VERTIGO PARVULA Sterki. PL 12, figs. 7, 9. 

"It is of about the size, shape and appearance of V. (Angus- 
tula) milium Gld., but ranges in quite another group, having 
a quite simple palatal wall and margin, and only 3 lamellae" 
(Sterki). 

The shell is minute, subcylindric, tapering very little up- 
wards, the summit obtuse; thin, subtransparent, slightly yel- 
lowish, smooth and glossy, becoming finely striate behind 
the outer lip. The whorls are moderately convex, the last 
whorl well rounded, slightly impressed behind the projection 
of the outer lip. The aperture is somewhat triangular, with 
three teeth : parietal lamella rather short and high ; columellar 
lamella short, steeply ascending inwardly; lower palatal fold 
rather high in front, rapidly becoming lower as it recedes, 
penetrating to the dorsal side. Peristome very little everted, 
slightly thickened, and having a distinct callus ridge within. 
The outer lip projects forward and is slightly bent inward 
above the middle. 

Length 1.55, diam. 0.85 mm. ; barely 5 whorls. 

Ohio: Summit Co., A. Pettingell. Also found by A. G. 
Wetherby in the mountains of North Carolina, according to 
Dr. Sterki. 

V. [ertigo] parvula STERKI, Nautilus iii, April, 1890, p. 136. 

Figures and description are from the type specimen, no. 
270 Sterki coll. It is one of the rarest species, known by the 
small size, cylindric shape and three well-developed teeth, 
the lower palatal being quite long and not marked externally 
by an impression. The auricle or point of the outer lip pro- 
jects well forward but is not much bent inward. 

It is much smaller than V. tridentata, and more cylindric 
and lighter colored than V. perryi, with relatively larger teeth. 
It is quite distinct from all of our species. 

The name parvula has been used in Pupa by Deshayes, 1864, 
but as there is some doubt as to whether his species is a 
Vertigo, I do not think it necessary to change the name of the 
form under consideration. 



106 AMERICAN SPECIES OP VERTIGO. 

16. VERTIGO TRIDENTATA Wolf. PL 12, figs. 1, 2, 3. 

"Shell narrowly ovate, amber-colored, highly polished; 
whorls 5, smooth, with three teeth in the mouth, one on the 
middle of the lower lip, and one on each side, forming a regu- 
lar triangle" (Wolf). 

The shape varies from ovate to tapering oblong. It is 
honey-yellow, shading to somewhat browner below, paler 
above; surface smooth, with only faint indications of striae, 
glossy. The last whorl is somewhat flattened externally over 
the lower palatal fold, and has a rather narrow but generally 
distinct crest behind the lip. The outer lip projects forward 
and slightly inward near the middle. Parietal lamella high, 
rather short. Columellar lamella blunt, directed downward. 
Lower palatal fold strongly developed. Upper palatal fold 
quite small or sometimes wanting. These folds stand on a 
more or less distinct palatal callus. Angular lamella and 
basal fold are never developed. 

Length 2.2, diam. 1.1 mm.; 5% whorls. 

Length 2, diam. 1.1 mm. 

Length 1.85, diam. 1.1 mm.; 4% whorls. 

New York : Troy, Aldrich ; Mohawk, G. H. Clapp ; Syracuse, 
pleistocene, Burnett Smith. New Jersey: Princeton, A. D. 
Brown ; near Clementon, Bayard Long. Pennsylvania : Phila- 
delphia; York Furnace, A. P. Brown; Northampton Co., B. 
Long. Ohio : New Philadelphia, Sterki. Indiana : Lafayette, 
Tippecanoe Co., Satterthwait. Illinois: Canton, J. Wolf, 
type loc. Also reported from Ontario, Minnesota, Michigan, 
Kansas, Colorado, New Braunfels, Texas, etc. 

Vertigo tridentata WOLF, Amer. Journ. of Conch, v, 
May 5, 1870, p. 198, pi. 17, f. 1. STERKI, Proc. U. S. Nat. 
Mus. xi, 1888, p. 375, no. 15, pi. 42, f . 4 ; Nautilus iii, April, 
1890, p. 135; xxix, 123 (Geneva, 0.). PLEAS, Naut. vii, 68 
(Henry Co., Ind.). HANHAM, Naut. xi, 111 (Quebec). 
DANIELS, 27th Ann. Rep. Dep. Geol. and Nat. Ees. Indiana, 
1902, 632 (Danville and Dunreith, Ind.). HANNA, Naut. 
xxiii, 95 (Douglas Co., Kansas). WALKER, Moll, of Michigan, 
1906, p. 518, f. 153 (Ann Arbor and Grand Rapids). J. 



AMERICAN SPECIES OF VERTIGO. 107 

HENDERSON, Univ. of Colo. Studies iv, p. 172 (South Park, 
Colo.). SARGENT, Naut. ix, 89 (Clearwater, Minn.). JOHN- 
SON, Fauna of New England 13, 1915, p. 214 (Maine). 

The light color, tapering form, absence of basal and angular 
teeth and the small size or sometimes absence of an upper 
palatal, distinguish this from other American species. Most 
specimens have the upper palatal fold developed, either dis- 
tinct though small, or as a trace; yet in some it is wholly 
absent. The type is no. 58008 A. N. S. P, figured in A. J. C. 
Figs. 1-3 represent specimens of the type lot. 

The species was placed in the synonymy of V. ovata by Mr. 
Binney, but Dr. Sterki, in 1888, called attention to its dis- 
tinctive characters, which are now generally admitted. The 
two species are not closely related. 

Mr. Wolf found it "abundant in shady copses on green 
weeds, climbing as high as three feet from the ground. I col- 
lected 12,000 from standing weeds and not one from the 
ground, although it was searched well to find them." 

V. perryi differs from V. tridentata by the smaller size, 
more fragile, distinctly greenish shell, with broader, more 
rounded aperture, smaller teeth and dark-edged peristome. 

17. VERTIGO MARKI Gulick. PL 13, fig. 17. 

The shell is shortly rimate, ovate, the spire convexly conic, 
the summit obtuse. Surface nearly smooth; color yellowish 
(bleached) . The whorls are moderately convex, the last some- 
what flattened over the lower palatal region, having a low, 
white crest close to the lip. Aperture with 4 teeth : a strong, 
moderately long parietal lamella, a low, massive columellar 
lamella, a small upper and larger lower palatal fold, both 
tuberculiform. The palatal callus is strongly developed. 
Peristome is slightly expanded, a little prominent and bent 
in above the middle of the outer margin. 

Length 2.05, diam. 1.1 mm.; 5 whorls (type). 

Bermuda: Paynter's Vale (Gulick, S. Brown) and Bailey 
Bay road cut ( Verrill) . 

Vertigo marki GULICK, Proc. A. N. S. Phila. 1904, p. 414, 
pi. 36, f. 7. VERRILL, Trans. Conn. Acad. xii, 1907, p. 170. 



108 AMERICAN SPECIES OF VERTIGO. 

It appears rather closely related to V. tridentata from which 
it differs chiefly by the narrower aperture, further contracted 
by a much stronger palatal callus. Description and figure 
from the type, no. 85574 A. N. S. P. Other specimens were 
taken by S. Brown in leaf mould, Paynter's Vale, near the type 
locality; but while doubtless recent they are bleached. It is 
far less abundant than V. numellata in the same places. 

( West Coast species of the V. pygmcea group ) . 
18. VERTIGO COLUMBIANA Sterki. PL 9, figs. 12, 13. 

Shell minute, cylindric-oval, perforate, thin, pale corneous- 
brown (grayish cream-buff), somewhat transparent, glossy and 
weakly striatulate. Whorls nearly 5, convex, the last ex- 
panded in a very low crest very close to the lip, not noticeably 
constricted in front of the crest. Aperture truncate-oval, 
4-toothed, the peristome thin, hardly expanded; parietal la- 
mella short and high, columellar a little smaller, lower palatal 
a rather short fold, about twice as long as the upper palatal 
which is smaller, shorter, almost tuberculif orm ; all the teeth 
are white, and the palatals show through the outside wall. 

Length 1.9, diam. 1.1 mm. (type). 

Length 2.05, diam. 1.2 mm. 

Vancouver Island, George W. Taylor, type loc. (no. 68881 
A. N. S.). Washington: Olympia and Tacoma, H. Hemphill; 
Seattle, Hemphill, P. B. Randolph; L. Quiniault, Chehalis 
Co., S. S. Berry. Oregon: Douglas county, F. H. Andrus. 
St. Paul Island, Bering Sea, Dall. 

Vertigo columbiana STERKI, Nautilus, vi, 1892, p. 5 (name 
only). PILSBRY & VANATTA, Proc. A. N. S. Phila., 1900, p. 
602, pi. 23, fig. 11. DALL, Alaska Land and Fresh Water 
Mollusks, 1905, p. 30. Pupa columbiana, STERKI, PILSBRY, 
Nautilus xi, 1898, p. 119 ; Class. Cat., p. 21, no. 212. 

The surface is decidedly less striate than in V. coloradensis, 
V. concinnula or V. gouldii. It is like that of V. ventricosa. 
There is no angular lamella and no basal fold in any of the 
specimens; but only a few have been seen from each locality. 
The single specimen from Olympia is very short, 1.4 x 1 mm. 



AMERICAN SPECIES OF VERTIGO. 109 

(fig. 13). The species was mentioned in lists by Dr. Sterki 
as early as 1892, but it was described from no. 68881 A. N. 
S. P., in 1900. 

I have hesitated whether to rank V. columbiana as a western 
race of V. ventricosa, but as I have seen only one or two from 
each of the localities, it is left distinct temporarily. There 
is no "auricle" or incurved point of the outer lip, such as 
most specimens of ventricosa show, and none of the examples 
seen shows a basal tooth; yet in ventricosa this is sometimes 
lacking. If columbiana is not a race of V. ventricosa, it is 
certainly very near akin. PL 9, fig. 12 represents the type 
specimen, no. 68881 A. N. S. P. I have not seen the St. Paul 
Island specimens. 

V. columbiana utahensis Sterki (Nautilus vi, p. 5, name 
only; Pils. & Van., P. A. N. S. 1900, 603, pi. 23, fig. 10; Pils. 
& Ferr. 1910, p. 144) is identical, I believe, with V. colo- 
radensis, and has no direct relationship with V. columbiana. 
The type, no. 109009 A. N. S. P., is figured, pi. 12, fig. 12. 

19. VERTIGO ANDRUSIANA Pils. PL 11, figs. 9, 10, 11. 

The shell is rimate, imperf orate, cylindric-oblong, with con- 
vexly conic, obtuse summit; cinnamon-brown, becoming paler 
upwards, the initial whorl gray; somewhat glossy, without 
noticeable striation, most minutely granulose. The whorls 
are moderately convex, the last having a rather low but distinct 
crest followed by a wide, shallow contraction behind the lip ; 
an impressed line over the upper palatal fold terminates at 
the lip, which is slightly bent in at this point. The aperture 
has (four to) six teeth: a short, high parietal lamella, a small, 
tubercular angular lamella (sometimes very inconspicuous), 
a stout, short, columellar lamella which ascends a little in- 
wardly, a small, tubercular basal fold (sometimes wanting) 
and two well developed palatal folds, the lower being longer. 
The palatal folds stand upon a thin, light-colored callus. The 
outer lip is scarcely expanded, the basal slightly so, the colu- 
mellar margin dilated. 

Length 2.46, diam. 1.3 mm.; 5y 2 whorls (type, fig. 10). 

Length 2.35, diam. 1.3 mm.; 5% whorls. 

Length 2.3, diam. 1.3 mm. 



110 AMERICAN SPECIES OF VERTIGO. 

Oregon : Douglas Co., F. H. Andrus, type loc. Also north- 
ward, to Chehalis Co., Washington. 

Vertigo cmdrusiwna PILSBRY, Proc. A. N. S. Phila. 1899, 
p. 315, fig. 3 ; Nautilus xvii, p. 131. 

It stands very near V. pygm&a, but is slightly longer, the 
crest and palatal callus are less developed, and there is an 
angular lamella in the most fully developed examples, which 
however have the other teeth smaller than in pygnuea. 

The type specimen is redescribed above and drawn in pi. 
11, fig. 10. In the original description and figure the angular 
lamella was overlooked; the shell was rolled too far towards 
the right under the monocular microscope used, and this small 
tooth was not seen. 

There are ten shells in the original lot, but perhaps only 
the type and another are absolutely mature. Both have the 
angular and the basal teeth distinct. Two other shells are 
very nearly adult. In both the angular lamella is represented 
by a slight thickening, of the color of the shell, and only notice- 
able in a basal view, and neither of them has a basal fold 
(fig. 11). It is likely that the basal is either wanting or 
present in adult examples of the species. 

Similar specimens were taken by Mr. S. S. Berry at Lake 
Quinault, Chehalis Co., Washington ; one before me has a very 
low angular lamella but no basal fold. The spire has whitish 
streaks. 

A series of some thousands of specimens was taken by Mr. 
John A. Allen "about clumps of bushes in a meadow," 
Oswego, Clackamas Co., Oregon. The shells (pi. 11, fig. 9) are 
all smaller than the type lot, variable in size and shape, and 
with the crest weak, or in the shorter individuals wanting. 
Among many examined, none has an angular lamella. The 
basal fold is occasionally developed, but more frequently 
absent. There is a distinct if thin palatal callus except in 
the quite small individuals. The color is usually darker than 
cinnamon brown or russet, often with light streaks on the spire. 

Length 2.15, diam. 1.2 mm. ; 514 whorls. 

Length 1.85, diam. 1.2 mm. ; 4% whorls. 

While this form has characters of V. columbiana, V. pygmtea 



AMERICAN SPECIES OP VERTIGO. Ill 

and V. a. sanbernardinensis, it appears most closely related 
to V. andrusiana. 

19a. Vertigo andrusiana sanbernardinensis n. subsp. PL 11, 
figs. 6, 7, 8. 

The shell is shortly rimate, imperforate, oblong, tapering 
very little upwards, terminating in a rounded, obtuse summit ; 
cinnamon-brown, moderately glossy, weakly, irregularly stri- 
ate. The whorls are rather convex, the last slightly flattened 
over the lower palatal fold, a trifle swollen in front of the 
flattening, but not crested, often having an impressed 
line parallel to the suture over the upper palatal fold. 
Teeth four to six: parietal lamella short, compressed, a 
small, tubercular angular lamella, even with or in advance of 
its outer end (sometimes wanting). Columellar lamella short 
but rather massive. Lower palatal fold strong and rather 
long, the upper palatal very much smaller and shorter. 
(Basal fold small and tubercular, at the junction of basal and 
columellar margins, and not present in the form selected as 
typical). There is no noticeable callus between the palatal 
folds. The outer lip expands very little; it is straightened 
and slightly thickened internally above the middle. Colu- 
mellar lip is narrowly reflected. 

Length 2.25, diam. 1.3 mm.; 5 whorls. 

Length 2.2, diam. 1.2 mm. 

Length 2.15, diam. 1.2 mm.; 5 whorls (type). 

Length 1.9, diam. 1.1 mm. ; 4% whorls. 

California: San Bernardino Mts., 7550 to 7750 ft., in the 
cienaga below Bluff Lake (type loc.), that north of the lake, 
and Bluff Lake meadow, everywhere associated with the local 
races of V. modest a, all collected by S. S. Berry. 

This form stands close to V. andrusiana, but there is no crest 
or contraction behind the lip, and no trace of a palatal callus. 
It appears to represent one extreme of a series leading through 
andrusiana to pygm&a. Whether the California form re- 
quires subspecific separation from andrusiana is open to doubt, 
but there is certainly some difference typically. It differs 
from V. columbiana by the much darker color, by being less 



112 AMERICAN SPECIES OF VERTIGO. 

thin, and typically by having more teeth. V. binneyana is 
a paler, slightly more slender species, having a more distinct 
crest, and a deeper external impression over the palatal folds. 
All of these forms appear to be closely related, and are dis- 
criminated with some difficulty. 

The type is no. 118419 A. N. S. P. A paratype has been 
placed in the collection of S. S. Berry. 

In the topotypic lot the following tooth-mutations were 
found in adult specimens : 

1. Angular, parietal, columellar, basal, upper and lower 
palatals. 

2. Angular, parietal, columellar, upper and lower palatals 
(type). 

3. ... parietal, columellar basal, upper and lower palatals. 

4. ... parietal, columellar, . . . upper and lower palatals. 
The second arrangement is much the more abundant, the 

first, or complete tooth formula, and the third being rare, 
one of each out of 27 specimens. 

The presence of a basal fold is therefore exceptional in 
shells from around Bluff Lake. 

20. VERTIGO STERKH n. sp. PI. 11, figs. 2 to 5. 

The shell is imperforate, with a curved umbilical crevice, 
ovate, chestnut-brown, glossy, slightly striate, but becoming 
distinctly striate behind the lip. The whorls are moderately 
convex, the last without any trace of a crest or contraction be- 
hind the lip, having a short, inconspicuous impression behind 
the auricle. The aperture has three teeth (or sometimes two) . 
The parietal lamella is low and short. Columellar lamella 
small, obtuse and deeply placed. The lower palatal fold is 
small, tuberculiform (and wanting in some examples). There 
is no palatal callus. The peristome is thin, brown, and well 
expanded, having a slight prominence inwardly above the 
middle of the outer margin, above which it is narrower. 

Length 2.1, diam. 1.25 mm. ; 4i/ 2 whorls. Type, fig. 3. 

Length 2, diam. 1.23 mm. 

Length 1.8, diam. 12 mm. 

California : Funston Meadow, Kern Eiver, Tulare Co., type 



AMERICAN SPECIES OF VERTIGO. 113 

loc.; Onion Valley, Kearsarge Pass, Inyo Co., J. H. Ferriss 
and E. Hand, 1916. 

V. dalliana is a more conic, thinner shell, of different color, 
having more rapidly increasing, more convex whorls, and a 
larger aperture. Moreover, adult specimens of V. sterkii are 
always toothed. V. andrusiana is generally larger, and so 
far as seen, it always has much larger teeth. V. tridentata is 
a far paler, more slender shell, with smaller aperture, stronger 
teeth and a distinct auricle. 

In both of the localities V. sterkii occurred associated with 
V. modesta castanea a form also characterized by degenera- 
tion of the teeth. 

Group of V. coloradensis. 

Subcylindric, distinctly striate forms, the striation strong- 
est on the penult whorl, related to the modesta and to the 
ventricosa and gouldii groups. Typically they are mountain 
snails 6000 ft. up ; but V. hannai and V. arthuri are from 
the lower plains eastward. 

Key to species. 

1. 2 to 2.3 mm. long ; a low crest ; parietal and columellar 
lamellae, sometimes an angular; 2 long palatal folds. 
Alpine in Colo., N. M., Ariz. V. concinnula, no. 23. 
1.5 to 1.8 mm. long. 2. 

2. No angular lamella. 3. 
An angular lamella present ; crest wanting or very weak. 5. 

3. No noticeable palatal callus; a low crest. 1.7-1.9 mm. long. 
Mountain forms. 4. 
A strong palatal callus ; 1-1-3 teeth ; 1.5 x 0.8 mm. Western 
Dakota. V. arthuri, no. 24. 

4. Teeth 1-1-2, no basal fold. V. coloradensis, no. 22. 
Teeth 1-1-3, a basal present. V. c. basidens, no. 22a. 

5. Teeth 2-1-2, no basal fold. V. c. arizonensis, no. 22&. 
Teeth 2-1-3, a basal present. 6. 

6. Diameter 1 mm. or more; striation irregular. Western 
Kansas. V. hannai, no. 21. 
Diam. under 1 mm. Santa Catalina Mts., Arizona. 

V. c. inserta, no. 22c. 



114 AMERICAN SPECIES OF VERTIGO. 

21. VERTIGO HANNAI Pilsbry, n. n. PI. 12, fig. 12. 

"Shell light brown; ovate in outline. Lines of growth 
faint and oblique. Whorls, four and a half, well rounded, 
and the sutures well impressed. Apex smooth and white and 
obtusely pointed. Peristome thin and sharp, slightly ex- 
panded and the ends connected across the body whorl by a 
thin deposit of callus, almost no indentation in the upper 
palatal wall. Aperture semicircular and with six teeth. Two 
on the parietal wall, both of which are lamellar in shape, and 
the angle tooth is the smaller of the two, one columellar in 
the center of that wall of the aperture. This tooth is bifid, 
that portion toward the apex of the shell being the larger. 
One basal tooth, small and nodule-like. Two palatals, both of 
which are lamellar in shape, the lower one of which is the 
larger. Variation in the large series of this species is slight " 
(H. & J.). 

Length 1.77, diam. 1.04 mm. 

Length 1.74, diam. 1.04 mm. 

Length 1.63, diam. 1.04 mm. 

Length 1.55, diam. 1.04 mm. 

Kansas: Phillips County, along Prairie Dog creek between 
Norton and the Eepublican river, Pleistocene, Hanna and 
Johnston; type in U. S. N. M. no. 226396. 

Vertigo martini HANNA and JOHNSTON, Kansas University 
Science BuUetin vii, no. 3, Jan., 1913, p. 120, pi. 18, f. 3. 
Not V. martini Sayn, 1911. 

"This and Vertigo ovata are the only species found in this 
part of the country with two teeth on the parietal wall. The 
latter species, however, is much the larger and more ovate in 
outline. The size of martmi is about the same as that of 
gouldi from the same deposits, but that species is more cylin- 
drical and the angle tooth is never developed. The size is 
somewhat less than that of Vertigo tridentata, a recent shell 
of Eastern Kansas in which the basal tooth is absent" 
(H. & J.). 

The entire absence or merely indistinct trace of a crest be- 
hind the lip, the slighter "auricle" of the latter, and the 



AMERICAN SPECIES OP VERTIGO. 115 

shorter palatal folds, as well as the somewhat shorter, wider 
shape, separate this species from V. binneyana, which is other- 
wise similar. It does not agree in detail with any of the 
forms of V. coloradensis, though closely related to them. The 
striation is of irregularly, rather widely spaced wrinkles, as 
strong as in V. coloradensis but decidedly less regular and 
less crowded than on the penult whorl of that species. The 
outer lip is not at all expanded, rather blunt. The parietal 
lamella is rather long, as in V. coloradensis. 

The figure is from the type, one of no. 226396 U. S. N. M. 
As the name had been used by Sayn for a Pleistocene species 
of France, that of Mr. Hanna has been substituted. 

22. VERTIGO COLORADENSIS (Cockerell). PL 12, fig. 13. 

"Shell brown, shiny, thinnish, translucent enough to show 
teeth through (body whorl) from outside, striate, especially on 
penultimate whorl. Outline oblong-oval, barrel-shaped, apex 
blunt. Whorls four. Aperture pyriform. Peristome brown, 
thick, continuous by a well-marked callus on parietal wall. 
Outer lip not constricted; a crest is indicated behind peris- 
tome, but not well developed. The teeth within the aperture 
are brown, one long one on parietal wall, one on columellar, 
and two, the lower one largest, on outer wall. Length 1%, 
diam. 1 mm. (Ckll.). 

Length 1.75, diam. 1 mm. 

Colorado: near Swift Creek, Custer Co., Cockerell, type loc. 




FIG. 1. VERTIGO COLORADENSIS CKLL. 

Utah : Box Elder canyon at 4500 ft., H. Hemphill. Arizona : 
Pine Canyon, at 7500 ft., and head of Cave Creek canyon. 



116 AMERICAN SPECIES OP VERTIGO. 

8000 ft., Chiricahua Mts., Ferriss. 

Pupa coloradensis COCKERELL, Journ. of Conch., Leeds, vi, 
1889, p. 63 (name only) ; British Naturalist, 1891, p. 100 
(description) ; and in Binney, Fourth Supplement to Terr. 
Moll., v, Bull. M. C. Z., xxii, no. 4, p. 191. Vertigo colora- 
densis Ckll., STERKI, Nautilus, vi, 1892, p. 5. COCKERELL, 
Nautilus, x, 1897, p. 134. PILSBRY & VANATTA, Proc. A. N. S. 
Phila. 1900, p. 603, fig. 2. PILSBRY, Nautilus xvi, p. 58 (copy 
of orig. desc.). Vertigo columbiana utahensis Sterki, in 
PILSBRY & VANATTA, Proc. A. N. S. Phila. 1900, p. 603, pi. 
23, fig. 10. 

Professor Cockerell's description and figure are given above. 
The type specimen is said to be in the British Museum. It 
differs from V. concinnula by the smaller size, from V. gouldii 
by the absence of a basal fold and the coarser striation ; typi- 
cally it is also a trifle smaller than usual in gouldn, measuring 
1.75 mm. long, 1 wide, while gouldii is generally 1.85 to 2mm. 
long. Prof. Cockerell's measurement, 1% mm., was certainly 
only approximate. The columellar lamella is a little oblique, 
ascending somewhat as it enters. Perhaps it could be ranked 
as a subspecies of gouldii, but I think it sufficiently distinct. 

V. columbiana utahensis from Box Elder canyon, northern 
Utah, is the same thing. The type is drawn in pi. 12, fig. 13. 

A few specimens from the Chiricahua Mountains are slightly 
larger, 1.9 x 1.1 mm. 

Dr. Sterki has given me the following notes on a specimen 
received from Cockerell through Binney. ' ' Subcylindrical to 
somewhat barrel-shaped, perforate, whorls over 5, the last nar- 
rowed; aperture very small, 0.5 mm. high; a slight, rounded 
crest behind the margin and a slight flattening or even im- 
pression over the palatals; at the auricle a slight, small, im- 
pressed groove just behind the margin and barely above the 
upper palatal plica. Peristome not or very little thickened; 
inside there is a distinct, rather thin callus into which the 
two palatal plicae merge, the lower rather long, the upper 
much shorter; no suprapalatal. Parietal moderately large; 
a very small angular; columellar not large but well formed, 
apparently not lamellar. 1.7 x 1 mm. ' ' 



AMERICAN SPECIES OF VERTIGO. 117 

22a. Vertigo coloradensis basidens Pils. & Van. PL 12, fig. 15. 

Cylindric-oblong, with strongly striate middle whorls, and 
with a parietal lamella only on the parietal wall, as in 
V. coloradensis ; but having a small, short basal fold within 
the junction of eolumellar with basal margins; the lower 
palatal fold is especially long; there is a more or less distinct 
callus running upward from the outer end of the upper palatal 
fold ; finally, there is a wide and more or less prominent crest 
behind the lip. Length 1.8, diam. 0.95 mm. 

New Mexico: Bland, Bernalillo Co., Ashnmn, type loc. 
Colorado: Rio Blanco, Cockerell. Montana: Ward, L. E. 
Daniels. British Columbia: Field, S. Brown. 

V. coloradensis basidens PILSBRY & VANATTA, Proc. A. N. S. 
Phila. 1900, p. 604. 

The palatal folds, especially the lower, are decidedly longer 
than in V. gouldi. The crest is rather broad and sometimes 
decidedly prominent, more so than in any closely related form. 

Two specimens from Field measure : 1.75 x 1.05 and 1.95 x 
1 mm. Those from Ward, Mont., have the crest especially 
strong and light-colored, and the callus above the upper 
palatal fold is quite heavy. 

22&. Vertigo coloradensis arizonensis Pils. & Van. PL 12, 

figs. 14, 16. 

Shell cylindric-oval, rimate, very small; very densely and 
sharply but most minutely striate ; light brown. Whorls con- 
vex, the last tapering below, the last half whorl narrow as 
though pinched at base, flattened over the position of the 
palatal folds, then rising in a low, hardly noticeable crest, 
obsolete except near the base. Aperture irregularly truncate- 
oval, the peristome well expanded, brown. Denticles 5, the 
parietal lamella high and strong, a minute angular lamella 
standing near its outer end. Columellar lamella entering 
obliquely, ascending a little. Upper and lower palatal folds 
very long, rising conically in the middle, distinctly showing 
through from the outside, the lower fold being a little stronger 
and more immersed, its position marked by a depression out- 
side. There is no palatal callus. 



118 AMERICAN SPECIES OF VEETIGO. 

Length 1.8, diam. 0.9 mm. ; 5% whorls. 

Length 1.7, diam. 0.9 mm. ; fully 5 whorls. 

Arizona : Mt. Mingus, near Jerome, at about 8500 ft., type 
loc., and on Oak Creek, 40 miles from Jerome, Yavapai Co., 
Ashmun; Bill Williams Mt., Coconino Co., Ferriss; Graham 
Mts., and Black River, Graham Co., Ferriss; Dragoon and 
Chiricahua Mts., Cochise Co., Ferriss & Pilsbry. New Mexico : 
Grants, Valencia Co., A. & J. Bailey, and Bland, Bernalillo 
Co., E. H. Ashmun; on Willow, Whitewater and Silver creeks, 
Mogollon Mts., Ferriss & Daniels; everywhere along the crest 
of the Black Mts., Ferriss & Pilsbry. 

Vertigo coloradensis arizonensis PILSBRY & VANATTA, Proc. 
A. N. S. Phila. 1900, p. 604, pi. 23, f. 9. PILSBRY, Nautilus 
xix, 130 (Grants, N. M.). 

This race, of which large numbers have been studied from 
many localities, is somewhat narrower than V. coloradensis, 
and invariably has an angular lamella when adult. The 
palatal folds are long, and there is no basal or subcolumellar 
denticle. The penult whorl is usually more coarsely and dis- 
tinctly striate than the others. 

22c. Vertigo coloradensis inserta n. subsp. PI. 12, figs. 10, 11. 

The shell is similar to V. c. arizonensis in size, shape and 
sculpture, and in possessing a small angular lamella; but the 
parietal and columellar lamellae are larger and thicker, and 
there is a basal fold. 

Length 1.85, diam. 0.9 mm. ; 5 whorls. 

Length 1.7, diam. 0.9 mm. 

Arizona: many places in the Santa Catalina Mts., the type 
locality being Bear Wallow; J. H. Ferriss. 

It appears to be of general occurrence in the Santa Cata- 
linas, the localities being between 8500 and 9500 ft. V. c. 
arizonensis, which inhabits the same zone in ranges north, west 
and east, was not found in the Santa Catalinas. One quite 
short specimen from Rustler Park, high in the Chiricahuas, 
measuring 1.6 x 0.95 mm. appears to be the same race. 

It differs from V. c. basidens by having an angular lamella. 



AMERICAN SPECIES OP VERTIGO. 119 

23. VERTIGO CONCINNULA Ckll. PL 10, figs. 9, 10. 

Shell ovoid-cylindrical in outline, slightly tapering toward 
the blunt apex ; solid and somewhat opaque, so that the palatal 
folds are usually only dimly seen through from the outside. 
Surface shining, irregularly, obliquely striate, more strongly 
so on the penult whorl. Whorls 5, apical 2 whitish, the rest 
cinnamon (often with numerous irregularly scattered spots 
and flecks of very light buff). Whorls quite convex, the last 
slightly ascending toward the aperture, its latter half very de- 
cidedly flattened on the outer-inferior portion, this part bear- 
ing a moderate or low wavelike crest or ridge behind the lip, 
and then slightly constricted. Umbilical rimation short, im- 
perf orate. Aperture rounded, truncate above; peristome a 
little expanded ; parietal wall bearing a rather strong entering 
parietal lamella in the middle, and usually a smaller angular 
lamella to the right of its outer end ; columella with a strong, 
deep-seated entering lamella ; outer wall with two rather low, 
long palatal folds, the lower one longer. Length 2.1, diam. 
1.2 mm. 

Colorado: Ouster and Summit counties, 6,000-10,000 ft. 
(Cockerell), and other places noted below. New Mexico: 
Beulah, Sapello canyon, T. D. A. Cockerell; Bland, Jemez 
Mts. and Capitan Mts., Ashmun ; Cloudcroft, Sacramento Mts., 
H. L. Viereck; Mogollon Mts., on Turkey, Willow and Silver 
Creeks, Ferriss and Daniels. Arizona: Mt. Mingus, near 
Jerome, Ashmun. 

Vertigo calif arnica Rowell, INGERSOLL, Bull. U. S. Geol. 
Surv. Terr., i, p. 128 (1875) ; Ann. Rep. U. S. Geol. and Geogr. 
Survey Terr, for 1874, Hayden, p. 392, 1876. No description. 
Not of Bowell. Vertigo ingersolli Ancey MSS. in COCKERELL, 
J. of Conch., Leeds, vi, 1889, p. 64 (name only, substituted 
for P. calif ornica Ing. non Row.). STERKI, Nautilus, vi, 1892, 
p. 5, with varieties haydeni Anc. and accedens Anc. (names 
only). COCKERELL, Nautilus, x, 1897, p. 135 (identity with 
concinnula affirmed from part of original lot). Pupa inger- 
solli Ancey MS., COCKERELL, British Naturalist, 1891, p. 101 ; 
reprinted in Nautilus xvi, p. 59. Vertigo concinnula COCKER- 



120 AMERICAN SPECIES OP VERTIGO. 

ELL, Nautilus, x, 1897, p. 135. PILSBRY and VANATTA, Proe. 
A. N. S. Phila. 1900, p. 599, pi. 23, f. 8. J. HENDERSON, Univ. 
of Colo. Studies iv, p. 172 (Animas Valley, Rio La Plata and 
Cunningham Gulch, Cockerell, Naut. x, 135. Summit and 
Custer Counties and Black Lake Creek, Cockerell. "Vertigo 
calif ornica" Ing., Blue River Valley, Los Pinos Agency, S. W. 
of Los Pinos ; Howardsville, Animas Valley and Rio La Plata, 
Ingersoll, 1874, 392. Eldora, Henderson). WHEELER, Nau- 
tilus xxv, 124 (Monte Sano, Madison Co., Ala.). Pupa con- 
cinnula Ckll., PILSBRY, Nautilus, xi, 1898, p. 119; Class. Cat. 
L. Sh. Amer., p. 21 ; Nautilus, xii, 1899, p. 103. 

The dull, rather opaque shell, cylindrical and small, with 
long palatal folds and parietal lamella, separate this from 
V. modesta and its varieties. It approaches V. modesta parie- 
talis, which, however, is larger and smoother, with shorter 
parietal lamellae and palatal folds. The form of modesta 
from Labrador agrees with concmnula in having the penult 
whorl distinctly striate. The larger size, more cylindrical 
shape and presence of an angular lamella distinguish con- 
cinnula from V. coloradensis. V. c. inserta is distinctly 
smaller than c&ncinnula. In Colorado, according to Professor 
Cockerell, it occurs at higher elevations than V. coloradensis, 
between 6,000 and 10,000 feet; but in the Mogollon Mountains, 
N. M., concinnula is found in the same zone with V. colora- 
densis arizonensis. 

This species was first collected by Ernest Ingersoll, who 
identified it with the West Coast V. calif or nica. Ancey de- 
tected the error, and named it Vertigo ingersolU in MS. In 
1891 Professor Cockerell published a brief descriptive note on 
V. ingersolU: "It is allied to coloradensis, but 2 mm. long, 
cylindrical, dull brown, with a half whorl more, and a double 
lamella on the parietal wall." As the number of whorls of 
V. coloradensis had not been stated, and there are two lamellae, 
not a "double lamella," on the parietal wall, I have held this 
note to be insufficient to establish a species (Nautilus xvi, 59). 
The first recognizable description is that of concmmda 
Cockerell, 1897. Since ingersolU is known solely by Cocker- 
ell's several notes, no description of Mr. Ancey 's type having 



AMERICAN SPECIES OF VERTIGO. 



121 



been published, it seems undesirable to revive that name. The 
type of V. concinnula, from near Brush Creek, Ouster Co., 
10,000 ft., is drawn in fig. 9. The two varieties, haydeni and 
accedens, have never been defined. 

There are some forms approaching corpulenta, but con- 
cinnula is more distinctly striate and has longer palatal folds. 

The specimens from Arizona and New Mexico are of a 
clearer, more translucent cinnamon color than those seen from 
Colorado; the crest is generally somewhat better developed. 
Length 2 to 2.3 mm. These southern examples form a transi- 





FIG. 1. VERTIGO CONCINNULA CKLL. 

tion to V. modesta insculpta, the only difference being the 
smaller size and generally longer palatal folds of concinnula. 
According to Mr. Wheeler, the record from Alabama was 
based upon an identification by the writer; but the place is 
so remote from the known range of concinnula, and the en- 
vironment so diverse, that possibly the identification might be 
revised if the specimen was at hand. 

24. VERTIGO ARTHURI (Martens). 

"Shell ovate, striatulate, perforate, of 5 rather swollen 
whorls ; the aperture triangular, outer margin thickened with- 
in ; 2 palatal folds, 2 columellars, the upper one larger, and 
1 strong parietal. Length 1.5, diam. 0.8, apert. 0.5 mm." 
(Marts.) 

Vertigo bollesiana Morse var. arthuri von MARTENS, SB. 
Ges. Nat. Freunde, Berlin, nr. 9, Nov., 1882, p. 140. 

"Little Missouri, Dakota, coll. by Arthur Krause. It dif- 
fers from the type form of the species [V. bollesiana] by the 



122 AMERICAN SPECIES OF VERTIGO. 

strongly thickened outer margin and the somewhat stronger 
dentition" (Marts.). 

The locality is at or near Medora or Little Missouri station 
of the Northern Pacific R. R., in Billings county, North 
Dakota. It occurred with. Euconulus fulvus, Valloma pul- 
chella and gracilicosta, PupUla muscorum and blandi, Gastro- 
copta armifera, "P. pentodon var." (? = #. holzingeri), and 
Succmea lineata W. G. B. V. Z>. arthuri is not known to 
American conchologists, but the description suggests V. colora- 
densis basidens. 

Dr. V. Sterki has sent a note on a Vertigo collected by Mr. 
A. W. Hanham at Winnipeg, Manitoba, which I suspect to be 
identical with arthuri. "Like coloradensis, though one speci- 
men has somewhat of a rounded crest over the palate. There 
is a strong, white callus in the palate, thickest where the 
short upper palatal plica merges into it, and thinner at the 
lower palatal. It does not extend up to the suture or to the 
base. 

"Length 1.6, diam. 0.9 mm. 

"Length 1.5, diam. 0.8 mm." 

Group of Vertigo modesta. 

Moderately large species, 2 to 3 mm. long, without sharp 
striation (except in V. m. insculpta), and never having a basal 
tooth, the tooth formula varying from 0-0-0 to 2-1-2. 

It is a circumpolar Arctic and alpine group, comprising 
many races with few teeth or none. The collections at hand 
are deficient in Siberian and Greenland Vertigines, and I 
am therefore leaving as species various forms which may 
prove to be merely local races of V. modesta, such as V. hoppii, 
V. krausseana and V. arctica. The arctic and subarctic forms 
have teeth as follows: 

Angular. Parietal. Columellar. Palatal. 

V. krausseana .... 1 or trace. 

V. arctica 1 1 Oorl 

V. a. extima 

V. m. ultima 

V. hoppii 1 or 1 or 1 or 

V. modesta ..lorO 1 1 2 






AMERICAN SPECIES OF VERTIGO. 123 

It may turn out that the slight differences in size and teeth 
are insufficient for separating these forms specifically, and the 
subordination of all under V. modesta would not surprise me, 
when series of specimens from Lapland, Siberia, and Green- 
land can be brought together for comparison with the better 
known boreal American forms of V. modesta. 

Key to American species. 

Shell cylindric-oblong or cylindric-ovate ; to 5 teeth. 

V. modesta and allies, nos. 25-29. 
V. rowelli, no. 32. 

Shell ovate, the spire strongly tapering ; California. 
5 small teeth. V. occidentalis, no. 25/. 

Without teeth. V. dalliana, no. 30. 

25. VERTIGO MODESTA Say. PI. 10, figs. 1, 2; page 124, figs. 
1, 2, 2a. 

"Shell dextral, suboval, minutely wrinkled; apex obtuse; 
whorls six; umbilicus distinct; aperture obliquely subovate; 
labium with a prominent compressed semioval tooth equidis- 
tant from the extremities of the labrum, and a somewhat conic 
one rather below the middle of the columella ; labrum not re- 
flected, joining the preceding whorl at its upper extremity 
with a curve; bidentate, lower tooth placed opposite to that 
of the middle of the labium, the other smaller and placed a 
little above. Length less than one-tenth of an inch. Inhabits 
the Northwest Territory" (Say). 

The cylindric-oblong shell is from tawny to cinnamon 
colored, glossy, rather weakly striate, the striation more dis- 
tinct on the middle whorls. The last whorl has a weak crest 
behind the obtuse, brown outer lip, which expands very little, 
and is not noticeably caught in to form a sinulus. Teeth four, 
white : the parietal and columellar lamellae and lower palatal 
fold subequal, short; the upper palatal fold smaller. Length 
2.6, diam. 1.3 mm. ; 5% whorls. 

Labrador, westward, over the crest of the Rocky Mts. 
(Field, B. C.) to Victoria and Nanaimo. Reported locally 
in Maine, Vermont and Connecticut. Loess of Iowa (Iowa 



124 



AMERICAN SPECIES OP VERTIGO. 




10 



AMERICAN SPECIES OF VERTIGO. 125 

City, Des Moines) and Kansas (Phillips Co.). Alaska, abun- 
dant. The typical modesta replaced in the Rocky Mountain 
system and California by various weakly differentiated races. 

EXPLANATION OP FIGURES, p. 124. 
Fig. 1. Vertigo modesta (Say). Photographic copy of 

Gould 's original figure of Pupa decora. 
Figs. 2, 2a. Vertigo modesta (Say). Loess, Iowa City, Iowa, 

no. 11657. 
Fig. 3. Vertigo modesta (Say), form resembling V. arctica 

Wallenb. Length 2.6 mm., Norton Bay, Alaska, no. 79561. 
Figs. 4, 4&. Vertigo modesta ultima Pils. Norton Bay, 

Alaska, no. 79562. 
Figs. 5, 5a. Vertigo modesta parietalis Anc. Wickersham's, 

Miller Peak, Huachuca Mts., Arizona, no. 97509. 
Figs. 6, 6a. Vertigo hoppii Moller. Photographic copies of 

Moerch's figures. 

Fig. 7. Vertigo occidentalis St. Type specimen. 
Figs. 8, 8b. Vertigo modesta parietalis Anc. Bluff Lake, 

San Bernardino Mts., Cal., no. 104659. 
Fig. 8a. V. m. parietalis Anc. Cienaga below Bluff Lake, 

no. 105167. 
Fig. 9. Vertigo modesta castanea St. Mouth of Big Arroyo, 

Kern R., Tulare Co., Cal., no. 115204. 
Fig. 9a. V. m. castanea. Wood's Creek, Tulare Co., no. 

115203. 
Fig. 9b. V. m. castanea. Onion Valley, Kearsarge Pass, 

Inyo Co., Cal., no. 115199. 
Fig. 10. Vertigo modesta corpulenta (Morse). Photographic 

copy of Morse's figure. 

Pupa modesta SAY, Long's Second Expedition, appendix, 
1824, p. 259, pi. 15, fig. 5. Vertigo modesta (Say), PILSBRY 
& VANATTA, Proc. A. N. S. Phila., 1900, f. 600, pi. 23, f. 2, 3, 
6. WHITEAVES, Ottawa Naturalist 1905, 171 (Kananaskis, 
Alberta). JOHNSON, Fauna of New England, 13, Mollusca, 
1915, p. 213 (Cape Elizabeth, Maine; Stonington, Conn.). 
HANNA & JOHNSTON, Univ. of Kansas Science Bull, vii, 1913, 
p. 120 (Pleistocene of Phillips Co., Kansas) . Ball, Alaska, 
Land and Freshwater Mollusks, p. 29, Harriman Alaska 
Exped. xiii, 1905 (in Alaska at Killisnoo, Chilkat and Chilkoot 
valleys, Pyramid Island, Portage Bay, Dyea, Klukwan, Point 
Romanof at the Yukon delta; St. George Island! St. Paul 



126 AMERICAN SPECIES OF VERTIGO. 

Island! Unalaska! Booluk Island, Unalga Pass! Akutan 
Island! Popof Island, Shumagins, St. Paul, Kadiak Island! 
Orca, Prince William Sound! Yakutat Bay! Berg Inlet, 
Glacier Bay ! Muir Inlet !). Pupa decora GOULD, Proc. Boston 
Soc. N. H. ii, 1848, p. 263, fig. in text (region of Lake Su- 
perior) . BINNEY, Terr. Moll, v, 1878, p. 201, pi. 71, f. 3 
(Fort Resolution, Great Slave Lake). Vertigo decora Gld., 
MORSE, Amer. Naturalist i, 1868, p. 670 (Ascutney, Vt., L. L. 
Thaxter). REINHARDT, S. B. Ges. Nat. Freunde, Berlin, 1883, 
p. 39 (Portage Bay, Killisnoo, Katlrachia, Kluguan, lower 
Deja valley and on Pyramid Island, Krauze). 

An imperforate cylindric-oblong or somewhat ovate shell, 
with short, rather small teeth arranged in form of a cross, 
or not infrequently the upper palatal fold is wanting, es- 
pecially in examples from the loess of Iowa. It is widely 
distributed in the Dominion of Canada and Alaska, and in 
the loess formation of Iowa and Kansas. 

V. modesta differs from typical V. m. corpulenta by the 
more cylindric shell, with one whorl more, but intermediate 
individuals or lots occur in the West. No specimens having 
a distinct angular lamella are known from east of the Rocky 
Mountains. 

There are numerous forms and mutations, some of them 
apparently subspecies characteristic of definite areas; others, 
such as parietatis, often occur associated with various races 
in the same colonies. The subspecific taxonomy is more or 
less arbitrary, and the number of forms worth recognition 
by name will vary with the material studied and the observer 
until collections fairly covering the range of the species are 
available. 

"This is the most abundant and widely distributed species 
in the north country. ' ' The type locality of modesta was 
somewhere near or west of the western end of Lake Superior. 
P. decora also came from the region of Lake Superior. The 
two names were evidently applied to exactly the same race. 
A photographic copy of Gould's figure of P. decora is given 
on p. 124, fig. 1. 

It is a variable snail. A specimen from Labrador meas- 



AMERICAN SPECIES OP VERTIGO. 127 

ures, length 1.9, diam. 1.2 mm., and is closely striate on the 
penult whorl. 

Specimens from the loess of Iowa are quite variable in 
form and size. Those figured measure : 

Length 2.65, diam. 1.45 mm. (page 124, fig. 2a, Iowa City). 

Length 2.4, diam. 1.35 mm. (page 124, fig. 2, Iowa City). 

Length 2.3, diam. 1.2 mm. (plate 10, fig. 2, Des Moines). 

The smallest of these, pi. 10, fig. 2, is rather strongly striate, 
much as in V. concinnula. 

It has also been found by Mr. A. A. Hinkley in loess at 
New Harmony, Indiana, according to Dr. Sterki. 

In a series from Dyea valley, Alaska, the size varies from 
2.4 x 1.4 to 2.65 x 1.4 mm. The crest is rather strong. The 
teeth vary from typical to slightly larger, and in some of 
them a small angular lamella is added. The lot is inter- 
mediate between modesta and mut. parietalis, or rather, it in- 
cludes both forms. A specimen was figured in Proc. A. N. S. 
Phila. 1900, pi. 23, fig. 2. 

Specimens taken on Norton Bay, Alaska (R. C. McGregor) 
are deficient in teeth, an upper palatal fold being absent 
(page 124, fig. 3), the parietal and columellar lamellae small. 
Length 2.4 to 2.6, diam. 1.4 mm. It agrees rather closely 
with the description and figures of V. arctica Wallenb., de- 
scribed from Lapland, and I have no doubt is identical with 
the form listed as arctica from Port Clarence by Westerlund 
(Vega-Exped. Vet. Arbeten, iv, 163) ; yet I can but regard it 
as a form of modesta, in view of the fact that it is in the 
V. modesta area, and that a similar deficiency in teeth occurs 
in forms of modesta found in the loess of Iowa and in Cali- 
fornia. I do not here consider the question whether V. 
arctica is distinct from modesta for the reason that I have no 
Lapland examples of arctica. 

Pupa borealis Morelet is generally admitted to be identical 
with decora (modesta). A translation of the description 
follows: Shell rimate, ovate-oblong, glossy, diaphanous, ful- 
vous-corneous, regularly striate under the lens; whorls 6, a 
little convex, the last compressed at base, forming a moder- 
ate excavation. Aperture somewhat rounded oval, moderate, 



128 AMERICAN SPECIES OF VERTIGO. 

four-toothed: 1 pliciform tooth deep on the parietal wall, 1 
columellar, the rest smaller, in the palate. Peristome simple, 
straight, the columellar margin shortly dilated above. Length 
3, diam. 1.5 mm. Kamchatka (Morelet). 

Pupa borealis MORELET, Journ. de Conchyl. vii, 1858, p. 9. 
PPR., Monogr. vi, p. 311. W. G. BINNEY, Terr. Moll, v, 
1878, p. 211; Third Supplement, Bull. M. C. Z. xix, 1890, 
p. 185, fig. in text ; Man. Amer. Land Sh. p. 483. 

Binney's figure of a specimen from Petropaulovski shows 
no upper palatal fold, and might be considered identical with 
V. arctica-, yet the teeth are so variable in these derivatives 
of modesta that a single specimen is inconclusive. 

Wallenberg's figures of the form he calls V. shuttleworthiana 
from Lapland look a good deal like modesta, the shells being 
much larger than the true shuttleworthiana. See Malak. Bl. 
v, pi. 1. 

25a. Vertigo modesta ultima n. subsp. Page 124, figs. 4, 4a. 

On the north shore of Norton Sound Mr. R. C. McGregor 
obtained an entirely toothless form, having the shape and 
striation of V. modesta, and probably merely a terminal mem- 
ber of the mutation-series represented by the three-toothed 
form found elsewhere on Norton Sound. Like the latter, it 
is not distinguishable from the terminal forms of V. m. cas- 
taneai except by the remote locality. It is smaller than 
V. arctica extima of central Siberia and differs from V. 
krausseana by having no teeth and by the somewhat greater 
size. 

Length 2.6, diam. 1.4 mm. 

Length 2.5, diam. 1.5 mm. 

25&. Vertigo modesta mut. parietalis Ancey. PI. 10, fig. 4. 

" Shell having two teeth on the parietal wall" (Ancey). 
The shape varies from that of corpulenta to about that of 
modesta ; whorls about 5. Teeth 5, there being an angular 
lamella; and the others are larger than in typical corpulenta. 
The surface is somewhat striate, as in modesta and corpulenta. 
Length 2.45, diam. 1.3 mm. ; 5% whorls. 



AMERICAN SPECIES OF VERTIGO. 129 

Rocky Mountain region etc., the type locality Ogden 
canyon, Utah. 

Pupa corpulenta Morse var. parietalis ANCEY, Conchologists 
Exchange ii, Dec. 1887, p. 80 (Ogden canyon, Utah). Vertigo 
modesta parietalis Anc., PILSBRY & VANATTA, Proc. A. N. S. 
Phila. 1900, p. 601, pi. 23, f. 1. HENDERSON, Univ. of Colo. 
Studies iv, p. 172 (Delta and Ouster counties, Black Lake 
Creek, Cockerell; North Park, Barber). COCKERELL, Nau- 
tilus xxv, 59 (Tolland, Colo.). BERRY, Nautilus xxix, p. 127 
(Big Snowy Mts., Mont.). 

In the Sierra Nevada counties of California V. modesta and 
parietalis appear to be rather abundant. They were collected 
in the valleys of the San Joaquin and King's rivers, Bear 
and Fish Creeks, Fresno county, in many places by Mr. 
Ferriss in 1917. In some lots four-toothed forms occurred 
with parietalis, the latter in the majority. 

Lots from Pumice Flats, San Joaquin river, Bear Creek and 
Grouse Meadow, Kings river, are quite distinctly striate, much 
as in the Arizonian insculpta. In one lot from Simpson's 
meadow, Kings river, there are apparently adult shells having 
5, 4 and 2 teeth (columellar and lower palatal). By individ- 
uals these specimens could be referred to parietalis, modesta 
and castanea. 

Other Californian forms of the species are noticed under 
occidentalis and castanea. 

The size and shape are variable in the same lots. The 
shorter shells having 5 whorls or even less, the larger fully 
5~y 2 whorls. 

Length 2.4, diam. 1.4 mm., Miller Pk., Huachucas (p. 124, 
fig. 5). 

Length 2.55, diam. 1.3 mm., Miller Pk., Huachucas (p. 124, 
fig. 5a). 

Length 2.4, diarn. 1.3 mm., Boise Co., Idaho. 

Length 2.7, diam. 1.35 mm., Boise Co., Idaho. 

This form is far more generally distributed than the typical 
corpulenta. In some places, as in in Ogden Canyon, the type 
locality, the two occur together; but in most localities all of 
the adult shells are parietalis. As forms with the parietalis 



130 AMERICAN SPECIES OF VERTIGO. 

teeth, occur in some places associated with both corpulenta 
and with shells having the contour of typical modesta, the sub- 
specific status can hardly be allowed this form. I conclude 
that parietalis is the more primitive stock, and corpulenta a 
mutation thereof, which has not obtained so wide a distribu- 
tion. Typical V. modesta is a more cylindric form which has 
mutated from long parietalis in the same way by loss of the 
angular tooth; but as the five-toothed stock is not found east 
of the Rocky Mountains, where modesta has a wide range, the 
racial distinction may perhaps be retained. 

Fig. 4 is from an Ogden Canyon specimen. 

Professor Cockerell has reported a form from Tolland,. 
Gilpin Co., Colorado, under the name V. m. parietalis, with 
the following note "Rather small for parietalis, but over 
2 mm. long; palatal plicae long, as is concinnula; shell clear 
chestnut ; aperture strongly elbowed above. This is apparently 
a distinct race, between parietalis and concinnula, but hardly 
recognizable by a separate name. Ancey's name' ingersolli 
certainly included such forms as this, and could be so re- 
stricted without much risk of error" (Nautilus vol. 25, Sept., 
1911, p. 59). 

See also under V. m. castanea, p. 134, for notes on parietalis 
forms of the San Bernardino Mts., CaL, figured on p. 124, 
figs. 8, So, 86. 

25c. Vertigo modesta corpulenta (Morse). PL 10, fig. 3. 

Shell rimate perforate, elongate ovate, finely striated, 
polished, translucent, dark olive brown, apex round, obtuse; 
whorls four, convex, tumid, wider at the base, aperture large, 
sub-circular, with four obtuse teeth, one on the parietal mar- 
gin, one on the columellar margin, and two on the labrum; 
peristome slightly thickened and reflected. Length, .10 inch ; 
breadth .06 inch (Morse). 

Nevada: Little Valley, Washoe Co., type loc., on the east 
slope of the Sierra Nevada, 6500 ft. above the sea, R. E. 
Stretch. 

Isthmia corpulenta MORSE, Am. Lye. Nat. Hist. N. Y., viii, 
1865, p. 210, f. 7. Pupa corpulenta Morse, BINNEY, Land and 



AMERICAN SPECIES OP VERTIGO. 131 

F.-W. shells of N. A. i, 1869, p. 238; Terr. Moll, v, p. 201; 
Man. Amer. L. Sh., 1885, p. 172, f. 163. Vertigo modesta 
corpulent a (Morse), PILSBRY & VANATTA, Proc. A. N. S. 
Phila. 1900, p. 601, pi. 23, f. 7. HENDERSON, Univ. of Colo- 
rado Studies iv, p. 172 (Colorado: Los Pinos Agency, Inger- 
sollj Center and Delta counties, CklL). 

Decidedly more obese than V. modesta, of only about 4% 
whorls. The crest behind the lip is distinct, and the four 
teeth are short, the lower palatal being tubercular or very 
short. Usual length 2.2, diam. 1.35 mm., to 2.45 x 1.4 mm. 
A photographic copy of Morse's figure is given on page 124, 
fig. 10. 

It is not known whether corpidenta occurs as a pure race. 
In a few small lots examined there are no parietalis, but all 
of the large lots available containing corpulenta, have 
parietalis also. 

25d. Vertigo modesta insculpta n. subsp. PL 10, figs. 12, 13. 

The shell is similar to V. m. parietalis in teeth, but differs 
by the distinct, rather sharp striation of the penult and ante- 
penult whorls, sometimes also the last. Length 2.6, diam. 
1.35 mm. 

Arizona: Bill Williams Mt., Coconino Co., Ferriss; Santa 
Catalina Mts. in many localities, 9000-9500 ft., the types from 
Mt. Lemon, 9500 ft. ; White Mts., Apache Co. ; Blue River and 
Rim of Blue Mts., Graham Co.; Chiricahua Mts., at head of 
Cave Creek and Long Park, about 8000 ft., Ferriss. New 
Mexico : Black Range, in the forest zone, Ferriss & Pilsbry. 

It differs from the southern form of V. concinnula by the 
larger size and typically shorter palatal folds, yet there is a 
certain amount of intergradation in both characters. In the 
Black Range, especially, there are often short specimens, down 
to 2.25 mm. long. Each of the ranges which ascends into 
the zone of coniferous forest appears to have either a smaller 
(concinnula) or a larger (inscidpta) stock, in the average 
distinguishable, though individuals are often perplexing. The 
distinction is rather finely drawn, yet such as it is, has been 
reached after examining some thousands of fresh shells. 



132 AMERICAN SPECIES OP VERTIGO. 

Specimens from Bill Williams Mt., in northern Arizona are 
particularly sharply striate. They were recorded as V. con- 
cinula in Proc. A. N. S. Phila. 1911, 197, but on account of 
their size, 2.4 to 2.5 mm. long, 1.4 wide, they appear more 
properly placed here. The locality is far north of others 
known, yet the whole intervening region is still unexplored 
for shells. 

The specimens from the Chiricahua range, which were form- 
erly recorded as V. modesta parietalis and V. concinnula ap- 
pear to be better placed here. They measure 2.4 to 2.5 
TTITTI. long. 

At several places in the Santa Catalinas Mr. Ferriss found 
beautiful albino specimens. They have the translucent wax- 
like texture of Gastrocopta pentodon or corticaria. In every 
case they occurred in colonies of cinnamon-brown shells. 

25e. Vertigo modesta castanea Sterki. PL 10, figs. 5, 6. 

' ' Shell oblong or cylindric-oval, glossy, somewhat translu- 
cent; chestnut, sometimes with some whitish stripes. Whorls 
4%-5, the last with a moderate crest behind the lip. Teeth 
very small, placed as in corpulenta, the lower palatal largest, 
columellar usually developed, parietal very small or obsolete, 
upper palatal wanting or minute. Alt. 2.3, diam. 1.4 mm." 
(P. & V.). 

California: Lake Co., HemphUl- Fish Camp, Fresno Co., 
H. Hemphill, type loc. ; Ranger, Panther creek, Wood 's creek, 
Funston meadow on Kern River, Babb Creek falls, Rae Lake, 
Tulare Co., Ferriss ; Wawona meadow, H. Lowe ; Onion valley, 
Kearsarge Pass, Inyo Co., Ferriss Holcomb meadows, east 
of Sugar Loaf Peak, at 8300 ft., San Bernardino Mts., S. B. 
Co., 8. S. Berry. 

Vertigo castanea STERKI, Nautilus vi, 1892, p. 5 (Lake Co. ; 
name only). Pupa castanea Sterki, PILSBRY, Nautilus xi, 
1898, p. 119 (name only). Vertigo modesta castanea Sterki, 
PILSBRY and VANATTA, Proc. A. N. S. Phila. 1900, p. 602, pi. 
23, f. 4, 5 (Fish Camp). LOWE, Nautilus xxx, p. 95 (Wawona 
Meadow, Cal.). 

This form was recorded from Lake Co. by Dr. Sterki, but 



AMERICAN SPECIES OP VERTIGO. 133 

without description. It was first described and figured from 
Fresno Co. specimens, the type being no. 11655 A. N. S. P. 
(pi. 10, fig. 6). 

The typical form has three teeth, parietal, columellar and 
lower palatal; but there are also specimens in the same lot 
which lack the palatal, and one without any teeth. 

In a lot from Wood 's creek, Tulare Co., there are specimens 
having 3 teeth, 4 teeth (a minute angular lamella being 
added), and no teeth (page 124, fig. 9<z), though the shells 
seem otherwise to be equally mature. As the teeth appear 
before the lip is fully formed, this variation is clearly not a 
matter of age. 

A very minute but distinct upper palatal fold is seen in 
some specimens, as no. 115204, from mouth of big arroyo, Kern 
Eiver, Tulare Co. (page 124, fig. 9). Others of the same lot 
have three teeth, and in others a minute angular lamella is 
developed also. The teeth are best developed in specimens 
from Eae Lake (pi. 10, fig. 5), some of which would be re- 
ferred to parietalis were they not associated with others more 
or less deficient in teeth, or without teeth, but otherwise 
similar. In this lot the shell is largely whitish, or is streaked 
with white. 

Some lots contain among normally proportioned shells, a 
few short examples such as page 124, fig. 9&, but having the 
crest and lip of adult character. Two from Onion valley, 
Kearsarge Pass measure: 

Length 2.5, diam. 1.4 mm. (normal). 

Length 2.3, diam. 1.5 mm. (page 124, fig. 9&). 

The more cylindric shape and the distinct crest behind the 
lip distinguish this race from V. occidentalis. I have not 
seen specimens from Lake Co., but the other localities are well 
represented in the collection of the Academy. 

V. m. easterner in some of its forms, does not seem distin- 
guishable from V. arctica and V. m. ultima so far as I can see ; 
yet as it has some different mutations, the localities are re- 
mote, and none have been found in the intervening territory, 
it may belong to an independent, parallel line of differentiation 
of the modesta-parietalis stock. On the other hand, there may 



134 AMERICAN SPECIES OP VERTIGO. 

possibly be unknown intermediate links in the distribution 
between Alaska and California. 

Perplexing forms are found in the occidentalis territory of 
the upper San Bernardinos, where Mr. S. S. Berry has col- 
lected many specimens, such as those listed by him as V. m. 
parietalis (Nautilus xxx, 38). Figs. 8, 80, 86, on page 124 
represent specimens having the subcylindric shape of 
parietalis; there is a very small crest close behind the lip 
(smaller than in parietalis, but larger than in typical occiden- 
tals'). The teeth vary from quite as small as in the type of 
occidentalis, or as in the most fully toothed forms of V. m. 
castanea, to as large as in some parietalis; and in some ex- 
amples which seem to be adult, there is no angular lamella. 
The columellar lamella is usually deeply immersed. The base 
is minutely perforate, or in some specimens scarcely so. 
Length 2.5, diam. 1.35 mm., or slightly smaller. It seems to 
be an intermediate stage between parietalis, castanea and 
occidentalis, some individuals resembling one, others another 
of these races. 

25/. VERTIGO OCCIDENTALS Sterki. PI. 10, fig. 11; page 124, 
fig. 7. 

The shell is perforate, and shortly rimate, globosely ovate, 
the spire being convexly conic, summit obtuse ; decidedly wider 
at the last whorl than at the penult ; of a dull hazel color, some- 
what glossy, subopaque when adult, of 4% rather convex 
whorls, the last having a small external dent over the lower 
palatal fold, but no ridge or crest behind the peristome. The 
aperture is rather large, obstructed by five teeth : parietal la- 
mella short, a very small angular lamella near it; columellar 
lamella about equal to the parietal, rather deeply placed ; both 
palatal folds short, the lower slightly larger. No palatal 
callus. The outer lip is thin, narrowly expanded, projecting 
forward above the middle ; columellar margin broadly dilated 
and reflected. 

Length 2, diam. 1.25 mm. ; 4y 3 whorls (type, p. 124, fig. 7) . 

Length 2.2, diam. 1.4 mm., aperture 0.95 mm.; 4% whorls 
(topotype). 



AMERICAN SPECIES OP VERTIGO. 135 

California: San Bernardino Mts., at Bluff Lake, 7,550 ft., 
type loc. ; also Bluff Lake cienaga, in the cienaga just north, 
and along the "New England Trail," 7500 ft. ; cienaga west of 
Green Valley, 6900 ft., S. S. Berry. 

Vertigo occidentalis STERKI, Nautilus xxi, Dec., 1907, p. 90. 
PILSBRY, Nautilus xxi, Apr., 1908, p. 133, pi. 11, f. 5. 
BERRY, Nautilus xxiii, Nov., 1909, p. 75. 

The original description was from a single specimen, which 
seems not quite mature, drawn in page 124, fig. 7 (no. 1860 
coll. Berry). An older shell from the type locality, pi. 10, 
fig. 11, has the teeth somewhat stronger, particularly the colu- 
mellar lamella, which is a very low nodule in the type specimen. 

The more conic shape and more distinctly perforate base, 
as well as the smaller teeth, distinguish occidentalis from 
V. m. parietalis-, the columellar lamella, too, appears to be a 
little more immersed than in parietalis, and there is no crest 
behind the peristome. 

I have not been able to satisfy myself about the status of this 
snail. On account of the ovate, tapering shape, I am dis- 
posed to give it specific rank, as Dr. Sterki has done ; yet there 
appear to be some equivocal forms connecting with V. m. 
parietalis, which influenced me to use the subspecific form. 
No doubt the examination of large series from the original 
locality will clear up this uncertainty either demonstrating 
intergradation or a constant difference. 

26. VERTIGO HOPPII (Moller). Page 124, figs. 6, 6a. 

"Shell dextral, cylindric, obtuse, smooth; columella two- 
toothed. Length 1.2 lines" (Moller). 

"Shell subperforate, cylindric-ovate, thin, very delicately 
striatulate, with a rather glossy, pellucid brown (often white) 
epidermis; spire convexly-conic. Whorls 5 or 7%, convex, 
suture deep, ascending in front; columella with an obtuse 
fold ; parietal wall having a compressed, almost median fold ; 
aperture subtriangular, the peristome a little reflected, the 
outer lip moderately arched, with a very obsolete denticle in 
front of the middle. Length 2%, diam. 1 mm." (Moerch). 

Greenland: Nisik, in the Kusukfjord; Igaliko; different 



136 AMERICAN SPECIES OF VERTIGO. 

places in Amaraglik, in the small willow thickets by the ruins ; 
Ekalluit at about 400-500 ft. above the sea (Holler). 

P. [upa] hoppn MOLLER, Index Molluscorum Groenlandiae, 
1842, p. 4. PFEIFFER in Kiister, Conchyl. Cab. p. 163, pi. 19, 
f. 29, 30; Monogr. ii, 328; iii, 536; viii, 366. BINNEY, Bost. 
Journ. vii, p. 147, pi. 7, f. 2. WESTERLUND, Malak. Blatter 
xxii, 1870, p. 57. W. G. BINNEY, Terr. Moll. vol. v, 1878, p. 
198, f. 102? (Greenland; Anticosti I.). Pupa (Vertigo) 
hoppii Moll., MOERCH, Amer. Journ. Conch, iv, 1868, p. 30, 
pi. 3, f. 6-9. Vertigo hoppii Moller, DALL. Alaska, Land and 
Fresh Water Moll., p. 29. Pupa steenbuchn BECK, Verz. Kiel., 
1847, p. 76 (nomen nudum). 

I have not seen this species, which evidently stands close to 
V. modesta, differing by the more reduced palatal folds. 
Morch notes that "it looks most allied to Pupa arctica 
Wallenb. ' ' He states that ' * the figure of Kiister is very bad. ' ' 
Those he gives, from pen drawings by Moller, do not look much 
better. Some show parietal and columellar lamellae, others 
none. Two of these figures are copied in my figs. 6, 6a. 

A var. with "the shell hyaline, glossy, white, destitute of 
epidermis," based upon "several live specimens with and 
without teeth, at Amaraglik, ' ' is noted by Morch. 

The localities Ungava, Labrador, and Anticosti Island are 
doubtful, and may perhaps refer to some form of V. modesta 
Say. 

Westerlund states that an example of this species from 
Greenland, in the Berlin Museum, is much larger than P. 
arctica Wallenb., almost larger than P. laevigata Kok., has a 
far more rounded aperture, and there is no trace of a tooth 
in the palate. 

27. VERTIGO KRAUSEANA (Reinhardt). 

Shell dextral, long-ovate, rimate, reddish brown, but little 
shining, the surface closely and finely striate under the lens. 
Whorls 5, convex, parted by a deep suture, the last three of 
about equal width, but gradually and regularly increasing in 
height, the last whorl forming about two-fifths the total height 
of the shell, not ascending to the aperture. The aperture is as 



AMERICAN SPECIES OF VERTIGO. 137 

high as wide, the columellar border nearly vertical, reflected 
above, dilated inwardly; outer margin somewhat angular to- 
wards the insertion, the basal margin semicircular. On the 
parietal wall a weak, short tooth stands deep within, behind 
the callus connecting the lip-margins, while the columella is 
toothless. Behind the aperture there is sometimes a very weak 
indication of a callus [crest] parallel to the outer margin. 
Length 2.2, diam. 1.2 mm. (Reinh.). 

Siberia: Chukchi Peninsula, at Pooten (Poot) ; branches of 
St. Lawrence Bay; Ratmanof Harbor (Krause). Alaska, at 
Chilkat Inlet. 

P. [upa] krauseana REINH ARDT, S. B. Ges. Nat. Freunde, 
Berlin, 1883, p. 38. Vertigo krauseana Reinh., DALL, Alaska, 
Land and Fresh Water Moll., p. 32 (Chilkat Inlet?). 

"Only a few examples show a weak, tubercular eleyation 
on the columella. The species stands near the Greenlandic 
P. hoppei Moller, in dentition, but the latter is distinguished 
by the more plump, ventricose shape, paler color, smoother 
and glossy surface and greater transparency" (Reinh.). 

This appears likely to prove merely a race of V. modesta, 
but as it is not known to me autoptically and is smaller than 
any Alaskan specimens I have seen, it is here allowed specific 
rank. I have seen modesta and modesta castanea as short as 
2.3 mm., but they were associated with larger specimens. 

28. VERTIGO DALLIANA (Sterki). PI. 11, fig. 1. 

Shell conic or ovate-conic, of greenish-horn color, transpar- 
ent, finely irregularly striate in the lines of growth, polished ; 
whorls 4%, well rounded, with deep suture, rather rapidly in- 
creasing, the last occupying about % of altit., towards the aper- 
ture somewhat ascending on the penultimate. Aperture la- 
teral, somewhat oblique, subovate with just perceptibly flat- 
tened palatal margin; margins approximate, the ends pro- 
tracted; peristome shortly but decidedly expanded, with a 
very fine thread-like lip near the margin, the same continuing 
as a very fine callus on the apertural wall inside of the line 
connecting the ends of the margins ; palatal wall quite simple ; 
no lamellae (Sterki). 



138 AMERICAN SPECIES OF VERTIGO. 

Length 2.1, diam. 1.35 mm. 

California: near Clear Lake, Lake Co., Hemphill. 

Pupa dalliana STERKI, Nautilus iv, June, 1890, p. 19 ; Au- 
gust, 1890, p. 39, pi. 1, f. 2. W. G. BINNEY, Fourth Suppl. 
Terr. Moll, v, in Bull. Mus. Comp. Zool. xxii, 1892, p. 195, fig. 
in text. Vertigo (Haplopupa) dalliana Sterki, PILSBRY, 
Nautilus xi, 1908, p. 119. 

"The specimens before me were fifteen, fresh, remarkably 
uniform in their whole appearance ; all were more or less cov- 
ered with a dark brown, hard crust of slime and dirt, generally 
thickest around the aperture. Doubtless this coating is done 
'purposely' by the animals, as in many other species also. 
When cleaned, it shows about the size and shape of a well- 
grown Vertigo ovata, Say, but by a good eye or under a glass 
is at once recognized as something else, by the rounded aper- 
ture and the absence of lamellae" (Sterki). 

The color is grayish-olive. There is a deep and rather long 
crevice but no umbilical perforation. It resembles V. m. occi- 
dentalis closely in shape, but is slightly smaller, greener in 
hue, and toothless. V. m. castanea has a more oval, less conic 
shape. V. dalliana is perhaps a toothless member of the V. 
modesta group, the end product of a tooth-degeneration series. 
In the modesta and calif ornica groups there are many forms 
showing various stages of this process. - V. m. occidentalis ap- 
pears to be the most nearly related species. 

By a typographical or other error the length was given as 
1.2 mm. instead of 2.1, in the original description. The type 
specimen is here figured, no. 416 Sterki collection. 

Group of Vertigo calif ornica. 

Nearctula STERKI, Nautilus vi, 1892, p. 5, type V. calif ornica. 

Vertigines of cylindric shape, without crest or grooves over 
the palatal folds, and with no palatal callus ; typically having 
parietal and columellar lamellae, upper and lower palatal 
folds, but in some forms the teeth have been partly lost, only 
the parietal remaining, or sometimes the aperture is wholly 
toothless. 

These forms are approached so closely by some of the 



AMERICAN SPECIES OP VERTIGO. 139 

V. modesta group that a distinction may be impracticable; 
yet the want of any trace of a crest behind the lip may serve 
to distinguish the present series from most of the modesta 
group. Moreover, the two groups differ so widely in zonal 
distribution that a different ancestry is suggested: the cali- 
fornica group inhabits low levels, while the modesta group is 
restricted to the colder Canadian and Transition zones, at 
higher elevations in California, descending to low levels only 
in the north. 

Key to species and subspecies. 

1. Sculpture of rather strong oblique riblets. 2. 
Surface striate but not distinctly ribbed; about 2.5 mm. 
long. Four well developed teeth. V. rowelli, no. 30. 

2. Aperture having 4 teeth, at least the columellar, parietal 
and lower palatal well developed. 3. 
Aperture having 3 small teeth. Monterey. 

V. c. trinotata, no. 29a. 
Aperture having 1 tooth, the parietal, or none. 6. 

3. About 2.5 x 1.3 mm. ; parietal lamella not entering deeply. 
San Francisco. V. californica, no. 29. 
Shell smaller or narrower; parietal lamella long, entering 
rather deeply. Monterey southward, and the islands. 4. 

4. Shell 2 to 2.5 mm. long. V. c. elongata, no. 29d. 
Shell 1.75 to 2 mm. long; whorls short. 5. 

5. Sculpture strongly developed. S. Catalina and S. 
Clemente. V. c. catalinaria, no. 29e. 
Sculpture weaker and irregular, Cypress Point. 

V. c. cupressicola, no. 29/. 

6. San Diego southward. V. c. diegoensis, no. 29&. 
Placer county. V. c. cyclops, no. 29c. 

29. VERTIGO CALIFORNICA (Rowell). PL 9, figs. 1, 2. 

' ' Shell rimate subperf orate, elongate ovate, thin, dark horn 
colored, with oblique rib-like striae ; apex obtuse ; deep suture ; 
with 5-6 convex whorls, the last a little compressed at the 
aperture; aperture oblique, suborbicular, armed with four 
white denticles: one lamelliform, strongly developed, slightly 



140 AMERICAN SPECIES OP VERTIGO. 

twisted, on the parietal wall, one on the columella, and two 
deeply seated within or near the base of the aperture ; peris- 
tome slightly expanded, columella margin somewhat reflected. 
Long. 2V2, diam. 1 mm." (Rowell). 

California : San Francisco, Rowell and others. 

Pupa californica ROWELL, Ann. Lye. Nat. Hist, of N. Y., 
vii, Jan., 1861, p. 287. BLAND, Ann. Lye. viii, 1865, p. 166, 
fig. 12. BINNEY, Terr. Moll. vol. v, 1878, p. 202, f. 107. 
STERKI, Nautilus iv, May, 1890, p. 8. 

The figures are from paratypes received from Mr. Rowell. 

V. californica differs from all other American Vertigos by 
its sculpture. The initial 1% whorls are smooth (showing the 
usual minute granulation under the microscope) ; the next 
whorl is finely but sharply rib-striate, after which the riblets 
become coarse, more oblique, and on the last whorl about 
one-tenth of a mm. apart. There is no trace of a crest or con- 
traction behind the narrowly expanded outer lip ; columellar 
border is well dilated. Both ends of the lip are somewhat 
calloused in fully adult specimens. 

In the typical subspecies the four teeth are well developed 
as a rule, but in some individuals the upper palatal fold is 
quite small, or even wanting in rare individuals. Dr. Sterki 
mentions seeing traces of an angular lamella in some specimens. 

Length 2.65, diam. 1.35 mm. ; 5% whorls. 

Length 2.5, diam. 1.3 mm. 

The size and shape are rather variable in a lot from San 
Francisco received from Dr. R. E. C. Stearns: length 2.6, 
diam. 1.4 mm., to length 2.25, diam. 1.25 mm. ; the shorter ones 
ovate, the longer cylindric. 

Besides the typical form, which is known to me from San 
Francisco only, the following local races have been defined : 

29a. Vertigo californica trinotata (Sterki). PI. 9, fig. 9. 

"In size not much different from the type, yet a little 
smaller, and more generally obovate ; the striae are less coarse ; 
the peristome is slightly but distinctly expanded. There is 
no superior palatal lamella, and the three present are small, 
the columellar even a trace, or wanting entirely" (Sterki). 

Length 2.25, diam. 1.15 mm. ; nearly 5 whorls. 



AMERICAN SPECIES OP VERTIGO. 141 

California : Monterey, Hemphill. 

[Pupa calif ornica] var. trmotata STERKI, Nautilus iv, June, 
1890, p. 18. 

The figure is from a paratype. 

29&. Vertigo calif ornica diegoensis (Sterki). PL 9, figs. 8, 11. 

1 * The diminution of the lamellae is going on ; none but the 
apertural is left in this variety, and even that is quite small 
or a mere trace. In size and shape the specimens are not 
much different from the Monterey form, which is an inter- 
mediate one" (Sterki). 

California: San Diego, H. Hemphill, type loc. Lower 
California: San Ramon, C. R. Orcutt. 

[Pupa calif ornica} var. diegoensis STERKI, Nautilus iv, 
June, 1890, p. 18. Vertigo diegensis St., BERRY, Nautilus 
xxx, p. 83 (False Bay near Asher Station). 

In the specimens from San Ramon the parietal lamella is 
sometimes well developed, and there is very rarely a tuber- 
cular palatal fold, as in pi. 9, fig. 8. A few examples have 
no teeth whatever. The least decadent individuals of this 
race have reached about the same stage of tooth-reduction as 
V. c. cyclops, and some specimens are not distinguishable from 
that ; yet as the localities are remote and in regions physically 
diverse, the resemblance is evidently not due to direct relation- 
ship, but to parallel evolution. 

Length 2.65, diam. 1.35 mm. ; 5^4 whorls. 

Length 2.35, diam. 1.25 mm. ; 5 whorls. 

29c. Vertigo calif ornica cyclops (Sterki). PL 9, fig. 3. 

"Large, conic or ovate conic, or turriculate, umbilicated, 
rib-like striae rather strong ; whorls 5, well rounded, with deep 
suture, the last occupying more than y 2 alt.; aperture sub- 
ovate or nearly circular, margins much approximate and the 
ends protracted, peristome shortly but decidedly expanded; 
lamella one, apertural [parietal], small. Alt. 2.5; diam. 1.5 
mill. For its size, rounded aperture and single lamella I 
would name this form var. cyclops" (Sterki). 

Length 2.7, diam. 1.45 mm. ; 5 whorls. 



142 AMERICAN SPECIES OP VERTIGO. 

California: Rocklin, Placer Co. (25 miles northeast of 
Sacramento), Hemphill. 

[Pupa californica] var. cyclops STERKI, Nautilus iv, June, 
1890, p. 18. 

A paratype is figured. It is slightly larger than Dr. Sterki 's 
type. ' ' Some specimens have no columellar lamella, and some 
have a small palatal fold" (Sterki, in letter). 

29d. Vertigo calif ornica elongata (Sterki). PI. 9, fig. 4. 

"A little smaller and generally more cylindrical than the 
type ; a part are even long cylindrical having the appearance 
of an Isthmia [Truncatellina]. The coloration is somewhat 
paler and the lamellae are well formed" (Sterki). 

Length 2.5, diam. 1.2 mm.; 6 whorls. 

Length 2.25, diam. 1.1 mm. ; 5y 2 whorls. 

Length 2.1, diam. 1.2 mm. ; 5 whorls. 

California: San Clemente Island, Hemphill, type loc. 
Santa Barbara I., Hemphill. 

[Pupa californica] elongata STERKI, Nautilus iv, May, 
1890, p. 8. 

Fig. 4 is from a paratype. The parietal lamella is de- 
cidedly longer than in V. californica, entering deeply. From 
the elongate form figured it varies, in the same lots, to much 
shorter forms. Specimens from Santa Barbara are 2.3 to 
2.4 mm. long, or even smaller, 2 to 2.25 x 1.15 mm. 

This race is intermediate between V. californica and V. c. 
catalinaria. 

290. Vertigo californica catalinaria (Sterki). PI. 9, figs. 5, 6. 

' * Small, rather short, pale horn colored ; shell thin, delicate ; 
rib-like striae less numerous and relatively larger; the whorls 
are less high, which gives the shell a different appearance. 
All lamellae are present and well formed, especially the aper- 
tural [parietal]" (Sterki). 

Length 1.75, diam. 1.05 mm.; 5% whorls. 

California: Santa Catalina I., Hemphill, type loc.; also 
San Clemente I. and Santa Barbara I., Hemphill. 

[Pupa californica] var. catalinaria STERKI, Nautilus iv, 
May, 1890, p. 9. 



AMERICAN SPECIES OP VERTIGO. 143 

' ' One peculiarity is that in about one-third of the examples 
a part of the shell is wanting, always on the side of the aper- 
ture, so that 3 or even 4 whorls are opened. This can hardly 
be accidental, and probably that part of the thin shell is worn 
off by friction in moving" (Sterki). 

This race has four well-developed teeth, the parietal lamella 
being long, as in V. c. elongata, from which it differs by the 
smaller size and shorter whorls; it is also more fragile. A 
paratype is figured (fig. 6), and a longer shell from Santa 
Barbara Island (fig. 5). 

The Santa Barbara Island specimens are somewhat perplex- 
ing. Some are typical catalwaria, 1.75 x 1 mm., but also up 
to 2.15 Trim, long, while there are also stouter shells with them, 
2 to 2.25 mm. long, 1.15 wide, which seem referable to 
elongata. If it were not for these examples, I would consider 
catalinaria specifically distinct. 

29 /. Vertigo calif ornica cupressicola Sterki, n. subsp. PI. 9, 
fig. 10. 

Small, cylindric, with the riblets distant, irregular and 
partly obsolete ; apertural lamellae and folds relatively larger 
than in californica, the parietal lamella long. Length 1.83, 
diam. 1.1 mm. ; barely 5 whorls. 

California: Cypress Point, Monterey, S. S. Berry, J. C. 
Paine. Type 118835 A. N. S. P. 

Closely related to V. c. catalinaria, but differing by the 
weaker, irregular sculpture. In some examples the spire 
tapers a little more than in the figured type. 

30. VERTIGO ROWELLII (Newc.). PL 9, fig. 7. 

Shell perforate, oblong-ovate, dark horn-colored, shining, 
translucent, finely striated; apex obtuse; whorls 5, convex; 
aperture truncately ovate, armed with 4 teeth : one prominent 
and plicate on the columella, 3 deeply seated within the 
aperture; peristome slightly reflected. Long. 2, lat. 1 mm. 
(Newc.). 

California: near Oakland, Newcomb, type loc. ; cienaga 
north of Bluff Lake, San Bernardino Mts., S. S. Berry. 
Oregon : Douglas county, F. H. Andrus. 



144 AMERICAN SPECIES OF VERTIGO. 

Pupa rowelln NEWCOMB, Ann. Lyceum of Nat. Hist. N. Y., 
vii, April, 1860, p. 146. BLAND, Ann. Lye. viii, p. 166, fig. 11. 
-W. G. BINNEY, Terr. Moll, v, 1878, p. 202, fig. 106 (near 
Oakland, Monterey, San Bernardino, El Dorado Co.) ; Man. 
Amer. L. Shells, p. 483 (Alameda Co.). Vertigo (Nearctula) 
rowelli Newc., STERKI, Nautilus vi, 1892, p. 5. 

With the teeth like V. califomica, this species differs by 
its smoother surface. Under a lens there are delicate, irregu- 
larly spaced thread-like striae corresponding to the far stronger 
riblets of V. califomica. The color is somewhat brownish 
olive. Specimens from Douglas county, Oregon measure: 

Length 2.7, diam. 1.5 mm. 

Length 2.65, diam. 1.35 mm. 

A single example was found in a lot of V. a. sanbernardi- 
nensis from the San Bernardino Mts. It measures 2.48 x 
1.35 mm. 

Subgenus VERTILLARIA n. subg. 

Shell oblong, with the columellar lamella in form of a blunt 
vertical plate, other teeth short; outer lip biarcuate. In the 
type species there is no angular lamella or basal fold, and 
the upper palatal fold is minute or wanting. 

Type V. oscariana. Austroriparian, from Florida to Texas. 

The blunt, vertical columellar lamella, parallel to the axis, 
is entirely unlike other American species of Vertigo, being 
comparable to that of Vertilla angustior. I suspect Vertilla 
may be the most nearly related form, though the American 
species has the lower palatal fold much larger than the upper, 
and there is no angular lamella. The deep external furrow 
of the last whorl suggests that an ancestral form had a long 
lower palatal fold, though the present species is rather de- 
generate in teeth. There is no external crest and no palatal 
callus. 

31. VERTIGO OSCARIANA Sterki. PL 8, figs. 8, 10, 11. 

"This is the most peculiar of our species. It is the size of 
milium, but oblong with either end nearly equally pointed, 
the last whorl being considerably narrowed and flattened to- 



PUPILLID^E 



PLATE 6 




12 



13 



PLATE 7 



13 




PUPILLID>E 



PLATE 8 




PUPILLID^E 



PLATE 9 




6 



10 



11 



12 



13 



PLATE 1O 




11 



12 



13 



SECOND SERIES: PULMONATA. 

MANUAL 



OF 



CONCHOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED BY 



GEORGE W. TRYON, JR. 

CONTINUED BY 

HENRY A. PILSBRY, Sc.D 



D EPARTMENT OF USCAi ^ 

SCIENCES OF PHILADELPHIA. 



Part 



PHILADELPHIA 

'UBLISPIED BY THE CONCHOLOGICAL DEPARTMENT 
ACADEMY OP NATURAL SCIENCES 
OF PHILADELPHIA 



AMERICAN SPECIES OF VERTIGO. 145 

wards the subtriangular small aperture. Shell thin, delicate, 
of pale horn color, as is the palatal wall and margin; the 
latter simple and straight, with a very slight, thin callus inside, 
lamellse 3, whitish, rather small: one apertural [parietal], one 
columellar, longitudinal, and the inferior palatal; sometimes 
there is also a very small superior palatal. Length 1.5, diam. 
0.8 mm." (Sterki). 

Length 1.45, diam. 0.8 mm. ; nearly 5 whorls. 

Length 1.53, diam. 0.85 mm. 

Florida: Mosquito Island, Volusia Co., type loc., Oscar B. 
and G. W. Webster; Grassy Key, Eaybon. Alabama: Ever- 
green, Conecuh Co., H. H. Smith. Louisiana: S.-W. cor. 
Madison Parish, Bayou Macon, C. B. Moore. Arkansas : Keller 
Place Landing, Ouachita R., Calhoun Co., C. B. Moore. Texas : 
Guadalupe B. near New Braunfels, Ferriss and Pilsbry. 

Vertigo ascariana STERKI, Proc. A. N. S. Phila. 1890, p. 33 ; 
reprinted in BINNEY, 4th Suppl. Terr. Moll, v, Bull. M. C. 
Zool. xxii, 1892, p. 197, fig. , Nautilus iii, 1890, p. 136 ; iv, 
1890, p. 39, pi. 1, f. 5. PILSBRY, Proc. A. N. S. Phila. 1906, 
p. 147. CLAPP, Nautilus xxviii, 1915, p. 137, pi. 6, f. 8. 

The surface is delicately but rather sharply and regularly 
striate, most conspicuously so on the penult whorl. The colu- 
mellar lamella is rather blunt and thick, the lower end vertical, 
the upper slanting slightly inward. The parietal lamella is 
short and rather high. Figured from the type, no. 60463 
A. N. S. 

It varies in degree of inflation, as the figures show. Dr. 
Clapp has noted that the specimens from Evergreen, Alabama, 
are nearly smooth, showing only obsolete striation. One meas- 
ures 1.45 x 0.8 mm. The Texan form is also nearly smooth, 
1.7 x 9 mm., and some have a thin palatal callus. All of the 
localities given are for specimens in coll. A. N. S. 

It is an isolated species, without near relatives in our fauna. 

Subgenus ANGUSTULA Sterki. 

Angustula STERKI, Proc. U. S. Nat. Mus. xi, 1888, p. 378, 
for V. milium and V. venetzii; Nachrbl. d. m. Ges. 1889, p. 
P. & V., Proc. A. N. S. Phila. 1900, p. 597. 



146 AMERICAN SPECIES OP VERTIGO. 

Angular, parietal and columellar lamellae are strongly de- 
veloped, the parietal long, columellar crescentic, its inner end 
curving downward. Palatal folds strong, the lower entering to 
the dorsal side, its inner end curving downward. Type V. 
milium. 

The peculiarities of Vertigo milium were first fully worked 
out by Doctor Sterki, who considered it related to the Euro- 
pean V. angustior (venetzii) ; but the relation seems to be one 
of partial parallelism rather than of direct affinity. I may 
mention that Dr. Sterki now considers the relationship 
doubtful. 

The two species differ as follows : 

V. milwm. V. angustior. 

Dextral. Sinistral. 

Columellar lamella running Columellar fold spirally as- 
downward inwardly. cending inwardly. 
Lower palatal fold entering Upper palatal fold entering 
deeply. deeply. 

Angustula has some features in common with the Cali- 
fornian species of Sterkia, particularly the well-separated 
angular and parietal lamellae and the descending inner end 
of the columellar lamella; but there is probably no direct 
relationship. 

Dr. 0. Boettger has considered V. ovatula, of the German 
upper Oligocene and lower Miocene, to be closely related to 
V. mitium. I have not seen that species, but as the form 
of the columellar lamella as figured is entirely different in 
V. ovatula and V. milium, the former evidently does not be- 
long to Angustula. 

Two species are known: V. milium, no. 32, in which there 
is a moderate swelling behind the outer lip, and V. bermu- 
densis, no. 33, having a very high, massive crest. 

32. VERTIGO MILIUM (Old.). PL 13, figs. 1 to 7. 

' ' Shell minute, of a globosely-oval form, color a light chest- 
nut; whorls four or somewhat more, obviously wrinkled 
obliquely, rather convex ; apex bluntly rounded ; suture deep ; 



AMERICAN SPECIES OP VERTIGO. 147 

aperture half the width of the last whorl, heart-shaped, the 
apex being at the right posterior angle ; the transverse margin 
is nearly direct; the outer lip is scalloped by an indentation 
of the lip ; the remainder of the margin is regularly arcuated ; 
lip white, slightly everted ; throat with six teeth, two of which 
are on the transverse lip, equidistant ; one, with a tubercle at 
its base, is on the middle of the columella, and nearly at right 
angles with the preceding, and is the largest; a fourth is on 
the indenture of the outer lip, directed between the two on 
the transverse lip ; and two smaller ones, more retired within 
the shell, are equidistant between the two last mentioned; 
umbilicus large and deep. Length less than 1/30 of an inch : 
breadth 1/40 of an inch" (Gould). 

Maine to Florida, west to South Dakota, Grand County, 
Colorado and southeastern Arizona; Tampico, Mexico; 
Jamaica. Type locality Oak Island, Chelsea, near Boston. 

Pupa milium GOULD, Boston Journal of Natural History 
iii, pt. 3, 1840, p. 402, pi. 3, f . 23 ; iv, p. 359, 1843 ; in Binney, 
Terr. Moll, ii, p. 337, pi. 71, f . 1 ; Invert. Mass. p. 187, f . 118. 
ADAMS, Vermont Moll., p. 157. Vertigo milium Gld., W. G. 
BINNEY, Terr. Moll, v, 1878, p. 215, pi. 71, f. 1 ; Man. Amer. 
Land Shells, 1885, p. 332, f. 361. MORSE, Amer. Nat. i, 1868, 
p. 669, f. 65, 66. STERKI, Proc. U. S. Nat. Mus. xi, 1888, p. 
377, pi. 42, f. 10, 13; Nautilus xxix, 123 (Geneva, 0.). 
WALKER, Moll, of Michigan, 1906, p. 515, f. 146, 147 (Newaygo, 
Kent, Bay, Oakland, Washtenaw and Lenawee counties) ; Nau- 
tilus xxxi, 54, 56 (Chester Shoals and Palm Beach, Fla.). 
JOHNSON, Fauna of New England 13, 1915, p. 215 (Me., 
Vt., Mass.). DANIELS, 27th Ann. Rep. Dep. Geol. and Nat. 
Res. Indiana, 1902, p. 632 (Steuben Co., Brookville, Lawrence- 
burg, Dunreith, Ind.). OVER, Nautilus xxix, 91 (Chamber- 
lain, Brule Co., S. Dak.). VANATTA, Naut. xxviii, 11 (Sussex 
Co., N. J.). PILSBRY, Nautilus xiv, 74 (Cape May, N. J.) ; 
xxv, 35 (Atlantic City, N. J.). HANNA, Nautilus xxiii, 95 
(Douglas Co., Ks.). HINKLEY, Nautilus xxi, 77 (Tampico). 
HANHAM, Nautilus x, 101 (Quebec). CHADWICK, Nautilus 
xix, 58 (near Milwaukee). CKLL., Nautilus iii, 21 (Kremm- 
ling, Colo.). SMITH, Nautilus xx, 90 (Otsego Co., N. Y.). 



148 AMERICAN SPECIES OP VERTIGO. 

STUPAKOFF, Nautilus vii, 135 (Allegheny Co., Pa.). PRIME, 
Nautilus viii, 70 (Long Island, N. Y.). GARDNER, Naut. viii, 
76 (Long Island). SARGENT, Naut. ix, 89 (Clearwater, 
Minn.). PLEAS, Naut. vii, 68 (Henry Co., Ind.). Pupa 
millium SOWERBY, Conch. Icon, xx, pi. 18, f. 164. 

Gould's description and figure are not quite correct in some 
points ; evidently the columellar lamella and lower palatal fold 
were not fully seen. The color of the shell is cinnamon or 
paler. The last whorl has an external impression over the 
lower palatal fold; there is a swelling in front of the im- 
pression and below a deeper impression which runs to the lip, 
over the upper palatal fold. The angular lamella is high, 
short, and situated inward from the insertion of the outer 
lip. The parietal is high and long, entering deeply. The 
high columellar lamella enters horizontally at first, then 
turns downward, being crescent-shaped. (Its downward con- 
tinuation was mistaken by Gould for * ' a tubercle at its base " ) . 
The upper palatal fold is long and high, slightly curved. 
Lower palatal is a little immersed, high, thin and enters to 
the dorsal side, where it curves downward (fig. 5). Both 
palatal folds are rather thick and tapering at their outer ends. 
The basal fold is somewhat immersed, short and high. There 
is sometimes a small, tubercular suprapalatal fold. The outer 
lip is somewhat expanded and strongly biarcuate. The 
parietal callus is generally rather thick. 

Length 1.75, diam. 1 mm. ; 5 whorls. Cleveland, Ohio, fig. 1. 

Length 1.38, diam. 0.8 mm. ; 5 whorls. Cleveland, 0., fig. 3. 

Length 1.65, diam. 0.85 mm. Vermont, fig. 7. 

Length 1.4, diam. 0.9 mm.; 4% whorls. Volusia Co., Fla., 
fig. 6. 

V. mUium shows but little variation in the teeth, though in 
some examples the lower palatal fold emerges further than in 
others, and the degree of curvature of its inner end varies 
somewhat. The size and contour of the shell, however, are 
quite variable, as will be seen by reference to the measure- 
ments and figures. In one lot I found the length from 1.38 
to 1.75 mm. 

The shortest specimens seen (but generally associated with 



AMERICAN SPECIES OF VERTIGO. 149 

others of normal shape) are from the Gulf States. In a speci- 
men from ' l Mt. Taylor, ' ' an artificial mound on the St. Johns 
River, south of Volusia, Florida, the diameter is two-thirds 
of the length (pi. 13, fig. 6). Examples from Arizona and 
Tampico, Mexico, appear to be normal in shape and teeth. 

Specimens from Brownstown, St. Ann's, Jamaica, have 
strongly developed teeth, the inner end of the lower palatal 
fold scarcely bent downward, the shape normal. Length 1.2 
to 1.25 mm. 

33. VERTIGO BERMUDENSIS n. sp. PL 13, figs. 8, 9, 10, 12. 

The shell is minute, oblong, gray, irregularly rather weakly 
striate but not polished on account of a microscopic granula- 
tion. Whorls rather strongly convex, the suture appearing 
margined by translucence of the shell; last whorl convex in 
front but becoming flattened and tapering downwards dorsally. 
Some distance behind the aperture there is a very high, mas- 
sive, rounded crest, followed by a deep contraction, in which 
there is a pit behind the prominence of the outer lip. The 
aperture is piriform, having teeth arranged as in V. milium. 
The angular lamella is rather long and emerges further than 
the parietal, which is very long and curved, entering spirally. 
The columellar lamella is somewhat immersed (sometimes in- 
conspicuous in a front view) ; it enters horizontally, then turns 
downward along the axis, much as figured for V. milium. 
The stout upper palatal fold emerges to the palatal callus. 
The lower palatal arises farther inward, is high and lamellar, 
penetrating to the dorsal side where it turns downward 
slightly. The basal fold is absent, or represented by an in- 
distinct callus. The peristome is well expanded and projects 
forward in a rounded "auricle" above the middle of the 
outer margin. 

Length 1.45, diam. 0.8 mm. ; 5 whorls. 

Bermuda: around Church Cave, Paynter's Vale, near 
Tucker's Town (S. Brown, Arthur Haycock). 

This species has about the size and shape of V. milium, 
from which it differs chiefly by the enormous development of 
the oblique crest behind the lip. The much larger V. numel- 



150 VERTIGO OF JAPAN AND EASTERN ASIA. 

lata has a similar crest, the lower palatal fold is long, also, and 
the angular and parietal lamellae similar in position; yet it 
has not the peculiarly shaped columellar lamella of Angustula. 
Pupa eyriesii Drouet, of Guyana, which I have placed in 
the genus Sterkia, has about the size of this species, but as 
nothing was said of the extraordinary crest, it appears to be 
quite different. I have not seen it. 

II. JAPAN AND EASTERN ASIA. 

Vertigo appears to be generally distributed in Japan, the 
maritime province of Siberia and northward to Bering Strait, 
though there are as yet but few records, widely scattered over 
a vast area. The insular species are known by shells brought 
together by Mr. Y. Hirase of Kyoto. For those of the main- 
land we depend upon records which leave much to be de- 
sired in point of illustration, and in some cases, possibly, 
of precision. 

The Japanese species (except V. dedecora) are quite similar 
to forms of North America and Europe, and belong to the 
same groups. Like Cochlicopa and a few other Holarctic 
genera, they mingle in a snail fauna which is otherwise chiefly 
Oriental in its affinities. V. dedecora of the Bonins is related 
to Tertiary species of Europe. 

The mainland species of eastern Siberia are not known to me 
by specimens, and all but one are boreal forms also found in 
Europe or America. The following have been reported. 

V. denudata (Mouss.) Vladivostok. No. 40. 

V. alpestris (Alder). Vladivostok, Amur Valley. See 
no. 73. 

V. borealis (Morel.), Kamchatka (= V. modesta, no. 25). 

V. Jcrausseana (Reinh.). Chukchi Peninsula (and Alaska?). 
No. 27. 

V. arctica (Wallenb.) Chukchi Peninsula? No. 66. 

The last three species, as represented in the Chukchi Pen- 
insula and Alaska are in need of rigid comparison, as it does 
not seem likely that three very similar forms in the same 
region are specifically distinct. 

They belong properly to the cireumpolar fauna. 



VERTIGO OF JAPAN AND EASTERN ASIA. 151 

Key to Japanese species of Vertigo. 

1. Angular lamella curving into and connected by a ridge 
with the upper termination of the peristome (subgenus 
Ptychalaea) . V. dedecora, no. 41. 
Angular lamella, if present, not connected with the upper 
termination of the peristome (Vertigo s. str.). 2. 

2. Having a strong palatal callus and an external crest. 3. 
Without a palatal callus; no external crest. 6. 

3. Shell finely but sharply striate, 1.8 x 1.15 mm. ; no angu- 
lar lamella present. V. hachijoensis, no. 36. 
Shell smoothish, striation very weak if present. 4. 

4. An angular lamella present; basal tooth present, in a 
subcolumellar position ; outer lip noticeably bent in near 
the middle. 5. 
No angular lamella ; crest very close to the lip. 

V. kushiroensis, no. 37. 

5. 2.15 x 1.4 mm. ; a suprapalatal nodule present ; crest 
strong. V. eogea, no. 34. 
1.75 x 1 mm. ; no suprapalatal tooth. 

V. hydrophila, no. 35. 

6. Outer lip straightened in the middle ; smoothish, aperture 
with 4 small teeth ; 1.65 to 1.8 mm. long. 

V. hirasei, no. 38. 
Outer lip biarcuate. 7. 

7. Smoothish, with 1-1-2 teeth. V. japonica, no. 39. 
Striate, with 2-1-2 teeth. V. j. tosana, no. 39&. 

Subgenus VERTIGO s. str. 
34. VERTIGO EOGEA n. sp. PI. 14, figs. 1, 2, 3. 

The shell is rimate, broadly ovate, auburn, fading some- 
what above, glossy, nearly smooth, but under the microscope 
some very weak striae are seen. The whorls are moderately 
convex, the last having a distinct, oblique impression over 
the palatal folds, followed by a rather strong, somewhat 
sinuous crest, which is light-colored and striate; in the con- 
cavity between crest and lip there is a linear impression run- 
ning to the projection of the lip. The aperture has a well 



152 VERTIGO OP JAPAN AND EASTERN ASIA. 

marked sinulus. The parietal lamella is strong and rather 
long ; angular lamella small and short. The columellar lamella 
is situated high, ascends inwardly, and is rather large and 
massive. The two strong palatal folds are rather short and 
near together. The basal fold is smaller, at the junction 
of basal and columellar margins. There is a low supra- 
palatal nodule. The palatal and basal folds are united by 
a strong buff callus. The outer lip is strongly biarcuate, a 
little expanded, the columellar margin being rather widely 
reflected. 

Length 2.15, diam. 1.4 mm.; nearly 5 whorls. 

Japan: Akkeshi, Kushiro, Yesso (Y. Hirase). Type and 
paratypes no. 87899 A. N. S. P., from no. 1307 Hirase coll. 
Also Kashima, Harima (Hirase). 

I formerly identified these specimens with V. hydrophila, 
but they differ by the larger size, paler color, by the develop- 
ment of a suprapalatal fold, and by the shape, if Reinhardt's 
figures are correct. Possibly a comparison of specimens may 
show that it is a subspecies of V. hydrophila. It has some 
resemblance to V. pygmaea. 

A race from Kashima, Harima, in central Japan, differs by 
the less swollen spire and the very small size of the basal 
fold. Length 2.05, diam. 1.3 mm. (Fig. 3). 

35. VERTIGO HYDROPHILA (Reinh.). PL 14, figs. 4, 5. 

Shell dextral, ovate, rimate, glossy, glabrous, striatulate un- 
der the lens, corneous, pellucid; spire conic, obtuse; 4 1 / 
slightly convex whorls, rapidly increasing, the penult large, 
the last rounded, not compressed at the base. Aperture trans- 
versely cordate, 5 to 6 dentate : a strong parietal lamella and 
another [angular] obsolete; 2 columellars, the upper one 
stronger; 2 palatals. The palatal and columellar teeth are 
united by a glossy, ivory-like callus, which appears rufous 
externally. Peristome acute, a little expanded, the margins 
remote, joined by a thin callus, the right margin a little 
produced and bent inward, the columellar margin a little 
reflected. Length 1.75, diam. 1 mm. (Reinhardt). 

Japan: Hakodate, Yesso (Dr. Hilgendorf). 



VERTIGO OF JAPAN AND EASTERN ASIA. 153 

Pupa (Vertigo) hydrophila- O. REINHARDT, Jahrb. d. Malak. 
Ges. iv, 1877, p. 323, pi. 11, f . 6. Pupa hydrophila Reinhardt, 
KOBELT, Fauna Moll. Extramar. Jap., 1879, p. 61, pi. 1, f. 16. 

"Belongs in the kinship of P. antivertigo and stands es- 
pecially near P. ovata Say, which differs, however, by the 
somewhat slimmer shape, the last whorl somewhat compressed 
at base, and by the somewhat stronger teeth" (Reinhardt). 

I have not seen this species. The second cohimellar tooth 
described is what would now be considered a basal fold, sub- 
columellar in position. 

36. VERTIGO HACHIJOENSIS n. sp. PL 14, figs. 6, 7. 

The shell is rimate, rather shortly oval, sayal brown, 
opaque. The surface is very finely, sharply striate on the 
penult and next earlier whorls, more coarsely and less sharply 
so on the last whorl. The whorls are moderately convex, the 
last not compressed below or impressed over the palatal region ; 
it expands into a rounded crest close to the lip. The aperture 
is ovate, with four well developed teeth: parietal lamella 
rather long ; columellar lamella somewhat smaller ; two rather 
small palatal folds, the lower one larger, as usual. In old 
specimens, such as the figured type, there is also a very low, 
almost inmersed basal fold (but in others this is wanting). 
There is a rather thick palatal callus. The outer lip is 
straightened and very slightly incurved in the middle, its edge 
a trifle expanded. Length 1.8, diam. 1.15 mm.; 4% whorls. 

Japan: Hachijojima, Izu, Y. Hirase. Type and para- 
types no. 83394 A. N. S. P. 

It is distinguished from other Japanese species having a 
crest and a palatal callus by the distinctly striate shell. It 
was mentioned without description in Proc. A. N. S. Phila., 
1905, p. 718. 

37. VERTIGO KUSHIROENSIS Pils. & Hir. PI. 14, figs. 8, 9, 10. 
The shell is rimate, ovate, auburn. Surface glossy, nearly 

smooth, with only weak traces of striation. The whorls are 
convex, the last not impressed over the palatal region, having 
a strong, opaque buff crest behind the lip. The aperture is 



154 VERTIGO OF JAPAN AND EASTERN ASIA. 

ovate, with 4 or 5 white teeth: parietal lamella moderately 
developed; columellar lamella directed downward, ascending 
a little within; two well developed palatal folds, the lower 
longer; a small basal fold (but wanting in some examples). 
There is a distinct whitish palatal callus. The outer lip 
is straightened in the middle but not inflexed; lower part 
of the peristome is expanded. 

Length 1.8, diam. 1.1 mm. ; barely 5 whorls (type, figs. 8, 10) . 

Length 1.75, diam. 1 mm. (fig. 9). 

Japan : Akkeshi, Kushiro, in northeastern Yesso, Y. Hirase. 
Type and paratype no. 90223 A. N. S. P. 

Vertigo hirasei kushiroensis PILSBRY & HIRASE, Proc. A. 
N. S. Phila., 1905, p. 718. 

This species differs from V. hydrophila by having no angu- 
lar lamella; from V. hirasei by the development of a strong 
crest, as well as by the stronger, darker shell. 

38. VERTIGO HIRASEI Pils. PI. 14, figs. 13, 14. 

The shell is rimate, cylindric-ovate, ochraceous-buff, im- 
perfectly transparent, thin. Surface glossy, with faint traces 
of striae, becoming finely more distinctly striate a short dis- 
tance behind the lip. The summit is very obtuse. Whorls 
are rather weakly convex, the last not compressed below, with- 
out external impressions or crest. The aperture is ovate, 
having 4 small teeth: the parietal lamella is oblique, about 
0.2 mm. long; columellar lamella short, ascending a little in- 
wardly. Lower palatal fold short, but longer and more im- 
mersed than the very small upper palatal. There is no trace 
of a palatal callus. The peristome is thin, strongly arched 
above, straightened in the middle; it is very slightly ex- 
panded in the lower part. In profile view (fig. 14) it is 
seen to bend well forward in a rounded lobe above the middle 
of the outer margin. Length 1.65, diam. 1 mm. ; fully 4% 
whorls. 

Japan: Yanagawa, Chikugo, in Kiushiu, Y. Hirase. Type 
and paratypes no. 79738 A. N. S. P. 

Vertigo hirasei PILSBRY, Nautilus xiv, March, 1901, p. 128 ; 
Proc. A. N. S. Phila., 1901, p. 484, pi. 28, f. 53. Vertigo 



VERTIGO OF JAPAN AND EASTERN ASIA. 155 

hirasei glans PILSBRY & HIRASE, Proc. A. N. S. Phila., 1904, p. 
631 (Okinoerabushima). 

The absence of angular and basal teeth and small size of 
the others, the thin lip, without palatal callus and not in- 
flexed, and the absence of a crest, are characteristic of this 
species. 

V. h. glans. The characters I relied upon in character- 
izing V. h. glans have little value, and I doubt whether it is 
really separable. The shell is more egg-shaped and a trifle 
smaller than the type specimen of V. hirasei, and there is no 
upper palatal fold. 

Length 1.5, diam. 0.85 mm. (pi. 14, fig. 15, type specimen). 

Specimens of a race of V. hirasei were sent by Mr. Hirase 
from Miyakejima, an island of Izu. The teeth are larger than 
in the Kyushu form. It probably is another subspecies, but 
the specimens are not in sufficiently good condition for 
decision (pi. 14, figs. 11, 12). 

28a. Vertigo hirasei okinoerabuensis Pils. & Hir. PL 14, fig. 16. 

The shell is longer than V. hirasei, of paler tint, with the 
lower palatal fold longer ; upper palatal present. Length 1.8, 
diam. 0.95 mm. ; nearly 5 whorls. 

Okinoerabushima, an island of Osumi, Hirase. Type and 
paratypes no. 87690 A. N. S. P. 

Vertigo hirasei okinoerabuensis PILS. & HIR., Proc. A. N. S. 
Phila., 1904, p. 631. 

The type is figured. A rather weakly differentiated form. 

39. VERTIGO JAPONICA Pils. & Hir. PL 15, figs. 1, 2. 

The shell is perforate and rimate, oblong, the summit ob- 
tuse, auburn, thin; surface smoothish, showing some weak 
striae under the microscope, the last whorl distinctly striate 
behind the lip. The whorls are convex, parted by a well 
impressed suture, the last half of the last whorl tapering 
towards the base, having a rather deep furrow running to 
the auricle, but no crest. Aperture ovate, with four well- 
developed teeth: parietal lamella large and rather long; 
columellar lamella smaller, ascending a little inwardly; two 



156 VERTIGO OP JAPAN AND EASTERN ASIA. 

palatal plicae well developed, the lower one somewhat more 
immersed. No palatal callus. The outer lip is biarcuate, 
a conspicuous entering angle at the junction of the arcs ; edge 
blunt and expanded. Length 1.65 diam. 0.95 mm. ; 5 whorls. 

Japan : Ikusagawa, jima, Y. Hirase. Type and paratypes 
no. 85746 A. N. S. P. 

Vertigo japonica PILSBRY & HIRASE, Nautilus xvii, Feb., 
1904, p. 118. 

It differs from V. hirasei by the entering point of the outer 
lip ; from V. hydrophila by the same character and by having 
fewer teeth. 

39&. Vertigo japonica coreana n. subsp. PL 15, fig. 4. 

Differs from V. japonica by the decidedly longer lower 
palatal fold. Length 1.53, diam. 0.9 mm. 

Korea: Island of Ko-je, Y. Hirase. Type and paratypes 
no. 95772 A. N. S. P. 

Some "dead" specimens from Fusan, Korea, partly young 
and all more or less broken, look like V. japonica, but their 
identity is not certain. 

39&. Vertigo japonica tosana n. subsp. PI. 15, fig. 3. 

The shell is distinctly striate ; there is a low angular lamella ; 
other teeth as in V. japonica. Length 1.65, diam. 0.95 mm. 

Japan: Irazuyama, Tosa, Y. Hirase. Type no. 86486 A. 
N. S. P. 

40. VERTIGO DENUDATA (Mousson). PI. 15, fig. 8. 

Shell very minute, perforate, long-ovate, denuded of epi- 
dermis, smoothish, slightly striatulate, subopaque, whitish. 
Spire convex, rather obtuse at summit; suture impressed. 
Whorl 5~y 2 to 6, convex, the penult a little larger, the last 
whorl somewhat tapering, not compressed below. Aperture 
nearly vertical, semicircular; peristome acute, a little re- 
flected, bordered by a wrinkle outside; margins connected; 
outer margin curved at the insertion, almost forming a right 
angle. Within there are two distant parietal teeth, one 
columellar and one palatal, sometimes a second small one being 



VERTIGO OF JAPAN AND EASTERN ASIA. 157 

added. Length 1.2, diam. 1 mm. Ratio of aperture [to spire] 
1 :1 (Mouss.). 

Siberia : Vladivostok, Louis Graeser. 

Pupa (Vertigo) denudata MOUSSON, Journ. de Conch, xxxv, 
1887, p. 19, pi. 1, f. 6. 

' ' Taken at the same point as P. alpestris, but it is not to be 
confused with that. It is always denuded, with little trans- 
lucence, whitish, while the other species is corneous and dis- 
tinctly translucent. P. denudata has % to 1 whorl more than 
alpestris, its whorls are less close, giving it a longer, slimmer 
shape; the aperture is more regularly rounded, the outer 
margin not concave" (Mousson). 

The presence of two teeth on the parietal wall, and of a 
crest ("perist. . . . extus ruga circumdatum") seem to show 
affinity with the V. hydrophUa group, but the figure is not 
sufficiently exact for a critical comparison with the Japanese 
species. Moreover, the dimensions given are clearly erroneous 

Subgenus PTYCHALAEA Boettger. 

Ptychalaea BOETTGER, Jahrb. Nassau. Ver. Naturkunde, 
Jahrg. 42, 1889, p. 293. Type by orig. des. V. flexidens 
(Bas.). 

Similar to Vertigo except that the angular lamella connects 
by a curved callous ridge with the termination of the outer 
lip; there is a rounded crest behind the peristome. Type 
V. flexidens Eeuss. 

Distribution, Lower Miocene to Pliocene of Europe ; recent 
in the Bonin Islands. 

The type of this subgenus, V. flexidem (pi. 15, fig. 15), has 
an infraparietal lamella and a suprapalatal denticle. The 
angular lamella and the rest of the structure agree exactly 
with the recent species, and I believe that there can be no 
doubt that the latter belong to Ptychalaea. The Bonin 
Islands have a highly peculiar snail fauna, doubtless evolved 
on a far larger land-mass which included all of the present 
islands. This fauna appears to have been derived from the 
continent not later than the middle Tertiary, and probably 
earlier. In Europe the species of Ptychalaea had about the 



158 VERTIGO OP JAPAN AND EASTERN ASIA. 

distribution and nearly the duration of the Gastrocopts, which 
are also related to those of the Bonin Islands. 
The Tertiary species are enumerated on page 220. 

41. VERTIGO DEDECORA (Pilsbry). PI. 15, figs. 5, 6, 7, 9. 

The shell is minute, with a distinct circular perforation, and 
a rather long umbilical chink, obesely oval, tawny, the surface 
having very weak but somewhat coarse striae and very little 
gloss. The whorls are moderately convex, the last one flat- 
tened over the palatal folds, rising in a rounded, lighter- 
colored crest behind the peristome, this crest being very promi- 
nent at the base. The parietal lamella is very long, entering 
deeply, not emerging to the edge of parietal callus. Angu- 
lar lamella is much shorter, connected by a callus with the 
termination of the outer lip. The columellar lamella is hori- 
zontal and deeply immersed. Upper palatal fold is larger and 
longer than the lower, and it emerges nearly to the peristome. 
Basal fold is short and somewhat immersed. There is a dis- 
tinct palatal callus. The peristome expands and is of a 
tawny color, the margins united by a distinct parietal callus. 
The outer margin is slightly impressed in the middle, a 
slight groove preceding the impression. 

Length 1.5, diam. 1 mm. ; 5 whorls. 

Length 1.6, diam. 1.05 mm. ; 5 whorls. 

Bonin Islands (Ogasawara-jima) : Hahajima, Y. Hirase. 
Type no. 82583 A. N. S. P. 

Nesopupa dedecora PILSBRY, Proc. A. N. S. Phila., Apr. 23, 
1902, p. 31. 

The compact, broadly oval figure, strong crest, long parietal 
lamella and the union of angular lamella with the peristome 
are the more prominent features of this species. 

410. V. d. tamagonari Pils. & Hir. PI. 15, figs. 10, 11. The 
shell is somewhat more globose and often smaller than typical 
dedecora :; the crest is not quite so prominent. Chichijima, 
Ogasawara. 

Length 1.4, diam. 0.95 mm. (type). 

Length 1.5, diam. 1 mm. 

Nesopupa tamagonari PILSBRY & HIRASE, Nautilus xvii, Feb., 
1904, p. 118. 



EUROPEAN SPECIES OF VERTIGO. 159 

Comparison of the whole series now at hand shows that this 
is merely a slight variant from another island, doubtfully 
separable from the original dedecora. 

III. PAUEARCTIC SPECIES, EUROPE, ASIA AND NORTHERN 

AFRICA. 

The European Vertigos have never been monographically 
studied. Westerlund's several works are synoptical compila- 
tions not really aiming to be critical monographs, I presume ; 
and the extensive and valuable papers of Gredler and 
Boettger, as well as the several faunal manuals, such as those 
of Clessin and others, cover only part of the ground. Wester- 
lund has carefully described many new forms from Sweden, 
where the genus appears to have an extraordinary develop- 
ment; but they have not been figured, and most of them are 
to be found in very few collections, or in but one, many 
being practically unknown outside of Sweden. 

Most of the species of Central Europe have been figured by 
Kobelt, Geyer and others, in lithographs, and by Steenberg 
in admirable pen drawings, among the best figures of Vertigo 
ever published. Having no special knowledge of European 
species, and possessing few of them, the author has compiled 
most of the following account from the original publications. 

NORTH AFRICAN SPECIES are all supposed to be distinct from 
those of Europe. They are still known only by Bourguignat 's 
work of over 50 years ago. 

V. discheilia Bgt, no. 49. V. briobia Bgt,, no. 59. 

V. maresi Bgt., no. 50. V. microlena Bgt., no. 60. 

V. aprica Bgt., no. 57. V. latasteana Bgt. & Let., no. 45. 
V. codia Bgt., no. 58. 

Westerlund has constructed a partial key to Palaearctic 
species, which is recast below with some small alterations. 

1. Shell dextral. 2. 
Shell sinistral. 9. 

2. Aperture toothless ; neck scarcely impressed in front, 
without an encircling crest; no internal palatal callus: 



160 EUROPEAN SPECIES OF VERTIGO. 

V. genesii, 78; dic<za, 63; celata, 64; inermis, 62; eggeri, 

66 ; regularis, 65. 

Aperture toothed. 3. 

3. No crest behind the peristome. 4. 
Having a crest. 6. 

4. No palatal callus, the palatal folds standing free. 5. 
Palatal folds united in front by a palatal callus. 

V. eremia, 61. 

5. One parietal lamella or none: V. daliaca, 69; pineticola, 
70 ; krauseana, 26 ; hebraica ; gemma, 71 ; ovoidea, 72 ; 
arctica, 67; tatrica, 67&; alpestris, 74; achetta, 76; Tiek^, 
74a ; haeusleri, 52a ; schultzii, 75 ; helvetica, 74d ; leontina, 
74b ; buttneri, 77. 

Having two lamellae on the parietal wall: thibetica, 
vol. xxiv, p. 138. 

6. No palatal callus between the palatal folds. 7. 
Palatal folds united by a palatal callus. 8. 

7. No impression behind the outer lip, crest weak : V. decora, 
25 ; rwmebynesis, 68 ; graellsiaiia,, 56. 

An impression between the strong crest and the edge of 
outer lip: V. antivertigo, 45; lilljeborgi, 46; kusteriana, 
536 ; latasteana, 45. 

8. One parietal lamella : V. moulinsiana, 53 ; pachygaster, 
54; pygmcea, 52; loroisiana, 55; aprica, 57; codia, 58; 
briobia, 59 ; microlena, 60. 

Two lamellae on parietal wall : F. sieversi, 73 ; substriata, 
51 ; reneana, 45c ; bcmdoni, 47 ; discheilia, 49 ; maresi, 50. 

9. Upper palatal fold not longer than the lower when 
present. 10. 
Upper palatal fold very long, entering to the dorsal side ; 
columellar lamella spirally ascending, blunt : V. angustior, 
81 ; callista, 82. 

10. 5 to 10 teeth: V. pusilla, 42; erlandi, 44; collina, 43. 
2 or 3 teeth : T. otostoma, 79 ; clevei, 80. 

Group of V. pusilla. 

These are merely sinistral species, otherwise like the 
vertigo group. 



EUROPEAN SPECIES OF VERTIGO. 161 

42. VERTIGO PUSILLA Miiller. PL 16, figs. 1, 2, 3. 

Shell subfusiform, with, somewhat of a quadrangular out- 
line, thin and semitransparent, very glossy, horn-color, with 
a faint tinge of yellow, very slightly and remotely striate in 
the line of growth; periphery rounded, with a tendency to 
angularity ; epidermis thin ; whorls 4% or 5, very convex and 
cylindrical, gradually increasing in size, the penultimate whorl 
as broad as the last, which occupies about two-fifths of the 
shell; spire shortish, but rather tapering, and blunt at the 
point ; suture very deep ; mouth semioval, contracted or sinu- 
ous in the middle of the outer edge; teeth six or seven, viz. 
two on the pillar [parietal wall], two on the pillar lip (the 
inner one of which is always larger, and the outside one 
tubercular and placed in the angle where the outer lip 
joins), and two or three within the outer lip (the third, when 
it is present, placed near the pillar lip and being a mere 
tubercle:) outer lip rather thick and slightly reflected, 
strengthened by a strong rib both outside and inside, which 
is situate near the opening of the mouth and is yellowish- 
white ; outer edge rather abruptly inflected ; inner lip slightly 
thickened in full grown specimens; umbilicus small and 
narrow, contracted by a rather sharp and gibbous crest or 
ridge at the base of the shell. L. 0.07. B. 0.045 inch 
(Jeffreys). 

Europe; Caucasus; Asia Minor. 

Vertigo pusilla MULLER, Vermium terr. et fluv. Hist., ii, 
1774, p. 124. A. MULLER, Wiegm. Archiv f. Naturg. 1838, i, 
p. 211, pi. 4, f. 5. ROSSMAESSLER, Iconogr. ii, pt. 10, p. 29, 
pi. 49, f. 649.--JEFFREYS, Brit. Conch, i, p. 263. MOQUIN- 
TANDON, Moll. France ii, p. 409, pi. 29, f . 12-14. Pupa pusilla 
(Mull.) PFEIFFER, Monographia ii, p. 364; iii, 560; iv, 686; vi, 
336 ; viii, 404. KUSTER, Conchyl. Cab. p. 129, pi. 16, f . 38-40. 
WESTERLUND, Fauna iii, 1887, p. 141 ; Acta Soc. pro fauna 
et flora Fennica xiii, no. 7, p. 70 (dist. in Norway, Sweden, 
Finland). Helix vertigo GMELIN, Syst. Nat. (13), p. 3664. 
Pupa vertigo DRAP., Hist. Moll. France, p. 61, pi. 3, f . 34, 35. 
Vertigo Keterostropha Leach, TURTON, Manual, 1831, p. 105. 
LEACH, Synopsis Moll. G. B. 1852, p. 94. 



162 EUROPEAN SPECIES OP VERTIGO. 

With teeth somewhat as in V. antivertigo, this species differs 
by the more oblong shape and sinistral coil. Westerlund 
gives the length as 1% to 2y 2 mm. ; those I have measured are 
mainly between 1.9 and 2.1 mm. long; the shortest adult 
1.6 x 1 mm. 

It is locally distributed in England and Ireland but not 
in Scotland, according to Jeffreys. 

42a. Vertigo pusilla tumida Westerlund. Shell ventri- 
cose-ovate, obtuse, the base swollen on the right; brown, stri- 
atulate ; whorls 4% to 5, convex, the last pale anteriorly, with 
a very thin callous, the base a little compressed. Aperture 
broadly, unequally cordate, 4 to 6 toothed: parietal 2 or 1, 
columellar 2 or 1, palatal 3. Length 1%, diam. 1% mm. 
Sweden, at Pehrsborg near Ronneby among rotten, damp 
leaves in a beech wood (type loc.) and Borgholm; Norway, 
Malmoen ; Denmark, Linaa Vesterskov. ( Westerlund) . 

Vertigo tumida WESTERL., Malak. Blatter xiv, 1867, p. 203.. 
Pupa tumida WESTERL., Fauna iii, 1887, p. 141 ; Acta Soc.. 
fauna et flora Fennica xiii, no. 7, p. 71. 

Said to differ from V. pusilla by the smaller, blunt and ven- 
tricose shell, with different shaped aperture, darker color and 
smaller umbilical crevice. 

"I am not sure that it is more than a dwarf variety or 
form of V. pusilla. The two specimens sent by Dr. Wester- 
lund differ from each other in the number of teeth, one speci- 
men having 5 and the other 7 teeth. He describes tumida 
as 6- dentate, and V. pusilla as 6-8 dentate." (Jeffreys, Ann. 
Mag. N. H. 5th ser., ii, 1878, p. 381). Mr. J. W. Taylor 
states that Westerlund sent him a specimen labelled Vertigo 
pusilla Mull. var. tumida West. (Journ. of Conch, v, 357). 

43. VERTIGO COLLINA Westerlund. 

Shell [sinistral] rimate, ovate, striatulate, corneous. Spire 
subconic, short, rather obtuse. Whorls 5%, somewhat flat, 
united by a lightly impressed suture, not descending at the 
aperture, the last whorl a little impressed in the middle in 
front, strongly sloping below, as viewed from the side. Aper- 
ture oblique, subcordate, with 2 lamelliform, parallel palatal 



EUROPEAN SPECIES OP VERTIGO. 163 

teeth, not reaching the margin ; peristome simple, unexpanded, 
the outer margin forming two well curved circles; connected 
with the columellar margin by a rather strong callus. 
Length 2, diam. 1% mm. (WesterL). 

Sweden: Pehrsborg near Ronneby, among rotten leaves in 
a beech wood. 

Pupa collina WESTERLUND, Nova Acta R. Soc. Sci. Upsal. 
(3), viii, 1871, p. 100; Acta Soc. pro fauna et flora Fennica 
xiii, no. 7, p. 71. 

44. VERTIGO ERLANDI (Westerlund). 

Shell [sinistral], circularly perforate, oblong-conoid, corn- 
eous-brown, delicately striatulate. Whorls 6, regularly in- 
creasing, convex, the suture oblique as far as to the aperture, 
the last whorl with no transverse callus, smooth. Aperture 
sinuate-oval, the parietal margin very oblique, with 1 enter- 
ing parietal lamella ; 2 conic columellar teeth ; 3 rather thick, 
long, palatal folds, impressed in the middle, or abrupt, the 
1st emerging, corresponding to an external furrow, the 3d 
basal. Outer margin much longer, formed of two arcs, the 
upper one stronger, produced. Length 2, diam. 1 mm. 
(Westerl.). 

Sweden: Dalbyo, prov. Sodermanland. 

Pupa (Vertilla) erlandi WESTERL., Acta Soc. pro fauna et 
flora Fennica, xiii, no. 7, 1897, p. 194. 

Group of V. antivertigo. 

Ovate, glossy, usually rather dark colored species with 
6 to 10 teeth, 2 or 3 on the parietal wall (or in V. moulwisia/ria 
4-6 teeth, 1 or 2 on parietal wall). 

This group of fully toothed forms of Europe and north 
Africa is equivalent to what is called the group of V. ovata 
in America. V. callosa of the central European Oligocene be- 
longs to the same group, and is probably ancestral to the 
recent European species. 

45. VERTIGO ANTIVERTIGO (Drap.). PL 16, figs. 4, 5, 6. 

The shell is rimate, oval, ventricose, nearly smooth, glossy, 



164 EUROPEAN SPECIES OF VERTIGO. 

amber-brown or nearly chestnut, slightly transparent, out- 
lines very convex. Whorls rather convex, the last somewhat 
compressed below, with an impression over the lower palatal 
fold, a moderately developed, opaque crest behind the peris- 
tome; and a very deep impression between the crest and the 
point of the outer lip. Aperture having 6 principal and 
usually several smaller teeth: parietal lamella rather long; 
angular and infraparietal short and smaller. Columellar 
lamella large, ascending inwardly. Upper and lower palatal 
folds strong, the lower longer. Basal fold stout, in a sub- 
columellar position. Usually there are small suprapalatal and 
infrapalatal denticles. Peristome thin, a little expanded, the 
outer margin biarcuate, with a median entering angle. Palatal 
callus well developed. 

Length 1.95, diam. 1.2 mm.; 5 whorls (Lyons). 

Length 2.25, diam. 1.4 mm. (Lausanne). 

Nearly all Europe; Transcaucasus ; Armenia, Talysch 
Turkestan (Westerlund). 

Pupa antivertigo DRAPARNAUD, Tabl. Syst. Moll. France, 
1801, p. 57; Hist. Nat. Moll. Fr., p. 60, pi. 3, f. 32, 33 (dans 
la Bresse; en Languedoc). PFR., Monogr. Hel. Viv. ii, 361; 

11, 558 ; iv, 685 ; vi, 332 ; viii, 404. KUSTER, Syst. Conchyl. Cab. 
p. 125, pi. 16, f. 27-30. WESTERLUND, Fauna iii, 1887, p. 140; 
Nova Acta R. Soc. Sci. Upsal. (3), viii, 1871, p. 89, with varr. 
ferox, seminulum. Vertigo antivertigo Drap., NOBRE, Moll. 
Portugal, p. 241. MOQUIN-TANDON, Moll. France, 1855, p. 
407, with var. octodentata Stud. BOETTGER, Jahrb. Nassau. 
Ver. Nat. 42, 1889, p. 299 (recent and Pleistocene distribu- 
tion). WEISS, Nachrbl, D. M. Ges. 1894, p. 156, pleistocene 
of Taubach, with ver. ferox. 

Alaea palustris Leach MS., JEFFREYS, Trans. Linn. Soc. 
Lond., xvi, pt. 2, 1830, p. 360. 

Turbo sexdentatus MONTAGU, Test. Brit. 1803, p. 337, pi. 

12, f. 8. Vertigo sexdentata WAGNER in Chemnitz, Conchyl. 
Cab. xii, p. 175, pi. 235, f . 4124. 

Vertigo 8 dentata STUDER, Kurzes Verzeich., in Naturwiss. 
Anzeiger allg. Schweiz. Ges. Naturwiss. iii, May 1, 1820, p. 89, 
based upon Drap. pi. 3, f. 32, 33 (near Bern and Ringgen- 



EUROPEAN SPECIES OF VERTIGO. 165 

berg). Pupa vertigo var. P. 7 dentata HARTMANN, Neue 
Alpina i, 1821, p. 219, based upon Drap. pi. 3, f. 32, 33, Pupa 
antivergio [sic] . Vertigo septemdenta FERUSSAC, Tabl. Syst. 
p. 64, based upon P. antivertigo Drap., V. 8 dentata Studer, 
T. sex dentatus Mont. ROSSMAESSLER, Iconogr. ii, pt. 10, p. 
28, pi. 49, f. 647. 

Distinct from other species of its region by the three teeth 
on the parietal wall, the obesely oval shape and the deep angle 
between the two arcs of the outer lip. It is very closely re- 
lated to the American V. ovata, but is smaller with less im- 
pressed suture. Boettger, however, considered ovata a variety 
of antivertigo (Jahrb. Nassau. Ver. Nat. 1889, p. 300) ; and 
probably, if the areas of the two were not so widely separated, 
this view might be generally entertained. 

Westerlund gives the size as length 2-2%, diam. 1%-1%. 
This indicates a somewhat larger size than any I have seen. 

According to Jeffreys it is found "under stones and logs 
of wood as well as at the roots of grass, and on moss, flags 
and water plants, in marshy places and at the sides of streams 
and canals." 

Many of the minor modifications have been named, but 
probably few if any of them have racial value, so far as one 
may judge by the literature. It has not been stated that any 
occur in pure colonies. Moreover, some forms appear to have 
been named more than once. 

Westerlund recognizes the following forms : sexdentata Mont., 
2 columellar teeth, the upper wanting; octodentata Hartm., 
with 3 palatal teeth; ferox West., aperture 9 to 10 toothed, 
2 or 3 parietal, 3 columellar, 4 palatals, two lamelliform, two 
minute; or with 3-2-4 or 3-2-5 teeth (= var. novemdentata 
Sandberger, Vorwelt, 1875, p. 795). Sterki noted antivertigo 
mit 10 Zahnen, Schleitheim, Switzerland, Nachrbl. 1881, p. 
37. This form occurs also in Germany. 

Var. seminulum West. Shell small, oval; whorls 4; aper- 
ture 5 to 6 toothed : 2-2-2. Length 1%, diam. 1 mm. Oester- 
gotland, Sweden. 

Pollonera defines the following forms: typica, teeth 2-3-2; 
irregularis, 2-2-3 ; aequidentata, 2-2-2 ; padana, 2-1-3 ; cisalpina, 



166 EUROPEAN SPECIES OF VERTIGO. 

3-2-2. All from Piedmont (Atti R. Accad. Sci. Torino, xx, 
1885, p. 686). 

Var. major Locard. Alt. 2-21/4, diam. 114 mm.; 5 folds. 
Uriage (Isere). (Locard, Ann. Soc. Agricult. Lyon (5), ii, 

1880, p. 855). 

Var. novemplicata Loc. Shell conforming to the type but 
with a third fold on the penult whorl; this fold is very low 
and situated near the eolumellar margin, ordinarily quite im- 
mersed. Alluvium of the Rhone at Lyons (Locard, t. c., p. 
856). 

Var. cornea Loc. Pale corneous, subtransparent. Environs 
of Lyons (Locard, 1. c.). 

Pfeiffer (Monographia ii, 361) mentioned, as a doubtful 
small variety of V. antivertigo, a Pupa pusilla Bivona fils, 
Monogr. p. 14; Arad. et Magg., Catal., p. 131. "T. valde 
parva, ovato-conica, apice obtuse, corneo-ruf escente ; anfr. 4 
5 convexis, suturis impressis; aperture 6- vel 7 dentata, 
latere dextro sinuosa ; margine reflexo. Long. y 2 lin." (Ar. et 
Magg.). It is from the river Oreto, Sicily. The name is 
preoccupied. Benoit (Nuovo Catal. Conch, terr. e fluv. Sicilia 

1881, p. 99) remarks that two varieties of Pupa antivertigo 
have been observed in Sicily, the larger one living at Madonie, 
the smaller on the banks of the Oreto river. If really as small 
as stated, this is evidently a distinct species. 

Vertigo sinuata Mousson (pi. 16, fig. 7). Shell small, dex- 
tral, rimate, globose-ovate, slightly striatulate, pellucid, glossy, 
pale corneous. Spire convexly conic, the apex obtuse, suture 
somewhat impressed. Whorls 5, rapidly increasing, the penult 
largest, inflated, the last ascending a little, tapering below, 
more distinctly striate, in the middle of its alt. acutely im- 
pressed and having a transverse wrinkle. Aperture small, 
vertical, irregularly semioval, conspicuously sinuate above on 
the right, internally 7-toothed: 2 distinct and a third rudi- 
mentary parietals. 1 long in the post-sinual angle, 1 deep, 
elongate and another shorter in the lower margin, finally a 
seventh, immersed, on the columella. Peristome rather ob- 
tuse, a little reflected, labiate within; margins somewhat ap- 
proximate and converging, joined by a somewhat callous la- 
mella, the right bisinuate, the angle projecting strongly into 






EUROPEAN SPECIES OP VERTIGO. 167 

the aperture, a little produced forward. Length 0.9, diam. 0.6 
mm. (Mouss.). Banks of the Araxis river, Transcaucasus. 

Vertigo sinuata MOUSSON, Journ. de Conchyl. 1873, p. 213, 
pi. 8, f. 10. Pupa sinuata Mouss., J. de C. 1876, p. 40. PPR., 
Monogr. viii, p. 405. 

If the dimensions are correctly given by Mousson, this 
must be distinct from V. antivertigo, of which it has been 
considered a synonym. 



Vertigo reneana Servain. Shell rimate-perforate, 
ovate-globose, ventricose, corneous, glossy, nearly smooth. 
Spire convexly oblong, the apex large. Whorls 5, convex, 
slowly increasing, parted by a deep suture ; the last whorl 
minute, rounded, straight above, crested-tuberculose below, 
the protuberance joined to the gibbosity preceding the peris- 
tome; gibbous behind the margin, the swelling paler, strong, 
transverse, stronger below, and concave in front of the swell- 
ing below. Aperture ventrical, subtrigonal, narrowed and 
indistinctly angular below, externally sinuate, 7 plicate, as 
follows: 3 parietals, of which the median is stronger; 2 colu- 
mellars, the upper one thicker, and 2 nearly equal palatals, 
the upper one attaining the margin. Peristome lipped within, 
somewhat expanded and slightly reflected, the outer margin 
sinuated within, columellar margins obliquely straightened, 
margins remote. Length 2!/2, diam. 13/2, aperture % x y 2 
mm. (Servain). 

Spain: drift of the Ebre at Saragossa. Vertigo reneana 
SERVAIN, Etude Moll. Esp. Port. 1880, p. 126. 

Resembles V. maresi of Algeria in form and size, but it 
differs by the aperture having 7 teeth and by the characters 
of the last whorl (Servain). It has not been differentiated 
satisfactorily from antivertigo. 

46. VERTIGO LILLJEBORGI (Westerlund). 

Shell ventricose, ovate, strongly glossy, very finely striate, 
chestnut horn-color. Whorls 5, rather rapidly increasing, 
convex, the last but little higher than the penult, double as 
high as the next earlier whorl, a little ascending in front. 
Suture but slightly oblique. Aperture quite piriform, or 
obliquely cordate, with 1 parietal tooth (sometimes with an- 



168 EUROPEAN SPECIES OF VERTIGO. 

other punctiform one), 2 columellar teeth, the lower very 
small, often wanting; 2 short, high, equal, immersed teeth 
in the palate, bounded by a reddish brown streak in front. 
Peristome weak, expanded, the margins delicately united; 
outer margin not impressed, scarce}^ produced angularly for- 
ward. Length 2 to 21/4, diam. 1% to 1% mm. (WesterL). 

Sweden and Norway, type loc. on the southern shore of 
Tresjon lake, near Ronneby. Western Ireland: Ballyna- 
hinch, Co. Galway ; Connemara. 

Vertigo moidinsiana JEFFREYS, Brit. Conch, i, 1862, p. 
255 (not of Dupuy), with var. bidentata, p. 256. Vertigo 
modesta WESTERLUND, (Efvers af K. Vet. Akad. Forh, 1865, 
p. 556; Malak. Bl. xiii, 1866, p. 45 (not of Say, 1824). Pupa 
modesta A. West., PFR., Monogr. vi, 332. Vertigo lilljeborgi 
WESTERLUND, Coll. Typ. Moll. Suecia, 1868, No. 60 ; Synopsis, 
1897, p. 119, with var. merita, p. 119, and var. globula, p. 120. 
R. A. PHILLIPS, The Irish Naturalist, May, xvii, 1908, p. 89, 
pi. 3, f. 13, 14; p. 92, figs. G, H. (history of the species in 
Ireland). Pupa Ittljeborgi WESTERLUND, Expose crit., Nova 
Acta Eeg. Soc. Sci. Upsal. (3), viii, 1871, p. 90 (dist. in 
Sweden) ; Fauna iii, 1887, p. 136. 

V. lilljeborgi, compared with V. moulinsiana, "is much 
smaller, more glossy, its whorls are more tumid, and its thinner 
lip lacks the broad, almost colorless margin of the latter. 
The habits of the two animals also appear to be quite differ- 
ent, for, as has been shown, V. moulinsiana, although inhabit- 
ing marshes, avoids during both summer and winter anything 
in the nature of damp or decaying matter ; while the favorite, 
if not only, habitat, in this country at least, of V. lillje~borgi, 
is among the decaying roots and stems of aquatic plants cast 
up on lake shores' 7 (R. A. Phillips). 

Westerlund's localities in Sweden are also lake shores. 
Among other notes he states that the present species has so 
much resemblance to P. antivertigo Drap. in form, color, size, 
that he at first glance took it to be the young of this species, 
which also occurs on nearly all of our lake shores. It differs 
from antivertigo by the yellowish, not reddish brown color, 
the more convex whorls, brighter gloss, the shape of the neck 



EUROPEAN SPECIES OF VERTIGO. 169 

and the aperture. In antwertigo the neck has a swollen 
callus, cut above by a scratch-like impression, the furrow 
between the callus and the somewhat reflected peristome deep 
and narrow; it has also a differently shaped aperture and 
more numerous teeth, 6 to 10. 

Var. merita Westerlund. Callus of the last whorl extremely 
thin, rufous or nearly obsolete ; no groove behind the aperture ; 
teeth 1-1-3 (2), very small, nodiform. Sweden at Oestersund, 
Finland at Rissalanranta (Westerl.). 

Var. globula Westerlund. Shell subglobose, extremely ven- 
tricose, brownish-chestnut, striatulate; 4 very convex whorls, 
the penult largest, 3 times as large as the antepenult, larger 
and especially wider than the last; last whorl dilated anter- 
iorly, having a callus [crest] of the same color, and a small 
groove on the back. Aperture with 1-1-2 teeth, the parietal 
small, deeply placed, columellar basal, palatals punctiform, 
distant, submarginal. Length 2, diam. 2 mm. Sweden, in 
the province Sodermanland (Westerlund). 

Var. bidentata Jeffreys. Labial or palatal teeth wanting. 
If this named variety is really a form of lilljeborgi, it should 
be noted that the name is anterior. 

47. VERTIGO BAUDONI Massot. 

Ovoid-globose; 5 quite convex whorls, the suture much im- 
pressed ; aperture half-round, very obtuse at base ; 2 immersed 
superior folds of which that in the middle is largest ; 2 deeply 
placed columellars; 3 palatals, the upper one short, the two 
others reaching the peristome ; peristome continuous, ex- 
panded, rather thick, with an external crest ; shell yellowish 
rufous-brown, ornamented with spaced, regular epidermal 
expansions. Length 2, diam. 1 mm. (Locard). 

France: Tautavel (Pyrenees-Orien tales). 

Vertigo baudoni MASSOT, Enum. Moll. Pyr.-Or., 1872, p. 
72 (not seen). LOCARD, Ann. Soc. Agricult., Sci., Ind., de 
Lyon, (7),iii, 1896, p. 221. 

This little-known form should be recognized by the 7 teeth 
and the sculpture ; it may be related to V. substriata. 



170 EUROPEAN SPECIES OP VERTIGO. 

48. VERTIGO LATASTEANA Let. et Bgt. 

Shell very minute, slightly rimate, swollen ventricose, sub- 
ovate-spherical, fragile, diaphanous, corneous, smooth; spire 
short, very obtuse; whorls 5, convex (the embryonic small, 
median relatively ample and turgid, the last smallest, dimin- 
ishing and tapering) , rapidly increasing, separated by a rather 
deep suture; last whorl small, less wide than the penult, ex- 
ternally compressed, scrobiculate, elegantly marked with a 
lip [crest] behind the peristome, compressed at base and angu- 
lar around the umbilical crevice; above ascending to the in- 
sertion of the lip. Aperture suboblique, very small, con- 
tracted, subtriangular and 5-toothed as follows: 2 long and 
rather strong parietal teeth; 1 robust columellar; 2 lamelli- 
form palatals, reaching the outer margin, the upper one 
stronger and forming a deep external pit. Peristome paler, 
rather thick, spreading throughout; columella very short; 
outer margin sinuated in the position of the external pit, 
and produced forward in a sort of little beak; columellar 
margin dilated; the margins remote. Length 2, diam. 1 mm. 
(L.&B.}. 

Tunis : debris of the Oued Sidi-Aich. 

V. [ertigo] latasteana LETOURNEUX et BOURGUIGNAT, Prodr. 
Malac. Tunisie, 1887, pp. 109, 166. 

This very small species, remarkable for the contraction of 
its aperture and the spherical shape, cannot be assimilated 
with any of the living Vertigos. We see only V. milne- 
edwardsi of the hill of Sansan which resembles it not only 
in respect to the denticulation but in shape and contours 
(L.&B.). 

49. VERTIGO DISCHEILIA Bourguignat. PL 16, figs. 8, 9. 
Shell very minute, rimate, globose-oblong, glossy, subpel- 

lucid, corneous, obliquely substriatulate under the lens. Spire 
tapering, obtuse, the apex smooth, paler, obtuse. Whorls 5, 
convex, rather rapidly increasing, separated by an impressed 
suture; the last small, tapering, a little compressed basally, 
straightened in front. Aperture a little oblique, lunate, half- 
round, ringent, with many folds : 2 parietals, the lower deeper 



EUROPEAN SPECIES OF VERTIGO. 171 

within, lamellif orm ; 2 columellar, or often 3, the upper one 
stronger; 2 lamellif orm palatals, forming external furrows; 
finally, in adult shells, very often a dentiform fold at the 
insertion of the outer lip. Peristome a little expanded, acute, 
with a whitish thickening deep within, and encircled ex- 
ternally at the periphery with a thick and strong whitish lip 
[crest] . Outer margin sinuous above, arcuate anteriorly, the 
margins joined by a rather strong callus. Length 2%, diam. 
11/4 mm. (Bgt.). 

Algeria: debris of the Harrach at Algiers, of the Oued 
Tademit near Oran, and the Safsaf near Philippeville ; type 
loc., fossil in Pleistocene deposits of the Oued Tademit, 15 
leagues S.-W. of Djelfa. Tunis: debris of the Oued Sidi- 
Aich (Bgt). 

Vertigo discheilia BOURGUIGNAT, Paleont. Algerie, 1862, p. 
78, pi. 4, f. 3, 4; Malac. Alger. ii, 1864, p. 105, 313, pi. 6, 
f. 45-47. LETOURNEUX et BOURGUIGNAT, Prodr. Malac. 
Tunisie, 1887, p. 109. 

Var. unilabiata Bourguignat. Like the type but without 
external peristomial callus. Drift of the Harrach. 

50. VERTIGO MARESI Bourguignat. PL 16, figs. 10, 11. 

Shell pygmy, rimate, globuloid, glossy, pellucid, fulvous, 
smooth. Spire very obtuse, the apex very obtuse, paler. 
Whorls 5, convex, rapidly increasing, separated by a deep 
suture, the penult largest, inflated-globuloid ; the last small, 
contracted, tapering, a little compressed basally, straightened 
in front, often in some specimens a little swollen and angu- 
late-flattened at the suture. Aperture lunate, somewhat half- 
rotund, ringent, many-folded: 2 lamellif orm parietal folds, 
the inferior deeper and stronger ; 2 or often 3 columellars, the 
upper one stronger and the lower remote and very minute; 
2 palatals, lamelliform, forming furrows externally. Peris- 
tome simple, acute, slightly expanded, slightly thickened 
within, and externally encircled by a more or less strong, 
vinaceous-fulvous, very rarely paler lip [crest] ; outer margin 
sinuous above, slightly arching forward, the margins joined 
by a thin callus. Length 2, diam. 1% mm. (Bgt.). 



172 EUROPEAN SPECIES OF VERTIGO. 

Algeria : Oued Tademit, 15 leagues S.-W. of D jelf a, origin- 
ally found fossil, but later also living in the same valley, in 
humid places under stones (Mares). Alluvium of the Isser 
(Letourneux). 

Vertigo maresi BOURGUIGNAT, Paleontologie de 1'Algerie, 
1862, p. 79, pi. 4, f . 6-8 ; Malac. Alg., ii, 1864, p. 106, 313, pi. 
6, f. 48-50. HANOTEAU & LETOURNEUX, La Kabylie, i, 1872, 
p. 228. 

Group of V. substriata. 

Strongly striate or rib-striate, with teeth as in the anti- 
vertigo group, 1-1-2 to 3-1-2 or 3-1-3. According to Jeffreys 
the half grown shell has a spiral columellar lamella. 

51. VERTIGO SUBSTRIATA Jeffreys. PI. 17, fig. 10. 

" Shell oval or subfusiform, rather thin, and semitrans- 
parent, glossy, pale yellowish-horn-color, very strongly and 
obliquely striate and almost ribbed in the line of growth, but 
less so on the body whorl, which is faintly striate spirally, 
periphery rounded : epidermis rather thick : whorls 4%, very 
convex or cylindrical, and suddenly increasing in bulk, the 
penultimate whorl slightly exceeding in breadth the last, which 
occupies about one-half of the shell: spire short, very abrupt 
and bluntly pointed: suture remarkably deep: mouth semi- 
oval, contracted or sinuous in the middle of the outer edge; 
teeth from four to six, viz. from one to three (usually two) 
on the pillar [parietal wall] , one on the pillar lip, and two or 
three on the inside of the outer lip, the last springing from 
a white rib ; in half grown specimens the pillar lip has a spiral 
or longitudinal fold. Outer lip thin and slightly reflected, 
strengthened [externally] by a strong rib, which is placed 
very near the opening the mouth; outer edge abruptly in- 
flected, inner lip thickened in the adult ; umbilicus small and 
narrow, contracted by a keel or ridge at the base of the shell. 
L. 0.065. B. 0.04 inch." (Jeffreys). 

Great Britain, "from Skye to Devon as well as throughout 
Ireland" (Jeffreys). Scandinavia, Denmark and Germany, 
south to Switzerland and the Austrian Tyrol; Caucasus. 



EUROPEAN SPECIES OP VERTIGO. 173 

Type loc., Barnstaple, Devonshire. " Under stones, among 
dead and decaying leaves and at the roots of grass in woods 
and moist places." 

Al<za substriata JEFFREYS, Trans. Linnean Soc. London, xvi, 
1830, p. 515 ; Vertigo substrwta JEFFREYS, British Conchology 
i, p. 261. KOBELT, Iconogr. (2), viii, p. 89, f. 1524, 1525. 
STANDEN, Journ. of Conch, xi, 1905, p. 200, monstr. sinis- 
trorsum. Vertigo (Alaa) substriata Jeffr., BOETTGER, Jahrb. 
Nassau. Ver. Nat. 1889, p. 303 (Pleistocene and recent dis- 
tribution). STEENBERG, Danmarks Fauna, Landsnegle, 1911, 
p. 162, f. 133. 

Pupa substriata Jeffr., KUSTER, Syst. Conch. Cab. p. 
180, pi. 21, f . 22, 23. PFR., Monogr. ii, 363 ; iii, 559 ; iv, 685 ; 
vi, 334; viii, 407. GREDLER, Nachrbl. D. M. Ges. 1872, p. 70, 
with var. sextana. WESTERLUND, Expose crit., in Nova Acta. 
Reg. Soc. Sci. Upsal. (3), viii, 1871, p. 92 (many localities in 
Sweden and Norway), with var. monas. Pupa (Vertigo) sub- 
striata Jeffr., GEYER, Jabresh. Ver. vaterl. Naturk. Wurttem- 
berg, 36, 1907, p. 424; occurrence in Germany. 

Pupa curta HELD, Isis, 1837, p. 304 (Bavaria). 

Vertigo 5-dentata and V. 6-dentata STUDER, in Coxe, 
Travels in Switzerland, 1789, iii, p. 388, names only; = sub- 
striata according to Jeffreys, Linn. Trans, xvi, 516. 

The short, keg-like shape, the 6 or 7 teeth and especially 
the very strong striation, amply distinguish this species. 
Length 1.75, diam. 1.1 mm., or slightly smaller. Steenberg, 
who has published the best figure of the species, gives the 
length of Danish examples as 1.5 to 1.8 mm. 

Var. mitis Boettger. PI. 17, fig. 4. Form, sculpture and 
position of teeth as in substriata, V. sieversi being less strongly 
sculptured, more glossy and turrited. There is a similar weak 
transverse keel preceding the aperture as in typical specimens 
from Yorkshire. The outer parietal tooth is always very 
small, a feature observed here and there also in specimens 
from Dalarne, Sweden, and in other localities of the species. 
The Caucasian form is indistinguishable from specimens of 
the last-named locality. Alt. 1%, diam. 1 mm. Caucasia: 



174 EUROPEAN SPECIES OF VERTIGO. 

Abastuman and Kasbek. Pupa (Vertigo) substriata var. 
mitis BTTG., Jahrb. d. m. Ges. vii, 1880, p. 140, pi. 4, f. 7. 

Var. monas Westerlund. Very small, oval, very finely 
striate, dark colored, whorls 4%, cervical callus yellowish- 
red. Length !%-!%, diam. 1-1% mm. Sweden: Ronneby 
and Mt. Mosseberg in prov. Westergotland. Wurtemberg at 
Hohin Wittlingen (Westerlund). 

Var. sextana Gredler. The lower tooth on the columella 
is wanting, the palatal teeth weaker, the crest and projection 
around the umbilicus are scarcely indicated, the shell is 
thicker and more opaque, browner, somewhat larger, also the 
striation is less close (Gredler) . Tyrol : Sexten valley (Puster- 
thal), under stones. 

Form viridana Lindholm. Shell pale greenish, the neck 
encircled anteriorly with a whitish callus. Bobylsk, near 
Lachta, northwest of Petrograd, at the extreme eastern end 
of the gulf of Finland (Alaea substriata f. viridana Lindh., 
Nachrbl. d. m. Ges. vol. 42, 1910, p. 35). 

Group of V. pygmaea. 

Similar to the V. antivertigo group, but with fewer teeth, 
a single lamella on the parietal wall; the shell smoothish or 
weakly striate. Europe and northern Africa, North America. 

Only V. pygnuea and V. moulinsiana can be considered 
well known species, most of the others being known by the 
original descriptions only. 

52. VERTIGO PYGMAEA (Drap.). PI. 17, figs. 15, 16, 17. 

Shell extremely small, oval-cylindric and obtuse at the 
summit, of a more or less deep brown, smooth and little shin- 
ing. Spire of 5 progressively increasing whorls. Aperture 
scarcely longer than wide, and nearly round, furnished with 4 
teeth, of which the superior is acute, two deeply placed in- 
ferior, and finally one on the columellar margin. Often a 
fifth is found in the base of the aperture. The lateral margin 
is slightly angular in the middle. Peristome reflected be- 
low. Umbilical crevice quite pronounced. Lives under the 
hedges (Drap.). 






EUROPEAN SPECIES OF VERTIGO. 175 

Europe; Caucasus and Transcaucasia; Talysch region; 
Asia Minor. Ponto Delgado, S. Miguel, Azores, probably 
imported. 

Pupa pygmaea DRAPARNAUD, Tableau Moll. terr. et fluv. de 
la France, 1801, p. 57 ; Hist. Nat. Moll. terr. et fluv. France, 
p. 60, pi. 3, f. 30, 31. PPEIFFER, Monogr. Hel. Viv. ii, 362; 
iii, 559; iv, 685; vi, 334; viii, 405 (see for older references). 
KiisTER, Syst. Conch. Cab. p. 127, pi. 16, f. 31-34. GREDLER, 
Tyrol's Land- und Siisswasser-Conchylien, Verh. Zool.-bot. 
Vereins in Wien. vi, 1856, p. 126, with var. athesina and 
sarena. WESTERLUND, Fauna, 1887, p. 137. WOLLASTON, 
Testacea Atlantica, p. 47. Pupa pygmea and var. 5 dentata 
HARTMANN, Neue Alpina i, 1821, p. 219. 

Vertigo (Ala-ea) pygmaea (Drap.) BOETTGER,- Jahrb. Nassau. 
Ver. Nat., 1889, p. 305 (Pleistocene and recent distribution). 
ROSSMAESSLER, Iconogr., pt. x, 1839, p. 29, fig. 648. 
MOQUIN-TANDON, Moll. France, ii, p. 405, pi. 28, f. 37-42 with 
var. quadridentata. STEENBERG, Danmarks fauna, Land- 
snegle, 1911, p. 161, f. 132, with forms quadridentata and 
quinquedentata Studer, sexdentata and septemdentata, new 
forms. GEYER, Unsere Land- und Siisswasser-Mollusken, 
1909, p. 55, pi. 5, f. 26, 27; 6, f. 1. 

Vertigo similis FER., Tabl. Syst. p. 64. 

Vertigo 4 et 5 dentata STUDER, Kurzes Verzeichniss der bis 
jetzt in unserm Vaterlande entdeckten Conchylien, Natur- 
wissensch. Anzeiger der allg. Schweizerischen Gesell. f. d. 
gesammten Naturwiss., May 1, 1820, p. 89 (based upon Pupa 
pygmcea Drap., 3:30, 31). 

Helix Isthmia cylindrica GRAY, Lond. Medical Repository, 
xv, 1821, p. 239, based upon Drap. pi. 3, f. 30, 31. Vertigo 
vulgaris LEACH, in TURTON, Manual of land and fresh-water 
shells of the Brit. Is., 1831, p. 103 (as synonym of V. pygmaea). 

A fuller description and American references may be found 
on p. 96. I can see no difference between European and 
American specimens. It is perhaps the most generally dis- 
tributed Vertigo in Europe, from Portugal and the British 
Isles eastward. 

Form similis Ferussac (Tabl. Syst. p. 64) had no definition 



176 EUROPEAN SPECIES OP VERTIGO. 

further than "4 dentata" and the citation of "Vertigo 4 a 5 
dentata, Studer, Catal."; this last being repeated also under 
V. pygmaea. It was taken up by Fitzinger as "Vertigo 
pygmaea var. similis mihi" without definition (Beitrage zur 
Landeskunde Oesterreich 's, iii, 1833, p. 109). Ferussac's in- 
tention was to segregate the form lacking the basal fold, which 
was subsequently named quadridens West., as follows. 

Form quadridens Westerlund. Aperture with 4 teeth, lack- 
ing the lower columellar (Pupa pygmaa v. quadridens, West., 
Expose critique, Nova Acta Beg. Soc. Sci. Upsaliensis (3) viii, 
1871, p. 93). No special locality is given for this, which ap- 
pears to be not subspecifically separable from pygmaea. Many 
Scandinavian localities are mentioned for the latter. Var. 
quadridens has been reported from numerous places, as far 
south as the Balkans. 

Var. sexplicata Locard. Similar to the type but there are 
6 apertural folds ; the upper fold of the penult whorl is split, 
giving rise to a second obsolete superior fold. Debris of the 
Rhone north of Lyons. (V. p. var. sexplicata Loc., Fauna 
Mai. Bassin Rhone, in Ann. Soc. d'Agrieult. Lyon, (5), ii, 
1880, p. 853). 

Var. rubella Loc. shell similar to the type, with 5 folds, but 
of deep, reddish color. Environs of Grenoble (Locard, I. c.). 

Var. cornea Loc. shell similar to the type, with 5 folds, but 
of pale, subtransparent corneous color. Environs of Lyons 
and Grenoble (Locard, I. c.). This appears not to differ from 
pallida, which is earlier. 

Form pallida Jeffreys. Shell thinner and of a lighter color 
(V. pygmaea Jeffr., Brit. Conch, ii, 1862, p. 257, with var. 
pallida on p. 258). 

Form ausonia de Stefani. Upper palatal fold elongated. 
Italy. (Bull. Soc. Malac. Ital. ix, 1883, not seen. WesterL, 
Fauna iii, p. 137). 

Form minor Westerlund. 1% to 1% x 1 mm. Caucasus 
(Fauna iii, 1887, p. 137). 

Var. callicarens (Bttg. Hesse). A form without a trans- 
verse callus preceding the mouth. Near Greussen, Germany 



EUROPEAN SPECIES OF VERTIGO. 177 

(Hesse). Neighborhood of Piazza, in the Val Brembana, Italy 
(Andreae). 

Pupa pygmaea var. callicarens Bttg., HESSE, Nachrbl. d M. 
Ges. 1881, p. 7. P. p. var. callocarens ANDREAE, Nachrbl. 
1883, p. 135. 

Var. athesina Gredler. The impression behind the cervical 
callus [crest] inconspicuous; in the palate a small third 
tooth near the insertion of the outer lip, which is often more 
elevated than the lower one of the columella, and relative 
to size appearing to stand in an inverse proportion to the 
latter. I first found [this variety] in several examples at 
Bad Bergfall, near Clang in the Pusterthal, under stones. 
Further I have noticed it, more or less characteristically de- 
veloped, in the whole Etsch river region, especially at Klausen, 
Botzen, Salurna, Naturna, etc. [Tyrol] (Gredler). 

Var. sarena Gredler. Shell larger ; aperture 8-toothed : the 
lower tooth on the columella split into two smaller ones; in 
the palate 4 instead of 2 teeth, of which the 1st and 3d are 
noticeably larger, the 2d and 4th bluntly conic. [Tyrol] In 
the Sarnthal, in the village of the same name, under stones 
associated with var. athesina (Gredler). 

Dr. D. Geyer (Nachrichtsblatt d. deutschen Malak. Ges., 
44, 1912, p. 117-124), in an article on what might be called 
over-grown snails, discusses the individuals sometimes met 
with (a) having the last whorl abnormally inflated, (&) the 
shell relatively elongated by a whorl or two, with weakening 
of apertural structures, or (c) with these peculiarities com- 
bined ; noting various instances. Elongation of the shell is 
said to be commonest in V. pygmaea ; frequently a sixth whorl 
is added, the peristome being left sharp, without the char- 
acteristic armature. Others have the aperture finished, but 
without teeth or folds. He believes that Pupa haeusleri and 
Vertigo heldi are such forms of Vertigo pygmaea. In the 
river valleys of the south German Jura, where these rare 
snails appear to be restricted, they always occur associated 
in the river debris with V. pygmaea, never without it, though 
pygmaea occurs also without either haeusleri or heldi. 

Similar cases of gigantism are known in CoMicopa (Proc. 
Malac. Soc. Lond., xii, p. 312), Azeca and other genera. 



178 EUROPEAN SPECIES OF VERTIGO. 

The description of P. haeusleri follows. It is a little-known 
form, placed by Westerlund near alpestris and heldi. 

52a. Pupa haeusleri Sterki. PI. 17, fig. 9. Shell perforate, 
cylindric, rather obtuse at the apex; rufous-brown, the early 
whorls paler ; irregularly striatulate, glossy, pellucid. Whorls 
6, slowly increasing, convex, the suture rather deep, a little 
ascending before the aperture. Aperture nearly i/ 3 the length, 
% oval or subtriangular, three plicate: one conic tooth on 
the columella, two lamelliform in the palate. Peristome sim- 
ple, straight, acute, the palatal margin having no callus or 
impression either outside or within. Length 2.5, diam. 
scarcely 1.2 mm. (Sterki). 

Switzerland : drift debris of the lower Aar in the vicinity 
of Brugg, Dr. Hausler. 

Pupa haeusleri STERKI, Nachrbl. d. Malak. Ges. 1883, p. 72 y 

fig. 

At first glance, especially on account of the similar color, 
the specimens have a resemblance to P. strmta Gredler, not 
considering the strong fold on the parietal wall of that species. 
The lack of this fold, in combination with the shape is con- 
spicuous and characteristic for our species as belonging to 
Vertigo (Sterki). 

53. VERTIGO MOULINSIANA Dupuy. PI. 17, figs. 1, 2, 3. 

Shell dextral, minute, ovate, ventricose, obtuse at apex, 
smoothish, subperforate. Aperture semiovate, 4-toothed: 
1 tooth on the parietal wall, another on the columella, and two 
palatals, the lower one longer. Whorls 4, parted by a dis- 
tinct suture, the last doubly larger than all the others to- 
gether. Eather solid, glossy, subpellucid and of a uniform 
fulvous color. Length 2y 2 to 3, diam. 2 mm. (Dupuy). 

Marshy places near Lyons, type loc. Spain, France, Sicily, 
east to the Caucasus at Poti; north to Germany, Denmark, 
the south of England and Ireland; Pleistocene of Oester- 
gotland, southern Sweden. Living upon plants growing 
around swamps. 

Pupa moulinsiana DUPUY, Cat. extramar. Gall. Test. 1849, 
no. 284; Hist. Nat. Moll. France, p. 415, pi. 20, f. 11. 
WESTERLUND, Fauna, iii, 1885, p. 136; Malak. Blatter, 1875, 
p. 132. Alaea mmilinsiana Dupuy, CLESSIN, Die Mollusken- 



EUROPEAN SPECIES OP VERTIGO. 179 

fauna Oesterreich-Ungarns u. der Schweiz, 1887, p. 274, f. 
173. 

Vertigo moulinsiana JEFFREYS, Ann. Mag. Nat. Hist. (5), 
ii, 1878, p. 380 (description of living animal, etc.) ; (4), xix, 
p. 432. PHILLIPS, The Irish Naturalist, 1908, p. 89-93, pi. 3, 
f. 1-12 (distribution, variation, etc.). ODHNER, Arkiv for 
Kemi, Mineral, och Geol. iii, no. 33, 1910, p. 4-8, fig. 1 (occur- 
rence in Pleistocene of Sweden, and general distribution). 
STELFOX, Proc. Malac. Soc. London x, 1912, p, 43. GROVES, 
Trans. Herts Nat. Hist. Soc. i, 1880, plate ; Journ. of Conch, 
iv, p. 85. PINI, Atti Soc. Ital. Sci. Nat. xix, 1876, p. 493. 
MOQUIN-TANDON, Hist. Nat. Moll. France ii, 1855, p. 403, with 
var. personata. BOETTGER, Jahrb. Nassau. Ver. Nat., 42, 1889, 
p. 306 (Pleistocene and recent distribution). KOBELT, 
Iconogr. (2), viii, p. 94, f. 1537. TOMLIN & BOWELL, Journ, 
of Conch, xii, 1908, p. 212, f. A-F, pi. 3, f. 1-12. Vertigo 
moulinsii MOQ.-TAND., Moll. Fr., Atlas, 1855, p. 51, ref. to 
pi. 28, f. 31-33. 

Pupa desmoulimiana Dupuy, JEFFREYS, Ann. Mag. N. H. 
(2), xv, 1855, p. 29 (Clarens; Payerne; Visp; marsh west 
of Lausanne, Switzerland). Vertigo desmoulinsi GERMAIN, 
Encycl. Sci., Moll, de la France et des reg. voisins, 1913, p. 188. 

Pupa laevigata Kokeil, in GALLENSTEIN, Conch, v. Karn- 
then 1852, p. 80. PFR., Monogr. iv, 681 (marshy meadows 
near Klageiifurt, Carinthia). 

Pupa charpentieri Shuttleworth MS., KUSTER, Conch. Cab. 
p. 129, pi. 16, f. 41-43 (Heidelberg, Bern u. Toulouse). 
PFR., Monogr. iii, p. 555; iv, 681. 

Vertigo limbata Partiot MS., ace. to MOQUIN-TANDON, MolL 
France ii, 1855, p. 403. 

The ovate shell is of a slightly transparent cinnamon color, 
glossy, with only weak traces of striae. Behind the lip there 
is an opaque, rounded crest, in front of this is a pit behind 
the "auricle," or projection of the outer lip. The parietal 
lamella is high but not long; columellar lamella stouter; 2 
palatals are short, well-developed. The small basal fold, 
when present, approaches a subeolumellar position. There 
is a strong palatal callus, which sometimes is weakly denti- 



180 EUROPEAN SPECIES OF VERTIGO. 

culate or uneven. The expanded peristome is distinctly 
caught in at the middle of the outer margin. 

Length 2.6, diam. 1.6 mm. (Lyons, fig. 1). 

Length 2.3, diam. 1.45 mm. (Wicken Fen, Cambridge, figs. 
2,3). 

Having examined many specimens from England, France 
and Hungary, I am unable to agree with Dr. Boettger that 
this species is identical with V. ventricosa (Morse). The 
European species is constantly much larger, has a stronger 
palatal callus and more expanded peristome with a deeper 
indentation behind the outer lip. 

Variations in the teeth have served for the definition of 
several varieties, which have little basis in nature. Among 
specimens from Lyons (received from Terver, one of the ori- 
ginal collectors according to Dupuy), the form with a basal 
fold, var. personata, is common ; also specimens having the dis- 
tinct trace of an angular lamella, and some weak infrapalatal 
nodules as in " var. octodentata. ' ' 

An account of the habits of V. moulinsiana has been given 
by Mr. K. A. Phillips, who on October, 1907, discovered a 
colony in a marsh by the river Barrow, near Tinnahinch, Co. 
Carlow, Ireland. "I first perceived it resting on the stems 
and leaves of the tall grass, Glyceria aquatica and other 
plants, and a further search revealed some dozens of specimens 
on the leaves and t T vigs of a hawthorn bush at the edges of the 
marsh. During the second week in January I was again at 
Tinnahinch and revisited the marsh. Expecting to find the 
animals wintering, like some other Vertigoes, in the shelter 
of hollow stems and other plant-remains, I spent some time 
examining the marsh debris without finding a trace of them ; 
at last quite unexpectedly, I caught sight of a specimen on 
the trunk of a small Alder; this gave me a clue to their true 
winter habitat and I was not long in discovering numbers of 
the hardy little creatures hibernating, during the coldest 
spell of weather we had last winter on the most exposed 
parts of the lower branches at from two to three feet above 
the ground. They are gregarious and apparently viviparous, 
as there were several small communities of from four to 



EUROPEAN SPECIES OF VERTIGO. 181 

about twenty individuals huddled together, many of them 
bearing young ones of two or three whorls on the backs of 
their shells. Later on I took some handfuls of loose, dry 
leaves that had got caught by the forking of the branches, 
and found that each curled-up leaf contained several speci- 
mens both adult and juvenile. In no case did I find one 
among leaves that were wet or in a position to retain moisture. 
Its constant companion in both these situations was Succinea 
putris. ' ' 

Var. personata Moquin-Tandon is described as having the 
shell a little longer; aperture with 2 columellar folds, the 
peristome interrupted. Toulouse (Partiot). 

Var. ventrosa Heynemann. PL 17, figs. 5, 6. Shell sub- 
perforate, very shortly ovate, smooth, glossy, chestnut- 
fulvous; spire conic, rather obtuse. Whorls 4, somewhat 
convex, the last slightly compressed at the base. Aperture 
obliquely cordate, 6 or 5 toothed: 2 or 1 parietal teeth, 2 
columellar, 2 palatal. Peristome a little expanded, the mar- 
gins joined by a thin callus, right margin sinuous, impressed 
outside above the middle. Length 2*4, diam. 1% mm., ap. 
% mm. long. (Heynemann). 

On reeds of a pond near Frankfort a. M. 

Vertigo ventrosa HEYNEMANN, Malak. Blatter ix, 1862, p. 
11, pi. 1, f. 6-8 (shell and teeth). Pupa ventrosa PFEIFFER, 
Monogr. viii, p. 406. 

Distinguished by the presence of an angular lamella and a 
basal fold; the former and often the latter being absent in 
typical moulinsiana. 

Westerlund's var. octodentata (Fauna Europaea 1878, p. 
195; Fauna Pal. Eeg. iii, 1887, p. 137) has 2-2-2 teeth, and 
two little denticles in the angle between columella and basal 
margin. Apparently an individual variation of ventrosa. 
Correctly the formula would be 2-1-3, as the second colu- 
mellar tooth is really the basal fold. 

530. V. moulinsiana kiisteriana (Westerlund). The shell 
is most like P. moulinsiana in form and size, but is very dif- 
ferent by the more ventricose shape, far blunter apex, the 
peristome much more broadly reflected, more strongly bulg- 



182 EUROPEAN SPECIES OF VERTIGO. 

ing out above the middle, also in the middle angularly pro- 
duced forward and externally deeply impressed; by the free 
palatal teeth, the upper one large, thickened anteriorly, emer- 
ging to the margin, the lower one broadly distant from the 
margin; 1 parietal and 1 columellar tooth (Westerl.) 

Germany: Mergentheim, Wiirtemberg (coll. Kuster). 

Pupa kusteriana. WESTERLUND, Malak. Blatter, 1875, p. 133 ; 
Fauna iii, p. 136. PFR., Monogr. viii, 407. 

54. VERTIGO PACHYGASTER Jensen. 

Shell rimate, ventricose, irregularly, finely striate, under a 
strong lens showing very fine spiral lines, glossy, greenish- 
brown. Spire broadly conic. Whorls 4%, convex, rapidly 
increasing, the penult double the height and wider than the 
preceding; in front much lower than, and behind equal to 
the last whorl. Last whorl has a strong brown-reddish 
callus behind the peristome. Aperture half-oval, 4-toothed: 
teeth 1-1-2, fold-like, short, white. Peristome simple, thin, 
of the same color, the margins separated, outer margin neither 
produced forward or impressed, regularly curved. Length 
13/4, diam. 11/5 to 1% mm. (Westerl.). 

Norway: Skien. 

V. [ertigo] pachyg osier JENSEN, Indberetning om en i Som- 
meren 1870 foretagen Reise i Kristiania og Kr.-sands Stift, 
etc., 1872 (p. 69 of separate copy), in Nyt Magazin for 
Naturvidenskaberne, xix, 1873, p. 171. Pupa gmvida 
WESTERLUND, Fauna Moll. terr. et fluv. Sveciae, Norwegise 
et Danise, 1873, p. 610; Malak. Bl. xxii, 1875, p. 328. 
PFEIFFER, Monogr. viii, 399. Vertigo pachygastra Jensen, 
WESTERL., Synopsis Moll, extramar. Pal., 1897, p. 120. 

The name of this species was changed by Westerlund on 
account of the earlier Pupa pachyg aster, which is an Abida-, 
it does not appear to be preoccupied in Vertigo, and has been 
restored in his later Synopsis. 

55. VERTIGO LOROISIANA (Bourguignat). PL 16, fig. 15. 

Shell dextral, minute, ovate-cylindric, rimate, rather fragile, 
smooth, but under the microscope some fine striae are visible ; 



EUROPEAN SPECIES OP VERTIGO. 183 

of a corneous-fulvous tint. Summit obtuse, smooth, and of a 
paler tint. 6 convex whorls, regularly increasing, separated 
by a deep suture. Aperture slightly oblique, rounded-lunate, 
having 4 strong, elevated denticles: one in the middle of 
the convexity of the penult whorl, a second on the columellar, 
2 other palatals on the internal callus of the peristome. 
Peristome double, the first, interior, is indicated within by a 
whitish callus, and externally by a swelling; the second peris- 
tome is simple, acute and a little reflected. Between the two 
peristomes the last whorl is a little constricted, this being 
very marked at the base. The margins are united by a nearly 
invisible callus of the same color as the rest of the shell. 
Length 2y 2 , diam. 1 mm. (Bgt.). 

France: around Vannes (Morbihan), under stones in shady 
places. 

Pupa loroisiana BOURGUIGNAT, Malac. terr. et fluv de la 
Bretagne, 1860, p. 65, pi. 2, f. 7-9. Pupa laroisiana Bgt., 
WESTERLUND, Fauna iii, 1887, p. 138. Vertigo laroisiana Bgt., 
WESTERLUND, Synopsis, 1897, p. 121. 

The strong palatal callus and external crest, the rather 
elongate shape and four teeth appear to be the salient charac- 
ters of this form, which is more slender than the related 
V. moulinsiana according to Bourguignat's account, by which 
it is solely known. 

56. VERTIGO GRAELLSIANA Servain. 

Shell minute, rimate-perforate, oblong, very glossy, smooth, 
subdiaphanous, reddish-corneous. Spire rather long, obtuse 
at summit, the apex large. Whorls 5, convex, regularly and 
slowly increasing, separated by a rather deep suture; the 
last whorl small, rounded, straight above, angular below, 
around the perforation, the angle confluent with a swelling 
behind the peristome; contracted and then gibbous behind 
the lip, the gibbosity paler, strongly convex, transverse, join- 
ing the basal angle. Aperture vertical, semiovate or sub- 
trigonal, wide above, narrowed and angular below, 4-plicate, 
as follows : one median, lamellif orm parietal ; one strong colu- 
mellar ; and two equal palatals, in the middle of the convexity 



184 EUROPEAN SPECIES OP VERTIGO. 

of the outer margin, and visible outside. Peristome lipped 
within, patulous and reflected throughout, the outer margin 
sinuated; columellar margin oblique, somewhat straightened; 
margins approaching. Length 2%, diam. 1^4 mm., apert. 
y 2 mm. high, 14 wide (Servain). 

Spain: drift debris of the Ebre at Saragossa and of the 
Xenil at Grenada. 

Vertigo graellsiana SERVAIN, Etude sur les Moll. rec. en 
Espagne et en Portugal, 1880, p. 124. 

How this differs from V. moulinsiana, has not been made 
clear. 

57. VERTIGO AFRICA Bourguignat. PL 16, fig. 14. 

Shell rimate, small, globose-oblong, obtuse, shining, sub- 
pellucid, fulvous-corneous, under the lens very sharply sub- 
striatulate. Spire tapering, obtuse, apex smooth, minute, 
obtuse; whorls 6, convex, regularly increasing, separated by 
an impressed suture, the last slightly larger, tapering, the 
base compressed, anteriorly ascending very little. Aperture 
slightly oblique, semioblong, 4-plicate : 1 parietal, 1 columellar 
and 2 palatal, of which the lower is more immersed. Peris- 
tome acute, with a whitish thickening within, expanded, es- 
pecially at the base, externally encircled by a whitish lip 
[crest] ; outer margin sinuous, arcuate in front and forming 
a sulcus externally at the periphery ; the margins joined by a 
scarcely visible callous. Length 3%, diam. 2 mm. (Bgt.). 

Algeria: Alluvium of the Oued Isser, near Algiers, Let- 
ourneux. 

Vertigo aprica BGT., Malac. de 1'Algerie ii, 1864, p. 101, 
pi. 6, f. 36-38. 

58. VERTIGO CODIA Bourguignat. PI. 16, fig. 12. 

Shell minute, perforate, globuloid, ventricose, glossy, sub- 
pellucid, smooth, corneous. Spire obtusely tapering, the apex 
paler, obtuse. "Whorls 5^ to 6, rapidly increasing, separated 
by a deep suture, the penult longest, inflated, rotund; the 
last tapering, compressed at the base, very little ascending 
in front. Aperture slightly oblique, lunate, trigonal, nar- 



EUROPEAN SPECIES OF VERTIGO. 185 

rowed at base, 4-plicate: 1 lamelliform parietal, 1 dentiform, 
deeply placed columellar and 2 palatals, the upper one emerg- 
ing, dentiform, the lower immersed, lamelliform. Peristome 
white lipped within, expanded especially on the basal and 
columellar margins. Outer lip sinuous and anteriorly arcu- 
ate, having a little sulcus at the periphery; the lip margins 
united by a rather strong callus. Length 2%, diam. 2 mm. 
(Bgt.). 

Algeria : alluvium of the Oued Mazaf ran and the Harrach, 
near Algiers. 

Vertigo codia BGT., Malac. Algerie ii, 1864, p. 103, pi. 6, 
f. 39-41. 

Vertigo codia differs from V. aprica by the globulose form, 
ventricose, not oblong, by the much more rapid increase, the 
far more swollen penult whorl, smaller, more tapering last 
whorl, the triangular aperture, etc. (Bgt.). 

59. VERTIGO BRIOBIA Bourguignat. 

Shell very minute, rimate, oblong, glossy, subpellucid, 
smooth, corneous; spire obtusely tapering, apex obtuse. 
Whorls 5, a little convex, regularly increasing, separated by 
an impressed suture, the last somewhat compressed basally, 
straightened at the aperture. Aperture slightly oblique, a 
little lunate, oblong-subtrigonal, 4-plicate; 1 lamelliform 
parietal, 1 dentiform columellar and 2 palatals, the lower 
stronger. Peristome straight, acute, white-lipped within, 
lightly expanded at the columellar margin, and outside en- 
circled with a paler lip [crest], swollen below; margins 
joined by a callus. Length 2, diam. 1 mm. (Bgt.). 

Algeria : debris of the Harrach near Algiers. 

Vertigo ~briobw, BGT., Malac. de 1'Algerie ii, 1864, p. 313. 

60. VERTIGO MICROLENA Bourguignat. PL 16, fig. 13. 

Shell rimate, very minute, oblong-globose, glossy, subpel- 
lucid, corneous, smooth; spire obtusely tapering, the apex 
minute, paler. Whorls 5, convex, rather rapidly increasing, 
separated by an impressed suture, the last slightly larger, 
tapering, somewhat flattened at the periphery behind the 



186 EUROPEAN SPECIES OF VERTIGO. 

outer lip, constricted, especially in the lower part (as well 
as having a hanging swelling, as though a blister had been 
raised) ; at the base lightly compressed and straightened in 
front. Aperture vertical, lunate, semirotund, ringent, 5- 
plicate: 1 strong parietal, 2 columellar (the upper lamelli- 
form, strong, the lower small, dentiform), and 2 strong 
palatals. Peristome expanded, white-lipped deep within, ex- 
ternally at the periphery encircled with a whitish lip [crest] 
which is swollen below. Outer margin straight; margins 
joined by a callus. Length 2, diam. 114 mm. (Bgt.). 

Algeria: debris of the ravine of Chabet-Beinan, near Cape 
Caxine, west of Algiers ; debris of the Harrach at Algiers. 

Vertigo microlena BOURGUIGNAT, Malac. Algerie ii, 1864, 
p. 104, pi. 6, f. 42-44. 1 Pupa pygm&a MORELET, Journ. de 
Conchyl. iv, 1853, p. 292. 

61. VERTIGO EREMIA (Westerlund). 

Shell tumid-ovate, ventricose, brown; smooth, the last 
whorl rugose-striate anteriorly. Whorls 5, convex, the penult 
twice as large as the antepenult, the last swollen outwardly, 
a little shorter. Aperture somewhat semirotund, 3-toothed, 
the teeth rather strong, 1-1-1 ; parietal tooth compressed, 
columellar tooth conic, palatal tooth thick, tuberculiform, 
united with a strong white palatal callus. Margins of the 
peristome united by a callus joined to the apertural callus, 
the outer margin arched above, somewhat straightened below. 
Length 2, diam. 1% mm. (Westerl.). 

Sweden : Medelpad at Ange. 

Pupa eremia WESTERL., Acta Soc. pro fauna et flora 
Fennica, xiii, no. 7, 1897, p. 67. 

Described from one specimen. Appears to resemble the 
American V. tridentata, but there is no crest behind the 
peristome. 

Group of V. modesta. 

The "auricle" or lip-point is but little developed; teeth 
1-1-2 to 0-0-0, small when present, and with a tendency to be 
reduced or lost in many of the species. Chiefly boreal or 
mountain forms. 



EUROPEAN SPECIES OF VERTIGO. 187 

62. VERTIGO INERMIS (Westerlund). 

Shell umbilical e-perf orate, ovate, very finely striate, glossy, 
reddish , horn-color, obtuse at apex. Whorls 5, convex, the 
penult (viewed from the right side) half as high in front as 
behind, the last double as high in front as behind on account 
of the deep suture strongly ascending, to the aperture; last 
whorl nearly horizontal at the base, almost equally as wide as 
the penult. Aperture distinctly lateral, oval, narrowed to- 
wards the base, toothless; margin very weakly lipped within, 
a broad, thin deposit of callus between the margins. Length 
2, diam. 1% mm. (West.). 

Siberia: north of Seliwarinskoje, about 70 N. Lat. 

Pupa inermis WESTERLUND, Siberiens Land- och Sotvatten- 
Mollusker, in Kongl. Sv. Vet. Akad. Handlingar, xiv, no. 12, 
1877, p. 103 ; Fauna iii, 1887, p. 130. Vertigo inermis WEST., 
Synopsis, 1897, p. 115. 

63. VERTIGO DIC^A (Westerlund). 

Shell cylindric-ovate, seen to be densely striatulate under 
a strong lens, yellowish corneous. Whorls regularly increas- 
ing, a little convex, separated by a rather deep, oblique suture, 
ascending below, back of the last whorl obliquely flattened a 
little, strongly constricted, dilated on both sides in front. 
Aperture semiovate, toothless, parietal wall oblique, outer mar- 
gin regularly arcuate, columella dilated, spreading. Length, 
1%, diam. 1% mm. (Westerl.). 

Sweden : Dalbyo, Sodermanland. 

Pupa diccza WEST., Acta Soc. pro Fauna et Flora Fennica, 
xiii, 1897, no. 7, p. 64. 

64. VERTIGO CELATA (Westerlund). 

Shell perforate, cylindric-conic, apex obtusely conic ; densely 
striatulate rufous-corneous. Whorls 6i/2> slowly increasing, 
very convex, or swollen-convex, separated by a deep suture 
which is subhorizontal throughout, penult and antepenult 
whorls equal, the last scarcely or slightly larger, rotund at 
base. Aperture small, semioval, wholly toothless. Peristome 



188 EUROPEAN SPECIES OP VERTIGO. 

simple, thin, the outer margin arcuate, unexpanded, colu- 
mellar margin a little straightened, strongly reflected. Length 
2%, diam. 1 mm. (Westerlund, 1894). 

Sweden, the special locality unknown. 

Pupa (Alaea) celata WESTERL., Nachrblatt D. M. Ges. 1894, 
p. 173. 

Belongs to the little group of P. genesi Gr., eggeri Gr. and 
inermis W., differing from all by its shape, the number and 
the different growth of the whorls (Westerl.). 

Westerlund subsequently (1897) stated that this descrip- 
tion, being inaccurate, was to be deleted and the following 
substituted. It appears likely that the two descriptions ap- 
ply to two species; but as neither has been figured, and no 
specimens are at hand, this surmise cannot be controlled. 

Vertigo celata W. Shell subperspectively perforate, obese, 
cylindric-ovate, obtuse, brown-reddish, whitish at apex, densely 
striate under a lens. Whorls 5%, convex, separated by an 
impressed suture which is oblique above, horizontal below ; the 
three last very slowly increasing, the antepenult nearly double 
the height of the preceding, the last two almost equal; last 
whorl almost equal in height in front and behind, the back 
regularly convex. Aperture semiovate, toothless, parietal 
margin subhorizontal ; margins unexpanded, the outer regu- 
larly arcuate. Length 2, diam. 1% mm. (Westerlund, Acta 
Soc. pro fauna et flora Fennica, xiii, no. 7, 1897, p. 64). 

Sweden: Dalbyo, Sodermanland. 

65. VERTIGO REGULARIS (Westerlund). 

Shell openly perforate, cylindric, very obtuse, smooth, 
rufous-corneous. Whorls 6, somewhat tumidly convex, the 
upper ones very narrow, the antepenult one-third higher than 
the preceding, equal to the penult, the last a third shorter 
than the penult, at the back scarcely, in front distinctly higher, 
rotund at base ; suture immersed [ ? impressed] , nearly hori- 
zontal throughout. Aperture toothless, truncate-ovate, the 
parietal wall a little oblique, the margins distant and separ- 
ated, equally curved, the outer margin straight, columellar 
margin shortly reflected. Length 2, diam. 1% mm. (West.). 



EUROPEAN SPECIES OF VERTIGO. 189 

Turkestan: confluence of the Eadonak with the Bartang 
river (A. Kasnakow). 

P.[upa] (Vertigo) regularis WEST., Annuaire Mus. ZooL 
Ac. Imp. Sci. St.-Petersb., iii, 1898, p. 166. 

"Belated to P. celata of middle Sweden." One specimen. 

66. VERTIGO EGGERI (Gredler). 

Shell umbilicate, ovate-conic, the spire much narrowed, un- 
evenly substriatulate, glossy, brown-rufous. Whorls 5, con- 
vex, rapidly increasing, the last somewhat widened, shortly 
ascending at the insertion. Aperture ample, semioval, tooth- 
less. Peristome a little reflected, whitish bordered, the outer 
margin subangularly impressed and more produced in the 
middle, arcuate above at the insertion, the margins joined 
by an indistinct parietal callus. Length 2%, width 1% 
mm. (Gredler). 

Tyrol: in the so-called Barental bei Steinegg, near Bozen, 
under damp moss. 

Pupa (Vertigo) eggeri GREDLER, Nachrbl. D. M. Ges. xxii, 
1890, p. 41. 

V. eggeri has much greater dimensions than V. geiiesii, and 
differs moreover by the totally different habitus, by its conic 
spire and more distinct striation, but it approaches genesii 
in the strong convexity of the whorls, in the coloring (also 
that of the peristome), and in lacking denticulation (Gredler). 

67. VERTIGO ARCTICA (Wallenb.). PL 10, figs. 7, 8. 

Shell dextral, rimate, ovate, thin, smoothish, somewhat 
glossy, pellucid, brownish-tawny. Whorls 5 to 5%, convex, 
the last nearly two-fifths the altitude, rounded at base, an- 
teriorly having a somewhat swollen crest. Aperture slightly 
oblique, semiovate or piriform, obstructed by 3 teeth : in the 
middle of the parietal wall, on the columella, and a smaller 
one in the palate (frequently wanting) ; peristome spreading, 
slightly labiate, the margins joined by a callus, the right 
margin very strongly curved above, columellar margin some- 
what dilated, spreading. Length 2.5, diam. 1.5 mm. 
(Wallenb.). 



190 EUROPEAN SPECIES OP VERTIGO. 

Lapland : Walli-corso, a narrow ravine between Walli-f altet 
and Gaskaiwo, at the upper limit of trees; also one of the 
islands near Quick jock (Wallenberg). Bohemia; at high ele- 
vations in the Tatra and Austrian Tyrol (as P. tirolensis). 
It has also been reported from the Chukchi Peninsula and 
Alaska, but these specimens may be referable to V. modest a. 

Pupa arctica WALLENBERG, Malak. BL, v, 1858, pp. 32, 99, 
pi. 1, f. 3&-c, 4. PFEIFFER, Monogr. vi, 325. REINHARDT, SB. 
Ges. Nat. Fr., Berlin, 1883, pp. 32, 38 (Chukchi Penins.; 
Emma Harbor) . WESTERLUND, Vega-Exped. Vet. lakttag. iv, 
1887, pp. 152, 155, 157, 163 (Lapland, Siberia, Port Clarence, 
Alaska). Vertigo arctica Wallenb., WESTERLUND, Mai. Bl. 
xiv, 1867, pp. 201, 202. BABOR & NOVAK, Nachrbl. 1909, p. 
147. KOBELT, Iconogr. (2), viii, p. 93, f. 1535. Pupa tiro- 
lensis GREDLER, Verh. Zool-bot. Ges. in Wien, xix, 1869, p. 912. 

Wallenberg states that out of 20 specimens apparently adult, 
only 2 had the palatal fold developed. 

It has never been made clear how V. arctica differs from 
weak-toothed races of V. modesta; especially since Wester- 
lund, who should certainly know V. arctica, identified it from 
Port Clarence, Alaska. Specimens possibly referable to V. 
arctica, collected at Norton Sound, Alaska by Mr. R. C. Mc- 
Gregor, have three teeth as in V. arctica, differing from typi- 
cal V. modesta by lacking an upper palatal fold, and com- 
pared with typical modesta, by the smaller size of the teeth. 
The example figured measures, length 2.65, diam. 1.35 mm. 
(page 124, fig. 3). I have not seen specimens of arctica from 
Lapland or any part of Europe, and cannot therefore indi- 
cate how it differs from V. modesta, if at all. 

Westerlund placed Pupa tirolensis in the synonymy of 
arctica after comparison of a specimen received from Gredler, 
and Kobelt has referred Pupa tatrica to the same species. 
The descriptions of these forms follow. 

67 a. Vertigo arctica extima (West.). 

Shell broadly perforate, ovate-oblong, brown-corneous, 
slightly shining, smooth; whorls 5%, convex, rather slowly 
and regularly increasing, rounded at base, encircled with a 



EUROPEAN SPECIES OP VERTIGO. 191 

thin callus of the same color behind the aperture; suture 
strongly ascending anteriorly. Aperture little oblique, semi- 
ovate, rounded at the base, entirely toothless; peristome 
slightly spreading, the right margin strongly curved above; 
columellar margin a little straightened, reflected-spreading 
above; the right margin thinly labiate within. Length 2% 
to 3, diam. iy 2 to 1% mm. (Westerlund). 

Siberia: Baklanowskij, Jenesei, lat. 64 25' N. 

Pupa (Vertigo) 'arctica Wallenb., var. extima WESTERLUND, 
Nachr. d. Malak. Ges., viii, 1876, p. 99; Sibiriens Land- och 
Sotvatten-mollusker, p. 42, in Kongl. Svenska Vet. Akad. 
Handl. xiv, no. 12, 1877. 

The shell appears to be larger than other toothless forms 
of the V. modesta stock, about the size assigned for V. hoppii. 

67&. Vertigo arctica tatrica (Hazay). 

Shell small, dextral, cylindric, obtuse, corneous-buff, glossy. 
Aperture semiovate, 2 or 3 toothed. Whorls 5%. Alt. 2, 
diam. 1 mm. 

The small shell has a long-cylindric shape, is glossy, trans- 
lucent, of yellowish horn-color; spire blunted. The 5% con- 
vex whorls increase very slowly, the last being scarcely notice- 
ably wider than the penult, occupying scarcely % the length 
of the shell. Aperture semiovate, having 2 or 3 denticles: 
one sharp, lamelliform, stands on the parietal wall, a smaller 
on the columella and a small punctiform tooth on the inner 
wall [of the outer lip] ; the last is often lacking. The outer 
margin is evenly curved, not impressed (Hazay). 

Carpathians: Hohen Tatra, in Kotlina valley, on old tree 
trunks. 

Pupa tatrica HAZAY, Jahrb. d. Malak. Ges. xii, 1885, p. 
32 ; Eszaki Karpat. p. 356. 

This species stands nearest to Pupa leantina Gredl., which 
is said to have 2 denticles, but differs by its conic shell with 
only 41/2 whorls and the impressed peristome. It is distin- 
guished from Pupa arctica Wallenberg, which also has 3 
denticles, by the shape, increase of the whorls, the peristome 
not bent in (Hazay). 



192 EUROPEAN SPECIES OF VERTIGO. 

67 'c. V. arctica tirolensis (Gredler). 

Shell umbilicate ovate, obtuse at the apex, obsoletely stri- 
ate, glossy, pellucid, rufous-corneous. 5 convex whorls. 
Aperture oblique, cordate, bidentate, the columellar tooth 
short, acute, the parietal plicifonn. Peristome a little ex- 
panded, furcate, margins joined by a very thin callus, the 
right margin slightly inflected in the middle and somewhat 
projecting forward. Length 1%, diam. % lines (Gredler). 

Tyrol : Rodlerberg near the Peitler Kof el, at the upper limit 
of trees, under stones. 

Pupa tirolensis GREDLER, Verh. k. k. zool.-bot. Ges. Wien, 
xix, 1869, p. 912. 

68. VERTIGO RONNEBYENSIS (Westerlund). PL 17, fig. 14. 

Shell deeply perforate, long-ovate, regularly finely striate, 
very glossy, reddish-brown. Whorls 5%, convex, the last 
about equal to the penult, which is a third higher than the 
preceding whorl, which is double the height of the next 
earlier; last whorl has a transverse callus of the same color 
near the aperture. Suture very oblique, ascending to the 
aperture. Aperture quite obliquely piriform, excised by the 
very oblique parietal wall, 4-toothed: 1 parietal lamella, 1 
conic tooth at the lower end of the sharply emerging, dark- 
colored columella; 2 short, widely separated, deeply im- 
mersed palatal folds. Margins delicately united, the outer 
margin weakly arcuate, nearly straight, the columellar margin 
broadly reflected. Length 2%. to 2y 2 , diam. 1% to 1% mm. 
(WesterL). 

Sweden: Pehrsborg, near Ronneby, under rotten beech 
leaves. Northern Germany, Bohemia. 

Pupa ronnebyensis WESTERLUND, Expose Grit., in Nova 
Acta Reg. Soc. Sci. Upsal. (3) viii, 1871, p. 94; Fauna iii, p. 
135. D. GOLDFUSS, Nachrbl. d. m. Ges. xxvi, 1894, p. 216; 
xxvii, 1895, p. 100 (Tegel near Berlin; Cladow, Neumark; 
near Landsberg a. Warthe, Spechthausen bei Eberswalde, 
Oderberg in der Mark and from the Oscher Walde (Kreis 
Schwetz) in West Prussia). Vertigo ronnebyensis West., 



EUROPEAN SPECIES OF VERTIGO. 193 

MERKEL, Nachrbl. d. m. Ges. 1887, p. 13-16 (Tegel bei Berlin). 
BABOR & NOVAK, Nachrbl. 1909, p. 147 (Bohemia). 
GEYER, Unsere Land- und Susswasser-Mollusken, 1909, p. 55, 
pi. 5, f. 18, 19 (dist. in Germany). 

This species approaches P. alpestris by the aperture pro- 
vided with 4 teeth, but it differs considerably by the shell be- 
ing of dark color, oblong-ovoid, and noticeably larger, nearly 
as large as a P. edentula of 6 whorls, the palatal teeth short 
and higher, etc. (Westerlund). 

The figure is from Geyer. 

69. VERTIGO DALIACA (Westerlund). 

Shell oblong-cylindric, with rounded summit, smooth, red- 
dish horn-colored, somewhat glossy. Whorls 5, somewhat 
convex, the first two small, the second half as long as the 
third, the three last regularly increasing, the last slighHy and 
very slowly ascending, rounded, not as wide as the penult. 
Suture oblique. Aperture rounded, with one strong, conic, 
acute tooth high on the columella ; parietal margin but slightly 
oblique ; the margins equally arcuate, outer margin somewhat 
angular above. Length 1%, diam. 1% mm. (Westerl.). 

Sweden : Dalsland. 

Pupa daliaca WESTERLUND, Fauna Pal. Reg. Binnen- 
conchylien, iii, 1887, p. 131. 

Var. dalecarlica Westerlund. Shell cylindric, brown, slightly 
striatulate ; whorls 5 to 6, slowly and regularly increasing, the 
last 2 subequal, a little convex, the last rotund at the base, 
the suture oblique in the middle, subhorizontal at the aperture. 
Aperture semiovate : 1 very short, white, deeply placed parie- 
tal tooth, one in the palate nearly at the base, nodiform, im- 
mersed, rarely with 1 or 2 punctiform, brown, very obsolete 
teeth in the middle of the palate outwardly. Columellar 
margin of the peristome subvertical, reflected, the outer mar- 
gin strongly curved above, then obliquely straightened, un- 
expanded. Length ~Ly 2 to 2, diam. 1 mm. Sweden in the 
province Dalarne, at Leksand and at Fu par. Mora. (Pupa 
daliaca var. dalecarlica Westerlund, Acta Soc. pro fauna et 
flora Fennica, xiii, no. 7, p. 64, 1897). 



194 EUROPEAN SPECIES OP VERTIGO. 

70. VERTIGO PINETICOLA (Westerlund). 

Shell rimate, ovate-cylindric, finely striate, reddish-brown, 
glossy. Spire produced, gradually tapering. 5 whorls, slowly 
increasing, convex, the last scarcely ascending in front. 
Suture deep, nearly horizontal. Aperture oblique, piriform, 
with one tooth below on the columella; parietal margin very 
oblique; peristome somewhat expanded. Length 2^-2%,, 
diam. 1% nim. (Westerl.). 

Sweden: Tenhult, prov. Smaland (E. Hemberg). 

Pupa pineiicola WESTERLUND, Expose Critique, in Nova 
Acta Reg. Soc. Sci. Upsaliensis, (3), viii, 1871, p. 97; Fauna 
iii, 1887, p. 131. 

71. VERTIGO GEMMA Westerlund. 

Shell cylindric-oblong, slowly tapering above the middle,, 
obtuse, chestnut, glossy, smoothish. Whorls 6, convex, the 
last two wide, slightly unequal, the upper ones narrow (the 
penult more than double the alt. of the preceding), last 
whorl in front dilated below, the base gibbous-crested ; suture 
oblique above, horizontal below. Aperture (margins not as 
yet wholly complete) sub triangular, narrowed below, 1 or 2 
toothed: palatal tooth nodiform, below the middle of the 
margin, a very minute denticle above the middle. Peristome 
simple, the outer margin angularly produced above the middle, 
very strongly arcuate above, a little straightened below the 
middle. Columella straight, vertical, thickened at base, the 
columellar margin dilated and reflected above, in front of the 
columella, below the columella strongly arched backward. 
Palatal and cervical callus wanting, between the penult and 
last whorls there is a transverse white line, from a thin in- 
ternal lip. Length 2%, diam. 1% mm. (Westerlund). 

Sweden: Dalbyo, Sodermanland. 

Pupa gemma WESTERLUND, Acta Soc. pro fauna et flora 
Fennica, xiii, no. 7, p. 65, 1897. 

Described from one specimen. 



EUROPEAN SPECIES OF VERTIGO. 195 

72. VERTIGO OVOIDEA (Westerlund). 

Shell ovate, ventricose, widest in the middle, reddish-yellow, 
finely striate, rather glossy. Whorls 5, convex, the upper 
ones rapidly increasing in width, the last equal in height to 
the penult in front, behind strongly sloping, far lower and less 
wide than the penult. Suture very oblique above, nearly 
horizontal below. Aperture semiovate, with three small tuber- 
cular teeth, one each on the parietal wall, columella and deep 
in the palate. Parietal wall scarcely oblique ; peristome sim- 
ple, acute, the columellar margin somewhat reflected above, 
margins delicately joined. Length 2, diam. li/4 mm. 
(WesterL). 

Sweden: Tenhult, Prov. Smaland (E. Hemberg). 

Pupa ovoidea WESTERLUND, Expose Critique, in Nova Acta 
Reg. Soc. Sci. Upsal. (3) viii, 1871, p. 96; Fauna iii, p. 131. 
PFR., Monogr. viii, 393. 

73. VERTIGO SIEVERSI (Bttg.). PI. 17, figs. 11, 12. 

Nearest related to V. siibstriata Jeffr., but ovate-turrited, 
not shortly ovate, the sculpture slighter. Shell small, ovate- 
turrite, somewhat glossy, rather silky, corneous-olivaceous, the 
apex obtuse. Whorls 5, convex, very delicately but distinctly 
striate, the last scarcely one-third the total length, encircled 
anteriorly with an annular broad but not strong callus, either 
fulvous or whitish, externally not or slightly impressed. 
Aperture truncate-ovate, 6-toothed: two pliciform palatals, 
two subequal columellars, and two parietals, the inner one 
larger. Peristome a little expanded, pale, somewhat thick- 
ened, sublabiate, the margins connected by a thin callus. 
Outer margin a little projecting and slightly impressed in the 
middle. Length 1% to 2%, diam. 1 mm. (Bttg.). 

Caucasia: Tabizhuri (Dr. Sievers) ; debris of the Psekup 
(Rosen). 

Pupa (Vertigo) pygnuza Drap. var. nitidula MOUSSON, 
Journ. de Conch. 1876, p. 143. Pupa (Vertigo) sieversi 
BOETTGER, Jahrb. d. m. Ges. vi, 1879, p. 407, pi. 10, f. 6; 
with var. punctulum, pi. 10, f. 7; Jahrb. vii, 1880, p. 141, 
with var. punctum [sic] and var. subalpestris. Vertigo 



196 EUROPEAN SPECIES OP VERTIGO. 

sieversi Bttgr. var. subalpestris BOETTGER, Bericht Senckenb. 
Nat. Ges. 1889, p. 25. Pupa sieversi Bttg., ROSEN, Nachr. d. 
m. Ges. vol. 39, 1907, p. 205. 

Constantly of a more elongate turrited form [than V. 
pygm&a], by the striation and position of the parietal tooth 
most nearly related to V. substriata (Bttg.). 

Var. punctulum Bttg. PL 17, fig. 13. Smaller, more 
ovate. Mauglis; debris of the Kura at Borshom (Sievers) ; 
also Martkopi and Abastuman (Leder). 

In all the specimens from Abastuman the lower columellar 
tooth of typical P. sieversi is wanting (Bttg.). 

Var. subalpestris (Bttg.). Differs from V. alpestris chiefly 
by the very small size, alt. 1%, diam. % mm., the more ovate 
shape, the somewhat stronger sculpture and the stronger de- 
velopment of the upper parietal denticle. 

Caucasia: on the Kasbek (type loc.) ; forest region of the 
southeastern and northern spurs of the Oschten-Fischt moun- 
tain-group (Leder). 

Dr. Boettger states that specimens from the last locality 
are somewhat larger, length 1%, diam. 1 mm.; the second 
parietal tooth is altogether lacking, and in these characters 
and the size this form is so similar to the true V. alpestris 
that without the most careful attention it would be mistaken 
for that. Besides the differential features noted above, sub- 
alpestris has the weak indication of a narrow transverse cal- 
lus in the base of the mouth, near the peristome. Were it 
not for the transitions to sieversi from the Kasbek, subalpestris 
would be united with the real alpestris. 

Vertigo sieversi was first described as P. pygmaea var. 
nitidula, by Mousson ; and I am not sure that his name should 
be rejected. Boettger gives no reason for doing so. There is, 
however, a Tornatella nitidula of Lamarck, which in modern 
nomenclature will become Pupa nitidula. Mousson 's descrip- 
tion follows. 

Pupa (Vertigo) pygmaea Drap. var. nitidula Mousson. 
Smaller, 1.5 mm. long, 0.9 diam., brown, glossy, the base not 
compressed; 2 minute palatal teeth, not produced, visible ex- 
ternally, columellar tooth single small. This appears to me 



EUROPEAN SPECIES OF VERTIGO. 197 

to be only a variety of V. pygmaeus, which in Europe also 
varies considerably. Its smaller size, slightly less cylindric 
shape, the more rounded base, the weakness of the teeth, of 
which the 2 palatals are more immersed but visible from 
the outside, and the single, rudimentary columellar distin- 
guish it from the type (Mouss.). 

Transcaucasia : Tabizhuri, Dr. Sievers. 

74. VERTIGO ALPESTRIS Alder. PL 18, figs. 1, 2. 

"Shell subcylindrical, thin and semitransparent, very 
glossy, pale yellowish-horn-color, closely and rather strongly 
striate in the line of growth; periphery rounded: epidermis 
thin. Whorls 4%, convex, but slightly compressed. Spire 
short, abrupt and bluntly pointed. Suture excessively deep. 
Mouth semioval and subangular, owing to the outward com- 
pression of the periphery: teeth four, viz.: one sharp and 
prominent tooth on the middle of the pillar [parietal wall], 
one strong and also prominent and thick tooth on the pillar 
lip, and two lamellae or plate-like teeth which are placed at 
some little distance within the outer lip, but not on any rib 
or callous fold as in V. pygm&a,; the labial teeth are visible 
on the outside, owing to the thinness and transparency of the 
shell. Outer lip rather thick very slightly reflected, not 
strengthened by any rib either outside or inside, outer edge 
abruptly reflected; inner lip somewhat thickened in adult 
specimen. Umbilicus small and narrow, but rather deep. 
L. 0.07. B. 0.04 inch (Jeffreys}. 

Length 2, diam. 1.1 mm. 

Length 1.9, diam. 1 mm. 

England: Lipwood, near Haydon Bridge, Northumberland, 
on an old wall, type loc. ; Clithero, Lancashire, etc. The Alps, 
Switzerland, the Lower Harz, etc. Norway, Sweden and 
Lapland ; in Siberia east to the Amur valley and Vladivostok. 

Vertigo alpestris Fer., ALDER, Trans. Nat. Hist. Soc. of 
Northumberland, etc., ii, 1838, p. 340. GRAY, Turton's 
Manual, p. 202, pi. 12, f. 141. JEFFREYS, British Conch., i, p. 
259. BOETTGER, Jahrb. Nassau. Ver. Nat. 42, 1889, p. 307 
(Pleistocene and recent dist.) KOBELT, Iconogr. (2), viii, p. 



198 EUROPEAN SPECIES OF VERTIGO. 

91, fig. 1531. DEAN and KENDALL, Journ. of Conch, xii, 1908, 
p. 209 (distribution in England, and frequent association with 
V. pusilla). 

Pupa alpestris Fer., PFR., Monogr. viii, 1877, p. 396. 
MOUSSON, Journ. de Conch. 1887, p. 19 (Wladiwostock). 
WESTERLUND, Fauna, iii, 1887, p. 132 ; Kongl. Sv. Vet. Akad. 
Handlinger, xiv, no. 12, 1877, p. 44 (Jenissei, Siberia, 6 m. s. 
of Podk. Tunguska, 61 N. lat. Lapland N. to Quickjock, 
67-68 N. lat.). Pupa alpestris Alder, STEEKI, Nachrbl. d. 
Malak. Ges. 1883, p. 73 (many localities in Switzerland), with 
var. elongata. 

Pupa shuttleworthiana Charpentier MS., PFEIFFER, Zeitschr. 
f. Malak. 1847, p. 148 (Bex, Switzerland) ; Monogr. Hel. Viv. 
ii, 355; iii, 555; iv, 680; vi, 327. KUESTER, Conchyl. Cab., 
Pupa, p. 128, pi. 16, f. 35-37. WALLENBERG, Malak. Blatter 
v, 1858, p. 101, pi. 1, f. 5 (Quickjock, Lulea-Lapland). 
SCHRENCK, Amurl. Moll., 1867, p. 657. GREDLER, Yerh. ZooL- 
bot. Vereins Wien, 1856, p. 128; Nachrbl. d. m. Ges. 1879, 
p. 112, with var. mitis (occurrence in the Tyrol). SCHRENK, 
Reisen u. Forsch. im Amurlande ii, p. 657 (Krasnoyarsk, 
around Irkutsk, Kultuk on L. Baikal, lower Amur above 
Kidsi). Vertigo shuttleworthiana Charpentier, KOBELT, 
Iconogr. (2), viii, p. 94, f. 1536. 

Pupa pygmaea var. alpestris MAACK, Bull. Acad. St. Pet. 
xi, 1853, no. 263, 264. 

Pupa (Vertigo) parcedentata (in part) SANDBERGER, Verh. 
Phys.-Med. Ges. Wiirzburg, n. F., xx, 1887, with varieties 
quadridens, pi. 8, f . la-c ; tridens, f . 2 ; and adversidens, f . 3. 

The total absence of a crest or palatal callus and the some- 
what straightened but not in-bent outer lip are characteristic. 
Wallenberg found the Lapland specimens a little larger than 
those of England, 2.2 x 1.2 mm. 

Var. mitis Westerlund. Aperture bidentate, one parietal 
and one columellar tooth; or tridentate, one parietal, one 
columellar and one lower palatal. Sweden. (Pupa alpestris 
var. mitis Westerl., Fauna Europaea, fasc. 1, 1876, p. 197; 
Pupa alpestris forma mitis, bidentata and tridentata, WESTER- 
LUND, Fauna iii, 1887, p. 132). 



EUROPEAN SPECIES OF VERTIGO. 199 

It has been reported from Transylvania by Clessin, and has 
been described as P. parcedentata var. tridens (see pi. 18, fig. 
13) and perhaps var. adversidens (pi. 18, fig. 16) by Sand- 
berger, who records these, together with var. quadridens 
(pi. 18, fig. 14, = typical alpestris) from the Rhine valley loess. 

It may be presumed that Westerlund did not intend the 
terms bidentata and tridentata as names, but merely descrip- 
tive of the forms he included under var. mitis. 

Var. elongata Sterki. A few examples of a beautiful, large, 
cylindric form, of brownish color, rather smooth, from Trous 
(Canton Graubiinden) [Switzerland] are in the Mousson col- 
lection, collected by Blauner, and labelled pygmaea (Sterki, 
Nachrbl. d. Malak. Ges. 1883, p. 73). 

The identity of Pupa shuttleivorthiana (pi. 18, fig. 3, copied 
from Steenberg) with V. alpestris has been affirmed by 
Jeffreys (British Conchology, p. 261) and by Boettger 
(Nachrbl. d. Malak. Ges. 1882, p. 19), who declared them 
absolutely identical. Kobelt, in the Iconvgraphie, remarks 
that "in the local faunas this species has frequently been 
misunderstood ; it is hardly anything other than a four-toothed 
race of V. alpestris.'" He states that the original locality is 
Bex, Switzerland, and it has also been reported from Belte- 
berga, Sweden, by Westerlund. 

74&. Vertigo alpestris heldi (Clessin). PL 18, fig. 4. 

Shell rimate, turrited, irregularly and very finely striate, 
of reddish-brown color, glossy. Whorls 6, slowly increasing, 
rather convex ; the first 3 form a blunt summit which is about 
% the length of the shell; the last 3 are of nearly equal 
height and form the remaining cylindric part of the shell. 
The last whorl is neither calloused nor contracted preceding 
the aperture Aperture about 14 the length of shell, arcuately 
convex, somewhat impressed on the outer side, the impression 
running as a groove-like depression for a short distance on the 
last whorl. Toothed, the teeth reddish, very weak and placed 
deep in the throat ; 1 tooth on the middle of the parietal wall, 
1 on the columella, 2 very weakly developed, frequently want- 
ing, on the palatal wall. Peristome continuous, somewhat 



200 EUROPEAN SPECIES OF VERTIGO. 

expanded, little thickened. Length 4-4.5, diam. 1 mm. 
(Clessin, 1877). 

Germany : drift debris of the Danube, Regensburg, Bavaria 
type loc. Giinzburg on the Danube, Schonthal on the Jagst 
and Neckarthailfingen on the Neckar (Goldfuss). Ireland? 

Pupa (Vertigo) heldi CLESSIN, Nachrbl. d. Malak. Ges. 
1877, p. 49; Deutsche Exk.-Moll.-Fauna, edit. 2, p. 266, fig. 
156. OBERNDORFER, Nachrbl. 1898, p. 69. Pupa (Alaea) 
heldi CLESSIN, Berichte naturwiss. Vereines zu Regensburg 
xiii, Heft fur 1910-1911, p. 83. Pupa pygmaea var. elongata 
GEYER, Jahresh. Ver. Vaterl. Naturk. Wiirttemberg 1894, p. 
107; identical with P. heldi, same Jahresh. vol. 36, 1907, p. 
424. Debris of the Danube, Neckar, etc. 

? Vertigo heldi TOMLIN, Irish Naturalist xii, 1903, p. 110; 
Journ. of Conch, x, 1903, p. 307; Proc. Malac. Soc. London 
xii, 313 (an abnormal V. pygmaea). B. B. WOODWARD, t. c., 
p. 361 (doubts the identification). 

Clessin found only dead specimens in the Danube debris, but 
thought it must live not far away. It was found living in the 
Wurttemberg Jura. Dr. Geyer believes heldi to be an over- 
grown V. pygmaea. He notes that "full-grown specimens 
with completed peristome and teeth are rare. Contrary to 
the other Vertigos, the palatal teeth appear first, and the 
parietal tooth may even continue absent, this condition lead- 
ing to P. haeussleri Sterki. ' ' 

V. heldi was first reported from Ireland (Ballintoy, Co. 
Antrim) by Mr. B. Tomlin, whose specimen was examined by 
Dr. Boettger, who believed heldi to be a variety of alpestris. 
In his second note, attention is called to the smooth and glossy 
surface of the Irish shell while heldi is described as striate- 
"Placed side by side with a typical V. alpestris it differs in 
possessing an extra whorl and larger dimensions in every way, 
in its reddish-brown color, in its feebler dentition, and in the 
shape of the mouth, the outer margin of which slopes very 
obliquely from right to left, thus producing the sort of 
channel at the top to which Clessin refers." Subsequently 
Mr. Tomlin thought the Irish shell an abnormal V. pygmaea. 
This harmonizes with the ideas of Geyer, noticed under 
V. pygmaea. 



EUROPEAN SPECIES OF VERTIGO. 201 

74&. Vertigo alpestris leontina (Gredler). PL 18, figs. 5, 6. 

The shell is very small, subperforate, ovate-cylindric, obtuse, 
very delicately striate, very glossy, pellucid, whitish-corneous. 
Whorls 41/2, convex. Aperture semioval, 5-plicate, plicae un- 
equal ; 1 parietal, 1 columellar, 3 very small palatals, the lower 
one punctiform, the median and upper lamelliform. Peris- 
tome a little reflected, thickened, white, margins joined by a 
thin callus, the right margin sinuous. Length %, diam. % 
of a line (Gredler}. 

Tyrol : Lienz, scarce, at the foot of the Tristacherwand. 

Pupa leontina GREDLER, Verh. zool-bot. Ver. Wien 1856, 
p. 127, pi. 2, f. 4. WESTERLUND, Malak. Bl. xxii, p. 128 (iden- 
tical with Pupa schultzii Phil.). PFR., Monographia iv, 680. 

Among the Tyrol Pupas this new species stands next to 
P. shuttlewvrthiana [alpestris}, but it is smaller, lighter 
colored, more transparent, and has half a whorl less. The 
palatal folds (though the specimen is full grown) are far 
more weakly developed and differ in number ( Gredler) . 

Pupa isarica Westerlund. PI. 18, figs. 7, 8. 

2-toothed, without any trace of palatal folds, though full 
grown, 1.7 x 1.1 mm. Bavaria : drift debris of the Isar, one 
specimen. 

Pupa leontina Gredler, CLESSIN, Deutsche Excursions- 
Mollusken-Fauna, 1876, p. 216, f. 122, not of Gredler. Pupa 
isarica WESTERLUND, Fauna Pal. Reg. Binnenconch. iii, 1887, 
p. 134 (based upon Clessin's note and figure) ; Synops. Moll, 
extramar. Reg. Pal., 1897, index p. 11. Vertigo isarica West., 
KOBELT, Iconogr. (2) viii, p. 95, f. 1539. 

Except by lacking palatal folds, this form agrees with V. 
leontina. It is known by a single example, possibly abnormal. 

74c. Vertigo alpestris helvetica (Westerlund). 

Shell rimate-p erf orate, long-cylindric, with conically taper- 
ing apex, light reddish horn-color, very finely striate, whorls 
6!/2, somewhat convex, regularly increasing, rather high, the 
last whorl small, but little more than 1/4 the total length, an- 
teriorly impressed at the base, then tubercularly blunt keeled 



202 EUROPEAN SPECIES OF VERTIGO. 

below, behind strongly sloping. Aperture oblique, narrow, 
triangular, with 1 parietal lamella, 1 columellar lamella, in 
the middle, receding below. Peristome simple, thin; outer 
margin arcuate above, then obliquely produced to the left; 
the columellar margin short, somewhat oblique towards the 
right. Length 2%, diam. 1*4 mm. (Westerlund). 

Switzerland : based on a single specimen without definite 
locality. 

Pupa alpestris var. helvetica WESTERLUND, Fauna Europaea 
1876, p. 198. Pupa helvetica WESTERL., Fauna Pal. Reg. etc., 
iii, 1887, p. 134. 

75. VERTIGO SCHULTZII (Philippi). PL 18, fig. 9. 

Shell minute, rimate, subcylindric, obtuse, very smooth, 
glossy, fulvous ; whorls cylindric, aperture rounded, the throat 
5-toothed, single teeth on parietal wall and columella, 3 in 
the palate, the lower one smallest. Alt. 1, diam. V 12 lines; 
whorls 5 to 6 (Philippi). 

Sicily: Palermo, Dr. Schultz. 

Pupa (Vertigo?) schultzii PHILIPPI, Zeitschr. f. Malak. 1844, 
p. 106. Pupa schultzii KUSTER, Conchyl. Cab. p. 93, pi. 13, 
f. 3, 4. PFR., Monogr. ii, 359. Vertigo schultzii Phil., 
KOBELT, Iconogr. (2), viii, p. 95, f. 1538 (copied from Kiister). 

A very remarkable species of the form of P. dilucida Ziegl., 
but with the whorls more strongly convex, and without trace 
of striae. By this it compares best with Vertigo pygmaea, 
though far more slender, with more strongly convex whorls. 
The parietal fold emerges very far, while the second palatal 
fold is only weakly developed (PhUippi). 

A lost species, possibly a form of alpestris as surmised by 
Kobelt, though the locality and very smooth surface are 
against that view. 

76. VERTIGO ACHEILA Servain. 

Shell very minute, perforate, ovate, more convex on the 
left than on the right side, very glossy, smooth, diaphanous, 
corneous. Spire obtuse. Whorls 5, a little convex, slowly 
increasing, separated by a rather deep suture, the last whorl 









EUROPEAN SPECIES OF VERTIGO. 203 

moderate, convex, angular below around the perforation, above 
straight and slightly descending to the insertion of the lip, 
nearly % the total alt., thicker but not swollen behind the 
aperture. Aperture slightly oblique, trigonal, below angu- 
larly narrowed and transversely straightly thickened, 4- 
plicate, as follows: 1 strong parietal, 1 columellar at the 
upper part of the columella, and two approximate palatals, 
visible externally, the upper one stronger and reaching the 
margin. Peristome lipped within, spreading and a little re- 
flected, the outer margin a little straightened, thicker in the 
middle; the columellar margin somewhat straightened, in an 
oblique direction from left to right; margins rather remote. 
Length 1%, diam. 11/4, alt. ap. %, width y 2 mm. (Servain). 

Spain : drift debris of the Ebre at Saragossa. 

Vertigo acKeila SERVAIN, Etude Moll. rec. en Espagne et 
en Portugal, 1880, p. 125. 

"The columellar margin has an obliquely rectilinear direc- 
tion from right to left, rendering the aperture triangular and 
quite angular at the base. The aperture is, moreover, char- 
acterized by a horizontal thickening interiorly in the lower 
part, altogether identical with that which has been remarked 
in the pupa microtragus of Turkey" (Servain). 

It appears to be known only by Servain 's description. 

77. VERTIGO BUTTNERI Siemaschko. 

Shell very minute, ovate, somewhat obtuse, very elegantly 
striate, subrimate. Aperture semiovate, peristome white- 
lipped; parietal wall and palate 1-toothed. Length 2, diam. 
1.5 mm. ; whorls 4% (Siem.). 

Courland: Parvecia (H. Biittner). 

Vertigo buttneri SIEMASCHKO, Bull, de la Classe Physico- 
Math. de 1'Acad. Imp. Sci. St. Petersb., vii, no. 159, 1849, p. 
234. 

This small reddish-brown snail has great resemblance to 
Pupa ligranata, but it is a true Vertigo ; lightly striate, above 
obtuse, the aperture half-round, with thickened white margins, 
outwardly a little angular, and two-toothed. Of the teeth, 
one stands above, the other below, obliquely opposite; on the 



204 EUROPEAN SPECIES OP VERTIGO. 

right from the latter one sees sometimes another quite small 
denticle which stands nearer the outer lip. The greatest 
diameter is equal to the height of the last whorl (Siem.). 

So far as I know, this species has not been recognized by 
any subsequent author; yet it may be known under another 
name. 

78. VERTIGO GENESII (Gredler). PL 18, figs. 10-12, 17, 18. 

The shell is very small, ovate, obtuse, indistinctly, spaced 
striate, glossy purplish brown. Whorls 4%, rather convex, 
high, rapidly increasing, joined by a somewhat impressed 
suture, the penult large, almost ventricose. Umbilical open- 
ing moderate. Aperture semirotund, nearly quadratic, with- 
out any folds. Peristome scarcely expanded, thickened lip- 
like, bordered with bluish black, the margins connected by a 
very weak callus, the right margin arched at the insertion. 
Length %, diam. y 2 li ne [about 1.5 x 1 mm.] . 

Animal raven-black, the tentacles short, contracted in the 
middle, the anterior pair not indistinct (Pupa ?) . Bears itself 
quite timidly, but promptly takes courage again (Gredler). 

Tyrol : above St. Jenesien, near Botzen, at 5000 ft. (Gredler). 
Also in Germany, near Kisslegg, southern Wiirtemberg 
(Geyer). Sweden: "Westergotland (Linnarsson). 

Pupa genesn GREDLER, Tirol's Land- und Siisswasser- 
Conch., Verh. z.-b. Ver. Wien vii, 1856, p. 122, pi. 2, f. 3 
PPR., Monogr. iv, 662. Alaea genesii CLESSIN, Corresp.-Bl. 
Zool. Min. Vereins Regensburg, 1877 ; Moll, fauna Oesterreich- 
Ungarns, 1887, p. 279. Pupa genesii WESTERLUND, Nova Acta 
Reg. Soc. Sci. Upsaliensis, (3), viii, 1871, p. 97; Fauna, iii, 
p. 130 ( Oestergotland ) . Pupa (Vertigo} parcedentata var. 
genesn SANDBERGER, 1887. Vertigo genesii GEYER, Nachrbl. ' 
d. m. Ges. 1912, p. 125. STEUSLOFF, Archiv Ver. Fr. Naturg. 
in Mecklenburg, 61 Jahr. 1907, p. 68, 71 (dist. in quaternary). 
PupUla genesn Gredl., KOBELT, Iconogr. (2), viii, p. 82, 
f. 1512. 

Originally described as toothless, this species varies to forms 
with 4 teeth. It has been considered a toothless variety of 
V. parcedentata by Sandberger and Boettger, and if this view 



EUROPEAN SPECIES OP VERTIGO. 205 

is supported, the latter will stand as V. genesii parcedentata, 
as genesii was the first to be described. 

Kobelt placed the species in Pupilla on account of Gredler's 
note on the tentacles. 

D. Geyer, who collected V. genesii at the original locality, 
under the guidance of Gredler, writes as follows : 

"Vertigo genesii Gredler is not invariably toothless ('ohne 
alle Fatten' Gredler), but 4-toothed examples occur with the 
toothless ones; 1 tooth on the parietal wall, 1 on the colu- 
mella and 2 in the palate, showing through outside. These 
last two develop last, and examples about to become complete 
therefore appear to be transitional, with only 2 or 3 teeth. 
V. genesii is therefore similar in dentition to Pupilla mus- 
corum, steini f triplicates and others. The station of the snail 
is at the Sal ten near Jenesien (so the name of the village is 
pronounced, and so it appears on the charts not 'St. 
Genesien'), in the neighborhood of Bozen. The snail is not 
restricted to rocks, as Gredler stated, but also lives on wood, 
in humid forest openings. 

"V. genesii is not confined to the Salten and Tyrol, but 
also lives in Germany. Years ago I found 4 specimens in a 
meadow near Kisslegg in the Wiirtemberian Allgau, and a 
single 4-toothed Pupa in the debris of the Aach near Wolfegg, 
9 kilom. northwest of Kisslegg. * * * I see now that these 
German specimens, also from a humid but far lower station 
(at Salten about 1200, at Kisslegg 647 m.), agree completely 
with those of the Tyrol, being only a little lighter in color." 

Geyer further states that genesii and parcedentata "show 
the same variability in dentition, both having to 4 teeth, 
but in size, contour, growth and number of whorls they 
differ from one another decidedly." Yet it must be noted 
that Geyer compares with Sandberger's fig. 1, which Boettger 
considered to be really V. alpestris. Geyer considers that 
parcedentata is a synonym of alpestris; but the small form 
without columellar or upper palatal folds, which is really the 
typical parcedentata, appears to support the view of Sand- 
berger and Boettger, that parcedentata is specifically not 
separable from genesii. 



206 EUROPEAN SPECIES OF VERTIGO. 

An allied but larger form has been found in the Kartitsch 
Valley, Tyrol, by Wiedemayr (Zeits. Ferdinandeums fur 
Tirol u. Vorarlberg, Innsbruck, 44, 1900, p. 167). 

Vertigo pygmaea var. concinna Scott = V. levenensis Scott 
has been placed by Kennard and Woodward (1917) in the 
synonymy of V. parcedentata. Dr. Boettger considered speci- 
mens submitted to him to be identical with genesii. The 
original description follows. 

Rather smoother and more glossy than the typical Vertigo 
pygm&a. It differs chiefly, however, in the mouth being un- 
furnished with teeth. In this respect it agrees Avith V. eden- 
tula, but the form of the shell is certainly that of V. pygmcea, 

(Scott). 

Scotland : Kirkland marl, Leven ; Pleistocene. 

V. pygnuza var. concinna T. SCOTT, Proc. Roy. Phys. Soc. 
Edinburgh, x, p. 335, footnote, 1890; Scottish Naturalist, 
April, 1891, p. 53 (not seen). Vertigo levenensis SCOTT, 
Scottish Naturalist 1891, p. 141, n. n. for V. p. concinna 
(not seen). CF. KENNARD & WOODWARD Proc. Malac. Soc. 
Lond. vii, 119. 

The name cancinna was quite unnecessarily changed to 
levenensis on account of Pupa concinna Lowe, a Madeiran 
snail belonging to a different subfamily. 

TSa. Vertigo martini Sayn. A very small species of the size 
and nearly the shape of the short varieties of V. pygmaea, 
but having the aperture absolutely without folds or teeth. I 
do not see that there is anything in the French fauna which 
is comparable, and the Pliocene fossil forms having the aper- 
ture toothless, such as V. crossei of Hauterive, are all much 
larger and of a different shape (Sayn). 

France : Pleistocene ; marnes inferieures de Pont-Neuf , Ain 
river near its confluence with the Rhone. 

V. [ertigo] martini SAYN, Ann. Soc. Linn, de Lyon Annee 
1911, Iviii, p. 246. 

A. S. Kennard and B. B. Woodward have suggested that 
this is probably identical with Vertigo parcedentata (Al. 
Braun) (Proc. Geologists' Asso., xxviii, 1917, p. 170). It 



EUROPEAN SPECIES OF VERTIGO. 207 

will be noted that the description of V. martini, such as it is, 
agrees with that of V. co-ncinna levenensis. Its distinctness 
from V. genesii remains to be established, if it is not identical. 

78&. Vertigo genesii parcedentata (Al. Br., Sbgr.). PL 18, 
figs. 15, 19. 

The small, narrowly ovate shell has a narrow umbilical 
crevice and blunt summit; consists of 4% to 5 convex whorls 
separated by a deeply impressed suture, the last whorl form- 
ing half of the total length. First whorl entirely smooth, the 
rest having rather oblique striae, of which there are several 
finer between two stronger striae. The aperture is semi- 
ovate, obliquely truncate above, with weakly reflected mar- 
gins which are connected by a very thin callus; the right 
margin being weakly arched forward. It has either one to 
four teeth [one or two; see below] or is wholly toothless. 
Length 1.8 to 2.2, diam. 1 mm. (Sandberger) . 

Germany: Pleistocene and Alluvium of the Rhine valley, 
etc. 

Pupa or Vertigo parcedentata A. BRAUN, Amtl. Bericht 
iiber die XX Versammlung deutscher Naturforscher u. Aerzte 
zu Main, 1842, p. 143 (Wiesbaden; no description). Pupa 
(Pupilla) parcedentata A. Braun, SANDBERGER, Vorwelt, p. 
876, pi. 36, f. 25, 1875; Verh. Phys.-Med. Ges. Wiirzburg xx, 
1887, pp. 229-235, pi. 8 ; with, on p. 232, varieties quadridens, 
fig. 1 ; tridens, fig. 2 ; adversidens, fig. 3 ; bidens, fig. 4 ; glandi- 
cula, fig. 5; genesii, figs. 6-8. Vertigo (Alaea) parcedentata 
(Al. Br.) BOETTGER, Jahrb. Nassau. Ver. Nat. xxxxii, 1889, p. 
308. Vertigo parcedentata Al. Br., KENNARD and WOODWARD, 
Proc. Geol. Asso. xxviii, 1917, p. 170. (Pleistocene and 
Holocene, Northampton, Middlesex, Fifeshire) ; Proc. Malac. 
Soc. London vii, 1906, p. 119 (identity of V. levenensis with 
parcedentata var. genesii) . Pupa adversidens and Pupa glan- 
dicula SANDBERGER, Verh. Phys.-Med. Ges. Wiirzburg, n. F., 
xix, 1886, p. 319, names only. Pupa (Vertigo) parcedentata 
Al. Br., WiisT., Zeitschr. deutschen Geol. Gesell, vol. 59, 1907, 
p. 121 ; Zeitschr. f . Naturwiss, vol. 75, 1902, pp. 316, 320, pi. 
6, f. 26-32. 



208 EUROPEAN SPECIES OF VERTIGO. 

The first description of this snail, so far as I can learn, was 
that of Sandberger in 1875; from this it appears that the 
form with a weak parietal tooth as well as one opposite it in 
the palate [lower palatal] , is to be regarded as typical. It is 
that which he figures. Boettger states that according to his 
understanding typical parcedentata forms have 1 or 2 blunt 
denticles, and occur in his environs in the younger Middle- 
Pleistocene loess of the ' ' Erbenheimer Talchens bei Wiesbaden 
und von Schierstein und im alten alluvium des Grossen Bruchs 
bei Traisa in der Prov. Starkenburg. " Outside of this region 
it is known from the loesses of Heidingsfeld, Wurzburg and 
Regensburg, and the toothless form has also been recognized 
in England and Scotland, perhaps also in France. 

According to Boettger, the earliest Pleistocene forms known 
appear to have been toothless; afterward variable, partly 
toothed forms in the later Pleistocene and Alluvium, finally 
extinction of the toothed parcedentata stock, which Boettger 
considered to belong to lower elevations, the toothless genesii 
alone surviving, and only in Alpine and high northern regions. 

In his paper of 1887 Sandberger described the following 
varieties, the first three probably referable to V. alpestris. 

Var. quadridens (pi. 18, fig. 14). Aperture armed with one 
mammilliform parietal tooth, one columellar and two on the 
throat [palate], of which the anterior is smaller. 

Var. tridens (pi. 18, fig. 13). Anterior tooth of the throat 
obsolete or wanting, the rest present. 

Var. adversidens (pi. 18, fig. 16). No columellar tooth, the 
rest are apparent. 

Var. bidens (pi. 18, fig. 15). Columellar and anterior 
tooth of the throat wanting. This form is the typical 
parcedentata. 

Var. glandicula (pi. 18, fig. 19). Only the parietal tooth 
visible. 

Var. genesii (pi. 18, figs. 10-12). Toothless; bright brown 
in life, subpellucid, the margins of aperture violacous-brown 
[ genesii Gredler] . 

Dr. Boettger considers the relatively large, 4-toothed loess 
form which Sandberger described as var. quadridens to be 



EUROPEAN SPECIES OF VERTIGO. 209 

y. alpestris (Alder). He states farther that he knows of no 
form of parcedentata-genesii having more than two teeth, none 
having a columellar or an upper palatal. Sandberger's varie- 
ties quadridens, tridens and adversidens would therefore be 
forms of alpestris. Boettger further remarks that recent 
forms of V. alpestris (Aid.) with only one palatal fold, or 
without them, are not rare, and have been described by 
Westerlund as var. tridentata and bidentata. Cf. notes under 
alpestris var. mitis West. 

See also Geyer's notes under V. genesii, from which it ap- 
pears that the living form may be either toothed or toothless. 
At present the distinctions, if any, between genesii and parce- 
dentata remain to be formulated. The present usage appears 
to be this: the recent specimens are called genesii and the 
fossil (Pleistocene or Holocene) are called parcedentata. 

Group of V. otostoma. 

Sinistral species with few, small teeth, arranged 1-1-1 or 
1-1-0. 

79. VERTIGO OTOSTOMA Westerlund. 

Shell [sinistral], cylindric, finely striate, yellowish-brown, 
glossy. Whorls 6y 2 to 7, slowly increasing, convex, the two 
penult nearly equal, the last much higher, having two pale 
longitudinal furrows anteriorly. Aperture small, ear-shaped, 
with only 2 low white folds and 1 very small tooth in the 
lower part of the palate. Peristome very oblique, margins 
connected, the outer margin long, angularly drawn forward 
above the middle, not impressed, strongly arcuate above, pass- 
ing into the weakly sigmoid columella in a short arc. Length 
22/3, diam. 1% mm. (West.). 

Sweden : Tenhult, prov. Smaland. 

Pupa otostoma, WEST., Nova Acta R. Soc. Sci. Upsal. (3) 
viii, 1871, p. 100; Fauna iii, 1887, p. 142. PFR., Monogr. 
viii, 410. 

The special locality of this "ganz eigenthumlich und hochst 
merkwiirdig" shell is Augustenborg, a high hill opposite the 
road to the estate of Tenhult, iy 2 Swedish miles from the town 



210 EUROPEAN SPECIES OF VERTIGO. 

of Jonkoping, southward, where it occurs with V. pineticola, 
avoided, arctica and others of the genus, upon twigs and fallen 
leaves. 

80. VERTIGO CLEVEI Westerlund. 

Shell [sinistral] obesely ovate; whorls 5%, convex, the 
penult inflated below, the last scarcely higher, much narrower,, 
impressed at the aperture. Aperture truncate-ovate, the outer 
margin arcuate above, straight in the middle, oblique; basal 
margin rounded. Teeth only 2, one each on the parietal 
wall and the columella. Length 1%, diam. 1 mm. (West.). 

Sweden: Sala, in a place called Grona gangen (Prof. P. T. 
Cleve). 

Pupa clevei WEST., Fauna Pal. Reg. Binnen-conch. iii, 1887,, 
p. 142. 

Subgenus VERTILLA Moquin-Tandon. 

Vertilla M.-T., Hist. Nat. Moll. terr. et fluv. France ii, 
1855, p. 408, for V. plicata and V. pusilla. 

Angular and parietal lamellae well developed, the angular 
not connected with the peristome, parietal long; columellar 
lamella subvertical, spirally entering above. Upper palatal 
fold very long, its inner end curving down ; the lower palatal 
short or wanting. Type V. angustior Jeffr. (V. plicata Moq.). 

This group differs from Augustula by having the columellar 
lamella of entirely different form, and the upper palatal fold 
penetrating deeply, its inner end decurved and somewhat 
hook-like (as in pi. 5, fig. 17). In Angustula and most species 
of Vertigo the upper palatal fold is shorter than the lower. 
While Angustula and Vertilla show a certain parallelism or 
convergence, there appears to be no direct or close relationship. 
Vertigo angulifera Boettger, a dextral species of the German 
Miocene, is said to be closely related to V. angustior, and prob- 
ably is ancestral to the recent species. 

Vertilla was proposed for sinistral species of Vertigo, in- 
cluding the type of the genus ; and the name has been exten- 
sively used by European authors in this sense. The sinistral 
forms have obviously been derived from several dextral stocks 



EUROPEAN SPECIES OP VERTIGO. 211 

independently, so that Vertilla, as used by Boettger, Wester- 
lund and others is heterogeneous. The name may properly 
be restricted to V. angustior and its immediate relatives. 

81. VERTIGO ANGUSTIOR Jeffreys. PI. 5, figs. 13, 16-18. 

Shell subfusiform or barrel-shaped, narrower in proportion 
than V. pusilla, rather solid, but semitransparent, glossy, light 
horn-colour, strongly, obliquely and rather closely striate in the 
line of growth ; periphery compressed and somewhat angular : 
epidermis thin: whorls 4%, rather convex, but compressed, 
gradually increasing in size, the penultimate one a trifle 
broader than the last, which occupies about two-fifths of the 
shell, the first or upper whorl smooth and shining: spire 
rather short, abrupt, and blunt at the point: suture rather 
deep, mouth subtriangular, and very narrow in consequence 
of the great contraction or sinuosity of the outer edge in the 
middle as well as towards the base ; teeth four or five, viz. two 
on the pillar [parietal wall] , the outer one of which is a little 
in advance of the other; one on the pillar lip [columella], 
which is sunk deep within the mouth and resembles a strong 
curved plate more than a tooth; and one thick and promi- 
nent tooth inside the outer lip, with rarely a small tubercle by 
the side of it. Outer lip exceedingly thick and scarcely in- 
flected, strengthened outside and inside by a strong rib which 
is situate near the rim and is yellowish- white ; the inside rib 
remarkably thick and increasing the contraction of the mouth ; 
inner lip consisting of a slight deposit on the columella. 
Umbilicus very small, narrow and indistinct, being much con- 
tracted by a sharp and gibbous keel or crest at the base of the 
shell. L. 0.06. B. 0.035 inch (Jeffreys). 

Vertigo angustior JEFFREYS, Trans. Linn. Soc. London xvi, 
pt. 2, 1830, p. 361 (rejectamenta of a small stream at Marino, 
near Swansea) ; British Conchology i, p. 265. STEENBERG, 
Danmarks Fauna, Landsnegle, 1911, p. 166, f. 137. Pupa 
angustior Jeffr., PFEIFFER, Monogr. iii, 560 ; iv, 686 ; vi, 336. 
WESTERLUND, Fauna Europaea Moll. Extramar. Prodr. ii, 
1878, p. 200, with var. product a and gothorum, p. 201 ; Fauna 
iii, p. 142, with var. tiana Mich., producta W., gothorum W. 



212 EUROPEAN SPECIES OF VERTIGO. 

Acta Soc. pro founa et flora Fennica, xiii, no. 7, p. 71 (dis- 
tribution, etc.). 

Turbo vertigo MONTAGU, Testacea Brit. 1803, p. 363, pi. 12, 
f. 6. Not Helix vertigo Gmelin. Vertigo vertigo ISSEL, Moll. 
Pisa, 1866, p. 23. 

Pupa ocsensis HALEVATS, Res. Wiss. Erforsch, Balatonsees, 
iv, pt. 2, p. 60, pi. 3, f. 10. Upper Pontic beds, Oecs, Lake 
Balaton (considered by 0. Boettger a var. of V. angustula). 

Vertigo venetzii Charpentier MS., FER., Prodr. p. 65 (nude 
name). CHARPENTIER, Cat. Moll. Suisse, p. 18, pi. 2, f. 11. 
EOSSMAESSLER, leonogr. pt. 10, p. 30, f. 650. STERKI, Proc. 
U. S. Nat. Mus. 1888, p. 380, pi. 42, f. 11, 12. Pupa venetzii 
Charp., PFR. Monogr. ii, 364. 

Vertigo hamata HELD, Isis, 1837, p. 304. 

Vertigo plicata A. MULLER, Wiegmann 's Archiv f iir naturg. 
1838, i, p. 210, pi. 4, f. 6 (Neuhaldensleben). 

Vertigo nana MICHAUD, Complement de 1'hist, nat. des moll., 
etc., 1831, p. 71, pi. 15, f. 24, 25 (Lyons). 

V. angustior is readily known by the strongly striate shell 
tapering towards both ends, the vertical columellar lamella and 
the very long and strong upper palatal fold, decurved at its 
inner end ; the lower palatal being either quite weak or want- 
ing. The palatal callus is very strong. 

Var. nana Mich. 2 parietal teeth ; columellar margin less 
thickened, with scarcely noticeable teeth. Upper palatal tooth 
short, the lower rudimentary. Southern France at Lyons; 
northern Italy (Westerlund). 

Michaud's description and figure are not complete for de- 
tails of structure, and as far as I can see pertain to practically 
typical V. angustior. Numerous specimens from Lyons ap- 
pear to be practically typical. Westerlund, however, defines 
nana as a variety. Dupuy says of specimens received from 
Michaud that the upper palatal fold is less strong, and the 
lower less distinct than in what he had figured as Pupa 
venetzii (Hist. Nat. Moll. France, p. 121). 

Var. producta Westerlund. Elongate ovate, chestnut- 
colored, 5% to 6 convex whorls, the last with straight back, 
very oblique, with a strong white tubercle at the base, pro- 



EUROPEAN SPECIES OF VERTIGO. 213 

duced forward, almost equally bipartite by the deep longi- 
tudinal furrow. Length 2, diam. 114 mm. (Westerlund). 
Sweden : Borgholm. 

Var. gothorum Westerlund. Shortly oval, rather ventri- 
cose, the 4^ to 5 whorls but little convex, the last longitudin- 
ally impressed in front, a little tubercular below. (Margins 
of the peristome united by a callus. Length 1%, diam. 1 
mm. (Westerlund). Sweden: "Bellevue," Carlshamn. 

82. VERTIGO CALLISTA Westerlund. 

Shell cylindric, chestnut-brown or rufous brown, densely 
striatulate ; whorls 6, convex, the antepenult and preceding en- 
tirely cylindric, separated by a deep suture, the penult whorl 
largest, the back of the last whorl strongly and straightly slop- 
ing, divided into two parts by a deep transverse furrow, the 
upper, larger part narrow, cylindric, the other very short basal 
part with a strong white protuberance near the aperture, from 
which it is separated by a wide furrow. Suture very oblique, 
not ascending to the aperture. Aperture placed to the left, 
suboval, with about 2 [parietal] teeth or folds: one shorter, 
more exterior, marginal, the other deeply entering, nearly 
continuous with the former ; palatal fold long, strongly curved 
within ; columella calloused, and the lip with a rufous callus 
deep within the outer margin. Peristome unexpanded, the 
outer margin strongly arcuate above, and angularly produced 
above the middle. Length 1%, diam. % mm. (WesterL). 

Sweden: Dalbyo, prov. Sodermanland. 

Pupa callista WESTERLUND, Acta Soc. pro fauna et flora 
Fennica, xiii, no. 7, 1897, p. 72. 

Vndescribed or uncertain species. 

Vertigo lucida Jan. T. dextrosa, conico-cylindrica, obtusa, 
diaphana, lucidissima ; apertura ovalis; labium columellae 
coarctatum; perist. edentulum. (Jan Mantissa p. 3), north- 
ern Italy. 

Vertigo dinii Stefani, = Truncatellina. 

Pupa (Vertigo) danica WesterL, Cat. der Conchyl.-Samm- 
lung Fr. Paetel, 1873, p. 107. Dania, nude name. 



214 TERTIARY VERTIGININAE. 

Vertigo guadalupensis Fer., Tabl. Syst. p. 64, no. 6 "(4 
dentata) . Habit. La Guadeloupe. ' ' 

Vertigo cylindrica J. Colbeau, Ann. Soc. Malac. Belgique 
iii, 1868, p. 97, pi. 2, fig. 7. Shell cylindro-conic, yellowish 
brown, glossy, the growth-striae quite noticeable, 5-6 whorls, 
the first very small, the later ones nearly equal ; aperture small, 
oval, longer than wide, without teeth. Length 1%, diam. 
about 1 mm. 

Colbeau remarks that the unique example of this shell he 
formerly regarded as a variety or anomaly of V. pygm&a, not 
adult. The aperture is relatively smaller than in V. 
muscorum [Drap., non L.] and edentula-, it is larger and more 
glossy than muscorum, smaller and darker colored than eden- 
tula. From Belgium. The figure is copied, pi. 17, fig. 8. 
May be a Truncatellina or an abnormal V. pygmaea. 

Vertigo sultrovhiformis de Gregorio (pi. 17, fig. 7). Shell 
very minute, pupoid, subtrochiform, the last whorl a little 
angular, aperture small, subrotund. Length 2 mm. Perhaps 
this is a variety of the preceding ["Pupa muscorum," a 
Truncatellina] having the last whorl broken and the spire 
a little wider. I have only one specimen (Vertigo subtrochi- 
formis de Greg., Annales de Geol. et de Paleont., 32 livr., Feb., 
1907, p. 8, pi. 1, f. 17). 

Italy : Abano, found in mud from a hot spring. The figure 
is obviously inexact, and looks like anything but a Vertigo. 
Generic position dubious; though not so absolutely hopeless 
as some other generic references in the same paper, such as 
the * ' Glandina ' ' and ' ' Melanopsis. ' ' 

IV. FOSSIL SPECIES OF VERTIGO AND OTHER VERTIGININAE. 

Vertigininae are common as Pleistocene fossils, which in this 
work are included with the recent species. 

A few Eocene species described as Pupa are probably Ver- 
tigininae, but the only one of which the apertural character 
is known has no teeth. As this is undoubtedly a secondary 
condition, such a form could not be ancestral to the modern 
genera, and it was probably an end-product of evolution from 
some toothed genus of the Palaeocene or Mesozoic. 



TERTIARY VERTIGININAE. 215 

Undoubted species of Vertigo, having teeth like V. moulin- 
siana or alpestris, and of quite modern aspect, appeared in 
the Lower Oligocene (amber of the Baltic coast) ; and in the 
Middle and Upper Oligocene and Miocene of central Europe 
the species became somewhat numerous. Most of these forms 
differ only in specific characters from living species, and the 
ancestors of the latter are probably among them. In Italy 
and southern France, Pliocene species occur, some closest to 
Miocene, others to recent forms. There has thus been a suc- 
cession of similar species in Europe from Oligocene to recent 
times. 

The subgenus VertUla appeared in one or two species in 
the German Lower Miocene. 

Ptychalaea, in the recent fauna known only from the Bonin 
Islands, is known by several European species from the Lower 
Miocene to Pliocene. 

Glandicula occurs in the Upper Oligocene and Lower 
Miocene, Germany and France, then becoming extinct, so far 
as known. Pseudelix and Enneopupa are other genera of the 
German Upper Oligocene which left no descendants. 

In America Vertigo is possibly represented by two Middle 
Eocene species, which from their shape and size appear refer- 
able to this genus rather than to Pupilla ; yet as the apertural 
characters are unknown, and they are from a far lower horizon 
than any indubitable Vertigo, little stress is to be placed upon 
the generic reference. The general appearance is that of the 
two French Eocene species following. Perhaps we have to 
do with an unrecognized Eocene genus, and for this reason the 
species are here listed as Pupae, as originally described. 

List of Tertiary species of Vertigo. 

PUPA ARENULA White. 12th Ann. Rep. U. S. Geol. and 
Geogr. Surv. Terr, for 1878, pt. 1, p. 46, pi. 19, f. 8a, &. Upper 
Green River Group [= Bridger] Valley of Henry's Fork 
southward from Green River City, Wyoming. 

PUPA ATAVUNCULA White. Loc. cit., pi. 19, f. 9a. Same 
locality and horizon. 



216 TERTIARY VERTIGININAE. 

PUPA BONNETI Cossm. Ann. Soc. Roy. Zool. et Mai. Belg. 
xli, 1906, p. 283. Eocene: Sparnacien inferieur, Grauves. 

PUPA HEBERTI Fougeroux et Denainvilliers. Journ. de 
Conchyl. 1875, p. 75, pi. 3, f. 7. Upper Eocene : Calcaire de 
Provins, Saint-Parres near Nogent-sur-Seine. 

VERTIGO HAUCHECORNEI Klebs. Jahrb. Preuss. geol. Landes- 
anst. u. Bergakademie zu Berlin, 1885, (1886), p. 382, pi. 
17, f. 5. Koken, Die Vorwelt, p. 463, f. 112. Lower Oligocene, 
in amber, Prussian coast. 

VERTIGO KUNOWII Klebs. Jahrb. Preuss. geol. Landesanst, 
1885 (1886), p. 386, pi. 17, f. 6. Lower Oligocene, in amber 
of the Prussian coast. Resembles genesii. 

VERTIGO MINOR Boettger. Vertigo callosa var. minor 
BTTG., Jahrb. d. geol. Reichsanst. xx, 1870, p. 296, pi. 13, f. 7. 
V. (Alaea) minor BTTG., Jahrb. Nassau. 1889, p. 305; 
Tuchoric, Bohemia, Upper Oligocene. 

VERTIGO DEFRANCII (Brong.). Pupa defrancii BRONGNIART, 
Mem. du Museum, xv, 1810, p. 378, pi. 22, f. 19. DESHAYES, 
An. s. Vert, ii, 1864, p. 857, pi. 56, f. 37-39. 

Aquitanian : Meulieres superieures, Milon, Palaiseau, Mont- 
morency. 

VERTIGO MOENANA Zinndorf. 37-42 Bericht ueber die 
Thatigkeit des Offenbacher Ver. Naturk. 1901, p. 136, pi. 5, 
f . 9. Upper Middle Oligocene, Offenbach a. M. 

VERTIGO ELSHEIMENSIS Bttg. Jahrb. Nassau. Ver. Nat. 42, 
1889, p. 295, pi. 7, f. 2. Middle Oligocene, Elsheim in Rhein- 
hessen. 

VERTIGO CALLOSA (Reuss). Pupa callosa REUSS, Paleon- 
tographica ii, 1849, p. 30, pi. 3, f. 7. SANDBERGER, Vorwelt, 
p. 400, pi. 24, f. 19. Vertigo (Alaea) callosa BOETTGER, Jahrb. 
Nassau. Ver. Nat. 42, 1889, p. 296. Upper Oligocene, 
Tuchoric, Bohemia. 

V. c. mut. maxima Bttg., t. c., 1889, p. 296, pi. 7, f. 3. 
Upper Oligocene, Hochheim. 

V. c. mut. convergent Bttg., t. c., p. 297, pi. 7, f. 4. Old 
Lower Miocene, Affenstein, near Frankfort a. M. 

V. c. mut. alleodus (Sandberger). Pupa alleodus SDBGR., 
Mainzer Becken p. 58, pi. 35, f. 10. V. c. mut. alleodus BTTG., 
t. c., p. 298. Lower Miocene, Appenheim, Rheinhessen, etc. 



TERTIARY VERTIGININAE. 217 

V. c. mut. quinquedentata Joos, Jahrb. Nassau. Ver. Nat. 
64, 1911, p. 66. Miocene, Budenheim. 

Vertigo callosa cyrenarum Zinnd. Vertigo (Alaea) callosa 
(Rss.) var. cyrenarum ZINNDORF. 37-42 Bericht Offenbacher 
Ver. Naturk. 1901, p. 136. Cyrena marl, Upper Middle Oligo- 
cene, Offenbach, a. M. 

Vertigo callosa divergens Flach. Verb. Phys.-Med. Ges. zu 
Wiirzburg, n. F., xxiv, 1890, p. 8, pi. 3, f. 4. Joos, Nachrbl. 
d. m. Ges. 1912, 39. Pupa (Vertigo) cardiostoma var. diverg- 
ens CLESSIN, Ber. naturw. Ver. Regensburg 1893, iv, p. 9, 
no. 38, pi. 1, f. 6. 

VERTIGO CARDIOSTOMA (Sandberger). Pupa (Vertigo) 
cardiostoma SDBGR., Vorwelt, 1875, p. 600, no description. 
CLESSIN, Regensburger Correspondenzblatt 1877, p. 37; 
Malak. Blatter viii, p. 86. Vertigo (Ptychalaea) cardiostoma 
BOETTGER, Jahrb. Nassau. Ver. Nat., 42, 1889, p. 294. FLACH 
Verh. Phys-med. Ges. Wiirzburg, 1890, p. 8, pi. 3, f. 5. 
Vertigo (Alaea) cardiostoma Joos, Nachrbl. d. m. Ges. 1912, 
p. 39. Upper Miocene, Undorf, near Regensburg, Bavaria; 
Steinheim. 

Placed by Boettger in Ptychalaea, by Joos in Alaea. 

VERTIGO PROTRACTA (Sandberger). Pupa protracta SDBGR., 
Vorwelt, 1875, p. 400. Vertigo (Alaea) protracta BOETTGER, 
Jahrb. Nassau. Ver. Nat. 1889, p. 300, pi. 7, f. 5. Upper 
Oligocene, Hochheim. 

VERTIGO OVATULA (Sandberger). Pupa ovatula SDBGR., 
Vorwelt, 1875, p. 400. V. (Alaea) ovatula BOETTGER, Jahrb. 
Nassau., 1889, p. 301, pi. 7, f. 6. Upper Oligocene, Hochheim. 

V. o. mut. miLiiformis Boettger. Bericht Senck. Ges. 1884, 
p. 270, pi. 4, f . 9 ; Jahrb. Nassau, 1889, p. 302. Older Lower 
Miocene, near Niederrad. 

V. o. mut. hydrobiarum Boettger, Jahrb. Nassau, 1889, p. 
302, pi. 7, f. 7. Newer Lower Miocene, Mosbach-Biebrich. 
Var. mosbachiensis Bttg., t. c., p. 303, pi. 7, f. 8; with the 
preceding. 

Dr. Boettger considers this species to be closely related to 
V. milium; but there is nothing in the description or figures 
to show that it has a columellar lamella of the Angustula type, 



218 TERTIARY VERTIGININAE. 

so that the relationship claimed is quite uncertain, and appears 
to me improbable. 

VERTIGO KOCHI Boettger. Jahrb. Nassau. Ver. Nat. 42, 1889, 
p. 304, pi. 7, f. 9. Upper Oligocene, Hochheim. 

VERTIGO DIVERSIDENS (Sandberger). Pupa diversidens 
SDBGR., Vorwelt, 1875, p. 549, pi. 29, f. 23. Vertigo diversi- 
dens BOURGUIGNAT, Malac. Sansan p. 84, f. 124-127; Vertigo 
presbytera Bgt. mentioned as a synonym. Middle Miocene: 
Sansan. 

VERTIGO TUCHORICENSIS (new name). Pupa microstoma 
REUSS, Sitzungsber. Math.-Nat. Cl. k. Akad. Wiss. Wien, xlii, 
1861, p. 73, pi. 2, f. 8. SANDBERGER, Vorwelt, p. 438, pi. 24, 
f. 17. Vertigo microstoma Reuss, KLIKA, Tert. Land- und 
Susswasser-Conch. des nordwesten Bohmen, p. 98, f. 93, in 
Archiv naturwiss. Landesdurchforschung von Bohmen, vii, no. 
4, 1891. Lower Miocene : Lipen, Bohemia. This is not Pupa 
microstoma Mlldff., 1881, or Pupa microstoma- Pfr., 1852. 

VERTIGO ( ?) GLOBULUS Desh. Pupa globulus Dh., An. s. 
Vert, ii, 1864, p. 860, pi. 56, f . 16-18. Lower Miocene : Cal- 
caire de Beauce. Perhaps a Gastrocopta. It has been re- 
ferred by Oppenheim to his genus Paracraticula. 

VERTIGO (?) ANODON (Desh.). Pupa anodon DESHAYES, 
An. s. Vert. Bassin Paris, ii, 1864, p. 849, pi. 56, f. 22-24. 
Lower Miocene : Calcaire de Beauce. 

VERTIGO (?) PARVULA (Desh.). Pupa parvula Dh., An. s. 
Vert, ii, 1864, p. 852, pi. 56, f . 31-33. Lower Miocene : Cal- 
caire de Beauce. 

VERTIGO COARCTATA (Desh.). Pupa coarctata Dh., An. s. 
Vert, ii, 1864, p. 856, pi. 56, f . 13-15. Lower Miocene : Cal- 
caire de Beauce. 

VERTIGO DIEZI (Clessin). Pupa (Alaea) diezi CLESSIN, 
Berichte Naturwiss. Ver. Regensburg, xiii, 1912, p. 107. 
Middle Miocene, Undorf, near Regensburg, Bavaria. 

VERTIGO UNDORFENSIS (Clessin). Pupa (Alaea) undorf- 
ensis CL., t. c., p. 108. Middle Miocene, Undorf. 

VERTIGO GLOBUS, new name. Pupa (Alaea) globula CL, 
t. c., p. 108 (Not Pupa globulus Deshayes). Middle Miocene, 
Undorf. 






TERTIARY VERTIGINTNAE. 219 

VERTIGO MUSCICOLA Clessin. Pupa (Alaea) muscicola 
CLESSIN, t. c., p. 108. Middle Miocene, Undorf. 

VERTIGO INFLATULA (new name). Pupa (Alaea) inflata 
CLESSIN, t. c., p. 109 (not Pupa inflata Wagner, 1827). 
Middle Miocene, Undorf. 

VERTIGO MINIMA Clessin. Pupa (Alaea) minima CLESSIN, 
t. c., p. 109. Middle Miocene, Undorf. The name minima 
has been used in Pupa in a varietal sense. 

VERTIGO CYCLOPHORA Bgt. Malacol. de Colline de Sansan 
1881, p. 83, pi. 5, f . 120-123. Middle Miocene : Sansan. The 
following 12 species are from the same formation. 

VERTIGO CAMPANEA Bgt., t. c., p. 85, pi. 5, f . 140-143. 

VERTIGO SANSANICA Bgt., t. c., p. 87, pi. 5, f. 128-131. 

VERTIGO LCEMODONTA Bgt., t. c., p. 88, pi. 5, f. 132-135. 

VERTIGO CALLOSTOMA Bgt., t. c., p. 89, pi. 5, f . 136, 137. 

VERTIGO CODIOLENA Bgt., t. c., p. 91, pi. 5, f. 144-147. 

VERTIGO MILNE-EDWARDSI Bgt., t. c., p. 92, pi. 5, f . 148-151. 

VERTIGO BOTHRIOCHEILA Bgt., t. c., p. 93, pi. 5, f. 152-155. 

VERTIGO RAGIA Bgt., t. c., p. 95, pi. 5, f. 156-159. 

VERTIGO TRIODONTA Bgt., t. c., p. 96, pi. 5, f. 160-163. 

VERTIGO RHYNCHOSTOMA Bgt., t. c., p. 97, pi. 5, f. 164-167. 

VERTIGO ONIXIODON Bgt., t. c., p. 98, pi. 5, f. 168-171. 

VERTIGO MICRONIXIA Bgt., t. c., p. 100, pi. 5, f. 172-175. 

VERTIGO PERAPERTA (new name). Pupa (Alaea) aperta 
Sandb. Ms., K. MILLER, Jahresh. Ver. Vaterl. Naturk. Wiirt- 
temberg, vol. 56, 1900, p. 397, pi. 7, f. 13. Joos, Nachrbl. 
1912, p. 40, pi. 2, f. 7. (Not Pupa aperta Marts., Mai. Bl. x, 
1863, p. 180). Upper Miocene, Steinheim. 

It is said by Joos to be closely related to V. angulifera Bttg., 
which appears to be a Vertilla. The palatal fold is described 
by Joos as about 5 mm. long, an obvious mistake, in a shell 
1.4 mm. long ; probably 0.5 mm. was intended. 

VERTIGO PSEUDANTIVERTIGO Paladilhe. Revue des Sciences 
Naturelles, ii, 1873, p. 207, pi. 2, f. 25-27. Pliocene, near 
Montpellier. 

VERTIGO ( ?) BLEICHERI Paladilhe. Eev. Sci. Nat. ii, 1873, 
p. 51, pi. 2, f. 19-21. Pliocene, Montpellier. 

VERTIGO MYRMIDO Michaud. Ann. Soc. Linn, de Lyon, Rev. 



220 TERTIARY VERTIGININAE. 

Ser., ii, 1855, p. 47, pi. 5, f. 14, 15. Pupa (V.) myrmido 
SANDBERGER, Vorwelt, p. 723, pi. 27, f . 22. Pliocene, Hauterive. 

VERTIGO CROSSEI Michaud. Journ. de Conchyl. x, 1862, p. 77, 
pi. 4, fig. 3. Pliocene, Hauterive (Drome). A toothless form 
of which the position is dubious; perhaps a Columella. See 
Locard, Ann. Mus. d'Hist. Nat. Lyon, ii, 1878, p. 230. 

VERTIGO SCHLOSSERI (Cossmann). Pupa (Vertigo) oviformis 
SCHLOSSER, Jahrb. k.-k. Geol. Eeichsanstalt 1907, Ivii, p. 757, 
pi. 17, f . 5. Not Pupa oviformis Michaud. P. [upa] schlosseri 
COSSMANN, Revue Grit. Paleozool. 1908, p. 257. Pliocene, 
Eichkogel near Modling. 

VERTIGO GLOBOSA Sacco. Fauna Malac. All. il. Piemonte, 
1885, p. 31, pi. 2, f. 3. Alaea globosa SACCO, I Moll. Terr. 
Terz. Piemonte xxii, p. 71, pi. 6, f. 8. Pliocene, Tassarolo. 

V. g. tassaroliana Sacco. Riv. Fauna malac. foss. terr. lac., 
1887, p. 174. Alaea g. var. tassaroliana SACCO, I moll. terr. 
Terz. Piem. xxii, p. 71, pi. 6, f. 9. Tassarolo, Fossano. 

Subgenus PTYCHALAEA Boettger. 

VERTIGO (PTYCHALAEA) FLEXIDENS (Ess.). Pupa flexidens 
REUSS, Sitzungs-Ber. K. Akad. Wiss. Wien, Math.-Nat. Cl., 
xxxii, p. 74, pi. 2, f. 9. SANDBERGER, Vorwelt p. 439, pi. 24, 
f. 18. V. (P.) flexidens BOETTGER, Jahrb. Nassau. Ver. Nat. 
xxxxii, 1889, p. 293, pi. 7, f. 1. Lower Miocene, Appenheim in 
Rheinhessen; Tuchoric and Lipen, Bohemia; Saucats, Bor- 
deaux (var. girondica Boettger, t. c., p. 294). WENZ, Nachrbl. 
d. m. Ges. 44, 1912, p. 190, f. 2. KLIKA, Arch. Naturwiss. 
Landesdurchforschung von Bohmen, vii, no. 4, 1891, p. 95, 
f. 90. 

VERTIGO (PTYCHALAEA?) TRIGONOSTOMA (Al. Braun). Pupa 
trigonostoma Al. Br., in Walchner's Geognosie, p. 1185. 
SANDBERGER, Mainzer Becken p. 59, pi. 5, f. 17. V. (Ptycho- 
chilus) trigonostoma BTTG., Jahrb. Nassau. Ver. Nat. 1889, 
292. Upper Oligocene, Hochheim. 

VERTIGO (PTYCHALAEA) BLUMI Boettger. V. (Ptychochilus) 
Uumi BTTG., Bericht Senck. Ges. 1884, p. 268, pi. 4, f. 8. 
Lower Miocene, Schleusenkammer bei Niederrad. 

VERTIGO (PTYCHALAEA) PRISCILLA Paladilhe. Vertigo pris- 



TERTIARY VERTIGININAE. 221 

cilia PALAD., Revue Sci. Nat. (Montpellier), ii, 1873, p. 53, pi. 
2, f . 22-24. Pliocene, Montpellier. 

VERTIGO (PTYCHALAEA) CAPELLINII Sacco. Vertigo (Scara- 
bella) capellinn SACCO, 1885. Scarabella capellinii SACCO, I 
Moll. Terr. Terz. Piemonte, xxii, p. 71, pi. 6, f. 7. 

Piedmont Upper Pliocene, Villafranchian stage, Fossano and 
Tassarolo. 

V. c. ligustica Sacco. V. (S.) capellinii var. ligustica 
SACCO, 1887. Scarabella c. L, SACCO, 1. c. Tassarolo. 

Submenus VERTILLA Moquin-Tandon. 

VERTIGO ANGULIFERA Boettger. Bericht Senck. Nat. Ges. 
1884, p. 271, pi. 4, f . 10 ; Jahrb. Nassau. Ver. Nat. 42, 1889, 
p. 310. Lower Miocene of the Schleusenkammer near Nieder- 
rad. Placed in Alaea by Boettger, but probably a dextral 
species of Vertilla. 

Genus GLANDICULA Sandberger. 

Glandicula SANDBERGER, Land- und Siisswasser-Conchyl. der 
Vorwelt, p. 401, 1875; monotype Pupa (Glandicula) tiarula 
A. Brn. Paracraticula OPPENHEIM Denkschr. k. Akad. Wiss., 
Wien, vol. 57, 1890, p. 126. 

Acorn-shaped, with deeply impressed suture, the post- 
embryonic whorls having sculpture of distant, oblique ribs; 
aperture triangular, the outer lip inflexed; typically with 
3-1-2 teeth, the angular and parietal lamellae emerging to the 
margin. 

Type G. tiarula (A. Brn.), pi. 5, figs. 14, 15. Upper Oli- 
gocene: Hochheim, Landsehneckenkalk, 

GLANDICULA TIARULA (A. Brn.). Vertigo tiarula A. Braun, 
Walchner's Geogn. 2 edit., p. 1135. Pupa tiarula Sandberger, 
Mainzer Becken p. 60, pi. 5, f . 18 ; Vorwelt, p. 401, pi. 23, f . 
17. Vertigo (Glandicula) tiarula Boettger, Jahrb. Nassau. 
Ver. Nat. Jahrg. 42, 1889, p. 290. 

Boettger has given a good account of this rare snail, which 
has not been figured in sufficient detail. He believes its 
affinity to be with Ptychochilus (Lyropupa), mentioning such 
forms as perlonga Pse. and lyrata Gld., evidently influenced 



222 TERTIARY VERTIGININAE. 






by the sculpture, a character of no great importance. I have 
not seen specimens. 

GLANDICULA TURCICA (Desh.). Pupa turcica Deshayes, 
An. s. Vert. Bassin Paris ii, 1864, p. 861, pi. 57, f. 1-3. Lower 
Miocene: Calcaire de Beauce. 

GLANDICULA (PARACRATICULA) UMBRA (Oppenh.), Pupa 
(Paracraticula) umbra OPPENHEIM, Denkschr. k. Akad. Wiss., 
Wien, vol. 57, 1890, p. 126, pi. 3, f. 10. Eocene, Pugniello, 
Italy. This species is the type of Paracraticula Oppenh. 
Referred to Glandicula with some doubt. 

Genus ENNEOPUPA Boettger. 

Enneopupa BTTG., Jahrb. Nassau. Ver. Nat, xxxii, 1889, 
p. 288. Monotype Pupa cylindrella Al. Br. 

Shell cylindric, of few (5-6) costulate whorls, the neck 
having a groove parallel to the suture ; aperture oblique ; peris- 
tome continuous, subobsolete ; angular lamella distinct but low, 
(1 or) 2 deeply placed, not marginal parietal lamellae separ- j 
ated from the angular; 2 columellar lamellae; (3 or) 4 long 
palatal folds. 

ENNEOPUPA CYLINDRELLA (Al. Braun). PL 5, fig. 12. 
Pupa cylindrella Al. Br., Walchner's Geogn. p. 1136. Sand- 
berger, Mainzer Becken, p. 50, pi. 5, f . 9 ; Vorwelt p. 393, pi. 
23, f. 7. Vertigo (Enneopupa) cylindrella BTTG., t. c., p. 
288. Upper Oligocene of Hochheim. 

Sandberger's figures are copied in pi. 5, fig. 12. 

Genus PSEUDELIX Boettger. 

Pseudelix BTTG., Jahrb. Nassauischen Vereins f. Natur- 
kunde, Jahrg. 42, 1889, p. 316, type Pupa microhelix Sbgr. 

Shell with a small perforation, globose-conic, heliciform, the 
apex rather obtuse. Whorls 4-5, nearly smooth. Aperture 
oblique, semilunar, with 1 to 3 compressed teeth, not deeply 
placed, the parietal lamella always distinct ; margins expanded, 
somewhat labiate. 

Type: Pupa microhelix Sbgr., from the Upper Oligocene 
landshell chalk of Hochheim (pi. 5, fig. 8). 

This group is compared by Boettger with the American Pupa 



TERTIARY VERTIGININAE. 223 

conoidea Nc., which has some resemblance in figure and teeth. 
The entirely different and peculiar sculpture of conoidea, 
which belongs to the genus Bothriopupa (Vol. XXIV, p. 231), 
together with other differential features, shows that the sup- 
posed affinity is improbable. Prof. Boettger had not seen a 
specimen of conoidea. Pseudelix has a somewhat specialized 
shape, and appears to have left no survivors or no closely 
related genera. 

The nomenclature of the two species is not clear to me, 
Boettger ? s synopsis is followed. 

PSEUDELIX MICROHELIX (Sbgr.). Pupa microhelix SAND- 
BERGER, Mainzer Becken, p. 61, pi. 35, f . 26 ; Vorwelt, p. 402, 
pi. 23, f. 18. 

PSEUDELIX COMES Boettger. Jahrb. Nassau. Ver. Naturk. 
1889, p. 317. Helix piipuU Al. Braun, in Walchner's 
Geognosie, 2d ed., p. 1139. Pupa microhelix in part, SAND- 
BERGER, Mainzer Becken p. 61, pi. 3, f. 8. Upper Oligocene 
landshell chalk of Hochheim. 

Genus Carychiopsis Sandberger. 

Fusulina SANDBERGER, olim. Carychiopsis SANDBERGER, 
Land und Siisswasser-Conch. der Vorwelt, pp. 169, 170 (1872), 
type Pupa dohrni [dhorni] Deshayes. 

A group of minute, slender Pupoid snails, chiefly Lower 
Eocene, contains the species Pupa remiensis, alternans, ~bige- 
minata, dhorni ( dohrni Dh., Sandberger) and coarctata, all 
of Deshayes (Anim. s. Vert. Bassin Paris, ii, 1864, pp. 853- 
856). Some of these have been referred to Vertigo by subse- 
quent authors, but in the opinion of Sandberger, which appears 
well founded, they have no special affinity with that genus, but 
belong near Carychium. 

Pupa interferens Desh. may possibly also belong to this 
group, though somewhat different. It has the general appear- 
ance of Gastrocopta. 

Carychiopsis costulata Sandberger ', t. c., p. 371, of the Lower 
Miocene of Germany and Bohemia, is the latest known member 
of the group. Pupa schwageri Reuss, Sitzungsber. d. k. Akad. 
"Wiss. Wien Ivii, p. 82 is synonymous. 






224 STAURODON. 

Genus STAURODON Lowe. 

Staurodon LOWE, Ann. and Mag. of Nat. Hist. (2), ix, 1852, 
p. 278, for Pupa saxicola and P. seminulum. 

The shell is minute, oblong, the last whorl without crest. 
Peristome continuous by a strong parietal callus. Teeth ar- 
ranged as in Vertigo except that the angular lamella is tuber- 
culiform, marginal upon- the parietal callus close! to the 
termination of the outer lip, with or without a short con- 
tinuation inward. 

Type Staurodon saxicola (Lowe). Distribution, Madeira, 
See pi. 15, figs. 12-14, 

The emergence of the angular lamella and its union with 
the outer lip is probably an ancestral character, found also 
in Ptychalaea and some other Vertiginine groups, as well as 
in other subfamilies, such as Gastrocoptinae. In Staurodon 
this lamella has been much shortened, often running inward 
less than shown in pi. 15, fig. 14. 

The genus has probably existed on Madeira since the early 
Tertiary, as it appears to have no near relatives in the Euro- 
pean series from the Oligocene on. The strong parietal callus 
is doubtless a later development. Many snails of arid regions 
show more or less similar strucures. 

In his publication of 1852, Lowe places only the species 
Pupa saxicola and P. seminulum under Staurodon, but in his 
differential diagnosis of saxicola he mentions P. (Vertigo) 
alpestris and P. pygmaea in order to point out how saxicola 
differs from them, but without stating or implying that they 
belong to Staurodon. It appears from this that either saxi- 
cola or seminulum must be the type of Staurodon. In 1854 
Lowe designated P. pygmaea Drap. the type of Staurodon, 
a course clearly barred by the fact that he had not included 
this species in his group in 1852. 

1. STAURODON SAXICOLA (Lowe). PL 15, figs. 13, 14. 

The shell is shortly rimate, cinnamon-brown, oblong, taper- 
ing slightly upwards, very finely but distinctly striate, the 
striation strongest on the penult and antepenult whorls. The 






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SECOND SERIES: PULMONATA. 



MANUAL 



OF 



CONCtiOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED BY 

GEORGE W. TRYON, JR. 

CONTINUED BY 

HENRY A. PILSBRY, Sc.D., 

SPECIAL CURATOR DEPARTMENT OF MOLLUSCA, ACADEMY OF NATURAL 
SCIENCES OF PHILADELPHIA. 



Part 



PHILADELPHIA : 

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES 

OF PHILADELPHIA. 



STAURODON. 225 

iirst whorl smooth. The whorls are moderately convex, the 
last without grooves or crest behind the lip. The aperture 
is rounded-piriform, containing 5 teeth. The angular lamella 
is a blunt, callous tubercle close to and in adults connected 
with the outer termination of the lip. The parietal lamella 
is long, high, and stands remote from the edge of the parietal 
callus. Columellar lamella short, entering horizontally. 
There are two rather short but high palatal folds, the lower 
one longer. No palatal callus. The peristome is slightly ex- 
panded, thick, of lighter tint than the shell; outer margin a 
little straightened in the middle, connected with the columellar 
margin by a thick, straight, cord-like callus across the parietal 
wall, bearing the angular lamella. 

Length 1.7 to 1.8, diam. 1 mm. ; 4 1 / r 4 1 / whorls. 

Madeira: beneath stones and scoriae in dry, rocky spots of 
low elevation, in the south of Madeira; Praia Bay, west of 
Funehal, under loose pieces of basalt at the top of the cliff 
at the eastern end (Wollaston, Leacock) ; Feijaa dos Asnos 
and Cabo Girao (Baron Paiva). Subfossil in calcareous 
deposits near Canical (Wollaston). 

Pupa saxicola LOWE, Ann. Mag. N. H. ix, 1852, p. 278; 
P. /. S. 1854, p. 214. PFEIPPER, Monogr. Hel. Viv. iii, 1853, 
p. 559. KUSTER, Conchyl. Cabinet, Pupa, p. 182, pi. 21, f. 26, 
27. ALBERS, Malacol. Maderensis p. 62, pi. 16, f. 13, 14. 
PAIVA, Monogr. Moll. Ins. Mader. 1867, p. 139. WOLLASTON, 
Testacea Atlantica, 1878, p. 237. 

Pupa seminulum LOWE, Ann. Mag. N. H. ix, 1852, p. 278; 
Syn. p. 14. PFR.,. Monogr. iii, 559. 

The cord-like parietal callus and the structure of the angu- 
lar tubercle are characteristic. 

la. 8. saxicola seminulum Lowe (PL 15, fig. 12). Differs 
from P. saxicola by the smaller, even very minute size, the 
whorls more swollen, suture deeper, shape more obese, the 
aperture triplicate, the folds unequal, arranged in a triangle, 
the ventral largest, the columellar and palatal smaller and 
snbequal; the subangularly somewhat straightened lip bent, 
inward; the upper denticle somewhat distinct inward (Lowe). 



226 LYROPUPA. 

The figure represents a shell from the Lowe collection. 
It is smaller than saxicola, length 1.55, diam. 0.8 mm. In 
his second description Lowe gives the length as 1*4 to 1%, 
diam. % mm. His description, translated above, was from 
a shell without the upper palatal fold, perhaps not fully 
mature. Wollaston remarks that "the P. semmulum Lowe, 
which was detected by Mr. Leacock at the Cabo Garajao 
or Brazen Head, does not appear to me to differ specifically, 
in any single particular, from the present species [i. e., 
saxicola] . ' ' 

Genus LYROPUPA Pilsbry. 
By H. A. Pilsbry and C. Montague Cooke. 

Lyropwpa PILS., Proc. A. N. S. Phila. 1900, p. 432 (as a sub- 
genus of Nesopupa) , type Pupa lyrata Gould. ANCEY, Proc. 
Malac. Soc. London vi, 1904, p. 124 (as a genus). 

The shell is openly rimate, often perforate, the shape from 
broadly ovate to cylindric ; embryonic 1% whorls microscopi- 
cally granulose, sometimes spirally striate, the following 
whorls abruptly becoming strongly sculptured with axial or 
oblique ribs narrower than their intervals. Aperture trun- 
cate above, rounded below, the peristome continuous, free and 
expanded. Angular lamella emerging to the margin ; parietal 
strong, somewhat immersed. Columellar deeply placed, 
strongly developed, axial or oblique. Two palatals the upper 
very long, emerging to the lip, defining a sinulus. A deeply 
placed basal fold often developed. Axis hollow and rather 
large. Immature stages toothless. 

Type Pupa lyrata Gould. Distribution, Hawaiian Islands. 

There are no inferior tentacles. The head has rounded 
lateral processes and a somewhat prominent median convexity 
above. The sole is almost white in species examined, the back 
and face pigmented. 

Important specific characters are found in the form and ar- 
rangement of the palatal folds, but they are so deeply im- 
mersed that the shell must be broken to see them. Holding it 



LYBOPUPA. 227 

in the fingers under a dissecting microscope, the front of the 
last whorl and the whole columellar side of the aperture can 
be broken away with a needle, leaving the entire inside of the 
outer wall of the last half whorl exposed. In fossil shells it 
breaks off (as figured in pi. 25, figs. 2, 14, 15) . The piece may 
then be stuck upon a slip of card for convenient examination 
under the microscope. Care must be taken to break away 
enough of the basal wall to show the basal fold, which is often 
hard to see. A bull 's eye or other condenser is useful. 

In comparing specimens with the drawings it must be re- 
membered that the apparent shape of the tubercle under the 
inner end of the upper palatal fold changes somewhat with, 
the angle of vision. 

The large, hollow axis is broadly open in the young, as in 
pi. 26, fig. 8, L. spaldingi. 

Lyropupa lives on or near the ground. The sinistral species 
inhabit the humid forest zone where they may be found on 
mossy stones, fallen twigs and dead leaves. The dextral species 
live in dryer open country, under stones and logs. Only dex- 
tral forms are known from Niihau and Kahoolawe. 

Lyropupae are abundant in Pleistocene and later deposits; 
on all the islands. They formerly occupied a far greater 
area than at present. The forest species occurred at lower 
levels than now, as the Achatinellas did. A good part of the- 
low country species are now extinct, and others which are 
abundant as fossils are rare and local as living snails. 

Twenty-two species are now recognized, nine of them de- 
fined by former authors, Gould, Pease, Ball and Ancey. One 
species, Vertigo striatula Pease, has not been recognized with, 
certainty. The type specimens of all the others have been 
examined in the preparation of this work. 

DISTRIBUTION. 16 of the 22 species are confined to single 
islands, so far as known. 6 are common to two or more islands,, 
but in most cases represented by special races on each. A 
narrower view of species than we have taken would increase 
the apparent insular endemicity but would not show inter- 
island relationships so well. 



228 LYROPUPA. 

L. perlonga, Oahu, a subspecies in Niihau and Kauai. 

L. lyrata, Oahu, a variety in Kauai. 

L. micra, Oahu, a subspecies in Molokai. 

L. ovdtula, Oahu, a subspecies in Molokai and Hawaii. 

L. rhabdota, Molokai, a subspecies in Lanai and Maui. 

L. kahoolavensis, Molokai, Lanai, Kahoolawe and Hawaii. 

L. kahoolavetisis shows very little variation throughout its 
range. It inhabits more islands than any other species, and is 
one of the most primitive. It occurs 011 4 islands as a fossil 
only. 

Except in the case of L. ovatula, the islands inhabited by 
one species are always contiguous. Probably some race of 
ovatula (also of kahoolavensis) will be found on Maui, where 
the Pupillidae are much less known than on other islands. 
If so the single exceptional case will be eliminated. 

Niihau and Kahoolawe each have but one species also found 
on the contiguous islands. 

Species of the islands from Molokai east are closely related, 
but Molokai also connects with Oahu, having three fossil sub- 
species of specific stocks which are widely spread on Oahu. 
Otherwise the species of the eastern group of islands all differ 
from those of Oahu and the western islands. 

In general, the distribution of Lyropupa agrees with Lept- 
^chatima. The orderly sequence of species and subspecies 
from island to island, as well as upon the single islands, sug- 
gests migration over continuous land, and gives no ground for 
the hypothesis of carriage by wind, birds or other accidental 
or unusual means, which would surely have produced a less 
harmonic distribution. 

In the following table r signifies recent, / fossil forms; fr 
denotes forms chiefly known as fossils, but also rarely found 
living. 



LYROPUPA. 



229> 





1 

8 


*eS 

a 

A 


1 


Molokai. 


1 


Kahoolawe. 


1 


: | 

1 


L. lyrata 


















uncifera 






fr 












" var 




r 












* 


fossilis 






f 












L. microthauma 


















L rh abdota 


















pluris 


















lanaiensis 










r 


























L. thaanumi 


















L. prisca 


. 














f 


L. clathratula 
















r 


L. truncata 
















r 


L. spaldineri 






r 












T t_ M 

L. mirabilis 






r 












L. antiqua 






f 












L. hawaiiensis 
















f 


L. Hparn.-i 


















sinulifera 


















L. anceyana 
















r 


L. scabra 














f 




L. kahoolavensis 








f 





f 


? 


f 


puukolekolensis 








r 










L. perlonga 






fr 












interrupta 






f 












filicostata 


f 


f 














L. raicra 






fr 












percostata 






f 












maunaloae 








f 










L. ovatula 






f 












kona 








f 








f 


L. plagioptvi .. 






f 












L. cyrta 
















f 


L. cubana 




9 


' 












L. thaumasia 




1 

































Key to species of Lyropupa. 

I. Shell sinistral. 

a 1 . Angular lamella long and high; surface minutely striate 
spirally (in fresh shells). 
& 1 . Species of Oahu. 

c 1 . Small, length about 2 mm. or less. 

L. microthauma, no. 2. 



230 LYROPUPA. 

c 2 . Larger, length over 2.2 mm. 

d 1 . Inner end of lower palatal fold straight. 

L. lyrata, no. 1. 

d 2 . Inner end of lower palatal fold hooked 
downward. L. lyrata uncifera, no. la. 
b 2 . Species of Molokai, Lanai, Maui and Hawaii. 
c 1 . Summit abruptly truncate ; Hawaii. 

L. truncata, no. 7. 
c 2 . Summit strongly convex;, obtuse. 

d 1 . Eibs rather close, 18 to 19 on the last 
whorl ; Maui. L. thaanumi, no. 4. 

d 2 . Ribs rather distant, about 15 on the last 
whorl; Molokai. 

L. rhabdota pluris, no. 3a. 
c 3 . Summit conic with obtuse apex. 

d 1 . Spiral threads distant; about 12 ribs on 
the last whorl. L. clathratula, 110. 6. 

d 2 . Spiral threads close or obsolete. 
e 1 . Molokai, Lanai and Maui. 

L. rhabdota, no. 3. 

e 2 . Hawaii. L. prisca, no. 5. 

a 2 . Angular lamella short and low, scarcely penetrating be- 
yond the outer end of the parietal lamella; later whorls 
with no spiral striation. 

b 1 . Lower palatal fold interrupted, a short outer and a 
high, tubercular inner portion. Oahu. 

L. mirabilis, no. 9. 

b 2 . Lower palatal fold continuous, penetrating more 
deeply than the upper, and usually stronger. 

c 1 . Back with 3 sulci; shell pale, fusiform; Oahu. 

L. spaldmgi, no. 8. 

c 2 . Back with one broad sulcus; color brown (in 
recent shells) . 

d 1 . Basal fold strong; Hawaii. 

e 1 . Whorls very convex, the last with 
12 to 14 ribs; length 2.25 mm. 

L. anceyana, no. 13. 



LYROPUPA. 231 

e~. "Whorls less convex, with more ribs ; 
length 2.4 to 2.9 mm. 

L. hawaiiensis, no. 11. 

d 2 . Basal fold rather weak or wanting ; Oahu, 
Molokai, Lanai and Maid. 

e 1 . 15 to 17 ribs on the last whorl. 

f 1 . Palatal folds rather long; 
Oahu. L. antiqua, no. 10. 

f 2 . Palatal folds short; Maui. 

L. scabra, no. 14. 

e 2 . 19 to 25 ribs on the last whorl; 

length 2 to 2.3 mm. ; Molokai, Lanai. 

L. sparna, no. 12. 

II. Shell dextral (Section Mirapupa). 
Lower palatal fold continuous, rather long, entering as far 
as the upper palatal, its inner end slightly enlarged and 
turned downward (pi. 22, figs. 2, 4) ; shells having the ex- 
ternal appearance of L. perlonga. Molokai, Lanai, Kahoo- 
lawe and Hawaii. L. kahoolavensis, no. 15. 

Lower palatal fold very short or reduced to a low, callous 
pad or tubercle; one, or usually two strong tubercles or 
short folds below the inner end of the upper palatal fold, 
forming a vertical barrier (pi. 25, figs. 14, 15). 

& 1 . Form cylindric, the last three whorls flattened in the 
middle ; three impressions on the back ; ribs narrow, 
generally flexuous. 

c 1 . Back having a hump below the middle of the 
last whorl; shell generally under 2 mm. long 

L. micra, no. 17. 
c 2 . No dorsal or basi-dorsal hump. 

d 1 . Narrow, flexuous ribs, mostly continuous 
and rather widely spaced ; Oahu. 

L. perlonga, no. 15. 
d 2 . Very irregular, closer ribs ; Oahu. 

L. p. interrupt^ no. 16a. 

d*. crowded, slender riblets; Kauai and 

Niihau. L. p. filicostata, no. 16&. 



232 LYROPUPA. 

fe 2 . Form ovate or cylindric-tapering, or cylindric with 
conic summit; whorls convex; ribs strong, arcuate, 
about 15-16 on the last whorl; diam. equal to or ex- 
ceeding half the length. 

c 1 . Last half whorl having deep impressions; 
length 2 to 2.3 mm. 
d 1 . Species of Oahu. 

e 1 . A prominent, straight basal crest 
(pi. 24, fig. 2) ; lower palatal fold 
a diffuse tubercle. 

L. ovatida, no. 18. 

e 2 . No distinctly differentiated basal 
crest; lower palatal a short, high, 
oblique fold. 

L. plagioptyx, no. 19. 

d 2 . Species of Hawaii. Lower palatal and 
basal folds developed as oblong tubercles ; 
back humped at base. 

L. o. kona, no. 18. 

c 2 . Last half whorl having two slight impressions, 
sometimes obsolete; lower palatal a very low, 
callous tubercle ; length 2.75 mm. Hawaii. 

L. cyrta, no. 20. 

Z> 8 . Form broadly oval, compact, the Avhorls short, con- 
vex, the last whorl with 25-30 ribs, and one or two 
impressions on the back ; length 1.9-2.2 mm. 

c 1 . Diameter about 70 per cent of the length; 5 
whorls ; aperture contained 2% times in length 
of the shell. L. cubana, no. 21. 

c 2 . Diameter about 58 to 65 per cent of the length ; 
5% whorls ; aperture scarcely one- third length 
of shell. L. thawmasia, no. 22. 

Group of L. lyrata. 

Sinistral Lyropupae characterized by a strong, long, angu- 
lar lamella, and by minute spiral striaB between and on the 
strong ribs. There are two series belonging to this group. 



LYROPUPA. 233 

Species from Kauai and Oahu have the surface of the em- 
bryonic whorls wrinkled-granulose (except L. microthauma) . 
Those from Molokai, Lanai, Maui and Hawaii have the em- 
bryonic whorls minutely, closely, spirally striate. 

The two Oahuaii species are quite distinct, but L. lyrata 
has numerous local races or forms not yet fully worked out. 
L. trmicata of Hawaii is one of the most characteristic of the 
genus, but the other known species of Molokai, Lanai, Maui 
and Hawaii are closely related and by no means conspicuously 
differentiated. The following key is given as an alternative to 
that on p. 229 ; it is based upon more natural characters, but 
partly not to be observed in worn or fossil specimens. 

a 1 . Embryonic whorls minutely granulose-wrinkled ; Oahu. 

L. lyrata, no. 1. 
a 2 . Embryonic whorls minutely striate spirally. 

5 l . Later whorls very closely striate spirally. 
c 1 . Summit bluntly rounded. 

d 1 . Ribs numerous; Maui. 

L. thaanumi, no. 4. 
d 2 . Bibs rather widely spaced ; Molokai. 

L. rhabdota pluris, no. 3a. 
c 2 . Summit subconic. 

d 1 . Length 1.8 to 2.1 mm.; Oahu. 

L. microthauma, no. 2. 

d 2 . Length 2.5 to 2.7 mm. ; Molokai, Lanai 
and Maui. L. rhabdota, no. 3. 

d*. Length 2.7 to 3.1 mm. ; Hawaii. 

L. prisca, no. 5. 
b 2 . Later whorls distantly striate spirally; Hawaii. 

c 1 . Summit subconic. L. clathratula, 110. 6. 

c 2 . Summit abruptly truncate. L. truncata, no. 7. 

1. LYROPUPA LYRATA (Gld.). PI. 19, figs. 1 to 6. 

"The shell is small, most generally simstral, chestnut 
colored, widely umbilicate ; 5 convex whorls, with about 20 
neatly clathrate flexuous riblets ; aperture somewhat orbicular, 
bell-shaped, posteriorly armed with two lamella-like teeth and 



234 LYROPUPA. 

two others in the throat. Length 22/200, diam. 13/200 inch" 
[2.75 x 1.62 mm.] (Gould). 

Hawaiian Islands (Gould). Oahu: very abundant over 
nearly the whole length of the Koolau Mountains (Thaanum, 
Spalding, Gouveia, Emerson, Cooke, Pilsbry). Type and 
paratypes no. 219, G. 2687, Museum of the University of 
New York. 

Pupa lyrata GOULD, Proc. Bost. Soc., i, 1843, p. 139 ; Boston 
Journ. of Nat. Hist., iv, 1844, pi. 16, f . 16 ; Otia Conch., 1862, 
p. 189 (Habitat, Maui; "length 1/10, diam. 1/20 -f- inch") 
[2.54 x 1.27 mm.]. PFR., Monographia Hel. Viv., iii, p. 561 
nee Pupa lyrata PFR., Monographia Hel. Viv. iv, p. 686 nee 
BOETTGER, in von Martens' Conchologische Mittheilungen, i, 
1881, p. 61, pi. 12, f. 17. Vertigo lyrata H. and A. ADAMS, 
Gen. Rec. Moll., ii, p. 172. PEASE, Proc. Zool. Soc., London, 
1871, p. 474 (Oahu). Nesopupa (Lyropupa) lyrata, PILSBRY, 
Proc. A. N. S. Phila., 1900, p. 432. non Lyropupa lyrata 
ANCEY, Proc. Malac. Soc. London, vi, 1904, p. 124. Pupa 
magdalenae ANCEY, Mem. Soc. Zool. France, v, 1892, p. 716 
(Palama, Baldwin). Pupa lyrata SYKES, Fauna Hawaiensis, 
ii, 1900, p. 294. Lyropupa carbonaria ANCEY, Proc. Malac. 
Soc. London, vi, 1904, p. 125, pi. 7, f. 21 (Nuuanu). 

The specimens in the Albany Museum., the original material 
on which Gould based his species, bear the habitat Kauai. 
Gould later, in Otia Conchologica, gave Maui as the habitat, 
and the measurements in this description are smaller than 
originally given. Both of these localities are incorrect. The 
Lyropupae from Maui and from Kauai have been compared 
with Gould's specimens and found to belong to different 
species or subspecies. P. lyrata was doubtless from Oahu, 
where typical examples have been taken in Nuuanu valley. 

Gould's type lot is heterogeneous. This accounts for his 
expression "plerumque sinistrorsa" and for the discordant 
measurements of his two descriptions. There is one specimen 
of the widely spread Nesopupa newcombi in the lot, and two 
of the dextral Lyropupa perlonga, a common Oahuan shell. 
The larger one of these was probably the original of Gould's 
left hand figure, which represents a dextral shell. His other 



LYROPUPA. 235 

three figures show a sinistral shell, which may be taken as 
the type. This shell is drawn in pi. 19, fig. 4 ; or at least so 
we assume, as only this one specimen had the aperture well 
cleaned when the lot was examined. 

The spire is convexly conic. The ribs are strong through- 
out, 17 on the last whorl (Gould probably doubled the number 
seen on the face to reach the number "ad 20"). There is 
a rather deep external impression about a half whorl long 
over the upper palatal, and a shorter one, diverging forward, 
over the lower palatal. The lower palatal fold is rather long, 
as seen in the mouth. It becomes stronger towards the inner 
end, which penetrates slightly beyond the upper palatal fold. 
It is not hooked downward at the inner end, and not inter- 
rupted, though in some specimens of the lot there is a sub- 
median depression of the crest of the fold. Fig. 5 represents 
the palatal region of an example of the Gould series. Some 
others have the lower palatal fold somewhat shorter and 
lower anteriorly ( fig. 6 ) . 

Length 2.7, diam. 1.6 mm.; 5% whorls (type). 

Length 2.4, diam. 1.6 mm. (another of type lot). 

The contour varies, as the measurements indicate, the shorter 
examples being shaped like pi. 19, fig. 1, which represents a 
paratype of magdalenae, 

With this typical, strongly ribbed form of lyrata there are 
other specimens, probably collected from a different colony or 
locality, in which the ribs are weak and irregularly developed 
or partly wanting on the last whorl or two (pi. 19, figs. 8, 9). 
These may be called L. lyrata form gouldi. 

Pease's locality for this species, "Oahu," was based doubt- 
less on correctly named specimens. Boettger's description of 
lyrata was founded on specimens of another species from 
Hawaii. These are undoubtedly the same specimens used by 
Pfeiffer for his description. Ancey's description of lyrata 
was based on specimens of an unnamed species for which the 
name L. anceyana is proposed. 

Pupa magdalenae Ancey (type no. 18744 Bishop Mus., para- 
types 18745 B. Mus. and 119455 A. N. S. P.) is not separable 



236 LYROPUPA. 

from lymta, in our opinion. It agrees in shape with the 
shorter examples of the Gould series. The only difference is 
that the lower palatal fold has a rather deep depression be- 
tween the inner tubercle and the rather high outer portion; 
but this is a variable feature in the type lot of lyrata. Two 
paratypes of magdalenae are drawn in pi. 19, figs. 1, 2. The 
paratypes have 17 to 19 costae on the last whorl, with the inter- 
stices minutely, spirally, deciduously striate ; just back of the 
aperture are two shallow sulci which further back unite, form- 
ing a rather broad sulcus extending for nearly half the length 
of the whorl ; the embryonic whorls, in immature specimens, 
are minutely granulose, the granules arranged in transverse 
wrinkles; at about the middle of the second whorl the costae 
suddenly appear. The angular lamella is strong, high, thin 
and long and terminates at the margin of the aperture; the 
parietal lamella is more deeply seated, strong, slightly bent 
outwards and is nearly half a whorl in length ; the columellar 
lamella is strong, lunate, axially seated on the columella and 
extending to the base ; the lower palatal is rather short, very 
deeply seated, corresponding to the lower outer sulcus ; upper 
palatal long, elevated in back, low in front, terminating almost 
on the margin of the peristome ; with the angular it forms a 
well-defined sinulus; there is no indication of a basal fold. 
There are from 5 to 5*4 whorls in adult specimens. Ancey's 
paratypes measure : Length 2.56 to 2.7, diam. 1.56 to 1.65, aper- 
ture 0.95 to 1.0 mm. 

L. carbonaria Ancey appears, from the entire type lot (now 
no. 18752 Bishop Museum), to be synonymous with lyrata. A 
paratype is drawn in pi. 19, fig. 3. It is somewhat more 
elongate than the Palama magdalenae, representing the longer 
phase of variation, as that does the shorter. It is similar to 
Gould's type specimen of Urata, which probably came from 
the same valley, Nuuanu. The palatal folds (pi. 19, fig. 6) 
are exactly as in some specimens of lyrata though the lower 
palatal is shorter than in the lyrata selected for the figure. 

la. Lyropupa lyrata nncifera C. & P., n. subsp. PL 19, figs. 
12, 13. The shell is slightly smaller than that of the typical 
form, and more cylindrical in outline, with 17 to 18 strong, 
hardly flexuous costae. In its fossil state, the fine deciduous 
spiral striae are absent. There is a light-colored peripheral 
zone on the last whorl between tawny sutural and basal zones. 
Whorls 5, convex and separated by a deep suture; the last 
tapering towards the base. Parietal lamella more deeply 



LYROPUPA. 237 

seated and closer to the angular than in the typical form. 
Lower palatal fold converging towards the upper inwardly, 
its inner end enlarged and curving towards the base. 

Length 2.4, diam. 1.4 mm. ; 5% whorls. 

Length 2.55, diam. 1.5 mm.; 5y 2 whorls (type). 

Length 2.2, diam. 1.35 mm. ; 5 whorls. 

Oahu: Coral bluff 1% miles west of Kahuku (Cooke and 
Pilsbry, type loc.) ; Malaekahana, Kaipapau, Laie and Punaluu 
(Cooke). Fossil in Pleistocene and later deposits. 

The essential character of this northwestern race is the hook- 
like shape of the lower palatal fold. In L. lyrata it is nearly 
straight. 

In its typical form the race appears to be extinct, yet there 
are some living forms which approach it and are referred here 
provisionally. 

Recent specimens from Kamanaiki, Kalihi, show some ap- 
proach to the fossil form. The lower palatal has an enlarged, 
slightly decurved inner end, which is almost separated from 
the low outer portion. Those collected living in Glen Ada, 
Nuuanu, have the lower palatal fold strongly hooked down- 
ward (pi. 19, iig. 10). The external shape varies from some- 
what conic to that of uncifera. The variations and races of 
lyrata will not be understood until specimens from a large 
number of places are opened for study of the palatal folds. 
It will probably be found necessary to recognize a number of 
local races. 

Kauaian race. A slightly larger race which appears closely 
related to this subspecies though not identical, is found in 
a fossil state on Kauai at Limahuli (Dole, Cooke) and Haena 
(Cooke) . The shells agree with uncifera in having the parietal 
lamella very deeply seated and close to the angular lamella, 
but the lower palatal fold is low and rather wide, and its 
inner end descends very little. One of the specimens meas- 
ures: Length 2.62, diam. 1.45, alt. apert. 0.95 mm.; whorla 
5 Va- A specimen from Limahuli is drawn in pi. 20, fig. 1. 

Ife. Lyropupa, lyrata fossilis C. & P., n. subsp. PL 19, figs. 
7, 11. The shell is longer and narrower in proportion to its 



238 LYROPUPA. 

length, than L. lyrata and more cylindrical in outline,* 
the apical whorls are granulose-wrinkled, the rest of the whorls, 
strongly sculptured, the last with about 15 hardly flexuous 
eostffi. Lower palatal fold very short. 

Length 2.8, diam. 1.47, apert. 1.0 mm.; whorls 5*/ (type). 

Length 2.6, diam. 1.55 mm. 

Oahu: Very abundant in pleistocene deposits, Manoa, 
(Thaanum, Cooke) ; "Waimanalu (single specimen, Cooke). 
Type 11039 Bishop Museum ; paratypes 45279 Bishop Museum 
and 119462 A. N. S. P. 

The external features are about as in uncifera, but the 
lower palatal fold is somewhat more reduced than in any 
other form. 

2. LYROPUPA MICROTHAUMA Ancey. PL 21, figs. 2, 3. 

" Shell smaller than the preceding [mirabUis] and less 
cylindrical, hence more conoidal-oblong ; whorls 5, with the 
upper margin extended less outwardly, angular lamella more 
elongate and distinct, the upper palatal elongate, almost 
abruptly truncate near the external margin, the lower more 
elongate, scarcely drop-shaped, not at all parallel to the pre- 
ceding, but diverging gradually forward. Length 1.8, diam, 
1, apert. 0.66 mm." (Ancey). 

Oahu: Nuuanu, type loc. (Thaanum) ; numerous localities 
in the Koolau range from Keawaawa to Kaipapau (Spalding, 
Gouveia, Bridwell, Cooke) ; Waianae range, Pukaloa (Spald- 
ing, Cooke). Holotype 18750 Bishop Museum and paratypes 
18751 Bishop Museum and Thaanum Coll. 

Lyropupa microthauma ANCEY, Proc. Malac. Soc., London, 
vi, 1904, p. 126, pi. vii, f. 20. 

L. microthauma is somewhat intermediate between the group 
of L. lyrata and that of L. mirabilis. The embryonic whorls 
are minutely spirally striate, a character found in most of the 
species of the first named group. The angular lamella is 
shorter than in the other species of this group but higher 
and more developed than in any species of the second group. 
The columellar lamella is very deeply, almost axially seated. 
It is rather strong and viewed from above slightly semilunate- 



LYROPUPA. 239 

in outline. The abrupt termination of the upper palatal fold 
at the lip, and its slight enlargement there, are special features 
of the species. The lower palatal fold is shorter and stouter 
than the upper, and penetrates more deeply. In a few speci- 
mens the last whorl is distinctly closely and minutely striate 
spirally, but in most specimens the minute spiral striae are 
visible only on the ribs and have entirely disappeared from 
the interstices. 

Specimens from the type locality have from 19 to 21 rather 
strong slightly oblique ribs on the last whorl. Those from 
Kaipapau have 23 to 24. 

In a Kaipapau shell opened the lower palatal fold is some- 
what longer than in the Nuuanu preparation figured, and is 
curved, the concavity towards the base, the inner end being a 
little stronger. 

Length 1.96, diam. 1.04, apert. 0.65 mm. ; 6 whorls (type loc., 
head of Nuuanu Valley). 

Length 1.9, diam. 1.04, apert. 0.64 mm.; 5% whorls 
(Kaipapau). 

Length 1.92, diam. 1.1, apert. 0.65 mm.; 5% whorls 
(Pukaloa, Waianae Range). 

This species has never been found abundantly in any local- 
ity. It is usually taken in loose, damp talus more or less 
covered with dead leaves. It is probably distributed over the 
whole length of the Koolau Mts., as it has been found by a 
number of collectors in rather widely separated colonies. In 
the Waianae Mts. it has only been taken, on two occasions, 
from one of the little valleys. 

3. LYROPUPA RHABDOTA C. & P., n. sp. PL 20, fig. 2. 

The shell is sinistral, rirnate-perforate, subcylindrical, with 
slightly convex and tapering outlines, brussels brown, except 
for a rather broad lighter zone at the middle of the last 
whorl, hardly translucent, dull. Sculpture of strong, slightly 
oblique ribs, 12 to 14 on the last whorl, the interstices and 
ribs spirally striate with minute, close, raised threads. Spire 
tapering above to a slightly conical, obtuse summit. The in- 
dividual whorls very convex and separated by a deep suture. 



240 LYBOPUPA. 



Whorls 51/2, 1% embryonic, nearly smooth, marked by minute, 
close, raised, spiral lines, the transverse ribs beginning abruptly 
at this point. The last whorl tapers downward, and the latter 
part ascends slowly towards the aperture; there is a broad, 
shallow furrow extending for nearly % its length and ending 
just back of the lip. The aperture is perpendicular, trun- 
cate above, rounded below, peristome continuous. The inner 
half of the parietal margin is appressecl to the pemiltimate 
whorl, the outer half free. Angular lamella strong, erect, 
long, slightly curved outwards in cross section, extending to 
the outer margin; parietal lamella strong, somewhat curved 
spirally. Columellar lamella very deeply seated, obliquely 
placed on the columella, short. Upper palatal fold long, ex- 
tending nearly to the margin of the aperture, slightly modi- 
fied in front by the angular, with which it forms an almost 
complete sinulus. Lower palatal long, low, much more deeply 
seated than the upper and almost parallel with it. Peristome 
more or less expanded on all sides, thin at the margin. 

Length 2.7, diam. 1.53, apert. 1.0 mm. 

Molokai: Pelekunu, type loc. (Forbes), Wailau Pali and 
Kamalo (Cooke), Kalamaula (Thaanum, Pilsbry and Cooke), 
fossil at the base of the Kalaupapa cliffs (Cooke) and top of 
Mauna Loa (Pilsbry and Cooke). Type 11040, Bishop 
Museum; Paratypes 24871 B. M. and 119456 A. N. S. P. 

There can be no doubt that this species is distributed over 
the whole of the wooded portion of Molokai. Unfortunately 
it has never been found in any number in any one locality 
and in none of the lots are there more than half a dozen 
specimens in most of them only 1 to 3. 

This species is, at first glance, very similar to some of the 
forms of lyra-ta from Oahu. It is easily separated by the dif- 
ferent sculpture of the embryonic whorls and the longer lower 
palatal fold. 

3a. L. rhabdota plnris P. & C., n. subsp. PL 20, figs. 3, 4, 5. 

This short form of L. rhabdota is found along the pipe-line 
trail, upper Kaunakakai. The summit is obtuse and rounded, 
about as in L. thaa-numi. The spiral striae are weaker, less 
regular than in typical L. rhabdota. 



LYROPUPA. 241 

Length 2.25, diam. 1.3, aperture 0.8 mm. ; 5^4 whorls. 

Length 2.15, diam. 1.4, aperture 0.9 mm. 

The form from the western ravine of upper Kamalo (figs. 
3, 5) is similar but slightly larger. 

Length 2.45, diam. 1.5, apert. 0.95 mm. ; 5% whorls. 

Length 2.4, diam. 1.4 mm. ; 5% whorls. 

These forms were collected by the authors in some abund- 
ance among dead leaves in humid, shady places, with L. h. 
sparna. They differ from L. thcuinumi by the smaller size and 
fewer ribs. 

A still smaller race of L. rhabdota was found in Mapulehu 
Valley (Cooke) . Except for their smaller size the shells agree 
perfectly with the larger and more widely distributed typical 
form. Length 2.1, diam. 1.28, apert. 0.75 mm. ; 5 whorls. 

'3b. L. rfwbdota lanaiensis Cooke, n. subsp. PL 20, fig. 6. 

Specimens from Lanai differ slightly from the Molokai form. 
The shells are slightly smaller, more conical in outline, taper- 
ing more gradually towards a somewhat acute summit; they 
are darker colored and with the pale zone only indicated near 
the aperture. There are 16 to 18 ribs on the last whorl ; the 
embryonic whorls, under a strong lens, are distinctly, spirally 
striate; the columellar fold enters obliquely, is short, almost 
nodule-like when viewed from above, and there is usually a 
minute basal fold, situated deep within. Length 2.5, diam. 
1.5, apert. 0.92 mm. ; 5% whorls. 

Lanai: (Thaanum, Spalding, Forbes). Type 11041, Bishop 
Museum ; Paratypes 34182, Bishop Museum, and 119451 A. N. 
S. P. 

3r. L. rhabdota baldwiniana Cooke, n. subsp. PL 20, figs. 7, 8. 
Specimens from West Maui agree very closely with the type 
form in shape, color, etc., but are slightly smaller. They differ 
mainly in that the embryonic whorls are much more finely 
stria,te spirally, and in the presence of a minute basal fold and 
the more axially seated columellar fold. There are usually 
about two more ribs on the last whorl, which is furnished 
with from 15 to 17. Length 2.5, diam. 1.5, apert. 1.0 mm.; 
514 whorls. 



242 LYROPUPA. 

West Maui: lao (type loc.), and Waikapu (Cooke) ; Hono- 
kawai (Thaanum) ; Maunahoama (Forbes and Cooke) ; Hona- 
kahua (Fleming). Type 11042 Bishop Museum; paratypes 
38949 Bishop Museum and 119457 A. N. S. P. 

From L. r. lanaiensis it is separated by its more convex and 
swollen whorls, lighter color and much more finely spirally 
striate embryonic whorls. 

4. LYROPUPA THAANUMI C. & P., n. sp. PL 20, figs. 12, 13. 

The shell is sinistral, rimate-perforate, cylindrical, mars 
brown, with a very indistinct, broad, pale zone just above the 
periphery of the last whorl, chiefly visible on the ribs which 
are there dirty white ; rather solid, opaque, dull. Sculpture of 
strong ribs, 18 or 19 on the last whorl, the interstices with 
minute, close, deciduous spiral striae, only visible with a strong 
lens. Spire nearly cylindrical, with a rounded, blunt almost 
truncate summit, the first two whorls nearly immersed in the 
third ; last three whorls rather strongly convex, separated by a 
deep narrow suture. Whorls 5y 2 , the embryonic iy 2 when 
viewed under a strong lens (in adult specimens) are nearly 
smooth though indistinctly spirally striate. The last whorl 
ascends slightly near the aperture, tapering towards and con- 
tracted at the base, furnished with a broad, shallow furrow 
for a little more than half its length; the furrow bifurcates 
indistinctly near its termination, the upper branch extending 
to the margin of the lip. Aperture perpendicular, nearly 
quadrangular in outline, with rounded corners, and base, the 
peristome continuous, the inner half of the upper margin ap- 
pressed to the penult whorl, the outer free. The angular 
lamella is long, strong, its free edge slightly curved outwards, 
extending to the outer margin of the aperture. Parietal 
lamella strong, very deeply seated, bent outwards ; columellar 
lamella very deeply and almost axially seated. Upper palatal 
fold is long, high within, becoming low in front, emerging 
nearly to the margin of the aperture. Lower palatal fold 
very deeply seated and parallel to the upper, a little curved, 
extending deeper than the upper palatal, and enlarged, nodule- 
like, below the inner end of the latter. 



LYROPUPA. 243 

Length 2.75, diam. 1.67, aperture 1 mm. (type). 

Length 2.5 mm. 

East Maui : Auwahi, type loc. (Thaanum) ; Ukulele (Forbes) . 
Type 11043 Bishop Museum ; paratypes 36896 Bishop Museum, 
119452 A. N. S. P. and 7597 Thaanum collection. 

This species is related to L. rhabdota and its subspecies. It 
differs, however, in its larger size, more cylindric outline and 
the depressed, rounded summit. The same character separates 
it from L. prisca, which appears to be the most closely allied 
species. L. thaawumi is also related to L. truncata, but is 
easily separated by its less flattened apex, thicker shell and 
especially by the finer and closer spiral striae on the last whorl. 

5. LYROPUPA PRISCA (Ancey). PI. 20, figs. 9, 10, 11. 

The shell is sinistral, rimate-perforate, subcylindrical, in its 
fossil state pinkish cinnamon with a narrow nearly white band 
above the periphery on the last two whorls, thick, opaque,, 
strongly costate, with 15 to 17 slightly oblique ribs on the last 
whorl; the interstices very minutely, closely, spirally striate 
(but in its fossil state these striae are usually absent). Spire 
subcylindrical, tapering gradually above to a rather obtuse- 
apex; individual whorls convex, separated by a deep suture.. 
Whorls 5% to 5%, the embryonic 1% whorls (under a strong 
lens) nearly smooth, only traces of minute close, spiral striae 
remaining. The last whorl ascends slightly towards the aper- 
ture, and, tapers very gradually to the base; it has two dis- 
tinct nearly parallel shallow sulci which unite further in back 
into a broad shallow furrow. Aperture perpendicular, roughly 
triangular with broadly rounded base, its margins continuous. 
Angular lamella long and strong, emerging to the margin. 
Parietal lamella thinner than in most of the species and de- 
cidedly slanting outwards. Columellar lamella strong, deeply 
seated, obliquely entering, high in front, low in back, running 
into the columella above. Upper palatal fold long, extending 
nearly to the margin, high within, very low where opposite the 
angular. Lower palatal fold shorter and broader, deeply 
seated, nearly parallel to the upper, and penetrating inward 
beyond it, either simple or divided by a median depression 



244 LYROPUPA. 

(figs. 9, 10). There is a very small and very deeply placed 
basal fold, at least in some specimens. Peristome very slightly 
expanded. 

Length 3, diam. 1.62, apert. 1.1 mm. 

Length 2.7, diam. 1.6 mm. 

Hawaii : In pleistocene deposits at Mana and Palihoukapapa 
(Henshaw, Thaanum). Type 18746 Bishop Museum; para- 
types 18756 Bishop Museum, 119453 A. N. S. P., also in 
Henshaw and Thaanum colls. 

Lyropupa magdalenae var. prisca ANCEY, Journal of Mala- 
cology, London, xi, 1904, p. 68, pi. 5, f. 19. ? Pupa lyrata 
Gould, BOETTGER in v. Marts., Conchol. Mittheil., i, 1881, p. 
61, pi. 12, f. 17 (Auf der Iiisel Hawaii). 

The above description is not drawn from Ancey 's type speci- 
men but from another from Mana, as the former is not in per- 
fect condition. Ancey gave Palihoukapapa as the type local- 
ity but as this species is extremely rare in the deposits and 
common in those from Mana, I am of the opinion that the 
former locality is an error. Two nearly equal lots of earth 
from each locality were picked over and the Mana lot contained 
nearly 200 specimens of prisca to 1 from Palihoukapapa. From 
the number of specimens of prisca in the Ancey collection I 
have no doubt but that his came from the Mana deposit. 

L. prisca is in no way closely related to L. magdalence, the 
surface sculpture of the embryonic whorls is enough to separ- 
ate the two as species. It differs from the other species from 
Hawaii in having close, fine, spiral striae in the interstices be- 
tween the costa? in the most perfectly preserved examples; 
these stria? are almost or entirely absent in most the specimens 
examined, due to the fossil condition of the shells. Ancey 's 
measurements in the Journal of Malacology are somewhat con- 
fusing; his type measures, length 2.8, diam. 1.5 mm. 

Dr. Boettger's description and figure of what he identified as 
Pupa lyrata appears to have been based upon recent specimens 
of this or some very closely related Lyropupa. He placed 
V. striatula Pease in the synonymy of lyrata, but without hav- 
ing any knowledge of it other than the original description. 

L. thaanumi of Maui is very near akin to prisca; but the 



LYROPUPA. 245 

blunter summit together with the distribution 011 a different 
island incline us to separate it specifically. 

6. LYROPUPA CLATHRATULA Ancey. PL 21, figs. 1. 

" Shell sinistral, oblong, rimate-perf orate, opaque, with a 
somewhat silky sheen, deep purplish-black, unicolorous, some- 
what thin ; sculptured, except on the apex, with acute, oblique,, 
equidistant lamella-like lirae (12 011 the last whorl) ; in addition 
encircled and slightly clathratulate in the interstices with very 
slightly prominent elevated membranous lines, especially on 
the last two whorls. Spire ovate, obtuse. Whorls 5, convex, 
separated by an impressed suture, under a strong lens marked 
with lines of growth, the last furnished near the aperture with 
a not very distinct impression parallel to the suture, scarcely 
angulate about the umbilicus, ovate, a little tapering. Aper- 
ture suboblique, appressed, irregularly ovate, externally angu- 
late above, obstructed by 5 lamellae or plicae, namely : 2 parietal 
lamellae, one of which [parietal] is submedian, free, and the 
second shorter, more externally seated, united with the margin ; 
a deeply seated serniluiiate columellar, visible when viewed 
obliquely ; 2 long palatal, the upper almost reaching the mar- 
gin, the lower more deeply seated, corresponding to the shal- 
low, external sulcus. Peristome slightly expanded, purplish- 
brown, with the margins approaching above, but hardly con- 
tinuous. Length 2.25, diam. 1.5 mm." (Ancey). 

Hawaii: Olaa (Thaanum). Type 18742, paratypes 18768 
Bishop Museum, 119436 A. N. S. P. and Thaanum coll. 

Lyropupa clathratula ANCEY: Proc. Malac. Soc. London, 
vi, 1904, p. 125, pi. vii, f. 19 ; note bottom of p. 68, Journal of 
Malacology, xi, 1904. 

L. clathratula may ultimately prove to be a synonym of 
L. striatula. It is somewhat closely related to L. truncata; 
both have few and widely separated spiral striae ; in the latter 
species, however, the apex is much blunter and there are more 
apertural teeth. From the other species of Lyropupa it is 
most easily distinguished by the stronger and more distant 
spiral threads. The embryonic whorls are minutely, closely 
and regularly spirally striate. The columellar lamella is 



246 LYROPUPA. 

rather obliquely seated. In most of Ancey's specimens the 
peristome is continuous, the portion which is appressed to the 
penultimate whorl being very thin. The lower palatal fold 
is shorter than the upper and penetrates inward beyond it. 
Length 2.45, diam. 1.5 mm. The original account follows of 

LYROPUPA STRIATULA (Pease). "Shell cylindrical, oblong, 
somewhat solid, sinistral, umbilicate, longitudinally obliquely 
and flexuously strongly costate, with the interstices delicately 
transversely striate; whorls 5, rotundly convex, frequently 
rotund-angulate in the middle, the last scarcely extended, con- 
centrically broadly sulcate at its middle, rotund-angulate 
about the umbilicus, compressed at the base; apex obtuse; 
suture well impressed ; aperture subquadrangular, rotund at 
base, contracted by 3 lamellae, 2 on the parietal wall, the first 
[angular] large, united with the lip, the second median in- 
ternal; one on the superior portion of the columellar margin, 
attached to the labium ; peristome continuous, somewhat thick- 
ened, slightly expanded and reflexed, the outer lip posteriorly 
sinuous; reddish chestnut, with whitish ribs. 

' ' Length 2.5, diam. 1.5 mm. ' ' (Pease ) . 

Hawaii (Pease). 

Vertigo striatula PEASE, Proc. Zool. Soc., London, 1871, 
p. 461. 

The type of this species is not to be found in Pease's collec- 
tion in the Museum of Comparative Zoology. Pease men- 
tioned 3 lamellae, 2 on the parietal wall and 1 columellar 
lamella joined to the labium, that is, the inner lip ; implying 
that none were seen within the outer lip. As no known 
Hawaiian Pupillid has this structure, we assume that for the 
moment he forgot that the shell was sinistral, and the phrase 
"in margine columellari superne [lamella] 1 ad labium 
juncta" really applies to the outer lip and its upper palatal 
fold. It is quite natural that Pease, a careless observer, over- 
looked the real columellar lamella and the lower palatal fold, 
as these are deeply immersed. This hypothesis makes the 
description intelligible, and increases the probability that 
-strmtula is identical with L. clathratula. Since the point can- 
not be demonstrated beyond doubt, we think it best to ignore 
V. striatulci as a lost species unless new evidence is found. 



LYROPUPA. 247 

7. LYROPUPA TRUNCATA Cooke. PL 21, fig. 4. 

The shell is rimate; sinistral, cylindrical, dark brown, 
elathratulate, with about 14 well developed lamelliform costae 
011 the last whorl (between the costse 5-8 minute stride) and 
with 3-4 raised spiral striae on each whorl. Spire cylindrical ; 
apex almost flat, giving the shell a truncate appearance. 
Suture simple depressed. Whorls 5, the embryonic without 
transverse costae but with numerous minute close spiral striae, 
increasing rather rapidly; the rest convex, clathratulate, in- 
creasing slowly; the last slightly narrower than the penulti- 
mate, with two indistinct shallow, longitudinal depressions 
corresponding to the palatal plicae. Aperture perpendicular 
quadrate-ovate, with 7 folds, viz. : the angular lamella slightly 
arcuate, extending nearly to the margin of the peristome; 
the parietal a little deeper-seated, strongly developed; and, 
a minute dentiform infraparietal lamella; a deeply seated 
obliquely entering, but well developed columellar lamella; 
three nearly parallel palatal plicae, the upper extending nearly 
to the margin of the peristome, the lower more deeply seated 
and in addition to these a very short, deeply seated basal plica. 
Peristome continuous, slightly expanded. 

Length 2.6, diam. 1.66, aperture 1.1 mm. 

Hawaii: Kohala Mts. (Thaanum). Type 15411 Bishop 
Museum ; paratype in Thaanum coll. 

Lyropupa truncata COOKE, Occasional Papers of the B. P. 
Bishop Museum, iii, 1908, p. 211, text figure. 

An extremely rare species of which only a very few speci- 
mens have been taken. The only close relative known at 
present is L. clathratula, from which it is easily distinguished 
by its much blunter summit and more numerous apertural 
teeth. Unlike all of the other Lyropupas except clathratula, 
the ribs are thin with membranous edges. The spiral striae 
are further apart than in any other species. 

Section Lyropupilla, n. sect. 

Sinistral Lyropupas with a low, short angular lamella, and 
without spiral striation on the lower whorls. Type Lyropupa 
spaldingi. 



248 LYROPUPA. 

8. LYROPUPA SPALDINGI n. sp. PL 21, figs. 10, 12, 13. 

The shell is sinistral, rimate-perf orate, somewhat ovately 
fusiform, cartridge buff, somewhat thick, translucent with a 
silky sheen, marked with strong slightly flexuous costee, about 
17 on the last whorl, the interstices are nearly smooth, in- 
distinctly flexuously striate. Spire convexly conic, tapering 
gradually to a slightly blunt apex, the upper whorls convex, 
the last two very convex, slightly angulate above the periphery', 
separated by a deep suture. Whorls 6%, the embryonic 
minutely granulose-wrinkled, the costee appearing abruptly 
at about the middle of the second whorl ; last whorl tapering 
strongly towards the base, somewhat inverted-conic in outline, 
with a rather acute base, deeply concentrically tri-sulcate 
back of the aperture, the upper sulcus long and broad, ex- 
tending for nearly % the length of the whorl, and reaching 
almost to the lip. Aperture perpendicular, somewhat quad- 
rangular in outline with much rounded corners, the margin 
continuous, free. Angular lamella short, blunt, extending to 
the margin ; parietal lamella deeply seated, its outer end ex- 
tending just beyond the inner end of the angular, strong, long, 
its free margin bent outwards, flexuous ; infraparietal usually 
present, very deeply seated and nearly nodule-like or short 
lamella-like ; columellar lamella very deeply and axially seated, 
nodular, viewed from above lunate with the horns pointing 
outwards; upper palatal fold long, high in back, low in front, 
extending to the inner margin of the peristome ; lower palatal 
deeply seated, long, parallel to the upper; there is present a 
short, low, deeply and concentrically seated basal fold which 
terminates almost at the base of the columellar lamella. Peris- 
tome free, continuous, its outer margin expanded. 

Length 2.82, diam. 1.53, apert. 0.85 mm. 

Oahu: Waianae Mts. from Palehua to Mokuleia; type local- 
ity Puu Kaua (Cooke). Type 11048, Bishop Museum, para- 
types 37192, and 119470 A. N. S. P. 

This is by far the most abundant species of Lyropupa found 
on the Waianae Mts., and its habitat extends over the whole 
length of the range. It is especially abundant in talus, but is 



LYBOPCPA. 219 

alao found on dead leaves and twigs. It is usually taken in 
rather open country bat has also been found in damp forest 
patches as well as in talus in open situations. A basal riew 
of a young: specimen is drawn in pL 26, fig. 8. 

It is easily separated from L. mtrvbHis by its lighter color. 
fewer and more widely spaeed eosta*, its broader outline, the 
continuous lower palatal fold and the additional lamella? in 
the aperture. 



9. LYBOPCPA iimAiuiJK Ancey. PL 21. figs. 5. 6, 7. 

"Shell sinistraL dark brown, eylindrieally oblong, except 
for the apex regularly and obliquely sculptured with distant 
acute lira*, scarcely shining, rimate-perfdrate. Spire obtuse. 
Whorls 6, regularly increasing, convex, with an impressed 
suture; the last inflated in front of the aperture, afterwards 
broadly slightly constricted, hardly attenuate, impressed at 
its middle, somewhat compressed at the umbilicus. Aperture 
scarcely oblique slightly projecting towards the right at the 
base and externally towards the outer and upper margin. 
extended, armed on the parietal wall with a marginal and 
short dentiform lamella near the upper angle: a second, the 
parietal, much larger, deeply seated and nearly median; a 
deeply seated strong eohuuellar fold (only seen when viewed 
obliquely}, and 2 palatal lamelbe. the lower of which is gutti- 
form. short deeply seated, the upper long anil extending to 
the margin. 

Length 2.5, diam. 1.33. apert. (alt.) 0.75 mm/" {Ancey). 

Oahu (Aneey) ; Popouwehu Waianae Mts, (Spalding, Cooke, 
Pilsbry). Type Ig747 Bishop Museum. 

Pupa mtrabdi* AXCEY, BulL Soc, MaL France, vii, 1890. 
p. 339; Mem. Soc. ZooL France, v, 1892, pp. 716. Lyro- 
pupa mirabHif AXCEY, Proc. Malac. Soc^ London, vi, 1904, p. 
126, pL vii, f. IB. 

Ancey described this species from a single specimen and 
without exact locality. As his specimen came from the aper- 
ture of a specimen of Ack. mHstflixa, the Waianae Range is 
undoubtedly the original locality. L. mirabHis is distributed 
nearly the whole range. It is very abundant among dead 



250 LYROPUPA. 

leaves and debris in the woods on the western ridge of Popou- 
wela, with Endodonta and other small shells. 

In the type specimen there are 6% whorls, the embryonic 
whorls are wrinkled-granulose. The last whorl has 22 slightly 
oblique, slightly flexuous ribs. The angular lamella is short, 
low, blunt ; the parietal is deeply seated, curved, its edge bent 
outwards, hardly emerging further than the inner termination 
of the angular; the columellar is strong, deeply and axially 
seated on the columella, nodule-like viewed from the back. 
The lower palatal fold is short and rather high. Some dis- 
tance beyond it there is a strong, oblique, white tubercle below 
the inner end of the upper palatal plica. A very weak basal 
appears below it (fig. 6). The type specimen, according 
to my method of measuring, has the following dimensions: 
length 2.38, diam. 1.22, apert. (vert.) 0.75 mm. The speci- 
mens figured are from Popouwela. Figure 5 measures : length 
2.3, diam. 1.28, apert, (vert.) 0.75 mm. 6 whorls. 

10. LYROPUPA ANTIQUA C. & P., n. sp. PL 21, figs. 8, 9, 11. 

The shell is oblong, somewhat cylindric, slowly tapering to 
the apex, the whorls rather narrow and convex; sculpture of 
strong, somewhat oblique ribs, 15 to 17 on the last whorl. 
The last whorl has a broad and deep spiral concavity somewhat 
below the middle. Armature of the aperture is outwardly as 
in L. mirabilis, the angular lamella low and short, the parietal 
deeply placed, becoming strong deep within, flaring towards 
the upper palatal. The columellar lamella is rather massive. 
The upper palatal fold is relatively rather short. Lower 
palatal very deeply placed, low in front, where it is parallel 
with the upper, then stronger and turning downward, pene- 
trating well beyond the upper palatal (pi. 21, fig. 8). 

Length 2.8, diam. 1.4, aperture 0.8 mm. ; 6% whorls. 

Oahu : Manoa, in pleistocene deposits along the Upper Manoa 
Road (Thaanum, Cooke). Type 11047, Bishop Museum; para- 
types 45280, Bishop Museum and 119472 A. N. S. P. 

This species is easily separated from L. mirabilis by its 
larger size, more numerous whorls and fewer ribs on the last 
whorl. Internally it differs by the continuous, strongly bent 
lower palatal fold. 



LYROPUPA. 251 

It is the only species of the mirabilis group so far reported 
from the Koolau Mountains. It is abundant in the deposits 
from the upper portion of the road, but entirely absent from 
the lower deposits less than half a mile distant. It appears 
to be rather local, but it may have had a wider extension be- 
fore the destruction of the lower forest zone. 

11. LYROPUPA HAWAIIENSIS (Ancey). PL 22, figs. 12, 14. 

The shell is sinistral, rimate-perforate, oblong, thick, opaque, 
in its fossil state brownish, marked with strong transverse, 
slightly flexuous ribs, 19 to 22 on the last whorl, the interstices 
minutely flexuously striate, parallel to the ribs. Spire with 
convex outlines, rapidly tapering above; apex rather obtuse. 
The individual whorls are rather strongly convex, separ- 
ated by a rather narrowly impressed suture. Whorls 
5% to 6, the embryonic 1% nearly smooth, under a 
strong lens very minutely granulose, the transverse costse ap- 
pearing abruptly at about the middle of the second whorl. 
The last whorl has a somewhat flattened base ; furnished with 
a rather broad shallow sulcus on about the last % of its length, 
the sulcus ending just behind the expanded peristome. Aper- 
ture perpendicular, indistinctly triangular in outline, with 
very rounded corners; peristome continuous, with free mar- 
gins. Angular lamella short, blunt and low, extending to the 
margin; parietal strong, very deeply seated, a little curved 
concentrically, not very long, oblique, its free margin flaring 
outward. Sometimes there is an indistinct nodular, deeply 
seated infraparietal present. Columellar lamella short, nodu- 
lar, high, deeply seated and very obliquely entering; upper 
palatal fold rather short, low and not emerging to the lip. 
Lower palatal is somewhat higher and more deeply seated, 
nearly parallel to the upper, which reaches about to the middle 
of its length. Basal fold transverse, deeply seated, blunt, 
Peristome continuous, expanded. Length 2.58, diam. 1.45, 
apert. 0.95 mm. (fig. 14). 

Hawaii: Palihoukapapa and Mana (Henshaw, Thaanum) ; 
fossil. Type 18748 Bishop Museum, paratypes 36656 Bishop 
Museum and 119468 A. N. S. P. Also in Thaaiium and Hen- 
shaw collections. 



252 LYROPUPA. 

Lyropupa mirabilis var. hawcwiensis ANCEY, Journal of 
Malacology, xi, 1904, p. 68, pi. 5, f. 19. 

The above description is based on a specimen from the ori- 
ginal material from which Ancey's type came, and may be 
considered a paratype. It agrees with Ancey's type which 
is slightly broken. 

This species is superficially close to L. mirabilis. It is here 
considered distinct on account of the fewer costae on the last 
whorl, the relative proportions of the lower palatal fold, which 
is not interrupted, to the upper, and the presence of a basal 
fold. The possession of this fold shows some relationship to 
L. anceyana from which it is easily separated by its oblong, 
less ovate form, the less convex whorls and more numerous 
ribs on the last whorl. 

Ancey's description follows: "Separated from the type 
(from the Waianae Mts., Oahu) as the shell is usually a little 
larger, more robust, with pale costulas; with a stronger and 
more elongate lower palatal lamelliform tooth ; whorls 6. 
Length 2.5, diam. 1.25, apert. (long.) 0.75 mm. There are 
twenty to twenty-two costulse on the last whorl, while in the 
type specimen there are about twenty-two to twenty-four. ' ' 

According to our methods, Ancey's type specimens has the 
following measurements: Length 2.45,_ diam. 1.35, apert. 
0.85 mm. 

12. LYROPUPA SPARNA C. & P., n. sp. PI. 22, figs. 6, 7, 10, 11. 

The shell is subcylindric, with rather long terminal cone, 
walnut brown or darker, with sculpture of rather short narrow 
ribs, about 20 on the last whorl, intervals microscopically 
granulose. There is a deep median sulcus on the last half 
whorl. Aperture is somewhat triangular, with well-expanded, 
pale, continuous peristome. The angular lamella is a low 
ridge or reduced to a mere swelling. Parietal high but short. 
The columellar lamella is short, thick, rounded, somewhat 
transverse to the columella. The upper palatal fold is low 
and not very long; lower palatal deeply placed, somewhat 
stronger than the upper, and penetrates much further in; it 
is usually a little curved. The basal fold is a low rounded 
or oblong tubercle. 



LYROPUPA. 



253 



Length 2.05, diam. 1.15, aperture 0.75 mm. (type). 

Length 2.1, diam. 1.33, aperture 0.75 mm. 

Length 2.2, diam. 1.2, aperture 0.8 mm. 

Molokai: Kalihi, type loc. (Pilsbry & Cooke), Kalamaula 
(Thaanum), Kawela. (Cooke), also fossil at Kaluaaha, Kalau- 
papa and Kaiehu (Cooke); Lanai: Mahana and Kaiholena 
(Forbes). Type 33627 Bishop Museum (fig. 6) ; para type 
108919 A. N. S. P. (figs. 7, 10, 11). 

L. sparna differs from L. mirabilis by the palatal armature. 
It resembles L. antique, in this respect, but the shell is smaller 
and the ribs thinner and more numerous. L. hawaiiensis is 
a larger, more robust shell, with less triangular aperture and 
a stronger basal fold, but closely related. It occurred along 
the pipe-line trail in some abundance among dead leaves. 

As far as known, only two specimens have been collected on 
Lanai, one from each of the localities mentioned above. These 
two specimens agree very closely with those from Molokai. 
One of them measures, length 2.1, diam. 1.25 mm., 5% whorls. 

12a. L. sparna simdifera P. & C. PL 22, fig. 13. 

The angular lamella is developed as a distinct though small 
plate, and the upper palatal fold is somewhat stronger than 
in sparna>, the sinulus being therefore better defined. The 
lower palatal fold is strong and deeply placed, more than 
half of its length beyond the inner end of the upper palatal. 

Length 2.2, diam. 1.1 mm. 

Molokai : Western ravine of Kamalo ; also in small numbers 
along the pipe-line trail with L. sparna (Cooke & Pilsbry). 

13. LYROPUPA ANCEYANA C. & P., n. sp. PL 26, figs. 3, 6. 

The shell is sinistral, ovate, chestnut-brown, scarcely shin- 
ing. Sculpture of strong, rather narrow, equidistant, oblique 
ribs, slightly flexuous on the last whorl, where they are often 
more prominent above the middle, and number 12 to 14; 
minute striae of growth may be seen between the ribs, but 
no spiral strise. The first 1% whorls are microscopically 
granulose. The spire is oblong-tapering, the summit obtuse. 
The whorls are very convex, the suture deep and undulating. 



254 LYROPUPA. 

The last whorl is impressed by a moderately deep and wide 
sulcus parallel with the suture, the base being narrowly 
rounded and prominent. The aperture is rounded below, 
truncate above, the sinulus not very well denned. The angu- 
lar lamella is short and very low. The parietal lamella is 
median, oblique, deeply placed, its deeply penetrating inner 
part stronger. The columellar lamella, rather deep within, 
appears as a plate with arcuate outline ; the upper end recedes, 
the plate standing obliquely. The upper palatal fold is low, 
slender, and does not penetrate far. Lower palatal is more 
deeply placed, very strong, somewhat curved, and penetrates 
to the back. There is a strong but short, obliquely axial basal 
fold. The peristome is whitish, somewhat thickened and well 
expanded, continuous. 

Length 2.25, diam. 1.4, aperture 0.72 mm. ; 5% whorls. 

Hawaii: Olaa (Thaanum). Type 11050 Bishop Museum; 
paratypes 18769 Bishop Museum, 119469 A. N. S. P. and 838 
Thaanum Coll. 

Lyropupa lyrata (Gld.) ANCEY, Proc. Malac. Soc. London, 
vi, 1904, p. 124; not Pupa lyrata Gould. 

This species is based on specimens believed by Mr. Ancey 
to be Gould's Pupa lyrata. It is well distinguished from 
allied forms by the very convex whorls, the unusually strong 
lower palatal fold and the short, high, deeply placed basal 
fold, stronger than in any related form. It is not nearly re- 
lated to L. lyrata, which differs by the well-developed angular 
lamella and the spiral striation. 

L. anceyana stands close to L. hawaiensis, but it is smaller 
with the ribs more widely spaced, the spire more tapering, and 
the whorls are conspicuously more convex. 

14. LYROPUPA SCABRA P. & C., n. sp. PI. 26, figs. 1, 2. 

The shell is perforate and rimate, tapers above, last two 
whorls forming a short, cylindric portion; russet with sculp- 
ture of well-spaced ribs, about 16 on the last whorl. Last two- 
thirds of the last whorl has a deep and wide sulcus. The 
aperture is dark within with pale brown peristome and teeth. 
Angular lamella short, rather low. Parietal strong. Colu- 



LYROPUPA. 255 

mellar lamella strong, almost transversely placed upon the 
columella. Upper palatal fold small, shorter than the lower, 
and not quite reaching the inner margin of the peristome; 
the entire length of both seen in a front view. 

Length 2.8, diam. 1.6, apert. 1.1 mm.; 5% whorls. 

East Maui: Ukulele, type loc. (Forbes). W. Maui: Mauna- 
hooma (Cooke). Type 11049 Bishop Museum; paratypes 
12639, Bishop Museum and 119465 A. N. S. P. 

Only five adult specimens of this race have been seen, four 
from the type locality, one from West Maui. 

The shell is less fusiform than L. spaldwgi, darker colored, 
the base less produced, and the palatal folds shorter; but it 
differs conspicuously by the single broad and deep furrow on 
the back of the last whorl. 

Section Mirapupa C. & P., n. sect. 

The shell is dextral ; the angular lamella is high and rather 
long; the surface is ribbed and minutely striate spirally (when 
unworn). Other characters as in Lyropupa. Type Lyro- 
pupa perlonga (Pse.). 

Typically the lower palatal fold is short, and there are two 
tubercles forming a vertical barrier under the inner end of 
the upper palatal fold; but some species have the palatals as 
in typical Lyropupae. 

Except in being dextral, these shells resemble the typical 
Lyropupae. 

On all of the Hawaiian islands, these are Lyropupae chiefly 
of the arid lower zone, probably nowhere found in the zone 
of humid forest. They live under stones. Owing evidently 
to increasing aridity since the Pleistocene, many forms have 
become extinct, and the others are local and infrequently found 
as living snails. They are among the most abundant Pleisto- 
cene and Holoceue fossils, in many places to be collected by 
hundreds or by thousands. 

The peculiar feature of most Mirapupae is the inner barrier 
formed of two short folds or tubercles below the inner end of 
the upper palatal fold, as in pi. 25, fig. 15 and other figures. 
These internal tubercles are the enlarged and isolated inner 



256 LYROPUPA. 

ends of the lower palatal and basal folds, which are otherwise 
reduced or sometimes absent. The structure is strictly com- 
parable to that of some Abidas, such as are illustrated in vol. 
24, pi. 44, fig. 6. L. kahoolavensis is a relatively primitive 
species not showing 1 this specialization. Among the sinistral 
Lyropupa* there is similar interruption of the lower palatal 
fold in L. mirabilis. 

15. LYROPUPA KAHOOLAVENSIS P. & C., n. sp. PL 22, figs. 1 to 
4, 8, 9. 

The shell is cylindric, obtuse at the ends, cinnamon-brown, 
very thin. Earlier whorls are strongly convex, the last two 
flattened in the middle, bulging below the deep suture. Em- 
bryonic 1% whorls are granose-vermiculate, the next whorl 
closely obliquely ribbed. The last 3 whorls, forming the cylin- 
dric portion, have widely spaced ribs, at first continuous and 
oblique (retractive), but on the last two whorls they are 
subvertical and widely spaced on the upper half of the whorl, 
interrupted in the middle, and on the lower half more oblique 
and irregular, and more numerous; minute spiral striae may 
be traced in the intervals. The last whorl has three impres- 
sions parallel to the suture, the upper long, reaching the lip, 
the others short, the third one being basal. The aperture has 
the usual squarish form. Peristome continuous, expanded, 
whitish. Lamellae of parietal wall as in L. perlonga except 
that the parietal does not penetrate so deeply. The columellar 
lamella is very deeply placed, vertical, but the lower end bends 
a little outward, the upper end bending inward. The upper 
palatal fold enters about one-third of a whorl. The lower 
palatal is immersed, continuous, bent downward and enlarg- 
ing at the inner end, which is even with that of the upper 
palatal. Below it there is a tubercular basal. 

Length 2.15, diam. 1.05 mm. ; 5% whorls. Type, Hakioawa 
Bay. 

Length 2.3 mm. ; 51/2 whorls. 

Length 2.4, diam. 1.2 mm. Hanakanaea or Smuggler's Bay. 

Length 2.6, diam. 1.2 mm. Kanapou. 

Length 2.55, diam. 1.15 mm. Kanapou. 



LYROPUPA. 257 

Length 2.75, diam. 1.15 mm. ; 6 whorls. Kaiiapou. 

Length 2.15, diam. 1.15 mm. ; 5 whorls. Lanai. 

Length 2.5, diam. 1.2 mm. ; 6 whorls. Maunalei, Lanai. 

Length 2.2, diam. 1.1 mm.; 5% whorls (Kona crater, 
Hawaii). 

Kahoolawe: Hakioawa (type loc., figs. 1-3, 9), Kanapou and 
Hanakanaea Bays (Cooke, Stokes and Pilsbry) ; probably 
wherever fossils occur, as these places are near the north, 
southeast and western angles of the island. 

Lanai : Maunalei (Forbes) . 

Molokai: Summit of Mauna Loa and at Moomomi (Cooke 
and Pilsbry) ; Mauna Loa near the shifting sands and Kaiehu, 
near Moomomi (Cooke). 

Hawaii: Kona crater (Thwing; pi. 22, f. 4, 8). 

This species, which seems to be widely distributed on 
Kahoolawe, and the only Lyropupa known from that island, 
differs from other Mirapupas by the better developed, longer 
and continuous lower palatal fold. The enlarged inner end 
of this fold represents the iipper of the two tubercles of the 
inner barrier of the other species. The Kahoolawe species 
is also characterized by the sculpture and thin shell. No 
specimens were collected alive, though some appear fresh. 
The island is now a mere desert, many square miles without 
humus or herbage of any kind. At the third locality it was 
found but a few feet above sea level ; at the others in aeolian 
material at higher levels, up to several hundred feet above 
the sea. 

Specimens from Hawaii (no. 119466 A. N. S., 12438 Bishop 
Mus.) are somewhat more solid than those of Kahoolawe. In 
the specimen figured there are about 20 ribs on the last whorl 
(pi. 22, figs. 4, 8). 

In Molokai this species is somewhat widely distributed on 
the treeless western part, as a fossil of Pleistocene and more 
recent age, sometimes in aeolian deposits of calcareous sand, 
but on Mauna Loa under stones where they lived. Usually 
the lower palatal fold is shorter, its inner half higher than 
typical, and the tubercle below it is generally strongly de- 
veloped (pi. 26, fig. 13, Mauna Loa, near the shifting sands). 



258 LYEOPUPA. 

On Lanai, as on Kahoolawe, this is the only Lyropupa known 
to exist. 

While some slight insular differentiation has been noted 
above, it is too variable to allow us to speak of races on the 
different islands. Cases of similar distribution have been re- 
corded in Leptachatma and the Tornatellinidae. 

L. kahoolavensis puukolekolemis P. & C., n. subsp. PI. 26, 
figs. 9, 12. 

Recent specimens from Puukolekole, Molokai, collected by 
Thaanum, are tawny or a little darker, irregularly sculptured, 
numerous riblets being intercalated on the lower half of the 
last whorl or two. The sinulus is more distinct than in Oahuan 
perlonga as the upper palatal fold emerges nearly to the edge. 
The lower palatal fold is a rather large callous pad, as seen 
in the mouth. It has a lower connection backward with the 
high tubercle below the inner end of the upper palatal. There 
is a quite weakly indicated basal callus, and the usual tubercle 
deep within. 

Length 2.1, diam. 1.2 mm.; 5 whorls (fig. 9). 

Length 2.25, diam. 1.25 mm. ; 5 whorls. 

This form has a lower palatal fold intermediate in develop- 
ment between kahoolavensis and perlonga, but the sculpture 
is practically as in the latter. It is more specialized than the 
fossil kahoolavensis, as the connection between the anterior 
and posterior portions of the lower palatal is low, and except 
in certain lights the fold appears interrupted. 

16. LYBOPUPA PERLONGA (Pease). PL 23, figs. 1-6, 13, 14; 
pi. 24, figs. 6, 7. 

"Shell elongate, cylindrical, rimate-perf orate, dextral, pale- 
straw color, longitudinally thread-like costate, the costae 
oblique, remote, flexuous; spire obtuse; suture impressed; 
whorls 5, somewhat flat, the last concentrically sulcate at the 
middle, indented near the base; the base compressed, rotund- 
ate-angulate about the umbilicus; aperture somewhat quad- 
rangular, slightly extended, contracted at the base, rotund, pos- 
teriorly bilamellate, the upper lip sinuous above, unilamellate ; 



LYROPUPA. 259 

peristome continuous, scarcely everted. Length 2.5, diam. 
1mm." (Pease). 

Oahu : (Pease) . Very abundant in pleistocene and holocene 
deposits on the costal plain of Oahu, Diamond Head eastward ; 
Kailua ; living examples from Koko Head. 

Vertigo perlonga PEASE, Proc. Zool. Soc. London, 1871, p. 
462. Pupa perlonga PFEIFFER, Mon. Hel. Viv., viii, p. 377. 
BOETTGER, in von Martens, Conchologische Mittheilungen, i, 
1880, p. 60, pi. 12, f. 16. Nesopupa (Lyropupa) perkmga, 
PILSBRY, Proc. A. N. S. Phila., 1900, p. 432. Lyropupa, per- 
longa ANCEY, Proc. Malac. Soc. London, vi, 1904, p. 126. 

This is the most widely distributed and commonest Oahuan 
species of Lyropupa. Numerous specimens of the typical 
form or of one of its races are found in most of the fossil 
deposits adjacent to the Koolau range. Living examples are 
only occasionally found and are hardly ever abundant ; as far 
as known, less than a dozen lots have been reported. In every 
case where the junior author found living examples they were 
taken in open country under dead sticks and stones. Only 
once were they found in any number, and that was near the 
top of Koko Head, at an elevation of about 1,000 ft. 

Pease's unique type specimen, No. 48063 Mus. Comp. Zool., 
is drawn in pi. 23, fig. 1, with the aperture enlarged, fig. 2. 
It is a fossil shell, a good deal worn, the ribs worn from the 
upper part, giving it a more tapering outline than it origin- 
ally had. There are about 17 narrow ribs on the last whorl, 
0.17 mm. apart on the face of the whorl, closer behind the lip, 
part of them extending over the strong basal ridge, which 
has an obtuse hump in the middle, preceded and followed by 
dimples, visible in basal view. The outer edge of the lip is 
broken away. The angular lamella is long, parietal lamella 
higher, entering deeply, emerging to the middle of the angular* 
The columellar lamella is seen as a broad, vertical plate, in 
front view. Upper palatal fold emerging to the lip. Lower 
palatal fold a low, callous nodule. Internal barrier, so far as 
it is visible in the mouth, as in pi. 24, fig. 7. Length 2.1, diam. 
1.05 mm. 

The dimensions do not agree fully with Pease 's description, 



260 LYROPUPA. 

but his measurement was doubtless only approximate. He 
gave the locality Oahu, and the label (in Anthony's hand) 
gives Nnuanu. It was probably from some Holocene deposit 
at a low level. 

Specimens from the bench of calcareous sandstone east of 
Diamond Head agree closely with the type specimen (pi. 23, 
figs. 3, 4, 5; also pi. 24, figs. 6, 7, showing the lamellae and 
an inside view of the palatal region ) . 

The summit is blunt ; the embryonic whorls are coarsely 
wrinkled-granulose ; the first three whorls increasing rapidly, 
the third whorl swollen, as broad as the last two; the later 
two or three whorls are slightly swollen below the suture, the 
last whorl tapering and narrowing below ; there are about 18 
costae on the last whorl, the back of which is trisulcate, the 
uppermost sulcus long, shallow and broad, lightly indicated 
on the penult whorl, the two lower being short, dimple-like; 
the margin of the peristome is free; the angular lamella is 
long and low, extending to the margin; the parietal more 
deeply seated, emerging to the middle of the angular; the 
columellar lamella is deeply seated, its upper part vertical, the 
lower end sloping towards the aperture somewhat. In shape 
it is a semi-circular plate, slightly concave on the side next 
the aperture. The upper palatal fold is long, slender, its 
crest only slightly sinuated where approached by the angular 
and parietal lamellae. The lower palatal fold is nodule-like, 
low and short; beyond it, further in, there is a strong, tri- 
angular or comma-shaped barrier, a lower callus below it (pi. 
24, fig. 7, interior view of palatal wall). Specimens from the 
raised bench of consolidated coral sand east of Diamond Head 
measure : 

Length 2.45, diarn. 1.05, aperture 0.8 mm. ; 5y 2 whorls. 

Length 2.25, diam. 1.1 mm. 

Length 2, diam. 1 mm. ; 5 whorls. 

The same typical form is found on Diamond Head near 
the lighthouse, and on the floor of the crater. All of these 
are fossil. 

Living specimens collected 011 Koko Head (pi. 23, fig. 6; pi. 
26, figs. 4, 7) are brussels brown in color, they are slightly 



LYBOPUPA. 261 

smaller than the typical perl&nga, with closer costee, and have 
distinct though minute close spiral striae in the intercostal 
spaces. There are about 20 riblets on the last whorl. 

The lower palatal fold is comma-shaped (pi. 26, fig. 4) in 
some examples, but in others there is a very low anterior 
callous pad, connected with the higher posterior portion (pi. 
26, fig. 7). This may be a more mature phase. Length 2.15, 
diam. 1.2 mm. The shell of nearly 3 whorls is drawn in pi. 23, 
figs. 13, 14; diam. 1 mm. 

At Kaelepulu, Kailua, on a low lime-rock bluff about a quar- 
ter mile from the shore, this species is abundant and variable, 
extreme specimens measuring : 

Length 2.4, diam. 0.96 mm. ; 6 whorls (pi. 24, fig. 10). 

Length 1.9, diam. 1 mm. ; nearly 5 whorls (pi. 24, fig. 9). 

It occurs with L. ovatula, L. micra and L. m. percostata, 
also with other land shells, all now extinct in that place. The 
lower palatal fold is reduced to a very low, small rounded 
callus, difficult to see. The same form is found on the neck 
of Mokapu peninsula, likewise extinct. 

Form cyUndrata, pi. 23, fig. 8. Specimens from a few locali- 
ties differ somewhat from the typical form. The shells are 
longer, more cylindrical in outline and the third whorl is 
usually slightly more swollen and wider than the lower whorls. 
The lower palatal fold is shorter than in the forms described 
above. 

Length 2.6, diam. 1.05, apert. 0.73 mm. ; 5% whorls. Oahu : 
Makua ; type loc. 

Length 2.7, diam. 1.1, apert. 0.8 mm.; 6*4 whorls. Oahu: 
Diamond Head, sea-cliffs. 

16a. Lyropiipa perlonga ititerrupta P. & C., n. subsp. PL 22, 
figs. 5 ; pi. 25, figs. 1 to 4, 10. 

The shell is cylindric, cinnamon or somewhat darker, the 
whorls flattened, swollen along the suture, with sculpture of 
irregular ribs, on the last whorl or two interrupted in the 
middle. Impressions on the last half whorl are strongly de- 
veloped. The parietal lamella is very long, curving at its 
inner end . Columellar lamella is crescentic, both ends curving 



262 LYROPUPA. 

towards the aperture. The internal palatal tubercles are 
strongly developed, the lower palatal fold otherwise very low, 
in form of a rounded callus pad (fig. 2). 

Length 2.25, diam. 1.1 mm. ; 5% whorls ; type. 

Length 2, diam. 1.2 mm. ; 5 whorls. 

Length 1.8, diam. 1.1 mm. ; 4% whorls. 

Oahu: abundant in debris of the "coral bluff" 1% miles 
west of Kahuku (Cooke & Pilsbry) ; Maleakahana (Cooke). 

It is readily distinguishable by the irregular sculpture. The 
internal structure is much as in ovatula, but it has not the 
strongly differentiated basal crest of that. The parietal 
lamella penetrates much further than in Diamond Head 
perlonga, and the columellar plate is larger and more curved. 

Occasional individuals are very short but of about the dia- 
meter of the longer ones, as in the third measurement given 
above. Sometimes there is an indistinct dorsal hump near the 
base of the last whorl. 

A small form, resembling the smallest found at Kahuku, has 
been taken at Maleakahana (pi. 25, fig. 10) ; dune deposit west 
of the Laie stream, between the highway and the sea (Cooke 
& Pilsbry), on the north shore, near the northwestern point 
of Oahu. 

There is an inconspicuous, rounded basi-dorsal prominence ; 
the lower palatal is rather strongly developed. It is wider 
than L. micra of the same length, but approaches it somewhat 
by the incipient hump. Length 2, diam. 1.1 mm. 

16&. L. perlonga filocostata C. & P., n. subsp. PL 23, fig. 12, 
Specimens from Kauai and Niihau differ from those from 
the other islands in that the surface is crowded with slender, 
oblique, flexuous costee (sometimes quite irregular), about 28 
to 30 on the last whorl. All the specimens examined are fossil. 
In some of these the delicate spiral striae are still present. 
The lower palatal fold is low but comparatively strongly 
developed, and appears to be weakly united with the strong, 
oblique tubercle further in. There is a weak basal fold, fol- 
lowed within by another callus tubercle. 

Length 2.15, diam. 1.15, apert. 0.8 mm.; 5^ whorls; Kauai: 
Limahuli, type (Cooke). 



LYROPUPA. 263 

Length 2.5, diam. 1.18, apert. 0.75 mm.; Niihau: Kiekie 

(Stokes). 

17. LYROPUPA MICRA C. & P., n. sp. PI. 23, fig. 7; pi. 25, 
figs. 5, 6, 7. 

The shell is smaller than L. perlonga, the whorls nearly fiat ; 
ochraceous-tawny, with sculpture of many low riblets, which 
are somewhat irregular and unevenly spaced. The last half- 
whorl has the usual three impressions well developed, and be- 
low the middle there is a vertical ridge or hump, running to 
the base, about a third of a whorl behind the outer lip. The 
sinulus is small and rounded. The parietal lamella is not 
very long, but penetrates beyond the strong columellar lamella, 
which is straight and vertical. The upper palatal lamella 
penetrates to the dorsal side. Lower palatal is deeply im- 
mersed, short but large. The basal fold is narrow and longer, 
but also deeply placed. There is a vertical barrier below the 
end of the upper palatal, composed of two strong tubercles 
separated by a rather narrow sinus. 

Length 1.9, diam. 1.1 mm.; 5 whorls (Kaelepulu, type). 

Length 1.9, diam. 1.05, aperture 0.65 mm.; 5 whorls 
(Lualualei). 

Length 1.8, diam. 0.95, aperture 0.65 mm.; 5 whorls 
(Rocky Hill). 

Oahu: Kaelepulu, Kailua, on a lime-rock bench about % 
mile from the north shore (Pilsbry, type loc.). Eocky Hill, 
Manoa valley (Cooke; fig. 7) ; and in the Waianae region at 
Lualualei (Cooke) and Kawaihapai (Cooke and Pilsbry). Liv- 
ing at Lualualei, elsewhere fossil. A subspecies in Molokai. 

L. micra is more slender than L. thaumasia. The dorsal 
hump, while variable in prominence, appears to be constant 
in the great number seen. Both on Oahu and 011 the dry 
western part of Molokai it occurs with larger species of the 
perlonga group. The living specimens from Lualualei are 
cinnamon-colored. 

The specimens from Kaelepulu, Kailua (pi. 25, figs. 5, 6, 7), 
have the sculpture weaker than most of those from Eocky 
Hill though there are often prominent, thin laminae near the 



264 LYROPUPA. 

base. The Rocky Hill example figured, pi. 23, fig. 7, is one 
of the most strongly sculptured. 

17a. L. micro, percostata P. & C., n. subsp. PL 25, figs. 11, 12. 

The ribs are more prominent than in L. micra, more separ- 
ated, often in pairs. There is a prominent dorsal hump, below 
the middle of the last whorl. The parietal lamella is deeply 
placed, high within, its free edge flaring towards the suture. 
Columellar lamella immersed, vertical, nearly straight, the 
ends very weakly bent forward. Upper palatal fold about, 
one-third of a whorl long. Lower palatal a small, low and 
indistinct rounded callus, deeply immersed. Basal fold 
deep within, blunt and elongate. There are well-developed 
tubercles below the inner end of the upper palatal fold, be- 
yond the lower palatal and basal. 

Length 1.8, diam. 0.9 mm.; 5 whorls (type). 

Length 2 mm. ; 5% whorls. 

Oahu : Kaelepulu, Kailua, on a lime-rock bluff about a 
quarter mile from the shore ( Pilsbry ) . 

Chiefly distinguished by its sculpture. This and the less 
strongly ribbed L. micra are abundant in the same Holocene 
deposit, without intergrading forms. 

176. L. micra maunaloae P. & C., n. subsp. PI. 25, figs. 8, 9. 

The Molokai form is somewhat more sharply sculptured than 
that of Oahu, and there are low protractive waves below the 
suture on the last whorl or two, or the riblets are grouped by 
fascicles there. 

Length 1.95, diam. 9 mm. (type, fig. 9). 

Length 1.75 mm. 

Length 1.9, diam. 0.97 mm. ; 5 whorls. 

Molokai: summit of Mauna Loa, and at Moomomi on the 
north shore, near sea level (Cooke & Pilsbry) ; near the shift- 
ing sands, Mauna Loa (Cooke). Pleistocene ?, and on the 
summit of Mauna Loa later, within the human period, though 
now extinct. 

Some of the examples from Moomomi are a little larger 
2 to 2.1 mm. long. 



LYROPUPA. 265 

18. LYROPUPA OVATULA C. & P., n. sp. PL 23, fig. 11 ; pi. 24, 
tigs. 1 to 5. 

The shell is of shorter, wider shape than L. perlonga, taper- 
ing more upwards. Whorls convex, the first 1% sinoothish, 
the rest sculptured with curved ribs stronger than in L. per- 
l&nga, subequally spaced, retractive on the spire, more nearly 
vertical on the last whorl, where there are about 16, standing 
nearly 0.2 mm. apart on the face of the whorl. The last 
whorl has three impressions on the last half : a long, deep and 
wide impression below the middle, the others shorter and 
sharper. The base has a prominent, straight crest (pi. 24, 
fig. 2), divided by the inferior impression, the ribs passing 
over it being small or obsolete. Its posterior end forms a 
rounded hump. The angular and parietal lamellae are longer 
than in L. perlonga. The vertical columellar lamella is 
crescentic, the concavity towards the aperture, and much more 
curved than in L. perlonga. The upper palatal fold emerges, 
and penetrates to the dorsal side. Below its inner end there 
are two callous nodules, as usual (pi. 24, fig. 4). There is a 
low but rather strong lower palatal fold, and a small basal 
fold. The peristome is rather well expanded. 

Length 2.25, diam. 1.3 mm. ; 5y 2 whorls. Kailua. 

Length 2.1, diam. 1.25 mm. Kailua. 

Length 2.2, diam. 1.28, ap. .85 mm.; 5 whorls. Manoa, 
pi. 23, fig. 11. 

Oahu : Kaelepulu, Kailua, in crevices and along the base of a 
low lime-rock bluff about 14 m ^ e from the shore (Pilsbry) ; 
Rocky Hill, Waimanalo and Koko Head (Cooke). It has only 
been found fossil. 

The peculiar basal crest is the most conspicuous external 
feature of this species. Internally, the longer parietal lamella 
and the markedly crescentic columellar lamella distinguish it. 
At Kaelepulu the type locality (pi. 24, figs. 1-3, 5) it occurs 
with L. perlonga, L. micra etc., all being abundant. 

The specimens are all dead, apparently being Holocene 
fossils. They vary in color from vinaceous-cinnamon to cart- 
ridge-buff, or of the former tint with a cartridge-buff median 



266 LYROPUPA. 

zone. The shape varies from cylindric to more tapering. 
Spiral striation is not perceptible in most specimens, but it is 
faintly visible on some. 

18&. Lyropupa ovatula kona P. & C., n. sp. PL 26, figs. 10, 
11, 14, 5. 

The shell is shortly cylindric, the upper third conic, apex 
obtuse ; whorls convex, the last contracted below the periphery. 
Cinnamon-colored. The first 1% whorls are smooth (micro- 
scopically pitted), the rest with coarse sculpture of well-spaced 
ribs, quite oblique, retractive on the spire, nearly vertical on 
the last whorl, where they are usually interrupted or partly 
so in the middle, and about 16 in number. The intercostal 
intervals are nearly smooth, without spiral striae. The last 
whorl has a conspicuous constriction in the middle, stronger 
on its last half; there is an inconspicuous furrow (sometimes 
obsolete) over the position of the lower palatal fold, and a 
short basal furrow. Behind the basal and lower palatal fur- 
rows, and below the main constriction, there is a small in- 
flation or hump. The aperture is wide, with continuous, ex- 
panded peristome. Angular lamella long; parietal lamella 
high, not emerging. The columellar lamella is very deeply 
placed. The upper palatal fold defines a nearly closed sinulus. 
Lower palatal is very deeply placed, a rather strong, oblong 
tubercle. Basal fold, below the preceding, similar but nar- 
rower (or sometimes shorter). Below the inner end of the 
upper palatal there is a barrier formed of two erect tubercles, 
representing the inner ends of the lower palatal and basal 
folds (pi. 26, fig. 10). 

Length 2.3, diam. 1.25 mm. ; 5y 2 whorls (type, Huehue). 

Length 2.1, diam. 1.25 mm. (Huehue). 

Length 2.25, diam. 1.3 mm. (Kapulehu). 

Hawaii: North Kona at Huehue (type loc.) and Kapulehu 
(Gouveia). Molokai: Moomomi (Cooke & Pilsbry). Only 
found fossil. 

The Hawaiian L. cyrta differs by the more regular shape of 
the last whorl. L. kahoolavensis has a continuous lower palatal 
fold, among other differences. L. o. kona is nearly related to 



LYROPUPA. 267 

L. micro, and especially L. ovatula. It is larger than the 
former, with far stronger sculpture. L. ovatula of Oahu is 
very closely related, but the prominent basal crest and hump 
distinguish it. The lower palatal and basal folds are stronger 
in L. o. kona. 

The chief variations noticed among the numerous specimens 
are in shape (see measurements) and strength of the ribs, as 
shown in the figures. 

A single specimen (pi. 26, fig. 5) was among Lyropupae 
taken by the authors on Molokai at Moomomi on the north 
coast, at the end of the escarpment running north from Mauna 
Loa. It measures, length 2.25, diam. 1.4 mm., and appears 
to differ in no respect from those of the Kona slope of Hawaii. 

Probably the species will be found in Maui when fossil 
material from that island comes to hand. 

19. LYROPUPA PLAGIOPTYX P. & C., n. sp. PL 24, figs. 8, 11, 12. 

The shell is ovate, rather solid, the whorls strongly convex, 
the greatest convexity below the suture, with sculpture of 
regular, strong, arcuate ribs, about 15 on the last whorl. The 
last whorl tapers downwards. Its last half is swollen and 
strongly ribbed below the suture, encircled in the middle with 
a rather deep sulcus, crossed by the ribs. Below this is a 
short, sharp lateral sulcus (over the lower palatal fold), and 
on the basal convexity are two very short impressions. The 
ribs diminish in size and are increased in number on ,the basal 
half of the last half whorl. Aperture as usual in the group 
externally. The columellar lamella is strongly crescentic, as 
in L. ovatula. The crest of the upper palatal fold is uneven. 
Below its inner end there is a small but high and short vertical 
tubercle. The lower palatal fold is immersed but visible in a 
front view, short, high and oblique, its outer end sloping 
steeply downward. 

Length 2.15, diam. 1.3 mm. 

Oahu : Kawaihapai, on a steep, wooded hillside about 500 
ft. above the plain and perhaps % mile from the sea (Cooke 
& Pilsbry, 1913). 

This species, found only dead in the surface debris, is re- 



268 LYROPUPA. 

lated to L. ova-tula, but it differs by not having a distinctly 
differentiated basal crest and especially by the internal struc- 
ture of the palatal region. The tubercle below the inner 
end of the upper palatal fold is of different shape, and the 
lower palatal fold is a high and oblique lamina in L. plagioptyx, 
a lower, diffuse tubercle in L. ova-tula-. 

The locality is at the western end of the Waiauae range. 

20. LYROPUPA CYRTA C. & P., n. sp. PL 23, figs. 9, 10. 

The shell is stouter than L. perlonga, the outlines more con- 
vex, the spire tapering more upwards. The whorls are regu- 
larly and strongly convex, the last having only two sulci, and 
these very shallow or even scarcely discernible. The ribs are 
stronger than in perlonga , not flexuous, and number about 15 
on the last whorl. They pass over the narrowly rounded base. 

The lower palatal and basal folds are deeply placed, the 
former a short, rather spreading callous tubercle, the latter 
narrower and longer. Two strong tubercles stand beyond 
their inner ends (pi. 25, fig. 15). 

Length 2.75, diam. 1.45, aperture 1 mm. ; 5i/ 2 whorls. 

Hawaii: fossil at Mana, type loc., and Palihoukapapa (Hen- 
shaw, Thaanum). 

This species resembles L. ovatula, but it is larger with far 
less distinct sulci on the last whorl and no differentiated basal 
crest. 

21. LYROPUPA CUBANA (Ball). PL 25, figs. 13, 16, 17. 

"Shell pinkish brown, paler toward the apex, small, short, 
stout, dextral, five-whorled, apex rather bluntly rounded, sides 
subparallel, base full and rounded; spiral sculpture of fine 
striae, most prominent between the ribs on the last whorl and 
obsolete on the early whorls; they do not overrun the ribs; 
transverse sculpture of ( on the last whorl about twenty-five) 
strong, sigmoid, squarish, narrow, elevated ribs, running from 
suture to suture, and separated by much wider interspaces; 
these ribs are closer and finer toward the apex of the spire, 
they resemble when best developed those of Pupa lyratct- 
Gould ; last whorl subconstricted before the aperture is 



LYROPUPA. 269 

formed; aperture obscurely trilobate, margin well reflected, 
whitish; primary lamellae, except A and B, set rather more 
deeply in the throat than usual. 

Formula ACE according to Dr. Sterki's arrangement. 
The primary A [angular lamella] is stronger and more tri- 
angular at the aperture than the others, but becomes a thin, 
narrow, sharp lamella internally; the secondary 2 [parietal 
lamella] does not come so near the aperture as A [angular 
lamella ] , but further back is a little higher than the latter, and 
the two run parallel like rails on a track, but somewhat irregu- 
larly. From the aperture no lamella can be observed on the 
pillar, but the primary C, [lower palatal fold] which fall* 
short of reaching the margin and is comparatively weak to- 
ward the aperture, in the depths of the throat, is higher and 
stronger, forming a prominent, high, thin lamella. The pri- 
mary E [upper palatal fold] is prominent at the angle of the 
outer lip, and projects toward A [angular lamella] ; deeper 
in it becomes lower and weaker, and still further in it projects 
in a high triangular plate stronger than any of the others at 
that depth. There are no indications of any other lamellae 
or callosities. The margin of the aperture is set somewhat 
obliquely to the axis (see fig. 16) . Max. Ion. of shell 1.75 ; max. 
diam. 1.37 mm. Habitat, Cuba" (DalL). 

Vertigo cubana DALL, Proc. II. S. Nat. Mus. xiii, 1890, pi. 1, 
figs. 1, 2. 

* ' A single specimen of this remarkable little species is in the 
U. S. National Museum, mollusk register 87645. It was re- 
ceived from the collection of the late Dr. Shurtleff, now be- 
longing to Wesleyan University, Middletown, Conn., and was 
obtained by him from the late Prof. C. B. Adams, or at least 
still occupies the tiny glass tube, stopped with sealing-wax, in 
which Professor Adams secured his minutiae, and a tiny label 
on which is written "Pupa .... Cuba" in Professor Adams' 
well-known chirography ' ' (DalL ) . 

There can be little doubt that the locality Cuba was an 
error. Professor Adams received and described land shells 
from the Hawaiian Islands, and as V. cubana differs very little 
from L. thaumasia, a Hawaiian origin appears probably, al- 



270 LYROPUPA. 

though up to this time no other Hawaiian specimens have been- 
turned up. The species was referred to costata Pse. as a 
synonym by the senior author (Proc. A. N. S. Phila. 1900, p.. 
432, footnote 4), upon comparison of the figures with examples 
of L. thaumasia which Pease sent out as costata ; but there are 
some small differences in shape which cause us now to leave 
them apart. 

The type specimen of cubana has lamellae and folds like 
those of thaumasia. In both species the broad, milk-white 
columellar lamella may be seen in an oblique view in the 
aperture. The lower palatal fold is short and rather high. 
The internal barrier is not visible in the mouth, but externally 
the impressions and the light markings due to the plicae are 
exactly as in thaumasia. The specimen measures : length 1.9, 
diam. 1.3 mm., alt. of aperture 0.8 mm. Figs. 16, 17 are 
copies of the original illustrations; fig. 13 a direct face view 
of the same specimen. 

22. LYROPUPA THAUMASIA C. & P., n. sp. PL 24, figs. 13 to 15 ;. 
pi. 25, fig. 14. 

The shell is small, broadly oval, perforate and rimate, tawny, 
solid. Outlines of the spire are strongly convex. The whorls 
are short, convex, the embryonic l 1 /^ very minutely granulate, 
the rest with sculpture of moderately strong, oblique, some- 
what arcuate ribs about half as wide as their intervals, about 
27-30 on the last whorl, which ascends slightly near the aper- 
ture. There are minute spiral striae between the ribs. The 
last half whorl has a rather shallow sulcus (sometimes scarcely 
noticeable) parallel with the suture, and nearer the base a 
shorter broader excavation preceded by an inflation. The base 
is narrow, rounded and straight (as viewed from below), 
passing posteriorly into the baso-dorsal inflation mentioned. 
The aperture is but slightly oblique, squarish, the peristome* 
well expanded, continuous, though adhering to the preceding 
whorl for a short distance above the columella. The angular 
lamella is rather long and straight; its edge curves slightly 
outward, to form a nearly entire circular sinulus with the 
upper palatal plica. The parietal lamella is very long, curved 



LYROPUPA. 271 

at the inner end, its free edge being reflected towards the 
suture. The very deeply placed columellar lamella is not 
visible in a frontal view; it is stout and semicircular. The 
upper palatal fold is long, its free edge having two con- 
cavities where the angular and the parietal lamellae approach 
it ; its inner end curves downward a little, terminated on a very 
low semicircular callous rim which bounds the cavity formed 
by the posterior inflation of the basal crest. Upon this rim 
stand two short, high oblique folds, upper and lower, converg- 
ing towards each other inwardly. The lower palatal fold is 
short and rather high. Externally the palatal folds are visible 
as buff markings, the upper palatal and upper oblique fold 
showing as a decurved band, the lower oblique fold and lower 
palatal as indistinct spots (pi. 25, fig. 14). 

Length 1.95, diam. 1.18, aperture 0.73 mm.; 5% whorls 
(type). 

Length 1.95, diam. 1.13, aperture 0.66 mm. Limahuli, fossil. 

Length 2.02, diam. 1.3 mm. Ekaula. 

Length 2, diam. 1.25 mm. ; 5% whorls. 

Length 1.9, diam. 1.25, aperture 0.6 mm. ; 5% whorls. 

Kauai: Hanakapiai (type loc.) ; Ekaula and Olokele 
(Cooke) ; also fossil at Limahuli (Dole and Cooke). Type 
11061 Bishop Museum ; paratypes 15575 Bishop Museum and 
119449 A. N. S. P. 

This species is well distinguished by its short, compact 
contour, numerous strong ribs, the absence of any trace of a 
third or subbasal impression on the back, and by the strongly 
developed apertural processes. 

L. thaumasm is closely related to L. cubana, and may prove 
to be connected by intermediate forms. On comparison of 
the type specimens there is no difference in color, sculpture 
or convexity of the whorls, but cubana is clearly wider than 
the shortest thaumasia at hand. In thaumasia of the same 
length as cubana, 1.9 mm., there is nearly a half whorl more, 
and the aperture is distinctly smaller. Its length, measured 
vertically is slightly less than one-third that of the shell, while 
in cubana it is decidedly more than that, contained 2^ times 
in length of shell. There is no difference in the apertural 
parts so far as visible in the mouth. 



272 LYROPUPA. 

In L. thaumasm the diameter of the shell is about 58 to 
65 per cent of the length ; in L. cubana it is about 70 per cent. 

The specimens from Bkaula (4) and Olokele (1) are slightly 
larger than the typical form, more globose in outline, with 
considerably weaker palatal plicae and with shallower and less 
distinct sulci on the last whorl. Fossil specimens from Lima- 
huli agree very closely with the typical form. 

Pease distributed this species as his Vertigo costata -, but 
the description of that shell does not apply well to this. 

Undetermined species. 

"VERTIGO COSTATA Pse. Shell cylindrical, oblong, some- 
what solid, dextral, rimate-perforate, longitudinally flexuously 
strongly cost ate, reddish; whorls 4, rotundate-convex, the last 
strongly concentrically bisulcate, compressed at the base, apex 
obtuse ; suture strongly impressed ; aperture bell-shaped, 
rotund at the base, furnished with 4 lamellae, 2 on the parietal 
wall, the first united with the lip, the second median, entering, 
1 on the basal margin, 1 on the lip posteriorly; inner lip 
flexuous ; peristome thin, with the margins not joined. Length 
2.0, diam. 1.0 mm. Hawaii." (Pease.) 

Vertigo costata PEASE, Proc. Zool. Soc. London, 1871, p. 461. 
Pupa costata PPR., Monographia Hel. Viv., viii, p. 399. 
BOETTGER, in von Martens, Conchologische Mittheilungen, i, 
1880, p. 59. ANCEY, Mem. Soc. Zool. France, 1892, v, p. 710. 

This species could not be found in the Pease collection at 
Cambridge, and the junior author, in the course of many years' 
collecting has never seen a specimen meeting all the require- 
ments of the description, which gives characters both of 
Nesopupa and Lyropupa ; the 4 whorls and disjoined margins 
of the aperture would place it in Nesopupa, but the deep 
suture, strong, flexuous costse and the position of the upper 
palatal fold on the outer lip show that costata has certain 
claims to be ranked as a Lyropupa. 

Specimens sent by Pease as costata to the Academy of 
Natural Sciences and others sent to A. D. Brown (now in the 
museum of the Academy) are Lyropupa ihaumasia; but this 
species is far broader than Pease's measurements indicate, the 



PTYCHAL^A. 273 

peristome is continuous, the dorsal impressions are not deep, 
and it could hardly have been described as flexuose forte 
costata. 

It was these specimens from Pease which the senior author 
referred to as Nesopupa (Lyropupa] costata, in Proc. A. N. S. 
Phila., 1900, p. 432. Other authors who have mentioned the 
species knew it only by Pease's description. 

At present therefore V. costata appears to be a lost species 
of uncertain genus ; unless the type comes to light it must be 
dropped as unrecognizable. 

Genus PTYCHAL^A Boettger. 

This volume, pp. 157, 220. 

This group was proposed for a Lower Miocene species of 
Europe ; a recent species of the Benin Islands appears to agree, 
but the writer has not seen the fossil genotype. 

They differ from Nesopupa chiefly by having a strong crest 
behind the lip, as in many Vertigos. This is not usually a 
character of much importance, but it is absent in the long 
series of species referred to Nesopupa. Boettger himself made 
a distinction between Ptychalosa and Ptychochilns (= Neso- 
pupa), both of which he recognized in the European Tertiary. 

We now consider Ptychalaea incorrectly placed in Vertigo. 
We are strongly disposed to subordinate Nesopupa, Costigo 
and Protiesopupa to Ptychalaea as subgenera ; yet specimens of 
the fossil type of the latter are not at hand, and species inter- 
mediate between Ptychalaea, and Nesopupa have not been 
found. At present we admit several genera, Nesopupa, 
Pronesopupa and Costigo, among the Polynesian and Oriental 
Vertigininae as a temporary expedient rather than as an ex- 
pression of definite conviction. It is quite probable that a 
study of the dentition would yield characters of value in 
this inquiry. 

1. PTYCHALAEA DEDECORA (Pils.). P. 158. 

The fossil species P. flexidms Reuss, and possibly ca-pellinn 
Sacco belong here. The other species enumerated under 
Ptychal&a on p. 220 belong to Nesopupa. 



274 NESOPUPA. 

Genus NESOPUPA Pilsbry. 

PtychochHus BOETTGER, Conchol. Mittheil., i, 1881, p. 47- 
type P. tantilla Gld. Not Ptychocheilus Agassiz, Amer. Journ. 
Sci. and Arts xix, 1855, p. 227. Not Ptychochilus, Jordan, 
Bull. U. S. Nat. Mus. no. 10, 1877, p. 58. 

Nesopupa Pils., Proc. Acad. Nat. Sci. Phila. 1900, p. 432. 

The shell is minute, rimate, oval or ovate, the surface usually 
opaque and dull, pitted, striate or ribbed. Aperture with 
angular, parietal and columellar lamellae and usually palatal 
folds; lip expanded. Type N. tantilla (Gld.). 

Animal similar to Vertigo in lacking inferior tentacles. Eye 
tentacles cylindric, rather stout, not swollen distally, the eyes 
central in the ends as seen from above. 

Distribution : islands of the Pacific, Oriental and Ethiopian 
regions, St. Helena. 

Nesopupa is closely related to Ptychalaea (see above) and 
to Lyropupa, also to Costigo, Pronesopupa and CampotemuS) 
which we believe to be groups derived from various Nesopupae. 
The Madeiran Staurod&n* is also closer to Nesopupa than to 
Vertigo. 

As tertiary fossils Nesopupae are known from the Upper 
Oligocene to Pliocene of Europe. N. trigonvstoma (A. Brn.), 
N. Uumi (Bttg.), JV. priscUla (Palad.) (see p. 220) apparently 
belong here, appearing to be most related to the section Indo- 
pupa. Central Europe at that time had numerous Oriental 
genera of land snails. 

Inhabiting widely separated island groups, there have been 
several nearly independent centers of evolution, making the 
construction of a phylogenetic classification exceptionally dif- 
ficult. Thus, the Hawaiian Vertigininae though greatly 
varied, show few points of contact with those of other regions. 
The forms of Polynesia and Micronesia are a homogeneous 
group, but a few closely related species occur in the Moluccas 
and Mauritius. The Oriental region series is closely related 
to the Ethiopian group. 

While the interrelations of Nesopupid groups of restricted 
areas are fairly clear, it is not so when those of different f aunal 



NESOPUPA. 



275 



units are to be combined in one classification. Different com- 
binations of characters, and parallel but not identical stages 
in their evolution are to be considered. Under these condi- 
tions it has been thought most convenient for those concerned 
to treat the species by faunal rather than by taxononic groups. 
The following synopsis of classification is therefore an arrange- 
ment for convenience, in which groups of different faunas in 
a similar stage of evolution but not directly related are some- 
times juxtaposed. 

Synopsis of Sections of Nesopupa. 

a 1 . Angular lamella long, entering rather deeply, anteriorly 
curved, joining the peristome. 

Inner end of the columellar lamella turning downward I. 

Inner end of columellar lamella straight or upturned II. 

a 2 . Angular lamella not entering deeply, straight, joining the 

peristome III.. 

a 3 . Angular lamella very short, nodular or shortly lamellar, 

sometimes not emerging to the peristome IV.. 

I. 

Nesopiipa, typical section. The angular lamella is long, 
strong, curving to join the outer lip near its insertion; the 
inner end of the columellar lamella turns downward ; there are 
at least 5 well developed teeth. The surface is more or less 
pitted-granulose, with spaced cuticular riblets, often deciduous. 

This Section includes all of the Nesopupse of Polynesia and 
Micronesia which we have seen, N. moluccana, N. gonioplax 
of Mauritius, and probably the Melanesian species. 



N. tantiUa (Gld.). Tahiti. 
N. pleurophora (Shutt.). Tahiti. 
N. armata (Pse.). Borabora. 
N. paivae (Crosse). Gambier Is. 
N. dentifera (Pse.). Cook Is. 
N. godeffroyi (Bttg.). Samoa Is. 
N. tongana (Bttg.). Tonga Is. 
N. vitiana (Bttg.). Viti Is. 



N. norfolkensis (Sykes). Norfolk I. 
N. lifouana (Gass.). Lifu. 
N. mariei (Crosse). New Caledonia. 
N. quadrasi (Mlldff.). Guam. 
N. eapensis (Bttg.). Yap, Carolines. 
N. ponapica (Mlldff.) . Ponape, ' ' 
N. moluccana ( Bttg. ) . Amboina. . 
N. gonioplax ( Pils. ) . Mauritius. 



276 NESOPUPA. 

la. Section Nesopi^pilla P. & C. A collateral group of the 
Hawaiian Islands, differing by the sculpture of smooth rib- 
striae without cuticular edges or pitting of the surface, having 
one or two furrows behind the lip, over the palatal folds; 
lamellae as in typical Nesopupa. 

N. plicif era Anc. Oahu. N. baldwini Anc. Maui. 

N. waianaensis C. & P. Oahu. N. b. centralis Anc. Hawaii. 

N. litoralis C. & P. Oahu. N. b. subcostata C. & P. Molokai. 

N. dispersa C. & P. Oahu to Hawaii. N. b. lanaiensis C. & P. Lanai. 

II. 

The angular lamella is long, strong, curving to join the lip 
insertion as in Nesopupa, but the columellar lamella is straight 
or slightly curved upward at the inner end, being less special- 
ized than in Nesopupa. Here are placed several sections not 
directly related. 

Ila. Section Cocopupa P. & C. Surface minutely pitted, 
without riblets. 

N. cocosensis (Dall.). Cocos Island. 

lib. Section Nesodagys C. & P. Surface with spaced ribs 
with cuticular edges and rugose intervals; angular lamella 
strong in front. 

N. wesleyana Anc. Hawaii to Oahu. N. w. rhadina C. & P. Molokai, 

N. w. gouveiae C. & P. Hawaii. Maui, Lanai and Oahu. 

N. w. tryphera C. & P. Kauai to N. thaanumi Anc. Oahu, etc. 
Molokai. 

lie. Section Indopupa P. & C. Similar, but with sculp- 
ture like typical Nesopupa; the angular lamella low in front. 
Oriental Region. 

N. moreleti (A. D. Br.). Borneo, N. filosa (Th. & Stol.). Burma. 

Philippines. N. brevicostis (Bs.). Barrackpore. 

N. moellendorffi. (Bttg.). Philip- N. salemensis (Blf.). Kalryen Hills. 

pines. N. cinghalensis (Crude.). Ceylon. 

lid. Section Afripu-pa P. & C. Sculpture of close rib-striae 
without cuticular edges. The last three species referred to 
this section with doubt. 



NESOPUPA. 277 

N.griqualandica (M.&P.) S. Africa. N. (?) iota (Preston). Brit. E. 
N. farquhari (Pils.). S.Africa. Africa. 

N. bisuleata (Jick.). Abyssinia. N. ( ?) miera (Pils.). Mauritius. 

N. b. rhodesiana (Pils.) . Rhodesia. N. ( ?) nannodes (Q. & M.) . Philip- 
pines. 

He. Section Helenopupa P. & C. Surface smooth. 

N. turtoni (Smith) . St. Helena. 

III. 

Section Insulipupa P. & C. Angular lamella low through- 
out, straight, joining the outer lip; surface as in Nesopupa, 
pitted; form rather cylindric. Oriental and Ethiopian Re- 
gions, Mascarene Islands. The last four species are placed in 
this section with some doubt. 

N. malayana (Issel). Borneo, N. raegalomastoma (Malz.). W. 

Philippines. Africa. 

N. barraekporensis (Oude). Bar- 1ST. (?) com ore nsis (Pils.). Comoros. 

rackpore. N. (?) incerta (G. Nev.). Bourbon. 

N. corrugata (Prest.). Rhodesia. N.( I )ventricosa(H. Ad.). Mauritius. 

N. minutalis (Morel.). Comoros. N. ( ?) eelebensis (T. C.). Celebes. 

IV. 

Angular lamella small, short, or not emerging to the lip 
insertion; inner end of the columellar lamella not turning 
downward. 

IVa. Section Infranesopupa C. & P. Striae without cuti- 
cular edges. Hawaiian Islands. 

N. limatula C. & P. Maui. N. subcentralis C. & P. Hawaii. 

N. dubitabilis C. & P. Molokai. N. bishopi C. & P. Maui. 

N. d. kaalaensis C. & P. Oahu. N. f orbesi C. & P. Hawaii. 

N. anceyana C. & P. Hawaii. N. infrequens C. & P. Kauai. 

IVb. Section Limbatipupa C. & P. Striae or riblets having 
cuticular edges. Hawaiian Islands. 

N. neweombi (Pfr.). Hawaii to N. kauaiensis Anc. Kauai. 

Kauai (numerous varieties). N. singularis C. & P. Oahu, Maui. 

N. oahuensis C. & P. Oahu. N. alloia C. & P. Kauai. 

IV. Section . Delicate lines of growth ; angular lamella 
remote from the lip insertion. 

N. proscripta (Smith) . Christmas Island, near Java. 



278 HAWAIIAN NESOPUPAE. 

HAWAIIAN NESOPUPILLID GROUPS. 

By C. Montague Cooke and H. A. Pilsbry. 

The Hawaiian species of this genus cannot readily be classi- 
fied with those of other regions. The characters elsewhere 
used for defining groups do not apply. They appear to have 
had an independent evolution. A pitted surface, so commonly 
seen in other islands, is not found here. There has been 
mature adaptive radiation; besides the terrestrial forms there 
are many living on the bark of trees, others on foliage ; some 
in relatively dry, others in very humid habitats. 

In arboreal, and especially in folicolous species there is a 
tendency towards degeneration of the teeth, which culminates 
in the species referred to Pronesopupa, which may be geneti- 
cally related to the Limbatipupae. This tendency may be 
traceable to the absence or rarity of predaceous insects, which 
are probably much more numerous on the ground than on 
bark and foliage. 

The authors have seen all of the Hawaiian Nesopupae and 
Pronesopupae except "Vertigo" bacca Pease. The types of 
the species of Ancey and of Cooke & Pilsbry are in the Bishop 
Museum, paratypes or topotypes of all but JV. alloia and 
N. forbesi are in the museum of the Academy. 

Section NesopupiUa P. & C., new section. 

While closely related to the typical Nesopupae by the form 
of the angular and columellar lamellae, these shells differ in 
sculpture and by the sulci of the last whorl. 

They are minute shells with nearly smooth, minutely granu- 
lose embryonic whorls, the third, fourth and last whorls costu- 
late (except in E. Maui and Hawaii specimens of N. baldwmi) ; 
costae without membranous edges, rather closely and evenly 
spaced. There are two nearly parallel sulci on the back of 
the last whorl, separated by a rounded ridge. The angular 
lamella is long, slightly sinuous, extending to the margin of 
the aperture ; palatal folds long, the upper emerging nearly to 
the margin ; columellar lamella rather short, ending abruptly 
in a swelling, its inner termination deflected downwards, al- 



HAWAIIAN NESOPUPAE. 279 

most at a right angle, and extending to the base of the 
columella. Type N. waianaensis. 

They are usually found close to the ground in rather damp 
places on stones, dead leaves, fallen twigs and the like. 

The unidentified Vertigo bacca Pease is like these species in 
having the last whorl concentrically bisulcate, but nothing 
is said of palatal folds. The very brief and incomplete de- 
scription follows. 

"Vertigo bacca Pse. T. cylindracea, abbreviata, tenuius- 
cula, dextrorsa, perforata, laevigata; apex obtusus; anfr. 4, 
rotundati, ultimus concentrice bisulcatus; apertura fere cir- 
cularis, in pariete aperturali bilamellata, columella subuniden- 
tata ; labrum vix eversum ; pallide fusca. 

"Hab. Kalaparia, insulae Hawaii" (Pease, Proc. Zool. Soc. 
London, 1871, p. 462). 

No dimensions were given. Subsequent authors have added 
nothing to the above account. Pease further remarks: "The 
above description was drawn up several years since from speci- 
mens collected at Kalapana, district of Puna, Island of Hawaii ; 
as they have been lost, I furnished the precise locality to en- 
able collectors to recover the type." 

Key to the Species of Nesopupilla. 

a 1 . Angular lamella extending to outer margin of peristome or 
united to the outer lip-insertion by a thick callus. 

6 1 . Angular lamella terminating on the parietal wall, 
united to the lip insertion by a thick callus; only the 
lower sulcus on the back distinctly impressed. 

N. plicifera, no. 1. 

fr 2 . Angular lamella united to the outer margin of the 

peristome ; two furrows behind the lip clearly defined. 

c 1 . Lower palatal fold accompanied by a short fold 

near its outer end. N. waianaensis, no. 2. 

c 2 . Lower palatal fold not accompanied by a short 

lamella : Spire tapering near the summit. 

d l . Whorls weakly convex; length of shell 

more than 1.7 mm. N. litoralis, no. 3. 



280 HAWAIIAN NESOPUPAE. 

d 2 . Whorls convex; length of shell less than 
1.65 mm. N. dispersa, no. 4. 

a 2 . Angular lamella terminating on the parietal margin near 
the insertion of the outer lip, not united to the lip insertion 
by a thick callus. N. baldwwi, no. 5. 

1. NESOPUPA PLICIPEBA Ancey. PL 27, figs. 1, 2, 3. 

" Shell small, ovate, dark reddish brown, somewhat thin, 
slightly shining, except for the apex obliquely and closely pli- 
cate, plicae smooth, distinctly perforate. Spire obtuse, ovate 
or cylindrically ovate. Whorls 5, noticeably slightly convex, 
suture impressed and simple, the last whorl tapering at the 
base, concentrically sulcately impressed, with the sulcus cor- 
responding to the lower palatal lamella, slightly compressed 
about the umbilical crevice. Aperture hardly oblique, reced- 
ing slightly at the base, ovately truncate, furnished with 
lamellae, as follows : two parietal, of which the upper angular 
extends to the upper margin, and is connected with the outer 
margin; strongly elongated; the lower [parietal lamella] is 
large, a little deeper seated, extending slightly spirally. One 
small, acute, tooth-like, columellar. Finally, two elongated 
palatals, parallel, the upper almost reaching the margin. 
Peristome brown, thickened, narrowly expanded, with distant 
margins, united by an inconspicuous callus, the upper sub- 
angularly produced outwardly. Length 1.5, diam. hardly 1, 
aperture 0.5 mm. long. Oahu, Nuuanu (Thaanum) " (Ancey). 

Nesopupa plicifera Anc. Proc. Malac. Soc. London, vi, 1904, 
p. 122, pi. vii, f. 14. Type 18703, paratypes 18740 Bishop 
Museum. 

Ancey ? s material consists of the type and 4 paratypes. The 
angular lamella is slightly curved and is united with the 
junction of the outer lip by a thick callus, forming, with the 
upper palatal fold, an almost circular sinulus. In immature 
specimens of 4% whorls, from the type locality on the slopes 
of upper Nuuanu, just south of the Pali, the embryonic 
whorls are almost smooth, under a high-power lens very 
minutely granulose for the first 1% whorls, then, almost ab- 
ruptly rather strong, sharp, oblique costae set in. In adult 



HAWAIIAN NESOPUPAE. 281 

specimens carefully compared with the type but with better 
preserved surfaces, the third and fourth whorls are distinctly 
eostate ; on the last whorl the costae are at first well developed 
near the suture, becoming weaker below; the interstices are 
very minutely striate with lines of growth. The base is com- 
pressed and prominent. There is a narrow, rather deep con- 
centric sulcus on the back of the last whorl, just above the 
lower palatal fold, but no twisted sulcus just in back and 
parallel with the peristome, and merely a flattening over the 
upper palatal fold. In the holotype the costag are about .055 
mm. apart on penultimate whorl and the shell has the follow- 
ing measurements: length 1.53, diam. 0.95, greatest length of 
apert. 0.61 mm. There are about 5% whorls and the outer 
margin of the aperture is not flattened. This is not the case 
in most of the adults examined ; for nearly all the fully adult 
specimens have 5% whorls, and the outer margin of the aper- 
ture is slightly flattened. 

The typical form is distributed over the whole Koolau 
Mountain range of Oahu. It is exceedingly abundant, if one 
has the patience to look for these minute forms, and can 
nearly always be found on the dead leaves of ieie (Freycinetia) 
wherever this plant occurs. It is occasionally found on other 
dead leaves, but rarely on stones. In one or two isolated cases 
the junior author has found typical specimens under moss on 
trunks of living trees. A single example of the typical form 
from the Waianae Mts. is in the Bishop Museum collection. 

Occurring with typical examples are a few slightly larger 
specimens from upper Kaliuwaa and Kaipapau, Oahu. They 
differ only in that the outline is slightly more cylindrical and 
the shells are slightly longer. One with 5% whorls measures : 
Length 1.71, diam. 0.92, apert. 0.6 mm. in greatest length. 

2. NESOPUPA WAIANAENSIS C. & P., n. sp. PL 27, figs. 4, 5, 6. 

The shell is minute, perforate (umbilicus minute, deep, semi- 
circular in outline), ovately conic, snuff-brown, distinctly cos- 
tulate except on the embryonic whorls, shining, quite thin, 
somewhat translucent. Spire with convex outlines, and 
rounded apex ; individual whorls convex, separated by a rather 



282 HAWAIIAN NESOPUPAE. 

deep suture. Whorls 5, the first rather large, nearly smooth, 
under a high-power lens very minutely wrinkled ; the second 
slightly narrower than the first, becoming costulate rather 
abruptly near its termination ; the rest of the whorls increasing 
slowly and evenly, regularly and strongly costulate; riblets 
strongly oblique, with rounded edges, on the penultimate whorl 
about 0.055 and on the last whorl about 0.14 mm. apart (or 
partly obsolete) ; the interstices microscopically wrinkled, the 
wrinkles short; last whorl slightly broader than the penult, 
tapering to the base, scarcely ascending near the aperture, 
bisulcate, the sulci deep, corresponding to the palatal plicae, 
the lower strongly curved, deep, broad, remote from the peris- 
tome, the upper arched, not quite so deep, nearly reaching 
the peristome, the two separated by a convex ridge. Aperture 
irregularly truncate-ovate, small, slightly oblique, contracted, 
its outer margin slightly sinuous, its lip insertions converging, 
nearly united by a thick callus. Angular lamella long, slightly 
curved, slender, not very high, extending to the margin of 
the outer lip just before its insertion. The parietal lamella 
is deeper seated, stronger and slopes outward. The columellar 
lamella is strong, at first horizontally entering, then descend- 
ing around the columella. The upper palatal fold is long, 
lamella-like, nearly reaching the peristome, with a depression 
just in back of its outer end. The lower palatal fold is very 
deeply seated, with a dentiform nodule close to and below its 
outer end. The outer margin of the peristome is slightly thick- 
ened within, scarcely expanded, the columellar margin 
broadly expanded. 

Length 1.41, diam. 0.8, diag. length of aperture 0.5 mm. 
Cotype. 

Length 1.41, diam. 0.85, diag. length of aperture 0.52 mm. 
Cotype. 

Oahu: Waianae Mts. at Pukaloa, in the open valley under 
stones near the "Hunter's Cabin/' type loc. : Lualualei, near 
the head of the valley (Cooke) ; Kawaihapai at the western 
end of the range, fossil in soil of the plain and slopes of the 
foothills (Cooke & Pilsbry). 

This species is closely related to N. plicifera. The angular 



HAWAIIAN NESOPUPAE. 283 

lamella is proportionally slightly weaker than that of 
JV T . pUcifera and is attached to the outer lip rather than the 
parietal wall ; the lower palatal fold is accompanied by a denti- 
form, nodule which does not appear to be developed in any 
specimens of pUcifera; the lip insertions converge more and 
are nearly united by a thicker callus. Externally it has two 
strong furrows on the last whorl. 

3. NESOPUPA LITOBALIS C. & P., n. sp. PI. 28, fig. 1. 

The shell is minute, perforate and shortly rimate, cylindrical, 
the upper third convexly conic, whitish in its fossil state, the 
last four whorls obliquely costate. Spire nearly cylindrical 
with slightly convex outlines, tapering above to a rather blunt 
apex, individual whorls only weakly convex, separated by an 
impressed suture. Whorls 5% to 5%, the first and second of 
nearly equal width, the rest increasing very slowly and regu- 
larly ; the embryonic shell, under a strong lens, almost smooth 
for about 11/2 whorls, the costae at first rather weak, blunt, the 
rest of the whorls evenly costate, costae oblique, with blunt 
edges, about 0.07 mm. apart on the penult and 0.11 apart on 
the last whorl, the interstices nearly smooth. Last whorl 
nearly cylindrical, tapering gradually towards the base, as- 
cending very slightly to the aperture, bisulcate on the back, 
the sulci corresponding to the palatal plicae, the lower sulcus 
shallow, broad, strongly arched, terminating at a slight dis- 
tance from the peristome, the upper, shallower, wide, nearly 
parallel almost reaching the peristome, the two separated by 
a rounded ridge. Aperture truncate-ovate, rather small, con- 
tracted within, its outer margin very slightly sinuous, slightly 
oblique, perpendicular, the lip-insertions united by a thick 
callus. Angular lamella strong, long, slightly diverging from 
the parietal, and nearly equal to the latter in height, extend- 
ing nearly to the margin of the peristome and united with it 
just outside of the lip insertion. Parietal lamella strong, quite 
oblique, deeply penetrating. Columellar lamella rather strong, 
at first transverse to the axis, then abruptly turning down- 
ward at right-angles. The upper palatal fold is long, strong, 
extending to the inner margin of the peristome, with a de- 



284 HAWAIIAN NESOPUPAE. 

pression just back of its prominent outer end. The lower 
palatal is deeply immersed, higher within, slightly sinuous. 
The outer and basal margins of the peristome are uniformly, 
slightly thickened within, not expanded ; the columellar mar- 
gin is broadly expanded. 

Length 1.83, diam. 1 ; greatest length apert. 0.61. Cotype, 
B. M. 

Length 1.75, diam. 1.03 ; greatest length apert 0.63. Cotype, 
A. N. S. 

Oahu: Ewa, type loc., very abundant under stones on the 
coral plain below "Sisal" (Cooke) ; Diamond Head (Cooke 
& Pilsbry) : found only fossil. Cotypes 11065, Bishop Museum, 
and 44694 A. N. S. P. 

The typical form of this species is easily distinguished from 
N. waicmaensis by its larger size, more cylindrical form and 
less convex whorls. Prom N. plicifera it is recognized by its 
more tapering summit, but especially by the thickened parietal 
callus, and the angular lamella terminating on the outer lip 
rather than on the parietal callus, though it is very near the 
junction of the two. 

4. NESOPUPA DISPERSA C. & P., n. sp. PL 27, figs. 7, 8. 

The shell is minute, perforate (umbilicus, extremely minute, 
open, deep) sub-cylindrical, whitish or brownish in its fossil 
state ; the last three whorls distinctly costulate ; opaque. 
Spire sub-cylindrical, extended, with slightly convex tapering 
outlines, rather obtuse at the summit, individual whorls moder- 
ately convex, separated by a rather shallow suture. Whorls 
5%, the embryonic large, the second slightly wider than the 
third near its initial stage, nearly smooth, under a strong lens 
very minutely punctate, the riblets beginning rather abruptly 
near the end of the second whorl ; rest of the whorls increasing 
slowly and regularly, distinctly finely and evenly costate, the 
costae slightly oblique, about 0.07 apart on the penult and 0.09 
on the last Avhorl, with the interstices minutely granulose ; last 
whorl slightly broader than the penult, subcylindric, tapering 
gradually toward the base, its last third straight, hardly as- 
cending at the aperture, bisulcate behind the aperture, the 



HAWAIIAN NESOPUPAE. 285 

sulci corresponding to the palatal plicae ; the lower sulcus is 
strongly arched, shallow, broad, terminating at a little distance 
back of the aperture, the upper longer, shallower, not as 
strongly curved, extending nearly to the peristome. Aperture 
truncate-ovate, scarcely oblique, its outer margin slightly 
sinuous, lip insertions converging somewhat, sometimes united 
by a thin callus, thickened at and near the lip insertion. 
Angular lamella long, high, of nearly uniform height, nearly 
parallel to the parietal, extending to the margin of the outer 
lip ; parietal lamella more deeply seated, strong, long and 
somewhat oblique. Columellar lamella strong, deeply seated, 
at first transverse to the columella, then abruptly turned down- 
wards almost at a right angle and extending to the base of 
the columella. Upper palatal fold strong, long, with a slight 
depression back of its outer end, its margin nearly touching 
that of the angular lamella; extending nearly to the margin 
of the peristome. The lower palatal fold is deeply immersed, 
long, strong within, tapering gradually to its anterior end. 
Peristome with the outer and basal margins uniformly thick- 
ened within, the columellar margin narrowly, triangularly 
arched above the umbilicus. 

Length 1.53, diam. 0.86, greatest length of apert. 0.5 mm. 

Oahu: Makua, type loc. (Forbes & Cooke), and Lualualei 
(Cooke), in the Waianae Mts. ; Kailua (Pilsbry, Cooke), 
Waimanalo, Rocky Hill, Manoa, Kahala, Malaekahana, Kahuku 
and Kawailoa (Cooke) in the Koolau Mts. 

Molokai : Kalaeokailio Pt., Mauna Loa, Shifting Sands and 
Moomomi (Cooke & Pilsbry). 

Lanai : Mahana Gulch a single living specimen, and Western 
End (Forbes). 

Kahoolawe: Kanapou, (Stokes, Forbes) and Schooner Bay 
(Pilsbry and Cooke). 

East Maui: Kanaio and Auwahi (Fleming). 

Hawaii: Puuwaawaa (Thurston), Kona (Thwing, Gouveia), 
Huehue, Kapulehu and Waiaha (Gouveia). All specimens 
fossil from more or less recent beds, except the single fresh 
specimen from Mahana Gulch, Lanai. Type 11066 Bishop 
Museum, para types 42942 Bishop Museum and 44707 A. N. S. P. 



286 HAWAIIAN NESOPUPAE. 

A very variable and widely distributed little species. Speci- 
mens from different colonies and islands vary considerably 
from the typical form described above. The most important 
races may be briefly described as follows : 

a. The shells from Rocky Hill and Manoa, Oahu, have 
slightly blunter apices than the typical lot. 

b. Those from Kahuku and Kailua are slightly smaller but 
agree in other characters. 

c. Specimens from Kalaeokailio Pt., Molokai, are slightly 
longer and narrower in proportion to their length. 

d. Lanaian and East Mauian specimens are almost identical 
with the type lot. 

e. Specimens from Kahoolawe have more convex whorls and 
the outlines of the spire are also more convex. 

/. The shells from Hawaii are slightly smaller, cylindrical 
in outline and with fewer and closer costee. 

Shells from some of the different deposits have the follow- 
ing measurements : 

a. Length 1.53, diam. 0.86, greatest length apert. 0.55 mm. 
Rocky Hill, Oahu. 

a. Length 1.6, diam. 0.9 mm. Diamond Head, Oahu. 

&. Length 1.35, diam. 0.8, greatest length apert. 0.5 mm. ^ 
Kahuku, Oahu. 

&. Length 1.38, diam. 0.8, greatest length apert. 0.5 
Kailua, Oahu. 

c. Length 1.6, diam. 0.83, greatest length apert. 0.55 mm. ; 
Kalaeokailio Pt., Molokai. 

c. Length 1.55, diam. 0.9 mm. Moomomi, Molokai. 

d. Length 1.5, diam. 0.86 ; greatest length apert. 0.52 mm. 
Western End Lanai. 

d. Length 1.45, diam. 0.82, greatest length apert. 0.55 mm. ; 
Kanaia, E. Maui. 

e. Length 1.53, diam. 0.86, greatest length apert. 0.55 mm. ; 
Kanapou, Kahoolawe. 

/. Length 1.43, diam. 0.75, greatest length apert. 0.52 mm. - r 
Puuwaawaa, Hawaii. 



HAWAIIAN NESOPUPAE. 287 

5. NESOPUPA BALDWINI Ancey. PI. 27, figs. 9, 10. 

" Shell shortly cylindrical-oblong, openly yet minutely per- 
forate, brownish red, with a silky sheen, almost destitute of 
sculpture except for very thin lines of growth hardly visible 
under a strong lens. Spire obtuse. Whorls 5y 2 , convex, the 
first three increasing rapidly in diameter, the rest nearly equal, 
with an impressed suture. The last oblong, scarcely swollen, 
slightly attenuate at the base, contracted back of the aperture 
by two shallow sulci parallel to the suture ; slightly ascending 
in front. Aperture vertical, outwardly dilated angulate above 
the middle, truncate ovate, armed with plicse or lamellae as 
follows : parietals 2, of which one, the angular, is elongate, but 
shorter than the next ; the other, nearly median, more deeply 
seated, long; one columellar, acute, dentiform; and two elon- 
gate palatals, corresponding to the exterior sulci, parallel, of 
which the upper almost reaches the margin. Peristome nar- 
row, unicolorous, brown, slightly thickened, except at the colu- 
mella, slightly expanded, with quite distant margins, the ex- 
terior of which is slightly sinuous above. Length 1.5, diam. 
0.8, alt. of apert. about 0.5 mm. Molokai (Baldwin) ; Kaupa- 
kulua, Maui (Baldwin.) (Ancey.) 

E. Maui; Kaupakalua (Baldwin), Kailua (Cooke), Mahiku 
(Forbes). W. Maui: Hahakea, Waihee (Cooke), Honokahau 
(Forbes). Molokai: abundant on the wooded eastern part 
(Thaanum, Forbes, Pilsbry, Cooke). Lanai: "top of Moun- 
tain" (Forbes). Hawaii: Keehia (Thaanum), Laaloa 
(Gouveia). Holotype 18698 Bishop Museum. 

Nesopupa Baldwini ANCEY, Proc. Malac. Soc. London, vi, 
1904, p. 122, pi. 7, fig. 13, with var. centralis Ancey. 

This species and N. plieifera, are certainly very closely re- 
lated. In baldwini the shell is darker colored, with slightly 
more convex outlines, the surface is smoother, the costulse 
much weaker, closer and finer ; the lip insertions are more re- 
mote, the angular lamella not being united with the outer lip 
by a thick callus. The columellar lamella is transverse to 
the columella, ending within in a rather prominent knob. 

In the typical form from Kaupakalua, Maui, the rib sculp- 



288 HAWAIIAN NESOPUPAE. 

ture of the group is almost completely wanting. The furrows 
behind the lip are only weakly impressed, the Upper one 
wholly lateral, the lower extending to the dorsal line. Both 
angular and parietal lamellae are long, the former reaching 
the thickened edge of the parietal callus near the insertion of 
the lip, the parietal lamella somewhat immersed, regularly 
curved in a basal view. The deeply placed columellar lamella 
is much thickened at the inner end, but is hardly bent down- 
ward. Lower palatal fold is deeply immersed, as usual. The 
upper palatal does not have a depression preceding the outer 
end, such as is seen in the Oahuan species. Ancey 's type has 
the following dimensions: length 1.6, diam. 0.98, greatest 
length of aperture 0.67 mm. These measurements are slightly 
greater than Ancey gave, but the specimen was labelled type 
in his handwriting. A paratype measures: length 1.65, 
diam. 1 mm. 

Specimens from West Maui are more like Molokai shells 
than those of East Maui. 

Var. centralis Ancey. A little smaller and less cylindric, 
whorls 5, otherwise scarcely differing from the type (Ancey). 

Hawaii: Olaa (Thaanum). 

Very close to the typical Mauian form. In an Olaa speci- 
men from Thaanum (no. 79991 A. N. S. P.) the surface 
shows no ribs, but there are some striae on the last fourth of 
a whorl. The upper palatal fold is indicated externally by a 
slight impression, the lower by a shallow, broad concavity, both 
being quite short for this species. It measures, length 1.65, 
diam. 1.05 mm. 

Specimens from Keehia, Hawaii, collected by Mr. Thaanum 
are almost exactly similar to Ancey 's type specimen. They 
are less distinctly costulate than those of Lanai, West Maui 
and Molokai. 

5&. Nesopupa baldwini subcostata P. & C., 11. subsp. PL 27, 
figs. 11, 12. 

The surface is distinctly and closely costulate, the riblets 
not very strong, sometimes irregularly developed. Sulcus 
over the lower palatal fold well marked, the upper one shallow 



HAWAIIAN NESOPUPAE. 289 

or nearly obsolete. Angular and parietal lamellae about as 
long as in the typical form, nearly straight, the parietal emerg- 
ing further than typical. 

Length 1.6, diam. 1 mm. 

Length 1.48, diam. 0.95 mm. 

Widely distributed on the wooded part of Molokai, the typ3 
from upper Kaunakakai, along the pipe-line trail. It occurs 
as far west as Puunea, north of Meyer's lake. 

5Z>. Nesopupa bald-mini lanaiensis P. & C., n. subsp. PL 27, 
figs. 13, 14, 15. 

The surface has weak ribs, or largely obsolete, last whorl 
with both sulci d-eep, surface strongly convex between them; 
base swollen. Angular and parietal lamellae somewhat longer 
than in typical baldwini. Columellar lamella with the inner 
knob somewhat turned down. 

Length 1.5, diam. 0.9 mm. ; 5% whorls. 

Lanai. 

Section Infranexopupa C. & P., new section. 

The shells are minute, dextral (except for N. infrequens) ; 
angle lamella short, not reaching the margin of the aperture, 
parallel to the parietal ; upper palatal plica much shorter than 
the lower, neither of which is accompanied by a sulcus on the 
back of the last whorl ; with a slight swelling just back of and 
parallel to the margin of the aperture ; columellar lamella 
obliquely seated, not descending, when viewed from above 
slightly sigmoid. Type N. limatula. 

They are usually found on the fronds of ferns or leaves of 
low plants, but occasionally on the trunks of trees. 

Key to Species of Section Infranesopupa. 

a\ Shell sinistral. N. infrequens, no. 12. 

a 2 . Shell dextral. 

& 1 . Adult shells nearly 2.1 mm. in length. 

c 1 . Surface, between striae, covered with minute 
anastomosing wrinkles. Maui. 

N. bishopi, no. 10. 



290 HAWAIIAN NESOPUPAE. 

c 2 . Surface, between striae, minutely granulose. 
Hawaii. N. forbesi, no. 11. 

b 2 . Adult shells less than 1.9 mm. in length. 

c 1 . Aperture of , adult shells always furnished with 
a minute basal fold. N. d. kaalaensis, no. la. 
c 2 . Basal fold usually absent, only present in very 
rare individual instances. 

d l . Adult shells minute, less than 1.4 mm. in 
length. Molokai. N. dubitabttis, no. 7. 
d 2 . Adult shells more than 1.5 mm. in length. 
Maui, Hawaii. 

e 1 . Palatal plicae very short. Hawaii. 
/*. Shells broadly ovate to glo- 
bosely ovate, parietal and 
angular lamellae well separ- 
ated. N. anc-eyana, no. 8. 
/ 2 . Shells nearly cylindric, angu- 
lar lamella usually dentiform, 
placed close to the parietal. 

N. subcentralis, no. 9. 
e 2 . Palatal plicae longer. Maui. 

N. limatula, no. 6.. 

6. NESOPUPA LIMATULA C. & P., n. sp. PI. 28, figs. 6, 10. 

The shell is minute, sub-perforate (perforation very minute, 
almost closed), ovate, the last two whorls hazel, apical whorls 
much paler ; rather thin ; translucent with a silky sheen, micro- 
sopically striate. Spire with convex outlines and a rather ob- 
tuse apex; individual whorls convex, separated by a rather 
deep suture. Whorls 5 to 5^4, the embryonic 1% whorls con- 
vex, almost smooth, under the microscope very minutely punc- 
tate, viewed with oblique light the points arranged in spiral 
lines, glossy; the remaining whorls faintly striate, striae 
slightly flexuous, very low, evenly spaced, the interstices and 
striae covered with a fine network of minute wrinkles only seen 
under the microscope. The last whorl is rotund at the back,, 
ascending slightly near the aperture, a little contracted behind 



HAWAIIAN NESOPUPAE. 291 

the lip, the contraction preceded by a low, wide swelling 
parallel to the margin of the aperture, near and at the base. 
Aperture truncate-ovate, scarcely oblique, nearly perpendi- 
cular, the outer margin slightly rounded, the columellar mar- 
gin nearly straight, margins united by a very thin minutely 
punctate parietal callus. Angular lamella short, upright, not 
extending to the margin of the aperture. Parietal lamella 
long, strong, bent obliquely outwards. Columellar lamella 
strong, slanting downwards, deeply seated, entering almost 
horizontally in front, its inner end ascending; the upper and 
lower palatal plicae rather long, slightly converging backwards, 
extending to a little distance within the margin of the aper- 
ture; the lower one quite stout. Peristome thin, expanded 
at the columella and base, the upper outer portion erect. 

Length 1.65, diarn. 1.1, aperture (diag.) 0.65 mm. 

E. Maui : Ainahou and Palikou in Haleakala Crater (Cooke) 
and Ukulele (Forbes, Cooke). All specimens taken on the 
fronds of living ferns. W. Maui: Maunahooma (Forbes and 
Cooke) Honokaoo (Forbes). Type 11067 Bishop Museum,, 
paratypes 38489 Bishop Museum and 44692 A. N. S. P. 

Apparently a widely dispersed species on East and West 
Maui. All the specimens from E. Maui, were found at over- 
6,000 ft. elevation; those from W. Maui 2,000 ft. elevation.. 
The single specimen collected by Mr. Forbes at Ukulele has a 
rather short, lamella-like fold between the upper and lower 
palatal plicae, otherwise it agrees perfectly with the typical 
specimens from Ainahou. A second specimen collected by 
Cooke from near the same spot is longer, slightly narrower 
with more convex outlines, lighter and uniformly colored and 
with short palatal plicae. This specimen measures : Length 1.9, 
diameter 1.07, aperture (diag.) 0.75 mm. 5!/4 whorls. West 
Maui specimens are slightly smaller, more compact and the 
apertural lamellae are not only more strongly developed but 
are also longer. A specimen from Maunahooma measures L. 
Length 1.55, diam. 1.1 mm. ; apert., greatest length 0.67 mm. 

7. NESOPUPA DUBITABILIS C. & P., n. sp. PI. 28, fig. 9. 

This shell is smaller than N. limatula, more shortly ovate- 



292 HAWAIIAN NESOPUPAE. 

in outline. Fully adult specimens have but 4% convex whorls. 
The surface of the shell is more closely, finer and more evenly 
striate. Length 1.38, diam. 0.95 ; apert. greatest length 0.55 mm. 

Molokai: Poholua (type loc.), 2,500 ft. elevation; Kamoku 
and Kawela (Cooke) ; Ualapue (Forbes) ; Western ravine, 
head of Kamalo (Cooke & Pilsbry). Type 11068, Bishop 
Museum, paratypes 35150 Bishop Museum and 44728 A.N.S.P. 

This species is undoubtedly closely related to N. limatula 
from which it is here considered as a separate species not only 
by its geographical distribution but by its much smaller size, 
form, etc. 

In an immature specimen with 3*4 whorls the shell is openly 
perforate, lighter colored than in the adults. The embryonic 
whorls, under the microscope, are very minutely and closely 
granulose, the granules do not appear to be arranged in any 
particular order. The delicate, slightly oblique striffi appear 
rather abruptly at about l^ whorls. 

Specimens from the western ravine at the head of Kamalo 
are somewhat larger : 

Length 1.55, diam. 1.05 mm. 

Length 1.5, diam. 1 mm. 

la. Nesopupa dubitabilis kaalaensis C. & P., n. subsp. PL 28, 
fig. 13. 

Specimens from Oahu are very similar to N. dubitabilis in 
size and form. They differ, however, in that adult specimens 
are always furnished with a minute basal fold slightly sub- 
columellar in position. This fold has not been observed in 
specimens from Molokai. It is somewhat immersed, low, short, 
lamella-like rather than nodular. 

Two gerontic specimens show considerable variation in their 
lamella character. In the first there is a rather long, low swell- 
ing between the two palatal plicae, and the parietal lamella is 
continued nearly to the margin of the aperture, ending in a 
low broad nodule. In the second there is a minute nodular 
infraparietal lamella. 

An average adult measures: Length 1.45, diameter 0.97; 
aperture, greatest length 0.92 mm. 



HAWAIIAN NESOPUPAE. 293 

Oahu: Waianae Range at Mokuleia (Cooke), and Kaala, 
eastern spur, about 2,500 ft. elevation under bark of lehua 
trees (Bridwell). Type 11069 Bishop Museum; para types 
42720 Bishop Museum and 44709 A. N. S. P. 

8. NESOPUPA ANCEYANA C. & P., n. sp. PL 28, figs. 2, 3. 

The shell is somewhat like that of N. limatula but longer, 
broader and usually with but 4% 5 whorls. Shell compact, 
broadly ovate to globosely ovate, antique brown to buckthorn 
brown, the lower whorls indistinctly, minutely striate, the 
strife and their interstices covered with microscopic, irregular 
hair-like wrinkles. Spire with convex outlines; whorls con- 
vex, separated by a rather deep suture. The embryonic 
whorls are microscopically punctate with a few indistinct 
spiral lines just above the suture; last whorl slightly swollen 
just back of the aperture and scarcely flattened over the palatal 
plicae. Aperture irregularly triangular, with broadly rounded 
angles, lip-insertions scarcely converging. Angular lamella 
short, lamella-like, rather widely separated from the parietal. 
Parietal lamella long, strong, and nearly perpendicular to the 
parietal wall; columellar lamella very short, deeply seated, 
horizontally entering, and slightly slanting downwards notice- 
ably turned upward at the inner end, and in old specimens 
continued up the columella as a low callous ridge. The two 
palatal plicae are very short, not approaching the peristome, 
nearly parallel and rather widely separated. 

Length 1.78, diam 1.15; apert,, greatest length 0.73 mm.; 
5 whorls. 

Length 1.53, diam. 1.1 ; apert., greatest length 0.7 mm. ; 
41^ whorls. 

Hawaii: Olaa (Thaanum, Ancey coll.; Lyman) ; Kilauea 
(Cooke), Piihonua and Humuula (type loe. ; Forbes). Type 
11072 Bishop Museum; para type 39300, Bishop Museum and 
44723 A. N. S. P. 

This species is most easily distinguished from N. limatula 
of Maui by the much shorter palatal folds; from N. sub- 
centralis by its broader and more ovate outlines, smoother 
surface, its shorter and broader aperture ; the rather distantly 



294 HAWAIIAN NESOPUPAE. 

spaced parietal and angular lamellae, the angular being usually 
slightly longer and stronger than in centralis, and also by the 
wider spaced and more nearly parallel palatal plicae. 

A single specimen in the Ancey collection from Olaa is 
almost identical with Humuula specimens. 

Near the edge of the crater of Kilauea specimens are very 
abundant on the low growing kukaineenee (Coprosma). They 
are smaller than the typical form, globosely ovate, and have 
but 414 whorls. One of these specimens measures: Length 
1.53, diam. 1.04, apart. 0.67 mm. The above measurements are 
of an average specimen. A number of slightly smaller speci- 
mens occur in most of the lots collected near Kilauea. At first 
glance they have the appearance of immature shells and are 
almost globular in outline. The peristome, however, is thick- 
ened. In these specimens the angular lamella is very short. 
One of them with 4% whorls measures : length 1.25, diam. 1.05, 
apert. 0.61 mm. 

In some specimens from the margin of Kilauea iki (fig. 3), 
the angular and parietal lamellae are quite close together, and 
the palatal folds are less widely spaced than in typical 
anceyanoi. 

Length 1.7, diam. 1.1 mm., aperture 0.66 mm. 

Length 1.5, diam. 1.1 mm. 

9. NESOPUPA SUBCENTRALIS C. & P., n. sp. PL 28, fig. 8. 

The shell is minute, sub-perforate (perforation very minute)', 
nearly cylindrical, in its fossil state pale brownish white, often 
with a broad lighter peripheral zone, rather thin, hardly 
translucent, under a strong lens closely, quite evenly but 
weakly obliquely striate. Spire with convex outlines. The 
summit convexly conic, apex obtuse ; individual whorls slightly 
convex and separated by a rather shallow suture. Whorls 5, 
the embryonic smooth for iy 2 whorls, the faint striae beginning 
rather abruptly at this point ; last whorl subcylindric, taper- 
ing towards the base, somewhat flattened over the palatal plicae, 
very slightly swollen back of outer margin of peristome. 
Aperture broadly truncate-ovate, hardly oblique, perpendicu- 
lar, lip-insertions remote. Angular lamella short, low, tooth- 



HAWAIIAN NESOPUPAE. 295 

like, deeply seated. Parietal lamella well developed, strong, 
perpendicular to the parietal wall. Columellar lamella short, 
horizontally entering, slanting downwards, inwardly, ending 
abruptly. Two palatal plicae rather short, lamella-like, nearly 
parallel and rather remote from the peristome, the lower one 
more immersed. Peristome slightly thickened within, ex- 
panded slightly except for the upper outer margin which is 
thin and erect. Length 1.7, diam. 0.98 ; apert., greatest length 
0.7 mm. 

Hawaii: Palihoukapapa (type loc.), very abundant in the 
fossil deposits (Henshaw, Thaanum) ; Mana (Thaanum), 
rather rare. Living specimens on the Flow of 1823, 7,000 ft. 
elevation (Forbes); Waihou; S. Kona, on ferns (Gouveia). 
Type 18699 Bishop Museum ; topotypes 36654, Bishop Museum ; 
44699 A. N. S. P. and in the Thaanum coll. 

Nesopupa baldwini var. centralis Anc., HENSHAW, Journ. of 
Malacology, xi, September 1904, p. 63. ANCEY, same Journal 
Sept., 1904, p. 67 ; not N. b. centralis AN GEY, Proc. Malac. Soc. 
Lond. vi, June, 1904, p. 122. 

This species is closely related to N. anceyana, but the contour 
is narrower, the palatal folds less widely separated, as are also 
the angular and parietal lamellae. 

Mr. Ancey named the Palihoukapapa specimens in his col- 
lection (now in the Bishop Museum) Nesopupa baldwini var. 
centralis, but the original locality of N. b. centralis was Olaa 
at the time it was described no Palihoukapapa material had 
been under examination. Moreover, a form of N. baldwini 
occurs at Olaa, and the present species has not been found 
there. The identification was apparently a hasty one, as 
N. baldwini and its varieties have dorsal sulci, lacking in this 
species. 

Living examples collected by Forbes at about 7,000 ft. eleva- 
tion, in the lava flow of 1823, are referable to this species. 
They are almost similar in size, though the living form has 
slightly more convex whorls; the color is brussels brown on 
the last whorl, gradually becoming paler above but none of 
the specimens have a pale peripheral zone. The surface has 
an oily sheen and under a strong lens is minutely closely 



296 HAWAIIAN NESOPUPAE. 

wrinkled ; the lamellae and plicae of the aperture have about the 
same development as in specimens from Palihoukapapa, except 
that the angular lamella is slightly longer and stronger. One 
of the specimens measures: length 1.83, diam. 0.98, apert. 
(diag.) 0.73 mm. 

There is but a single slight difference between these speci- 
mens and those collected by Gouveia at Waihou. In the latter 
the angular lamella is considerably longer, otherwise they 
agree perfectly. 

10. NESOPUPA BISHOPI C. & P., n. sp. PL 28, fig. 4. 

The shell is minute, perforate (perforation very minute, 
nearly closed), nearly cylindrical, argus brown, lighter colored 
above, rather thin, translucent, with an oily sheen, minutely 
very weakly striate. Spire with nearly parallel outlines below, 
tapering above to a somewhat obtuse apex. The individual 
whorls somewhat convex, separated by a rather shallow, suture. 
Whorls 5%, the embryonic 11/2 nearly smooth, under a strong 
lens minutely indistinctly punctate ; rest of the whorls faintly 
striate, striae weak, oblique, discontinous ; under the microscope 
the whole surface is .seen to be covered with minute short 
wrinkles; last whorl cylindrical, tapering towards the base, 
ascending slightly and slowly near the aperture. Flattened 
outside of the palatal plicae, having a very small impression 
over the lower one, and slightly swollen just before reaching 
the aperture. Aperture truncate-ovate, scarcely oblique, 
nearly perpendicular, the lip insertions remote, not united 
by a callus. Angular lamella deeply seated, weak, short, 
about one-half the height of the parietal ; parietal lamella 
stronger, emerging further, nearly perpendicular to the 
parietal wall ; columellar lamella weaker than in most species 
of this group, short, horizontally inserted on the columella 
and deeply seated. Two palatal plicae of about equal length, 
rather short, not approaching the peristome and nearly 
parallel, the upper emerging slightly further than the lower. 
Peristome very slightly thickened, the columellar margin ex- 
panded, the lower and outer margins nearly erect. Length 
2.08, diam. 1.1 ; apert., greatest length 0.8 mm. 



HAWAIIAN NESOPUPAE. 297 

E. Maui : Haleakala Crater, near Crystal Cave, under stones 
011 the floor of the Crater (Cooke). Type 12465 Bishop 
Museum, paratype 44696 A. N. S. P. 

As far as known only three specimens of this species have 
been collected. It differs principally from N. limatula, from 
the same island, by its larger size and weaker lamellae. 

11. NESOPUPA FORBESI C. & P., n. sp. PL 28, fig. 5. 

The shells are larger than any of the other species from 
Hawaii, thinner and more transparent, more glossy, orange- 
citrine and with very low almost evenly spaced striae, the 
interstices not wrinkled as in living examples of central/is or 
a-nceyana. Spire oblong with very convex outlines, tapering 
gradually to the apex. Whorls 5, the embryonic almost 
smooth, hardly granulose under the microscope, the evenly 
spaced rather distant low striae beginning almost abruptly 
near the middle of the second whorl. Last whorl ascending 
slightly, swollen back of the aperture and hardly flattened 
over the palatal plicae. Aperture irregularly trapezoidal with 
rounded angles, the lip-insertions remote, united by a thin 
callus. Angular lamella minute, almost dentiform, deeply 
seated ; parietal deeply seated, long, strong, perpendicular to 
the parietal wall; columellar rather strong, deeply seated, 
slanting downwards; two palatal plicae short, rather deeply 
seated, converging inwardly, the lower considerably larger 
than the upper. Peristome tinged with a darker color than 
the rest of the shell, erect, very faintly thickened within. 
Length 2.08, diam. 1.22 ; apert., greatest length 0.85 mm. 

Hawaii: Huumula (Forbes). Type 39288 Bishop Museum. 

Only two specimens of this rare species are known at present. 
Both were taken by Forbes on plants in a large kipuka in the 
1855 Flow, about half way between Halealoha and Ainahou, 
at about 5,000 ft. elevation. The size of this species easily 
distinguishes it from the other species from Hawaii. From 
JV. bishopi it is most easily separated by its lighter color, 
thinner shell more convex outlines, and the surface, which is 
very minutely granular rather than wrinkled in the inter- 
stices between the striae. 



298 HAWAIIAN NESOPUPAE. 

12. NESOPUPA INFREQUENS C. & P., n. sp. PL 28, fig. 7. 

The shell is simstral, minute, sub-perforate (perforation 
very minute), nearly cylindrical, buckthorn brown, thin, trans- 
lucent, hardly glossy ; under a strong lens the surface is very 
minutely, closely and obliquely wrinkled, the wrinkles hair- 
like, rather short, more or less anastomosed. Spire with con- 
vex outlines, gradually tapering towards the apex, individual 
whorls slightly convex, separated by a rather narrow suture. 
Whorls 5, the embryonic nearly smooth, under a strong lens 
minutely granulose, the granules apparently arranged in trans- 
verse rows, the minute striae gradually appearing at about the 
middle of the second whorl. The last whorl tapers gradually 
to the base, ascending slightly near the aperture, slightly flat- 
tened over the palatal plicae, and only faintly swollen back of 
the aperture. Aperture scarcely oblique, nearly perpendicu- 
lar, irregularly truncate-ovate, lip-insertions remote, united 
by a thin transparent callus. Angular lamella short, lamella- 
like, deeply seated; parietal strong, high, emerging further 
than the angular, perpendicular to the parietal wall; colu- 
mellar lamella rather short, strong, deeply seated, indistinctly 
slanting downwards; two palatal plicae not approaching the 
peristome, rather short, nearly parallel. Peristome very 
slightly thickened within, arched above the columella, slightly 
expanded on the lower margin, the outer margin erect. Length 
1.65, diam. 0.98, apert. (diag.) 0.65 mm. 

Kauai: Halemanu, on a tree trunk; also on ferns (Cooke), 
Holotype 15489 Bishop Museum. 

It is the only sinistral Hawaiian Nesopupa known. Besides 
its sinistral coil, it is easily recognized from the other species 
of Nesopupa by the slightly coarse anastomosing striae of the 
lower whorls. 

One of us (Cooke) recently collected probably about 40 
specimens in different localities, the extreme points probably 
six miles apart. It has a different habit from any of the 
true Nesopupae ; all the specimens taken in 1919 were on the 
fronds of ferns (Asplenium arnottii). A single one, the type, 
had previously been found on a tree. 



HAWAIIAN NESOPUPAE. 299 

Section Nesodagys new section. 

Surface like Limbatipupa, but there is a long, lamella-like 
angular lamella reaching the lip. 

The two species included in this section are more or less 
intermediate between Limbatipupa and Nesopupa. 
a 1 . Shell oblong to ovate, embryonic whorls granulose. 

P. wesleyana and sub-species, no. 13. 

a?. Shell short, almost globose, embryonic whorls minutely 
spirally striate. P. thaanumi, no. 14. 

13. NESOPUPA WESLEYANA Ancey. PL 29, figs. 1, 2. 

''Shell oblong, thin, perforate, scarcely shining, under a 
strong lens granulate, brown-corneous, ornamented with 
oblique and distant, more or less deciduous, membranous cos- 
tulse, slightly wavy on the last whorl. Spire oblong, slightly 
obtuse. Whorls 5, separated by an impressed suture, the last 
oblong, ascending slightly towards its end, tapering ^gradually 
towards the base, slightly dilated near the aperture. Aper- 
ture vertical, truncate-ovate, very slightly receding at the base, 
nearly equal to % of the length, brownish within, 5-plicate; 
peristome not continuous, expanded and reflexed throughout, 
slightly thickened, sublabiate, with a unicolorous brownish 
lip. Angular lamella arcuate, elongate, slightly bent ; parietal 
a little more deeply seated, nearly median in position; colu- 
mellar lamella acute, deeply seated; two elongate, parallel 
palatal, the lower more deeply seated. Length 2, diam. 1, 
aperture about 0.75 mm. ' ' ( Ancey ) . 

Hawaii: Hilo, 4 miles Olaa road, type locality (Thaanum, 
Cooke) ; Glenwood (Pilsbry) ; Waipio Pali (Henshaw, Thaa- 
num) ; Kainbow Falls (Cooke) ; Puna (Thurston) ; fossil at 
Puuwaawaa (Thurston), Huehue (Gouveia) Palihoukapapa 
(Thaanum, Henshaw). Kahoolawe; fossil at Kanapou 
(Forbes & Stokes). E. Maui: Kaupakalua (Baldwin), fossil 
at Kanaio and Auwahi (Fleming). W. Maui: Waiakapu and 
Waihee (Forbes), Hahakea and lao (Cooke). Oahu: Ahui- 
manu (Cooke), Kamanaiki (Gouveia) fossil at Kailua 
(Pilsbry). No typical specimens of this species have been 



300 HAWAIIAN NESOPUPAE. 

seen from Kauai. Type 18704, Bishop Museum; paratypes 
18707 Bishop Museum, 44727 A. N. S. P. and in Thaanum coll. 

Nesopupa wesleyana ANCEY, Proc. Malac. Soe. London, 
vi, June 1904, p. 123, pi. 7, fig. 16. 

This species is easily recognized from P. newcombi by its 
more cylindrical spire and the long, lamella-like angular 
lamella. Typical P. wesleyana is usually found on rather 
damp rocks, though it is occasionally taken on rather smooth- 
barked trunks of trees. It does not seem to possess as many 
varieties or races as P. newcombi and the few varieties, noted 
below, are much more constant in all their characters than 
those of P. newcombi. 

The embryonic whorls are rounded, scarcely lighter colored 
than the rest of the shell, under a strong lens very minutely 
granulose, the granules not appearing to be arranged in any 
regular manner. 

In the typical form the angular lamella is situated parallel 
to the parietal inwardly, curving outwards in front and unit- 
ing to the outer lip at a slight distance back of the outer 
margin. It descends rather abruptly in front but gradually 
at the inner end. Sometimes there is a slight depression or 
notch just back of outer end. This is very well represented 
in fossil specimens collected by Pilsbry at Kailua, Oahu. The 
parietal lamella is strong, rather deeply seated and nearly per- 
pendicular to the parietal wall. The columellar is deeply 
seated, perpendicular to the columella and rather long. The 
lower palatal is somewhat deeply seated, long, parallel to, 
but stronger than the upper. The upper palatal extends 
nearer to the margin than the lower, inwardly its free margin 
is slightly bent towards the lower palatal. Externally there is 
a slight impression over it. 

A somewhat larger fossil race was found by Messrs Thaanum 
and Henshaw at Palihoukapapa. The whorls are slightly 
flatter than in the typical form. One of the specimens meas- 
ures: Length 2.15, diam. 1.22, apert. (diag.) 0.86 mm. 5% 
whorls. 

The type specimen has the following dimensions: Length 



HAWAIIAN NESOPUPAE. 301 

1.95, diam. 1.15, apert. (diag.) 0.8 mm. The color of fresh 
typical specimens from Hilo, Hawaii is buckthorn-brown. 

130. Form gouveia C. & P. PL 29, fig. 4. 

A very small race of wesleyana was found by Gouveia at 
Hookena, and by Forbes at Kapua, both localities in South 
Kona, Hawaii. In this form the outline is narrowly ovate, 
tapering above. The aperture is truncately ovate, its outer 
margin hardly flattened. Whorls 4%. In cross section the 
upper palatal is only slightly bent ; the angular unites with the 
outer lip at a slight distance from the margin. Length 1.65, 
diam. 0.98, apert, (diag.) 0.73 mm. Type 11081 Bishop 
Museum, paratype 39429 Bishop Museum, and 44732 A. N. S. P. 

13&. Form tryphera C. & P. PL 29, fig. 3. 

A form which is usually smaller than the typical and which 
appears to occur in pure colonies, has the upper palatal fold, 
abruptly bent in the middle, the inner half descending towards 
the lower palatal fold. A few of the specimens from Oahu 
(Nuuanu, Makiki and Moanalua, coll. by Spalding), are nearly 
equal to the typical form in size, but the majority of the 
colonies are decidedly smaller. Embryonic whorls similar to 
those of the typical form. 

Typical examples of this form have been taken from the 
following localities: Kauai : Lihue. Oahu: Palolo (type loc.), 
Tantalus, Nuuanu and Kaipapau, in the Koolau Range; 
Palehua and Popouwela, in the Waianse Range. Molokai : 
Puunea and Mapulehu (Thaanum). Except for the larger 
form from Oahu mentioned above the examples are remark- 
ably uniform in size and other characters. Length 1.75, diam. 
1.1, apert. (diag.) 0.8 mm. 4% whorls (Palolo). Type 11082 
Bishop Museum, para types 15422 Bishop Museum and 44714 
A. N. S. P. 

13c. N. w. rhadina C. & P., n. subsp. PL 29, fig. 13. 

The shell is slender, oblong, usually tapering above, brussels 
brown, convex, closely and regularly marked with well-de- 
veloped membranous costse. Spire with convex outlines. 



302 HAWAIIAN NESOPUPAE. 

Whorls 5%, very convex. The embryonic whorls are more 
coarsely granulose than any of the forms noted in this sub- 
genus. The granules for the first half whorl are arranged 
longitudinally, giving the appearance of striae; at about the 
beginning of the second whorl the very faint transverse striae 
appear to be made up of uniting granules. These striae, gradu- 
ally become stronger with the growth of the shell, the granules 
persisting in the inter-costal spaces, gradually becoming 
fainter and disappear about the end of the second whorL 
Aperture truncate-ovate, the outer margin hardly flattened, 
the lip insertions slightly converging, furnished with 5 lamellae. 
Angular lamella somewhat thickened and calloused in front, 
slender and thin inwardly, lower than the parietal. Colu- 
mellar lamella slightly slanting downward. The upper palatal 
fold is equal to if not higher than the lower, not quite as 
deeply seated, and slightly shorter. Length : 2.32, diam. 1.15, 
apert. (diag.) 0.85 mm. Type 11083 Bishop Museum, para- 
types 35126 Bishop Museum and 44726 A. N. S. P. 

Molokai: Poholua, type locality (Cooke) ; Kilohana and 
Puukolekole, Kamalo (Pilsbry and Cooke) ; Kalihi, Puunea, 
Waikolu (Cooke) ; Mapulehu (Thaanum) ; fossil specimens 
from Mauna Loa, Moomomi, Kalaeokailio (Cooke). 

This subspecies was at first considered as a separate species, 
but on comparing the large series of both wesleyana and 
rhadina in the Bishop Museum collection it appears to be 
better to consider both forms as belonging to the same species. 
N. rhadina is easily recognized from wesleya/iM by its more 
convex whorls, darker color and narrower outlines. 

Living examples of this subspecies are only taken under 
the loose bark of trees or in moss growing on tree trunks. 
It is never found abundantly, due to the amount of time con- 
sumed in finding only a few specimens. The most usual 
habitat is under loose bark of the lehua (Metrosideros 
polymorpha) . 

On the flat east of the peak, Puukolekole, it varies in size 
and shape. 

Alt. 1.95, diam. 1.18 mm. 

Alt. 2.15, diam. 1.3 mm. ; decidedly ovate-conic. 



HAWAIIAN NBSOPUPAE. 303 

Alt. 2.25, diam. 1.3 mm.; subcylindric. 

In forest at the head of Kamalo there is a large, stout, cyl- 
indric form with the laminae of the surface more crowded, 
and generally having an interpalatal fold. Length 2.4, diam. 
1.25 mm. This may be called form kamaloemis (pi. 29, fig. 9) . 

Kauai : This sub-species has been taken in several localities 
and is probably well distributed over the whole of the moun- 
tainous area of the island. It is also occasionally found in 
some of the fossil deposits of the lowlands. The specimens 
from this Island are fairly typical. They are slightly more 
tightly coiled than the typical form and the outlines are a little 
more convex. A typical example, from back of Lihue, meas- 
ures: Length 2.3, diam. 1.1, apert. (diag.) 0.83 mm. A fossil 
specimen from Limahuli is decidedly smaller, more conical in 
outline and there are but 5 whorls. Length : 1.95, diam. 1.05, 
apert. (diag.) 0.75 mm. 

Oahu : This sub-species is widely distributed over the Koolau 
range but is not represented by a large series in any of the 
lots. From the Waiana? Mts. only a very few examples seem 
to have been taken. It is also found, though only in a few 
isolated cases abundantly, in a number of the fossil deposits 
from this island. Most of the specimens are considerably 
smaller than the typical form from Molokai. PL 29, fig. 7 
represents a fossil specimen from Kahuku coral bluff. 

Specimens taken from a few of the lots in the Bishop 
Museum collection showed the following measurements. 

Length 2.02, diam. 1.1, apert. (Diag.) 0.79 mm. 5 whorls; 
Manoa Palolo ridge (Cooke). 

Length 1.9, diam. 1.0, apert. (Diag.) 0.75 mm. 5 whorls; 
Waimano (Gouveia). 

Length 1.95, diam. 1.1, apert. (Diag.) 0.8 mm. 5 whorls; 
Waialaeiki ( Gouveia ) . 

Length 2.08, diam. 1.08, apert. (Diag.) 0.76 mm. 5% whorls; 
Popouwela, Waianae Mts. (Cooke). 

Length 1.8, diam. 0.95, apert. (Diag.) 0.7 mm. 5 whorls; 
Malaekahana, fossil (Cooke). 

Length 2.0, diam. 1.07, apert. (Diag.) 0.76 mm. 5 whorls; 
Kahuku, fossil (Cooke). 



304 HAWAIIAN NESOPUPAE. 

Length 2.25, diam. 1.13, apert. (Diag.) 0.85 mm. 5% whorls; 
Manoa, fossil (Cooke). 

Maui: Specimens of this subspecies from the highlands of 
West Maui, have the apices much eroded giving the appearance 
of a very blunt spire. The whorls are very convex and the 
outlines are almost cylindrical. The surface is covered with 
rather low, close, membranous costae. A well preserved speci- 
men (the apex of which is not much eroded) measures: Length 
2.35, diam. 1.22, apert. (diag.) 0.97 mm. 5y 2 whorls. 

Specimens from East Maui are similar to those from West 
Maui. 

Lanai: I have seen only two specimens of this sub-species 
from Lanai. Both were collected by Forbes and each came 
from a different locality. Both specimens appear to be more 
closely related to the Molokaian form than the Mauian. One 
of them measures: Length 2.15, diam. 1.15, apert. (diag.) 
0.86 mm. 5% whorls. 

No specimens of this subspecies have been seen from Hawaii, 
and so far as I know up to the present time it has not been 
taken on that island. 

14. NESOPUPA THAANUMI Ancey. PI. 28, figs. 11, 12. 

"Shell dextral, minutely rimate-perf orate, of a pleasing 
corneous buff color, ornamented with rather distant, weak 
and very deciduous, silvery shining, membranous costula?; 
somewhat conic-ovate, obtuse. Whorls 4^, convex, suture im- 
pressed ; the upper quite minute ; the last sub-saccate, shortly 
ovate, scarcely tapering at the base, ascending slightly to- 
wards its end, very lightly impressed outwardly back of the 
aperture. Aperture erect, truncate-oval, sub-rotund, ex- 
tended outwardly and bent in front (above the middle portion 
of the dextral margin), armed with strong white lamella as 
follows : an arcuate angular, elongate but short, extending to 
the margin ; a little more deeply seated parietal lamella, enter- 
ing deeply; a transverse, internal columellar; and 2 palatals, 
the upper seated a little nearer the margin, both elongate but 
not very long. Lip narrowly expanded, with quite distant 
insertions. Length 1.33, diam. 1.0, alt. apert. 0.5 mm. (Ancey). 



HAWAIIAN NESOPUPAE. 305 

Hawaii: Olaa (D. Thaanum), type 18701 Bishop Museum, 
topotypes 44678 A. N. S. P.; Glenwood, in moss on tree 
trunks, abundant (Thaanum, Pilsbry). 

Nesopupa thaanumi (ANCEY), Proc. Malac. Soc., London, vi, 
Tune 1904, p. 123. 

Unfortunately the holotype from the Ancey collection is 

yadly broken. This species is very abundant in the type 

ocality, where it is found on tree-trunks, living among and on 

-he fronds of a loose-growing species of moss. Under similar 

onditions this species has been found in nearly all the 

1 Hawaiian Islands. P. thaanumi is somewhat similar to P. 

uesleyana tryphera. Its shell is, however, smaller, more com- 

)actly coiled, much lighter colored and the aperture is more 

Crowded w r ith the five lamellae. It has apparently some re- 

ationship to the species included in Nesopupa ss. It differs 

Tom all the species of the latter by its more convex whorls 

ind especially by the membranous character of its rather dis- 

ant riblets. Immature specimens from the type locality have 

lie embryonic whorls very minutely spirally striate. The 

ingular lamella is strong, long and slightly sinuous, high in 

'ront, tapering gradually backward, and does not terminate 

>n nor is it joined to the outer lip, but extends to the margin 

>f the parietal wall. In this character it resembles species of 

\esopupa ss. with this distinction: that in thaanumi this 

amella is stronger, with a more sinuous margin and the outer 

ortion is not parallel to the parietal, but is distinctly bent 

owards the outer lip. The parietal lamella is only slightly 

:iore deeply seated than the angular, and emerges almost to 

he margin on the parietal wall. It is very strong, high and 

Dug, and is nearly perpendicular to the parietal wall. The 

olumellar lamella is very strong, horizontally seated, rather 

>ng, diminishing rather abruptly inward, there turning up- 

. r ards and accompanying the columella as a mere thread. The 

wo palatal folds are nearly parallel, about equal in height 

nd length ; the lower slightly more deeply seated ; the upper 

xtendirig nearly to the margin of the outer lip, with its free 

dge abruptly bent, almost at a right angle, towards the 

)wer palatal. 

-' 



306 HAWAIIAN NESOPUPAE. 

Length 1.4, diam. 1.0, apert. (diag.) 0.65 mm. Whorls 4% 
(topotype). 

Length 1.56, diam. 1.0, apert. (diag.) 0.63 mm. Whorls 4y 2 
(topotype). 

Ancey's type specimen, according to my method, measures 
1.53 mm. in length. 

This species has not yet been reported from Kauai. 

Oahu : widely distributed over the Koolau range ; up to this 
time none have been seen from the Waianae Mts. Shells of 
the different colonies differ but slightly from each other in 
size and otherwise agree very closely. Most of the specimens 
are slightly lighter colored than those from Hawaii. 

Length 1.51, diam. 1.0 apert. (diag.) 0.64 mm., 4% whorls; 
Waialae-iki. 

Length 1.4, diam. 0.98, apert. (diag.) 0.61 mm., 4% whorls;. 
Nuuanu. 

Length 1.53, diam. 1.02, apert. (diag.) 0.64 mm., 4% whorls;. 
Kaliuwaa. 

Lanai : I have been able to examine but two specimens of this 
species collected on Lanai. Both specimens agree closely with 
the typical form. One of them measures: length 1.47, diam. 
0.98, apert. (diag.) 0.65 mm., 4% whorls. 

Maui : This species is only known to have been taken from 
three localities on West Maui, and one from E. Maui. Like the 
specimens from the other islands, all the Maui specimens agree 
closely with the typical form. A specimen from Honokahau 
measures: length 1.43, diam. 1.02, apert. (diag.) 0.64 mm.,, 
whorls. 



Section LIMBATIPUPA C. & P., n. sect. 

The shells are minute, brown to greenish-brown; the free 
margins of the costulae membranous and sometimes slightly 
prolonged into sharp membranous points; the 5 primary 
lamellae and folds present in most species, the angular lamella 
either shortly lamelliform, nodular or absent. Type N. new- 
combi. 

The species composing this section are differentiated by their 



HAWAIIAN NESOPUPAE. 307 

sculpture and the short, lamella-like, or nodular angular 
lamella, which is sometimes lacking. 

The species are rather closely connected ; however, they seem 
to be sufficiently differentiated from each other, and may be 
easily recognized. N. newcombi is by far the most variable 
species of the family found in the Hawaiian Islands, and a 
number of quite distinct races have here been grouped under 
this name without subspecific rank. 

Key to the Species of the Section Limbatipupa. 

a 1 . Angular lamella absent or represented by a slightly swol- 
len callus. 

6 1 . Whorls 4, embryonic whorls minutely spirally striate, 
diameter about 70% of total length. 

N. alloia, no. 19. 

b 2 . Whorls 4%-5, embryonic whorls granulose, diameter 
about 60% of total length. N. singularis, no. 18, 

a 2 . Angular lamella nodular or short lamelliform. 
b 1 . Costa? prominent, thin, membranous. 

c 1 . Upper palatal fold short lamella-like. 

N. newcombi, no. 15.- 
c 2 . Upper palatal fold absent or nodular. 

N. oahuensis, no. 16,. 
b 2 . Costae low, with very narrow membranous margin. 

N. kauaiensis, no. 17. 

15. NESOPUPA NEWCOMBI (Pfr.). Page 309, fig. 1. 

The shell is minute, perforate (perforation very minute,, 
circular), ovate, brownish olive, thin, somewhat translucent, 
slightly shiny, uniformly and widely costulate, the riblets with 
membranous margins, about 15 on the last whorl. In some 
specimens they are slightly spiniferous just above periphery. 
Intercostal spaces minutely striate (striae short, parallel). 
Spire conic with convex outlines, apex obtuse; individual 
whorls convex, separated by a rather deep suture. Whorls 
4%, the embryonic 1% whorls are microscopically granulose, 
the granules apparently arranged in transverse rows, the mem- 
branous costa? appearing abruptly in about the middle of the.- 



308 HAWAIIAN NESOPUPAE. 

second whorl. The rest of the whorls are costate. Last whorl 
large, rotund, tapering towards the rounded base, scarcely 
ascending near the aperture. Aperture nearly circular, 
scarcely oblique, perpendicular, the lip insertions approaching 
and united by a thin callus. Angular lamella short, lamella- 
like, united to the outer lip by a thick callus for about half 
its height and not extending to the parietal margin. Parietal 
lamella more deeply seated, large, strong, nearly perpendicular 
to the parietal wall. Columellar lamella strong, transversely 
seated and slightly slanting downward. Two palatal plicae 
rather deeply seated, nearly parallel, the lower slightly 
stronger than the upper. Peristome expanded, its outer and 
lower margins thin, membranous ; very slightly thickened with- 
in; columellar margin dilated above the umbilicus. Length 
1.65, diam. 1.1, aperture, greatest length 0.79 mm. 

Explanation of figures, page 309. 

Fig. 1. Nesopupa newcombi (Pfr.). Typical. 18714BM. Wai- 
pio, Kauai. 

2. N. newcombi seminulum. 23375BM. Kapua, Hawaii. 

3. 3d, N. newcombi multidentata. Type, 15316BM. Glen 

Ada. 

4. N. newcombi angusta. Type, 20227BM. Kipu, Kauai. 
4a-4d. N. newcombi interrupta. 44760. Kaelepulu, Kai- 

lua. 

5. N. newcombi interrupta. Type, 42673BM. Waiaha, 

Hawaii. 

6. 60-. N. newcombi interrupta. l 1 /^ miles west of Ka- 

huku. 44761. 

7. N. newcombi disjuncta. Type, 17219BM. Mokuleia. 

8. 8a. N. newcombi gnampta. Type, 12526BM. Lua- 

kaha, Nuuanu. 

9. N. kauaiensis (Anc.). 20608BM. Kipu, below the 

Gap, Kauai. 

10. N. kauaiensis (Anc.). 20730BM. Kipu, slopes of 
Haupu, Kauai. 



HAWAIIAN NESOPUPAE. 



309 




310 HAWAIIAN NESOPUPAE. 

Hawaiian Islands (Pfeiffer) ; Hawaii (Boettger, Pease), 
Waipio Valley (Thaanum). Lanai, Maunalei Gulch (Forbes) ; 
Mahana, Kaihalena (Spalding). Molokai, Mapulehu, (Thaa- 
num and Cooke) ; Puulua and Kalihi (Cooke). Oahu, Palolo, 
Tantalus, Makiki (Cooke), Waialaeiki (Bridwell), Pauoa 
(Gouveia). 

The above is the distribution of the typical form as known 
at present. More or less distinct races of this species have 
been taken on all the main islands except Niihau. 

f Pupa newcombi PPR., Proc. Zool. Soc. London, 1852 
(1854), p. 69; Monographia Heliceorum Viventium, hi, 1853, 
p. 530. 

Pupa newcombi PFR., Monographia Heliceorum Viventium, 
iv, 1859, p. 682. KUESTER, Conchyl. Cab., Pupa, 1859, p. 172, 
pi. 20, f. 23, 24. BOETTGER, in von Martens, Conchologische 
Mittheilungen, i, 1881, p. 56, pi. 11, f. 12; pi. 12, f. 13. 
ANCEY, Memoires de la Societe Zoologique de France, v, 1892, 
p. 709. Nesopupa newcombi PILSBRY, Proc. A. N. S. Phila., 
1900, p. 433. 

Vertigo costulosa PEASE, Proc. Zool. Soc. London, 1871, p. 
462. Pupa costulosa PPR., Monographia Heliceorum Viven- 
tium, viii, p. 408. ANCEY, Mem. de la Soc. Zool. de France, 
v, 1892, p. 711. 

This is one of the most variable species of shells found on 
the Hawaiian Islands. The typical form, with five lamellae in 
the aperture, appears to be rather uncommon. Specimens with 
from 6 to 10 lamellae and folds are found on nearly all the 
islands of the group. In some cases these races are found in 
nearly pure colonies, in others individuals representing two or 
more races are found in a single colony. 

Nearly 350 specimens from 33 colonies were closely examined 
and among them 40 different lamella-formulae were noted. 
The presence and form of almost all the lamellae seem to be 
variable characters except in a very few races. The only 
teeth found in all the specimens were the columellar, lower 
and upper palatal. In one (probably pathological) specimen 
the parietal and angular lamellae were absent, and in another 
the parietal lamella was lacking. Except for extremely rare 



HAWAIIAN NESOPUPAE. 311 

specimens the presence of these two lamellae must be con- 
sidered as a constant character. 

Two distinct forms of the angular lamella were found; in 
the first this lamella is reduced to a nodular or a short 
lamella-like form. This is usually united to the outer lip by 
A thick callus sometimes of more than half its height. In 
most cases it does not extend to the margin of the parietal 
callus, while in others it not only reaches the margin but is 
continuous with it, forming a low descending ridge along the 
callus of the margin. In the second form the angular lamella 
is discontinuous. It is represented by a low nodule seated 
deep within and to the outside of the parietal lamella, followed 
by a short interval with no indication of any thickening, and 
terminating near the angle with the usual nodule, as in fig. 4&. 
Forms with this character of the lamella are abundant in the 
Pleistocene deposits on nearly all of the islands. 

The parietal lamella is always strong and usually slants 
slightly outwards, but is often perpendicular to the parietal 
wall ; in some cases the free margin is slightly sigmoid. The 
infra-parietal lamella is usually represented by a low nodule- 
like swelling, rather deeply seated. In a very few cases speci- 
mens were found in which this lamella is discontinuous as 
described in the case of the second form of the angular lamella. 
The supra-columellar and subcolumellar lamellae are only pres- 
ent in a few specimens. They are never lamelliform, but 
appear as low callous swellings. The columellar lamella is 
always strong, horizontally entering, and generally slanting 
slightly downwards and emerging nearly to the outer margin 
of the columella. In most cases it terminates rather abruptly 
at the back and follows the margin of the columella upward 
as a low thread-like swelling. In two races, one from Kauai 
and the other from Oahu, the inner portion of this lamella 
is turned abruptly downwards terminating at the base of the 
columella, as in typical Nesopupa. The subcolumellar lamella, 
when present, is usually represented by a low elongate swell- 
ing parallel to the basal fold. The basal fold when present is 
generally lamelliform and reaches about half the size of the 
lower palatal. An infrapalatal fold is found only occasion- 



312 HAWAIIAN NESOPUPAE. 

ally. It is more deeply seated than the outward termination 
of the lower palatal fold, and has a low lamella-like form. 
The lower palatal fold is always present; it is deeply seated 
and generally much stronger than the upper palatal. The 
interpalatal fold is usually low, lamelliform, and more deeply 
seated than either the upper or lower palatals. Like the 
lower, the upper palatal fold was present in every specimen 
examined. The suprapalatal fold, when present, is nodular 
in form and is ordinarily seated about half way between 
the termination of the upper palatal fold and the inner margin 
of the peristome. 

The outline of the shell varies from narrowly conic-ovate 
to broadly ovate. There are from 4 to 4% whorls in adult 
specimens. 

The number of costag on the last whorl varies from about 
15 to nearly 30. In each of the different colonies the number 
of costas is nearly uniform. 

Typical specimens of newcombi from Oahu are slightly 
larger than those examined from Hawaii. Otherwise they 
agree fairly closely. A specimen from Palolo has about 17 
costas on the last whorl. It measures: length 1.75, diam. 1.1 
apert. (diag.) 0.73 mm., 4i/ 2 whorls. Another specimen from 
Tantalus measures: length 1.83, diam. 1.2, apert. (diag.) 0.70 
mm., 4% whorls. In all the specimens from the latter locality 
there are from 25 to 28 costae on the last whorl. Lanai speci- 
mens are somewhat smaller than those from Hawaii. They 
have about 20 costae on the last whorl. The outline of the 
shell is globosely-ovate. An adult specimen with 4% whorls 
measures: length 1.53, diam. 1.1, apert. (diag.) 0.67 mm. 
Molokai specimens are very similar to those from Oahu. An 
adult shell with 414 whorls measures: length 1.72 diam. 1.05 
apert. (diag.) 0.73 mm. 

When typically developed the minor races can be deter- 
mined by the following key. 

a 1 . Columellar lamella straight or slightly upturned within. 
fe 1 . Angular lamella a short lamella or tubercle only. 
c 1 . Form rather broadly ovate-conic. 



HAWAIIAN NESOPUPAE. 313 

d\ With 5 or 6 teeth. 

Length about 1.75 mm., typical newcombi. 

Length about 1.5 mm., form seminulum. 

d~. With 9 or 10 teeth. form multidentata. 

c 2 . Form narrower. form angusta. 

b 2 . Angular lamella interrupted, having a low inward 

lamella. 

c 1 . Aperture with 5 or 6 teeth. subsp. interrupt d. 
c 2 . Aperture with 7 or 8 teeth. form disjuncta. 

a 2 . Inner end of the columellar lamella turned downward. 

subsp. gnampta. 
Pfeiffer originally described this species thus : 

"P. newcombi Pfr. T. subperforata, ovata, tenuis, longi- 
tudinaliter costata, haud nitens, diaphana, saturate fusca; 
spira inflata, apice acutiuscula; anfr. 4 convexi, ultimus % 
longitudinis subaequans, basi rotundatus; apertura obliqua, 
semicircularis, edentula; perist. tenue, vix expansiusculum, 
margine columellari subreflexo. Long. 2, diam. 1 mill. (Mus. 
Cuming. et Coll. Nr. 2.) Habitat in insulis Sandwich (New- 
comb)." 

This description appeared first in the Monographia Heli- 
ceoriim, later in Proceedings of the Zoological Society of 
London. It will be noted that he termed it edentulous. Sub- 
sequently, in the Manographia iv, 1859, he characterized his 
former description as incomplete, and reprinted it except that 
in place of the word edentula he substituted ' ' dentibus 4 pro- 
fundis coarctata : 1 parietal! compresso, 1 subtransverso ad 
columellam, 2 in palato ; ' ' the rest of the description, the meas- 
urements, habitat, and collector standing unchanged. 

It has been surmised that he confused two species, one tooth- 
less, like P. admodesta, the other dentate; but a more likely 
supposition is that he did not see the teeth at first. This is 
clearly what he implied by calling his first description "in- 
completus. ' ' 

The supposed types of newcombi in London, examined by 
one of us (Cooke) are typical specimens of N. wesleyana Anc. ; 
doubtless they had been substituted for the original examples. 
Boettger has described the true newcombi from specimens in 



314 HAWAIIAN NESOPUPAE. 

the Pfeiffer-Dohrn collection. The same author has stated 
that Vertigo costulosa Pease, according to specimens from 
Pease, is completely synonymous. Yet obviously Pease con- 
fused two species in his description, as he says that the last 
whorl is sometimes shortly bisulcate towards the lip. This is 
not true of any form of N. newcombi, and probably indicates 
that he had also a Nesopupilla. 

150. Form seminulum (Boettger). Page 309, fig. 2. 

' * The shell is smaller, narrower, more elongate, horn-brown, 
more often the angular lamella is less distinct. Length 1.5 
to 1.75; diam. 1 to 1.25 mm." (Boettger). 

Hawaii : (Boettger, Thurston, Gouveia) . W. Maui : Waihee, 
Lahaina (Forbes, Cooke). E. Maui: Alalele (Cooke). Molo- 
kai: Mapulehu (Thaanum). Oahu: Tantalus, Nuuanu and 
fossil at Rocky Hill (Cooke), Waiawa and Waialaeiki (Spald- 
ing). Kauai, various localities (Cooke). 

Pupa newcombi var. seminulum BOETTGER in von Martens, 
Conchologische Mittheilungen i, 1881, p. 58, pi. 12, f. 14. 
Not Pupa seminulum Lowe, 1852. 

Except for a slightly narrower form, as Boettger has already 
pointed out, this variety differs in no way from some of the 
various races of P. newcombi. In fact some of the narrower 
forms mentioned above are not proportionately broader than 
the measurements given by Boettger would indicate. The 
name is preoccupied, but as its value is rather uncertain it 
is not renamed at this time. 

A number of shells from different localities have the fol- 
lowing measurements: 

Length 1.5, diam. 1.0 mm. 4% whorls; Kapua, Hawaii; 
23375. 

Length 1.53, diam. 0.98 mm. 4% whorls; Puuwaawaa, 
Hawaii; 39784. 

Length 1.65, diam. 1.05 mm. 4% whorls ; Lahaina, W. Maui ; 
20967. 

Length 1.8, diam. 1.1 mm. 4% whorls; Waihee, W. Maui; 
39012. 



HAWAIIAN NESOPUPAE. 315 

Length. 1.53, diam. 0.98 mm. 414 whorls; (fossil) Rocky 
Hill, Oahu; 39832. 

Boettger mentions only five lamellae in the typical form of 
this species. In most of the races examined by us, a low 
lamelliform basal fold is present. 

15&. Farm multidentata. Page 309, figs, 3, 3a. 

An easily recognized form that is widely distributed on 
Oahu is somewhat larger than the typical form, Living ex- 
amples are only found on damp moss-covered stones. The 
aperture is usually furnished with 9 or 10 lamellae and folds, 
the infraparietal, suprapalatal and basal being present in all 
the specimens examined. The interpalatal was present in 
about 90% of the specimens, and the infrapalatal and sub- 
columellar in about 20%. Angular lamella not very long, 
lamelliform. 

Length 1.9, diam. 1.12, apert. (diag.) 0.83 mm.; 4% whorls. 
Glen Ada, Nuuanu. 

Length 1.95, diam. 1.3 mm. Glen Ada, type. 

15c. Form angusta. Page 309, fig. 4. 

The most abundant form on Kauai, and one which is nearly 
uniform in all its characters, is longer and slightly narrower 
than the typical form; the angle lamella is somewhat longer 
than in the form already described. A low lamelliform basal 
fold is always present. There are about 20 riblets on the last 
whorl and most of the shells are somewhat darker in color 
than the other forms. An adult specimen with 4% whorls 
measures: length 1.96, diam. 1.15, apert. (diag.) 0.73 mm. A 
similar form though slightly smaller is found on Oahu. An 
adult specimen with 4% whorls measures: length 1.83, diam. 
1.1, apert. (diag.) 0.73 mm. 

15d. JV. newcombi interrupta C. & P., n. subsp. Page 309, 

figs. 4or-6a. 

In the following races the angle lamella is discontinuous; 
there is a nodular outer and a lamelliform inner portion. 
From a number of fossil deposits of Kona, Hawaii, specimens 



316 HAWAIIAN NESOPUPAE. 

have been collected by Thwing and Gouveia. The infra- 
parietal and basal folds are usually present. One of the 
specimens from Waiaha, Hawaii, measures : Length 1.65, diam. 
0.98, apert. (diag.) 0.64 mm.; 4% whorls. 

A number of fossil specimens of this form were also found 
at Kanapou, Kahoolawe, by Forbes and Stokes. In these the 
infraparietal lamella was present in about half of the speci- 
mens; the basal fold was present in all. One of them meas- 
ures: length 1.85, diam. 1.07, apert. (diag.) 0.65; 4% whorls. 

In a fossil deposit at Kainaio, E. Maui, a few specimens were 
found by Fleming. These are the smallest so far as seen. In 
nearly all the infraparietal and basal folds are present. One 
measures: length 1.47, diam. 0.98, apert. (diag.) 0.61; 4 1 /4 
whorls. 

This form is exceedingly abundant in a fossil deposit on 
the northern slope of Mauna Loa, Molokai. In slightly more 
than half the specimens the infraparietal was present. The 
basal fold was noted in all but one example. In a few speci- 
mens infrapalatal or interpalatal plicae were seen, and in two 
instances both of them were present in the same specimen. An 
average shell measures: Length 1.7, diam. 1.12, apert. (diag.) 
0.67 ; 4y 2 whorls. 

In Oahu this form is exceedingly abundant in fossil de- 
posits, and living examples are not uncommon; most of the 
living newcombi from the Waianse Mts. belong to form 
interrupta. 

It sometimes varies extraordinarily in shape, 4a-d represent- 
ing selected specimens from Kaelepulu, Kailua where it is an 
abundant Holocene fossil. The inner portion of the angular 
lamella is sometimes wanting. Figs. 6, 6& are from an aver- 
age and a short example from the coral bluff west of Kahuku, 
showing variations in teeth. The abundance of this and 
other species of Pupillidae in this deposit is astonishing. Fig. 
6 measures, length 1.7, diam. 1.2 mm. 

Length 1.55, diam. 1.02 mm., whorls 4%; fossil, Kailua; 
(fossil). 

Length 1.59, diam. 1.1 mm., whorls 4 1 /* ; fossil, Kahuku ; 
(fossil). 



HAWAIIAN NESOPUPAE. 317 

Length 1.73, diam. 1.1 mm., whorls 4% ; fossil, Rocky Hill. 

Length 1.72, diam. 1.17 mm., whorls 4%; fossil, Ewa, 
Waiane Mts. 

Length 1.78, diam. 1.1 mm., whorls 4% ; fossil, Lualualei, 
Waianas Mts. 

Length 1.78, diam. 1.1 mm., whorls 4% ; fossil, Palolo. 

loe. Form disjuncta. Page 309, fig. 13. 

In another slightly distinct form, with a discontinuous an- 
gular lamella, a race which is widely distributed on Oahu, the 
infraparietal, inter-palatal, suprapalatal and subcolumellar, 
as well as the basal are almost uniformly present. No speci- 
mens of this race were found in fossil deposits. In a very few 
cases the supra- and subcolumellar infrapalatal folds were also 
noted. It is larger than form interrupts. 

Length 1.86, diam. 1.2 mm. ; whorls 4%, Mokuleia, Waiange 
Mts. 

Length 1.78, diam. 1.17 mm. ; whorls 4%, Moanalua. 

Length 1.65, diam. 1.1 mm. ; whorls 4%, Kalihi. 

15/. N. newcombi gnampta C. & P., n. subsp. Page 309, figs. 

14, 15. 

A widely distributed form of Oahu, is nearly typical except 
that the inner end of the columellar lamella turns abruptly 
downward forming a right-angle, with its inner leg terminat- 
ing at the base of the columella. One from Luakaha, Nuuanu, 
measures: length 1.65, diam. 1.05, apert. (diag.) 0.73 mm.; 
4% whorls. Specimens from Kauai, with a similar columellar 
lamella, are slightly larger, one of them measuring: length, 
1.78, diam. 1.1, aperture (diag.) 0.79 mm. 4% whorls. 

16. NESOPUPA OAHUENSIS C. & P., n. sp. PI. 29, figs. 11, 12. 

The shell is minute, perforate, broadly ovate-conic, buck- 
thorn-brown, thin, slightly translucent, rather dull, marked 
with distant, weak, thin, membranous riblets (about 15 to 18 
on the last whorl), the intercostal spaces minutely striate, 
striae parallel. Spire with slightly convex outlines, tapering 
above to a rather sharp apex. The individual whorls are very 
convex, separated by a deep suture. Whorls 4%, the em- 



318 HAWAIIAN NESOPUPAE. 

bryonic, under the microscope, minutely granulose, the minute, 
even, close striae appearing abruptly at about the middle of the 
second whorl; last whorl large, saccate, tapering towards the 
base and rounded about the margin of the perforation, ascend- 
ing slightly and slowly near the aperture. Aperture scarcely 
oblique, nearly perpendicular, irregularly truncate-ovate, 
slightly flattened on the right margin and arched towards the 
upper insertion. The lip insertions distant but somewhat con- 
verging and united by a thin callus. Angular lamella puncti- 
form, seated somewhat within the margin and united to the 
outer lip by a very thin callus. Parietal lamella more im- 
mersed, nearly perpendicular to the parietal wall, lamella-like, 
high and short. Columellar lamella small, short, deeply 
placed, nearly horizontal, the edge slightly slanting down- 
ward. Lower palatal fold rather deeply placed, short, rather 
high, lamella-like ; upper palatal, when present, not so deeply 
seated, weak, low, almost punctiform. Peristome expanded, 
the outer margin rather thin, the inner margin slightly thick- 
ened. Length 2.15, diam. 1.5, aperture 1 mm. 

Oahu: Nuuanu Valley at Luakuha falls, type locality 
(Cooke) Ahuimanu (Spalding). Type 11075 Bishop Museum; 
paratypes 12559 Bishop Museum and 44695 A. N. S. P. 

This species is very closely related to P. kauaiensis ; it differs, 
however, by the thinner and lighter colored shell ; the spire is 
shorter and broader and the aperture is larger than in the 
latter species; also, the membranous eostee are more distinct 
and stronger. In P. oahuensis the upper palatal fold is re- 
duced to a minute tubercle, hardly lamella-like in form. In 
most of the specimens examined it appears to be entirely 
lacking. 

This species has only been taken, so far as known, on damp 
rocks at the sides of waterfalls. Its larger size and shorter 
lamellae easily separate it from the various forms of P. new- 
combi. 

17. NESOPUPA KAUAIENSIS Ancey. PL 29, figs. 5, 6 ; page 309, 
figs. 9, 10. 

1 1 Shell sub-oblong, short, corneous-brown, obliquely wrinkled 



HAWAIIAN NESOPUPAE. 319 

with close and somewhat rough growth riblets, minutely per- 
forate, scarcely shining, somewhat thin. Spire obtuse, shortly 
subcylindrical but yet slightly conic-tapering. Whorls 5, con- 
vex, regularly increasing, suture impressed; the last shortly 
ovate, swollen, somewhat saccate, hardly ascending in front, 
not laterally compressed, rotund about the umbilicus. Aper- 
ture slightly oblique, not sinuate, truncate-subcircular, hardly 
effuse, furnished with tooth-like lamellae, as follows : two parie- 
tal, one of which, the angular, is small tubercle-like, and the 
second, parietal, median or nearly median, is elongate, more 
deeply seated, ending abruptly. An acute, small columellar; 
and 2 short, nearly equal palatals, slightly elongate, parallel. 
Peristome narrowly expanded and thickened on all sides, 
slightly more dilated at the columella, livid, with quite dis- 
tant margins. Length 2, diam. 1.33, apert. 0.75 mm." (Ancey). 

Kauai : Kipu (type loc.), Kipukai, Koloa, Halemanu, Haena, 
Wainiha and Kalalau (Cooke) ; Nounou Mts. (Dole). Type 
18700 Bishop Museum; topotypes 20733 Bishop Museum and 
44724 A. N. S. P. 

Nesopupa Kauaiensis ANCEY, Proc. Malac. Soc., London, vi, 
June 1904, p. 124, pi. 7, f. 17. 

The type specimen measures, length 2.15, diam. 1.4, apert. 
(diag.) 0.86 mm. Interstices of costas granulose, the granules 
arranged in rows, costoe granulose ; apical whorls eroded, the 
exact number could not be made out. 

The original specimens of this species were collected on 
the Kupu side of Haupu. Specimens in the Bishop Museum 
from near the original locality agree very closely with Ancey 's 
type now in the Bishop Museum. One of these specimens, 
carefully compared with the type, has 4% whorls, the color is 
argus-brown. The costae are lower, closer and more evenly 
spaced than in typical P. newcombi; their edges only very 
narrowly membranous. The embryonic whorls are distinctly 
granulose when viewed under a strong lens; the angular 
lamella is short, punctiform and seated close to the margin 
of a rather thick parietal callus. The parietal lamella is 
shorter than that of P. newcombi, deeply seated and nearly 
perpendicular to the parietal wall. The columellar lamella 



320 HAWAIIAN NESOPUPAE. 

is rather short, thickened and low; the two palatal are short, 
nearly parallel, the lower nearly twice as high and long as the 
upper. An average specimen measures : length 2.1, diain. 1.4, 
greatest length apert. 0.86 mm. At a somewhat higher ele- 
vation the shells are slightly larger. One such measures: 
length 2.6, diam. 1.55, apert. 1.04 mm. 

On the same ridge, but nearer the sea and at a slightly 
lower station, the shells are much smaller. Among thirty 
examples taken at random, 12 had the five normal lamellae 
in the aperture; in 18 there was in addition a more or less 
minute basal fold. In this small race of kauaiensis the speci- 
mens at first glance have the appearance of typical newcombi. 
They are most easily recognized by the less expanded and 
more heavily thickened peristome and the lower and less mem- 
branous ribs, of which there are about 30 on the last whorl. 
A specimen of this small race (p. 309, f. 9) measures: length 
1.63, diam. 1.16, aperture 0.73 mm. ; 4% whorls. This race is 
widely distributed on Kauai, in rather damp situations on 
rocks. 

18. NESOPUPA SINGULARS C. & P., n. sp. PI. 29, fig. 8. 

The shell is minute, perforate (perforation very minute, 
nearly circular) ovately conic, with a short base, clay color, 
thin, translucent, rather dull, the surface rather closely ribbed- 
striate, a few of the striae, especially on the back of the last 
whorl (rather distantly and more or less evenly spaced) are 
more strongly developed and furnished with membranous 
edges; under a microscope the whole surface covered with 
very minute wrinkles. Spire with convex outlines tapering 
gradually to the semi-globular apex; individual whorls very 
convex, separated by a deep suture. Whorls 4% ; the em- 
bryonic minutely granulose, the granules more or less ar- 
ranged in transverse rows, the striae beginning gradually 
at about 1% whorls. The last whorl is sub-saccate, rounded 
about the umbilicus, the last % nearly straight. Aperture 
hardly oblique, nearly perpendicular, almost a complete circle, 
lip-insertions nearly united, connected by a rather thick, 
short white callus. Angular lamella usually absent, sometimes 



HAWAIIAN NESOPUPAE. 321 

indicated by a low indistinct swelling ; parietal lamella strong, 
deeply seated, slanting outwards; columellar lamella strong, 
deeply seated, slanting downwards, the inner end abruptly 
turned upwards, following the columella as a thickened 
rounded ridge; two palatal folds rather short, parallel when 
seen through the shell from the back, rather remote from the 
peristome, of almost equal size. Peristome with a uniform 
whitish thickening within, the margin expanded, thin, of the 
<?olor of the shell. Length 1.57, diam. 0.9, aperture (diag.) 
0,58 mm. 

Oahu : Kaliuwaa, on the rocky walls of the canyon, near the 
waterfall (Cooke) ; West Maui ; Honokahau (Forbes). Type 
11077 Bishop Museum; paratypes 19842 Bishop Museum and 
44697 A. N. S. P. 

This species is separated from newcombi by its narrower 
outline, lighter color, circular mouth and the almost complete 
absence of an angle lamella. It is rather remarkable that 
the only two known colonies of this species should be from 
different islands. The species seems to be abundant where 
found, as 36 specimens were taken at Kaliuwaa and 13 at 
Honokahau. The Maui specimens are slightly larger with 
more convex outlines, and are darker-colored, with slightly 
more numerous and stronger developed membranous ribs than 
those from Oahn. One measures: length 1.56, diam. 0.95, 
apert. (diag.) 0.67 mm.; 4% whorls. 

19. NESOPUPA ALLOIA C. & P., n. sp. PI. 29, fig. 10. 

The shell is very minute, perforate, shortly ovate, isabella 
uolor, thin, slightly translucent, scarcely shining, marked with 
thin, distant rather high membranous costee (about 19 on the 
last whorl), the inter-costal spaces minutely granulose- 
wrinkled. Spire short, summit convely conic, the individual 
whorls convex, separated by a deep suture. Whorls 4, the 
first minutely granulose and very faintly spirally striate, the 
oblique, delicate riblets beginning at about the middle of the 
second whorl; the penult whorl is closely costulate, swollen; 
the last whorl convex, tapering towards the base, scarcely 
ascending near the aperture. Aperture rather large, truncate- 



322 COCOS ISLAND NESOPUPAE. 

oval, hardly oblique, slightly diagonal, lip-insertions distant,, 
hardly converging. There is no trace of an angular lamella ; 
parietal lamella very strong, upright, oblique, slanting out- 
ward; columellar lamella rather strong, deeply seated, hori- 
zontal, and perpendicular to the inner margin of the colu- 
mella; lower palatal fold short, lamella-like, rather deeply 
seated; upper palatal nearly equal and parallel to the lower, 
but not so deeply placed. Peristome expanded, its outer mar- 
gin very thin, membranous, the inner margin slightly thick- 
ened. Length 1.4, diam. 0.98, apert. (diag.) 0.62 mm. 

Kauai: Hanapepe falls, on Trichomanes (Heller). Type 
11078, paratype 20176 Bishop Museum. 

This most interesting little species was found by Mr. Forbes 
on mounted fern specimens collected by Heller, and now in 
the Bishop Museum herbarium. It is represented by two speci- 
mens which agree very closely, though one of them is slightly 
broken near the summit. This species was at first considered 
a not quite fully developed P. thaanumi. Comparison with 
nearly adult specimens of thaanumi, shows that the angular 
lamella is present in that species before the peristome is at 
all thickened. P. alloia is easily distinguished from P. singu- 
laris by its blunter summit, shorter and broader outline and 
fewer whorls. 

II. Cocos ISLAND SPECIES. 
Section Cocopupa P. & C., n. sect. 

The shell is minutely pitted with very little striation, the 
teeth typical for Nesopupa except that the inner end of the 
columellar lamella does not turn downward; last whorl not 
distinctly furrowed back of the lip. 

Cocos Island, in the Pacific about 550 miles southwest of 
Panama, has several species which have been referred to 
Ochroderma, a genus of the Caroline Islands ; but this relation- 
ship has not been fully established by anatomical comparison 
(see Vol. XVIII, p. 325). In the Galapagos Islands, however, 
there is a truly Polynesian snail, Tornatellides chathamensis 
(Vol. XXII, 201) ; so that the occurrence of the Polynesian 



COCOS ISLAND NESOPUPAE. 323 

group Nesopupa at the eastern rim of the Pacific is not un- 
precedented. 

20. NESOPUPA COCOSENSIS (DalL). PL 30, figs. 10, 11. 

"Shell small, reddish brown, rather pointed, with nearly 
five rounded whorls; apex paler, polished, rather blunt; last 
whorl much the largest; base rounded with a well-marked 
umbilical pit; aperture wider behind, the posterior part of 
the outer lip and the pillar lip broadly reflected, the anterior 
outer and basal margin narrow ; the pillar and outer lip united 
by a thin callus ; lamellae according to Sterki 's formula .ABDE, 
comprising one columellar and two parietal folds, and, on the 
outer wall well within the peristome, two narrow little-elevated 
ridges, of which the anterior is shorter. Axial length of shell 
2.2, of aperture .8, width of last whorl 1.5, of aperture .8 mm." 
(Dall.) 

Cocos Island, on leaves (Heller and Snodgrass). 

Vertigo cocosensis DALL, Proc. Acad. Nat. Sci. Phila. 1900, 
p. 98, pi. 8, f. 13. 

"The chief peculiarity of this species is that the surface, 
which looks silky under an ordinary hand lens, when more 
magnified is seen to be punctate all over, recalling V. variolosa 
Gould, of Florida, which, however, differs in form, size and 
dental armature" (DaM). 

By the color, sculpture and teeth this species appears most 
nearly related to those of Polynesia, but differs by the simpler 
columellar lamella, which does not curve down at its inner end. 
It is the only member of the genus found on the American 
side of the Pacific. 

The shell is ovate, deeply rimate, tawny, when not dulled by 
age, with a brighter cinnamon-rufous lip; densely and very 
minutely pitted, and with weak striae of growth. The last 
whorl has a slight impression over the upper palatal fold. 
The teeth are grayish-white. The angular lamella curves out- 
ward and in old individuals joins the termination of the outer 
lip, as in fig. 11. The high parietal lamella is somewhat 
curved. The columellar lamella slants downward somewhat, 
enters horizontally and is not turned down at the inner end. 



324 POLYNESIAN NESOPUPAE. 

The two palatal folds are well developed, not very long, either 
subequal, or the upper somewhat shorter ; the lower decidedly 
further in. The peristome is expanded and in old shells 
thickened. Its outer border is bent in a little in the region 
of the upper palatal fold. Parietal callus thin. 
Length 2.2, diam. 1.35 mm. ; 5 whorls. 

III. POLYNESIAN, MICRONESIAN AND MELANESIAN SPECIES. 
Section Nesopupa proper. 

Nesopupae with the surface more or less pitted or pitted- 
granose, striae with membranous edges often present. Angu- 
lar and parietal lamellae strong and long, the former curving 
outward to join the end of the outer lip; inner end of the 
columellar lamella bent downward ; palatal folds present. 

Nesopupae of the typical group inhabit high islands of all 
Polynesia and Micronesia, the East Indies and Mascarene 
Islands, probably also Melanesia, having thus a very wide 
distribution. 

All of the Polynesian species were considered varieties or 
subspecies of N. tantilla by Dr. Boettger, whose revision is 
the main work on them. In the absence of evidence of inter- 
gradation it seems more logical to recognize the various island 
forms as species. No doubt many more remain to be dis- 
covered, as no adequate search has been made for the minute 
shells of Polynesia. Probably arboreal species will be found, 
as in Hawaii. 

(Polynesia.) 

Beck mentioned two undescribed species. Alcua pitca-irnensis 
Beck, and Alaea millium Beck, Index Molluscorum 1837, p. 85. 
Pitcairn Island. Names and locality only. 

21. NESOPUPA TANTILLA (Gould). PL 30, figs. 1 to 4. 

Shell very minute, of an oval form, obtuse at summit, and 
composed of about four convex whorls, of which the upper ones 
are irregularly marked with delicate longitudinal bars, and 
the lower one, under a magnifier, appears to be roughened by 
minute, irregular granulations. Color whitish. Aperture 



POLYNESIAN NESOPUPAE. 325 

somewhat four-sided, rounded at the corners, the lip slightly 
expanding and the throat armed with five teeth, one on the 
columella, one at the base, one on the outer lip, and two un- 
equal approximate ones on the middle of the transverse lip. 
There is a small umbilical fissure. Length 1/15, width 1/20 
inch (Gould). 

Society Islands: Tahiti at 2000 ft. (Couthouy, U. S. Ex- 
ploring Exped.) Tahiti, Huaheine, Borabora and Maupiti 
(Garrett). Also reported from the Marquesas, Paumotu 
Archipelago, and as far west as Viti, but these forms require 
critical comparison. 

Pupa (Vertigo) tantilla GOULD, Proc. Bost. Soc. N. H. ii, 
1847, p. 197; Expedition Shells p. 33. Vertigo tantilla 
GOULD, U. S. Expl. Exped., xii, Mollusca and Shells, 1852, p. 
92, pi. 7, f. 105, #, &.- GARRETT, Journ. Acad. Nat. Sci. Phila. 
viii, 1881, p. 400 (Cook's or Harvey Islands) ; ix, p. 84; Bull. 
Soc. Malac. France, iv, 1887, p. 34 (Marquesas). Pupa tan- 
tilla Gld., PPR., Mon. Hel. Viv. iii, 557. BOETTGER, Conchol. 
Mittheil. i, 1881, p. 49, pi. 10, f. 1. "P. pazi Crosse" Ancey, 
Bull. Soc. Malac. France, iv, 1887, p. 35, footnote. 

This senior member of the group differs from its allies by 
the smaller number of teeth, 5, or when an interpalatal is 
developed, 6. The inner end of the columellar lamella turns 
down less than in related forms. 

The type specimen, 5505 U. S. N. M., labelled as the original 
of Gould's figure 103, is drawn in pi. 30, figs. 1, 2, 3. The sur- 
face is minutely confluent-pitted or pox-marked with traces of 
striae as usual in the Polynesian group. The last whorl has a 
rather short and shallow furrow over the upper palatal fold. 
Angular lamella high, straight within, then well curved out 
to join the lip. Parietal lamella curved. The columellar 
lamella is horizontal, its inner end curving downward, but far 
less than in N. pleurophora or armata. The upper and lower 
palatal folds are rather short, subequal, and between them a 
small interpalatal stands. It is hard to see, and evidently was 
overlooked by Gould; (but in other specimens it is wanting). 
There is no basal fold or subcolumellar lamella, but in oblique 
view a very weak, hardly noticeable trace of a subcolumellar 



326 POLYNESIAN NESOPUP^E. 

tubercle may be seen. The peristome is thickened a little, 
well expanded. 

Length 2.25, diam. 1.3 mm.; nearly 5 whorls (type). 

Length 2, diam. 1.35 mm. 

G-arrett, in his several lists, included dunkeri, armata, 
pleurophora and dentifera as synonyms of tantilla, which he 
says ranges from the Society to the Viti Islands, and is found 
beneath rotten wood, under stones, and among decaying leaves. 
Boettger gives tantilla an equally wide range, but he distin- 
guishes 5 varieties and 2 subspecies. In the absence of inter- 
grading individuals it appears more logical to rank part of 
these as species. Typical tantilla is at present known posi- 
tively from Tahiti only. Mr. Ancey mentioned a P. pazi 
Crosse, stating that it does not differ specifically from tantilla ; 
but I cannot find that Crosse described such a species. 

22. NESOPUPA PLEUROPHORA (Shuttleworth). PL 30, figs. 5, 6, 

Shell minute, rimate-perforate, shortly ovate, thin, slightly 
striate, beautifully ribbed with very thin, widely-spaced 
lamella?, sometimes vanishing, rufous chestnut, somewhat pel- 
lucid, slightly glossy. Spire rounded, obtuse. Whorls 5, 
strongly convex and somewhat inflated, the last rounded at 
the base. Suture deep. Aperture rounded auriform, with 
seven folds : angular lamella strong ; parietal strongly ele- 
vated, somewhat immersed ; two columellar lamellae, the upper 
strong, lower small ; three immersed palatal folds, the middle 
one smaller. Peristome expanded, callous-lipped, whitish, 
right margin angularly sinuous above, somewhat tuberculate 
below the sinus. 

Length about 2, diam. 1%, aperture % mm. (ShuttL). 

Marquesas Islands and Tahiti (Verreaux). 

Pupa pleurophora SHUTTLEWORTH, Bern. Mittheil., 1852, 
p. 296. PPR., Monogr. Hel. Viv., iii, 560. [Pupa tantilla] 
var. pleurophora Shuttlw., BOETTGER, Conchol. Mittheil., i, 
1881, p. 51, pi. 10, f. 3. Pupa dunkeri ZELEBOR, in Pfr., 
Monographia, vi, 1868, p. 333 (Taheiti). 

Tahiti may be selected as the type locality. Under the 
microscope the surface is irregularly granose or vermiculate- 



POLYNESIAN NESOPUP^E. 327 

granose, more or less noticeably striate, with sparse traces of 
cuticular riblets in some specimens, probably well developed 
in fresh shells. There are 8 teeth: angular and parietal 
lamellae strongly developed, columellar strong but deeply 
placed, turned down at its inner end ; there is a subcolumellar, 
usually invisible in direct front view; upper and lower pal- 
atals are subequal, not very long, and converge somewhat in- 
wardly; interpalatal smaller; and there is a small tubercular 
suprapalatal. 

Length 1.9, diam. 1.15 mm. 

Length 1.8, diam. 1.1 mm. 

It differs from N. tantilla by the smaller aperture, the more 
numerous teeth and by having the inner end of the columellar 
lamella turned more strongly downward, much as figured for 
N. armata. Still, it may prove to be a race of that species. 
N. armata has a much larger upper palatal fold, and appears 
to be quite distinct. 

Pupa dunkeri Zelebor, whose MS. description was printed 
by Pfeiffer, is supposed to be identical with pleurophora. 
The description follows. 






P. Dunkeri Zelebor. T. dextrorsum convoluta, ovato- 
cylindrica, apice obtuso, umbilici rima capaci, crassa, striis et 
granulis tan turn oculo armato conspicuis holosericeo-splendens, 
purpureo-fusca aut flavescente-olivacea ; spira sensim incres- 
cente, anfr. quatuor vel quinque valde convexis sutura pro- 
funda conjunctis; apertura semiovata plicis septem vel novem 
coarctata; plica magna et altera in radio, minima et duabus- 
magnis incrassatis profunde intrantibus, quae cum aperturae 
margine conjunctae valde prominent, superiore aperturae 
pariete, quae cum ora, vix conspicua dentiformi in pullis de- 
ficiunt in toro aperturae, collocatis; margine aperturae medi- 
ocre incrassato rotundato paulo reflexo, non conjuncto, rufes- 
cente-fusco, in speciminibus clarioribus obscure rufescente- 
violaceo. Alt. 2, lat. iy 8 mill. Habitat in insula Taheiti." 
(Zelebor in Pfr.) 

23. NESOPUPA ARMATA (Pease). PL 30, figs. 9, 12, 13. 

The shell is ovate, cinnamon-brown, darker behind the lip, 
with a tawny peristome and not much gloss. The surface is 
densely pitted or vermiculate-wrinkled, with irregular oblique 



328 POLYNESIAN NESOPUP.E. 

striation, which is most noticeable on the penult and next 
earlier whorls. The whorls are strongly convex, the last 
having a long but not very deep impression behind the lip, 
descending obliquely from the upper palatal position. The 
peristome is expanded and heavily thickened within, ex- 
cavated at the sinulus. The angular and parietal lamellae 
are much longer than in N. tantilla but similar in shape, the 
angular being straight within, then curving out to join the 
lip, the parietal well curved and entering deeply. There is a 
well-developed infraparietal lamella. The columellar lamella 
is deeply placed, ascending slowly within, then abruptly bent 
downward along the axis (fig. 9). Below it a tubercular 
subcolumellar nodule projects; this is probably a displaced 
basal fold. Within the outer lip there is a thin, high, diag- 
onally descending upper palatal fold and a small, tubercular 
suprapalatal. The lower palatal fold is immersed, basal in 
position, straight, and shorter than the upper. 

Length 2.2, diam. 1.3 mm. ; 5% whorls. 

Society Is.: Borabora (Garrett). 

Vertigo armata, PEASE, Proc. Zool. Soc. Lond., 1871, p. 461. 
Pupa armata Pease, PFEIFFER, Monographia, viii, p. 407 
(copied from Pease). 

Described from the type, figured from the type and another 
of the type lot, no. 48315 M. C. Z. It is well distinguished 
by the oblique, lamellar upper palatal fold, the wholly basal 
lower palatal, the long lamellae of the parietal wall and 
numerous teeth, 8 in all. Fresh specimens evidently have 
cuticular laminae on the striae. Pease says "interdum re- 
mote, tenuiter filocostata . . . filis albidis"; but these have 
been worn from the type specimens. N. tongana (Bttg.) ap- 
pears from the description to be somewhat similar. 

24. NESOPUPA PAIVAE (Crosse). PL 30, figs. 7, 8. 

"Shell not very deeply umbilicate, ovate-ventricose, thin, 
subpellucid, pale chestnut; spire with obtuse somewhat flat- 
tened apex. Whorls 41/2, very convex, the embryonic I 1 /! 
smooth, whitish, flattened, the rest ventricose, impressed with 
rather oblique, somewhat distant striae, easily seen without a 



POLYNESIAN NESOPUP^E. 329 

lens, the last whorl ascending, about equal to the spire, 
rounded-tapering at base. Aperture truncate-rounded, vio- 
laceous-fleshy, narrowed by 9 teeth: on the parietal wall 3, 
the first rather large, somewhat projecting forward and con- 
tinuous with the lip, the second largest, entering, the third 
minute; on the columellar margin 2 unequal teeth within; 
in the basal and outer margins 4 deeply placed teeth. Peri- 
stome somewhat expanded, reflected, the margins separated, 
columellar margin spreading, broadly expanded, the middle 
of the outer margin more or less bent in. 

"Length 1.75 to 2, diam. 0.75 mm." (Crosse). 

Gambier Islands (Crosse) ; Mangareva (Pease). Paumotu 
Group (Bttg.). 

Pupa paivae CROSSE, Journ. de Conchyl., xiii, 1865, p. 218, 
pi. 6, f. 5. PFEIPPER, Mon. Hel. Viv., vi, 333. Vertigo paivae 
Cr., PEASE, P. Z. S., 1871, p. 474, Mangareva. [Pupa tan- 
tttla] var. paivae Crosse, BOETTGER, Conchol. Mittheil., i, p. 
50, pi. 10, f. 2. 

Boettger, who had specimens (pi. 30, figs. 7, 8) given by 
Crosse to Pfeiffer, notes the discrepancy between Crosse 's 
term ovato-ventricosa and his measurement of diameter, % 
mm. He notes the relationship to pleurophora "so great 
that a specific separation from this, and therefore also from 
typical P. tantUla Gould, is evidently not to be thought of." 
He gives the following comparative description : 

Shell rather thick, pale brown, more distinctly costulate; 
last whorl more tapering and anteriorly dilated funnel-like. 
Aperture generally 7-toothed : 1 angular, 1 or 2 parietals, the 
inner one smallest, often wanting; columellars 2, the lower 
one small; palatals 3, equidistant and subequal. Peristome 
generally discontinuous, somewhat thickened. Length 1% to 
2, width iy 8 to 11/4 mm. 

25. NESOPUPA DENTIPERA (Pease). PL 30, fig. 14. 

"Shell cylindric, rather solid, perforate, dextral. longitu- 
dinally obliquely very finely striate, rufous- or yellowish- 
chestnut. Whorls 4, convexly rounded, the last shortly bi- 
sulcate behind the lip. Aperture nearly circular, biplicate 
posteriorly, columella 1-plicate, base 3 or 4-toothed; lip 



330 POLYNESIAN NESOPUP^E. 

slightly reflected, peristome a little thickened, not continuous. 
Alt. 1.75, diam. 1 mm. (Pse.). 

Cook or Hervey Is.: Aitutaki, in forests near the seashore 
(Garrett). 

Vertigo dentifera PEASE, Proc. Zool. Soc. London, 1871, p. 
462. Pupa dentifera Pse., PFEIFFER, Monogr., viii, p. 408. 
[Pupa tantilla] var. dentifera Pease, BOETTGER, Conchol. 
Mittheil., i, p. 52, pi. 10, f . 4. Vertigo tantiUa GARRETT, 
Journ. A. N. S. Phila., viii, 401. 

The original locality given by Pease, * ' Roratonga, ' ' was an 
error on the part of Pease. Garrett, who supplied the types 
to Pease, states that he collected them on Aitutaki. 

The shell is oblong, somewhat cylindroid, cinnamon-brown , 
opaque, with little gloss. Surface is densely pitted, with un- 
evenly developed oblique striae or traces of them, strongest 
below the suture. Whorls convex, the last having a long, 
somewhat oblique furrow in the upper palatal position, ter- 
minating at the lip. The peristome is well expanded and 
thickened within except at the sinulus, colored like the out- 
side. The rather long angular lamella is straight within, 
anteriorly curving outward and joining the lip some distance 
from its insertion. Parietal lamella about as long as the an- 
gular, more immersed, somewhat curved. The columellar 
lamella is deeply placed, small in a front view; below it is a 
small immersed subcolumellar lamella, visible in an oblique 
view. There are three rather short, deeply placed palatal 
folds, the interpalatal a little shorter, the lower palatal nearly 
basal in position. The upper palatal, foreshortened in the 
figure, is moderately long, laminiform. 

Length 2, diam. 1.15 mm. ; length of aperture 0.8 mm. ; 5 
whorls. 

This description and the figures from the two cotypes of 
the Pease collection, no. 48314 Mus. Comp. Zool. The shells 
were quite dirty, this doubtless accounting for the discrepan- 
cies between Pease's description and that now given. 

26. NESOPUPA GODEFFROYI (Boettger). PL 31, figs. 13, 14. 
"Shell smaller [than tantilla Gld.], more conic-ovate, the 



POLYNESIAN NESOPUP^. 331 

last whorl lower. Aperture smaller, semioval, 8-toothed, the 
teeth stronger, white; 1 prominent, angular lamella, arc- 
uately connected with the peristome; 2 parietals, the outer 
one curved below, the interior very small; 2 columellar la- 
mellae, compressed, high; 3 rather deeply placed palatals, 
equidistant and subequal. Peristome discontinuous, whitish- 
lipped, somewhat thickened at the receding sinulus, the middle 
of the outer margin produced forward. 

Length 1.75, diam. 1 mm." (Bttg). 

Samoa Islands. 

[Pupa tantilla] var. godeffroyi BTTG., Conchologische Mitt- 
heil., i, p. 53, pi. 10, f . 5. ? V. tantilla Gould, GABRETT, 
Proc. A. N. S. Phila., 1887, p. 138 (Samoa Is.). 

Although showing much peculiarity, this form may be 
viewed as a variety of P. tantilla Gld., as it approaches it 
very nearly in the dentition, and by its shell characters it 
may be recognized almost with certainty that a close blood 
relationship unites it to that species (Bttg.). 

27. NESOPUPA TONGANA (Boettger). PI. 31, figs. 15, 16, 17. 

Shell more conic [than tantilla] the whorls more distinctly 
and generally more distantly membranous-ribbed, the last 
with protracted base and anteriorly more distinctly dilated 
funnel-like, in the middle having a long longitudinal furrow. 
Aperture a little larger, always oblique, rhomboidal, little 
compressed basally, 6 to 8-toothed : one emerging sigmoid 
angular ; the rest of the teeth deeply placed, 2 parietals, 1 or 
2 columellars, 2 or 3 palatals, of which the upper is much the 
largest, lamelliform, inside approaching close to the larger 
parietal lamella, and entering deeply with it; the lower 1 or 
2 palatals small and very deeply placed. Peristome brownish 
lipped, somewhat thickened below the sinulus, the margins 
joined by a thin callus. 

Length 2i/ 8 to 2%, width 1% mm." (Bttg.). 

Tonga group : Tonga Tabu. 

Pupa tongana O. Semper in Museum Godeffroy Catal., v, 
1874, p. 89 (name and locality Tongatabu only). [Pupa tan- 
tilla} var. tongana 0. Semper, BOETTGER, Conchol. Mittheil., 
i, 1881, p. 54, pi. 10, f. 6 and 11, f. 7. 



332 POLYNESIAN NESOPUP^E. 

Dr. Boettger remarks that this gives the impression of a 
separate species, mentioning the long, deeply penetrating 
upper palatal fold and the funnel-like extended aperture; 
yet he ranks it as a subspecies of tantitta on account of the 
analogous upper palatal of tantilla var. dentifera Pse., and 
the near agreement in form of the last whorl in tantilla subsp. 
vitiana. 

28. NESOPUPA VITIANA (Boettger). PL 31, figs. 18 to 23. 

"Shell more solid [than tantilla Old.], more globulose, the 
spire almost exactly conic, the apex always more acute. Rib- 
lets more delicate and less distant. Otherwise it is more like 
subsp. tongana Semp. in the last whorl and aperture, but 5 
to 6-toothed : 1 angular, 1 parietal, 1 or 2 columellar lamellae, 
2 or 3 palatal folds, the upper one hardly lamelliform and 
not larger than the lower. Peristome with the margins either 
separated or joined by a very thin callus, distinctly labiate, 
a little thickened below the sinulus (Boettger). 

"Length 1.75 to 2, width 1.25 to 1.4 mm. Oneata; pi. 31, 
f. 18-21. 

"Length 1.75, width scarcely 1.25 mm. Viti Levu; pi. 31 y 
f. 22, 23. 

Viti group : Oneata, Vanua Balava, Kanathia and Viti 
Levu (Godeffroy Mus., and according to Alb. Mousson). 

[Pupa tantilla] subsp. vitiana BOETTGER, Conchpl. Mitt- 
heil., i, p. 55, pi. 11, f. 8-10. Pupa tantilla Gould, MOUSSON, 
Joum. de Conchyl., 1870, p. 127. ? [Vertigo tantilla] var. 
oneatensis of Boettger, according to ANCEY, Bull. Soc. Malac. 
France, iv, 1887, p. 34, footnote (Oneata, Viti). 

What appears chiefly to characterize the Viti Island form 
is the constant, ventricose-conic shape of the shell, which of 
course differs quite conspicuously from the purely ovate shape 
of typical P. tantilla Gould of Tahiti (Boettger). 

Ancey says that Boettger described a var. oneatensis of 
Vertigo tantilla, the type being in his collection ; but no such 
description exists to my knowledge, arid the form referred to 
is probably vitiana. 






MELANESIAN NESOPUP^. 333 

(Species of Melanesia.) 

Alaea annaensis Beck, Index Molluscorum, 1837, p. 85. I. 
Annaa. No description. 

Pupa condita n. sp., with. var. &, zonata, of Gassies, Journ. 
de Conch., xvii, 1869, p. 73, Art Island,' New Caledonia. This 
is said by Crosse (Journ. de Conchyl., 1894, p. 303) to be a 
marine shell of the genus Rissoa, in poor condition. 

29. NESOPUPA NORFOLKENSIS (Sykes). PI. 31, fig. 26. 

Shell subperforate, pyramidate-ovate, yellowish, somewhat 
shining, obliquely striate, apex rather obtuse. Whorls 5%, 
convex, the last (with the peristome) about half the length 
of the shell. Aperture ovate, 7-toothed: two parietal teeth, 
2 columellar teeth, 3 palatals, the upper one smallest. Peri- 
stome white, a little thickened, somewhat reflected, the outer 
margin forming an obtuse angle. Length 4, maximum diam. 
2.3 mm. (Sykes). 

Norfolk Island, in the crevices of the back of a Dracena 
(Brit. Mus.). 

Vertigo norfolkemis SYKES, Proc. Malac. Soc. London, iv, 
Oct., 1900, p. 145, pi. 13, f. 3. 

An interesting little form with tumid whorls. Six of the 
teeth are of about the same size, but the lower parietal and 
upper palatal are very small (Sykes). 

The figure shows a sinistral and rather strongly sculptured 
shell, with the lower (parietal) lamella longer than the upper 
(angular). 

30. NESOPUPA LIPOUANA (Gassies). PL 31, fig. 25. 

The shell is widely umbilicate, very small, finely and reg- 
ularly striate, shining, of reddish corneous, uniform color 
with an uneven zone of reddish-brown at the periphery of the 
last whorl. Spire conic-cylindric, with the summit obtuse, 
mamillate. Suture crenulate and rather deep. Whorls 6 to 
6!/2, convex, the last whorl forming one-third the total length. 
Aperture oblique, oval, contracted at base, having 4 folds and 
lamellae arranged as follows: one on the columella, descend- 



334 MELANESIAN NESOPUP^E. 

ing and entering deeply; two parietal, the first very small, 
running inward and descending, the second very large, 
placed far forward, projecting beyond the peristome, a little 
oblique towards the right side but perpendicular to the axis. 
Finally, the last tooth is placed within the base of the lower 
margin, and is a lamina which runs from right to left. The 
peristome is thick, reflected, excavated at the upper right 
margin opposite the large fold which projects beyond it, and 
which causes the peristome to form a quite distinct callous- 
projection in % of the right margin. 
Length 4, diam. 2^/2 mm. (Gassies). 

Loyalty group : Lifu. Lives under ferns and in the moss 
(R. P. Lambert). 

Pupa lifcwana GASSIES, Fauna 'Conch. N. Caled., part 2, 
1871, p. 98 ; Journal de Conch., xxi, 1873, p. 53, pi. 2, f. 8. 
SOWERBY, Conch. Icon., pi. 4, f. 28. PPEIPFEB, Mon. Hel. 
Viv., viii, 376. 

31. NESOPUPA MARIEI (Crosse). PL 31, fig. 24. 

Shell narrowly perforate-rimate, subglobose, ventricose, 
rather thin, subtranslucid, seen under a strong lens to be 
very delicately, subobliquely striatulate, fleshy-buff, unicol- 
ored. Spire moderately elevated, the apex obtuse. Suture 
impressed. Whorls 4%, convex, the last scarcely one-third 
the length, slightly contracted near the aperture. Aperture 
subvertical, lunate-rotund, colored like the outside. Peri- 
stome very strongly thickened, reflected, violaceous; the colu- 
mellar margin short, the basal rounded, the outer slightly 
sinuous, towards the insertion somewhat emarginate, taper- 
ing, somewhat scrobiculate externally. Parietal margin with 
two minute teeth, the larger near the insertion of the outer 
lip, the other more deeply placed, not conspicuous. 

Length 2, diam. scarcely 1.5 mm.; aperture 1.5 mm. long, 
scarcely % mm. wide (Crosse). 

New Caledonia: near Noumea (E. Marie). 

Pupa mariei CROSSE, Journ. de Conch., xix, 1871, p. 202;. 
xx, 1872, p. 358, pi. 16, f . 3 ; xlii, 1894, p. 303. 



MICRONESIAN NESOPUP^E. 335 

(Species of Micronesia.) 

32. NESOPUPA QUADRASI (Mlldff.). PI- 32, figs. 7, 8, 9, 10. 

Shell subperforate, oblong-ovate, thin, sculptured with very 
delicate, rather widely-spaced riblets, silky, brown. Whorls 
5, convex, slowly increasing, separated by a deep suture, the 
last slightly ascending in front, compressed at base, distinctly 
pitted behind the lip. Aperture nearly vertical, rounded- 
triangular, the peristome well expanded, with a brown thick- 
ening, the margins converging, right margin deeply sinuate, 
with a noduliform callus. Angular lamellar rather high, 
curving outward, and together with the nodule of the external 
lip forming an elliptical sinulus; 2 parietal lamellae, one 
strongly elevated, entering deeply, the other smaller; 2 colu- 
mellar lamellae; 3 deeply placed palatal plicae, of which the 
right one is rather long and lamelliform. 

Length 1.9, diam. 1 mm. (MUdff.}. 

Length 1.75, diam. 1.05 mm., aperture 0.8 mm. 

Marianne Islands: Guam (Quadras). 

Vertigo (Ptychochilus) quadrasi MOELLENDORFF, Nachr. d. 
Malak. Ges., xxvi, 1894, p. 17. 

This species is closely related to N. moreleti, but it differs 
by the presence of a small interpalatal fold, or in one speci- 
men two, and the subcolumellar lamella which is rather deeply 
placed (and may be a displaced basal fold). There is some- 
times a suprapalatal fold, as in N. moreleti, these specimens 
having 9 or even 10 teeth. The cuticular laminse are often 
fugacious, and the surface then is somewhat pitted. 

This species has been distributed by dealers under the un- 
published name Pupa marianarum Q. et M. 

33. NESOPUPA EAPENSIS (Boettger). PL 31, figs. 27, 28. 

"Differing from P. tantilla, Gould by the perforate-rimate 
shell, oblong-ovate, the whorls more rapidly increasing, less 
convex, the last nearly half the total length, hardly impressed 
externally towards the aperture. The aperture is larger, 
rather high, truncate-subovate, 5-toothed, the teeth not very 
strong; angular small, rather deeply placed, scarcely con- 



336 EAST INDIAN NESOPUP^E. 

nected with the margin; 1 parietal, 1 columellar, 2 palatals, 
opposed cross-like to the others, not very deeply placed. 
Peristome widely separated, a little reflected, somewhat thick- 
ened within, the outer margin not thickened at the sinulus 
and scarcely sinuate. 

" Length 2, width 1% mm." (Bttg.). 

Caroline Islands: Yap (Mus. Godeffroy). Pelew Islands 
(Kubary). 

Pupa eapensis BTTG., in Conchologische Mittheilungen, 
1881, p. 56, pi. 11, f. 11. Vertigo (Ptych&chH/iis) eapensis 
(Bttgr.), MOELLENDORFF, Journ. of Malac., vii, 1900, p. 112. 

34. NESOPUPA PONAPICA (Mlldff.). 

Shell perforate-rimate, ovate-oblong, delicately and dis- 
tantly costulate, silky, brown. Spire subcylindric below, 
conoid above, the apex obtuse. Whorls 5, moderately convex, 
the last somewhat compressed at base, with a deep, long ex- 
ternal impression behind the aperture. Aperture nearly ver- 
tical, rounded-trapezoidal, the peristome moderately ex- 
panded, having a reddish lip, externally distinctly sinuate, 
the margins joined by a very thin callus. Teeth 6, the colu- 
mellar receding; palatals 3, the lower two deeply placed, 
short, the third long, strong ; angular lamella strong, entering 
deeply ; parietal remote from the margin, approaching the 
angular. 

Length 1.75, diam. 1 mm. (Mttdff.). 

Caroline Islands: Mpomp, Ponape (Kubary). 

Vertigo (Ptychochilus) ponapica MOELLENDORFF, Journal 
of Malacology, vii, March, 1900, p. 113. 

* ' This minute shell belongs to the group of V. tant&la Gld., 
and might be considered, like the preceding form [eapensis], 
to be merely a representative subspecies of that species, 
widely distributed over Polynesia" (Mlldff.). 

IV. NESOPUP^E OF THE BAST INDIES AND PHILIPPINES. 

N. moluccana is a typical Nesopupa related to Polynesian 
species. The other known species belong to groups common 
to the rest of the Oriental region, or to Africa also. 



SECOND SERIES: PULMONATA. 



MANUAL 



OF 



CONCHOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED BY 

GEORGE W. TRYON, JR. 

CONTINUED BY 

HENRY A. PILSBRY, Sc.D., 

SPECIAL CURATOR DEPARTMENT OF MOLLUSCA, ACADEMY OF NATURAL 
SCIENCES OF PHILADELPHIA. 



Part 



PHILADELPHIA : 

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES 

OF PHILADELPHIA. 



EAST INDIAN NESOPUP^E. 337 

Key to Vertiginince of the East Indies. 

t 1 . Angular and parietal lamellae diverging forward, the for- 
mer connecting with the outer lip. 

& 1 . Angular lamella small and low; 5 or 6 teeth. 

N. malayana, 110. 39. 

fe 2 . Angular lamella strongly developed; 5 to 10 teeth. 
c 1 . Shell cylindric-oblong, palatals readily visible. 
d 1 . Irregularly striate with largely cutic- 
ular riblets at intervals, often fuga- 
cious; frequently minutely pitted. 
e 1 . Without interpalatal folds. 
/'. Length 1.8 to 2.15 mm. 

N. moreleti, no. 36. 
f~. Length 1.5 mm. 

N. moellendorffiana, no. 37. 
e 2 . With 1 or 2 interpalatal folds, 7 
to 10 teeth in all ; Guam. 

N. quadrasi, no. 32. 

d 2 . Having fine, close rib-striae without 
cuticular edges; length 1.4 to 1.6 mm.; 
Philippine Islands. N. nunnodes, no. 38. 
c 2 . Shell globose-ovate ; palatals, except the upper, 
deeply immersed ; 1.36 x 1.12 to 1.45 x 1.1 mm. 

N. molucccwa, no. 35. 

Angular lamella short, remote from the upper termina- 
tion of the outer lip. 

6 1 . Diam. less than half the length; 2.25x0.9 mm.; 
Christmas Island. N. proscripta, no. 41. 

Z> 2 . Diam. more than half the length; 2 x 1.2 mm. 

N. selebeiisis, no, 40. 

No angular lamella ; parietal and columellar lamellae pres- 
ent (Costigo). 

b\ One or two palatal folds; striate, with spaced hair- 
like riblets ; Moluccas. Costigo sapwruana, p. 366. 
b 2 . No palatal folds ; no cuticular riblets ; Busuanga. 

Costigo catomanica, p. 367. 






338 EAST INDIAN NESOPUPJE. 

Section Nesopupa proper. 
35. NESOPUPA MOLUCCANA (Boettger). PI. 32, figs. 11, 12. 

Shell small, with long, deep rima, globose-ovate, brown, 
silky. Spire short with convex sides, the apex obtuse. Whorls 
4, a little convex, separated by a distinct suture, striate and 
moreover ornamented with delicate, membranous, very widely 
separated riblets, the last whorl distinctly compressed at the 
base, ascending above, towards the aperture dilated funnel- 
like, and with a long, rectangular lateral impression; three- 
sevenths the total length of the shell. The aperture is large, 
subvertical, truncate-oval, slightly impressed at the sinulus, 
with 6 or 7 teeth. Peristome expanded, thickened below the 
sinulus, calloused, violaceous, the margins joined by a light 
callus. Angular lamella very strong, emerging, joined to the 
margin of the peristome, curved, parallel within with the- 
strong, receding parietal lamella; one acute palatal tooth 
deeper in the region of the sinulus more distinct, the others, 
2 or 3 parietals [palatals] and one columellar very deeply 
placed, scarcely visible. 

Length 1%, diam. 1%, alt. and width of aperture % mm. 
(Boettger). 

Moluccas: Ema, southern Amboina (Strubell). 

Vertigo (PtychochUus) moluccan<i BTTG., Bericht Senck. 
naturf. Ges., 1891, p. 269, pi. 3, figs. 11, 110. 

This species is remarkable for its short, broad shape, the 
very strong angular and parietal lamellae, and the deep im- 
mersion of the lower palatal and columellar. In the single 
specimen seen, a topotype having Strubell's label, the cutic- 
ular riblets are mainly wanting. The surface shows some 
granulation or pitting in places, as in the Philippine species. 
The last whorl has a horizontal impression behind the lip, 
over the upper palatal fold, and further back there is a flat- 
tening over the lower palatal, giving the base a pinched ap- 
pearance. The very large parietal lamella curves towards 
the columella inwardly. There is a small but erect supra- 
palatal fold, and below it, more immersed, the laminar upper 
palatal. A small very deeply immersed interpalatal fold and 



EAST INDIAN NESOPUP^. 339 

a larger lower palatal are partially concealed by the parietal 
lamella. The columellar lamella is far within, not visible in 
a front view. Its inner end turns abruptly downward and 
along the pillar. 
Length 1.45, diam. 1.1 mm. 

Section Indopupa n. sect. 

Aperture having a strong angular lamella connecting with 
the outer lip, and typically a somewhat pitted surface, as in 
Nesopupa, but the columellar lamella is horizontal or ascends 
slightly at the inner end. Type: N. filosa (Theob. & Stol.). 

36. NESOPUPA MOKELETI (Brown). PL 32, figs. 1, 2, 3, 6. 

Shell perforate, ovate, thin, costate, pellucid, tawny. Spire 
convex, the apex obtuse. Whorls 6, convex, the last more 
than one-third the length, ascending in front. Aperture 
nearly vertical, somewhat rounded, with fine lamelliform 
teeth: one columellar; two parietal, the left one largest, the 
right deeply incised in the middle; one basal, one palatal. 
Peristome white, strongly reflected, flexuous. 

Length 2*4, diam. 1 mm. (Brown). 

Labuan Island, on the north coast of Borneo (Brown). 
Philippines (Quadras). 

Pupa moreleti A. D. Brown, Journ. de Conch., xviii, 1870, 
p. 393. PFR., Monogr. Hel. Viv., viii, 391. E. A. SMITH, 
Ann. and Mag. N. H. (6), xiii, p. 458 (Bunguran, Natuna 
Group, Everett). 

As this species has not been figured and the description is 
somewhat inexact it has been misunderstood by some authors. 
The record from the Natuna group is uncertain, as Mr. Smith 
states that his examples did not agree fully with the descrip- 
tion of moreleti. 

N. moreleti stands close to N. moellendorffi Bttg., which is 
probably a small race of the same species. As specimens from 
Cebu are not at hand, and the size is less than in any of the 
series of moreleti seen, both forms are left standing for the 
present. 

The outline is cylindric-oblong, the last two whorls almost. 



340 EAST INDIAN NESOPUP^E. 

equal in diameter. The color is near olive-buff, the shell 
slightly translucent with but little gloss. There are narrow 
riblets at wide, unequal intervals, chiefly on the penult whorl, 
and under the microscope weak interstitial striae and some 
irregular pitting are visible. The whorls are strongly convex. 
The last whorl has a rather deep, horizontal impression be- 
hind the outer lip, over the upper palatal fold. The aperture 
has five to eight teeth. The angular lamella is strong and 
curves outward to join the outer margin. It is low in front, 
with a depression before the higher part inward; this seen 
foreshortened was what Brown referred to as medio valde 
in-ciso. The parietal lamella is very high, strong and long, 
also curved. Infraparietal small, elongate. Columellar la- 
mella is strong and ascends a little inwardly. There is a 
small transverse basal tubercle in the most fully developed 
shells, but wanting or very inconspicuous in some apparently 
adult. The two palatal folds are long, the lower one longer 
and more deeply immersed. In some individuals there is a 
small, tubercular suprapalatal. The peristome is well ex- 
panded and strongly thickened within, the callus whitish. 
Sinulus defined by a thickening in the outer lip. 

Length 2, diam. 1.2 mm. ; scarcely 5 whorls. 

Length 2.15, diam. 1.2, aperture 0.85 mm. ; 5y s whorls. 

The variation in number of teeth is due to the absence 
some examples of the basal, suprapalatal and infraparietal. 
Figures 1-3 represent specimens from A. D. Brown. There is 
a set of moreleti in the collection labeled "P. pnnctilucens 
Issel, Borneo," probably = [Pupa (Istmia)] punctilium 
Issel, Borneo, Paetel, Catalog, 4th edit., 1889, ii, p. 303, nude 
name. 

Shells from Penon de Bintuan agree well with the Labuan 
form, having 8 teeth, the developed sculpture well developed. 

Length 2.2, diam. 1.23 mm. ; 5 whorls. 

Small examples from Montalban have a suprapalatal and a 
small basal fold but no infraparietal. The riblets are few 
and weak. Length 1.8, diam. 1.15 to 1.2 mm. 

Both of the above lots were from the Quadras collection, 
and labeled Pit-pa moellendorffi Bttg. 






EAST INDIAN NESOPUP^. 341 

37. NESOPUPA MOELLENDORFFI (Boettger). PL 32, figs. 4, 5. 
Shell small, shortly punctate-rimate, conic-oval, thin, cor- 
neous-fulvous, silky. Spire conic with moderately convex 
sides, the apex rather acute. Whorls 4%, a little convex, 
separated by a deep suture, striatulate and ornamented with 
very oblique, distant, deciduous riblets, the last whorl two- 
fifths the altitude, very slightly ascending to the aperture, 
the middle part spirally impressed and somewhat compressed 
at the side below. Aperture oblique, somewhat heart-shaped, 
with 5 to 7 whitish teeth : on the parietal wall 2 or 3, the 
middle one lamelliform, receding, longer, the left tooth small, 
sometimes disappearing, the right or angular emerging and 
either contiguous to or connected with the upper lip margin ; 
upper columellar lamella strong, above the middle of the 
columella, the lower one small and sub-basal, sometimes dis- 
appearing; palatals two, distant, lamelliform. Peristome 
acute, little expanded, the middle of the right margin a little 
drawn forward. 

Length l 1 /^, diam. %, alt. aperture y 2 mm. (Boettger). 

Philippines : Mt. Licos, Cebu. Also lives on Siqui jor, Min- 
danao, Masbate, Luzon, Katanduanes. 

Ptychockilus moellendorffi BOETTGER, Bericht Senck. Ges., 
1890, p. 252, pi. 9, f. 4. Vertigo moellendorffi Bttg., MOEL- 
LENDORFF, Abh. nat. Ges. Gorlitz, xxii, 1898, p. 152. 

"Thinner shelled, narrower and more slender than all the 
rest of the species" according to Boettger, whose figures are 
copied. 

Very close to N. moreleti, probably not specifically distinct. 

38. NESOPUPA NANNODES (Quadras & Moellendorff), n. sp. 
PL 32, fig. 13. 

The shell is broadly rimate, oblong, the spire tapering 
slowly below, then rapidly to the obtuse apex; light brown. 
Sculpture of fine, close rib-striae, straight, oblique, a little 
paler and narrower than the intervals, the first l 1 /^ whorls 
smoothish. The whorls are strongly convex, parted by a deep 
suture, the last slightly pitted behind the lip over the upper 
palatal fold. The aperture is subvertical, 6-toothed : angular 



342 EAST INDIAN NESOPUP^E. 

lamella high, rather thin, curving outwardly to join the callus 
in the posterior termination of the lip. Parietal lamella 
larger; columellar lamella well developed, subhorizontal ; a 
small basal fold where the columella curves into the basal 
margin. The two palatal folds are strong, the upper one 
high but rather short, the lower more immersed, long, enter- 
ing deeply. The peristome is well expanded, thickened 
within, the outer lip rather strongly bent inward in the 
middle. 

Length 1.55, diam. 0.9 mm.; 4% whorls. 

Length 1.4, diam. 0.9 mm. 

Philippine Archipelago: Bohol (Quadras). 

Vertigo nannodes Quadr. et Mild., MOELLENDORFF, Abhandl. 
Naturforsch. Ges. Grorlitz, xxii, 1898, p. 152, no description. 

This species is distinguished by its fine, regular rib-striae 
and the absence of accessory teeth. It is also smaller than the 
related forms. The columellar lamella enters horizontally, its 
inner end not turned either up or down. The specimen de- 
scribed and figured is from Quadras. 

N. nannodes is similar to Afripupa in sculpture and prob- 
ably belongs to that section. 

Section Insulipupa P. & C., n. sect. 

Angular lamella straight, low, joining the outer lip but not 
penetrating much beyond the outer end of the parietal lamella. 
Columellar lamella straight or slightly turned up at its inner 
end. Type N. minutalis (Morel.). 

39. NESOPUPA MALAYANA (Issel.). PL 32, figs. 14, 15, 16. 

Shell very minute, rimate, cylindric-ovate, ornamented with 
oblique, rough, lamellar riblets under a strong lens ; corneous 
buff, translucid. Spire tapering, the apex obtuse. Whorls 6, 
a little convex, parted by an impressed suture, the last about 
one-third of the total length, slightly ascending at the aper- 
ture, a little compressed in the middle. Aperture nearly 
semioval, vertical, 6-toothed: two deep parietal teeth; two 
columellar, the lower one smaller; two lamelliform palatals. 
Peristome reflected, the right margin sinuate, columellar ob- 
lique, lightly arcuate (Issel). 



EAST INDIAN NESOPUP^E. 343 

Length 2*4, width 1% mm. (Issel). 

Borneo (Damon). Philippine Islands (Moellendorff, Qua- 
dras). 

Vertigo mdlayanus ISSEL, Mollusehi Borneensi, in Ann. 
Mus. Civ. Genova, vi, 1874, p. 416, pi. 5, f. 30-32. Stcmrodon 
moreleti (A. D. Brown), BOETTGER, Bericht Senekenb. Ges., 
1890, p. 252, not Pupa moreleti A. D. Brown. 

Figures 14, 15 are copied from Issel. He explains that the 
second columellar tooth is small and was inadvertently omitted 
by the draughtsman. 

I have provisionally identified this shell with specimens 
from Manila and Masbate, which are the Staurodo-n moreleti 
of Boettger and von Moellendorff, reported from Cebu, Busu- 
anga and Luzon, and in Moellendorff 's Verzeichnis from the 
entire Philippine Archipelago. But in material seen there is 
no basal or second columellar tooth (there being but 5 teeth), 
and the angular lamella is less erect than figured by Tappa- 
rone Canefri. These specimens are, in fact, not specifically 
distinct from N. barrackporensis (Gude). They differ from 
N. minutalis (Morel.) by being more rugose. A Manila ex- 
ample (pi. 32, fig. 16) is here described. 

The shell is cylindric-oblong, hazel or somewhat darker in 
color, somewhat glossy, rather coarsely striate, and under the 
microscope densely pitted or vermiculate-gran&se. The whorls 
are moderately convex. The aperture has five teeth. The 
angular lamella is low, diverging forward from the parietal, 
joining the outer margin. Parietal high and long, slightly 
curved. The columellar lamella is strong, nearly horizontal, 
but ascending a little inwardly. There is no trace of a sub- 
columellar or basal. The two rather short palatal folds are 
rather deeply placed. The peristome expands rather broadly 
but is hardly thickened within. There is no perceptible 
parietal callus. 

Length 2, diam. 1.1 mm. ; 5 whorls. 

40. NESOPUPA SELEBENSIS (Tapp. Can.). PL 31, fig. 2. 

Shell minute, subrimate, subovate, very thin, corneous- 
brown. Spire a little elevated, the apex obtuse. Whorls 5, 



344 EAST INDIAN NESOPUP^E. 

subconvex, ornamented with frequent membranous riblets, the 
last usually nearly smooth, somewhat compressed behind the 
aperture, about two-thirds the total length of the shell. Aper- 
ture rather large, subquadrate above. Peristome a little ex- 
panded and somewhat reflected throughout, the margins sep- 
arated. Apertural folds 4 or 5: angular minute, parietal 
larger, somewhat sinuous, diverging from the angular; colu- 
mellar moderate ; palatals 1 or 2, the upper one larger. 

Length 2, width 1.2 mm. (T. C.). 

Celebes: Macassar (Beccari). 

Pupa (Vertigo) selebensis TAPPARONE CANEPRI, Ann. Mus. 
Civ. Geneva, xx, 1884, p. 171, pi. 1, f. 12, 13. 

Systematic position doubtful. 

41. NESOPUPA PROSCRIPTA (E. A. Smith). PL 31, fig. 1. 

Shell minute, oblong, cylindric, umbilicate, brownish-cor- 
neous, striate with delicate lines of growth. Whorls 5, very 
convex, separated by a deep, slightly oblique suture, the last 
whorl slightly ascending, impressed behind the lip. Lip pale, 
expanded, somewhat reflected, the outer margin slightly pro- 
jecting inwardly in the middle. Aperture five-toothed: two 
unequal parietal teeth, a minute columellar, and two sub- 
equal, lamelliform palatals, distant from the edge of the lip. 

Length 2.25, diam. 0.9 mm. (Smith). 

Christmas Island (in the Indian Ocean south of the west- 
ern extremity of Java, lat. 10 25' S., Ion. 105 42' E;) : Fly- 
ing Fish Cove. Type in British Museum. 

Jaminea proscripta SMITH, Ann. and Mag. Nat. Hist. (7), 
xvi, Nov. 1905, p. 551, fig. 

This species is much more slender than N. selebensis, which 
from the description seems to have similar teeth. There is a 
similar form on Tenimber Is., I believe still undescribed, and 
known to me by imperfect shells only. These species have 
the angular lamella short. It is remote from, not running to 
the upper termination of the outer lip, thereby differing from 
N. malayana. 



NESOPUP^E OF INDIA AND CEYLON. 345 

V. SPECIES OP INDIA AND CEYLON. 

Three of these species, filosa, brevicostis and salemensis, 
belong near N. moreleti and its East Indian allies. N. barrack- 
porensis is not specifically separable from Philippine speci- 
mens referred to N. malayana-, described from. Borneo, and is 
also related to species of the Comoros and Africa. The teeth 
of N. cmghalensis have not been described or figured in suffi- 
cient detail for a decision, but it is probably near barrack- 
pore-usis. The Nesopupa fauna of the Oriental Region is thus 
quite homogeneous. 

Specimens of filosa and barrackporensis have been exam- 
ined, the accounts of the others being compiled. 

Angular lamella curved, rather high ; shell ovate. 

Length 2 mm., irregularly striate. N. filosa, no. 42. 

Length 1.5 to 1.75 mm. 

N. brevicostis, no. 43 ; N. salemensis, no. 44. 
Angular lamella low, straight ; shell cylindric ; length 2 mm. 

Seven teeth. N. cinghalensis, no. 45. 

Five teeth. N. barrackporensis, no. 46. 

Section Indopupa P. & C. 
42. NESOPUPA FILOSA (Theob. & Stol.). PL 31, figs. 9, 10. 

The shell is ovate, imperforate but shortly rimate, dull, 
cinnamon-buff, thin, having rather strong but unequal striae 
widely but unequally spaced, the intervals vermiculate-gra- 
nose. Outlines of spire convex, the summit obtuse; penult 
whorl large, strongly convex; last whorl having a slight 
impression behind the outer lip, over the upper palatal fold. 
Aperture with 5 teeth: angular lamella rather low in front, 
higher within, curved, joining the outer lip. Parietal lamella 
high, slightly sinuous, not entering very deeply. There is 
usually a vestigeal infraparietal tubercule, so low that it is 
readily overlooked. Columellar lamella rather deeply placed, 
subhorizontal. Two rather short palatal folds, the upper low, 
lower much higher. The peristome is narrowly reflected, 
colored like the shell, slightly thickened, thinner at the sin- 



346 NESOPUP^E OP INDIA AND CEYLON. 

ulus, which is not very well defined. Parietal callus scarcely 
noticeable. 

Length 2, diam. 1.25 mm. ; 4^2 whorls. 

Burma: Arakan coast, rather abundant (Theobald). 

Pupa filosa THEOBALD & STOLICZKA, Journ. Asiatic Soc. 
Bengal, xli, pt. 2, 1872, p. 333, pi. 11, f. 8. HANLEY & THEO- 
BALD, Conch. Indica, pi. 160, f . 5. Pupilla filosa Theob. & 
Stol., GUDE, Fauna Brit. India, Moll, ii, p. 287. Nesopupa 
filosa PILS., Man. Conch., xxiv, p. 140, 366, pi. 25, f. 10, 11. 

It is characterized by the large size of the last and penult 
whorls, giving it a somewhat cylindric outline, and the rather 
weak teeth of the large aperture, as compared with N. more- 
leti, which is related. There are only five distinct teeth, as 
the infraparietal is so small that it was overlooked by Theo- 
bald and Stoliczka, though present in their type lot. 

The figures represent the type (fig. 10) and a paratype 
from the lot of four in coll. Indian Museum. Before seeing 
these specimens the original figures had been copied on pi. 
25, figs. 10, 11 of Vol. xxiv, under the impression that it was 
a Gastrocopta. 

43. NESOPUPA BREVICOSTIS (Benson). PL 31, fig. 11. 

"Shell rimate-perf orate, cylindric-ovate, corneous, apex ob- 
tuse. Whorls 41/2, rapidly increasing in length, the last not 
ascending, one-third the length of the shell, upper whorls 
convex, remotely semicostulate above, last and penult whorls 
rather flattened, the lower half of the rest also silky, bearded. 
Aperture rounded-oval, 5 to 6 plicate: a short angular fold; 
an oblique, more deeply placed parietal; a single columellar; 
two or three palatals, deeply placed; peristome expanded, 
subreflected. 

"Length 1.5, diam. scarcely 1 mm." (Bens.). 

India: Barrackpore, Bengal (Bacon). 

Pupa brevicostis BENS., Ann. Mag. N. H. (2), iv, August 
1849, p. 127. KUSTER, Conchyl. Cab., Pupa, p. 136, pi. 17, 
f. 20-22. Pupilla brevicostis Bens., GUDE, Fauna British 
India Moll., ii, p. 288, f. 104. 

"Taken by Dr. J. F. Bacon on the trunk of a tamarind 



NESOPUP.E OP INDIA AND CEYLON. 347 

tree at the Cantonment of Barrackpore, near Calcutta, dur- 
ing the rainy season of 1848. Out of several individuals for- 
warded to me, overland, by letter in a quill, two reached me 
alive, and creeping about when supplied with moisture en- 
abled me to verify their affinities. The lower pair of tentacula 
is deficient or inconspicuous, as in Vertigo; the upper pair 
carry the eyes at their summits. The shell is often carried at 
an angle of 45" (Benson). 

Mr. Gude has figured a specimen from the British Museum 
measuring 1.75 x 1.1 mm. Fig. 11 is a copy of this. The type 
cannot be found. * ' The species is characterized by the lamel- 
late striae and the short spire as compared with the diam- 
eter" (Gude). 

44. NESOPUPA SALEMENSIS (Blanford). PL 31, figs. 7, 8. 

' ' Shell rimate, ovate-oblong, obliquely striate, tawny. Spire 
elevated, conoidal, the apex obtuse ; suture impressed. Whorls 
41/2, the upper ones somewhat convex, the last about two-fifths 
the total length, excavated in the middle, behind the aperture, 
arcuately ascending in front, compressed near the suture and 
below. Aperture reniform, columella receding, calloused 
above; two compressed parietal teeth. Peristome expanded, 
the outer lip flexuously excavated, one-toothed. 

"Length 1.6, diam. 1, alt. aperture 0.6 mm." (Blanf.). 

India: Kalryen Hills (Foote). 

Ennea salemensis BLANF., Journ. Asiatic Soc. Bengal, xxx, 
1861, p. 359, pi. 2, f. 8. Pupa salemensis Blanf., PFB., 
Monogr. Hel. Viv., vi, 1868, p. 318. HANLEY & THEOBALD. 
Conch. Indica, 1876, pi. 160, f. 9.Piipilla salemensis Blan- 
ford, GUDE, Fauna Brit. India, Moll., ii, p. 289. Pupa sale- 
manensis Sowerby, Conch. Icon., xx, 1876, Pupa, pi. 7, f. 62. 

Mr. Gude remarks that it appears much nearer brevicostis 
than to Pupa seriola with which Blanford compared it. The 
figures are copied from Blanford and Conchologia Indica. 

45. NESOPUPA CINGHALENSIS (Gude). PL 31, figs. 5, 6. 

' ' Shell narrowly perforate, almost rimate, cylindrical ovate, 
rather solid, very minutely striatulate, the earlier whorls 



348 NESOPUP^ OP INDIA AND CEYLON. 

corneous, the later ones rufous. Spire convex, apex obtuse r 
suture impressed. Whorls 5, slightly convex, the last flattened 
and scrobiculate behind the peristome, shortly ascending, 
slightly compressed and subangulated around the extremely 
narrow perforation of the umbilicus. Aperture oblique, 
broadly obovate, furnished with seven denticles, 2 compressed, 
elongate, on the parietal wall, 2 very minute and deeply 
seated, on the columella, 3 also very minute and deeply 
seated, on the palatal wall. Peristome rufous, slightly thick- 
ened and reflexed ; outer margin slightly inflexed, basal mar- 
gin rounded, columellar margin obliquely ascending and 
slightly dilated above. 

"Long. 2, diam. 1 mm." (Gude). 

Ceylon. Type in the British Museum. 

Pupilla cinghalensis GUDE, Fauna of British India, MolL T 
ii, 1914, p. 286, fig. 108. 

"This new species is based on a specimen from Ceylon, 
without precise indication of habitat, acquired by me some 
years ago at a sale. It differs from P. barrackporensis, its 
nearest ally, in having seven instead of four denticles in the 
aperture, and in having the peristome fulvous instead of 
white" (Gude). 

Section Insulipupa Pils. 
46. NESOPUPA BARRACKPORENSIS (Gude). PL 31, figs. 3, 4. 

This species was placed in Gastrocopta with a mark of 
doubt in Vol. xxiv, p. 135. Specimens sent from the Indian 
Museum, labeled by Nevill "Pupa (Vertigo) brevicostis = V. 
indica Pfeiffer," now at hand (pi. 31, figs. 3, 4), described 
below. They are apparently the Barrackpore examples 
noticed by Nevill in his Hand List Moll. Ind. Mus., i, p. 197, 
and which Mr. Gude surmised were barrackporensis (=> indica 
Pfr. not Benson). If so, the low angular lamella was over- 
looked by all who have described the species. Mr. Sowerby's 
two wretched figures (Conch. Ind. and Conch. Icon.) appear 
to show it indistinctly. In renaming the shell Mr. Gude 
added nothing to the definition or illustration, though he had 
the types and other specimens under examination, and must 



NESOPUP^E OP THE MASCARENE ISLANDS. 349 

have been aware of the inadequacy of the published data. 
The original description may be found in Vol. xxiv, p. 135. 

The shell is imperforate, shortly rimate, cylindric, cinna- 
mon-buff. First 1% whorls smoothish, the rest having un- 
even oblique striaB, mainly obsolete or blurred on the last two 
whorls, which have a minute vermiculate-granose sculpture, 
varying in distinctness on different specimens. The upper 
whorls are quite convex, the last two much less convex; last 
whorl having a broad, shallow impression behind the lip, over 
the upper palatal fold, both palatals showing through the 
shell as light spots. The squarish-oval aperture is longer 
than wide, having five teeth : angular lamella is quite low, 
diverging from the parietal and joining the outer lip. The 
parietal lamella is much the largest tooth. It is thin but 
high, directed obliquely and a trifle sinuous. The columellar 
lamella is situated high on the columella and rather deep 
within, ascending a little as it enters, and the edge is directed 
obliquely downward. The two palatals are well developed, 
the lower a little deeper within and longer. The lip is slightly 
lighter than the exterior, well expanded, with hardly any 
thickening. A parietal callus is scarcely visible. 

Length 2, diam. 1.12 mm. ; barely 5 whorls. 

These shells do not differ materially from the Philippine 
specimens which I have identified as malayana Issel ; yet as I 
have examined the types of neither species, and the original 
account of malayana is somewhat discrepant, the species are 
both allowed to stand provisionally. 

This Indian species has much stronger microscopic rugosity 
and less prominent oblique striae than N. minutalis. It differs 
from N. corrugata chiefly by the well-developed palatal folds, 
but it is also a little smaller than that species. 

VI. SPECIES OP THE MASCARENE ISLANDS AND COMOROS. 

The islands of the western Indian Ocean have representa- 
tives of three genera of three subfamilies of Pupillidae. One 
of these, the genus Gastrocopta,, has been considered in Vol. 
xxiv, p. 127. They are small, white-toothed shells in which 



350 NESOPUP^E OP THE MASCABENE ISLANDS. 

the angular and parietal lamellae are eoncrescent into one 
sinuous lamella. 

The Vertigininae have been very poorly described, and only 
two of the nine species are at hand for examination. One 
subgenus, Nesopupa, is common to these islands and Poly- 
nesia; another, Insulipupa, is common to the East Indies, 
India, the Mascarene Islands and Africa. Several other 
species, in the absence of specimens or adequate figures, can- 
not safely be classified. The species borbonica and desmazu- 
resi resemble the East Indian group Costigo in teeth, and 
have provisionally been placed there. 

Key to Species. 

a 1 . Aperture having both angular and parietal lamellae. 

ft 1 . Angular lamella strongly developed, rather long 
and high ; columellar lamella entering deeply, dilated 
and curving downward at the inner end (Nesopupa) . 
c 1 . Length 2.5 mm., about 8 teeth. 

N. gonioplax, no. 47, 
c 2 . Length 1.5 mm., at least 5 teeth. 

N. micro,, no. 48, 

b 2 . Angular lamella small and much lower than the 
parietal; columellar lamella subhorizontal, not 
turned down within (Insulipupa) . 

c 1 . Species of the Comoros, having 5 or 6 teeth. 
d 1 . 2x1 mm. N. minutatis, no. 49. 

d 2 . 1.5 x 0.75 mm. N. comorensis, no. 50. 
c 2 . Bourbon ; 4 to 5 teeth, 1 or 2 palatals. 

N. incerta, no. 51* 

c 3 . Mauritius; angular, parietal and columellar 
lamellae only, no palatals. N. ventricosa, no. 52. 
a 2 . Parietal lamella present, but no angular; 1 columellar, 2 
palatals. 

ft 1 . 2.5 x 1.33 mm. ; Bourbon. Costigo borbonica, p. 367. 

b 2 . 2.25 x 1 mm. ; Rodriguez. Costigo desmazuresi, p. 368. 

a 3 . No teeth on the parietal wall; columellar lamella, basal 

and two palatals present ; apex very obtuse ; length 2 mm. 

"Vertigo" praslinensis, no. 53. 



NESOPUP.E OP THE MASCARENE ISLANDS. 351 

Section Nesopupa proper. 

47. NESOPUPA GONIOPLAX n. sp. PL 33, figs. 8, 9, 10. 

The shell is ovate, cinnamon-brown, the peristome more 
reddish, the teeth paler. The surface has little gloss, and 
under the microscope shows some rather weak growth- 
wrinkles and a dense, very shallow pitting or minute mallea- 
tion. The whorls are convex, the last having behind the lip 
two furrows, one lateral, the other sub-basal, diverging for- 
ward, and corresponding to the upper and lower palatal 
folds. The aperture has subparallel sides, straight parietal 
callus and strongly curved base, the peristome well expanded, 
somewhat thickened, at the sinulus thinner and retracted. 
The angular lamella is very strong, curved, joining the outer 
lip, diverging forward from the very strong parietal lamella, 
the inner end of which is bent towards the columella. There 
is a small infraparietal, rather deep within. The columellar 
lamella enters nearly horizontally, then deep within on the 
axis after rising a little it turns downward forming a broad 
plate, only imperfectly seen from in front but prominent from 
the back (fig. 9). There is a small basal fold at the junc- 
tion of columellar and basal margins. The upper and lower 
palatal folds are sinuous, strong, and converge inwardly. A 
small suprapalatal stands above them, making 8 teeth in all. 

Length 2.5, diam. 1.55 mm. ; 5 whorls. 

Mauritius (Morelet collection from Nevill, type 64093 A. 
N. S.P.). 

The most remarkable character of this snail is the dilation 
and deflection of the columellar lamella. Nothing of this 
structure is seen in N. moreleti and others of that group, 
which are otherwise similar. Though the shells came from 
Nevill, according to Morelet 's label, I cannot find that either 
G. Nevill or H. Adams described the species. The following 
species is apparently its nearest affinity. 

48. NESOPUPA MICRA, new name. PI. 33, fig. 4. 

The shell is deeply rimate, ovate-cylindric, thin, very lightly 
striate, silky, brownish-corneous. Spire convex, apex obtuse. 



352 NESOPUP^E OF THE MASCARENE ISLANDS. 

Whorls 5, convex, the last obtusely angular around the um- 
bilical crevice, narrower and bisulcate behind the aperture. 
Aperture truncate-oval, vertical, with two parietal lamellae, 
the right-hand one prominent, left one remote. Peristome a 
little expanded, the right margin sinuous, provided with two 
entering folds. Length 1.5, diam. 0.75 mm. (H. Ad.). 

Mauritius: Bamboo (G. Nevill). 

Pupa (Pupilla) exigua H. ADAMS, Proc. Zool. Soc. Lond., 
1868, p. 13, pi. 4, f. 4. Not Pupa exigua Say, 1822. 

Adams did not mention a columellar lamella, but it is seen 
in his figure. The species is perhaps related to the preceding 
or to the Continental bisulcata group. It is smaller and nar- 
rower than N. gonioplax, with different sculpture. 

Section Insulipupa P. & C. 

Probably N. ventricosa H. Ad., which lacks palatal folds, 
belongs to this group. In that case Insulipupa would include 
the subgenus Pagodella? H. Adams, Proc. Zool. Soc. London, 
1867, p. 304, monotype Pupa ventricosa H. Ad. ; not Pagodetta 
Swainson, 1840. 

49. NESOPUPA MINUTALIS (Morelet). PL 33, figs. 1, 2, 3. 

Shell minute, rather deeply rimate, acutely subcylindric, 
corneous-tawny, somewhat shining, smooth, the apex obtusely 
conoid. Whorls 5%, a little convex. Aperture regularly 
oval, 6-plicate : one lamellif orm angular ; one transverse pari- 
etal; a rather prominent columellar; and three nearly equi- 
distant punctiform teeth in the outer margin. Peristome 
simple, a little expanded, the margins separated. Length 2, 
diam. 1 mm. (Morelet). 

Comoro Is. : Mayotte, on the islet of Dzaoudzi, on the bark 
of an Acacia. 

Pupa minutalis MORELET, Journ. de Conchyl., vol. 29, 1881, 
p. 231, pi. 10, f. 5. 

The form is cylindric with obtuse, rounded or convexly 
conic summit. The surface is rather dull with some irregular 
striation. The angular lamella is quite low, forwardly diverg- 
ing from the parietal and joining or almost joining the outer 






NESOPUP^E OP THE MASCARENE ISLANDS. 353 



lip. Parietal lamella high, rather long; columellar strong, 
siibhorizontally entering, its crest slanting downward. There 
are two quite short palatal folds, the lower one somewhat the 
larger and more immersed. The upper palatal is sometimes 
very small. According to Morelet ' ' trois denticules, visibles a 
la loupe, sont placees sur le cote libre du peristome, a des 
distances a peu pres egales Tune de 1'autre"; one of these 
must be a basal fold ; but no basal is to be seen in the speci- 
mens examined, and Morelet 's figure shows only parietal and 
columellar teeth. The outer lip is well expanded. In a series 
received from M. Marie it is not thickened, though whitish; 
in another set, also from Mayotte, there is a distinct though 
rather thin callous thickening, and the teeth are larger. 

Length 2, diam. 1.15 mm. ; slightly over 5 whorls. 

Length 1.8, diam. 1.1 mm. ; 5 whorls. 

This species, as Dr. Boettger has already noted, is related 
to the East Indian N. malayana. It differs from that species 
by the smoother surface, which has very little pitting or gran- 
ulation. 

50. NESOPUPA COMORENSIS n. n. PL 33, fig. 5. 

The shell is minute, deeply rimate, ovate, glossy, brownish- 
corneous, the apex rather obtuse. Whorls 5, convex, joined 
by an impressed suture, the last slightly ascending, somewhat 
compressed at the base. Aperture semioval, with two parietal 
lamellae, another columellar, and two small teeth in the basal 
margin. Peristome somewhat expanded, the margins sub- 
parallel. Length 1.5, diam. 0.75 mm. (Morelet). 

Comoro Is.: Anjouan (Bewsher) ; Mayotte (Marie). 

Pupa monas MORELET, Journ. de Conchyl., vol. 27, 1879, p. 
310, pi. 12, f . 4 ; vol. 31, 1883, p. 196. Not Pupa substriata 
var. monas Westerlund, 1871. 

As specific and varietal names are held to have the same 
status in nomenclature, the name of this species has to be 
changed. 

51. NESOPUPA INCERTA (G. Nevill). Not figured. 

Shell very closely resembling Vertigo (Pagodella) ventri* 



354 NESOPUPJE OP THE MASCARBNE ISLANDS. 

cosa H. Ad., from Mauritius; there are, however, two pari- 
etal teeth and within the outer lip a distinct, well-developed 
tooth, with occasionally another small, indistinct one close to 
it; the eolumella is slightly more dilated and subangulated 
(NeviU). 

Bourbon: near Salazie, in company with V. pupula Dh. 
(G. NeviU). 

Vertigo (PagodeUa) incerta G. NEVILL, Journ. Asiatic Soc. 
Bengal, vol. 39, pt. 2, 1870, p. 413. 

"This most perplexing of shells can only be distinguished 
from Pagodella ventricosa H. Ad. (Proc. Zool. Soc., 1867, p.. 
303) by the different dentition of the aperture; of the latter 
I found about 40 specimens, to all appearance full grown and 
in first-rate condition, some of them, to my mind, very old 
specimens; in none of them were there any signs of any 
teeth whatever within the outer margin of the aperture ! Of 
the Bourbon species, I only found 5 specimens, one evidently 
young, the other 4 full grown and all showing the peculiar 
characteristics pointed out in my description. Still the re- 
semblance is so striking that I think no naturalist would 
hesitate to avow that they must at no very remote period 
have had a common origin; there is, indeed, just the chance 
that at Mauritius a similar variety may exist, but, from the 
number of specimens I found there, I doubt it exceedingly. 
I have, however, written to Mr. Dupont at Mauritius to ask 
him to kindly examine all he can possibly procure of this, 
species, and to see if he can find any trace of the apertural 
teeth, which give such a different appearance to my V. in- 
certa' 1 (NeviU). 

It is curious that anyone could write so much about a shell 
and tell so little. 

52. NESOPUPA VENTBICOSA (H. Adams). PI. 33, fig. 6. 

Shell deeply rimate, ovate, rather thin, obliquely striat- 
ulate, pale brown ; spire convexly conic, the suture impressed. 
Whorls 5, somewhat convex, the last ventricose, rounded at 
base. Aperture semioval, provided with a compressed, en- 
tering parietal fold and a tooth at the insertion of the right 



NESOPUP^ OP THE MASCARENE ISLANDS. 355 

margin; peristome simple, slightly expanded, the margins 
joined by a callus, right margin somewhat sinuous, columel- 
lar margin dilated above. Length 2.5, diam. 1.75 mm. 
(H. Ad.}. 

Mauritius: The Moka ravines (G. Nevill). 

Pupa (Pagodella) ventricosa H. ADAMS, Proc. Zool. Soc. 
London, 1867, p. 304, pi. 19, f. 6. Pfr., Monogr., vi, p. 308. 
Pagodella ventricosa H. Ad., G. NEVILL, Journ. Asiatic 
Soc. Bengal, vol. 39, pt. 2, 1870, p. 413. 

G. Nevill writes of Pagodella ventricosa: "I found about 
40 specimens, to all appearance full grown and in first-rate 
condition, some of them, to my mind, very old specimens; in 
none of them were there any signs of any teeth whatever 
within the outer margin of the aperture." 

53. " VERTIGO " ( ? ) PRASLINENSIS ( G. Nevill ) . Not figured. 

Shell resembles Pupa lienardiana Crosse, from Rodriguez, 
but the spire is less produced and there is no trace of the 
conspicuous parietal tooth; apex remarkably obtuse; 4% 
whorls which increase very rapidly, the first three more or 
less cylindrical, the last one moderately convex ; smooth, even 
under the lens no trace of sculpture is discernible; aperture 
triangular, regularly rounded at base, external margin some- 
what obsoletely sinuate; a strongly developed, transverse 
columellar tooth, three other palatal teeth, equidistant, the 
middle one the largest, no parietal tooth (G. Nevill). 

Length 2 mm. 

Seychelles Group: Praslin, near the Protestant church, at 
the foot of a cocoanut tree (Nevill; type in Indian Mus.). 

Carychium n. sp., G. NEVILL, Proc. Zool. Soc. Lond., 1869, 
p. 65. Vertigo praslinensis G. NEVILL, Journ. Asiatic Soc. 
Bengal, vol. 50, pt. 2, 1881, p. 140. 

The position of this species is quite uncertain. It may 
possibly be a Gastrocopta or one of the Pupillinae. For a 
shell of this group or of Gastrocopta to have well-developed 
columellar and palatal teeth and no parietal is anomalous. 
In dead shells the latter sometimes scales off with the parietal 
callus. It appears that only one specimen was found. 



356 AFRICAN NESOPUP^E. 

VII. AFRICAN SPECIES. 

a 1 . Angular and parietal lamellae present. 

& 1 . Angular lamella rather high, strong and curved. 
c 1 . Length 3 mm.; nearly smooth. St. Helena. 

N. turtoni, no. 60. 

c 2 . Length less than 2 mm. ; closely rib-striate. 
'd l . Sculpture of rather coarse, very oblique 
striae ; shell ovate, white ; 1.5 x 1 mm. 
British E. Africa. N. iota, no. 57. 

d 2 . Sculpture of fine, close rib-striae; shell 
brown. 

e l . One broad median external fur- 
row behind the aperture; lower 
palatal and basal folds rather 
deeply immersed; length 1.4 to 
1.6 mm. S. Africa. 

N. griqualandic-a \, no. 54. 
e z . One very slight impression behind 
aperture; lower palatal but little 
immersed, basal minute, in sub- 
columellar position; length 1.6- 
1.65 mm. S. Africa. 

N. farquhari, no. 55. 
e s . One very slight impression be- 
hind aperture; palatals rather 
short ; no basal fold ; length 1.85 
mm. Rhodesia. 

A T . b. rhodesiana, no. 56a. 
e*. Two deep furrows behind aper- 
ture; no basal fold; length 1.87 
mm. Abyssinia. 

N. bisulcata, no. 56. 

b 2 . Angular lamella short and low; 2.25 to 2.4 mm. 
long. 

c 1 . Surface densely pitted; one palatal fold or 
none. Rhodesia. N. corrugata, no. 58. 

c 2 . Nianing, West Africa. 

AT. megalomastoma, no. 59. 



AFRICAN NESOPUPvE. 357 

a 2 . No angular lamella; parietal, columellar and two palatal 
teeth present; length 1.75 mm. Island S. Thome. 

Costigo nobrei, p. 368. 
a 3 . Aperture without teeth (Genus Negulus; Vol. XXVI). 

Section Afripupa P. & C., n. sect. 

Minute Nesopupa with teeth of the typical section except 
that the columellar lamella does not turn down at its inner 
end ; sculpture of close rib-strias without cuticular edges. 
Type N. griqualandica (M. & P.). 

54. NESOPUPA GRIQUALANDICA (Melv. & Pons.). PL 34, figs. 
1-4. 

Shell very minute, rimate, thin, rotund-cylindric, brown; 
whorls 5, rather swollen, compressed at the sutures, every- 
where closely, longitudinally, finely striate, the last whorl de- 
pressed-sulcate in the middle, dorsally towards the lip. Aper- 
ture ovate-rotund; peristome thickened, sinuated on the 
labial [labralj margin, plicate: two parietal folds, very 
prominent and recurved, a third marginal on the columella, 
dentiform. Length 1, width 0.75 mm. (Melv. and Pons.). 

British South Africa: Cape of Good Hope: Griqualand 
East (type loc., Sykes) ; Cradock (Farquhar) ; Port Eliza- 
beth (Ponsonby coll.). Natal: Pietermaritzburg, Dargle, 
Tongaat, Edendale (Burnup). Zululand: Dukuduku (Top- 
pin). Transvaal: Pretoria District (Parquhar) ; Heidelberg 
(Miss Livingston) ; Buiskop (Connolly). Rhodesia: Victoria 
Falls (Connolly). 

Pupa griqualandica MELVILL and PONSONBY, Ann. Mag. N. 
H. (6), xi, Jan. 1893, p. 3, fig. 9; (8), i, Jan. 1908, p. 76, pi. 
1, f. 8, 10. BURNUP, Ann. Mag. N. H. (8), vii, 1911, p. 405. 
Jaminia griqualandica (Melv. & Pons.), CONNOLLY, Ann. 
South African Museum, xi, pt. iii, 1912, p. 182. 

This species appears to differ from N. bisulcata (Jickeli) 
by the more deeply immersed lower palatal fold, the presence 
of a basal fold and the deeper external furrow of the last 
whorl over the upper palatal fold. It is also smaller than 
Jickeli records. 



358 AFRICAN NESOPUP^E. 

Melvill and Ponsonby's original description and figure are 
chiefly remarkable for what was omitted. The dimensions 
are grossly inaccurate. In their second account two species 
appear to have been in view, though in the main it relates to 
the true griquMandica. 

The shell is distinctly perforate, openly rimate, long-ovate, 
the spire with convex outlines and very obtuse apex. Color 
cinnamon, becoming a little darker towards the base and 
paler towards the apex. First 1% whorls smoothish (micro- 
scopically irregularly granulose), the rest with straight, close, 
somewhat oblique striae; the latter part of the last whorl 
having a broad median furrow running to the lip, convex 
above and below it, with a shorter impression at the base. 
The aperture is wider in the upper part, nearly closed by 6 
teeth: the angular lamella is subvertical, large, entering 
deeply, curving towards and joining the outer lip. Parietal 
lamella very long and high, oblique, directed towards a point 
above the lower palatal fold. The columellar lamella is 
rather deeply placed, not very high, and enters horizontally 
to the dorsal side, where the end tapers and turns upward 
but. very slightly. The upper palatal fold emerges to the 
peristome, and with the angular lamella defines a nearly 
closed, oval sinulus; it enters deeply, descending a little. 
The lower palatal is a long fold, deeply immersed, only par- 
tially visible in a front view. The basal fold is short, trans- 
verse to the cavity, deeply placed but visible from in front. 
The parietal callus is rather thin. 

Length 1.5, diam. 0.85 to 0.9 mm. ; 4% whorls. Pretoria. 

"Length 1.41, diam. 0.83 mm." (Burnup). 

"Length 1.6, diam. 0.99 mm." (Burnup, largest specimen). 

55. NESOPUPA FABQUHABI Pilsbry. PL 34, figs. 7, 8, 9. 

Shell perforate, openly rimate, long-ovate, walnut-brown, 
somewhat glossy. Spire convexly conic, the apex obtuse; 
first 1% whorls smooth, the rest closely, somewhat obliquely 
rib-striate, as in N. griqualandica; the latter part of the last 
whorl flattened and having a quite small and shallow im- 
pression behind the lip, over the upper palatal fold. Aper- 



AFRICAN NESOPUPyE. 359 

ture shaped about as in griqualandica, with 6 nearly white 
teeth. Angular lamella strong, slightly curved, joining the 
outer lip. Parietal lamella high, oblique, penetrating to the 
dorsal side. Columellar lamella deeply placed, strong, pene- 
trating to the dorsal side, where it is somewhat enlarged and 
deflected, then passing into a low callus obliquely ascending 
the axis. The upper palatal fold is strong, curved, emerging 
to the peristome, entering deeply. Lower palatal very high, 
long, somewhat immersed. Basal fold very low and small, 
on the basal end of the columella. The peristome is pale, 
well expanded and a little thickened, incurved and thicker 
in the middle of the outer margin. 

Length 1.6 to 1.65, diam. 0.9 mm. ; 5 whorls. 

South Africa: Grahamstown (J. Farquhar). 

Nesopupa farquhari PILS., Nautilus, xxxi, October 1917, 
p. 50. 

While related to N. griqualandica, this species differs in 
numerous particulars. The spire is somewhat longer; the 
last whorl is far less impressed behind the lip ; the angular 
lamella curves less; the lower palatal fold emerges much 
further, and is as prominent in a face view as the upper. 
Finally, the basal fold is even smaller than in griqualandica, 
and is upon the columella. In the other species it, stands 
deep within the base, in a face view being in front of the 
deeply immersed lower palatal fold. 

The Abyssinian N. bisulcata has not been directly com- 
pared with this species. By the description and figure it 
differs by being somewhat larger with no basal or subcolu- 
mellar tubercle, and the teeth do not appear to be as large as 
in farquhari. 

56. NBSOPUPA BISULCATA (Jickeli). PL 34, figs. 11, 13, 14. 

The umbilicate shell is ovate-conic, scarcely glossy, very 
finely and obliquely costate longitudinally (visible under a 
lens), brown. The conic spire has an obtuse apex. The 5 
convex whorls are separated by an impressed suture, the last 
whorl forming a third of the total length, is narrowed towards 
the base, with angular neck, and has two deep furrows. The* 



360 AFRICAN XE60PUPJE- 

vertieal aperture is rounded and contracted by 5 folds: the 
right one of the parietal wall is connected with the termina- 
tion of the outer lip; the second is more deeply placed* 
There is a high columellar lamella and two palatal folds, of 
which the upper emerges to the peristome and unites there- 
with, the second disappears before reaching it. The peri- 
stome is moderately expanded and thickened within, the 
right margin bent inward by the upper furrow which cor- 
responds to the upper palatal fold. Length Ivg, diam. \ f 
aperture % x % mm, (Jickeli) . 

Abyssinia: plateau of Rora-Beit-Andu. province of Ha- 
maszen. and on the Keren Mts., Bogos. under rotten leaves, 
wood and stones (Jickeli). 

Pupa bisulcata JICKELI, Malak. Blatter. 1873. p. 107; 
Fauna der Land- und Siisswasser-Mollusken Nord-Ost- 
Afrika's, in Xova Acta Acad. Caes. Leop.-Carol. Germ. Nat. 
Cur., voL 37, 1875, p. 119, pL 5, f. 10. 

No Abyssinian specimens are at hand, and as Mr. Burnup 
has observed. Jiekeli's figures do not agree in proportions 
with his measurements. The latter indicate a shell propor- 
tioned like the following form. 

56a. X. bisulcata rbodesiana n. subsp. PL 34, figs. 5, 6. 

The shell is more elongate and cylindric than in N. bisul- 
cata according to Jickeli *s figures, the penult whorl equalling 
the last in diameter. The whorls are strongly convex, the 
striation less regular and not so strong as in 3T. griqualandica. 
Latter part of the last whorl is a little flattened laterally 
and close behind the outer lip there is a short furrow above 
the middle. The teeth are much smaller than in griqualan- 
dica and farquhari, the palatals especially being shorter. No 
basal or subcolumellar fold is present. 

Length 1.85, diam. 1 mm,; 5 whorls. 

Rhodesia: Victoria Falls (Connolly). 

A specimen of this form was sent with N. griqualandica. 
It differs so much in contour from Jiekeli's figures that, tak- 
ing the locality and faunal zone into consideration, I am dis- 
posed to give it subspecifie standing. A T . bisulcata is figured 



AFRICAN NESOPUP2E. 361 

as evenly rib-striate, like N. griqualandica, while in the pres- 
ent form the sculpture is uneven. 

57. XESOPUPA( ?) IOTA (Preston). PL 34, fig. 6a. 

44 Shell minute, ovately pyramidal, thin, white; whorls 5, 
regularly increasing, the last ascending in front, sculptured 
with rather coarse, very oblique, transverse striae; suture 
jBomewhat deeply impressed, labrum sinuous, white, scarcely 
reflexed, the margins not joined, though it is continued for 
some distance along the parietal wall ; aperture almost irreg- 
ularly triangular, furnished with two oblique parietal lamellae, 
of which the lower is the coarser, a short lamella on the outer 
lip, a coarse basal denticle, and a coarse oblique lamella on 
the columella. 

"Alt. 1.5, diam. maj. 1 mm." (Preston). 

British East Africa: between Bumruti and Mount Kenia 
(collector unrecorded). 

Ennea iota PRESTON, Ann. Mag. N. H. (8), vii, May, 1911, 
p. 463, pL 11, f. 2. 

This appears, from the figure, to be a Nesopwpa. The 
sculpture seems more like the Indian than the other African 
species. It is said to be white ; if so when alive, it is unlike 
any of the known species. Not seen by the writer. 

Section Insulipupa P. & C. 

58. NESOPUPA CORRUGATA (Preston). PL 34, figs. 12, 15. 
Shell perforate, small, cylindrical, brownish-bronze color; 

whorls 5, rather convex, the first three rapidly increasing in 
both length and breadth, the last two in length only, corru- 
gated throughout, the third and fourth whorls being also ob- 
liquely, somewhat irregularly and distantly plicate; suture 
deeply impressed ; umbilicus very narrow ; columella whitish, 
outwardly and rather erectly expanded, obliquely descending ; 
labrum also whitish, somewhat sinuous, dilated below, erect 
and angulated above; aperture subquadrate, armed with a 
denticle just below the margin of the labrum, and below and 
to the left of this an oblique curved lamella on the parietal 






362 AFRICAN NESOPUP/E. 

wall, a slightly projecting denticle very internally situate on 
the upper portion of the columella, and a small, rather in- 
distinct basal denticle situated well within, and rather on the 
right-hand side of the shell. Alt. 2.25, diam. maj. 1 mm. 
(Preston) . 

Length 2.29, diam. 1.08 mm. (Burnup). 

Length 2.25, diam. 1.15 mm.; 5% whorls (Connolly coll.). 

Rhodesia: Rain Forest, Victoria Falls (M. Connolly). 

Jaminia corrugata PRESTON, Ann. Mag. Nat. Hist. (8), ix, 
Jan. 1912, p. 71, fig. 4. 

The initial l 1 /^ whorls appear smooth, but there is a very 
close microscopic pitted-granose sculpture. The next whorl 
has rather weak but coarse striation. On the following whorls 
the surface appears punctate under a lens; when more mag- 
nified it is seen to be very closely, irregularly pitted, the pits 
very irregular in shape. There are also unequal, oblique 
strise, conspicuous on the last whorl only immediately below 
the suture, but noticeable on the penult and earlier whorls. 
The striation is hardly strong enough to justify Preston's 
term "plicate". The angular lamella is rather low, straight, 
extending inward slightly beyond the anterior end of the 
parietal lamella, diverging from the latter and nearly join- 
ing the outer lip. The parietal lamella is high, oblique, and 
enters rather deeply. The columellar lamella is deeply placed, 
rather short, ascending very little as it enters. There is a 
very low but fairly long lower palatal fold in a specimen in 
Mr. Burnup 's collection (fig. 12), but none in the Connolly 
shell (fig. 15). The peristome is thin, rather well expanded; 
parietal callus very thin. 

The pitting, while similar to that of many Nesopupae, is 
unusually copious. By the characters of the teeth it is an 
Insidipupa, standing close to N. ~barrackporensis and mala- 
yana. Fig. 14 is from a drawing by Mr. Burnup; fig. 15, 

from a specimen lent by Major Connolly. 

i 

59. NESOPUPA MEGALOMASTOMA (Maltzan). Not figured. 

Differs from the Comoro Island type [of Pupa, minutalis 
Morel.] by the larger shell, the last whorl more lengthened- 



NESOPUP^: OP ST. HELENA. 363 

-effuse downward, more impressed transversely behind the 
aperture. Aperture larger for the length of the shell, the 
peristome more expanded. Length 2%, diam. 1% mm ; aper- 
ture 1 mm. high, % wide (Maltzan) . 

West Africa. Nianing, in forest. 

Pupa (Stawodon) minutalis Morel., var. megolomastoma 
MALTZAN, Nachr. d. Malak. Ges., xxii, April 1890, p. 48. 

The name was evidently intended to he megalomastoma, 
;and should be so corrected. 

VIII. ST. HELENA SPECIES. 
Section Helenopupa Pilsbry, n. sect. 

Lamellae and folds as in typical Nesopupa except that the 
columellar lamella curves slightly upward at its inner end; 
surface nearly smooth. 

Like Indopupa, Cocopupa and Insutipupa, this is less 
specialized than the typical Nesopupse. It is notable for the 
smooth surface. 

60. NESOPUPA TURTONI (EJ. A. Smith). PL 33, figs. 11, 14, 15. 

Shell minute cylindric, obtusely conic above, whitish, sub- 
rimate. Whorls 5 to 6, convex, slowly increasing, parted by 
a slightly oblique suture, striated with very oblique, very 
delicate lines, the last whorl hardly descending, with one pit 
in the middle behind the lip. Aperture small, irregularly 
subquadrate, hardly a third of the total length, having 6 
teeth within, three on the parietal wall, one columellar, two 
palatal; peristome white, slightly expanded and reflected, 
continuous, indented above. 

Length 3, diam. 1 mm.; aperture % mm. long and wide 
(Smith). 

St. Helena: Sugarloaf Quarry, common; extinct (Turton). 

Pupa turtoni SMITH, Proc. Zool. Soc. Lond., 1892, p. 268, 
pi. 22, f. 20, 20a, 

Like the St. Helena Endodonts, this species has a Poly- 
nesian aspect. 

In the best preserved specimens the surface is smooth ex- 



364 CAMPOL^MUS. 

cept for some very slight lines of growth, and of a somewhat 
transparent bluish-gray tint. There is a rather long flatten- 
ing of the last whorl, behind the outer lip, in the middle a 
very little impressed over the upper palatal fold. Other ex- 
amples are more chalky and opaque ; and when a little worn, 
long, narrow, external furrows (varying in depth) appear 
over the upper or over both palatals, often with a little pit 
above the labral end of the upper one. These furrows are the 
result of erosion, and not proper to the species in its perfect 
state. 

There is usually a triangular callus where the angular 
lamella joins the upper termination of the lip, which is more 
or less detached from the preceding whorl, and often more 
highly arched than in the example figured. The columellar 
lamella curves upward a little at its inner termination. 
There is a basal fold in form of a low tubercle, subcolumellar 
in position, and so deeply immersed that it is not visible in a 
face view; but often this is reduced to a slight callus which 
would hardly be noticed. 

Length 2.5, diam. 1.2 mm. ; 5% whorls. 

Length 2.2, diam. 1.2 mm.; 5*4 whorls. 

These notes are from two lots collected by Capt. Turton. 
Mr. Smith's measurements are obviously inexact, and do not 
agree with the proportions of his figures. 

Genus CAMPOL^MUS Pilsbry. 

Tomigerus sp., E. A. SMITH, Proc. Zool. Soc. London, 1892 r 
p. 267. 

Campolccmus PILS., Nautilus, vi, Dec. 1892, p. 96, for 
Tomigerus (1) perexilis Smith. 

Shell resembling Tomigerus in miniature; minute, imper- 
forate but with a long basal suture, obtusely conic, the last 
whorl distorted, ascending. Aperture slanting upward, with 
distinct sinulus bounded by the long,i emerging angular 
lamella and upper palatal fold; parietal and columellar 
lamellae and lower palatal fold very deeply immersed ; lip 
well expanded. 

While this snail is astonishingly like Tomigerus in form> 



CAMPOLuEMUS. 365 

yet the teeth are of Pupillid type, and it is believed that 
Nesopupa turtowi is perhaps the most nearly related of known 
species. Several other Pupillid groups, such as Boysia and 
Hypselostoma have the last whorl turned upward. 

1. CAMPOLAEMUS PEBEXILIS (Smith). PL 33, figs. 13, 16-19. 

' ' Shell dextral or sinistral, minute, obtusely pyramidal, im- 
perforate, whitish or pale tawny. Whorls 5, somewhat con- 
vex, slowly increasing, striate with delicate lines of growth, 
the last whorl strongly ascending in front, constricted behind 
the lip, furrowed and distorted, also furrowed basally, ap- 
*pressed to the preceding whorl. Spire very obtuse at apex. 
Aperture transversely ovate, having a nearly circular sinus 
above, narrowed within ; peristome continuous, more or less 
expanded, 

" Length 1.5, greatest diam. 1.5, least 1 mm.; aperture 0.5 
mm. long and wide" (Smith). 

Length 1.55, diam. 1.4 mm. ; 5 whorls. 

St. Helena : Side Path, and the sinistral form from Sugar- 
loaf Ridge; extinct (Capt. Turton). 

Tomigerus(l) perexUis B. A. SMITH, Proc. Zool. Soc., 
1892, p! 267, pi. 22, f. 19-19b. 

This shell is remarkable for the upwardly inclined aper- 
ture and distorted last whorl, with a long basal suture, as in 
the South American genus Tomigerus. 

The first 1% whorls are smooth and unusually wide; 
penult and antepenult whorls have unevenly spaced but 
rather strong, quite oblique striae ; on the last whorl these are 
much weaker. The emerging angular lamella converges 
towards the upper palatal fold, forming a shortly oval, nearly 
tubular sinulus. It penetrates deeply, as far as the anterior 
nd of the parietal lamella. The latter is deeply immersed, 
its forward end visible in a front view, deep within ; becom- 
ing a high lamella, it penetrates to the dorsal side. An infra- 
parietal lamella, more deeply placed, not visible in the aper- 
ture, penetrates about as far as the parietal. The anterior 
nd of the columellar lamella is visible in an oblique view in 
the aperture. It also becomes high and penetrates to the 



366 COSTIGO. 

dorsal side, turning downward at the inner end. The upper 
palatal fold emerges to the lip where it terminates in a 
rounded prominence. It penetrates very deeply, running ob- 
liquely downward. The lower palatal, which is situated far 
within, appears to consist of two rather short, oblique laminae 
meeting below in form of a wide V. The lip is expanded, 
thickened ; parietal callus strong. 

Mr. Smith's figure of the sinistral form referred to this 
species, copied in fig. 13, shows the sinulus less enclosed than 
in the dextral. In the typical form figured by Mr. Smith 
(fig. 17) the upper part of the peristome is some distance 
below the preceding suture. The external pit over the colu- 
mellar lamella and lower palatal fold is broad and rather 
deep. The length and diameter are given as equal by Mr. 
Smith ; in one measured the length exceeds the diameter. 

Mutation ascendens, pi. 33, figs. 16, 18, 19. Suture of the 
last whorl ascending to the preceding suture. Base more 
swollen, the pit smaller. Length 1.5, diam. 1.6 mm. 

Mut. cantrarius, pi. 33, fig. 13. Shell sinistral. Known to 
me by Mr. Smith's figure, here copied, which shows a less 
completely enclosed sinulus. 

Genus COSTIGO Boettger. 

Costigo BTTG., Bericht Senck. naturf . Ges., 1891, p. 270, as 
a section of Vertigo, for V. (c.) saparuana Bttg. 

"Differs from the section Alaea by the scarcely glossy 
shell, densely striate or costulate, the neck simple, without 
annular callus [crest], a single parietal tooth, no angular. 
Columellar tooth always present; palatals to 2. Moluccas 
and Philippines ' ' ( Bttg. ) . 

This group, which appears very nearly related to Neso- 
pupa, is not known to us by specimens. To the species de- 
scribed by Boettger are added a few Mascarene and African 
forms which agree in teeth. It is probably not a natural group 
in its present limits merely a convenience. 

1. COSTIGO SAPAEUANA (Boettger). PI. 31, fig. 12. 

Shell small, punctate-rimate, subacutely ovate, corneous- 



COSTIGO. 367 

apex rather acute. Whorls 5, convex, separated by a deep 
suture, very densely obliquely striatulate and ornamented 
with distant hair-like riblets, the last whorl somewhat en- 
larged, rounded basally, % the height of the shell, more 
strongly ribbed towards the aperture, and not ascending. 
Aperture ample, nearly vertical, semioval, slightly angular at 
the sinulus, 4-toothed, the teeth compressed; 1 parietal, 
simple, oblique, strong; 1 columellar, horizontal, in the 
middle of the columella; palatals either 2, widely separated, 
the lower one stronger, or only a lower fold. Peristome ob- 
tuse, narrowly expanded, colored like the outside, the mar- 
gins joined by a light callus, columellar margin somewhat 
dilated. 

Length 1.75 to 2, diam. 1.125 to 1.25, alt. and width aper- 
ture 0.75mm. (Bttg.). 

Moluccas: Sirisori on Saparua Island (Strubell). 

Vertigo (Costigo) sa^aruana BTTG., Bericht Senck. naturf. 
Ges., 1891, p. 270, pi. 3, f. 12, 12a. 

Distinguished from the externally similar genus Leuco- 
chttus by the single, entirely simple parietal tooth, from the 
oceanic Vertigine group Ptychochilus, which otherwise is very 
closely related, by the absolute lack of an angular tooth. 

2. COSTIGO CALAMIANICA (Moellendorff). Not figured. 

Vertigo calamianica Mlldff., from Busuanga, is mentioned 
as a new species of the section Costigo in von Moellendorff 's 
Verzeichnis, Abhandl. Nat. Ges. Gorlitz, xxii, 1898, p. 152. 
It is evidently the form alluded to by Boettger, following his 
account of C. saparuana, in the following terms: "A second 
species of Costigo lives on Busuanga, Catanduanes, Philip- 
pines ; it is a new species, for which I am indebted to Consul 
Dr. O. Fr. von Moellendorff, and which has no cuticular ribs 
in addition to the sharp striation, and in which palatal teeth 
are lacking." 

3. COSTIGO BORBONICA (H. Adams). PL 33, fig. 7. 

Shell deeply rimate, oblong-ovate, thin, nearly smooth, 
silky, rufous-brown. Spire convexly conic, the apex obtuse, 



368 COSTIGO. 

suture impressed. Whorls 5, a little convex, the last some- 
what compressed at base, swollen and pitted behind the aper- 
ture. Aperture truncate-oval, subvertical, four-toothed : one 
entering, compressed parietal tooth, one tooth deep on the 
columella, two in the palate, the upper one minute. Peri- 
stome a little expanded, white-lipped, the margins joined by 
a thin callus, right margin somewhat sinuous. Length 2 1 /, 
diam. 1% mm. (H. Ad.). 

Bourbon (G. Nevill). 

Vertigo (Alaea) borbonica H. AD., Proc. Zool. Soc. Lond., 
1868, p. 290, pi. 28, fig. 8. 

4. COSTIGO DESMAZURESI (Crosse). PL 33, fig. 12. 

Shell umbilicate, shortly cylindric, thin, translucent, ap- 
pearing smooth to the naked eye, but really marked with fine, 
slightly oblique striae visible only under a lens. Shell a little 
shining and of a light corneous-tawny color. Spire forming 
a short cone terminating in an obtuse summit. Suture well 
marked. Whorls 6, a little convex, the last whorl a little 
smaller than the spire, slightly tapering and rounded at the 
base. Aperture subvertical, of rounded semilunar shape and 
provided with 2 unequal spiral lamellae [palatal folds| situ- 
ated deep within. Peristome simple, narrowly expanded, a 
little reflected, and of a light rosy white ; margins separated ; 
parietal and columellar margins provided each with one in- 
ternal tooth ; these two teeth being placed at right angles to 
one another (Crosse). 

Length 2.25, diam. 1 mm. 

Rodriguez: Pointe aux Coraux (A. Desmazures). 

Pupa desmazuresi CROSSE, Journ. de Conchyl., xxi, 1873, 
p. 140; xxii, 1874, p. 227, pi. 8, f. 3. 

Crosse remarks that one of the palatal folds is more readily 
visible than the other, and it only is shown in the figure. 

5. COSTIGO NOBREI (Girard). PI. 34, fig. 10. 

Shell rimate, oval, very finely striate, as though vermie- 
ulate, corneous, subtransparent. Whorls 4-41^, very convex, 
with a deep suture, the last whorl contained about 2% time* 



COSTIGO. 369 

in the total length, very distinctly contracted near the aper- 
ture, ascending a little, and very distinctly compressed at the 
base. Peristome simple, reflected. Aperture subvertical, 
somewhat rounded, toothed: a parietal fold larger than the 
others, a columellar fold more deeply placed, and two teeth 
on the outer wall, situated still further within the aperture. 
Length about 1%, diam. 1 mm. (Girard). 

Island of San Thome: Binda, under the bark of trees, F. 
Newton. 

Pupa tiobrei GIRARD, Jonial de Sci., Math. Phys. e Nat. 
Acmd. Real Sci. de Lisboa, (2), iii, no. 10, 1893, p. Ill, pi. 1, 
f. 21. 

Compared by Girard with Pupa corpulenta, an American 
Vertigo. It has sculpture like the Nesopupa, apparently, but 
does not appear nearly related to known African forms. By 
the single lamella on the parietal wall it resembles Costigo. 



ADDITIONS AND COEEECTION8, VOL. XXIV 



PAGE 95, 12th line from bottom, the references to figures 
should read: PL 28, figs. 2, 3, Ecuador, and pi. 28, fig. l r 
Duran. 

GASTROCOPTA OP THE MASCARENE ISLANDS. Vol. XXIV, p, 
127. 

M. Germain has proposed a new genus, Falsopupa, for the 
species exigua H. Ad., microscopica Nev., lienardi (=> lieii- 
ardiana) Crosse and desmazuresi Crosse (Bull. Mus. Nat. 
d'Hist. Nat. Paris for 1918, no. 7, p. 521; this number re- 
ceived at the Academy Oct. 14, 1919). 

This list contains species of two genera. No diagnosis of 
the group was given, and no species was selected as type. I 
take Pupa lienardiana Crosse (Falsopupa lienardi Crosse of 
Germain's list) to be the genotype. If the views of the 
affinities of the Mascarene species expressed in Vol. XXIV, 
pp. 127-133, are upheld, Falsopupa will become a synonym 
of the typical section of Gastrocopta. 

Genus ABIDA, vol. XXIV, p. 262. 

Add to the synonymy : 

Pupa Draparnaud, Tableau des Moll. terr. et fluv. de la 
France, 1801, pp. 32, 56, for muscorum [Drap. not Linne], 
pygmaea, antivertigo, vertigo [= pusttla Mull.], umbilicata, 
marginata, doliolum, dolium, granum, avena, frumentum, 
secale, polyodon, variabilis, 4-dens, 3-dens, cinerea, and vari- 
ous species of Clausilia and Balea. Pupa frumentum may be 
selected as type. Not Pupa Bolten 1798, or Pupa Lamarck, 
1801. 

Deloplecta Agassiz, Nomenclator Zoologicus, 1842-6, Mol- 
lusca, p. 29, there said to "= Pupa". Agassiz, in the same 

(370) 



ADDITIONS AND CORRECTIONS. 371 

work, mentions Pupa Humph., Mus. Calonnianum (a nomen 
nudum), and Pupa Drapamaud. Deloplecta will be consid- 
ered equivalent to the latter, with the same type, Pupa fru~ 
mentum. Agassiz gives the reference "Charp., Cat. Moll. 
Suisse, 1837, ' ' but the name does not occur in that work. 

Pupella SWAINSON, Treatise on Malacology, 1840, pp. 183, 
334 (P. variabttis here designated as type). Not Pupetta 
B. St. Vincent, 1825 (Infusoria). 

PAGE 264. Under the caption "Radula", the reference to 
Proc. Malac. Soc. London should be Vol. II. 

PAGE 307, tenth line from bottom, for "Bull. Soc. d'Agric." 
read: Ann. Soc. d'Agric. The same correction should be 
made on pp. 311 and 314. 

"PUPA" PYRENAICA Boubee. r 

Pupa hospitii (p. 37) is said to be a synonym of Pupa* 
(ModiceUa) pyrenaica Boubee (B., H. & d'A-A., Fauna Mala- 
cologia del Pirineu Catala, i, 1818, p. 99). The writer has 
not seen Boubee 's publication. According to Westerlund,. 
Clausitia pyrenaica Boubee 1833 is identical with Pupa 
pyren&aria (Boub.) Mich., and P. pyrenaica Boubee 1839 is 
P. ring em (Calliaud) Mich. It is evidently Abida "pyrena- 
aria" which Bofill, Haas and d'Aguilar-Amat had in view. 
See Man. Conch. XXIV, pp. 278, 288. The status of Bou- 
bee 's species must be determined by reference to the original 
publication. 

VOLUME XXV. 

CHONDRINA GONIOSTOMA (Kiister). P. 33. Bofill, Haas and 
d'Aguilar-Amat consider Pupa angulata Fagot (p. 34) to be 
one extreme of this species, P. leptocheilos Fagot (p. 31) the 
other, the first having the mouth more angular, the second 
less so (Fauna Malacologia del Pirineu Catala, i, 1918, p. 95). 
As stated in the text, P. leptocheilos was proposed as a sub- 
stitute for tenuimarginata. 



372 ADDITIONS AND CORRECTIONS. 

CHONDRINA ARAGONICA (Fagot). P. 54. P. saltus (p. 51) 
is a form of this according to the authors mentioned above 
(op. cit., p. 97). 

VERTIGO ALABAMENSIS CONECUHENSIS, p. 80. The authority, 
Clapp, should be substituted for "n. subsp." 

VERTIGO OVATA Say. P. 82. 

This species occurs also in the southeastern U. S. Virginia, 
western North Carolina, Georgia and Florida (Sterki collec- 
tion) ; Florida at Snapper Creek hammock, Dade Co., Lower 
Matacumbe Key and Madeira hammock near the end of the 
peninsula (Geo. H. Clapp coll.). 

VERTIGO VENTRICOSA. P. 94. 

Dr. Sterki writes: "I believe that V. ventricosa is closely 
allied to V. ovata, not to V. pygm&a. I have repeatedly had 
forms in doubt between the two. Both have essentially the 
same formation of the palate, and there are small ovata with 
a slight palatal callus, occasionally no infraparietal and a 
barely vestigeal angular lamella." 

VERTIGO HEBARDI Van. P. 103. 

Additional localities are: Elliott's, Pumpkin, Lignum Vitae, 
No Name, Big Pine and Little Palo Alto keys, off south and 
southeastern Florida, Geo. H. Clapp collection. 

VERTIGO CONCINNUDA Ckll. P. 121. 

The shells illustrated in fig. 1 are from Willow Creek, 
Mogollon Mts., New Mexico; x 15. 

VERTIGO MODESTA (pp. 123-134). 

On p. 134 a race of V. modesta from near Bluff Lake in the 
San Bernardino Mts., California, was discussed. It was fig- 
ured on p. 124, figs. 8, 80, 8&. Without locality it would not 
be possible to separate some specimens from. V. m. castanea, 
while others would be as near V. m. parietalis. These ex- 



ADDITIONS AND CORRECTIONS. 373> 

tremes, however, are fully connected in the Bluff Lake series, 
which must thus be treated as a unit. As in the Santa Cata- 
lina race of this species (p. 132), albino shells occur in some 
abundance. As in many other cases, the albino mutation 
occurs in colonies with the brown shells, and there is no 
reason to believe that it does not interbreed with them, as has 
been demonstrated for a number of other such cases. Dr. 
Berry has described the albino mutation as V. m. micro- 
phasma. In the writer's opinion the brown shells found with 
them are of the same race ; but the name can be used for the 
San Bernardino race as a whole, with its several variations 
and mutations. Though certainly without definite differen- 
tial characters, it is not, as a whole, either typical castanea or 
typical parietalis. 

VERTIGO MODESTA MICROPHASMA Berry. Figs. 1-6. 

"The shell is cylindro-conic, rimate-umbilicate, thin, very 
pale horn-color, by transmitted light transparent and color- 
less. The surface is glossy and distinctly, irregularly, ob- 
liquely striate, especially on the upper whorls. The spire 
tapers from the last whorl, at first gradually, then more 
rapidly, to the obtuse apex. The whorls are strongly convex, 
the last with an indentation just back of the aperture over 
the lower palatal tooth, subsequently with a narrow, abrupt, 
axial constriction, then swollen to form a low, wave-like crest 
just back of and parallel to the lip. The aperture is rounded 
triangular, scarcely constricted on the outer margin, the peri- 
stome thickened and porcelain white in color, showing through 
the back of the shell as a white line, but the sharp lip scarcely 
reflected except over the columella. The posterior angle of 
the outer lip curves in rather sharply to the body whorl. The 
number of teeth varies from 2 to 5. The parietal and colu- 
mellar lamellae are always well developed. In addition there 
is almost always a well-developed lower palatal. A smaller, 
but variable upper palatal is frequently present, as also a 
minute angular lamella. All the teeth are porcelain-white in 
color. 



374 



ADDITIONS AND CORRECTIONS. 




1-6, VERTIGO MODESTA MICROPHASMA. 7, VERTIGO ALLYNIANA. 
8, V. ALLYNIANA mut. XENOS. 

"Length of type 2.6; diameter to lip edge 1.5; length of 
aperture 0.9 mm.; whorls 5. 

"Type: Cat. no. 2740 of the writer's collection, fig. 2. 
Paratypes in the Academy of Natural Sciences of Philadel- 
phia, California Academy of Sciences, Southwest Museum, 
United States National Museum, and the private collections 
of Mr. G-eorge H. Clapp and others. 

"Type Locality: 7,550 feet altitude, cienaga near Bluff 
Lake, San Bernardino Mountains, California; under sticks 
and logs at edge of forest; Nina G. Spaulding, G. E. Dole 



ADDITIONS AND CORRECTIONS. 375 

S. S. Berry, August, 1910; 59 specimens in this and 
neighboring cienagas. Also taken at 7,200 feet altitude, west 
slope of Falls Creek Canyon, near the narrows about one 
mile above Dobbs Cabin, Dollar Pass Trail, San Bernardino 
Mountains, California ; under small sticks and pine cones on 
springy slope ; G. E. Dole and S. S. Berry, Sept, 29, 1918 ; 
32 specimens" (Berry). 

Vertigo modesta microph-asma BERRY, Nautilus, xxxiii, Oct. 
1919, p. 48, figs. 1-6. 

1 * This very puzzling little mollusk is one of the most beau- 
tiful of American Vertigos. It is very close to V. modesta 
parietalis and may also be described as an albinistic race of 
that subspecies, but it is a protean form and some shells are 
equally close to V. modesta modesta or even to V. m. castanea, 
That it is more than a mere 'albino' of the recognized type 
is strongly evidenced by its occurrence in such abundance and 
at scattered localities, as also by the fact that its distribution 
is by no means coincident with that of any of the other forms 
mentioned. Nor, although usually associated, do the white or 
brown shells occur in any apparent regular ratio. At the 
second locality above cited diligent outlook yielded but three 
specimens of the brown parietalis. It is evidently a com- 
paratively recent offshoot from the parent stock, but the field 
evidence is that it already is a race with its peculiar char- 
acters heritable to a marked degree. 

1 i It seems rather remarkable that such features as the color, 
shell texture, and similar characters in this form should ex- 
hibit such constancy as compared with the variability shown 
in the development of the lamellae. In 39 specimens of the 
type lot now before me, 1 has only 2 teeth (columellar and 
parietal), 15 have 3 teeth (columellar, parietal, and lower 
palatal), 9 have 4 teeth (an upper palatal usually the one 
added), and 14 have a full set of 5 teeth. No mature speci- 
mens with fewer than 2 nor more than 5 teeth have been 
noted. This variation in a single well-defined colony (its 
members having, as shown by the other characters noted, an 
undoubtedly close phylogenetic relationship with one an- 
other) throws a valuable bit of light on the difficulty of at- 



376 ADDITIONS AND CORRECTIONS. 

tempting the separation of the various races of the modesta 
series by means of variations in the number of teeth alone. 
It chances that the specimen chosen as type is one of the 
3-toothed forms. 

' ' The animal is bluish-gray or slate in color, the body quite 
dark, the foot and peripheral portions much lighter and semi- 
transparent. A rough sketch of the cephalic region of one of 
the Falls Creek specimens is offered in fig. 6. 

"Whether the hereditary value of this race is that of a 
'form' or a subspecies can only be shown by the more de- 
tailed study which must be left for the future. Until then 
the personal equation must necessarily largely govern. In 
any case it will prove useful to have a name for it" (Berry), 

VERTIGO ALLYNIANA Berry. Fig. 7. 

"The shell is minute, short, robust, ovate-conic in outline, 
thin, dark reddish-brown in color, with only a dull gloss; 
weakly, irregularly striate. The spire tapers with increasing 
rapidity from the last whorl to the obtuse apex. The whorls 
are convex, the last having a shallow but distinct excavation 
in the palatal region and a weaker one over the upper palatal 
tooth, the latter extending to the lip, which thus becomes 
flattened or very slightly indented on its outer segment. The 
aperture is pyriform in outline, and would be rather small 
except for the quite flaring lip, which is little thickened and 
very fragile at the edge. There are 5 teeth constantly de- 
veloped in all the material examined. The parietal, columel- 
lar, and upper and lower palatal lamellae are well developed, 
and there is a distinct, though small angular lamella. The- 
columellar is situated well back in the aperture and quite 
high up on the pillar. The lower palatal is also rather deeply 
immersed. 

"Length of type 2.1; diameter to lip edge 1.3; length of 
aperture 0.81 mm. ; whorls 4%. 

"Type: Cat. no. 3764 of the writer's collection. Paratypes 
in the Academy of Natural Sciences of Philadelphia, and the 
private collection of Allyn G. Smith. 



ADDITIONS AND CORRECTIONS. 377 

"Type Locality: Donner Lake, California; A. G. Smith, 
May 30, 1916; 22 specimens" (Berry}. 

Vertigo aUyniana BERRY, Nautilus, vol. 33, Oct. 1919, p. 51, 
fig. 7. Vertigo allyniana xenos BERRY, op. cit., p. 52, f. 8. 

""I am not quite certain of the relationships of this small 
Vertigo. The texture of the shell, as well as the shape, are 
strongly reminiscent of V. occidentals Sterki, a more weakly- 
toothed species from the San Bernardino Mountains. None 
of the other species with which I am familiar require any 
special comparison. V. corpulenta (Morse) has a somewhat 
similar outline, but otherwise does not seem especially close" 
(Berry). 

Two paratypes measure : 

Length 2.18, diam. 1.4 mm. 

Length 2.1, diam. 1.27 mm. 

The holotype of V. a. xenos measured the same way : 

Length 2.1, diam. 1.45 mm. 

The latter form, based upon a single shell, found with 
typical attyniana, appears likely to be merely a shape muta- 
tion. Variation of contour equal to this is of common occur- 
rence in single colonies of many Vertigos, and in the absence 
of series sufficient to show that there is racial differentiation 
they are better left nameless. 

In washing a specimen of the typical allyniana the color 
and texture appear to be exactly as in the type of xenos. 

Mut. xenos (Berry), fig. 8. "With the preceding occurred 
a single specimen of a very similar form having the same 
number of teeth, but differing abruptly in its shorter, much 
more robust and swollen outline, its more transparent, glos- 
sier texture, and lighter brown color. The columellar tooth 
is placed distinctly further down on the pillar, and the re- 
maining lamellae differ slightly from those of the shells de- 
scribed above both in size and position. Length of type 2.0; 
diameter to lip edge 1.5 ; length of aperture 0.85 mm. ; whorls 
4i/ 2 . Type: Cat. no. 4128 of the writer's collection" (Berry). 

VERTIGO CALIFORNIA ELONGATA (Sterki). Page 142. 

The name el-ongata is several times preoccupied in both 
Pupa and Vertigo. It may stand as Vertigo calif ornica longa. 



378 ADDITIONS AND CORRECTIONS. 

VERTIGO ROWELLJ (Newc.). P. 143. 

Two specimens from Oakland, type locality, measure : 

Length 2.8, diam. 1.4 mm.; nearly 6 whorls. 

Length 2.48, diam. 1.3 mm. ; 5% whorls. 

The minute costulation is rather pronounced on the penult 
whorl. The low, rounded rib-striae are about as wide as their 
intervals. On the last whorl they become weak, more and 
irregularly spaced. The long, convexly tapering spire is 
rather turrited. 

VERTIGO OSCARIANA St. P. 144. 

Found at Ft. Lauderdale and BriekeH's hammock below 
Miami by Geo. H. Clapp. 

PAGE 146, 15th line, for "columellar fold" read columel- 
lar lamella. 

VERTIGO MILIUM Gld. P. 146. 

Further localities in south and southeastern Florida: Ft. 
Lauderdale, Brickell's Hammock, Snapper Creek Hammock, 
Snake Creek Hammock, Pumpkin and Big Pine Keys; Fla- 
mingo and Coot Bay, Cape Sable, Madeira Hammock (Geo. 
H. Clapp coll.). 

PAGE 184, 7th line, for "Grenada" read Granada. 

P. 212, 8th line, for "V. atwustiOa" read V. angustior. 

VERTIGO THIBETIC A (Bens.). 

Pupa thibetica Bens. (vol. xxiv, p. 138) has been placed in 
Vertigo by "Westerlund ; as it is in the Palaearctic fauna, and 
is said to be smooth, translucent and glossy, this reference is 
probably correct. It has not been figured. 

VERTIGO PUSILLA mut. ALBINA Taylor. Shell translucent- 
white. Found with the typical form at Portsalon, near Let- 
terkenny, Donegal County, Ireland (J. W. Taylor, Journ. of 
Conch., vii, 1893, p. 194). 



ADDITIONS AND CORRECTIONS. 379 

VERTIGO PUPAEFORMIS Pollonera. Vol. XXVI, pi. 5, fig. 10. 

Shell small, ovate-cylindric, very lightly striatulate ; whorls 
6, convex, the last one-third the total length, provided with a 
strong, transverse callus [crest] anteriorly. Aperture sub- 
ovate, strongly five-toothed : 1 parietal tooth, 2 columellar, the 
lower one smaller, and 2 pliciform palatals, the lower one 
larger. Length 2, diam. 1.25 mm. (Pollonera). 

Italy : Val Salice, colline di Torino, Pleistocene. 

Vertigo (Dexiogyra) pupaeformis POLLONERA, Mem. Reale 
Accad. Sci. Torino, (2), xxxviii, 1888, p. 30, fig. 22 of the 
plate. 

This form, which Pollonera states has not been found living 
in Piedmont or elsewhere, is said to differ from V. pygnuza 
by the less swollen, more cylindric form, the last whorl being 
less developed. 

VERTIGO (ALAEA) TROLLI Fischer and Wenz. Jahrb. nassau. 
Ver. Nat., vol. 67, 1914, p. 102, pi. 7, f. 27. Upper Miocene 
of Oppeln. 

VERTIGO ( ? GLANDICULA) GAALI Wenz. Senekenbergiana, i, 
no. 3, 1919, p. 67. New name for Pupa (LeucochUa) cf. 
larteti Gaal, Mitt. a. d. Jahrb. ungar. geol. Eeichsanst., xviii, 
1911, p. 68, pi. 2, f. 4. 

VERTIGO DIVERSIDENS (Sandb.), this volume, p. 218, includes 
the following according to Dollfus, Etude sur la Molasse de 
1'Armagnac, in Bull. Soc. Geol. France (4), xv, 1915, p. 362: 
V. laemodonta,, V. sansanica and V. callostoma Bgt. 

VERTIGO RHYNCHOSTOMA Bgt., this volume, p. 219, includes, 
according to Dollfus, op. cit., p. 363: V. onixiodon and V. 
micrcmixia Bgt. This species was found in the Noulet collec- 
tion under the name P. vascanensis Noulet Ms. 

PAGE 220. Vertigo trigonostoma, blumi and priscilla are 
to be removed from Ptychalaa and placed in Nesopupa, where 
they appear to be related to the section Indopupa. Vertigo 
capellinii (p. 221) belongs either to Nesopupa or to Ptycha- 
laa, probably the former. 



380 ADDITIONS AND CORRECTIONS. 

Paetel's Catalog contains many errors in spelling and 
authorities which are not worth noting here. The following 
are samples : 

[Pupa (Vertigo)} eurieysii Drouet. I. Trinidad. Paetel, 
Catalog der Conchylien-sammlung von Fr. Paetel, edit. 1883, 
p. 159. Error for Pupa eyriesii Drouet. 

[Pupa (Vertigo)} turgida Zglr. Illyr. Paetel, op. cit., p. 
160. 

[Pupa (Alaea)] achila Sow. Hispan. Paetel, op. cit., 4th 
edit., 1889, ii, p. 293. Error for Vertigo acheila Serv. 

VERTIGO MURCHISONIAE Moore, Quart. Journ. Geol. Soc. 
London, xxiii, 1867, p. 557, pi. 15, figs. 11, 12, from the Lias- 
sic deposit in Charter-House mine, Somerset, England, is a 
poorly preserved sinistral fossil, certainly not a Vertigo. 



EXPLANATION OF PLATES, VOL. XXV. 



Except when otherwise stated, the figures were drawn 
from specimens in the Academy of Natural Sciences. Those 
marked B. M. after the museum number are in the Bishop 
Museum, Honolulu. 

PLATE 1. 

FIGURE PAGE 

1. Chondrina megacheilos C. & J. Milan. 22754 7 

2. 3. Chondrina megacheilos C. & J. Como. 3843 7 

4, 5. Chondrina avenacea Brug. Metz. 22776 10 

6, Chondrina avenacea Brug. Sicily. 22769 10 

7, 8. Chondrina a. clienta West. Herkulesbad, Banat. 

66045 15 

9, 10. Chondrina a. riviana Schr. Riva. 101617 18 

Fig. below 7. Chondrina a. duplicata Kiister. After 

Kiister . . . 14 

Fig. below 8. Chondrina subhordeum West. After Kiis- 
ter 13 

11. Chondrina cereana Kiist. After Kiister 21 

12. Chondrina apuana Iss. After Del Prete 15 

13, 14. Chondrina oligodonta Del Pr. After Del Prete. . 19 

15. Chondrina cianensis Caz. After Caziot 20 

PLATE 2. 

1, 2, 7. Chondrina bigorrwmsis Ch. Bigorre. 22762 29 

3. Chondrina bigorriensis Ch. After Des Moulins 29 

4. Chondrina tenuimarginata Des M. Pyrenees. 3894. . 31 

5. Chondrina tenuimarginata Des M. After Des Moulins. 31 

6. Chondrina t. elongatissima Des M. After Des Moulins. 32 

8, 9. Chondrina b. moquiniana Kiist. After Kiister .... 30 
10-12. Chondrina arigonis Rm. After Rossmaessler .... 34 
13, 14. Chondrina goniostoma Kiist. After Rossmaessler. 33 
15, 16. Chondrina g. juliensis Bgt. After Bourguignat . . 34 
17, 18. Chondrina arigonis Rm. Pego. 22792 34 

PLATE 3. 

1. Chondrina arigonis Rm. Catalonia. 114976 34 

(381) 



382 EXPLANATION OP PLATES, VOL. XXV. 

FIGURE PAGE 

2, 3, 4. Chondrina k. ordunensis Pils. Pefia de Orduna. 

22791 39 

5. Chondrina hisitanica Em. After Eossmaessler 40 

6. Chondrina lusitanica Em. Portugal. 22795 40 

7. 8. Chondrina gigantea. After Eossmaessler 32 

9. Chondrina tingitana Kob. Tetuan. 115001 41 

10, 11. Chondrina calpica West. Gibraltar. 78391 41 

12. Chondrina gorbeana Pils. Pena de Gorbea. 22779.. 56 

13. Chondrina lallemantiana Bgt. After Bourguignat . . 43 

14. Chondrina letourneuxi Bgt. After Bourguignat 42 

PLATE 4. 

1. Chondrina similis Brug. Nice. 22751 57 

2. Chondrina similis Brug. Florence. 101600 57 

3. Chondrina similis Brug. Grasse. 22739 57 

4. Chondrina similis Brug. Marseilles. 22747 57 

5. Chondrina pallida amicta Parr. Nervi. 4942 63 

6. Chondrina pallida amicta Parr. Grasse. 44795 63 

7. Chondrina pallida Phil. After Eossmaessler 63 

8. Chondrina pallida amicta Parr. Spezia. 78390 63 

9. Chondrina psarolena Bgt. After Bourguignat 65 

10. Chondrina bergomensis Ch. Bergamo. 115007 .... 22 

11. Chondrina psarolena Bgt. After Eossmaessler 65 

12, 13. Chondrina circumplicata Mss. Lombardy. 114980. 22 

PLATE 5. 

1. Chondrina farinesii Des M. Pyrenees. 22787 45 

2. Chondrina farinesii Des M. La Preste. 67669 45 

3. Chondrina boettgeriana 01. After Clessin 49 

4. Chondrina farinesii dentiens Moq. After Moquin- 

Tandon 47 

5. Chondrina massotiana Bgt. After Bourguignat 52 

6. 7. Chondrina jumillensis Guir. After Bourguignat . . 49 

8. Pseudelix microhelix Sandb. After Sandberger 222 

9, 10. Chondrina obliterata Ch. After Kiister 52 

11. Chondrina m. penchinatiana Bgt. After Bourguignat. 53 

12. Enneopupa cylindrella Al. Br. After Sandberger . . 222 

13, 18. Vertigo angustior Jeffr. Lyons. 3801 211 

14, 15. Glandicula tiarula Al. Br. After Sandberger ... 221 

16, 17. Vertigo angustior Jeffr. Lausanne. 22901 211 

PLATE 6. 

1, 2. Vertigo ovata Say. Upper Eed Hook, N. Y. 48553. 82 



EXPLANATION OP PLATES, VOL. XXV. 383 

FIGURE PAGE 

3. Vertigo ovata Say. Oswego, Oregon. 111397 82 

4. 7. Vertigo ovata Say. Hamilton, Ontario. 62900 82 

5. 6. Vertigo o. mariposa Pils. Mariposa Co., Cal. 11644. 88 

8, 9. Vertigo morsei St. Lake James, Ind. 105714 81 

10, 13. Vertigo berryi Pils. San Bernardino Mts. 105166. 89 

11, 12. Vertigo o. diaboli Pils. Devil's River, Texas. 90437 88 

PLATE 7. 

1. Vertigo ventricosa Mse. Tignish, Prince Edwards I. 

106971 94 

2. Vertigo ventricosa Mse. Hebron, Maine. 87273 94 

3. Vertigo ventricosa Mse. Buekfield, Maine. 87294 ... 94 

4. 5. Vertigo gouldii Binn. Brookline, Mass. 82690 .... 98 

6. Vertigo elatior St. Woodland, Aroostook Co., Maine. 

79782 95 

7. Vertigo perryi St. Duxbury, Mass. 14830 103 

8. Vertigo gouldii Binn. Brookline, Mass. 82690 . 98 

9, 10. Vertigo bollesiana Mse. Buekfield, Maine. 87312.. 101 
11, 12. Vertigo pygmaea Drap. Upper Red Hook, N. Y. 

48554 96 

13-15. Vertigo nylanderi St. Woodland, Maine. 98331.. 100 

PLATE 8. 
[Pigs. 1-7x19; figs. 4-8, 10-14x25.] 

1, 2. Vertigo rugosula St. De Soto Parish, La. 87438 . . 77 

3. Vertigo rugosula St. Sullivan's I., S. C. 60462 77 

4. Vertigo hebardi Van. Long Key, Fla. 106359 103 

5. Vertigo oralis St. Tallapoosa R., 7 miles above We- 

tumpka, Ala. 91060 78 

6. 7. Vertigo oralis St. Volusia Co., Fla. 11654 78 

8, 10, 11. Vertigo oscariana St. Volusia Co., Fla. 60463. 144 
12. Vertigo a. conecuhensis Clapp. Evergreen, Ala. 

113414 80 

9, 13, 14. Vertigo alabamensis Clapp. Tuscaloosa Co., 

Ala. 113412 79 

PLATE 9. 

1, 2. Vertigo californica Rowell. Paratypes. San Fran- 
cisco. 59392 139 

3. Vertigo c. cyclops St. Rocklin, Cal. 79817 141 

4. Vertigo c. longa Pils. (elongata St.). S. Clemente I. 

57856 142, 377 

5. Vertigo c. catalinaria St. S. Barbara I. 113847 142 



384 EXPLANATION OF PLATES, VOL. XXV. 

FIGURE PAGE 

6. Vertigo c. catalinaria St. S. Catalina I. 62362 142 

7. Vertigo rowelli Newc. Douglas Co., Ore. 76372 143 

8. Vertigo c. diegoensis St. San Ramon. 64578 141 

9. Vertigo c. trinotata St. Monterey. 62363 140 

10. Vertigo c. cupressicola St. Cypress Point. 118835.. 143 

11. Vertigo c. diegoensis St. San Ramon. 64578 141 

12. Vertigo columbiana St. Vancouver I. 68881 108 

13. Vertigo columbiana St., var. Olympia. 11661 108 

PLATE 10. 

1. Vertigo modesta Say. Laggan, Alberta, 76375 123 

2. Vertigo modesta Say. Loess, Des Homes, la, 79797. . 123 

3. Vertigo m. corpulenta Mse. Ogden canvon, Utah. 

11663 130 

4. Vertigo m. parietalis Anc. Ogden canyon, Utah. 

11659 128 

5. Vertigo m. castanea St. Rae Lake, Cal. 115214 132 

6. Vertigo m. castanea St. Type, Fish Camp, Cal. 11655. 132 

7. 8. Vertigo arctica Wallenb. After Wallenbaum 189 

8. Vertigo concinnula Ckll. Type, Custer Co., Colo. 

59095 119 

10. Vertigo concinnula Ckll. Jemez Mts., N. M. 73587. . 119 

11. Vertigo occidentals St. Bluff Lake meadow, San 

Bernardino Mts. 105048 134 

12, 13. Vertigo m. insculpta Pils. Mt. Lemon, S. Cata- 

linas. 109538, 109547 131 

PLATE 11. 

1. Vertigo dalliana St. Type, 416 Sterki coll 137 

2-5. Vertigo sterkii Pils. Type and paratypes. 43706 . . 112 

6, 8. Vertigo a. sanbernardinensis Pils. Cienaga below 

Bluff Lake. 118419 Ill 

7. Vertigo a. sanbernardinensis Pils. 20 miles north of 

Yosemite. 114796 Ill 

9. Vertigo andrusiana Pils. Oswego, Ore. 11398a 109 

10. 11. Vertigo andrusiana Pils. (Fig. 10 the type.) 

Douglas Co., Ore. 76380 109 

12. Vertigo binneyana St. Paratype. Helena, Mont. 

11672 90 

PLATE 12. 

1-3. Vertigo tridentata Wolf. Fig. 2 the type. Canton, 

111. 58008 . 106 



EXPLANATION OF PLATES, VOL. XXV. 385 

FIGURE PAGE 

4, 5. Vertigo gouldii cristata St. Paratype. Quebec. 

119008 100 

6,, 8. Vertigo gouldii paradoxa St. Paratype. Woodland, 

Me. 119007 99 

7, 9. Vertigo parvula St. Type, 270 Sterki coll 105 

10. 11. Vertigo coloradensis inserta Pils. Bear Wallow. 

109559 118 

12. Vertigo hannai Pils. Type. U. S. N. M 114 

13. Vertigo coloradensis Ckll. (type of V. columbiana 

utahensis St.). Box Elder Co., Utah. 119009. . 115 
14, 16. Vertigo c. arizonensis P. & V. Type. 119010 ... 117 

15. Vertigo c. basidens P. & V. Bland, N. M. 79467 . . . 117 

PLATE 13. 

1-5. Vertigo miliuin Old. Near Cleveland, O. 95913 ... 146 

6. Vertigo milium Old. Mt, Taylor, Volusia Co., Fla. 

43707 146 

7. Vertigo miliuin Old. Vermont. 58204 146 

8. 10, 12. Vertigo bermudensis Pils. Type (f. 10, 12) 

and paratype. 105610 149 

9. Vertigo bermudensis Pils. 91159 149 

11. Vertigo neglecta Poey. After Poey 87 

13. Vertigo ovata var. S. Domingo. 43705 87 

14. 15. Vertigo numellata Gul. Bermuda. 91158 91 

16. Vertigo ovata var. Porto Rico. 3807 87 

17. Vertigo marki Gul. Type ; Bermuda. 85574 107 

PLATE 14. 

1, 2. Vertigo eogea Pils. Akkeshi. 89899 151 

3. Vertigo eogea Pils. Kashima. 83396 152 

4, 5. Vertigo hydrophila Reinh. After Reinhardt 152 

6, 7. Vertigo hachijoensis Pils. Type. 83394 153 

8-10. Vertigo kusbiroensis Pils. Type. 90223 153 

11. 12. Vertigo hirasei Pils., var. Miyakejima. 86484 . . . 155 
13, 14. Vertigo hirasei Pils. Type. 79738 154 

15. Vertigo hirasei glans Pils. Type. 82689 155 

16. Vertigo hirasei okinerabuensis Pils. Type. 87690 . . 155 

PLATE 15. 

1, 2. Vertigo japonica Pils. Type. 85746 155 

3. Vertigo j. tosana Pils. Irazuyama. 86486 156 

4. Vertigo j. coreana Pils. Fusan. 95772 156 

5. 6, 7, 9. Ptychalaea dedecora Pils. Hahajima. 82583.. 158 



386 EXPLANATION OF PLATES, VOL. XXV. 

FIGURE PAGE 

8. Vertigo denudata Mss. After Mousson , . 156 

10, 11. Ptychalaea d. tamagonari P. & H. Type. 85745. 158 

12. Staurodon s. seminulum Lowe. 97298 225 

13, 14. Staurodon saxicola Lowe. 4930 224 

15. Ptyehalaea flexidens Rss. After Wenz 157 

PLATE 16. 

1. Vertigo pusilla Mull. Lausanne. 22898 161 

2, 3. Vertigo pusilla Mull. Niedermodau. 123426 161 

4, 5. Vertigo antivertigo Drap. Lyons. 3797 163 

6. Vertigo antivertigo Drap. Calvados. 22890 163 

7. Vertigo sinuata Mouss. After Mousson 166 

8. 9. Vertigo discheilia Bgt. After Bourguignat 170 

10. 11. Vertigo maresi Bgt. After Bourguignat 171 

12. Vertigo codia Bgt. After Bourguignat 184 

13. Vertigo microlena Bgt. After Bourguignat 185 

14. Vertigo aprica Bgt. After Bourguignat 184 

15. Vertigo loroisiana Bgt. After Bourguignat 182 

PLATE 17. 

1. Vertigo moulinsiana Dup. Lyons. 22882 178 

2, 3. Vertigo moulinsiana Dup. Cambridge. 109428 ... 178 

4. Vertigo substriata mitis Bttg. After Boettger 173 

5, 6. Vertigo m. ventrosa Heyn. After Heynemann 181 

7. "Vertigo" subtrochiformis Greg. After de Gregorio. . 214 

8. "Vertigo" cylindrica Colb. After Colbeau 214 

9. Vertigo haeussleri St. After Sterki 178 

10. Vertigo substriata Jeffr. 109425 172 

11. 12. Vertigo sieversi Bttg. After Boettger 195 

13. Vertigo s. punctulum Bttg. After Boettger 196 

14. Vertigo ronnebyensis West. After Geyer 192 

15. 16, 17. Vertigo pygmaea Drap. Lyons. 3799 . . 174 

PLATE 18. 

1. Vertigo alpestris Aid. Halker, England. 109427 197 

2. Vertigo alpestris Aid. Lowenburg, Transylvania. 

114999 197 

3. Vertigo a. shuttleworthiana Ch. After Steenberg. . . . 199 

4. Vertigo heldii 01. ( ? = pygm8ea). After Geyer 199 

5. 6. Vertigo leontina Gredl. After Gredler 201 

7, 8. Vertigo isarica West, (leontina Clessin). After 

Clessin 201 

9. Vertigo schultzii Phil. After Kiister 202 



EXPLANATION OF PLATES, VOL. XXV. 387 

FIGURE PAGE 

10-12. Vertigo genesii Gredl. Topotype. After Sand- 
berger 204 

13. Vertigo alpestris (tridensSandb.). After Sandberger. 208 

14. Vertigo alpestris (quadridens Sandb.). After Sand- 

berger 208 

15. Vertigo genesii parcedentata (bidens). After Sand- 

berger 207 

16. Vertigo alpestris (adversidens Sandb.). After Sand- 

berger 208 

17, 18. Vertigo genesii Gredler. After Gredler 204 

19. Vertigo g. parcedentata (glandicula) Sandb. After 

Sandberger 207 

PLATE 19. 

I, 2. Lyropupa lyrata Gld. (Paratype of Pupa magda- 

lena Anc.) Palama, Oahu. 18745BM 233 

3. Lyropupa lyrata Old. (Paratype of Pupa carbonaria, 

Anc.) Nuuanu, Oahu. 18752BM 233 

4. Lyropupa lyrata Gld. Type. 219G, 2687 N. Y. State 

Museum 233 

5. Lyropupa lyrata Gld. Paratype, same museum 233 

6. Lyropupa lyrata Gld. Nuuanu. 108336 233 

7. Lyropupa 1. fossilis C. & P. Manoa. 119462 237 

8. 9. Lyropupa lyrata, form gouldi P. & C. Type and 

paratype. New York State Museum 235 

10. Lyropupa 1. uncifera P. & C. Glen Ada, 119422 ... 236 

II. Lyropupa 1. fossilis C. & P. Type. Manoa. 11039BM. 237 
12, 13. Lyropupa 1. uncifera C. & P. Cotypes. 1% miles 

west of Kahuku, Oahu 236 

PLATE 20. 

1. Lyropupa lyrata var. Limahuli, Kauai. 11038BM... 237 

2. Lyropupa rhabdota C. & P. Pelekunu, Molokai. 

11040BM 239 

3. 5. Lyropupa r. pluris P. & C. W. ravine upper Kamalo. 

119427 240 

4. Lyropupa r. pluris P. & C. Pipe-line trail, Kauna- 

kakai. 48625 24Q 

6. Lyropupa r. lanaiensis Cooke. Lanai. 11041BM .... 241 

7, 8. Lyropupa baldwiniana Cooke. lao, W. Maui. 

11042BM 241 

9. 10. Lyropupa prisca Anc. Mana. 119453 243 

11. Lyropupa prisca Anc. Mana. 11044BM 243: 



388 EXPLANATION OF PLATES, VOL. XXV. 

PIGURE 

12. Lyropupa thaanumi C. & P. Auwahi. 11043BM ... 242 

13. Lyropupa thaanurai C. & P. Auwahi. 119452 242 

PLATE 21. 

1. Lyropupa clathratula Anc. Olaa, Hawaii. 18768 .... 245 

2. Lyropupa microthauma Anc. Nuuanu. 11045BM . . . 238 

3. Lyropupa microthauma Anc. Nuuanu. 21562BM. .. 238 

4. Lyropupa truncata Cooke. Kohala Mts. 36996 247 

5. Lyropupa mirabilis Anc. Popouwela. 11046BM 249 

6. 7. Lyropupa mirabilis Anc. Popouwela. 108910 .... 249 
8, 9. Lyropupa antiqua C. & P. Manoa. 119472 250 

10. Lyropupa spaldingi C. & P. Puu Kaua. 119470 248 

11. Lyropupa antiqua. Type ; Manoa. 11047BM 250 

12,13. Lyropupa spaldingi C. & P. Puukaua. 11048BM. 248 

PLATE 22. 

1-3. 9. Lyropupa kahoolavensis P. & C. Type, fig. 1, and 

paratypes. Kahoolawe. 108871 256 

4, 8. Lyropupa kahoolavensis P. & C. Kona crater, Ha- 

waii. 119466 256 

5. Lyropupa perlonga interrupta P. &C. Kahuku. 119446. 261 

6. Lyropupa sparna C. & P. Kalihi. 33627BM 252 

7, 10, 11. Lyropupa sparna C. & P. Pipe-line, upper 

Kaunakakai. 108919 252 

12. Lyropupa hawaiiensis Anc. Palihoukapapa. 119468. 251 

13. Lyropupa sparna sdnulifera P. & C. Western ravine 

of Kamalo, Molokai. 119429 253 

14. Lyropupa hawaiiensis Anc. Palihoukapapa. 119468. 251 

PLATE 23. 

1,2. Lyropupa perlonga Pse. Type (a worn specimen). 

Nuuanu. 48063 Mus. Comp. Zool 258 

3-5. Lyropupa perlonga Pse. Typical. Bench of consoli- 
dated coral sand east of Diamond Head. 11052 
B. M., f. 3, and 119473, f . 4, 5 258 

6. Lyropupa perlonga Pse. Recent. Koko Head. 11053 

BM 258 

7. Lyropupa micra C. & P. Rocky Hill. 11056BM 263 

8. Lyropupa cylindrata C. & P. Makua. 11054BM 261 

9. Lyropupa cyrta C. & P. Mana. 11060BM 268 

10. Lyropupa cyrta C. & P. Mana. 119441 268 

11. Lyropupa ovatula C. & P. Manoa. 11059BM 265 



EXPLANATION OP PLATES, VOL. XXV. 389 

FIGURE PAGE 

12. Lyropupa filocostata C. & P. Limahuli, Kauai. 

11057BM 262 

13, 14. Lyropupa perlonga Pse. Embryo of Koko Head 

specimen 258 

PLATE 24. 

1-3. Lyropupa ovatula C. & P. Type. Kaelepulu. 

119434 265 

4. Lyropupa ovatula C. & P. Palatal wall. Manoa. 

119440 265 

5. Lyropupa ovatula C. & P. Kaelepulu. 119434 265 

6. 7. Lyropupa perlonga Pse. Coral bench east of Dia- 

mond Head. 119473 261 

8, 11, 12. Lyropupa plagioptyx P. & C. Kawaihapai. 

109942 267 

9, 10. Lyropupa perlonga Pse. Kaelepulu, Kailua. 119425. 261 
13, 14. Lyropupa thaumasia C. & P. Hanakupiai. 11061 

BM 270 

15. Lyropupa thaumasia C. & P. Hanakupiai. 119449.. 270 

PLATE 25. 

1-4. Lyropupa perlonga interrupta P. & C. West of Ka- 

huku. 119426 261 

5-7. Lyropupa micra C. & P. Type and paratype. Kaele- 
pulu, Kailua. 48626 263 

8, 9. Lyropupa micra maunalose P. & C. Summit of 

Mauna Loa, Molokai. 119447 264 

10. Lyropupa perlonga interrupta P. & C. Malaekahana, 

Oahu. 119463 261 

11, 12. Lyropupa micra percostata P. & C. Kaelepulu, 

Kailua. 48627 264 

13. Lyropupa cubana Dall. Type specimen, U. S. Nat. 

Mus 268 

14. Lyropupa thaumasia C. & P. Paratype. 119449 270 

15 Lyropupa cyrta C. & P. Mana, Hawaii. 119441 .... 268 

16, 17. Lyropupa cubana Dall. After Dall 268 

PLATE 26. 

1. Lyropupa scabra P. & C. Ukulele, E. Maui. 11049BM. 254 

2. Lyropupa. scabra P. & C. Ukulele, E. Maui. 119465. . 254 

3. 6. Lyropupa anceyana C. & P. Olaa, Hawaii. 11050 

BM . . 253 



390 EXPLANATION OP PLATES, VOL. XXV. 

FIGURE PAGE 

4, 7. Lyropupa perlonga Pse. Palatal folds. Koko Head. 

119437 260 

5. Lyropupa ovatula kona P. & C. Moomomi. 44762 . . . 266 

8. Lyropupa spaldingi, base of embryo. 11048BM 248 

9, 12. Lyropupa k. puukolekolensis P. & C. Puukolekole. 

119475 258 

10, 11, 14. Lyropupa ovatula kona P. & C. Huehue, Ha- 
waii. 44763 266 

13. Lyropupa kahoolavensis P. & C. Palatal folds. Mauna 

Loa, near shifting sands, Molokai. 44764 257 

PLATE 27. 

1-3. Nesopupa plicifera Anc. 11063, 15346, 19328BM. . . 280 

4-6. Nesopupa waiaiiaensis C. & P. 11064BM 281 

7, 8. Nesopupa dispersa C. & P. 11066BM 284 

9, 10. Nesopupa baldwini Anc. 18732BM 287 

11, 12. Nesopupa b. subcostata P. & C. 108885 288 

3, 14, 15. Nesopupa b. lanaiensis P. & C. 34521BM 289 

PLATE 28. 

1. Nesopupa litoralis C. & P. Ewa. 11065 283 

2. Nesopupa anceyana C. & P. Olaa, 11072 293 

3. Nesopupa anceyana C. & P. Kilauea-iki 293 

4. Nesopupa bishopi C. & P. 12465 296 

5. Nesopupa forbesi C. & P. 11074 297 

6, 10. Nesopupa limatula C. & P. 11067 290 

7. Nesopupa infrequens C. & P. 15489 298 

8. Nesopupa subcentralis C. & P. 11070 294 

9. Nesopupa dubitabilis C. & P. 11068 291 

10. Nesopupa limatula C. & P. 11067 290 

11, 12. Nesopupa thaanumi Anc. Olaa. 11084 304 

13. Nesopupa dubitabilis kaalaensis C. & P. 11069 292 

(All but fig. 3 from Bishop Museum specimens.) 

PLATE 29. 

1, 2. Nesopupa wesleyana Anc. 11079 299 

3. Nesopupa w. tryphera C. & P. 11082 301 

4. Nesopupa w. gouveije C. & P. 11081 301 

5. 6. Nesopupa kauaiensis Anc. 11076 318 

7. Nesopupa w. rhadina C. & P. Kahuku 303 

8. Nesopupa singularis C. & P. Kaliuwaa. 11077 320 

9. Nesopupa w. kamaloensis P. & C. Kamala. 108725. . . 303 

10. Nesopupa alloia C. & P. Kauai. 11078 321 



EXPLANATION OP PLATES, VOL. XXV. 391 

FIGURE PAGE 

11. 12. Nesopupa oahuensis C. & P. 11075 317 

13. Nesopupa w. rhadina C. & P. Type. 11083 301 

(All but figs. 5, 6, 7 and 9 from Bishop Museum 
specimens.) 

PLATE 30. 

1,2,3. Nesopupa tantilla (Gld.). Type. 5505USNM. . . 324 

4. Nesopupa tantilla (Gld.). 20665USNM 324 

5, 6. Nesopupa pleurophora (Sh.) . 22917 326 

7, 8. Nesopupa paivae (Crosse). After Boettger 328 

9, 12, 13. Nesopupa armata Pse. Type. 48315MCZ 327 

10, 11. Nesopupa cocosensis (Dall). Cocos Island 323 

12. 13. Nesopupa armata (Pse.). Type. 48315MCZ . . . . 327 

14. Nesopupa dentifera (Pse.). Cotypes. 48314MCZ . . . 329 

PLATE 31. 

1. Nesopupa proscripta (Smith). After Smith 344 

2. Nesopupa selebensis (T.-C.). After Tapperone-Canefri. 343 

3. 4. Nesopupa barrackporensis (Gude) 348 

5, 6. Nesopupa einghalensis (Gude). After Gude 347 

7. Nesopupa salemensis (Blanf.). Conch. Indica 347 

8. Nesopupa salemensis (Blanf.). J. A. S. Beng 347 

9, 10. Nesopupa filosa (Theob. & Stol.). Type and para- 
type, Indian Mus 345 

11. Nesopupa brevicostis (Bens.). After Gude 346 

12. Costigo saparuana Bttg. After Boettger 366 

13. 14. Nesopupa godeffroyi Bttg. After Boettger 330 

15, 16. Nesopupa tongana Bttg. After Boettger 331 

17. Nesopupa tongana Bttg. After Boettger 331 

18, 19, 20, 21. Nesopupa vitiana Bttg. Oneata. After 

Boettger 332 

22, 23. Nesopupa vitiana Bttg. Viti Levu. After Boettger 332 

24. Nesopupa mariei (Crosse). Journ. de Conch 334 

25. Nesopupa lifouana (Gass.). Journ. de Conch 333 

26. Nesopupa norfolkensis (Sykes). After Sykes 333 

27. 28. Nesopupa eapensis (Bttg.). After Boettger 335 

PLATE 32. 

1-3. Nesopupa moreleti (A.D.Br.). A.D.Brown. 64105. 33 
4,5. Nesopupa moellendorffi (Bttg.). After Boettger. .. 341 

6. Nesopupa moreleti (A. D. Br.) . 96488 339 

7, 8. Nesopupa quadras! Mlldff. 78395 335 

9, 10. Nesopupa quadras! Mlldff. 96014 335 



392 EXPLANATION OP PLATES, VOL. XXV. 

FIGURE PAGE. 

11. Nesopupa moluccana (Bttg.). 117139 338 

12. Nesopupa moluccana (Bttg.). After Boettger 338 

13. Nesopupa nannodes (Q. & M.). Bohol. 114970 341 

14, 15. Nesopupa malayana (Issel). After Issel 342 

16. Nesopupa malayana (Issel). Manila. 63973 342 

PLATE 33. 

I, 2, 3. Nesopupa minutalis Morel. 78394 352 

4. Nesopupa micra Pils. (exigua H. Ad.). P. Z. S 351 

5. Nesopupa comorensis Pils. (mo-nas Morel.). Journ.de 

Conch 353 

6. Nesopupa ventricosa (H. Ad.). P. Z. S 354 

7. Costigo borbonica (H. Ad.). P. Z. S 367 

8-10. Nesopupa gonioplax Pils. Type. 64093 351 

II. Nesopupa turtoni (Smith). St. Helena. 69935 .... 363 

12. Costigo desmazuresi (Cr.). Journ. de Conch 368 

13. Campolaemus perexilis (Smith), mut. contrarius. P. 

Z. S 366 

14, 15. Nesopupa turtoni (Sm.). St. Helena. 69935 363 

16, 18, 19. Campolaemus perexilis mut. ascendens 366 

17. Campolaemus perexilis (Sm.). P. Z. S 365 

PLATE 34. 

1-4. Nesopupa griqualandica (M.& P.). Pretoria. 114968. 357 

5, 6. Nesopupa rhodesiana Pils. Victoria Falls 360 

6#. Nesopupa iota Prest. After Preston 361 

7-9. Nesopupa farquhari Pils. 117282 . 358 

10. Costigo nobrei (Girard). After Girard 368 

11. 13, 14. Nesopupa bisulcata (Jick.). After Jickeli. . . 359 

12. Nesopupa corrugata (Prest.). Burnup delin 361 

15. Nesopupa corrugata (Prest.). Connolly coll 361 

DATES OF ISSUE OP VOL. XXV. 

Part 97, pp. 1-64, plates 1-5, November 5, 1918. 
Part 98, pp. 65-144, plates 6-10, February 20, 1919. 
Part 99, pp. 145-224, plates 11-18, June 30, 1919. 
Part 100, pp. 225-404, plates 19-34, April, 1920. 



INDEX 



ABIDA Leach 370 

abundans West 16 

aeeedens Anc 119, 121 

acheilaServ 202 

achilaSow 380 

adversidens Sandb 208 

aequidentata Poll 165 

affinis Arad 66 

APRIPUPA P. & C 276, 357 

alabamensis Clapp 79 

alaea Jeffr 69, 72 

albilabris Ziegl 8 

albina Tayl 378 

algesirae Kob 42 

alleodus Sandb 216 

Alloglossa Linds 1 

alloia C. & P 321 

allyniana Berry 376 

alpestris Aid 150, 197 

alternans Dh 223 

amicta Parr 63 

anceps Fagot 68 

anceyana C. & P 253, 293 

andrusiana Pils 109 

angulata Fag 34, 371 

angusta C. & P 315 

angustata West 53 

angustior Jeffr 211 

ANGUSTULA St 71, 145 

angustula 212, 378 

annaensis Beck 333 

anodon Dh 218 

antiqua C. & P 250 

antiquorum Ckll 84, 85, 86 

antivertigo Dr 163 

aperta Mill 219 



(393) 



approximans St 94, 95 

aprica Bgt 184 

apuana Iss 15 

apuna Iss., West 15 

aragonica Fagot 54, 372 

arcadica Reinh 16 

arctica Wallenb 189 

arenula White 215 

arigoi West 34 

arigonis Rm 34 

arizonensis P. & V 117 

armata Pease 327 

arthuri v. Marts 121 

ascendens Pils 366 

aseendens West 47 

atavuncula White 215 

athesina Gredl 175, 177 

aucta West 33 

aureacensis Loc 14 

ausonia Stef 176 

avena Drap 11 

avenacea Brag 10 

avenieuhrm Hartm 14 

avenoides West. 16 



badia Moq 31 

baldensis Parr 68 

baregiensis Bgt 35 

baldwiniana Cooke 241 

baldwini Anc 287 

barrackporensis Gude . . . 348 

basidens P. & V 117 

baudoni Mass. 169 

bergomensis Ch 22 

bermudensis Pils 149 

berryi Pils 89 



394 



INDEX. 



bidens Saiidb 208 

bidentata Jeffr 168, 169 

bigeminata Dh. 223 

biplicata Bgt 47, 51 

bisulcata Jick 359 

bleieheri Pal 219 

blumi Bttg 220, 274, 379 

boettgeriana Cl 49 

bollesiana Mse 101 

bonneti Cossm 216 

borbonica H. Ad 367 

borealis Morel 127 

bothriocheila Bgt 219 

bourgeaui Shutt 50 

brevicostis Bs 346 

briobia Bgt 185 

Bulimus cinereus Mort. . . 65 
Bulimus psarolenus B. . . 65 
biittneri Siem. . . 203 



calamanica Mlldff 367 

californica Ing 119, 120 

californica Rowell 139 

calpica West 41 

callicarens Bttg 176 

callista West 213 

callosa Reuss 216 

callosa St 97 

callostoma Bgt 219, 379 

campanea Bgt 219 

CAMPOL^EMUS Pils 364 

capellinii Sacco 221, 379 

carbonaria Anc 234 

cardiostoma Sandb 217 

CABYCHIOPSIS Sandb. . . . 223 

castanea St 132 

catalinaria St 142 

caziotiana Pils 10 

celata West 187 

centralis Anc 288 

centralis Fag 35 

cereana Mhlf 21 

cerealis Ziegl 12 

charpentieri Sh 179 



CHONDEINA Reich 1 

Chondrus Cuv 1, 5 

cianensis Caz 20 

cinerea Drap 58 

cinereus Mort 65 

cinghalensis Gude 347 

circumplicata Mss 22 

cisalpina Poll 165 

clathratula Anc 245 

clevei West 210 

clienta West 15 

coarctataDh 218,223 

cocchi Ben 13 

Cochlodonta Fer 1, 6 

COCOPUPA Pils 276, 322 

cocosensis Dall 323 

codia Bgt 184 

codiolena Bgt 219 

coUina West 162 

coloradensis Ckll 115 

columbiana St 108 

comes Bttg 223 

Qomorensis Pils 353 

concinna Scott 206 

concinnula Ckll 119, 372 

condita Gassies 333 

conecuhensis Clapp . . 80, 372 

confusa West 53 

consobrina Parr 30 

contorta Calc 66 

contrarius Pils 366 

convergens Bttg 216 

coreana Pils 156 

cornea Loc 166, 176 

corpulenta Mse 130 

corrugata Prest 361 

costata Pease 272 

COSTIGO Bttg 366 

costulata Sandb 223 

costulosa Pease 310 

crassata Bof 36 

cristata St 100 

crossei Mich 220 

cubanaDall 268 

cupressicola St 143 



INDEX. 



395 



curtaHeld 173 

-cyclophora Bgt 219 

cyclops St 141 

eyrenarum Zinn 217 

Neyrta C. & P 268 

cylindrata P. & C 261 

^ylindrella Al. Br 222 

<jylindrica Colb 214 

cylindrica Gray 175 

D 

daliaca West 193 

dalecarlica West 193 

dallianaSt. . . 137 

dalmatina Partsch 68 

danicaWest 213 

decora Gld 126 

dedecoraPils 273 

defrancii Brong 216 

deloplecta Ag 370 

denticus Settep 20 

dentiens Marts 64 

dentiens Moq 47 

dentifera Pease 329 

denudata Mss 156 

depressilabris Parr 68 

dertosensis Bof 36 

xiesmazuresi Crosse .... 368 

desmoulinsiana Jeffr. . . . 179 

desmoulinsi Germ 179 

Dexiogira De Betta 70 

Dexiogyra Stab 70 

dhorni Dh 223 

diaboli Pils 88 

dicaea West 187 

-diegoensis St 141 

diezi Cless 218 

dinii Stef 213 

discheilia Bgt 170 

disjuncta C. & P 317 

dispersa C. & P 284 

dissimilis West 60 

divergens Flach 217 

diversidens Sandb. . .218, 379 

dohrni Sandb. . . 223 



domicella West 54 

dubitabilis C. & P 291 

dunkeri Zelebor 326 

duplicata Kiister 14 

B 

eapensis Bttg 335 

eggeri Gredl 189 

elatior Paul 15 

elatior St 95 

elongata St 142, 199, 377 

elongatissima DesM 32 

elsheimensis Bttg 216 

Enneopupa Bttg 222 

eogea Pils 151 

eremia West 186 

erlandi West. 163 

eupora West 12 

eurieysii Paetel 380 

excelsa Issel 64 

exigua H. Ad 352 

extima West 190 

F 

Falsopupa Germ 370 

f arinesii Des M 45 

farinesi Mich 31 

farquhari Pils 358 

fasciata Caz 61 

ferruginea West 12 

filocostata C. & P 262 

filosa Theob. & Stol 345 

flexidens Rss 220, 273 

forbesi C. & P 297 

formosa Parr 68 

fossilis C. & P 237 

frumentum Boub 31 

Fusulina Sandb 224 

G 

gaali Wenz 379 

gallop rovincialis Marg. . . 10 

GASTROCOPTA 370 

gastrodes Ziegl 66 



396 



INDEX. 



gemma West 194 

genesiiGredl 204 

gigantea Moq 32 

girondica Bttg 220 

glandicula Sandb 208 

GLANDICULA Sandb 221 

glans Pils. & Hir 155 

globosa Sacco 220 

globula Cless 218 

globula West 168, 169 

globulusDh 218 

globusPils 218 

gnampta C. & P 317 

godeffroyi Bttg 330 

gonioplax Pils 351 

goniostoma Kiist 33, 371 

gorbeana Pils 56 

gothorum West 211, 213 

gouldi P. & C 235 

gouldii Binn 98 

gouveise C. & P 301 

gracilis Rm 29 

graellsiana Serv 183 

gratiosa West 37 

gravida West 182 

griqualandica M. & P. . . 357 

guadalupensis Fer 214 

guidoni Caz 61 

guiraonis Pils 51 

H 

hachijoensis Pils 153 

haeusleri St 177, 178 

hamata Held 212 

hannai Pils 114 

Haplopupa Pils 70 

hassiaca Pfr 11 

hauchecornei Klebs .... 216 

hawaiiensis Anc 251 

haydeni Anc 119, 121 

hebardi Van 103, 372 

heberti Foug. & Den. . . . 216 

heldiCl 177,199 

HELENOPUPA P. & C. 277,363 
helvetica West, . 201 



heptodonta Bisso 66 

heterostropha Lch 161 

hexodon C. B. Ad 8T 

hirasei Pils 154 

hoppii Moller 135 

hordeum Rm 24 

hornbeckii Villa 66 

hospitii Fag 37, 371 

hydrobiarum Bttg 217 

hydrophila Reinh 152 



ignota Fag 50 

ilendensis Fag. 56 

incerta Nevill 353 

inconspecta Parr 68 

indica Pfr 348 

INDOPUPA P. & C 276 

inermis West 187 

inflata Cless 219 

inflatula Pils 219 

INPRANESOPUPA C. & P. 

277, 289 

infrequens C. & P 298 

ingersolli Anc 119,120 

insculpta Pils 131 

inserta Pils 118 

INSULIPUPA P. & C. 

342, 348, 352, 361 

interf erens Dh 223 

interrupta P. & C. . .261, 315 

iota Preston 361 

irregularis Poll 165 

isabellae Caz 62 

isarica West 201 

isthmia Gray 69, 71 



japonica P. & H 155 

Juliana Issel 61 

juliensis Bgt 34 

jumillensis Guir 49 

jumillensis Rm 51 



INDEX. 



397 



kaalaensis C. & P 292 

kahoolavensis P. & C. ... 256 

kauaiensis Anc 318 

kobelti Hid 38 

kochi Bttg 218 

krauseana Reinh 136 

kiinowi Klebs 216 

kushiroensis P. & H 153 

kiisteriana West. . 181 



labiosa Parr 68 

labiosa Caz 10 

lamiodonta Bgt 379 

laevigata Kok 179 

laevigata West 61 

lallemantiana Bgt 43 

lanaiensis Cooke 241 

lanaiensis P. & C 289 

laroisiana West 183 

latasteana L. & B 170 

leoiitina Gredl 201 

lepida West 23 

lepta West 14 

leptocheilos Fag 31, 371 

leptochila Loc 31 

lessinica Ad 16 

letourneuxi Bgt 42 

levenensis Scott 206 

libanotica Trist 66 

lienardiana Or 370 

lienardi Germ 370 

lifouana Gass 333 

ligustica Sacco 221 

Hlljeborgi West . 167 

limatula C. & P 290 

limbata Part 179 

LIMBATIPUPA C. & P. 277, 306 

litoralis C. & P 283 

loemodonta Bgt 219 

longa Pils 377 

longini Fag 48 

Joroisiana Bgt 182 



lucana Brig 

lucida Jan 213 

lusitanica Rm 40 

lusitanica Tourn 4 

luzitanica Nobre 40 

lyrata Ancey 254 

lyrata Gld 233 

LYROPUPA Pils 226 

Lyropupilla P. & C 247 

M 

magdalenae Anc 234 

manotiana Bgt 55 

maresi Bgt 171 

mariei Crosse 334 

mariposa Pils 88 

marki Gul 107 

martini H. & J 114 

martini Sayn 206 

major Kiist 24, 26 

major Loc 166 

major Moq 61 

malayana Issel 342 

massotiana Bgt 52 

maunaloae P. & C 264 

maxima Bttg 216 

megacheiloides Cl 17 

megacheylos Dup 8 

megacheilos C. & J 7 

megalomastoma Malz. . . 362 
megolomastoma Malz. . . 363 

melanostoma Paul 15 

merita West 168, 169 

micra C. & P 263 

micra Pils 351 

microdon West 49 

microhelix Sandb 223 

microlena Bgt 185 

micronixia Bgt 219, 379 

microstoma Reuss 218 

millium Sowb 148 

milne-edwardsi Bgt 219 

miliiformis Bttg 217 

minima Cless 219 

minima Loc. . 14 



398 



INDEX. 



minor Bttg 216 

minor Menke 24 

minor Moq 61 

minor Pfr 58 

minor West 9 

minor West 176 

mirabilis Anc 249 

MIRAPUPA C. & P 255 

mitis Bttg 173 

mitis West 198 

modesta Say 123, 372 

modesta West 168 

MODICELLA Ads 1, 6, 44 

moellendorffi Bttg 341 

moenana Zinn 216 

moluccana Bttg 338 

monas Morel 353 

monas West 173, 174 

moquiniana Kiist 30 

moreleti Brown 339 

moreleti Bttg 343 

morsel St 81 

mortilleti v. Marts 65 

mosbachiensis Bttg 217 

moulinsiana Dup 178 

muhlfeldtii Kiist 24 

muhlfeldi West 26 

multicostulata Gutz 4 

multidentata C. & P. ... 315 

multidentata Strob 22 

murchisoniae Moore .... 380 

muscieola Cless 219 

myrmido Mich. . . 219 



nitidula Mss 195, 19S 

nobrei Girard 368 

norfolkensis Sykes 333 

novemplicata Loc 166 

numellata Gul 91 

nylanderi St 100 



oahuensis C. & P 317 

obesa Bgt 47 

obliterata Ch 52 

oblonga Parr 68 

obscura Miihlf . ....... 24, 26 

occidentals St 134 

ocsensis Hal 212 

octodentata Stud 164 

octodentata West 181 

okinoerabuensis P. & H. . 155 

oligodonta Del Pr 19 

olivetorum Loc 60 

oneatensis Bttg., Anc. . . 332 

onixiodon Bgt 219, 379 

oscariana St 144, 378 

otostoma West 209 

ovataSay 82, 372 

ovalis St 79 

ovatula C. & P 265 

ovatula Sandb 217 

oviformis Schl 220 

ovoidea West 195 

ovulum Pfr 86 

oviilum St. 79 



N 

nana Mich 211, 212 

nana P. & M 24 

nannodes Q. & M 341 

navasi Fag 48 

Nearctula St 70, 138 

neglecta Poey 87 

Nesodagys C. & P. . .276, 299 

NESOPUPA Pils 274, 351 

NESOPUPILLA P. & C. 276, 278 
newcombi Pfr. . 307 



pachygaster Jens 182 

pachygaster Sh 61 

padana Poll 165 

paivae Crosse 328 

pallida Phil 63 

pallida Jeffr 176 

palustris Jeffr 164 

Paracraticula Opp 221 

paradoxa St 99 

parcedentata Al. Br. ... 207 

parietalis Anc 128 



INDEX. 



399 



parrajena Orb., Paetel. . 68 

parvula Dh 218 

parvula St 105 

patula Mke 63 

paucidens West 12 

pazi Crosse 325 

penchinatiana Bgt 53 

percostata P. & C 264 

perexilis Sm 365 

perlonga Pse 258 

peraperta Pils 219 

perryi St 103 

personata Moq 179, 181 

pinetieola West 194 

pitcairnensis Beck 324 

plagioptyx P. & C 267 

plicifera Anc 280 

pluris P. & C 240 

ponapica Mlldff 336 

porcellata West 61 

praehistorica Nev 62 

praslinensis Nev 355 

presbytera Bttg 218 

prisca Anc 243 

priscilla Pal. . . .220, 274, 379 

producta West 211, 212 

prolongata Parr 68 

proscripta Smith 344 

protracta Sandb 217 

proxima Rni 68 

psarolena Bgt 65 

pseudantivertigo Pal. . . 219 

PSEUDELIX Bttg 222 

PTYCHAI^EA Bttg. . .273, 379 

Ptychochilus Bttg 274 

pulehella Bofill 54 

punctilium Paetel 340 

punctulum Bttg 195, 196 

punctum Bttg 195 

Pupa Drap 1, 370 

pupaeformis Poll 379 

PupellaSw 371 

pupula Al. Br 223 

pusilla Biv 166 

pusilla Des M 29, 30 



pusilla Mull. 161, 378 

puukolekolensis P. & C. . 258 

pygmaea Drap 96, 174 

pyrenaica Boub 371 

pyrenaica Par 31 

Q 

quadridens Sandb 208 

quadridens West 176 

quinquedentata Born ... 58 

quinquedentata Joos '. . . 217 

quinquedentata Stud. . . 173 

quinquelamellata Eisso. . 58 

quinqueplicata Miihlf . . . 24 

quinqueplicata P. & M. . . 58 

R 

ragia Bgt 219 

regularis West 188 

remiensis Dh 223 

reneana Serv 167 

rhabdota C. & P 239 

rhadina C. & P 301 

rhodesiana Pils 360 

rhynchostoma Bgt. . . 219, 379 

riviana Schr 18 

ronnebyensis West 192 

rowelli Newc 143, 378 

rubella Loc 176 

rufula Moq 8, 27 

rugosula St. 77 

S 

salemensis Blf 347 

saltusFag 51, 372 

sanbernardinensis Pils. . Ill 

sansaniea Bgt 219, 379 

saparuana Bttg 366 

sarena Gredl 175, 177 

saxicola Lowe 224 

scabraP. & C 254 

scalariformis Rm 68 

schista West 36 

schlosseri Cossm 220 

schultzii Phil. . 202 



400 



INDEX. 



schwageri Rss 223 

selebensis T.-C 343 

seminulum Bttg 314 

seminulum Lowe 225 

seminulum West. . . .164, 165 

septemdentata Fer 165 

septemdentata Bisso ... 11 

sexdentata Stud 173 

sexdentatus Mont 164 

sexplicata Bof 53 

sexplicata Loc 176 

sextana Gredl 174 

shuttleworthiana Ch. . . . 198 

sicula De Greg 13 

sieversi Bttg 195 

similis Brug 57 

similis Fer 175 

singularis C. & P 320 

sinistrorsa Anc 29 

sinistrorsum Stand 173 

sinuata Mss 166 

sinulifera P. & C 253 

Solatopupa Pils 57 

spaldingi C. & P 248 

sparna C. & P 252 

spelta Beck 23 

speluncae Bgt 47 

speluncarum Nev 62 

STAURODON Lowe 224 

steenbuchii Beck 136 

sterkii Pils 112 

striatula Parr 68 

striatula Pse 246 

subalpestris Bttg. . . . 195, 196 

subcarinata Bgt 47 

subeentralis C. & P 294 

subcereana West 12 

subcostata P. & C 288 

subhordeum West 13 

substriata Jeffr 172 

subtrochiformis Greg. . . 214 

superioris Pils 97 



tamagonari P. & H 158 



tantilla Gld . . 324 

tarraconensis Fag 48 

tassaroliana Sacco 220 

tatrica Haz 191 

tenuimarginata Des M. . . 31 

thaanumi Anc 304 

thaanumi C. & P 242 

thaumasia C. & P 270 

thibetica Bens 378 

tiarula Al. Br. 221 

tirolensis Gredl. .... 190, 192 

tirolica West 16 

Tomigerus 364 

tongana Bttg 331 

Torquilla Stud 1 

tosana Pils 156 

toscolana Schrod 9 

tournoueri Anc 4 

tournoueri Cotter 4 

transiens Cless 17 

transiens West 16 

tricolor Ant 8, 9 

tricolor Sowb 68 

tridens Sandb 208 

tridentata St 106 

trinotata St 140 

triodonta Bgt 219 

triphera C. & P 301 

trocnulus Sandb 4 

trolli F. & W 379 

truncata Cooke 247 

tuchoricensis Pils 218 

tumida West 162 

turgida Ziegl 68 

turgida Zglr 380 

turcica Dh 222 

turtoni Smith 363 

typica Poll 165 

TJ 

ultima Pils 128 

umbra Opp 222 

uncifera C. & P 236 

undorfensis Cless 218 

unifasciata Caz. 62 



INDEX. 



401 



unilabiata Bgt 171 

upsoni Calk 84 

utahensis St 109, 116 



variegella Ziegl 58, 60 

variegellus P. & M 60 

vasconensis Noulet 379 

vasconica Kob 38 

venetzii Ch 212 

ventilatoris Parr 26 

ventricosa Dup 29 

ventricosa H. Ad 354 

ventricosa Mse 92, 372 

ventrosa Heyn 181 

Yertigininae 68 

vertigo Gmel 161 



vertigo Mont 212 

VERTIGO Mull 69 

VEBTILLA Moq. . .71, 210, 221 

VERTILLARIA Pils 71, 144 

viridana Lindh 174 

vitianaBttg 332 

W 

waianaensis C. & P 281 

wesleyana Anc 299 



xenos Berry 377 

Z 

zonata Gass. . 333 



PUPILLID>E 



PLATE 19 




PUPILLID^E 



PLATE 20 




11 



13 



PUPILLID^E 



PLATE 21 




11 



12 



PUPILLID>E 



PLATE 22 




12 



PUPILLID/F. 



PLATE 23 




PUPILLID^E 



PLATE 24 




PUPrLLID>E 



PLATE 25 




17 



PUPILLID^E 



PLATE 26 




12 



V U f 1 L, L, i LJ OL 



fL,A TJfi 




PUPILLID^ 



PLATE 28 




IE 






PUPILLID^E 



PLATE 29 




13 



PUPILLID.E 



PLATE 30 




13 



14 



PUPILLID^ 



PLATE 31 




24 25 



PUPILLID^E 



PLATE 32 




16 



PUPILLID^ 



PLATE 33 




ie 



PUPILLID^ 



PLATE 34 




U. C. BERKELEY LIBRARIES