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SECOND SEEIES: PULMONATA.

MANUAL

OF

CONCHOLOGY

STRUCTURAL AND SYSTEMATIC

WITH ILLUSTRATIONS OF THE SPECIES

VOL. XXII
ACHATINELLID^E

BY

HENRY A. PILSBRY

ASSISTED BY

C. MONTAGUE COOKE j

GENEALOGY AND MIGRATIONS OF THE ACHATINELLID^J

BY

ALPHEUS HYATT

PHILADELPHIA :
Published by the Conchological Department

ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA

1912-1914

PUBLICATION COMMITTEE

SAMUEL GIBSON DIXON, M. D. , LL.D. , President of the Academy.
HENRY SKINNER, M. D., Sc. D., WITMER STONE, A. M., Sc.D.,.

HENRY A. PILSBRY, Sc. D., WILLIAM J. Fox,

EDWARD J. NOLAN, M.D.
Treasurer : S. RAYMOND ROBERTS.

A

CONTENTS.

PAGH

PREFACE v

INTRODUCTION—

Achatinellidos defined and compared with other families. ix

Structure and interpretation of the teeth x

Classification xiii

Color-patterns xix

Shape and structure of the shell xxiv

Conceptions of species and minor races xxv

Habits, stations, conditions of collecting, etcetera .... xxxiv

Supplementary observations on Hawaiian zoogeography. . xli

Remarks on the synonymy of Achatiriettidce 1

Historical notes on the literature of Achatinellidos .... lii

SYSTEMATIC PART—

Genus NEWCOMBIA Pfeiffer 1

Genus PARTULINA Pfeiffer 14

Section Perdicella Pease 15

Section Partulina s. str 23

Section Eburnella Pease 67

Section Baldwinia Ancey 90

Observations on Hawaiian Achatinellidse, by H. W. Henshaw. 91

Genus ACHATINELLA Swainson 117

Section Bulimella Pfeiffer 118

Section Achatinellastrum Pfeiffer 180

Section Achatinella s. str 274

APPENDIX I. Additional notes, undefined species etcetera . . 355

APPENDIX II. Genealogy and migrations of the Achatinellidse

in the Hawaiian Islands, by Alpheus Hyatt • 370

EXPLANATION OF PLATES . . , 400

INDEX 423

DATES OF ISSUE OF THE PARTS OF VOLUME XXII 428

(iii)

PREFACE.

This volume treats of the tree snails of the Hawaiian Islands
belonging to the family Achatindlidae ; those of the family Amas-
tridas (the genus Laminello) having been dealt with in Volume
XXI, and arboreal forms of Tornatettinida (Auriculella) remain-
ing to be considered in Volume XXIII of this series.

The Achatinellidse hold an important place in general biological
literature for the reason that a contribution of the first importance
to the doctrine of evolution arose from a study of these snails.
Gulick's theory of "divergent evolution through cumulative seg-
regation ' is now generally recognized as setting forth one of
the conditions invariable for the "origin of species,'1 having no
necessary connection with natural selection, Lamarckian factors
or mutation theory, but everywhere essential to speciation. For
this reason, among others, the account of the group has been
made somewhat more elaborate than usual in purely systematic
treatises, in order that zoologists may be in a position to form
intelligent ideas of the facts of distribution and variation of the
group. It will readily be understood that having to cover the
whole field, it was impossible to enter upon a detailed study of
any single group of forms in a restricted area. The necessary
limits of this work permit only brief discussion under each
species. For an exposition of the facts bearing on evolution in
general, a far more limited field should be chosen, two or three
adjacent valleys, where the data could be made nearly complete.
Even so, a certain amount of breeding under control would be
essential to solid conclusions. As matters now stand, we infer
where we should have exact knowledge.

The name of Dr. C. Montague Cooke is placed upon the title-
page of this volume in recognition of his collaboration in eluci-
dating the involved synonymy of the genus Achatinella. To the
systematist this is perhaps the more important work of the vol-

v

VI PREFACE.

ume. Indeed, the main features of this synonymy (excepting so
much as had been recognized by earlier naturalists) were original
with my colleague, having been worked out before we went over
the ground together in Honolulu. While for the accuracy of the
synonymy the senior author assumes equal responsibility with
Dr. Cooke, it would be unfair to hold the latter responsible for
the manner of presentation, or for the treatment of questions left
in abeyance for want of time or type material in Honolulu. It
was not practicable for my collaborator to go over the work in its
final form; and for this reason many statements of fact or opinion
throughout the text are made in the first person, especially when
based upon material we had not gone over together. The determi-
nation of all forms of Achatinetta believed to be new to science
was also the joint work of both authors, as indicated in the text.
Indebtedness to Dr. Cooke for various helpful suggestions rela-
tive to Partulina and other genera is acknowledged under the
respective species. I may also be permitted to recall the com-
radeship which made the months spent in the Islands among
the happiest of my life.

Information used in compiling the maps on pages 183, 277,
and 341 was largely supplied by Dr. Cooke and Mr. Spalding.

Professor Hyatt's MS. descriptive of tree snails was not utilized
in the preparation of this volume for the reason that his treat-
ment of species could not be reconciled with the views herein
expressed. Moreover, it is believed that had Hyatt visited the
Islands and seen the actual association of forms, his ideas must
have been very materially changed. Professor Hyatt's general
results in phylogeny and zoogeography, so far as worked out at
the time of his death, are given in an appendix ( pp. 370-399),
for comparison with the views developed in course of this study.

In order to keep this volume within reasonable size, an ac-
count of the soft anatomy of Achatinelloid snails and an appendix
to volume XXI (Amastrida) will form the first part of Vol.
XXIII. That volume will also treat of Auriculclla, Tomatellina
and their allies, and contain a general inaex of all Achatinelloid
snails.

The color nomenclature follows Ridgway's Color Standards and
Nomenclature, except in the first two numbers, which were pre-
pared before the publication of that work.

PREFACE. Vll

There remains the pleasant duty of thanking many friends
who assisted in the preparation of this book. Acknowledg-
ments are due the President and Trustees of the Bernice Pauahi
Bishop Museum, who aided the author's work in the Islands by
a generous grant. The Director and scientific staff of the
Museum furthered the investigation at every point.

Messrs. Irwin Spalding of Honolulu and D. Thaanum of
Hilo, Hawaii, placed their knowledge of Hawaiian shells and
their splendid collections at the disposal of the author with such
generosity as one meets but rarely in a lifetime.

To Professor Wm. Alanson Bryan, Messrs. R. A. Cooke, J. S.
Emerson, Hon. L. A. Thurston, Mr. W. D. Wilder and others
mentioned in the text, the author is deeply indebted for speci-
mens used in this work, and for other favors which aided inves-
tigations.

The open-hearted hospitality of Hon. George C. Cooke of the
Molokai Ranch enabled us to see far more of the Molokai fauna
in a limited time than would have been possible under less
ideal conditions.

Mr. E. A. Smith assisted with various notes on types in the
British Museum, many of them figured for the MANUAL, under
his critical eye.

Mr. H. W. Henshaw prepared an account of his observations
on Hawaiian Partulinas (pp. 91-105) containing much impor-
tant material.

Mr. C. W. Johnson lent a series of Gulick's type-specimens
from the Boston Society of Natural History. Like favors were
granted by Mr. Samuel Henshaw of the Museum of Compara-
tive Zoology, where Pease's types are preserved, and by Pro-
fessor G. D. Harris of Cornell University, custodian of the
Newcomb collection.

Finally it should be said that the production of this work
would not have been possible without the encouragement and
support of the President of the Academy of Natural Sciences,
Dr. Samuel G. Dixon, and of the Publication Committee, with
Mr. S. Raymond Roberts, having the work in charge.

My warmest thanks to all of these friends and colleagues.

H. A. P.

PHILADELPHIA, April, 1914.

INTRODUCTION.

Family ACHATINELLIDJ3 Tryon.

Achatinellida Tryon, 1884 (exclusive of Tornatellinida and

Amastrida).

Orthurethra with the kidney longer than the pericardium,
oblong and narrow. Surface of the lung plain. Genitalia
complicated by the presence of a long appendix provided with
a branch from the penial retractor muscle. Spermatheca em-
bedded in the prostrate gland, its duct long and simple ; albu-
men gland minute; prostate gland enormously developed,
composed of vermiform caeca. Jaw wanting or extremely thin,
indistinctly plaited vertically. Radula broad and short.
Teeth excessively numerous, arranged in V-shaped transverse
rows ; rake-like, composed of a narrow basal plate and a broad
recurved portion bearing numerous unequal cusps.

The shell is more or less glossy, dextral or sinistral, oblong
or ovate, minutely perforate or closed, composed of five to
seven whorls, the last convex ; aperture ovate ; outer lip thick-
ened or simple, sometimes expanding a little; columella usu-
ally bearing a strong, obliquely entering lamella in the last
whorl; internal axis sinuous. No entering lamella or tooth
on the parietal wall. Animal externally as in Helicidce. Foot
moderately broad, usually shorter than the shell. Viviparous..

Distribution, the Hawaiian Islands (except Kauai, Niihau
and perhaps Kahoolawe). Living on trees and other plants.

Studies of the soft anatomy with abundant material have
demonstrated, in my opinion, that the Achatinellidcz and
Amastrida are strongly distinct families. None of the many
species of both, dissected by other naturalists or by myself,
show the slightest approach in their differential characters.

The Achatinellidce in essential structure stand nearest to
the Amastridcr, Enidce and Ferussacida, and are somewhat

(ix)

X TEETH OF ACHATINELLID^E.

more remotely related to Partulidce. The structure of the
pallial organs and kidney is very similar in the first three of
these families. The male reproductive organs also, while dif-
fering in details, are much alike, having features widely pre-
valent in the lowest land snails, also in Onchidium, etc.

Partula, by lacking an appendix on the penis, and also by
the simple prostate gland and the somewhat different form
of kidney, stands a little apart.

By the structure of the female reproductive system, the
AchatinelUdce are most nearly akin to the Amastridce, but are
more evolved by the reduction of the albumen gland to a
minute rudiment, the prostate being at the same time enorm-
ously enlarged. The difference between the two families in
this character is one of degree rather than of kind.

From all other families mentioned above, the Achatinellidcz
differ strikingly in dentition. Those families have teeth of
normal shape, with cusps resembling and homologous with
these of the Helicidce, Lymnccidce, and most other Pulmonate
snails. The teeth of Achatineltidce and fornatellinidce differ
strikingly by having basal-plates of a different shape ;
and their cusps are not recognizably homologous with those
of other land snails. Moreover, there is no division into
lateral and marginal areas of the radula with different forms
of teeth.

Morphology of Achatinellid teeth.

It is well known that arboreal snails usually have the teeth
specially modified, more or less unlike those of their nearest
relatives among ground snails. Liguus and Polymita are not-
able examples, but in many others, such as Drymaus, Papuina,
etc., a process of change is evidently in progress. In all of
these genera, the cusps, though variously changed, are still
clearly homologous with those of normal teeth. Without com-
mitting ourselves to any special hypothesis of how such
changes come about, we cannot avoid the conclusion that in
some way the nature of the food or the conditions of grazing
lead to adaptive remodelling of the teeth in arboreal snails.
So long as the modification does not affect the homologies of

TEETH OP ACHATINELLID^. XI

the cusps, it is not necessarily of much systematic importance.
The amount of change in various groups is apparently some
indication of the length of time since arboreal habits were
assumed. In Achatinellidce and Tornatellinidce the modifica-
tion of the teeth is wholly unlike that usual in arboreal
snails of other families, in which the cusps are always broad-
ened. The Achatinellid type is really a further development
of the multicuspid teeth of small ground snails, and it would
seem, not an adaptation induced by arboreal life, though it
has obviously proved efficient in that station, in the absence
of any competitors.

The multicuspid cutting edges of the teeth, and the absence
of differentiation into laterals and marginals, cause me to
view the Achatinellid radula as one from which central and
lateral teeth have been eliminated, leaving only marginals.
The marginal teeth of Amastrida and Enidce, as of many
other snails, stand in somewhat oblique transverse series. If
the median field was eliminated, the transverse rows would
be broadly V-shaped. Pachnodus in the Enidcz, as figured
by F. Wiegmann, has somewhat such a radula as that we may
suppose the ancestors of Achatinella had. The teeth of
Pachnodus are very numerous, 375 to 393 in a row, the mar-
ginal fields have the rows strongly oblique, and the marginal
teeth are multicuspid. The teeth of Achatinella differ from
the marginals of Pachnodus, or of Leptachatina, chiefly by
the long and narrow basal-plate, whereas in most marginal
teeth this plate is very short and broad.

By a similar reduction, some species of Mesomphix have
practically eliminated the lateral teeth, while others retain a
few of them. The closely related Omphalina has the usual
development of laterals. The Agnatha and Agnathomorph
snails also have lost all laterals.

I believe, therefore, that all the teeth of AchatinelUdce have
been derived from the marginals of some unspecialized group
of ground snails having multicuspid marginal teeth. This
ancestral group was evidently also ancestral to the Amastrida,
in which Leptachatina still retains multicuspid marginal
teeth; but even the most primitive existing Amastrida have

Xll FOSSIL ACHATINELLHXE.

so far specialized the dentition that it seems to me quite im-
possible that Achatinella could have been derived from any
Amastrine genus now living.

The teeth of Physidce may be a similar evolution from the
marginal teeth of some Basommatophorous stock. They have
a remarkable superficial resemblance to those of the Acha-
tinellidce. The Athoracophoridce have also evolved teeth won-
derfully similar to those of Achatinella. This is evidently
a case of convergent evolution, as the relationship must be
remote.

Notwithstanding its strong differential characters, the
family is a very compact one, with remarkably little struc-
tural variety for a group so prolific in species. Most genera
of Helices show much greater specific diversity in characters
of the soft parts than I have been able to find in the whole
family of Achatinellidcz. There has been no adaptive radia-
tion, and with the possible exception of Newcombia, there
are no aged or phylogerontic branches.

This contrasts with the Amastridce, which have been adapted
to a variety of stations, humid, semiarid and arboreal, and in
which the shell varies from cylindric or turrited to discoidal,
with wide diversity in sculpture, color and solidity.

It appears that the AchatineUidcs are a still youthful group
in the full flower of their evolution, probably derived from
some plain terrestrial form which was rejuvenated by the dis-
covery of a new station — the leaves and branches of trees.

Fossil Achatinellidcz known up to this time are all of
Holocene, or perhaps in part of Pleistocene age. They are
far less numerous than fossil Amastridcz, probably because the
deposits wherein land shells are preserved lie mainly close
to the sea, while Achatinellidc? are shells of the mountain
forests. The known fossil species follow.

Achatinella phaeozona Gul. Kailua (p. 187).

Achatinella cccsia littoralis P. & C. Kahuku (p. 266).

Partulina montagui Pils. Manoa Valley (p. 66).

Partulina divighti occidentalis P. & C. Molokai (p. 361).

Newcombia philippiana Pfr. Molokai (p. 356).

Partulina confusa Sykes. Hawaii (p. 105).

CLASSIFICATION.

Partulina montagui is a very distinct species, but not in the
least primitive in character. Two other extinct . forms are
distinguishable races of living species, while the other three
found fossil do not seem to differ from modern shells.

Further extinct species are to be expected in comparatively
high beds, such as the Manoa and Palihoukapapa forest de-
posits. Many such must exist, and they may afford a good
deal of light on such obscure questions as the part taken by
Partulina in the ancient Oahuan fauna.

CLASSIFICATION OF ACHATINELLID^E.

It has been stated above that the genera and subordinate
groups of Achatinellidee are based entirely upon characters
of the shell. The soft anatomy, so far as known, is practically
alike in all. The genera are not groups of much systematic im-
portance— not more I should say, than the "sections" estab-
lished in my monograph on Partula; but they are obviously
natural groups, which have been found useful in dealing
with long series of species.

The family divides primarily into two groups, Partulina
and Achatinella. By their coloration and sculpture, New-
combia, Perdicella and Baldwinia are obviously derivatives
of Partulina. The Partulina group is more numerous in
species and far more varied in structure than Achatinella,
which consists of three very closely related sections. Ebur-
nella is a group of uncertain affinities, but apparently linked
to Partulina by certain species of Maui and Lanai. The ap-
proximate relationships of the groups are represented in the
following diagram.

Newcombia

Baldwinia Perdicella

| Bulimella

Partulinella

Partulina s. sir.

Eburnella Achatinellastrum

(Partulina) (Achatinella} Achatinella 8. sir'

Ancestral
Achatinellid

XIV CLASSIFICATION.

Key to Genera and Sections of Achatinellida.

a. Columella straight or only weakly folded.

6. Shell slender, turrite, usually sculptured; 3% em-
bryonic whorls, having coarse spirals, the last obli-
quely striped. NEWCOMBIA, p. 1.
61. Shell oblong or ovate.

c. Shell striped or tessellated, sometimes banded,
small, usually less than 17 mm. long.

Perdicella, p. 15.

c.1 Shell zigzag striped, banded or white, usually
well over 17 mm. long, or if so small, the last
whorl is unusually large. Baldivinia, p. 90.
a1. Columella strongly folded.

6. Shell dull or polished, usually with distinct spiral
striation ; the embryonic shell biconic, often striped ;
usually two or several embryos in the uterus at one
time. PARTULINA, p. 14.

c. Shell small, tessellated or striped, columellar
fold thin. Perdicella, p. 15.

c1. Shell larger, spirally striate, columellar fold
stout. Partulina, s. str., p. 23.

c2. Shell polished, often banded, columellar fold
stout. Eburnella, p. 67.

61. Shell polished; spiral striation weak or almost want-
ing; embryonic shell short and broad, its last whorl
never striped obliquely; only one well-developed
embryo in the uterus at a time. Oahu :

ACHATINELLA, p. 117..

c. Outer lip thickened by an internal callous
ridge ; shape oblong-conic or ovate, summit
obtuse. Bulimella, p. 118.

c1. Outer lip but little or not thickened within,
not expanded; summit conic, the embryonic
whorls not conspicuously flattened.

Achatinellastrum, p. 180.

ca. Outer lip thickened within ; shape globose-
conic ; embryonic whorls almost flat.

Achatinella, s. sir., p. 274.

CLASSIFICATION. XV

Systematic List of species and subspecies of Achatinellidce.

Numbers following the species refer to pages where they
are described. In Partulina, where there are several col-
lateral phyla, no linear arrangement can be made which will
show the affinities with Newcombia and Achatinella, and at
the same time preserve a logical sequence of groups and
species within the genus.

Genus NEWCOMBIA Pfeiffer.

(Molokai) (Molokai)

N. plicata High., Pfr. 2. N. pfeifferi Newc. 13, 355.

N. p. gemma Pfr. 3. N. p. ualapuensis Pils. 12.

N. sulcata Pfr. 5. N. p. cinnamomea Pfr. 10.

N. canaliculata Bald. 6. N. p. decorata Pils. 12.

N. c. wailauensis Pils. 7. N. p. honomuniensis Pils. 12.
N. philippiana Pfr. 8, 356. (Maui)

(+ perkinsi Sykes) N. cumingi Newc. 10.

Genus PARTULINA Pfeiffer.

Section Perdicella Pease.

(Molokai) P. mauiensis Pfr. 20.

P. helena Newc. 16, 356. P- zebrina Pfr. 20.

P. fulgurans Sykes 21.
(Maui) p. carinella Bald. 7, 358.

P. ornata Newc. 18. P. thwingi P. & C. 357.

P. zebra Newc. 19. P. kuhnsi Pils. 22.

Section Baldwinia Ancey.
(Hawaii)

P. confusa Sykes 105. P. h. fuscozonata P. & C. 365..

P. horneri Bald. 107, 365. P. physa Newc. 109.

P. h. Candida P. & C. 365. P. p. errans Pils. 111.

P. h. fuscospira P. & C. 365. P. p. konana P. & C. 365.

(Maui)
P. grisea Newc. 111. P. thaanumiana Pils. 112.

XVI CLASSIFICATION.

(Oahu)
P. dubia Newc. 113.

Section Partulina Pfr., s. sir. (Molokai).

P. virgulata High. 25, 358. P. v. kaluaahacola P. & C. 359.
P. v. halawaensis Borch. 27, P. subpolita Hy. & Pils. 359.
359.

Section Partulinella Hyatt (p. 392).
(Molokai)

P. tessellata Newc. 28, 360. P. theodorei Bald. 33, 360.

P. t. meyeri Borch. 29. P. dwightii Newc. 35, 360.

P. rufa Newc. 29, 360. tP. d. occidentalis P. & C. 361.

P. proxima Pse. 32, 360. P. d. mucida Bald. 34, 361.

P. p. schauinslandi Borch. 33. P. redfieldii Newc. 38, 362.

P. p. multistrigata Pils. 34, P. r. kamaloensis P. & C. 362.
360.

(Oahu)
t P. montagui Pils. 66.

(Lanai)
P. crassa Newc. 40, 362.

(Haul)

P. kaaeana Bald. 41. P. tappaniana C. B. Ad. 54.

P ustulata Gul. 47, 362. P. t. ampulla Gul. 57.

P. marmorata Gld. 42. P. t. eburnea Gul. 57.

P. plumbea Gul. 43. P. t. carnicolor Bald. 58.

P. winniei Bald. 44. P. nivea Bald. 59, 363.

P. perdix Rve. 45, 363. P. dolei Bald. 60, 364.

P. induta Gul. 48. P. lemmoni Bald. 61.

P. radiata Gld. 49, 363. P. terebra Newc. 61, 364.

* * * P. t. attenuata Pfr. 63.
P. splendida Newc. 51, 363. P. t. longior Pils. 63.
P. s. baileyana Gul. 52. P. t. lignaria Gul. 63.
P. gouldii Newc. 52. P. fusoidea Newc. 64.

• * * P. crocea Gul. 65.

CLASSIFICATION.

XV11

Section Eburnella Pease.

(Maui)

P. mutabilis Bald. 68.
P. porcellana Newc. 69.
P. p. flemingi Bald. 71.
P. p. wailuaensis Sykes 72.

P. variabilis Newc. 83.
P. v. lactea Gul. 86, 364.

P. mighelsiana Pfr. 77.
P. m. bella Eve. 79.

P. p. fulvicans Bald. 73.
P. nattii Bald. & Hartm. 73.
P. anceyana Bald. 75.
P. germana Newc. 76.

(Lanai)

P. semicarinata Newc. 86.
P. s. hayseldeni Bald.

(Molokai)

P. m. polita Newe. 80.

Genus ACHATINELLA Swainson.

Section Bulimella Pfeiffer (Oahu).

A. bulimoides spadicea P.&C.

A. abbreviata Rve. 123.

A. viridans Migh. 125.

A. tasniolata Pfr. 130.

A. byronii Wood 133.

A. b. rugosa Newc. 135.

A. b. waimanoensis P.&C. 137.

A. b. nigricans P. & C. 138.

A. lila Pils. 139.

A. pulcherrima Sw. 140.

A. p. nympha Gul. 144.

A. decipiens Newc. 145.

A. d. planospira Pfr. 147.

A. d. kaliuwaaensis P.&C. 150.

A. d. swainsoni Pfr. 150.

* * *

A. rosea Swains. 151.
A. bulimoides Sw. 154.
A. b. mistura P. & C. 156.

157.

A. b. obliqua Gul. 158.
A. b. ovata Newc. 160.
A. b. rotunda Gul. 163.
A. b. glabra Newc. 164.
A. elegans Newc. 166.
A. e. wheatleyana P. & C. 168.

A. fuscobasis Sm. 170.
A. f . lyonsiana Bald. 172.
A. f . wilderi Pils. 173.
A. pupukanioe P. & C. 174.
A. sowerbyana Pfr. 175.
A. s. thurstoni P. & C. 177.
A. s. laiensis P. & C. 178.
A. s. dextroversa P. & C. 179
A. s. roseoplica P. & C. 180.

XV111

CLASSIFICATION.

Section Achatinellastrum Pfr.
(Main range of Oahu)

A. phseozona Gul. 184.
A. buddii Newc. 187.
A. fulgens Newc. 190.
A. f . versipellis Gul. 196.
A. f . ampla Newc. 198.
A. solitaria Newc. 204.
A. stewartii Green 204.
A. s. producta Rve. 207.
A. vulpina Fer. 212.
A. v. colorata Rve. 224.
A. v. tricolor Smith 226.
A. bellula Smith 230.
A. b. multizonata Bald. 231.
A. casta Newc. 235.

A. casta margaretae P.&C. 240.

A. juncea Gul. 241.

A. papyracea Gul. 243.

A. juddii Bald. 244.

A. livida Swains. 246.

A. 1. emersoni Nc. 247.

A. 1. recta Nc. 248.

A. 1. herbacea Gul. 251.

A. curta Newc. 252.

A. dimorpha Gul. 258.

A. cassia Gul. 263.

t A. c. littoralis P. & C. 266.

A. c. cervina Gul. 267.

A. c. cognata Gul. 267.

(Waianae range)
A. spaldingi P. & C. 271. A. 1. gulickiana P. & C. 273.

A. lehuiensis Smith 271.

A. thaanumi P. & C. 273.

Section Achatinella Swains., s. sir.
(Main range of Oahu)

A. lorata Fer. 278.

A. 1. nobilis Rve. 283.

A. 1. pulchella Pfr. 284.

A. cestus Newc. 286.

A. vittata Rve. 289.

A. v. cinerea Sykes 291.

A. v. simulans Rve. 292.

A. turgida Newc. 294.

A. t. perplexa P. & C. 296.

A. t. ovum Pfr. 297.

A. t. simulacrum P. & C. 299.

A. t. cookei Bald. 300.

A. leucorraphe Gul. 301.

A. 1. irwini P. & C. 302.

A. swiftii Newc. 306.

A. s. chromatacrne P.&C. 316.

A. s. dolium Pfr. 316.

A. apexfulva Dixon 317.

A. a. vespertina Bald. 322.

A. a. alba Sykes 324.

A. a. apicata Newc. 324.

A. a. beata P. & C. 329.

A. a. aloha P. & C. 330.

A. decora Fer. 331.

A. valida Pfr. 334.

A. v. leucophaea Gul. 336.

A. v. cinerosa Pfr. 336.

A. v. kahukuensis P. & C. 338.

DISTRIBUTION, COLOR PATTERNS.

XIX

(Waianae range)

A. mustelina Migh. 342. A. concavospira Pfr. 351.

A. m. sordida Newc. 349. A. c. turbiniformis Gul. 353.

A. m. lymaniana Bald. 350.

Incertce sedis.
A. aptycha Pfr. 54, 145, 363.

DISTRIBUTION OF GENERA, SUBGENERA AND SPECIES.

PARTULINA

ACHATINELLA

cc

T3

C

ISLANDS

«!

hH

a

c6

^
'M

M

g

cs

c^

.S

ce

c3

3

1 i

<p

>^

o

'cu

fl

.rH

3

•I-H

'o

iD

4*3

^H

&

O
*

o

•rH

rW

1— 1

C3

~Ui

£

T3

rt

*H

S

a

c8

»v

^

-M
CJ

J3

te
'ct

-4-i

W

<D

eg

ei

^0

^

o

0

O

K

PH

PH

PQ

w

pq

<^

<5

EH

"NJiihau

Kauai ... ...

Oahu . . .•

1

1

13

17

11

43

Molokai

5

1

8

1

15

I f\ i~» O -|

1

2

3

JVIaui

1

8

14

2

6

31

Kahoolawe .....

Hawaii •

3

3

Total no. of species.

6

9

24

6

9

13

17

11

95

N. B. No species is common to two islands.

COLOR PATTERNS OF ACHATINELLID^:.

The primitive color pattern of arboreal Achatinellidtz may
have consisted of dark streaks in the direction of growth-
lines, on a light ground. This pattern is common to many
species of various genera of the family, and to some species
in practically all the minor groups. It is one of the common
patterns of coloration of snails in many families to have the
pigment thus deposited periodically instead of continuously.
In many Partulinas the streaks have become distinctly denned,
and lose their primitive relation to the growth-lines, becoming
zigzag or oblique, or they may be interrupted to form a tessel-

XX COLOR PATTERNS.

lated, marbled or mottled pattern. This is a line of pattern
evolution peculiar to Partulina, at its acme in such species as
P. proximo, and the Perdicellas, decadent in many Partulinas
and Newcombias.

In Achatinella the streaks, when present, are not sharply
defined and always run in harmony with growth-lines.

The spiral bands of Achatinellidcu belong to two band-
systems, entirely separate in inheritance. The genesis of
the irregularly placed bands is readily traceable in existing
species, but the origin of those of the four-band pattern is
unknown.

Irregularly placed bands. — Many species which have spiral
bands in the adult stage, are streaked when young. The
several stages from the streaked to the banded pattern char-
acterize various races or species ; so that it appears likely
that the continuous-banded pattern was never produced by
a single mutation from the streaked, but by a series of pro-
gressive or orthogenetic mutations. The sequence seems to
be somewhat thus :

(a) longitudinally streaked (pi. 60, figs. 4a-4c).

(b) streaks cut by light spiral lines or bands, leaving
streaked zones or spotted spiral lines (pi. 60, f. 6; pi.
56, f. 56).

(c) color of the dark zones or bands intensified or diffused,
losing the streaked or spotted pattern (pi. 56, f. 5e, 5/).

Many individuals are intermediate between a and b, or b
and c. Often instead of several or many pale or white bands
appearing, the upper half of each whorl loses the streaked pat-
tern, leaving it on the portion below the periphery, producing
a bicolored pattern of cuticle, as in pi. 57, figs. 2&, 15. This
is merely a special modification of stages & or c.

Judging from its sporadic occurrence, albinism may ap-
pear as a mutation at any stage of pattern-evolution; melan-
ism is perhaps more frequent in the later stages.

All stages from streaked to banded may be seen in some
species, such as Achatinella fulgens and Partulina redfieldi.
May species remain in the primitive stage, while in others
only the terminal pattern is seen.

COLOR PATTERNS.

The zoned or many-banded patterns are a higher (more
evolved) stage; the streaked are lower in the scale; but from
the co-existence of all stages in many species, it appears that
the evolution of one from the other is a rapid process.

The transition from longitudinal streaks to spiral bands in
these shells is entirely analogous to the transition from longi-
tudinal to transverse stripes in mammals. It is a common
phenomenon in mollusks, and examples can readily be found
in most faunas, exactly parallel to the process in Ackatinella.
Cf. Drymceus, Amphidromus, Helicostyla.

Spiral bands or lines, resulting from the evolution of stripes,
as described above, are indefinite and variable in number and
position.

The four-band pattern. - - The black-brown bands, which
stain the prismatic layer of the shell and often appear as spots
on the lip, have perfectly definite positions in Achatinellas-
trum, Bulimella, and a few Partulinas, when they occur at
all, and are evidently homologous throughout these groups.
See plate 38. These bands are four in number: 1, sub-
sutural ; 2 and 3, above and below the periphery ; and 4
around the columella. All are developed in pi. 38, figs. 4a,
6, 6a, 14. Often only bands 1 and 4 are developed, figs. 1,
la, 2-4, etc. Other forms may have bands 2 and 3, or various
other combinations. The bands of the four-banded or ' ' tetra-
tseniate ' ' system may be expressed by a formula, such as is
used for the five-banded or so-called pentataeniate Helices.
Thus pi. 38, fig. 6 has the formula 1234. Fig. 7, 0000. Fig.
7c, 0230. Fig. 18«, 1(234). Fig. 19c, 1(23)0. In Acha-
tinellastrum a particularly common formula is 1000. As in the
pentatasiiiate Helices of Europe, the bands vary through the
several possible combinations in presence or absence, and in
width when present; they are sometimes split, or confluent;
but their positions are invariable.

In inheritance, bands of this system are entirely independ-
ent of the variable or indefinitely-placed bands. They may
exist in combination with any stage of the other pattern,
from plain or streaked to many-banded; and even in melan-
istic shells they may be apparent as bands of slightly different
luster, visible only in a certain light.

XX11 COLOR PATTERNS.

I have elsewhere noted that in Liguus some forms show
an extremely interesting change in the pattern, whereby the
dark and the light markings exchange places, as in a photo-
graphic negative, without other noticeable change in the shell.
An entirely similar exchange is seen in some individuals of
certain AcTiatinellidcc •. Thus on plate 19, it will be seen that
fig. 10 has the normal, tetratamiate pattern; fig. 5 is a "nega-
tive ' ' of it, the black bands occupying the places of the white
ones in the other shell. A. crassidentata, pi. 30, fig. 23, is a
similar ' ' negative ' pattern. In Partulma, P. porcellana has
the normal four-banded pattern, while P. nattii has the
"negative' pattern.

The typical group of Achatinella (Apex) never has the
four-banded pattern. This is one of the most important dif-
ferences between this group on the one hand, and Bulimella
and Achatinella strum on the other.

Parallel and convergent evolution of patterns. — Since
many species have forms with primitive streaked pattern and
others with distinct bands, it appears that bands have many
times been evolved independently. Occasionally this has
resulted in the production of strikingly similar varieties as
terminal evolution products of several species. Thus, A. vit-
tata simulans, A. turgida simulacrum and A. leucorraphe
irwini are superficially similar forms, which as their geo-
graphic distribution and the less evolved stages of some of
them show, have descended from less similar streaked stocks.

In Achatinellastrum there are some forms of A. ccusia (be-
longing to the series of A. livida), which resemble forms of
the series of A. vulpina so closely that some authors have
actually united them specifically. Yet they inhabit opposite
ends of the Main Range of Oahu, and their similarity is
merely phenotypic.

Albino or melanic forms, belonging to different species or
races, are sometimes practically identical phenotypically, when
the differentiation in shape is slight or variable.

By the study of many colonies of a species, and of many
species, in various stages of evolution, it appears that in all
the genera of Achatincllidce the evolution of color-pattern

COLOR PATTERNS. XX111

has been orthogeiietic. Partulince and Achatinella of many
groups have followed the same path, beginning with streaked,
leading to banded patterns. While the mutations of any one
species may appear to be multifarious, it is clear that progress
has been in the direction indicated. It is significant that the
early neanic and (in Partulina) the late embryonic stages of
banded forms often show the earlier streaked pattern.

Some forms of Partulina have taken another course, em-
phasizing the vertical markings; but in some of these (such
as P. Helena, P. crassa, Hawaiian Baldwinias etc.), there are
forms which are evolving or have reached the banded stage.

The many-banded stage is the acme of pattern-evolution
in Achatinellidtf. So far as we know, it is capable of no
further modification except by decadence of pattern.

We may perhaps infer that the four-band pattern has
been inherited from the ancestral stock of AchaUnella. It
is now present in many species of Achatinellastrum and
Bulimella, and in some Mauian Partulinas. Sometimes it is
represented by the "negative' or complementary pattern
noticed on p. xxii, and very frequently it is imperfect, only
part of the bands present, or all wanting. It seems thus to
be decadent or in process of disappearing from the modern
forms. There seems some ground for the hypothesis that
in the present Achatinellid fauna we have evidence of two
successive cycles of color-pattern.

(1) In the earlier cycle there were four dark color bands
in definite positions. Of this pattern we see only the final
stage, frequently in decadence, or perhaps remaining only as
bands on the embryonic whorls. (2) The second cycle is now
in the mid-stage of evolution from a streaked or axially striped
pattern to one of many bands without definite arrangement.
In the ' ' Apex ' ' group and some others, all traces of the four-
band pattern has been lost; but in some Achatinellas and
Partulinas it lingers on with a later pattern usually super-
posed upon it.

There seems to be 110 evidence that the axially striped
Partulinas and their derivative groups, Perdicella, Newcombia
and Baldwinia, ever had the four-banded pattern.

XXIV STRUCTURE OF THE SHELL.

SHAPE AND STRUCTURE OF THE SHELL.

The shape of the shell has been varied much less in Acha-
tinellidcr than in Amastridce. Except in Newcombia, where
the spiral has been conspicuously lengthened, the ovate or
ovate-conic contour has been modified only in minor features.
In sculpture the diversity is of degree. Nearly all species
are polished, showing weak growth-lines and minute spirals,
often descending obliquely, and frequently faint or absent on
the later whorls. The later whorls of the embryonic shell are
always minutely striate spirally. In Partulina the spiral
stria? are often rather well developed ; and in Newcombia they
become prominent.

The axis of the shell is somewhat sinuous, and in the last
wiiorl it bears a spiral lamella except in Newcombia, Perdi-
cella and Baldwinia, where it is more or less degenerate.
This lamella is progressively absorbed as the shell grows.
In the embryonic stage it is represented only by a sinuation
of the columella. In both Achatinellidce and Amastridcc the
forms lacking a columellar lamella are obviously not directly
related, being terminal members of several evolutionary series.
Thus Perdicella, Newcombia and Baldwinia have more direct
affinity to Partulina than to one another. That the ancestral
stock of AchatinelUda, Amastridce and Tornatellinidce had a
columellar lamella seems a fair inference.

Professor Hyatt's idea that Partulina dubia is a representa-
tive of the ancestral stock of Partulina seems to me inad-
missible, as its weak columella is far more likely to be a recent
degenerative character than a primitive heritage, and it has
no other feature more primitive than other Partulinas.

The direction of coil of the shell and the dextral or sinistral
disposition of the organs are unusually variable in Achatinel-
lidcc ; far less so in Amastridcu, where sinistrality is unusual,
and in all cases a specific or group character. Many species
of Achatinellidce are invariably sinistral. In Newcombia all
are sinistral. Other species are as invariably dextral. In
such species a reversed specimen is as rare as among Ameri-
can land snails. A few species of Partulina and many of
Achatinella are indifferently dextral and sinistral, either in

THE SPECIES CONCEPTION. XXV

the same colony or less frequently in separate colonies. The
direction of coil is hereditary as a general rule. I have never
found a sinistral embryo in the uterus of a dextral mother.
Mr. Thaanum, whose experience has been very wide, states
that he has never taken both dextral and sinistral embryos
from one mother, in the few species of Partulina which are
indifferent in coil.

Frequently in adjacent colonies of the same race, sinistral
shells may predominate in one, dextral in another. In other
species, the prevalent direction of coil differs in different
parts of its area. See Partulina virgulata, p. 358, Acha-
tinella vulpina, and many other cases noted in the systematic
part of this work. It seems likely that, as Mr. Gulick thought,
dextral and sinistral snails find some mechanical difficulty in
mating; but experimental evidence is needed.

THE SPECIES CONCEPTION IN ACHATINELLA.

The conception of " species ': applied to Achatinella is in-
volved in more than the usual difficulty. We have to do with
evolution products of several grades, possibly of more than
one kind.

Grade I. — Races having certain seemingly permanent as-
sociations of characters (though usually with other variable
characters), and co-existing with allied races without form-
ing hybrid colonies.

Grade II, — Geographic races (i. e. with a definite and con-
sistent distribution), having moderately coherent associations
of characters, but blending with other like races through
hybrid or undifferentiated colonies where the geographic
ranges meet.

Grade III. — Forms characterized by the possession of one
or more special characters (usually of color), but which occur
for the most part in hybrid colonies with other diverse forms ;
the characters seeming to be freely interchangeable, and oc-
curring in different combinations, as in Mendelian hybrids.

The races included under (I) are species in the usual sense.
It may be supposed that the specific isolation is physiologic.
Thus, Achatinella bellula and A. vulpina are related species

XXVI THE SPECIES CONCEPTION.

occupying the same area without intergrading or hybrid
forms.

Races included under (II) may be illustrated by the group
composed of Achatinella phceozona, fulgens, stewartii and
vulpina. By reference to the map on p. 183, it will be seen
that the areas of these overlap slightly, but are in the main
separate. At the points of contact there are transitional
colonies which have every appearance of being hybrid groups,
composed of individuals having various combinations of the
characters of the adjacent races. It will readily be seen that
the "species' of this grade do not have the same value as
those of grade I, and they might be more logically considered
subspecies. Such treatment of the section Achatinellastrum
would reduce the species from 17 to 10, as explained on p. 181.
In the genus Achatinella it has been thought more practical
to recognize as "species' a certain number of races which
admittedly intergrade at their limits, than to make the species
conception so broad that no definite idea is conveyed. More-
over, the term subspecies is needed for subdivisions of lower
grade. Thus the Koolau forms of A. fulgens show slight
racial divergence, which is recognized by the term A. fulgens
versipellis.

In Achatinellastrum and Bulimella, then, it is frankly ad-
mitted that the species of this work are groups of two grades,
certainly differing widely in degree, quite likely differing in
kind. Related forms which inhabit the same district without
evidence of hybridizing may safely be put down as species
of the first grade. In other forms, which may appear equally
as distinct phenotypically, we find abundant evidence of
hybridizing where their areas overlap ; and these we rank as
species of the second grade. How fundamental this distinc-
tion is in animals generally we do not know, and it has not
yet been satisfactorily worked out from the experimental side.
Practically the grade of any given form is to be decided in
each case by field observations and abundant collections. On
pages 119 and 181 the real or first grade species of Bulimella
and Achatinellastrum are indicated. In the section Acha-
tinella s. str. ("Apex"), the conditions are very complex.

THE SPECIES CONCEPTION. XXvli

The species of the mustelina-decora group, and those of the
apexfulva-leucorraphc group (see p. 275 and diagram on
p. 278) are certainly very closely related while A. lorata
stands isolated. In Partulina most forms admitted as species
seem to belong to the first grade.

Species and subspecies of all grades are usually composed
of numerous color-forms of Grade III — that is, elementary
patterns, which may rarely occur in pure colonies, but almost
always in hybrid colonies comprising two or more patterns,
in an almost endless variety of combinations.

What may be considered "fluctuating variations", that is,
variations in the potency with which a pattern may be ex-
pressed, among individuals of a pure colony, are of great
amplitude in AchatineUidce. Thus, plate 60, figs. 17 to 17c
may be presumed to be individuals having the same gametic
constitution, but differing in the degree of potency of the
factor controlling the deposition of pigment. Other examples
are pi. 56, figs. 14 to 14d ; 13 to 13& ; figs. 1 to Ic. Most of
the pure colonies show such variation in greater or less degree.
Most shells having highly developed color-patterns show
similar fluctuation. Cf. Neritina, Liguus, etc.

In a hybrid colony there is often almost perfect segregation
of the color-patterns composing it, as in pi. 39, figs. 8 to 8e.
This is usually the case where only two or three patterns are
involved. Where the number is greater, the combinations be-
come numerous. We are getting into the realm of trihybrids
and multihybrids. Although we see only the phenotypes of
the gigantic Mendelian experiment, it is our belief that with
a good knowledge of the forms of any limited region, a trust-
worthy estimate of the elementary patterns present in a colony
not too complex may be formed, though of course it would be
absurd to say what the genotypic constitution of any individ-
ual of the colony might be.

The good segregation which prevails in hybrid colonies
leads collectors to assort their shells. The strikingly diverse
patterns were not unnaturally described as distinct species
by European conchologists, who were ignorant of the facts of
association. Moreover, before the Mendelian era, it seemed
improbable that such diverse patterns could be hybrids.

XXV111 THE SPECIES CONCEPTION.

From the results visible, it can hardly be doubted that the-
various components of color-pattern act as unit characters in
inheritance.

In species covering an extended area, there is usually dif-
ferentiation correllated with distribution. Thus in A. fulgens
the plumata pattern, streaked in delicate tints, is a con-
spicuous form in Niu, the eastern limit of the species where
green forms are not found (pi. 36, figs. 4 to 6e). Also in
the next valley, Wailupe (pi. 36, figs. 3, 3a). Further west,
in Waialae, the plumata pattern has become rare, though still
present in various combinations, and green forms are com-
mon (pi. 37, figs. 1 to 6&). In Palolo, still westward, the
green and yellow patterns are ascendant, and the plumata
marking is hardl}r represented except in the varia combina-
tion. The ancient pattern of black bands in definite positions
pervades the whole species, though the factor necessary for
the appearance of these bands is often absent. Also albinos,,
doubtless genotypically various, as usual, appear sporadically
throughout the range of the species.

This topic may be further illustrated by a diagram of the
conditions found in A. vulpina. If we represent the various,
color-patterns (or "elementary species", at least in part) by
their initials, the forms of the hybrid colonies of successive
districts, from the east westward, may be represented thus

Pauoa E. Nuuanu W. Nuuanu Kahauiki Halawa Waimano Manana

avco vceo ovl ovsa\ oval or v

(a, adust a • v, vulpina- c, ca-stanea; e, ernestina; o, olivacea;
v, virens-, s, suturdtis; a, analoga; 1, longispira).

See also p. 215. The formulas could be made more complex
by introducing various other characters. Thus, dextral in-
dividuals occur in the West, but are absent eastward. More-
over, several colonies represented by the same formula almost
always differ in the relative numbers of individuals of the
component patterns ; and adjacent colonies, sometimes on a
single tree or group of trees, may differ, some lacking pat-
terns common in other colonies of the district.

Many of the "species' of Gulick are based upon colonies

THE SPECIES CONCEPTION.

in which a certain pattern or patterns predominated, though
not to the entire exclusion of others. The same patterns may
be present though uncommon in colonies a few miles removed,
where other patterns take the leading role. Thus Gulick says
of the special pattern he called Apex tuberans, "the metro-
polis of this species is Kalaikoa ; it is also found in Ahonui,
and single specimens have been brought from Wahiawa and
Helemano. ' Any mutation having a recognizable pattern
(races of the third grade) could be treated as a "species'
by this method ; and in fact there are numerous unnamed
mutations neither more or less distinct than many of those
which have been named and denned. It is often a convenience
to have names for patterns in which the association of char-
acters is moderately stable. We may speak of the ernestina
pattern, the dunkeri pattern, etc., without committing our-
selves for or against the theory that these names stand for
races of systematic significance, or are in fact anything more
than character combinations which from their abundance
would seem to have a certain degree of cohesion.

A philosophic method of dealing with intraspecific differ-
entiation is one of the greatest present needs of systematic
zoology. Systematists can 110 more deal with species and sub-
species as their ultimate units than physicists with molecules
or even atoms. It is senseless to oppose the facts which our
vaster collections and more critical methods are forcing upon
us. The terms species and subspecies, though elastic, have a cer-
tain status recognized by those accustomed to deal with these
matters, even though the categories may not be rigidly formu-
lated. To enlarge the species conception to cover every group
of individuals having a recognizable special character, as some
naturalists have recently advocated, would be to reduce the
whole subject to chaos. Some English naturalists give a
special name to every form having a special character, such
as albinism, xanthism, a special pattern, a particularity of
shape, or the like. This is one method, doubtless far better
than none, of making intelligible record of such matters. We
are not prepared to name every individual of polymorphic
species, but in reasonable limits it seems useful to have names

XXX THE SPECIES CONCEPTION.

for groups of individuals having moderately coherent com-
binations of characters, and yet not properly to be ranked
as subspecies. In this sense we have used such terms
as ' * olivacea pattern " ; ' dunkeri pattern. ' In other cases
we have avoided naming the forms, using numbers in-
stead. The ideal (if impracticable) method would be to have
a name or symbol for each unit character, and express the
several combinations like a chemical formula; but although
almost all the world is breeding animals or plants at present,
it is not likely that we will ever have enough knowledge of
unit characters to apply this method generally in nomen-
clature !

Color-pattern is usually independent of direction of coil of
the shell, but in some cases, noted under various species, these
characters are linked in inheritance. See under Partulina
virgulata, p. 358, etc.

The facts indicate that mutations may appear anywhere
in the area of a species, and spread from the initial point as
far as time and conditions permit, mingling with the pre-
existing forms of the species. The population is thus becom-
ing constantly more complex. It occasionally happens that
one comes upon what appears to be a mutation in an early
stage of its career. Thus, near the foot of a small ridge in
Popouwela, there is a large and prolific colony of Acha-
tinella mustelina (pi. 63, figs. 10 to lOc). In the midst of
this colony, in an area of two or three rods square, we found
20 or 30 pure white individuals, most of them immature. Not
one was found in other parts of the colony, which three of
us looked over pretty thoroughly, or in any other colony in
the same district. The inference seems fair that the albino
form had very recently arisen in that place.

Many cases of special color-forms known from single or
very few trees are recorded in the descriptive part of this
work. Some of these are doubtless remains of races once
more widely spread, especially when they occur in decadent
forest; but others we can hardly doubt, are new mutations.
The collections made to-day very often show color-forms dif-
fering from those of the same valleys made sixty years ago;

THE SPECIES CONCEPTION. XXXI

but the comparison is vitiated by the uncertainty whether the
same colonies were found. The older collections were not
closely enough localized to afford evidence of progressive
change. In many cases it is certain that they were from
districts now deforested. The accurate record of colonies now
kept by the best collectors will permit of such comparisons in
the years to come, and if the snails are not exterminated, in-
valuable evidence for evolution will be available for Hawaiian
students of fifty years hence, or even less. Answers may be
found to such questions as the appearance of mutations, their
influence on the colony and their fate, and the permanency or
changes of hybrid and pure colonies under natural conditions,
etcetera. In this connection, suggestive observations were
made by Mr. Thaanum on a colony of Partulina proximo,, re-
visited after an interval of twelve years. In the interval, an
apparently new mutation — shells with pale greenish ground —
had dispossessed the normal white-ground shells which alone
were found twelve years before (see p. 360).

While there are good reasons for the belief that color
mutations of considerable amplitude are common in Acha-
tinella, it must be remembered that there is no actual evidence
bearing on the production of species of the first grade, in other
words, real species, by mutation.

Observations on the snails of many regions has led me to
believe that the well-defined species are often, perhaps usually,
rather complex populations of individuals differing in many
minor but heritable characters, such as size, minutiae of shape
and sculpture, number of teeth on the radula etcetera. In
Achatinella, as in Liguus, Polymita, Amphidromus etc., the
versatile character is the very conspicuous one of color, and
therefore has attracted far more notice than equally im-
portant but less noticeable characters in plainer snails.

One condition favoring intense speciation in the Hawaiian
Islands is that the snails breed and grow all the year round.
Hence, in a given time, their evolution should be at least
twice as fast as that of snails in temperate latitudes, in which
the organism is quiescent more than half of the time. This of
course applies to humid regions in the tropics in general.

XXX11 THE SPECIES CONCEPTION.

To the naturalist knowing these shells in the field, it would
seem absurd to claim that the colors are protective in any
ordinary sense, or that their evolution is controlled or directed
by any sort of selection. It is perfectly obvious that one
color or pattern is as good as another. A pattern predomin-
ating in one colony may be rare in the next, living under con-
ditions apparently the same, and perhaps not half a mile dis-
tant. White shells, the most conspicuous of all to my eye,
are as abundant as any others; though I would not give the
impression that any Achatinella is very conspicuous. Mr.
Perkins found no evidence that Achatinellidcz are ever eaten
by birds, or in fact by any animal but the imported rats
(Fauna Hawaiiensis I, vi, p. ccxxvi).

Weissmann's treatment of the subject of specific differen-
tiation of Hawaiian snails does not appear to be based upon a
sufficient knowledge of the facts. He considers the pheno-
mena due to adaptive variation of species in a new environ-
ment.1

I have elsewhere expressed the opinion that the opaque
coloring everywhere characterizing land shells living in ex-
posed situations is an adaptation to exclude light. The
corneous yellow, or light brown tints usual in ground
snails are more translucent. The particular patterns evolved
since arboreal life was assumed do not seem to affect the
existence of the species. In common with Dr. Cooke and
other observers, I have been unable to trace any constant
relation between coloration and physical conditions (except
as noted below), and there is certainly no connection between
color and the species of tree inhabited.

In the typical group of Achatinella (Apex], a certain cor-
relation seems to exist between the pattern and elevation,
banded forms occupying higher situations than streaked forms
of the same species, at least in many cases. I believe that
the relation is an indirect one. Banded patterns have been
evolved from streaked, as stated on p. xx; evolution in more
humid stations would naturally be more rapid than in lower,

irPhe Evolution Theory, translation by J. A. Thomson and Margaret K.
Thomson, London, 1904, II, 293-296.

SPECIFIC AREAS. XXxiii

less abundantly watered places, since resting periods would
be shorter, and the succession of generations would be faster.
I believe, therefore, that such forms as A. turgida simulacrum,
A. vittata simulans, A. leucorraphe irwini, are accelerated
races of species which at lower levels have lagged behind in
pattern-evolution.

While many conehologists may consider the treatment of
Achatinella in this work an extreme example of "lumping",
(since we recognize but 43 species in place of 171 described),
it really belongs to the splitting school. Both authors hold
that a considerable further reduction would have to be made
to make the species of equal value with most Oahuan species
of Leptachatina or Amastra ; that is, to make them all species
of the first grade.

The immense color-differentiation of Achatinella has not
been exaggerated by former naturalists, but its systematic
value has been overrated. While it is believed that the ' ' sub-
species" of this work are in large part comparable to groups
of equal rank elsewhere, the authors are prepared to find that
some of them may turn out to be undeserving of a trinomial
title. In other words, they may turn out to be forms of the
third grade, like many of the species of Gulick.

Extent of specific areas.

An idea of the size of the specific areas of Achatinella may
be gained from the maps on pp. 183 and 277 and the table on
p. 276. The colonies of one species may be spread over from
a dozen to fifty square miles, but actually occupy only a small
per cent of the territory. Some color-races are known from
very small areas — even down to one or two trees, as recorded
in the text. In general, the specific areas in Achatinella are
smaller than in Amastra, though there are exceptions. Some
Amastras of the high peaks of the main range are as yet known
from very limited areas. The species or races of the extremely
humid region on top are generally of very limited distribu-
tion, but of course the summit of the main ridge is only
known in spots, as there is no ridge trail.

XXXIV HABITS AND STATIONS.

HABITS AND STATIONS OP ACHATINELLID.E.

Achatinellas are nocturnal as a rule. By day they sit
dormant, usually attached by dried mucus, except in quite
wet places. They are not firmly attached, like Oxystyla in
dry weather, but securely enough to resist any but very violent
shaking. Forms which frequent the trunk and limbs of trees
often gather in crotches, in knotholes, or hide under loose
edges of bark. Others stick on the leaves, particularly in
the concave of a curled leaf. Often two or three roost in
company; and when one is found alone, there is usually a
companion not far away. Sometimes many gather in a com-
modious knothole.

Pubescent plants are avoided. Otherwise there seems to be
little selection, any species living on a considerable number
of plants. Dr. Cooke whose observations are recorded under
A. bellula multizonata, was unable to trace any connection
between color of snail and species of tree, and other observers
have reached the same conclusion. Various introduced plants,
particularly guava and lantana, are frequented by the snails.

In the axils of ieie one finds Achatinella in company with
Philonesia, Succinea and Auriculella. It is a favorite station
for A. abbreviata and A. lila, but is also frequented by many
other species. This plant reminds one of the tropical Ameri-
can bromeliads, as in the humid zone water collects in the axils
of the terminal bunches, deluging one as they are bent down.

The native bananas are good shell plants, both for Acha-
tinella and Laminella. They grow in small clumps in humid
ravines.

The kukui or candle-nut tree (Aleurites moluccana), so
often mentioned by Mr. Gulick, occupies a broad zone on the
lower mountain slopes. It is conspicuous from a distance by
the very light green foliage. The tree is commonly rather
rounded, with widely spreading, open limbs, branching much
like our Floridan gumbolimbo.

While Achatinella ascends thirty feet or more, most collect-
ing is from quite small trees and shrubs. The snails are hard
to see on such copiously leaved shrubs as mokihana and the

HABITS AND STATIONS. XXXV

like, and it takes some experience to find them. The eye
must be trained. I have had good collectors follow me ami
say that the best collecting was on the trees I had looked over !

I have never seen an Achatinella crawling on the ground,
though occasionally on returning over a trail, one finds shells
which had been dislodged some time before. Mr. Spalding
tells me that his experience is the same. See also pp. 99-102
for observations by Mr. Henshaw. These snails are certainly
more inert than our American Helices, which if dropped usu-
ally make off promptly ; yet it seems likely that by night they
wake up. It seemed to me that the young shells wandered
more widely than the adults. Certainly with us young snails
walk more in the day time and wander more than old ones.
In the head of Kamalo, Molokai, there was a forest of native
trees, now all dead and prostrate, doubtless from the destruc-
tion of the undergrowth by cattle or deer. It is now covered
with low lantana, standing in patches, groups or singly, and
bearing a copious population of Partulina redfieldi kama-
loensis. They are also on the dead trees. It does not seem
likely that they could have spread so universally from the
trees to the immigrant lantana without crawling on the
ground for short distances. Observations with a lantern
should be made in some such prolific colony. However this
may be, it is evident that migration is slow, and probably
for the greater part by way of the interlocking branches of
adjacent trees. Where the vegetation is dense, there is often
a pretty continuous highway; but the colonies of shells on
isolated trees or small groups of shrubs must often be segre-
gated from their fellows for many generations.

Mr. Perkins has mentioned a fact which I find is known to
all Island collectors, that a species may be found on one tree
or shrub, year after year, without spreading to neighboring
shrubs. This is true also of some arboreal Pupillidcc and
Endodontidce. It strikes the outsider as uncanny to be taken
to one certain tree to collect specimens of some snail which
either has never been found anywhere else, or nowhere else
in the neighborhood or district. To find that trees just as
good all around are barren of specimens is always a surprise.

XXXVI HABITS AND STATIONS.

The presence of special color-patterns on single trees or
small groups, in many instances, is an evidence of the slow-
ness of migration, as well as of the facility attending the
origin of varieties.

Food.- -The frequent presence of living snails on trees long
dead indicates that their food is cryptoganic. Moreover, the
leaves show no traces of grazing. The feces of Partulina
confusa examined by Mr. Henshaw were composed chiefly of
remains of fungi or algae (p. 103). The contents of stomachs
of Achatinella mustelina and several other species, examined
by Dr. A. Brown and myself, were recognized as fungi, often
with shreds or fibres probably of bark, which remained un-
digested in the intestine.

AchatinellidcF breed at all times of year. In any catch one
finds part of the individuals carrying embryos. The length
of life of the individual is not known.

The native Hawaiians claim that the tree shells have a
song, which they have fancifully supplied with words. Dr.
Newcomb (P. Z. S. 1853, p. 129) and Dr. N. B. Emerson
(Sacred Songs of the Hula, p. 121), and others have given
versions of this song. Mr. Perkins believes it to be the chirp-
ing of crickets.

Brief notes on the topography and conditions of collecting
may be useful to conchologists who have not visited these
islands. Those interested in the subject should consult the
Introduction of Fauna Hawaiiensis by Mr. B. C. L. Perkins.

The Achatinellge are not shells of the valleys but of the
ridges and upper ravines. The bottoms of the larger valleys
often lie below the zone of requisite humidity. This is doubt-
less much more generally true now than before the valleys
were so extensively deforested. Certainly at the present time
the ridges and their slopes are the chief natural highways of
the tree-snails, and the valleys are barriers, in varying de-
gree, to the spread of species along the ranges.

Since it is the valleys and not the ridges which figure in
topographic nomenclature, it follows that the usual locality
records by valleys are often less definite than we would de-
sire. For instance the locality "Nuuanu valley" might mean

HABITS AND STATIONS. XXXVli

either the northern or the southern slope, a matter of some
importance, since the Nuuanu-Kalihi ridge is a faunal unit
quite noticeably different from the Nuuanu-Pauoa ridge. The
locality records of Gulick are usually by valleys. Newcomb
generally named only the district, a far less definite indication.

Mr. Gulick has related to me that in making his rounds
from one mission station to another, he would engage Kanakas
to collect for him. His own collecting was mainly 011 the
lower slopes, reached from the valleys, in the kukui tree belt.
Much of this territory is now barren, by recession of the forest.

The relatively small number of land snails on the wind-
ward (Koolau or northern) side of the main range of Oahu
is not due to its ' ' rougher climate ? ' as supposed by Professor
Hyatt, but to the fact that the slopes are largely too pre-
cipitous to support forests, — often so steep that they are
practically barren. From the summit of the main ridge,
reached by a long climb up the southern slope, a gigantic
pali, ribbed by erosion, drops beneath your feet nearly to sea
level. Such vegetation as finds root-hold often cannot be
reached further than a few rods down from the summit, where
one ventures clinging to bushes for support. At the foot of
the pali there is a talus-slope, often with kukui and other
trees, but at this low level there are few tree-shells or none,
under present conditions.

Where long butresses extend out upon the Koolau side,
as in the Kailua region, and from Kahana northwestward,
these conditions are modified in many places ; and here the
forests were, or still are, rich in tree-snails, though the wood-
land limits have retreated far within their old boundaries.
Once forests with AchatinelUdcE and Endodontidce shaded the
plains far seaward from the lovely peak of Kaneohe, where
now dead shells may be picked up in plowed fields, or gathered
out of 'pockets' in the rocks. It has been the same in the
northwest. Forest-snails are found in the sand-dunes of
Kahuku, now far from where living tree-shells exist,

The poverty of the ocean side of the range is therefore
due chiefly to the small area of forested slopes and ridges,
owing to the colossal erosion of this side, and also to the prac-

XXXV111 HABITS AND STATIONS.

tically complete destruction of primaeval forest on the lower
levels.

The chief area of Achatinella is from the ridge down to
about the 1000 ft. contour on the southern or Kona slope.
The best published maps of Oahu give a wholly false idea of
the topography of the western half of the Main Range. The
fact is that the precipitation along the main mountain axis
has formed great amphitheatres along the lee side. The
ridges running westward or southwest from it are in part
broken down, lowered, and worn to knife-like crests. Several
miles further from the main axis, where erosion has been
less potent, the ridges broaden out and branch, so that to-
wards their lower ends there are more and wider high ridges
than further up. Far down, finally, they may be cultivated
on top. A map of this region, supplied by Mr. Spalding
from his note books, is given on page 277, in order that an
intelligent idea may be formed of the localities mentioned in
the text for the western species.

In the Waianae range (see p. 341) the conditions are
similar. Though it rises to a greater elevation than the Main
range, it has also been more extensively eroded. Broad val-
leys, separated by knife-like or interrupted butresses, have
been carved out on the southern (ocean) side, heading for the
greater part in precipitous slopes, mostly inaccessible. There
is a narrow fringe of forest, with tree-snails in places, around
the heads of these valleys, but the major part of the slopes
themselves are almost or quite barren, and too steep to sup-
port forest. The valleys of the inland side of the range
head up so near those of the Ocean side in some places that
only a narrow, wrall-like arete remains, as above the heads of
the Popouwela gulches. Standing on the wild-goat trail at
the summit, a stone dislodged on either side might bound
downward a thousand feet or more, so narrow is the ridge. At
Waianae Pass the range has been almost gnawed through.
Immediately west of this pass Mt. Kaala rises, the highest
point on Oahu — huge, flat-topped, — its summit drenched by
daily cloud or rain. West of Kaala the ridge narrows again.
At the northAvestern end of the range, where the foothills ap-

HABITS AND STATIONS. KXX1X

proach the ocean at and west of Kawaihapai, there is now
no forest on top, but Amastra, Leptachatina etc. may be found
on the wooded slopes. The chief forests of the range stand
on the inland slopes and butresses from the top down to per-
haps 1000 or 1500 feet. They are often prolific in Acha-
tinellas of the Mustelina group. Several species of Acha-
tinellastrum have also been found, but they are extremely
local, and among the rarest Oahuan snails. Amastra and
small shells are abundant.

The western half of Molokai is grass land, or barren where
the country is broken, and there are no living land shells.
The eastern half has the general structure of the main range
of Oahu. Western Maui is similar, but here the gulches radi-
ate from a central point. The Kohala region of Hawaii,
like West Maui, is deeply cut by magnificent gorges, as seen
from the sea. I regard it as one of the most important re-
gions to be explored.

Lanai, which I did not visit, has a very limited amount of
forest remaining, and this has been thoroughly worked by
Mr. Thaanum. Important work is still to be done in the
deposits of fossil shells.

While considerable climbing is involved in any serious col-
lecting in the Hawaiian group, the altitudes are not great.
In the Rocky Mountains our basal camps are usually above
the tops of the Oahuan mountains, and the shell country
goes up to over double the elevation. Yet the open, rocky
slopes of our western mountains render them vastly more
accessible than the Hawaiian mountains, where one does not
think of making any considerable ascent unless there is a trail.
I have gone through scrub oak thickets in Arizona as strenuous
as any fern or ieie patches in Oahu, but they are exceptional,
and can usually be avoided. In general, the slopes are more
precipitous in the Islands, many more of them are practically
inaccessible, and owing to the humid atmosphere and dense
jungle, far less ground can be covered. Cross country tramps
or short cuts are hardly thought of unless in a thoroughly
known district.

To the collector who has worked in the West Indies, the

xl HABITS AND STATIONS.

larger shells seem scarce and hard to find. Only small or
minute shells are really abundant, and these only in spots.
I would say that in Cuba, the same amount of work would
result in at least ten times as many large shells as in Oahu,
while the number of small ones would be about the same. In
southern Arizona, one could not expect more "live" shells in
a day's work than in Oahu or Molokai, if so many; and the
number of species would be smaller. Helicina, Succinea,
Philonesia, Tornatellinida: and Leptackatina are usually
rather abundant. Pupillidce and Endodonta often abundant,
but very much more local.

It is easy to understand the genesis of arboreal snails
when one sees the conditions in such humid forests as those
of Hawaii, where the rainfall totals one to two hundred
inches. Snails of all kind — HeUcinidce, Zonitidcc, Endo-
donts, Amastras, Pupillidcc — wander freely over the herb-
age and tree-trunks. There is no need for them to stay close
to the damp earth and under cover, as in dryer climates.

Achatinellas are usually cleaned by "squirting' without
cooking. A small nozzle attached to a faucet gives a good
stream for the purpose, or a syringe may be used. Hot water
changes the green color to a dirty olive or brown. The chemi-
cal relations of green and brown must be very intimate.
White shells or those with dull colors, like Partulina, Amas-
tra and many others, may be boiled without detriment to
the colors. As a rule, AchatinelUdce, Amastridcc and Auri-
culclla pull easily. Living Achatinellas are daintily clean and
bright; and of course no oil or anything else should be put
on the shells to improve or preserve the colors.

The numerical preponderance of arboreal Achatinellidce
in the Island faunas has been much exaggerated. The actual
number of species known is less than one hundred. The
genera Amastra and Leptachatina both outnumber the tree
shells. Leptachatina will eventually prove to be far the most
prolific genus in the fauna, in the opinion of Dr. Cooke, who
has given it special attention. Kauai and Maui will doubt-
less turn out many new species of Amastridcc, and in Maui,
Achatinellida also. Hawaii - - especially the Kona side — is

NOTES ON ZOOGEOGRAPHY. xli

yet very imperfectly known. The spectacular gorges of the
Kohala mountains appeal to me as a promising fielcl. More-
over, it is an important point from the zoogeographic stand-
point. On all the islands, the fossil beds promise good enter-
tainment.

The snail fauna, outside of Achatinelloid forms, has been
only imperfectly worked up, and many undescribed species
are now known. We cannot doubt that a large number re-
mains to be discovered. On the whole, it appears that the
work to be done in faunistics alone is much more than one
generation can accomplish.

Much in the preceding pages may seem trite to Hawaiian
naturalists, accustomed from boyhood to scale the rugged,
jungle-clad mountains of their exquisite islands in the quest
of shells, or wild goats. I wxmld not presume to write for
them upon many topics which have been touched on here, in
order that conchologists elsewhere may better understand the
Hawaiian faunas.

NOTES ON THE ZOOGEOGRAPHY OF ACHATINELLOID SNAILS.

(Supplemental to Vol. XXI, pp. xi-xix.)

Arguing from the distribution of the living snail fauna,
the theory was advocated in Vol. XXI that the islands were
formerly united, allowing land and fresh-water snails and
other land animals and plants to pass from one to another.
The exploitation of the deposits of fossil land shells lends ef-
fective support to this theory.

In the recent fauna, the species of Achatinellidge and
Amastridae are special to each island, though in a few cases
the difference between some species of two islands is small.
In the Pleistocene fauna the relationship was closer. Oahu
had a Partulina of Molokaian type (A. montagui). The
Amastra umbilicata group is found to be common to every-
one of the islands. A. umbilicata of Oahu and Molokai is
barely distinguishable from A. morticina of Maui and
Kahoolawe, and A. ultima of Hawaii is very closely related.

In Leptachatina, the group of forms having a posterior
tubercle and groove in the aperture, formerly known only by

xlii NOTES ON ZOOGEOGRAPHY.

one specimen from Kauai (L. fossilis, Vol. XXI, p. 61), is no;v
known from many beds on the north, west and south shores
of Oahu, from Molokai, Maui, Kahoolawe and Hawaii. This
group of very closely related forms, now extinct, was commo'i
to at least six of the islands in the Pleistocene. One species,
L. subcylindracea, was common to Oahu, Molokai and
Kahoolawe, and the forms of Maui and Hawaii are barely
distinguishable. The extinct forms of the L. oryza group
from Oahu to Hawaii are so very similar that we have doubts
of their specific distinction.

Upon the hypothesis that the snails had been distributed
by ocean drift, birds, etc., it would be expected that the num-
ber of species common to two or more islands would be in-
creasing by the occasional action of such agencies. The fact
is, that species common to two islands were more numerous
in the Pleistocene than they are today, even though the ex-
ploitation of the fossil deposits has only begun, and the recent
fauna has been studied for years. Such a condition is
exactly what we would expect if the islands had been united
in the late Tertiary, and their common faunal elements had
been distributed by known and normal modes.

The hypothesis that Achatinellidce (or rather Amastridce)
first come to Kauai, and thence to the other islands, as advo-
cated by Professor Hyatt (p. 371 et seq.) no doubt had its
inception in the orthodox view of the relative geologic age of
the islands. The biological evidence relied upon by Hyatt
has proved, now that the fauna is more fully known, to
indicate no such succession of faunas. Taking the fossil
species now known into the account, it becomes obvious that
all the islands have, or in the Pleistocene had, very closely
related faunas of the primitive groups: Leptachatina (cer-
tainly the most primitive genus of the family), Amastrella,
Cyclamastra. Between these closely related species it would
be absurd to claim that those of Kauai are more primitive
than those of Hawaii, or any other island. The same evolu-
tionary stage is equally represented by species on all the
islands. Hence, it seems likely that these common faunal
elements remain from a former time when the islands were

• NOTES ON ZOOGEOGRAPHY. xliU

united. In addition to these primitive Amastridce, almost
every non-Achatinellid genus of land and fresh water shell
found in the archipelago occurs on all the islands, excepting
the low and arid Niihau and Kahoolawe, where part are want-
ing. It appears highly improbable that so many genera
should have become so generally diffused by any ' ' accidental '
means of over-sea transportation. That Kauai has ever con-
tributed faunally to the other islands by means of drifting
trees or the like is the more improbable because such drift
would be against what current there is, and directly across
the course of the trades. The snail fauna of Kauai is
"primitive" only in lacking representatives of the Achatinel-
lid<z, and the arboreal genus Auriculella. The fauna of
Amastridcc and other families is on a par with the faunas of
the other islands. The absence of Achatinellidcc from Kauai
seems to be a rather strong argument, albeit negative, against
the view that the fauna of the entire group emanated from
Kauai.

Mr. Perkins, in discussing the dispersal of Achatinellidcc
states that ' * once, on Molokai, a young living Achatinella was
found attached to the feathers of the Drepanid bird Chloro-
drepanis. Frequently they become adherent to one's clothes
when passing through the brush. Doubtless in high winds
very young shells are sometimes carried to a distance in curled
up leaves in which they hide' (Fauna Hawaiiensis, Intro-
duction, p. Ixvi). While such means of travel must have
had some part in the distribution of Achatinella, we are in-
clined to believe it a small part, for the reason that
anomalies in distribution are very rare. If birds had carried
the snails about, there would be colonies of "erratics,' like
granite boulders in a glaciated limestone region, instead of
the thoroughly consistent distribution which is the rule on
every island.

It is now known that the family Achatinellidce differs in
important structural characters from the Amastridce. While
the two are allied, their separation must, if we judge by the
analogy of other groups of known age, date back to the early
Tertiary at least. Indeed I believe that no existing genus of

Xliv NOTES ON ZOOGEOGRAPHY.

Amastridce could have been ancestral to the AchatinelUdte.

The points of agreement between Kauaia and Achatinella
ph&ozona, noted by Hyatt (pp. 398-9), pertain to shell char-
acters of little or no phylogenetic significance, and leave the
important anatomical differences absolutely untouched. In
fact I have failed to find any characters in A. phccozona which
indicate it as an ancestor even of the species immediately
related — far less of the whole Oahuan genus Achatinella !

The absence of arboreal AchaUnettidcs from Kauai has
provoked comment because the forests seem well adapted to
tree shells. It is a grave difficulty from any point of view.
I am informed that Achatinella introduced there thrives.
Several hypotheses may be formulated. (1) It is possible
that the AchatinelUda reached the region later than Amas-
tridcc, and after Kauai had been isolated from the remainder.
This seems the simplest explanation. (2) Achatinellidcs
may have existed on Kauai, but become extinct from some
unknown cause, as the horse family in America became ex-
tinct. Or (3), it is possible that the events attending the
submergence of the old Hawaiian land did not allow tree
snails an opportunity to gain access to the new forests on the
Kauaian volcanos. However this may be, speculation upon
a negative character of this kind seems rather unprofitable
emplojonent in the present condition of our knowledge.

In my opinion there is no evidence whatever indicating
southeastern Oahu as the point of divergence of the snails
of that island. The geological features and the distribution
of species are both too ' ' mature ' to indicate one place more
than another in the Main Range as the initial point. In
Hawaii both the geological structure and the distribution of
Achatinellidce point to the Kohala mountains as the area from
which land snails have radiated. Species and varieties pro-
gressively diminish from this region down the east and west
coasts.

From studies on Amastridcc (Vol. XXI) I concluded that
the Waianae fauna had been independent of that of Koolau
from the early period when all the islands were connected
until late Pliocene or more likely Pleistocene. The tree shells

NOTES ON ZOOGEOGRAPHY.

fully support this conclusion. There are five species of
Achatinella, one Partuliiia and two Laminellas. Three of
these are referable to Koolau species, and the others are so
clearly allied to several diverse species, belonging to groups
widely spread in the Koolau range, that they must be deriva-
tives from them. There is no endemic Waianaean strain of
Achatinellida', and from all indications it seems that this
family was lacking there until Pleistocene time. The old
Kaala massif, like Kauai, had a fauna of Amastrida. In the
diagram no. 2, in Vol. XXI, p. xx, Kaala might better have
been isolated from the Oahu-Hawaii island. These conclu-
sions have support from the distribution of plants. Mr. Charles
N. Forbes states of Kaala and its range that "its flora is as
distinct from the main range of Oahu as is the flora of any
separate island of the group' (Oec. Pap. B. P. B. Mus. V,
no. 4, p. 13, 1913).

In the case of Maui, the two mountain masses are now
separated by a considerable width of low land, impassible
for land snails. It seems certain that the connection was
formerly more intimate. The number of identical or very
closely related species is quite considerable, and judging from
Mr. Thaanum's last expedition, it is likely to increase. The
isthmus must have been densely wooded, and probably it stood
at a decidedly greater elevation, to permit free exchange of
mountain snails. A rather recent subsidence of the whole
island is indicated by the absence of such an extensive pene-
plain as would result from the tremendous erosion of West
Maui, if that had been accomplished at the present level of
the island.

The Kauaian satellite island Niihau will doubtless prove to
have had all of the widely spread genera of Kauai — such as
Leptachatina, Succinea, Helicina, Tornatellina, Pupillidce,
Endodontidce etc. — when its Pleistocene deposits are ex-
amined. A general account of the island and list of the plants
has been given by Mr. C. N. Forbes from notes made by Mr.
J. F. G. Stokes (Occ. Pap. B. P. B. Mus. V, no. 3 with map).

Theories relating to the origin of the Hawaiian fauna have
been unduly influenced by the inferences of geologists con-

Xlvi NOTES ON ZOOGEOGRAPHY.

cerning the growth and history of the islands, especially as
to their relative age, and the idea that they were built up
from the sea bottom solely from materials ejected by the
volcanoes. This hypothesis presumes an enormous output
and a duration very long for the active life of a volcano.
Nowhere on land, where the geologic structure is accessible,
has any mass even remotely comparable in magnitude to the
Hawaiian ridge been piled up by purely volcanic agency.
While it is true that most volcanoes rise from low levels, it
is also noteworthy that those whose summits stand high
(18,000 to 30,000 feet above the general level of the
supporting sea bottom, in the case of the Hawaiian
ridge) are borne on the backs of great folded mountain
uplifts, as the Andes, or the Mexican plateau. Since
none of the known volcanic deposits are believed to be older
than tertiary the hypothesis also limits the time available for
the evolution of the peculiar Hawaiian fauna to a period far
shorter than experience has shown is probable. Elsewhere
we know that many groups of generic rank in the land shells
go back to the Oligocene and some to the Eocene. The dif-
ferentiation of the modern families must therefore have taken
place largely in the Mesozoic. It is not likely that so strongly
characterized a family as the Achatinellidce is much later.
The case would be different if Achatinellidce were known to
have existed on any of the continents, for then it might be
claimed that they had drifted to the islands. Personally, I
think the drift hypothesis, applied to these remote islands, is
about as credible as the idea that life was first brought to
earth on a meteorite. I can conceive that wood-boring beetles
might travel in the logs from America which I have seen
thrown upon the windward coast of Molokai — logs stripped
of bark and worn deep into the solid wood by the buffeting
of the waves — but my faith is too weak to believe that any
snail could get deep enough in the wood to make the journey
of two thousand miles dry, or that if it did, it would ever
get out when the log finally dried on a tropical beach. Only
arm-chair zoogeographers can hold to the hypothesis that the
land snails, Lymn&idce and Melaniidce of these islands were

NOTES ON ZOOGEOGRAPHY.

brought there by drifting logs, or sticking to the feet or
feathers of birds.

We have to account for the presence of representatives of
eight families of land and three of fresh-water snails.1 There
must have been at least sixteen, probably more, original
immigrants. The affinities of practically all are with Poly-
nesia, and much more remotely with the western border of
the Pacific, in no case with America. Under present condi-
tions, drift material can come only from America. This has
certainly been the case during the duration of the present
islands, as their coast topography shows. In all probability
the trades have blown and the current flowed as they do now,
since early geologic times.

The idea that the mollusks were carried by wind, as some
insects and seeds may have been, is hardly to be entertained..
The distance is too great.

The hypothesis that the Hawaiian volcanoes rise from
a preexisting mid-Pacific ridge, now lost by subsidence,,
gives room in time and space for the development of the
peculiar fauna. The tertiary history of the islands, on this
hypothesis, has been mainly one of subsidence along with
volcanic upbuilding. This subsidence has continued down to
comparatively recent times. Such topographic forms as Pearl
Harbor, Kaneohc Bay, Kahana Bay, can hardly be explained
except as valleys of subaerial erosion. On the northern coast
of Kahoolawe it may be noticed that the ravines and ridges
extend out into the sea in a succession of little bays and points,
although the present erosive action tends to wear away the
points and silt up the bays.

There is also, as many authors have noted, conclusive evi-
dence that on Oahu there has been a small Pleistocene eleva-
tion. This certainly amounted to 20 ft., as beds containing
recent marine shells indicate. According to Professor Hitch-
cock, the bluffs at Kahuku are of marine origin up to 60 ft.,

1 The following groups, now well established, are regarded as introduced
in the human period : Limacidce, PhilomytidcB, Helicidce, Opeas, Gcecilioides,
Planorbis, Viviparus and probably Ancylus and Musculium.

xlviii NOTES ON ZOOGEOGRAPHY.

above which they are consolidated dune formation. Some of
these elevated beds have been referred to by Doctor Dall as
Pliocene. This determination was not based upon actual
determinations of fossils, but, I gather, from general con-
siderations. My collections of marine fossils around Pearl
Harbor and east of Diamond Head show only recent species,
and lend no support to the idea that the raised beds are older
than Pleistocene.

The "raised reefs' I had opportunity to examine in west-
ern Molokai are consolidated calcareous sand. They are dune
deposits, as the irregular bedding clearly shows, although
they often contain sea shells, blown up from the shore. Not
much dependence is to be placed upon Hawaiian geology seen
from a steamer's deck. Upon the whole, it appears that in
Oahu there has been a very recent elevation, following a
subsidence of far greater amplitude.

A good summary of the physiography and geology may be
found in the earlier chapters of Hawaii and its Volcanoes,
by Dr. Charles H. Hitchcock, Honolulu, 1909.

Age of the land-shell beds.

In the Islands the distinction between Pleistocene and
Holocene deposits is a real and necessary one, since human oc-
cupation brought in factors profoundly affecting the physiog-
raphy of the whole lower zone, or on some islands the entire
area. It would also be inexact to call the Holocene beds "re-
cent", though some of them are apparently of no great antiq-
uity. If the terms are used loosely* in this book it is because
not enough work has been done to fix the age of the various
dune deposits definitely. The Manoa, Kailua and Kahuku
bluff deposits, and the beds of Mana, Hawaii, seem to be un-
doubtedly Holocene. The dune-covered beds of northwestern
Oahu, those of Moomomi, Molokai, and those of the neck of
Maui (which I have not seen), are doubtless older than the
first group, but may possibly belong also to the early human
period. The human remains in the Moomomi dunes, however,
are probably intrusive. The land shell breccia and tuff of
Diamond Head and other tuff cones of the Kona side of
Oahu, are undoubtedly Pleistocene.

NOTES ON ZOOGEOGRAPHY.

As yet, only the Acliatinellidce and Amastridcz of all these
beds have been thoroughly studied, and these are the groups
which everywhere show the greatest local speciation, there-
fore less useful for comparative studies of the beds than the
widely distributed small shells of other families.

The Diamond Head breccia contains a considerable num-
ber of extinct species. Dune sand, interbedded with land-
shell breccia, where the section has been fully exposed in the
sand quarry along the road, contains only recent marine shells.
I conclude therefore that the Diamond Head and Punch-
bowl land-shell deposits are Pleistocene, not Pliocene. A cer-
tain proportion of special species would naturally be ex-
pected in a region of such intense local endemicity as Oahu.
They are not of themselves an indication of age.

Whether any fossiliferous deposits older than Pleistocene
will be found on Oahu is doubtful. There is a possibility
of inland deposits, but if the movement of the island has
been mainly downward, as I believe, there is little probability
that marine Tertiary beds will be found near the surface.

Recent Climatic change.

That there has been a change from more humid to dryer
climate in many districts of all the islands, if not known by
historic evidences would be demonstrated by the restriction
of most land snails to higher levels than formerly occupied.
Sixty years ago the Achatiuellas were found in abundance
at half the elevations now inhabited by them. Still earlier
forest shells lived within a few feet of the present sea level, as
the Kailua and Kahuku deposits show. Forest shells,
Amastra and others, are found in many of the low deposits,
and it is obvious that the Pleistocene forests extended nearly
or quite to the sea on the northern and western coasts of
both Oahu and Molokai. The changes within the last cen-
tury are held to be due to deforestation by cattle, which by
destroying the underbrush cause the dessication of the forest
humus, and prevent reproduction of the native trees. In
Laiiai the wild goats have almost, and in Kahoolawe have
totally, completed the destruction of native forests. With

1 SYNONYMY OF ACHATINELLID/E.

loss of the forest there is less precipitation on the heated soil,
and what water falls is not conserved. Whether the earlier
destruction of low-lying forest was due to human agency is
not knOAvn, but in the absence of any other known cause
that seems likely.

The change from humid to arid conditions in the lower
zone seems to have been too rapid to admit of the evolution
of many arid country species. There is one Succinea which
lives under volcanic rocks in the most arid places; a species
of Bifidaria is often found in dry country, and a few Lept-
achatinas exist in similar places. In general however, one
finds little or nothing to encourage search on the slopes of the
lower zone.

The same tale of increasing aridity is heard from the tuff
cones of the Kona coast, where deposits of fossil forest-snails
occur on Koko Head, Diamond Head, Punchbowl, and others.
These desolate cones, furrowed with gullies, chill the traveller
approaching Honolulu with their austerity. Of herbage there
is now little, save for a thin line of dusty algarobas (Prosopis)
bordering the shore and straggling up the gulleys, and tufts
of dry grass which may harbor the dry country Succinea and
Bifidaria. In Pleistocene times, between periods of activity
and after the eruptions ceased, these cones were heavily
wooded, with a copious snail fauna. Amastra, Leptachatina,
Tornatellinidce, Lyropupa and Nesopupa, Endodonta and
Nesophila, Zonitidcc, etc. found congenial environment, if we
may judge by the abundance of individuals.

NOTES ON THE SYNONYMY OF ACHATINELLID^E.

The specific synonymy of Achatinellidcc has been discussed
by Pfeiffer in the various volumes of the Monographia Heli-
ceorum, by Newcomb (1858), Dr. W. D. Hartman (1888),
D. D. Baldwin (1893) and Mr. E. R. Sykes (1900), all of
whom have given synonymic lists of the species known to
them. Messrs. Gulick and Thwing have also made sugges-
tions. While the synonymy of the present work differs
widely from previous arrangements, it has not been thought
desirable to occupy space with criticism of the views of other

SYNONYMY OF ACHATINELLID^E. H

authors, except when occasionally some felicitous grouping
by one or another student inspired approving comment.

It would be quite impossible for an investigator knowing
these shells only by small museum series, already assorted,
and often imperfectly localized, to arrive at right conclusions.
Several factors combine to render the recognition of species
and races unusually arduous in Achatinella. The absence
of differentiation in structural characters is the chief hard-
ship. In any allied series the shape is about the same, and
the size is remarkably uniform throughout the genus. There
are no such extremes of size as in Partulina. Both shape
and size vary so much individually and with locality that they
cannot often be depended upon to separate allied species,
except in the average. There is very rarely any specific
sculptural modification ; and finally, the color and pattern are
kalaidoscopic in many races, and a special pattern is often
produced orthogenetically in several species.

Any polychromatic species is likely to produce individuals
of strange or unusual patterns, sometimes possibly as new
mutations, more often as a result of some unusual combina-
tion of color- factors in complex hybrid colonies. Such unique
or exceptional individuals may readily enough be referred to
their proper species if kept with their associates in life ; but
separated from their colonies, and without definite locality,
it is sometimes almost or quite impossible to tell what racial
stock they belong to. A number of the species described by
Dr. Pfeiffer from specimens sent by Mr. Frick were evi-
dently such exceptional, or as they say, 'freak' shells.
Any large collection contains shells which, if isolated and
described without locality, could be referred to their proper
species only with, the greatest difficulty if at all.

A further source of perplexity is the tendency of some
species to form melanistic or albinistic mutations. In the
absence of structural characters, such forms, when not ac-
companied by their associates, and especially if without exact
locality, may be difficult or even impossible to refer to their
proper species or race. This is particularly true of the Apex
or typical group of Achatinella.

Hi SYNONYMY OP ACHATINELLID^E.

Dissections of a considerable number of species indicate
that the soft anatomy shows even less specific differentiation
than the shells. Further work along this line should be done ;
but my impression is that very little assistance is to be ex-
pected from the soft parts in distinguishing closely related
species of Achatinella. I did not have time to test the value
of body or mantle color as a specific character. It is not
U\ely to be more constant than shell color.

Finally, we have to do with artifacts. Newcomb, Gulick
and others have commented upon the manufacture of "new
species ' ' of Achatinella, practised by some persons. These in-
dividuals were not actuated by a desire to advance science.
They were moved by cupidity, or a perverted sense of humor.
Perhaps all of the old collections contain some of these altered
or ' ' improved ' ' shells. The commonest alteration is a change
of hue by the use of hot water. Other patterns were altered
by scraping the very thin colored cuticle, sometimes adding
bands with colored ink, as in pi. 30, figs. 48, 48a, 51, 52. An
ingenious method practised years ago, which Dr. Cooke ob-
tained from an old resident of Honolulu, was to glue thread
around the suture or elsewhere, and hold the shell in the
smoke of a whale oil lamp. When the thread was soaked off,
some weird color-effects Avere produced.

The synonymy of Partulina is less involved than that of
Achatinella. The species differ much more in shape and size,
and are less variable in color and pattern. In other words,
there has been much more speciation. Moreover, the shells
have been collected less. Probably when Maui is more fully
explored some of the accepted species will be found to inter-
grade. The Amastridcu present 110 special problems in the
distinction of species. They are not more or less difficult
than ground-shells of other parts of the world.

HISTORICAL NOTES ON THE LITERATURE.

I.

The first Achatinellid shells brought to Europe, so far as
we know, were obtained by Captain George Dixon, who visited
the Hawaiian Islands in 1786 and 1787. They were strung

HISTORY OP ACHATINELLID^. 1m

on a lei or necklace, which seeins to have been made entirely
of Achatinella apexfulva and A. decora. It appears that
four specific names were based upon these specimens. The
shell described by Chemnitz as Turbo lugubris was doubtless
from the same lei. It was purchased in London by Spengler.
While the source of Lamarck's Monodonta seminigra is not
positively known (Delessert ascribing it to Captain Cook),
yet very likely it also was from Captain Dixon's lei. There
is no satisfactory evidence that Captain Cook's expedition
brought back any Achatinellidce.

The French corvettes Uranie and Physicienne visited the
Hawaiian group in 1819. They obtained 14 species of land
shells, which were worked up by Ferussac, at that time the
foremost authority on land shells. These shells were from at
least two sources: A. decora, lugubris and spirizona were
probably from a lei made in the Kawailoa-Helemano district,
in western Oahu. A. luteola has not been rediscovered. I
formerly thought it might be from the western slope of Hawaii
(Vol. XXI, pp. 321-2), but the little I could see of that
arid coast from the steamer's deck does not favor such a
theory. A. vulpina, gravida, lor at a, turrit ella, ventulus,
textilis, tristis, auricula were collected near Honolulu; ac-
cording to Dr. C. M. Cooke, probably in Pauoa valley, where
the same association still exists, by members of the expedi-
tion, perhaps in course of an ascent of Mt. Tantalus. En-
dodonta lamellosa and contorta were probably from the same
neighborhood. The latter is said to have been found on ferns,
a somewhat unusual station ; neither has yet been quite satis-
factorily identified.

The first American work on the genus was published by
Jacob Green in 1827, when Achatina stewartii was described
and figured. It was collected by C. S. Stewart, an American
missionary who spent the years 1823-5 in Oahu.

The voyage of Captain Byron in H. M. S. Blonde, 1824-5
supplied the progressive English naturalist William Swain-
son with several fine species from the Kawailoa-Helemano dis-
trict, where the stringing of shell leis or necklaces seems to
have been carried on extensively. From the study of these

liv HISTORY OF ACHATINELLID^.

shells Swainson in 1828 erected the genus Achatinella, with
the following species : A. pica, perversa, livida, bidimoides,
rosea, pulcherrima and acuta. The type of Helix ~byronii
Wood was also from the same source.

In 1845 Dr. J. W. Mighels described twelve species of
Achatinellidtz and many other Hawaiian shells, and in the
same year Dr. A. A. Gould published seven species, collected
by the Wilkes Exploring Expedition.

In the Proceedings of the Zoological Society of London
for 1845, p. 89, Doctor L. Pfeiffer offered some remarks on
Achatinella, which he here for the first time adopted as a
genus. He enumerated 18 species, described by Chemnitz,
Ferussac, Swainson and himself. A few additional species
were described in the following year, all from the Cuming
collection.

The monograph in Conchologia Iconica by Lovell Reeve,
published in April-May, 1850, brought the subject fairly up
to date. It was marred by erroneous identifications of some
of the earlier species, which were a source of error for many
years. 45 species are admitted and figured, 15 being de-
scribed as new from the Cuming collection; about half of
which are still considered valid. Some of the Cumingian
shells were received from Mighels ; others were doubtless from
Thomas Nuttall, who collected plants and shells in the Islands,
and who is known to have contributed Hawaiian marine shells
to Cuming 's collection. The figures of Reeve's monograph
are beautifully drawn and lithographed and well colored
in the original edition, but in separate copies of the mono-
graph I have seen the lithographs are coarse (from improper
removal of the varnish put on the stones to preserve the
drawings), and the coloring is crude.

II.

In the decade 1850-1860 the subject passed largely into the
hands of Hawaiian Americans. It was the era of the dis-
covery and definition of species. The quantity of work along
these lines accomplished in five or six years is marvellous.

HISTORY OF ACHATINELLID^E. lv

Dr. Wesley Newcomb, a physician-naturalist from New
York, collected extensively in Oahu about 1850-55 ; also in
Molokai and Lanai. His two chief papers, published in
1853-4, more than doubled the number of species known.
Newcomb was an excellent conchologist of the old school.
Most of the forms defined by him are still held to be valid
species or subspecies. He gave but little attention to small
or critical species, and his localities are given by districts or
islands.

In 1854 Doctor Pfeiffer enumerated the species then known,
122 in number, and classified them in seven sections. The
systematic classification of the group began with this sketch.
Numerous species were described by Pfeiffer from this time
to 1859, partly from specimens sent by Newcomb, but mainly
from shells collected by Mr. Frick. Many of these species
have proved difficult to recognize, and a large proportion of
them are synonymous with the earlier species of Newcomb
and others. This was no doubt due to the lack of adequate
material. Mr. Frick evidently assorted his specimens
shrewdly, to get the greatest number of "species", and
many of the forms described by Dr. Pfeiffer were based upon
one or two specimens of unusual color or shape. He gave
no definite localities.

Frick himself published only one paper 'Notes on
Hawaiian terrestrial Conchology (Sandwich Islands' Monthly
Magazine, I, May, 1856, pp. 137-140). He mentioned only
one Achatinellid shell by name, Achatinella gigantea-, but
numerous MS. names were given by him, some of which were
printed years later in Paetel's Catalogue, while others linger
only in the traditions of Island collectors. Among concho-
logists, Frick 's name is chiefly remembered for the difficul-
ties ensuing from his ignorant and unscientific methods. He
seems to have been an industrious collector.

Mr. G. S. Emerson began collecting Achatinellae in the
period of Newcomb and Gulick, and some of his finds were
recorded by both of these authors. A variey of A. livida was
named for him. Some time later, Mr. J. S. and Rev. 0. P.
Emerson carried on the work begun by their father. W. T.

M HISTORY OP ACHATINELLIIXE.

Alexander, better known for his work in other directions,
also contributed to Mr. Gulick's material.

John T. Gulick began collecting Achatinellid<z in 1850, at
the age of 18. Most of his collecting was done between that
time and 1853. His work differed from that preceding by
the careful attention given to locality and food-plants of the
snails. He was also the first to collect the small ground-
shells, especially Leptachatina, extensively, as he was the first
to appreciate their characters. Two descriptive papers were
based upon this most comprehensive and valuable of all the
early collections. The first was published in 1856 ; the second,
in collaboration with Mr. E. A. Smith, not until 1873, though
dealing with material collected in the early fifties.

Mr. Gulick's descriptive work, like his field work, was on
a higher plane than that of his contemporaries. He sought
to record the marvelous variation and differentiation of the
island fauna. The suggestive geographic relations of allied
forms made strong appeal to his imagination. One sees the
same trend of thought in C. B. Adams' Jamaican papers of
about the same date; and as everyone knows, Darwin had
been deeply impressed by similar phenomena observed in the
Galapagos.

Many of the species of Gulick are now given another inter-
pretation or value ; many have been confirmed by subsequent
investigations. His classification of the group (P. Z. S. 1873)
was a long stride forward, improving the arrangements of
Pfeiffer and von Martens in important details. Though this
work is concerned chiefly with systematics, allusion should
be made to Gulick's greatest intellectual service, his theory
of segregation in its several forms, as a necessary condition
of the evolution of species, originally suggested by his studies
on Oahuan Achatinellidse. First stated in 1872, it was fully
discussed in his ''Evolution, Racial and Habitudinal, Wash-
ington, 1905.

W. H. Pease, whose occupation as a surveyor took him
over Kauai and other islands, published several new species,
and in 1869 a classification and catalogue of AchatineUid(£.

From 1855 on there seems to have been little scientific col-

HISTORY OF ACHATINELLIME. Ivii

lecting for about twenty-five years, and scarcely anything
was published on AchatinelUda, if we except the two papers
by Gulick (1873), which belong properly with his earlier
work.

III.

Several papers on the soft anatomy of Achatinellidce by
Wm. G. Binney and T. Bland, published in 1873-6, were im-
portant as showing that there are two radically diverse types
of dentition in the group.

Mr. D. D. Baldwin and Mr. Thwing began their extensive
collections about this time, and in the later eighties and first
half of the next decade a number of younger men were en-
listed, most of whom contributed new material for Mr. Bald-
win's papers. Dr. C. M. Cooke's collection, which was studied
by Professor Hyatt, and extensively used in this work, was
begun at this time.

Dr. W. D. Hartman, whose material came from Newcomb,
Pease and Baldwin, published a catalogue of the group in
1888. It was marred by many errors of all kinds.

Mr. D. D. Baldwin's papers, published from 1886 to 1908
were the most important expression of the work of this period,
and stimulated the study of the fauna by local naturalists.
His Catalogue of 1893 though modest in form, has been widely
used and quoted on account of its reliability in classification
and geographic distribution. His descriptive papers are
lucid and well considered.

The Mollusca of Fauna Hawaiiensis, by Mr. E. R. Sykes,
(1900), contains a useful synopsis of the group, and a bibliog-
raphy of Hawaiian conchology.

The "Reprint of the original descriptions of the genus
Achatinella" by Mr. E. W. Thwing (Occasional Papers
Bishop Museum, 1907) has proved very useful to Hawaiian
naturalists. The grouping of the species and the suggestions
relative to synonymy involved a good deal of original work
on the part of the compiler.

Herr Fr. Borcherding has published two profusely il-
lustrated papers, AchatineUen-Fauna der Sandwich-Insel Mol-

HISTORY OF ACHATINELLID^E.

okai, Zoologica, vol. 19, 1906, and Monographic der auf der
Sandwichinsel Kauai lebenden Molluskengattung Carelia,
Abh. Senck. Naturforscli. Ges. vol. 32, 1910. The first was
based chiefly upon material collected by Mr. Meyer and his
sons, purchased by Professor H. Schauinsland. It contains
a lengthy history of the literature of Achatinellidae and an
excellent bibliography.

IV.

In the last six or eight years there has been a revival of
interest among Hawaiian naturalists, and a number of large
collections have been formed. The exact data preserved, the
large series obtained from a great number of localities and
the new localities exploited, give these collections great value,
whether for systematic or for biometric investigations. Sev-
eral papers by Dr. Cooke have been based upon this material,
and it has also been utilized in these volumes. The few
months spent by the senior author in the Islands did not
permit anything more than a superficial study of these collec-
tions. Much of what value this work possesses is due to
information derived from material brought together by the
competent and enthusiastic Hawaiian naturalists of the pres-
ent time.

Genus NEWCOMBIA Pfeiffer.

Newcombia PFR., Malakozoologische Blatter, i, 1854, p. 117,
for A. Helena, physa, ornata, cwningi, plicata, newcombiana, cere-
alis, elongata. — MARTENS, Die Heliceen, 1860, p. 249 (type A.
plicata Migh.) — PEASE, P. Z. S., 1869, p. 649 (exclusive of
Sect. 2). — SYKES, Fauna Hawaiiensis, p. 331.--BORCHERDING,
Zoologica, xix, Heft 48 ', p. 94. — Newcombiana THWING, Occas.
Papers B. P. B. Museum, iii, no. 1, p. 138, 1907.

Shell oblong-turrited, usually coarsely sculptured, the embry-
onic shell of 3-J- whorls having comparatively coarse spirals (8
to 12 spirals on the second whorl), the last embryonic whorl
longitudinally striped. Outer lip slightly expanded or not;
columella straight or weakly folded.

Type N. plicata Migh. Distribution, Molokai and Maui.

Newcombia, by its distinct, minute, decurrent, spiral striation,
and boldly striped last embryonic whorl, is clearly a specialized
group of Partulinse. It is here accepted as of generic value only
by reason of its considerable secondary modification in sculpture
and shape of the columella. Moreover those concerned with
Hawaiian faunas now generally use the group as a genus. It
is obviously not on a par, systematically, with the genera of
Amastrinee.

N. carinella is the only dextral species known.

Pfeiffer's list under Newcombia was heterogeneous, but con-
tained several species (cumingi, plicata and newcombiana) , now
included in the genus. Professor Von Martens, in 1860, selected
A. plicata as the type. His second section of the group contains
species of Laminella. Pease, in his classification of 1869, asso-
ciated species of Heteramastra with his Newcombias. These dis-
crepant forms have been eliminated by Hartman (1888), Bald-
win (1893), and subsequent authors.

The distribution of closely related forms on Molokai and
Maui shows that the genus was developed in nearly its modern
condition while these two islands formed a single area. It is
somewhat peculiar that no Newcombia has yet been found on
Lanai, and I anticipate the discovery of the genus there, either
recent or as a fossil.

£ NEWCOMBIA.

Key to Species of Newcombia.

a. Parietal wall making an angle with the columella, the latter

having a low fold or twist.

l>. Surface smooth to the eye, finely striate spirally under
a lens. Molokai. N. perkinsi, no. 6.

N. philippiana, no. 5.
/A Surface spirally ribbed or coarsely striate, sinistral.

Molokai.
c. Pale, the later whorls cream-colored; length 15 to

25 mm.

d. Spirals acute. N. plicata, no. 1.

d *. Spirals rounded or subobsolete.

N. p. gemma j no. la.
c1. Dark colored; length 12 to 18 mm.

d. Spirals acute. X. camilicu'lata, no. 2.

d1. Spirals obtuse, base darker. N. sukata, no. 3.
//'. Surface with fine beaded spirals, a small carina at peri-
phery; zigzag-striped, dark-colored, dextral. Maui.

N. carmeHa, no. 4.
c/1. Parietal wall continuous with the columella, which is not in

the least folded. Sinistral.

b. Longitudinal folds coarse and strong, rude spirals run-
ning over them. Molokai. .Y. ifciffrri. no. 9.
/A Longitudinal folds weak or wanting.

r. Very weak folds crossed by spiral threads, the inter-
vals of which are minutely wave-striolate spirally.

N. cinninyi, no. 7.

c1. Smoother; spiral threads weak or effaced, the minute
wavy striolation well developed. Molokai.

N. cinnamomea, no. 8.

c2. Wrinkled or puckered over spiral cords; no minute
striolation. Molokai. JV. c. ualapuensis, no. Sn.

Series of N. pi Ira (a.
1. N. PLICATA ('Mighels' Pfr.). PL 1, figs. 1, 2, 3, 6.

Shell sinistral, turrited, rather thin; sculptured with acute,
elevated, spiral liree and very fine longitudinal lines; brown-

NEWCOMBIA.

whitish, spire elongate, the apex rather obtuse. Whorls 6, a
little convex, those of the summit marbled with brown. Colu-
mellar fold obsolete. Aperture semioval ; peristome unex-
panded, acute. Length 15, diam. 6 mm.; aperture 6x3 mm.

Molokai (Mighels in Cuming coll.): Kalae (Meyer); Moanui
(Thaanum).

Achatinella plicata Mighels MS., PFR., Monographia Hel.
Viv., ii, p. 235 (1848).— REEVE, Conch. Icon., pi. 6, f. 44a, b.
(1850).— NEWCOMB, Ann. Lye. Nat. Hist, of N. Y., vi, 1858,
p. 312.— THWING, Reprint Orig. Descript. Achat., 1907, pi. 3,
f. 11. — Newcombia plicata Migh., BORCHERDING Zoologica, p. 94,
pi. 9, f. 1, la.—Bulimus liratus PFR., P. Z. S., 1851, p. 261,
under B. newcombianus Pfr. (1853), no description; new naiuo
for plicata ' Migh.' Pfr.

The shell is perforate. The apex is obtuse, rounded, smooth
in the first half-whorl; then spirals begin above the suture at
the end of 1 J whorls; then about 6 to 8 smooth, subequal spiral
cords. On subsequent whorls the cords become more acute,
and some interstitial threads appear. The growth-stride become
strongly developed and irregular. The third whorl is longi-
tudinally striped with brown or pale olive, the rest of the
whorls being uniform cream color. The peristome in fully
adult shells is narrowly expanded and thickened within. Colu-
niella weakly sinuous, its edge dilated in a triangular white
callus above.

Length 18, diam, 8.2 mm.; whorls 6.

Length 25, diam. 9 mm.; whorls 6J.

Newcomb described the animal as "rather shorter than the
shell; tentacles at their origin closely approximating, short and
strongly clubbed; bottom of foot, mantle, and posterior part
light grey, entirely mottled with fine bluish dots. Motions at
first sluggish and timid but soon becoming bold, rapid and
gliding.'

la. N. PLICATA GEMMA (Pfeiffer). PI. 1, figs. 7, 8, 10, 11.

Shell subimperforate, sinistral, oblong-turrite, rather solid,
striatulate and spirally lirate (the lirse rather flattened, closely

4 NEWCOMBIA.

sulcate), white; spire turrited, the apex rather acute; suture sub-
marginate. Whorls 7, the upper ones flat, obsoletely varie-
gated with brown; following whorls rather convex, the last
about two-fifths the total length, somewhat carinated in the
middle by a more acute cord. Columella lightly plicate.
Aperture a little oblique, obauriform; peristome nearly simple;
columellar margin subreflexed, a little expanded outwardly.
Length 17, diam. 6.5, aperture 7x3 mm. Sandwich Is., Frick
in Cuming coll. (Pfr.).

Molokai: Pohakupili (Baldwin); Halawa (Borcherding, for
N. costata).

Achatinella gemma PFR. , Malak. Blatter iv, 1857, p. 230;
Monogrp.phia iv, 560; P. Z. S., 1858, p. 22. — Newcombia gemma
Pfr., SYKES, Fauna Hawaiiensis, p. 331.

Achatinella (Newcombia) sulcata Pfr., BALDWIN, Catalogue,
1893, p. 8. — Newcombia sulcata Pfr., GWATKIN; Proc. A. N. S.,
Phila., 1895, p. 238 (dentition). — Newcombia costata BORCHER-
DING, Nachrbl. d. d. Malak. Gesell, 1901, p. 57; Zoologica, p.
95, pi. 9, f. 2, 2a.

The sculpture of the last whorl or two is more or less obsolete.
This form occurs associated with more strongly lirate shells, in
which the lirse are wider than in plicata, rounded, or more or
less marked with spiral striae. The gemma form is, in fact, only
an extreme phase, with obtuse lirse, of this race, which has been
called " sulcata Pfr. ' by Baldwin, and costata by Borcherding.
The description of the latter follows:

Newcombia costata [pi. 1, figs. 4, 5, copied from Borcherding].

Shell subperforate, sinistral, turrited, rather solid; encircled
with close, elevated line, more slender in the upper whorls,
then more rounded, and sculptured with very fine longitudinal
lines; last whorl with 7 rounded, flattened ribs above the mid-
dle, the ribs suddenly wreaker below the middle, and becoming
stronger again around the umbilicus. Color yellowish-white,
glossy. Spire regularly tapering; apex rather obtuse; suture
linear. Whorls 6, flatly convex, the upper flamed with yellow,
the last forming half the total length, rounded. Aperture in-
versely ear-shaped, white within. Columella slightly folded,

iNE \VCOMBIA. £>

white. Peristome expanded, with a white lip within; colu-
mellar margin recurved. Length 18, diam. 8, aperture 8x6
mm. Halawa, eastern Molokai (Borcherding*) .

In N. p. gemma, as in typical plicata, the Iira3 are variable in
arrangement and number, no two specimens being quite alike.
Figs. 8, 10, 11 represent typical gemma, while figs. 4, 5, 7 are
the " costata' ' type, intermediate between plicata and gemma.

2. N. SULCATA (Pfeiffer). PL 2, figs. 9, 10; pi. 14, figs 9,
10, 11.

Shell subperforate, sinistral, oblong-turrited, rather solid
striatulate, and encircled with close Iira3, which are compressed
in the upper whorls, then rounded; chestnut colored, somewhat
shining. Spire regularly tapering, the apex rather acute; suture
nearly simple. Whorls nearly 6, rather flat, the upper flamed
with white, the last about two- fifths the length, the base saccate,
of a deeper chestnut shade. Columella very lightly folded.
Aperture oblique, acuminate-oval; peristome thin, the colu-
mellar margin dilated above, reflexed, outer margin a little ex-
panded. Length 12J, diam. 5f mm. Aperture 5f mm. long,
3J wide. Sandwich Is., Frick in Cuming coll. (Pfr. ).

Molokai: Ahaina (D. Thaanum).

Achatinella sulcata PFR., Malak. BL, iv, 1857, p. 231; Mono-
graphia, iv, 560; P. Z. S., 1858, p. 22. — Newcombia sulcata Pfr.,
SYKES, Fauna Hawaiiensis, p. 332, — BORCHERDING, Zoologica,
p. 97, pi. 9, f. 4, 4a. — Not of Baldwin, Catalogue p. 8, or of
Gwatkin, Proc. A. N. S.. Phila., 1895, p. 238 (radula).

" This species can be confused with no other. The whorls
are regularly, obsoletely, transversely striate, increasing in
strength to the last whorls and disappear on the lower half of
the last whorl. The color is red-brown, becoming more intense
with the increase of the wrhorls, and on the last whorl it is quite
shining dark red-brown. The same analogy which exists be-
tween plicata and costata is here found between canaliculata and
sulcata.1 (Borcherding, from specimens in the Hartman collec-
tion, probably received from Newcomb. )

Collectors of the present generation seem not to have found
this species until it was taken by Mr. Thaanum at Ahaina.

6 NEWCOMBIA.

The sculpture is most like that of N. plicata gemma, which
differs by its much larger size and lighter color, N. sulcata hav-
ing the last whorl rich chestnut colored, the next earlier whorl
chestnut or reddish, and the preceding one flamed with white.
The spiral cords of the last whorl vary in number and prom-
inence. Specimens from Ahaina are drawn on pi. 14, figs. 9,
10, 11. Length 11. diam. 5 mm. The figures on plate 2 are
copied from Borcherding.

3. N. CANALICULATA (Baldwin). PI. 2, figs. 5, 6. 7.

"Shell sinistral, very minutely perforated, somewhat solid,
acuminately turreted, apex subacute; surface sculptured
throughout with numerous acute, spiral keels, which become
blunter as they approach the apex, the interstices between the
keels exhibiting under a lens very delicate growth striae. Color
brown, upper whorls tessellated with brown and white. Whorls
6, slightly convex, lower one somewhat flattened at the base;
suture lightly impressed. Aperture oblique, oval, livid white
or light brown within; peristome acute, very lightly thickened
within, expanded, columellar margin reflexed over the small
perforation, margined with light brown on both face and the
reverse; columella very slightly developed, plain and smooth.
Length 14, diam. 6^ mm.

"Animal wrhen extended in motion as long as the shell.
Mantle slate color, margined with brown. Foot light slate,
studded on the sides and head above with spots of deeper shade.
Tentacles short and slender, dark slate.' (Baldwin.)

Molokai: Halawa (Baldwin); Punkolekole (Meyer).

Achatinellacanaliciilata'BALVWis, Proc. A. N. S., Phila., 1905,
p. 226, pi. 10, f. 28, 29. — Newcombia c., Bald., BORCHERDING,
Zoologica, p. 96, pi. 9, f. 3, 3a.

Smaller and much darker than N. plicata, and with more
strongly developed, more acute spiral ribs than N. sulcata. The
spirals are at first equal and rounded, but become unequal and
acute on the later whorls, with some smaller threads in most of
the intervals, which are marked with growth-striaB. The color
is red-brown, indistinctly streaked along growth-lines with
whitish, the spirals also being whitish or pale at their summits.

NEWCOMBIA. 7

The third whorl, or the second, third and fourth whorls are
flamed with white. The figures represent cotypes. The largest
shell in the type lot of 12 measures length 14, diarn. 7.2 mm.
The smallest is 13.3 mm. long.

3«. N. c. wailauensis n. subsp. PI. 14, fig. 5.

Larger, more robust than canaliculata, lighter in color, the
intervals between the spirals flesh-pink; spirals narrower.
Whorls 6.

Length 17 to 17.5, diam. 8.7 mm.

Length 16.5, diam. 8 mm. (smallest).

Molokai: Wailau (D. Thaanum). Cotypes in coll. Bishop
Museum and A. N. S. P.

4. N. CARINELLA Baldwin. PI. 2, figs. 1, 2, 3.

The shell is dextral, minutely perforate, oblong-turrited,
moderately solid; surface lusterless, sculptured with rather
coarse and unequal folds and wrinkles along growth-lines, and
numerous, minutely beaded spiral threads, 10 to 12 on the
penult, whorl; on the last whorl the threads are smaller and
closer below the acute peripheral keel; second and third whorls
with many even, close, nearly smooth spiral threads. Marbled
and zigzag-flamed with dark red-brown and white, the pattern
becoming more confused on the later whorls; the smooth apex
yellowish or fleshy. Aperture whitish or dark within. Peri-
stome blunt, yellow at the slightly expanded edge, usually
with a reddish submargin. Columella weakly folded, white,
reflexed.

Length 14, diam. 7 mm. ; 5J whorls.

Length 14, diam. 6.3 mm.; 6 whorls.

"Length 15, diam. 6.5 mm.; 6 whorls' (Baldwin).

East Mani: Nahiku (Baldwin).

Newcombia carinella BALDWIN, Nautilus, xix, April, 1906, p.
136.

A beautiful species, somewhat variable in contour and mark-
ing, but otherwise constant in a lot of 21 received from Mr.
Baldwin. With the sculpture of N. cumingi it unites the shape
of the plicata group, and is thus a connecting or synthetic form.

8 NEWCOMBIA.

It differs from all other Newcombias by being dextral. Figured
from cotypes.

Series of N. philippiana.

5. N. PHILIPPIANA (Pfeiffer).

Shell sinistral, subimperforate, fusiform-turrited, rather thin,
longitudinally striate, decussated with somewhat wavy close
spiral strise; brown; spire slender, the apex rather acute; suture
margined. Whorls 6, the upper flat, streaked with corneous
and white, the last whorl more convex, about two-fifths the
total length. Columella nearly simple, not folded. Aperture
oblique, angularly semi-oval; peristome simple, unexpanded,
the margins joined by a very thin callus, the outer margin
bordered with brownish, columellar margin dilated above, sub-
adnate. Length 15 to 15J-, diam. 6 mm. ; aperture 7 mm. long,
3 wide (P/r.).

Var. b. Smaller, paler, a little more distinctly striated
spirally (Pfr. ).

Sandwich Islands (Frick, Cuming coll.).

Achatinella philippiana PFR., Malak. BL, iv, 1857, p. 89;
Monographia, iv, 559. — Newcombia p. , BORCHERDING, Zoologica,
p. 103.

This species is known by the original examples only.
Borcherding believes it to be an immature stage of N. perkinsi
Sykes, basing this opinion on the smaller size and thin shell as
described by Pfeiffer. This view, which seems quite probable,
was held by Baldwin and Thwing; but until young perkinsi can
be compared with the type of philippiana, and their identity
demonstrated, both may be let stand.

6. N. PERKINSI Sykes. PI. 7, figs. 18, 19.

" Shell sinistral, narrowly perforate, elongate-fusiform, solid,
ashy-brown, or elegantly marked with zigzag streaks of brown.
Spire slender, the apex obtuse, smooth. Whorls 6£, rather flat,
the last about half the length of the shell; suture margined.
Columella subplicate. Aperture a little oblique, semioval, sub-
angular at the base; peristome simple, the columellar margin

KEWCOiMBIA. y

dilated above, adnate. Length 25, diam. 7.5 mm.; alt. aper-
ture 7.5, width 4 mm ' (Sykes).

Molokai (Perkins, Hutchison): Makakupaia (Baldwin).

Newcombia perkinsi SYKES, Proc. Malac. Soc. , London, ii,
October, 1896, p. 130; Fauna Hawaiiensis, p. 332, pi. 11, f. 36,
1900. — BORCHERDING, Zoologica, p. 102, pi. 9, f. 6. — Achatinella
(Newcombia) philippiana Pfr. , BALDWIN, Catalogue, p. 8. —
THWING, Reprint Grig. Descript. Achat., pi. 3, f. 10.

"Specimens are to be found in some collections under the
name N. philippiana Pfr. The present species however is larger,
much more solid, the whorls are flatter, the coloring is lighter
and different, the columellar lip is more reflexed and the per-
foration more conspicuous ' (Sy Ices').

In unworn shells there is a faint longitudinal ribbing after
the smooth initial half whorl. Spiral striation sets in about the
middle of the second whorl, and color-flames appear on the
third whorl. The whole embryo consists of 3J whorls, the last
one usually profusely marked with zigzag brown streaks, and
more finely striate spirally than the other species. The post-
embryonic stages have no major spirals, but under a strong lens
are seen to be covered with a minute, irregular striolation, such
as has been described for N. cinnamomea, etc. This sculpture
is more or less effaced on the last whorl, often not discernible
there, especially in individuals having weak longitudinal folds.
The later whorls are mottled, marbled or streaked with white
and flesh-color, and the last whorl often has a peripheral angle
in front. The aperture is white within, with a brown lip-
border. The columella has a long, low fold or twist, and makes
an angle with the parietal wall, as in N. plicata; columellar re-
flection usually brown-tinted. Length 21, diam. 7.5 to 8 mm.,
6J whorls.

This species is apparently the most primitive existing New-
combia, having no trace of coarse spirals on the whorls. The
embryonic spirals also are finer than in other species, more as
in Perdicella. The coloration too is rather that of Perdicella.
Cf. N. philippiana Pfr. , which is probably an immature stage of
perkinsi.

10 NEWCOMBIA.

Series of N. cmningi.

7. N. CUMINGI (New comb). PL 3, figs. 1 to 6.

Shell sinistral, acuminate, turrited, with strongly marked
transversely oblique stria? and with longitudinal incremental
striae more or less developed. Color brown with undulations of
white at the upper whorls. Whorls 5, flatly convex; suture
moderately impressed, margined. Aperture oblong-ovate; colu-
mella slightly callous; outer lip thin, elliptical. Length .7,
diam. .2 inch (Newcomb).

East Maui: Haleakala (Newc. ). Makawao (Gulick, Bald-
win). W. Maui: Wailuku (Gulick).

Achatinella cmningi NEWC., Ann. Ly.c. N. H. of N. Y. vi, p.
25 (May, 1853); P. Z. S., 1853, p. 150, pi. 24, fig. 59 (1854).—
PFR., Monographia iv. 559. — THWING, Reprint Orig. Descript.
Achat., 1907, pi. 3, f. 12.

N. cumingi is imperforate. The embryonic shell of 3^ whorls
is spirally striate and flammulate as usual (pi. 2, fig. 13). The
following whorls have nearly obsolete longitudinal folds crossed
by crenulated spiral threads, usually coarsest in the peripheral
region of the last whorl, finer on the base; the intervals minutely
marked with spirally descending, wavy stria? (pi. 3, fig. 5, de-
tail from back of last whorl, Wailuku specimen). Very often
a peripheral thread is stronger, forming a low carina on the
front of the last whorl. The surface is lusterless, dull brown.
Aperture nearly wrhite with a brown border within the thin,
acute lip. The columella is long, straight, heavily white-
calloused, and continuous in direction with the parietal wall.

Length 21, diam. 7.2 mm.; 6 whorls. From Newcomb.

Length 20, diam. 7 mm. ; 6 whorls. Wailuku.

Length 19, diam. 6 mm. ; 6 whorls. Wailuku.

The types were from East Maui, but I see no difference be-
tween those before me from Makawao (pi. 3, fig. 4) and the
West-Mauian shells (Wailuku, pi. 3, figs. 1-3, 5).

8. N. CINNAMOMEA (Pfeiffer). PI. 3, figs. 7 to 11; pi. 14, figs.

1 to 4.

Shell imperforate, sinistral, fusiform-turrited, rather solid,

NEWCOMBIA. 11

opaque, longitudinally plicatulate, spirally sublirate and very
lightly granulated, cinnamon colored. Spire long, somewhat
rectilinear, the apex rather acute; suture nearly simple. Whorls
6, slightly convex, the upper marbled with brown and white,
last whorl about two-fifths the total length, tapering and chest-
nut-colored below the middle. Columella simple, receding.
Aperture slightly oblique, semioval, subangular at base; peris-
tome simple, unexpanded, acute. Length 19, diam. 5, aperture
7§ x 3 mm. Sandwich Is., Frick in Cuming coll. (Pfr.).

Molokai : Mapulehu (Baldwin, Thaanum) ; Makakupaia
(Perkins, Meyer); Honomuni, Moanui, Kupeke and Ahaino
(Thaanum).

Achatinella cinnamomea PFR., Malak. BL, iv, 1857, p. 230;
Monographia, iv, 559; P. Z. S., 1858, p. 22. — Newcombia
cinnamomea Pfr., BALDWIN, Catalogue, p. 8. — SYKES, Fauna
Hawaiiensis, p. 331. — Newcombia cumingii Newc. (in part),
BORCHERDING, Zoologica, p. 99, pi. 9, f. 7 (Makakupaia).

This form resembles N. cumingi closely in shape and micro-
scopic sculpture, but it differs by the weakness of the spiral
cords, which vary from weak to almost effaced. As this differ-
ential feature seems to hold in considerable series of both forms,
I think it best to retain the name cinnamomea for the Molokai
race, although the relationship to the Mauaian N. cumingi is so
close that perhaps cinnamomea might better be ranked as a sub-
species.

On account of the weakness of the coarse sculpture, the
minute, wavy, crenulated or granular striation (pi. 3, fig. 7) is
more distinct in this race than in Mauian cumingi. The axis is
often perforate. Immature shells are more strongly lirate than
the adults, and often could not be distinguished from N. cumingi.
N. cinnamomea is therefore to be viewed as a derivative of cumingi.

The fully formed embryo of 3J whorls, length 4.7 mm., has
a very heavily calloused columellar fold, but no lamella (pi. 1,
fig. 12). Younger embryos have the callus weaker, hardl}'1
noticeable in those of two whorls (pi. 1, fig. 9). All are per-
forate.

The types were supplied by Frick who gave no locality other
than the " Sandwich Islands.' In specimens collected by New-

12 KEWLOMBIA.

comb at about the same time — almost sixty years ago — the color
of the last whorl is a ricn cinnamon, between raw sienna and
ochraceous (of Ridgway's " Nomenclature of Colors''), fading
upwards, and darker at the base (pi. 3, figs. 8, 9). This was
evidently the coloration of Pfeiffer's type. Some of the
Mapulehu shells are of this color, others being darker, as in
those from Moanui.

At Moanui (pi. 14, fig. 1) the base of the last whorl is burnt
umber, the upper part drab or olive, the transition often abrupt
at the periphery. Frequently there are numerous umber bands
occupjdng half the space between periphery and suture. The
size is greater than at other localities up to length 23J, diam.
8J, and length 24, diam. 8 mm.

At Ahaino and Kupeke (pi. 14, figs. 2, 3, 4) the shells are
small, the microscopic granulo-striatkm well developed, other
sculpture obsolete. The last whorl is ochraceous with several
umber bands and basal tract, or olive with an umber base.
The largest shells are 18 mm. long, 6 wide, most being between
16 and 17 mm. long. This local race may be called var.
decorata.

Specimens from Honomuni (pi. 14, figs. 6, 7) are very much
lengthened, ochraceous, base a little darker, spire worn, dull
violet or purple. Columellar reflection very small, adnate; a
"false umbilicus' visible in basal view. Length 19, diam. 6
mm., whorls 6. This form may be called var. honomuniensis.

8ft. N. c. UALAPUENSIS n. subsp. PL 2, figs. 11, 12.

Similar to N. cumingi in having well-developed spiral cords,
but the surface between and over them is minutely but strongly
puckered or wrinkled longitudinally (fig. 12), and not spirally
striolate as in cumingi and cinnamomea.

Molokai: Ualapue (D. Thaanum), no. 94490 A. N. S. P.;
Kahaanui (Thaanum).

The upper whorls are considerably eroded in the type lot, but
the younger individuals show the striped pattern of cumingi.
It is a little smaller than N. cumingi, length of figured type 20,
diam. 6J mm.

This race seems about equally related to cumingi and ciniia-
moraea, with incipient characters of pfeifferi.

NEWCOMBIA. 13

9. N. PFEIFFERI (Newcomb). PL 2, figs. 4, 8, 14.

"Shell sinistral, acuminate, turrited, with the surface irregu-
larly cut up into furrows, ridges and tubercles by deep longi-
tudinal sulcations crossed by strongly developed transverse
striae. Color brown, with white longitudinal lines on the upper
whorls. Whorls 6, flatly convex; suture deep. Aperture
oblong-ovate. Columella plain and smooth; outer lip simple.
Length .65, diam. .25 inch' (Newc.).

Molokai (Newcomb): Puukolekole (Meyer); Kaluaaha
(Baldwin).

Achatinella pfeifferi NEWCOMB, Ann. Lye. Nat. Hist, of New
York vi, p. 25 (May, 1853); P. Z. S., 1853, p. 150, pi. 24, f.
58 (1854). — Newcombia pfeifferi Newc., SYKES, Fauna Hawaii-
ensis, p. 332. — Bulimus newcombianus PFR. , Monographia iii, p.
414 (1853); P. Z. S., 1851, p. 261 (December, 1853).— New-
combia newcombiana PFR., Malak. Bl. i, 1854, p. 119. — PEASE,
P. Z. S., 1869, p. 649. — A. newcombiana THWING, Reprint Orig.
Descript. Achat. , 1907, pi. 3, f. 9. — N. neivcombia Pfr., HART-
MAN, Proc. A. N. S. Phila., 1888, p. 41. — BORCHERDIXG,
Zoologica, p. 98, pi. 9, f. 5, 56.

Very distinct by the coarse longitudinal folds and rude spirals
of the last two whorls.

The initial half whorl is smooth; the next whorl has about 8
close, smooth spirals; on the following whorl these become
more widely spaced, and a color pattern of broad brown and
white longitudinal stripes appears, continuing to the end of the
embryonic stage, which comprises 3J- whorls. Weak folds then
begin on the neanic stage, gradually becoming stronger, the
color being indistinctly marbled with white on a dark brown
ground. The last two whorls are brown with the high points
of the ribs yellowish; the sculpture increasing in strength.
There is a minute sculpture of irregular, crenulated striae over
the coarser ribs and cords. The whorls are strongly convex.

Length 15, diam. 6 mm.; 5^ whorls.

Length 17, diam. 6 mm.; 6J whorls.

It varies in details of sculpture and in the degree of elongation.

14 PARTULINA.

Genus PARTULINA Pfeiffer.

Partulina PFR. , Malakozoologische Bliitter, i, p. 114, 1854
(for A. virgulata, tessellata, splendida, perdlx, goiddi, dubia). —
VON MARTENS, Die Heliceen, 1860, p. 243, type A. virgulota
Migh. — GULICK, P. Z. S., 1873, p. 90, same type. — SYKES,
Fauna Hawaiiensis, p. 311, same type.

Includes as sections, Perdicella, Eburnella and Bcddunnia.

Shell ovate-conic, usually perforate, and having distinct
minute sculpture of spirally descending rippled strife. Embryonic
shell after the first whorl spirally striate. Color pattern of
oblique stripes or spiral bands, the last embryonic whorl gen-
erally striped. Lip expanded more or less; columella folded or
almost simple.

Type Partulina virgulata (Migh.). Distribution, Molokai,
Lanai, Maui and Hawaii, one species on Oahu.

The Achatinellina of the islands from Molokai southeastward
form a group of common ancestry, characterized by their spiral
and decurrent sculpture, perforate axis, oblique stripes and
spiral striae on the early whorls, etc. Some or all of these
characters may fail in particular species, but their general prev-
alence leads us to believe them the heritage from a common
ancestral stock. The Oahuan Achatinellina do not possess the
sculpture and color-pattern noted above as characteristic of the
other islands, and evidently are more related inter se than to
any Molokai- Hawaii snails. Although the difference between
Oahuan and Molokai forms are sometimes elusive and not to be
laid down in formal diagnoses, yet it may be allowable to ex-
press, by a generic separation, the idea that we have to do with
two collateral stocks of arboreal snails, the one on Oahu, the
other on the islands eastward.

Newcombia has the color and sculpture pattern of typical
Partulina, but with other special modifications which make it
advisable to treat that group as a genus.

Several subdivisions or sections of Partulina have been named.
They are not of much systematic importance, and there are
some intermediate species; yet as they seem to be natural groups
we admit them here.

PARTULINA, SECTION PKRDICELLA. 15

Sections of Partulma.

a. Columellar fold prominent, heavily calloused.

/>. Spiral sculpture distinct, at least at the early whorls;
shell not brilliantly glossy.

Section Partulma s. str., species 8-35.

61. Spiral sculpture weak or obsolete; shell smooth and
glossy, light colored, uniform or banded.

Section Eburnella Pse. , species 36-47.
a1. Columellar fold wanting or weak, not calloused.

b. Spire slenderly conic; rather capacious forms; outer lip
narrowly expanded.

Section Baldwinia Ancey, species 48-53.
61. Shell small, oblong or oblong-conic; outer lip not ex-
panded.

Section Perdicella Pse., species 1-7.

Section PERDICELLA Pease.

Perdicella PEASE, P. Z. S., London, 1869, p. 648. — SYKES,
Fauna Hawaiiensis, Mollusca, p. 329, type A. helena Newc.

Small Partulintz having protractive stripes on the last embry-
onic whorl, later whorls striped or with a peripheral band;
sculpture Partulinoid; col umella nearly straight, or if developed
thecolumella fold is moderate, convex, scarcely or not truncate;
outer lip thin, not expanded. Type P. helena Nc.

Perdicella differs from Partulma by the unexpanded outer lip,
the weakness of the columella fold and the small size of the
shell. In coloration and sculpture it is altogether Partulinoid.
This affinity was recognized by C4ulick, who in his classification
of 1873 ranked Perdicella as a section of Pa-rial nia. Such species
as fulgurans, theodorei and winniei are directly intermediate be-
tween Perdicella and Partulina. Perdicella is not closely related
to Baldwinia, although in both the columellar fold has degener-
ated. It is a branch from the tessellata or marmorata series of
Partulinse.

Key to Species of Perdicella.

a. Molokai. Shell sinistral, usually with a light or dark band.

P. helena, no. 1.

16 PARTULINA, SECTION PERDICELLA.

a1. Maui.

b. Shell sinistral, with long, straight-sided spire, finely
zigzag-streaked or speckled, with a dark peripheral
band. P. ornata, no. 2.

b l. Shell dextral, oblong or ovate-conic.
c. Columellar fold distinct, convex.

d. Pattern of longitudinal chestnut lines, base
umber with a spiral umber band; col. fold
prominent; length 14 mm. P. zebra, no. 3.
d l. Boldly striped or zigzag-striped, without a spiral
band; columella white or pink, with a rather
strong fold.
e. Length about 13 mm. ; form narrow.

P. zebrina, no. 5.
e\ Length about 15-16 mm. ; form wider.

P. falgurans, no. 6.
c1. Columellar fold very weak.

d. Lip and columella brown; diam. not much over
half the length. P. mauiensis, no. 4.

d1. Columella white or tinged with purple-brown;
diam. nearly two-thirds the length.

P. kuhnsi, no. 7.

1. P. HELENA (Newcomb). PI. 4, figs. 1 to 7.

Shell sinistral, ovate-conical, with finely decussating stria? ;
rufous, alternating with broad longitudinal zigzag white lines
covering the entire shell ; the last whorl often encircled by a
white band. Whorls 5, rounded ; suture deep, simple. Aper-
ture ovate, columella slightly callous. Length 0.5, breadth 0.22
inch (Newcomb).

Molokai, on the Ti tree (Newcomb). Kamalo to Kalae
(Baldwin); Kalae and Makakupaia (Perkins); Kealia (Meyer).

Achatinella helena NEWC., Annals Lyceum of Nat. Hist, of N.
Y. vi, p. 27, April, 1853; P. Z. S., 1853, p. 151, pi. 24, f. 63;
1854, p. 311. — PFR., Monogr., iv, 561. — A. (Partulina) h.,
BALDWIN, Catalogue p. 6. — Perdicella helena Nc. , PEASE, P. Z. S.
1869, p. 648. — SYKES, Fauna Hawaiiensis, p. 330. — BORCHERD-
ING, Zoologica, p. 75, pi. 4, f. 17 (Kealia), and 18 (Maka-

PARTULINA, SECTION PERDICELLA. 17

kupaia). — THWING, Reprint Orig. Descript. Achat. p. 134, pi.
3, fig. 5.

Newcomb states that A. helena "is extremely limited in its
locality, which has been twice carefully searched by myself
without discovering larger specimens, dead or alive, that ap-
proached it in form. Several of the specimens contained young
in the oviducts.' It was taken " within the coil of the Ti tree
leaf, as it starts from the trunk. '

The shell is sinistral, perforate; summit obtuse. There are
fully 2J convex embryonic whorls, the initial half-whorl smooth,
the next two evenly, rather strongly striate spirally; last em-
bryonic whorl is variegated with broad, forwardly descending,
brown and white flames. The neanic and last whorls have a
much finer and very close spiral sculpture of minutely waved
or crinkled striae, and a variable pattern of zigzag streaks, inter-
rupted by a cream-white peripheral belt; the axis in a light
area. The outer lip is thin, columella very short, with a rather
strong callous fold, and broadly but shortly dilated.

Length 11.3, diam. 6.7, aperture 5.3 mm.; 5J- whorls.

Length 12.2, diam. 6.5, aperture 6 mm.; 5^ whorls.

Fig. 3 represents a typical specimen. Five specimens from
Newcomb, and others from Baldwin, the University of Wiscon-
sin, Cooke collection, etcetera, examined. There is considerable
variation in details of color-pattern, as in all related species.
Sometimes the stripes on the later whorls are smeared or partly
defaced, reduced to indistinct streaks. Occasionally the whitish
subperipheral belt does not appear until near the end of the last
whorl (fig. 4, Cooke coll.), or it may be entirely absent (fig. 2).
The color of the peripheral band, like the ground-tint of the
shell, varies from whitish to yellow.

Color var. balteata n. v. has a single brown belt at the peri-
phery, continuous or indistinctly interrupted, on a pale fleshy
or brown-tinted ground, the flames of the last embryonic whorl
faint (pi. 4, fig. 7).

A pretty color-form in the Cooke collection has the periphery
occupied by a series of dark spots, oblique in one example,
chevron-shaped in another, the flammules elsewhere reduced
to weak streaks or spots (fig. 6).

18 PARTULINA, SECTION PERDICELLA.

Var. minuscula Pfeiffer. PL 5, fig. 6. Shell subim perforate,
sinistral, ovate-turrite, rather thin, under a lens minutely decus-
sate, scarcely shining, brown-whitish; spire turrite-conic, the
apex slightly obtuse; suture simple. Whorls 5, very slightly
convex, the intermediate ones variegated with brown, last whorl
a little shorter than the spire, encircled with a brown band and
with a brown area on the rotund base. Columella scarcelv

V

plicate, semioval; peristome simple, acute, the columellar mar-
gin dilated above, reflexed. Length 10, diam. 5 mm. ; aper-
ture 4-| mm. long, 2f wide (Pfr. ).

Molokai Mountains at 4000 ft. (Perkins). Sandwich Islands
(Frick in Cuming coll., type loc. ).

Achatinella minuscula PFR., Malak. BL, iv, 1857, p. 231; P.
Z. S., 1858, p. 22, pi. 40, f. 10; Monographia, iv, 562.— Per-
dicella m., PEASE, P. Z. S., 1869, p. 648. — SYKES, Fauna
Hawaiiensis, p. 330. — Cf. BORCHERDING, Zoologica, p. 76.

Pfeiffer' s figure, which we copy, shows a color-pattern like
that of some immature specimens of Helena. Borcherding con-
siders it a synonym of that species, an opinion which I think
well-founded. It seems to be a wider shell than the dextral
P. zebrina Pfr., which it resembles closely in coloration. Mr.
Baldwin reported minuscula from Lahaina, Maui, evidently con-
fusing it with the quite different P. kufmsi.

2. P. ORNATA (Newcomb). PI. 4, figs. 8, 9, 10.

" Shell sinistral, acutely pyramidal, shining; whorls 6, plano-
convex, margined above; suture well-impressed; aperture sub-
quadrate in adult, subovate in immature shells; lip slightly
thickened at the edge; columella broad and flattened; surface
of shell covered with alternating undulations or zigzag markings
of white and black arranged longitudinally, with a subcentral
transverse black band, sometimes margined with a white one
below on the last whorl. Length 10, width 4 twentieths of an
in ch ' ( Newcomb ) .

East Maui: in a deep ravine back of Lahaina (Newcomb,
type loc.). West Maui: Mt. Helu, 4000 ft. (W. F. Kaae).

Achatinella ornata NEWC. , P. Z. S., 1853, p. 149, pi. 24, f.
55.— Pfr., Malak. Bl. 1854, p. 118; Monogr. iv, p. 561.—

PARTULINA, SECTION PEUDICELLA. 19

THWING, Reprint Orig. Descript. Achat. , p. 136, pi. 3, f. 7.-
Perdicella ornata Nc. , BALDWIN, Nautilus, xix, p. 113.

A rare species, of which twenty specimens from the Newcomb
and Gulick collections, taken 50 to 60 years ago, are before us,
all "dead" shells, and a smaller series of "live" shells from
Baldwin. The straightly pyramidal spire and subangular peri-
phery give the shell an aspect of its own. The suture is very
distinctly margined by a band defined by an impressed line in
some shells, but in most of those seen there is no trace of such
margination.

First 1J or 2 whorls are flesh or whitish flesh-colored; next
half whorl has broad flexuous or irregular flesh-brown and
white stripes; after which the brown markings become narrow,
zigzag, and on the last whorl they are often dislocated, spotted
or mottled, though sometimes distinctly striped as in Newcomb's
type figure. On the last whorl there is a subperipheral brown
belt in all the specimens seen. It is bordered below by a pale or
cream-colored belt, more or less distinct, The columella is
dilated above in a small triangle, calloused, and convex, ob-
liquely trancated far above the base.

Length 14.8, diam. 6.5, aperture 6 mm.; 6 whorls.

Length 14.8, diam. 6, aperture 5.8 mm.; 6J whorls.

3. P. ZEBRA (Newcomb).

"Shell dextral, conically elongate, shining, with microscopic
decussating striae; whorls 5J round, narrowly margined above,
suture well marked; aperture ovate; lip thin; columella short,
abruptly terminating in a large prominent plait; color of epi-
dermis yellowish white, alternating with longitudinal chestnut
lines; base of an umber color, with a revolving line of the same
color above. Length 11, width 5 twentieths of an inch' [I3f
X 6J mm. (Newc.)

East Maui (Newcomb).

Achatinella zebra NEWC. , Annals of the N. Y. Lyceum of
Natural History, vi, p. 142, Oct., 1855. — PFR., Monogr. ix, 537.

Described from a single specimen, and not figured. It seems
to resemble P. ornata in color-pattern, differing from zebrina Pfr.
by the basal band.

20 PARTULINA, SECTION PERDICELLA.

4. P. MAUIENSIS (Pfeiffer). PL 4, figs. 11, 12, 13, 14.

Shell subperf orate, ovate-oblong, rather thin, longitudinally,
irregularly striate, decussated with very close spiral striae visible
under a lens, hardly shining; whitish, variegated with serrated
brownish-corneous streaks. Spire regularly conic, apex obtuse,
suture shallow, slightly margined. Whorls 5J, rather flat, the
last nearly two-fifths the total length, somewhat tapering at the
base. Aperture little oblique, oblong. Columellar fold obso-
lete. Peristome simple, unexpanded, brown-bordered, the
columellar margin reflexed, subadnate. Length 13. diam. 6,
aperture 6x3 mm. (Pfr.).

West Maui (Newcomb); Makawao to Huelo (Baldwin).

Achatinella maniensis Newc. [typogr. err. for mauiensis]
PFEIFFER, P. Z. S., 1855, p. 207 (1856). — A. mauiensis Newc.,
PFR. , Monographia, iv, p. 563. — NEWCOMB, Amer. Jonrn. of
Conch., ii, 1866, p. 217, pi. 13, fig. 16.— THWING, Reprint
Orig. Descript. Achat. , p. 134, pi. 3, f. 6.

The shape and coloration are about the same as in P. zebrina
Pfr. , but the aperture is dark brown within (sometimes over-
laid with whitish), and the brown columella has only a low
fold situated high, and not heavily calloused, much as in some
Bulimuli. The whorls of the spire are sometimes distinctly
margined below the suture. Figured from shells received from
Baldwin. Length 12.2, diam. 6.2, aperture 6 mm.; 5 J whorls.

5. P. ZEBRINA (Pfeiffer). PI. 5, figs. 5, 8 to 12.

Shell imperforate, dextral, rather thin, smooth, very minutely
decussate under a lens; glossy; whitish, very elegantly marked
with blackish-chestnut zigzag streaks. Spire somewhat turrited,
apex obtuse, brown; suture with a thread-like margin. Whorls
5, moderately convex, the last nearly three-sevenths the length.
Aperture slightly oblique, truncate-oblong, lilac within; colu-
mellar fold above, twisted, callus, slightly prominent. Peri-
stome simple, unexpanded, brown-bordered. Length 12J,
diam. 6, aperture 6x3 mm. Sandwich Is., Frick in Cuming
coll. (Pfr.).

East Maui: Honomanu (Baldwin).

PARTULINA, SECTION PERDICELLA. 21

Achatinella zebrina PFR., P. Z. S., 1855, p. 202 (1856); Mono-
graphia, iv, 561. — A. zebra No., BALDWIN, Catalogue, p. 7.

''Related to A. ornata Nevvc. , but dextral and quite distinct
in the characters of the aperture ' (Pfr.).

This species is known to us by specimens collected by Mr.
Baldwin. About If whorls at first are uniform light reddish
brown. The next half-whorl has wide alternating, more or less
forwardly-descending stripes of white and dark brown. After
that, the stripes become narrower, more sharply defined, and
more or less irregular or dislocated, the light and dark stripes
being about equal in width. The ground remains white nearly
to the base in some shells, in others becoming coffee-tinted,
darkest at the base. The spiral striation of the embryonic shell
is extremely delicate. The interior is purplish in fully adult
shells. The columella has a rather heavy callous fold, situated
high, or sometimes heaviest at the lower end, but there is no
lamella properly speaking. The axis is imperforate. Length
13.2, diam. 7, aperture 6 mm. 5J whorls.

P. zebrina differs from P. mauiensis chiefly by the much
more prominently folded and calloused white or flesh-tinted
columella.

6. P. FULGURANS (Sykes). PL 5, figs. 1 to 4, 7.

Shell subperforate, dextral, ovate-turrited, glossy, closely
sculptured with spiral lines under a lens, whitish, very elegantly
marked with zigzag chestnut stripes. Suture moderately im-
pressed, the apex rather obtuse. Whorls 5^, plano-convex,
the last two-thirds the length of the shell. Aperture ovate-
piriform, lilac within; right margin of the peristome simple,
columellar margin somewhat reflexed; columellar fold twisted,
somewhat projecting, moderate, rapidly ascending. Length 16,
diam. 8, length aperture 8.1 mm. (Sykes).

East Maui: Makawao to Huelo (Baldwin).

Achatinella zebrina Pfr. , BALDWIN, Catalogue p. 8, not of
Pfeiffer. — Perdicella zebrina Pfr., BORCHERDING, Zoologica xix,
p. 77, pi. 4, f. 19. — Perdicella fidgurans SYKES, Fauna Haw-
aiiensis, Mollusca, p. 329, pi. 11, f. 5, 1900.

"This very pretty shell is akin to P. zebrina Pfr., but may

22 PARTULIXA, SECTION PERDICELLA.

be readily separated from it by its greater size, by being much
broader in proportion to the length, and by the color-pattern
being finer in design and more zigzag' (Sykes).

The embryonic shell is like that of P. zebr'ma, and the
sequence of patterns exactly similar. An almost fully formed
embryo is 4.2 mm. long, imperforate, with a straight, very
slightly calloused columella, 2^ whorls.

The columella resembles that of P. zebrina, and is much
more strongly twisted than in the East Mauian P. mauiensis.
The color-design is variable, but a majority of the shells before
us are similar to fig. 7. Mr. Sykes's original figure is copied,
fig. 4.

Length 15, diam. 8, aperture 7.6 mm.; 5i whorls.

Length 13, diam. 7.8, aperture 6.7 mm.; (not quite mature).

7. P. KUHNSI Pilsbry, n. sp. PI. 14, figs. 8, 12-15.

The shell is dextral, perforate, conic, thin; outlines of spire
straight; whorls moderately convex. Embryonic whorls white
or brownish, uniform or more frequently having festooned axial
stripes or two bands of lunate spots on the last half-whorl.
Subsequent whorls maculate with brown; the last whorl streaked
and maculate with brown, and encircled with white bands, of
which one on the upper surface, another below the periphery
and an umbilical area are the most constant. Other specimens
have the last embryonic whorl broadly striped with brown, last
whorl copiously zigzag-striped with chestnut on a white ground;
this pattern is sometimes interrupted by one or two white spiral
bands. Surface glossy, densely and minutely striate spirally,
the striation weak on the last whorl, Growth-striae weak.
Aperture white within; peristome thin, unexpanded; columella
white, or having a brown or purplish stain in the lower part,
nearly straight, but in oblique view a low fold is visible within;
sometimes the fold is obvious in a front view; outer edge tri-
angularly reflected.

Length 13.2, diam. 8 mm.; whorls 5-J.

Length 12, diam. 7.2 mm.

West Maui: Honokohua, type loc. ; Honokowai; Mt. Lihau:
Mt. Helu; Mauna Hoomaha; ridges above Lahaina; Hailaau;

PARTULINA. 23

Kapuna; Moomuku and Honolua (Thaanum). Cotypes in
coll. Bishop Museum and A. N. S. P.

Achatinella minuscula Pfr. , Thwing, Reprint Orig. Descript.
Achat., 1907, pi. 3, f. 8. — ? Achatinella minuscula Pfr., BALD-
WIN, Catalogue p. 6, 1893. Not A. minuscula Pfr.

"Not common anywhere, but a widely distributed species.
It is undoubtedly the Part, minuscula Pfr. of Baldwin's Cata-
logue' (Thaanum').

It differs from P. zebrina Pfr. and fulgurans Sykes by the
much more slender conic spire, which resembles that of P.
helena.

This is a polymorphic species in which several mutations
have been established in the same colony. The original pattern
was probably zigzag-striped, the derivatives differing by the
development of white zones and coalescence of the interrupted
stripes in a spiral direction. Figs. 12 to 15 represent shells
from the type locality, 12 and 14 being the prevalent color-
forms.

Shells from Honokowai (pi. 14, fig. 8) are snow-white above
the periphery, yellow below it, the periphery marked by a band
of chestnut spots. There is also a very narrow dark crescent
behind the columella and some indistinct dark streaks and
spiral lines on the base. Length 14.5, diam. 8.8 mm. I have
not seen specimens from the other localities given by Mr.
Thaanum.

This species is named in honor of Mr. D. B. Kuhns, to whose
ability in the field we owe many new and interesting Achatinel-

Section PARTULINA (restricted).

Larger, more capacious shells than the Perdicdlce, and having
the columellar fold well calloused, and the outer lip usually
well expanded, thickened within. Sculpture and color-patterns
about the same. They live on Molokai (species no. 8 to 15),
Lanai (species no. 16), and Maui (species 17 to 35).

24 PARTULINA, MOLOKAI.

SPECIES OF MOLOKAI.

Key to Achatinellinae of Molokai.

NOTE. — Yellow forms plain or with zigzag black stripes should
be looked for in Laminella, Vol. XXI, p. 345.

A. Shell elongate, dull, with distinct or strong spiral sculpture,
the columellar fold weak or wanting.

Newcombia, p. 1.

A1. Shell very glossy and smooth, porcelain-like, coloration
light and in bands, never longitudinal stripes or flames;
columellar fold strong.
a. Shell dextral.

b. Spire long; shell white, uniform or with gray or

light brown streaks, generally having a subperi-
pheral dark band or several bands; length 16-19
mm. mighelsiana, no. 45.

ft1. Spire shorter, concavely conic, the last whorl bulg-
ing; yellow, usually with a chocolate subsutural
line or band; length 13-18 mm. polita, no. 44.
62. Spire straightly conic, last whorl not bulging, color
various. bella, no. 42.

a1. Shell sinistral, obesely ovate-conic, yellow, the embry-
onic whorls wrhite with a dark band.

subpolita, no. 43.

A*. Shell ovate-conic or pyramidal, showing moderate sculpture
under a lens; last embryonic whorl usually variegated;
columellar fold well developed.
6. Embryonic whorls having a dark spiral band.

virgulata, no. 8.

6'. Embryonic whorls having vertical or protractive stripes,
or sometimes plain.

c. Ovate-conic, rather large, length usually 23 to

26 mm.

d. Surface dull, fine spiral sculpture being well
developed; banded or streaked.

rw/rt, no. 10; tessellata, no. 9.
dl. Surface glossy; striped or plain.

proxima, no. 11.

PARTULINA, MOLOKAI. 25

c . Narrower, pyramidal-conic, sinistral.

d. Small, length 11 to 13 mm. Helena, no. 1.

dl. Larger, length over 17 mm.

e. Longitudinally boldly striped throughout;

length 18 mm. theodorei, no. 12.

el. Suture bordered below by a dark band,

others at periphery and columella; lip

narrow, white; length 20 to 22 mm.

mucida, no. 13.

e2. Suture bordered by a white band or line;
lip and columella brown; length 21 to 25
mm. redfieldij no. 15.

es. No conspicuous sutural border; variously
streaked, often with a subperipheral white
band, rarely others; lip wide, both lip
and columella white; length 22 to 26 mm.

divightii, no. 14.

Series of P. virgulata.

Embryonic whorls bicolored, having an upper white and
lower dark spiral zone; later whorls varying from many-banded
to plain. Shell either dextral or sinistral.

While this group has the characteristic sculpture and form of
the other Partulinse, it is remarkably aberrant in the color-
pattern of the embryonic whorls.

8. P. VIRGULATA (Mighels). PL 6, figs. 1 to 6, 9, 10.

' ' Shell ovate-conic, light fawn color, beautifully adorned with
dark brown bands, more or less numerous; iniperf orate; whorls
5, convex; incremental strise delicate. Aperture oblong; lip
reflected, slightly inflected. Length 1 inch, diam. f inch '
(Mighels).

Embryonic whorls nearly 3, the first purple-brown, the next
bicolored, having a wide snow-white band below the suture.
A narrow, more or less distinct cream-white subsutural band
continues to the end. Spiral bands appear about the middle of
the first neanic whorl, are often pale at first, becoming dark
chestnut on the last whorl, which is variously marked with

26 PARTULINA, MOLOKAI.

lines and bands. Under a glass the spiral striation is dense
and deeply cut, the striae rippled and descending. The axis is
either closed or slightly open. Lip slightly expanded and well
thickened within. The colnmella is reflexed in a short, wide
triangle, and the columellar fold varies from distinct to weak.

Length 25 to 26, diam. 15.2, length aperture 14 mm.

Length 22.5, diam. 15, length aperture 13 mm.

Length 23, diam. 14, length aperture, 11.6 mm.

Length 21, diam. 12.3, length aperture 10.5 mm.

Molokai, in the eastern part, Ualapue, Kaluaaha, Mapulehn,
Waialua, Halawa and Pelekunu (Meyer).

Partula virgulata MIGHELS, Proc. Boston Soc. of Nat. History,
II, 1845, p. 20. — PFR., Monogr., iii, p. 454. — Achatinella virgu-
lata High., REEVE, Conch. Icon., pi. 1, figs. 3, 4, 5, 5b. — NEW-
COMB, P. Z. S., London, 1854, p. 311; Lye. N. H. of N. Y., vi,
1858, p. 312 (description of animal). — THWING, Reprints Orig.
Descript. Achat., pi. 2, f. 8. — Partulina virgulata Migh., BOR-
CHERDING, Zoologica, xix, Heft. 48, p. 50, pi. 1, figs. 1-20. —
Achatinella rohri PFR., P. Z. S., 1846, p. 38. — Bulimus rohri
PFR., Zeitschrift fur Malak., 1846, p. 115; Monographia, ii, p.
74. — Bulimus insignis Mighels on labels, REEVE, Conch. Icon.,
Achatinella, species no. 3. — Achatinella virgulata var. halawaensis
Baldwin in coll., BORCHERDING, Zoologica, xix, p. 52, pi. 1, f.
13-16, 1906.

This species is well distinguished by the broad band above
the suture on the embryonic whorls, usually purple-brown but
sometimes yellowish and quite pale, and the total absence of
flame markings. It is indifferently dextral or sinistral.

The typical form (pi. 6, figs. 1, 3), such as occurs at Mapu-
lehu, while varied a good deal, is rarely so dark as the forms
from the neighboring valley Kaluaaha, in which the light ground
is often reduced to narrow lines; or in Ualupue, the next valley
westward, the light ground is altogether extinguished (figs. 5, 6).
These dark shells are very glossy, with much fainter spiral striae
than the typical form, which has little or no gloss. Fig. 9 rep-
resents a form from Ualupue, fig. 10 from Pelekunu, after Bor-
cherding. Uniform purplish-flesh tinted examples, without
bands on the post-embryonic whorls, perhaps occur in all the

PARTULINA, MOLOKAI. 27

colonies. They are present in the lots of the typical form which
I have seen.

8a. Var. halawaensis Baldwin (pi. 6, figs. 7, 8, 11), which
occurs further east at Halawa, is white or pale straw-colored,
becoming darker towards the outer lip, without post-embryonic
bands, or with them narrow and few; lip brown-edged within.
Dark band on the embryonic shell normal, or rarely much
reduced. Contour long. Length 28, diam. 15.2, aperture
15 mm.

A peculiar form (pi. 6, fig. 2) from the University of Wis-
consin collection has the embryo white with a narrow chocolate
band, the last 2£ or 3 whorls fawn writh many narrow longi-
tudinal darker streaks and only faint traces of spiral bands.
The shell is very glossy and small, length 19.7 to 20.5 mm.
Exact locality of these specimens unknown, but Herr Borcher-
ding, who has given a magnificent plate of 40 figures to the forms
of virgulata, figures this race from Pelekunu, on the northern
water-shed, together with small, slender, nearly white examples.

It is evident that a good deal of local differentiation has
taken place among the colonies of virgulata.

A. rohri was acknowledged by Pfeiffer to be a synonym of
virgulata. It was thus described: "Shell perforate, ovate-
conic, rather solid, longitudinally striatulate, decussated by
very close spiral striae; whitish fulvous, variously ornamented
with narrow chestnut bands; spire conic, rather acute. Whorls
6, but slightly convex, the last about as long as the spire, com-
pressed in the middle. Columella twisted, callous. Aperture
subtetragonal, glossy white within; peristome slightly expanded,
labiate within, the margins subparallel, the right margin shortly
arcuate above, columellar margin reflexed, nearly closing the
perforation. Length 24, diam. 13, aperture 13 mm. long, 6f
mm. wide inside.'

Group of P. tessellata.

Embryonic whorls unicolored or having obliquely axial
stripes.

28 PARTULINA, MOLOKAI.

9. P. TESSELLATA (Newcomb). PL 6, figs. 12 to 21.

"Shell sinistral, ovate-oblong, solid, with minute decussating
striae, color white or fawrn-colored, variously striped or not with
black and chesnut bands, upper whorls always tessellated with
black and white; whorls convex, the last somewhat inflated;
aperture white or roseate, ovate, effuse below; columella short
and broadly callous; columellar lip broad and slightly reflected.
Length 1 to 1.1 inch; breadth 0.6 inch.

" Body light gray, mantle slate color ' ' (iVewc.).

Molokai (Newc. ): Kalae, Kealia, Kalawao, Kahanui, Maka-
kupaia, and Pelekunu valley (Meyer).

Achatinella tessellata NEWC., Ann. Lye. N. H. of N. Y. , vi, p.
19, May, 1853; P. Z. 8., 1853, p. 139, pi. 23, fig. 26; 1854, p.
311. — PFR., Monogr. , iv, p. 516. — SYKES; Fauna Hawaiiensis,
p. 319. — THWING, Reprint Orig. Descript. Achat, pi. 2, f. 6. —
Partulina tessellata Newc., BORCHERDING, Zoologica, p. 52, pi.
2, figs. 1 to 16, 1906.

Out of 57 specimens before me, 53 are sinistral. The figure
is stouter than in the following species, and it differs from
virgulata constantly by the obliquely striped last whorl of the
embryo. The first 1 to 1J whorls are some shade of brown;
then broad, slightly retractive stripes of opaque white and dark
brown alternate to the end of the embryonic shell. A clouded
or zigzag-mottled pattern ensues on the first neanic whorl. The
umbilicus is narrowly perforate, and the surface has little gloss
or none, the dense, wavy, spiral lineolation being well developed.

Length 26, diam. 17.3, aperture 14 mm.

Length 23, diam. 15.5, aperture 12 mm.

The typical form probably came from Makakupaia (pi. 6 figs.
16, 18, and fig. 12 received from Newcomb). The ground-color
of the later whorls is whitish, more or less streaked with fleshy,
with a varying arrangement of dark spiral bands and lines; lip
light liver-brown. Rarely the bands are absent, the later whorls
being profusely streaked (pi. 6, fig. 17).

At Kahanui most specimens have the post-embryonic whorls
uniform reddish-chocolate, by coalescence of the bands, or they
are banded with that color (figs. 13, 14, 15, 20), but some

PARTULINA, MOLOKAI. 29

specimens resemble the typical form in pattern. They occur
both sinistral and dextral, the latter rare.

From Kealia Borcherding figures shells with very weak or light
chestnut bands (fig. 19).

In the C. M. Cooke collection there is a set of nearly white
shells (fig. 21).

9a. P. T. MEYERI Borcherding. PL 7, figs. 1, 2.

'"Shell narrowly umbilicate, dextral, rather solid ovate-
conic, under a lens decussated regularly and very finely, glossy;
pale gray or grayish-brown, ornamented with light and darker
brown, or with gray and blackish streaks; spire conic, the apex
rather obtuse; suture crenulated, more or less margined, whorls
6, convex, the last \ the total length; upper whorls tessellated
with reddish-gray and yellowish-gray, the last w^horl rounded,
uncolored, or under the middle having narrow horn-colored
bands here and there. Aperture inversely ear-shaped, glossy
bluish within; peristome narrowly expanding and thickened,
with a reddish-brown internal lip. Columella twisted, calloused;
columellar margin reflexed, nearly covering the umbilicus.
Length 22 to 24, diam. 13, aperture 12 X 8 mm." (Borch-
erding. )

Molokai: Pelekunu (Meyer).

Partulina meyeri BORCHERDING, Nachbl. d. d. Malak. Ges.
1901, p. 55; Zoologica xix, p. 54, pi. 2, f. 17, 18.

Found in Pelekunu valley, where a large, dextral form of
tessellata reminding one of virgulata also occurs.

10. P. RUFA Newcomb. PL 7, figs. 3 to 11.

"Shell sinistral, conically ovate, solid, with decussating
strise; ground color externally and internally brownish red,
covered with an epidemis of a mottled brown and white, the
latter arranged on the central whorls in fine zigzag markings,
which are lost on the last whorl in a uniform grayish umber;
lower half of this whorl encircled by a broad white band,
whorls 6, flatly convex; suture plain, moderately impressed.
Columella strongly callous ; umbilicus open, small. Lip ex-
panded, reflected below. Length 0.9, breadth 0.5 inch'
(Newcomb).

30 PARTULINA, MOLOKAI.

Molokai (Newcomb): Kalae, Kaweeku, Kalamaula, Kahanui
and Makakupaia (Meyer), in the central part of the island.

Achatinella rufa NEWC., Ann. Lye. N. H. of N. Y. vi, p. 21,
May 1853; P. Z. S. 1853, p. 130, pi. 22, fig. 3; Ann. Lye. vi,
p. 324 (description of animal). — PFR., Monographia iv, 537. —
SYKES, Fauna Hawaiiensis p. 318. — Partulina rufa Nc.,
BORCHERDING, Zoologica xix, p. 54, pi. 3, f. 1-9. — Partulina
ides BORCHERDING, Nachrbl. d. d. Malak. Ges. 1901, p. 52;
Zoologica p. 56.

(1) Typical P. rufa (pi. 7, fig. 3) is a dull shell with the
growth-lines usually strongly marked, unequal, crossed by low,
usually wide and more or less granulose spirals mingled with
finer waved strife, the latter often predominating, especially be-
low the periphery. The sculpture is extremely variable, the
granulose spirals being very strong in some forms, such as that
described as idae, and in others much reduced, the finer decur-
rent striation then dominating. The summit is yellowish-
brown; last 1^ or 1 embryonic whorls are marked with protractive
yellowish-brown and white stripes; following neanic whorls have
a dense, confused zigzag pattern, often indistinct or illegible.
This may continue on the upper half of the last whorl, or it
may be replaced by a uniform or clouded tawny color, inter-
rupted by a white or yellowish belt just below the periphery.
The columellar fold is strong, calloused, brown tinted, the lip
of similar tint ; interior pink or dull brownish- violaceous.
Length 24, diam. 13.5, aperture 11.7 mm.

The original localit}^ on Molokai is uncertain. Borcherding
figures various forms from localities in the mountains south of
the northern peninsula, none of them just like the typical form,
represented by specimens before me received from Newcomb.
Borcherding tentatively suggests the view that P. rufa is "not
a pure species, but a bastard-form between P. tessellata Newc.
on the one side, and P. proximo, Pse. on the other.' While
this seems rather doubtful, and probably could be proved only
by breeding experiments, rvfa certainly unites characters of the
two species. The embryo has protractive stripes as in P.
proxima. Some very rare color-forms closely resemble proximo
in the later stages; but it differs from that species by the dull

PAETULINA, MOLOKAI. 31

surface, deeply sculptured spirally; proxima being glossy, with
weak spirals and very slight trace of the minute, wavy, spiral
lineolation of the dull Partulinas. PI. 7, figs. 6, 7 represent
two forms of rufa from Kaweeku, copied from Borcherding.

(2) There is a dull brown (reddish to olive brown) form, with
a few scattered light spots, much as in a common form of /'.
warmorata (pi. 7, fig. 4). Sometimes this form has a suit-
peripheral white zone (fig. 5); the sculpture either as in viir.
idie or without major spirals, clothed throughout with minute,
crinkled, spirally descending striae.

(3) Another form has zebra-stripes of white and brown, ex-
actly as in one form of proximo,, the surface having fine deeply
cut Partulina sculpture. Length 28.5, diam. 16.5, aperture 15
mm.; whorls 7 (fig. 8).

(4) Similar to var. ida, except that the spiral engraving is
close, nearly even, without granules. Last W7horl of the embryo
having very obliquely protractive brown stripes; following
whorls pale yellow, darker towards the base, or showing some
faint tawny stripes on the spire. Aperture pinkish white, colu-
mella and narrow lip pale flesh-pink. PI. 7, fig. 9, 10, 11.

Length 23, diam. 14.2, aperture 12 mm.
Length 17, diam. 10.2, aperture 8.2 mm.

(5) Partulina rufa var. idee Borcherding (pi. 7, figs. 13, 14,
15), from Kalae and Kealia, may easily be recognized, accord-
ing to Borcherding, by its compressed and ventricose shape and
the regularly granulose spiral sculpture (fig. 13). The last
whorl is more or less distinctly uniform yellowish-brown, or has
a yellowish-white belt below the middle, the upper whorls
being zigzag marked. In a series of 40 specimens before me I
find the characters so variable that I cannot look upon P. idee as
anything but a local race. The sculpture in particular inter-
grades perfectly with typical P. rufa.

In many specimens of idte the color above the periphery is
pale or even whitish, and on the base it becomes yellow, rich
brownish-yellow or olive, the change being gradual in some,
abrupt in other examples. In many of the pale forms the
stripes of the last embryonic whorl are faint or even not dis-
cernible.

32 PARTULINA, MOLOKAI.

11. P. PROXIMA (Pease). PL 9, figs. 1 to 9.

Shell sinistral, im perforate, oblong-ovate, ventricose: rather
solid; finely striated transversel}7, striae somewhat flexuous,
granulose and interrupted, more regular and conspicuous be-
neath the sutures. Whorls 6, convex, marginated, the last
being somewhat produced obliquely and flattened on the middle,
so as to give in some specimens a subangulated appearance at
the base. Aperture somewhat oblique, of an oblong-ovate
form; columellar fold strong. Color chestnut-brown, striped
and mottled irregularly with darker brown and white; columella
and inner edge of aperture purplish red (Pease}.

Molokai (Pse. ): Maunahui, Kahanui, Waikolu, Makakupaia,
Pelekunu, Makolelau and Kamalo (Meyer).

Helicter proximus PSE. , P. Z. S , 1862, p. 6. — Achatinella prox-
imo, Pse., PFR. , Monographia, vi, 1868, p. 168; viii, p. 217. —
HARTMAN, Proc. Acad. Nat. Sci., Phila., 1888, p. 27, pi. 1, f.
1, 2. — Partulina proxima Pse., BORCHERDING, Zoologica, xix, p.
63, pi. 4, f. 1-12.

"The above species from the island of Molokai appears to
represent the H. marmoratus and its varieties of the island of
Maui. It is, however, larger and heavier; the last whorl has
also a peculiar shape, in common with that of several of the
larger species found on Molokai. All the specimens I have
seen are sinistral, and the columella and edge of the aperture of
a deep red ' (Pease).

The first 1^ whorls are white; then very obliquely protract! ve
brown stripes appear, continuing to the end of the embryonic
stage, which comprises 3J whorls. The succeeding neanic stage
has stripes in general parallel to the growth-lines, but more or
less interrupted or irregular. This pattern may continue to and
upon the last whorl (figs. 1, 5) ; or the stripes may become
nebulous or reduced on the last two whorls. The ground-color
varies from glossy white to strongly brown tinted, or the last
whorl may be brown tinted towards the base, white above. In
some shells the stripes are very obliquely retractive on the later
whorls (fig. 2).

In another variety the stripes are tawny yellow on a white or
slightly coffee-tinted ground. The sutural margination is dis-

PARTULINA, MOLOKAI. 33

tinct (figs. 7, 8). According to Borcherding, this pattern comes
from Pelekunu.

Another form has close hair-like lines, of brown, cream, and
pale lilac, in harmony with growth-lines, over the whole post-
embryonic shell. It is glossy, with typical embryonic colora-
tion (fig. 9). Borcherding figures a shell of this pattern from
Makakupaia.

In fully adult shells the outer and basal lips expand slightly.
The lip and columella are brown in nearly all specimens, but
in some light ones it is a vinous brown. The interior varies
from nearly white to lilac or violaceous of various tints. The
surface is glossy, and the spiral strife weak or subobsolete.

Length 26.5, diam. 15.5, aperture 14 mm.; 6f whorls.

Length 24, diam. 14, aperture 13 mm. ; 6^ whorls.

Color- var. schauinslandi Borcherding. PL 9, figs. 10, 11.

The shell is very glossy, white or yellowish white, uniform
or with faint, pale buff lines along growth lines on the last one
or two whorls. Interior white or pink, the narrow lip roseate-
brownish or nearly white, collumellar fold fleshy brown. The
suture has a distinct narrow margin defined by an impressed line.
Length 24, diam. 13, aperture 12 mm. ; 6J whorls.

Young shells are sometimes white above, yellow below the
angular periphery. In some specimens there are faint stripes
on the last embryonic whorl. These are transitions from
schauinslandi to the paler forms of proximo, such as that from
Pelekunu valley, clearly showing its genesis from the latter,
the only difference being loss of color. This has been fully set
forth by Herr Borcherding, and is well shown by the large
series I have examined.

Molokai: Kaluahauoni and Waileia (Meyer).

12. P. THEODOREI (Baldwin). PI. 9, figs. 14, 15.

Shell sinistral, minutely perforate, rather thin, narrowly
pyramidal, glossy, with sculpture of rather rude growth lines
and minute, rather weakly incised, spiral stria?. Embryonic
whorls whitish, the last one usually marked with broad, protrac-
tive brown stripes. Following whorls have irregular purple-

34 PARTULINA, MOLCKAI.

brown flames or streaks on a white ground, in general harmony
with growth-lines, widening downwards, scarcely reaching to the
suture above. On the last whorl the streaks are often some-
what dislocated. Whorls nearly 7, but slightly convex, the
suture rather indistinctly margined below. Aperture small,
lilac within; lip narrow, not expanded or noticeably thickened,
brown-bordered within, columellar fold very low, brown.

Length 18.6, diam. 8.8, aperture 7.7 mm.

Length 18, diam. 8, aperture 7.3 mm.

Molokai: Kawela (Baldwin).

AchatineUa theodorii BALDWIN, Catalogue etc., 1893, p. 7,
(name only). — AchatineUa theodorei BALDWIN, Proc. A. N. S.,
Phila., 1895, p. 226, pi. 10, f. 27.— SYKES, Fauna Ha-
waiiensis p. 330. — Partulina theodorei Bald., BORCHERDING,
Zoologica xix, p. 65, pi. 4, f. 14, and f. 15, var.

Figured and described from the types. While closely related
to Partulina proximo, by the sculpture and coloration, this species
is constantly much narrower, in a considerable series seen.
The columellar fold is also less developed. P. proximo, does
not occur in Kawela valley, the type-locality of theodorei.

Mr. Sykes placed this species in Perdicella, a position ap-
proved by Professor Hyatt. Herr Borcherding groups it with
proxima Pse., in Partulina. It is one of several species connect-
ing Perdicella with Partulina.

Var. midtistrigata n.v. (pi. 9, figs. 12, 13, 16) differs by its
broader shape and much more numerous, narrow stripes. The
types are without exact locality but Borcherding figures a sim-
ilar shell from Makakupaia. Length 20.5, diam. 11, aperture
9.8 mm.

13. P. MUCIDA (Baldwin). PL 9. fig. 17.

Shell sinistral, minutely perforate, solid, ovate-pyramidal,
the spire straight-sided, ra.ther acute at apex. First 2 to 4
whorls tawny brown, the last two or three whorls slightly con-
vex, whitish, mottled and banded with flesh color, and having
purple brown bands speckled or mottled with white above and
below the suture, at the periphery and around the axial perfor-
ation. The surface is glossy, with low, rude growth-lines and

PARTULINA, MOLOKAI. 35

faint spiral striation. Aperture small, bluish white within, lip
slightly expanded, nearly white and well thickened within.
Columellar lamella strong, white with a light brown patch at
and above its insertion. Length 21 to 21.2, diam. 11.8, aper-
ture 9.6 to 10 mm. ; 6f to 7 whorls.

Molokai: Makakupaia (Baldwin).

Achatinella mucida BALDWIN, Proc. A. N. S., Phila., 1895, p.
222, pi. 10, f. 23. — SYKES, Fauna, p. 315. — Partulina muc/d«
BORCHERDING, Zoologica, xix, p. 73, pi. 6, f. 7, 8.

This small species, described and figured from the types, has
a peculiar mildewed or mouldy appearance from having the
dark markings overlaid with white. It is very closely related
to A. dwightiij but differs by its smaller size and details of col-
oration. No form of dwightii has a dark band along the suture
below. The figure represents the same shell figured by Bald-
win.

13a. Color-var. macrodon Borcherding. PI. 9, figs. 18, 19, 20.

Last embryonic whorl zigzag striped; following whorls white
with a faint flesh tint, having distinct purple-brown bands,
minutely flecked with white, at suture, periphery and umbili-
cus. Columellar lamella strongly calloused, projecting hori-
zontally. Length 20 to 22, diam. 11.5 to 12, aperture 9.5 to
10 mm. ; whorls 6f to 7.

Makakupaia (Meyer and others).

Partulina macrodon BORCHERDING, Nachrbl. d. d. Malak. Ges. ,
1901, p. 56; Zoologica, xix, p. 74, pi. 6, f. 9, 10.

14. P. DWIGHTI (Newcomb). PL 8, fig. 1 to 5.

"Shell oblong conical, sinistral, solid, surface covered with
longitudinal strire, cut across by very numerous, fine, slightly
undulating cross-stria?. Whorls 6, flatly convex, crenulated at
the sutures; suture simple, well marked. Aperture ovate; colu-
mella broad, short, and slightly twisted; outer lip expanded,
subreflected below. Subumbilicated, color a grayish white,
with numerous blotches and zigzag markings of brown, more
obscure on the last whorl. Aperture and lip of a dingy white.
Length 19, width 9^ twentieths of an inch [about 24 X 12 mm.]
(Newc. )

36 PARTULINA, MOLOKAI.

Molokai (Newcomb): Makakupaia, Kawela, (fig. 2), Mako-
lelau, Kamalo and Punkaeha, fig. 1, (Meyer); Kamalo
(Baldwin).

Achatinella dwightii NEWCOMB, Ann. Lye. N. H. of N. Y. vi,
p. 145, October, 1855; P. Z. S. 1855, p. 207; American Journal
of Conchology ii, p. 213, pi. 19. f. 9. — PFR., Monogr., iv, p.
517. — SYKES, Fauna Haw. p. 313.- -TnwiNG, Reprint Orig.
Descr. A chat. pi. 2. f. 7. — Partulina d. , BORCHERDISG, Zoologica
xix, p. 68, pi. 5, f. 1-8.

P. dwightii is characterized by the solidity, narrow form with
a high, straight-sided, acute spire, and the streaked, mildewed
appearance. Typically the last embryonic whorl has protractive
stripes and the following whorls of the spire show fleshy-brown
mottling or confused zigzag patterns on a pinkish or yellowish-
white ground. Last whorl streaked, usually paler below the
suture. The surface has rather rude, irregular but low growth -
lines, and on the last whorl there are regular, distinct, waved
spiral stria3 in shells from Newcomb (fig. 5). In most other
lots seen the spirals are quite weak or faint. The aperture is
small, and the rib is placed rather far within, so that there is an
unusually wide whitish lip. The outer edge often expands
noticeably. In fully adult shells the columellar lamella is
strong and white or nearly so. Some specimens are white with
only faint brown streaks and stains, the aperture delicately
lilac-tinted, lip white. In others the brown streaks are dark,
in part blackish, and nearly cover the last whorl.

Length 26, diam. 15, aperture 13 mm.

Length 25, diam. 13.5, aperture 12 mm.

Length 22, diam. 13, aperture 10.8 mm.

The shape is more straightly pyramidal than in A. redfieldi,
which also differs by its brown columellar fold and the narrower
and brown lip. Its range of patterns is also different.

14a. P. dwightii compta (Pease). PI. 8, figs. 6, 7, 8.

Shell sinistral, narrowly perforate, rather solid, oblong-conic;
spire obsoletely transversely delicately rugose-striate, the
wrinkles disappearing on the last whorl, and longitudinally im-
pressed with delicate, irregular striae. Spire turrited or oblong-

PARTULINA, MOLOKAI. 37

conic; suture impressed, whorls 6, convex, sometimes sub-
angular above, the last equal to half the length of the shell.
Aperture vertical, ovate; columellar fold superior, strong, white;
lip slightly expanded, callous within; leaden-gray, encircled
with lines or bands of reddish-brown, the last whorl ornamented
with a white band at the base. Length 25, diam. 13 mm.
(Pease).

Molokai (Pease); Kawela and Makakupaia (Meyer).

Partulina compta PEASE, Journ. de Conchyl. xvii, 1869, p.
175; P. Z. S. 1869, p. 647. — BORCHERDING, Zoologica xix, p.
57, pi. 3, f. 14-16, 18-20.

"This species, which comes from the island of Molokai, is
very closely related to P. splendida Newc., which lives on Maui.
It is more contracted at the base and sometimes has a peculiar
angulation of the upper part of the whorls. In coloration it is
nearer P. grisea Newc." (Pse.).

The ground-color is white or nearly so in the specimens fig-
ured by Borcherding and before me in the Academy, Cooke and
University of Wisconsin collections. Otherwise they agree well
with Pease's description. It would probably be more correct
to describe the shell as brown, pale or dark, cut by white bands
and lines. It differs from A. redfieldi by its white coiumella
and lip. The following race is almost fully connected with
compta by intermediate specimens, and may perhaps be found
to be superfluous. I retain it here because both Borcherding
and Hyatt have recognized it, though under different names.

146. P. dwightii concomitans Hyatt, n. var. PI. 8, figs. 9 to 13.

Similar to dwightii in the acutely pyramidal contour, solidity,
wide white lip, etc. ; but it differs by the predominance of chest-
nut-brown (rarely olive-brown) which forms nearly a solid
color on the last two whorls except that there is a subperipheral
white belt and sometimes a white line on the upper surface,
continuing on the penultimate whorl; the suture has a narrow
white edge.

Molokai (A. N. S. P. and Cooke collections): Makakupaia
(figs. 9, 10, 12), Kawela (fig. 11) and Makolelau (Meyer).

Partulina grisea Nc., BORCHERDING, Zoologica xix, p. 69, pi.

38 PARTULLXA, MOLOKAI.

5, f. 9-16. Not Achatinella grisea Newcomb. — ? Achatinella
redfieldi var. </, SYKES, Fauna Hawaiiensis, p. 318, pi. 11, f. 16
(Makakupaia and Kamalo, Perkins).

Borcherding figures a long series of this form, varying from
pale brown and gra}T to deep chestnut, all having a subperi-
pheral white belt. I have not seen gray examples and copy his
figure of one (pi. 8, fig. 9). Fig. 13 is the type, no. 106061,
A. N. S. P.

While dwighti and concomitans stand near grisea Nc. , they
differ from it decidedly by the pattern of the early whorls and
the stronger columellar lamella. The shell is also larger and
more solid than that of grisea, which is certainly distinct. We
cannot agree with Herr Borcherding that P. grisea occurs on
Molokai.

15. P. REDFIELDII (Newcomb.) PL 7, fig. 12, 16, 17; pi. 8,
figs. 14 to 20.

"Shell sinistral, elongate, conical, striated longitudinally,
color light fawn passing into deep chestnut, paler above, plain
or with transverse chestnut bands with obscure undulation upon
the third whorl only; a white band also traverses the suture,
whorls 6; suture well impressed, slight!}7 margined. Aperture
subovate; columella brown, flat and twisted; lip slightly re-
flected, the color of the columella. Length 1, breadth 45 inch
(New., 1853).

Animal light flesh color, mantle dark slate (Newc., 1854).

" Animal as long as the shell, grayish above, sprinkled slightly
with brown, tentacles of same color, mantle slate, bottom of
foot of a greenish gray ' (Newc. , 1858).

Molokai: Mapulehu (Baldwin); Ualapue (Thaanum); Maka-
kupaia and above Kamalo (Perkins); Kamoku, Kawela, Mako-
lelau, Kaluaaha and Ualapue (Meyer).

Achatinella redfieldii NEWC., Ann. Lye. N. H. of N. Y. vi, p.
22, April, 1853; p. 325; P. Z. S. 1853, p. 131, pi. 22, f. 5;.
1854, p. 311. — PFR., Monogr. iv, 523. — SYKES, Fauna Hawaii-
ensis p. 317, pi. 11, f. 15. — Partulina redfieldii Newc., BOR-
CHERDING, Zoologica xix, p. 71, pi. 6, f. 1-6.

The outlines of the spire are less straightly pyramidal than in

PARTULINA, MOLOKAI. 39

P. dwightii, which is usually more solid, often larger, with a
white columella and lip. Newcomb's description of the color-
pattern is not quite clear, owing to the failure to place a comma
after the word "bands' in his original description, reprinted
above. This was rectified by him in the P. Z. S., where in
both Latin and English versions he makes it plain that it is
only the "undulations' (or axial stripes) which are restricted
to the third whorl. Sixteen specimens in coll. A. N. S. P., in
two lots received from Newcomb, agree well with his account.
Some of them are figured on plate 8, figs. 15 to 18. It will be
seen that the color-pattern varies widely. In some specimens
the last two whorls are rich brown, uniform or varied with
spiral bands and lines, the spire paler above, suture white-
edged, lip and columellar fold brown. Other shells have a
light cream ground on which there are few or many brown
spiral lines and bands, or in place of them there may be dark
narrow streaks. The last embryonic whorl may have oblique
or zigzag stripes, but they are often lacking (pi. 8, figs. 15-18,
from shells received from Newcomb). Newcomb's type figure
is copied, pi. 8, fig. 14.

Length 25, diam. 14, aperture 13 mm.

Length 21, diam. 13, aperture 10.5 mm.; 6i whorls.

Length 24.5, diam. 13, aperture 11.3 mm.; 6J whorls.

There is a continuous intergradation between the dark brown,
rich orange-brown and longitudinally lineolate forms, all hav-
ing a snowy and usually rather broad subsutural band. Often
the tawny, longitudinally lineolate form has white or pale
brown spiral bands (figs. 15, 16), producing, when the bands
become numerous, the pattern shown in Newcomb's figure
(copied in pi. 8, fig. 14), which I take to be the typical pattern,
though Mr. Sykes has figured one of the dark, bandless shells
as typical. It is no great step from some shells of this pattern
to P. dwightii concomitans.

In the Cooke collection there are white specimens with the
lip very pale brown, apical whorls fleshy (pi. 7, fig. 17), with
others showing traces of the brown bands on the base and latter
part of the last whorl. A peculiar shell has narrow brown lines
near suture and on periphery, and a patch at the axis, on a

40 PARTULINA, LANAI.

pale brownish ground, the spire white, becoming fleshy above
(pi. 7, fig. 12).

Another form (no. 2068, Cooke coll. ) has a snow-white spire
and coffee-colored last whorl, aperture pink within, lip brown
(pi. 7, fig. 16).

In some examples there is a sutural margin banded by an
impressed line, as in P. proximo, schauinslandi. This is probably
a local race.

The color-forms of A. redfteldii may be arranged thus:
A. — Uniform dark or light brown with a white sutural border.
Ab. — The same, with darker spiral bands.

a. — Yellowish-brown with narrow streaks of darker

brown along growth lines.

ab. — The same cut by white or whitish spiral bands.
b. — White predominating, leaving only traces of dark

bands or none.

To what extent these patterns mingle in the same colony is
not known to me.

SPECIES OF LANAI.
16. P. CRASSA (Newcomb). PI. 18, figs. 1, 2.

The shell is usually dextral, narrowly umbilicate, acutely
ovate-conic, solid, chestnut (the shade varying) with zigzag
lines, dots and streaks of a whitish tint, and a white zone below
the periphery; spire typically a little attenuated above, the whorls
scarcely convex. Surface dull or slightly glossy, closely and
rather deeply engraved with spiral lines. Suture scarcely im-
pressed. Aperture ovate, the lip expanded, sometimes almost
reflected, white, very strongly thickened within, the callous
usually irregular. Columellar lamella obtuse but rather strong.
Columellar lip built forward, reflected.

Length 20, diam. 14 mm.; 6J whorls (typical).

Length 18, diam. 13 mm.

Length 19, diam. 12 mm.

Length 22, diam. 14 mm.

Lanai (Newcomb); near Koela (Perkins).

Achatinetta crassa NEWC., P. Z. S. 1853, p. 155, pi. 24, f. 71

PARTULINA, MAUL 41

(1854). — PFR., Monogr. iv, 527. — SYKES, Fauna Ifawaiiensis
p. 312.--THWING, Reprint Orig. Descript. Achat. pi. 2, 1 10.

This very distinct species is more closely related to P. dwightii
concomitans of Molokai than to any other existing form. In the
series of 70 specimens in the collection of the Academy only one
is sinistral. There is a good deal of variation in the extent of
light markings, but the subperipheral band seems rather con-
stant, being absent in only one example. The dark color varies
from chestnut to chocolate, and is more or less streaked with
lighter and darker. One specimen (pi. 18, fig. 2) has the
whole post-embryonic shell banded as in P. tessellata — a notable
advance in pattern, foreshadowed by a few others having several
basal bands.

The embryonic shell, 5 mm. long, has the first two whorls
light fleshy brown, whitish below the suture; some broad, indis-
tinct darker clouds then appear, changing on the last whorl to
zigzag axial stripes of dark brown and cream- white ground.
The axis is perforate, columella a little convex (pi. 18, figs. 3, 4).

SPECIES OF MAUL

The numerous species and varieties fall into three series: that
of P. marmorata, stout, ovate forms with streaked or striped
pattern; series of P. splendida, thinner shells with profuse spiral
bands and lines; and the series of P. tappaniana, mainly light
colored, rather narrow forms, uniform, or with spiral bands on
a white ground.

Series of P. marnwrata.

17. P. KAAEANA Baldwin. PI. 10, figs. 5, 6, 7.

"Shell sinistral, subperforated, solid, globose with a conical
spire, apex subacute; surface rather lusterless, covered with
rather coarse wavy growth lines, and under a lens exhibiting
close and delicate decussating spiral lines; nuclear whorls faintly
decussated. Color ashy brown, with a light brown band just
below the periphery which enters the aperture; whole surface
covered with minute longitudinal white flecks or streaks; apex
tessellated white and brown. Whorls 6, not margined above,
flatly convex; suture lightly impressed. Aperture a little

42 PARTULINA, MAUI.

oblique, oval, livid white, showing the outside coloring within.
Peristome acute, slightly thickened within, expanded, colu-
mellar margin reflexed, light brown on both face and the re-
verse. Columella white, terminating in a strong, plaited, pro-
jecting tooth.' (Baldwin1).

Length 21, diam. 13.8 mm.; whorls 6^.

Length 19.8, diam. 13 mm.

"Animal extended in motion longer than the shell. Mantle
brownish-black with outer edge bordered with a narrow white
line. Foot below and sides light slate color. Head above and
tentacles dark slate and granulated' (Baldwin).

West Maui; Mt. Helu, 4000 ft. elevation.

Partulina kaaeana BALDWIN, Nautilus xix, p. 113, February,
1906.

A dull brown shell, yellows-banded below the periphery,
speckled with whitish flecks, or with the last whorl having only
some indistinct flecks and streaks, dark on each side of the light
band. Another specimen is reddish brown writh many irregular
streaks and spots of white, but without the subperipheral light
belt. The last embryonic whorl is striped white and brown,
as usual. While related to marmorata, this is a quite distinct
species. Figured from the type lot.

"This shell was found on a mountain peak quite isolated
from the main mountain mass of West Maui. We dedicate it
to Mr. W. F. Kaae who seems to have been the only one in
quest of shells who has ventured to climb this lonely peak. He
found the shell in company with Perdicetta ornata Nc., a species
supposed long since to have become extinct,'1 (Baldwin').

18. P. MARMORATA (Gould). PL 10, figs. 1, 2, 3, 4.

"Shell dextral, rather solid, of an elongated, acutely conical
form, composed of 6 convex whorls which are somewhat
shouldered superiorly; the last large in proportion, and ven-
tricose. Surface delicately striated with lines of growth. Color
a dusky chestnut, marbled with white irregular and angular
markings, generally arranged somewhat in longitudinal stripes,
the whole seeming to be coated with a very thin, ash-colored
epidermis. Aperture moderately large, ovate, the lip slightly

PARTULINA, MAUI. 43

expanded; the columellar fold white, compressed, nearly trans-
verse, and standing out very prominently. Length f , breadth
Mnch" (Old.)

East Maui: foot of Mt. Haleakala (U. S. Exped.); Makawao
(Baldwin); Kula (Gulick).

Achatinella marmorata GLD., Proc. Bost. Soc. N. H. ii, p.
200, March, 1847; Expedition shells p. 35; U. S. Exploring
Exped., Mollusca, p. 85, pi. 7, figs. 94, 94a.--PFR., Monogr.
iii, p. 455. — NEWCOMB, Ann. Lye. N. H. of N. Y. vi, p. 311.
-TnwiNG, Reprint Orig. Descript. Achat. pi. 2, f. 13. — Achati-
nella adamsi NEWCOMB, Ann. Lye. vi, p. 19 (May, 1853).
— A. adamsii NEWCOMB, P. Z. S. 1853, p. 137, pi. 22, f. 20.

Perforate, dextral, rather solid, gray-brown, with scattered
white spots or irregular stripes dark-bordered on the left side.
Last embryonic W7horl having wide, strongly protractive white
and brown stripes. Surface dull, without gloss, by reason of
the dense, microscopic, crinkled spiral striae. Gould's figure is
copied, fig. 1.

Length 25, diam. 14.3, aperture 13 mm.; 6^ whorls.

Length 21.2, diam. 13.3, aperture 11.3 mm.; 6 whorls.

Length 21, diam. 12.2, aperture 11 mm.; 6-J whorls.

Newcomb described the animal as "light pea green, strongly
granulated, one-third longer than the shell, tentacles light slate;
mantle and base of foot same color as the body. ' He included
as synomous: A. adamsi Newc., A. induta Gul., A. plumbea
Gul., A. ustulata Gul. Mr. Thwing states that "in over one
thousand examples only one sinistral specimen was found.'

liA. adamsii. — Shell dextral, conically acuminate, whorls 6,
finely striated transversely, roughly striated longitudinally;
suture deeply impressed, lip expanded, columella terminating
in a strong, abruptly twisted plait. Interior of shell of a leaden
blue color, exterior dark chestnut, irregularly mottled and
striped with white. Length 18, diam. 9-twentieths of an inch.'
A. adamsii is terrestrial in its habits.

19. P. PLUMBEA (Gulick). PI. 10, figs. 8 to 12.

"Shell dextral, sometimes perforate, ovate-conic, solid, shin-
ing, irregularly striated, very finely decussated; lead-colored

44 PARTULINA, MAUL

with a white band on the periphery of the last whorl; apex sub-
acute; spire conical; suture marginate, well impressed. Whorls
6-J, very convex. Columellar fold central, white, strong. Aper-
ture slightly oblique, sinuately oval, bluish- white within; peri-
stome brown, thickened within, with dextral margin arcuate,
scarcely reflected; columellar margin dilated, white, sometimes
slightly detached; parietal margin wanting. Length 23, breadth
13 mm.; length of body-whorl 16J mm. Average weight 6
grains; least wr eight 4.3 grains.

Var. b. Globose, conic, spire shortened, with concave out-
lines, body-whorl inflated.

"Var. c. Without the white band on the body-whorl'

East Maui : Kula, on the trunks of the aiea and kukui.
(E. Bailey).

Achatinella plwnbea GUL., Ann. Lye. N. H. of N. Y.? iv, p.
213, pi. 7, f. 39, 1856.

Very closely related to mannorata, perhaps not always dis-
tinguishable, but polished, typically belted with white, the
band rarely wanting, and without the special marking of mar-
morata. The typical form from Gulick (pi. 10, figs. 9, 10) is
usually speckled on the spire with white on a fawn-brown
ground. A lot from Baldwin, also from Kula (pi. 10, figs. 11,
12) is drab with darker and lighter streaks, the shell smaller.
Another lot from Baldwin, locality not stated, is fawn with

*/ j

narrow brown streaking, with or without a belt, sometimes with
some scattered whitish dots (fig. 8).

Large specimens from Gulick measure 25 mm. long, 14.5 wide.
The smaller drab form is 22.5 x 12 mm.

20. P. WINNIEI Baldwin. PI. 13, figs. 1, 2, 3.

Shell sinistral, subperforate, rather thin but moderately
strong, long-conic, somewhat glossy. First 1| or 2 whorls of
the embryonic shell flesh-colored, next half whorl boldly
marked with broad brown and white axial stripes. Following
neanic and adult whorls cream-white, copiously striped with
dark chestnut, the stripes more or less irregular and unequal.
Growth-lines and spiral sculpture rather weak. A sutural mar-

PARTULINA, MAUI. 45

gin is defined by an impressed line. Aperture flesh-colored
within, the lip narrowly thickened, a trifle expanded, white.
Columellar fold rather strong, white and oblique.

Length 16.5, diam. 8, aperture 7.8 mm.; whorls 6J.

Length 15, diam. 7.8 mm.; whorls 6.

Length 16, diam. 8 mm.

West Maui: Kahakuloa.

Partulina winniei BALDWIN, Nautilus, xxii, p. 67, November,
1908.

"Named in honor of Miss Winnie of Wailuku, Maui.'
Figured and described from cotypes sent by Mr. Baldwin, who
states that it is the counterpart of Partulina theodorei Bald, of
Molokai. There is a strong resemblance, but I am inclined to
think winniei a diminutive relative of P. perdix. It approaches
Perdicella in structure of the columella.

21. P. PERDIX (Reeve). PL 10, figs. 14, 15, 16; pi. 13, figs.
10 to 13.

"Shell ovate, ventricose, rather thin, dextral, spire exserted,
somewhat obtuse at the apex, whorls 6, convex, margined
around the upper part, obliquely sculptured with flexuous im-
pressed lines, columella plicately twisted, rather callous,
whitish, painted and varigated throughout with olive-brown,
sutures and columella white' (Reeve).

W. Maui: Lahaina (Baldwin, here selected as type loc. );
Olinda at 4000 ft. (Perkins); Honokowai (D. Thaanum).

Achatinella perdix REEVE, Conch. Icon, vi, pi. 6, f. 43 a, b
(May, 1850). — SYKES, Fauna Hawaiiensis p. 315. --BALDWIN,
Catalogue p. 7.--THWING, Reprint Orig. Descript. Achat. pi.
2, f. 20.

Achatinella undosa GULICK, Ann. Lye. N. H. of N. Y., vi
p. 20/>. pi. 7, f. 33, 1856.

The apex (1 to 1^ whorls) is brownish white; the next one
or 1 J wrhorls have broad protractive alternating stripes of snow-
wrhite and dark brown. The neanic whorls following have
narrow stripes of the same colors, running with the growth-
lines. This pattern may continue upon the last whorl, or the
stripes may become wider and bisinuate or angular there, with

46 PARTULINA, MAUL

a streak of darker brown on the left side of each white stripe.
In some shells the white is reduced to irregular spots on the last
whorl. These patterns are figured. The surface is rather
glossy, showing fine spirals under the lens; suture margined
with a white cord. Axis imperforate. Length 22, diam. 12^,
aperture 10 mm. ; 6f whorls.

The shade of brown varies a good deal in different lots.
Lahaina specimens are figured, pi. 10, figs. 14 to 16, and pi.
13, fig. 13. In pi. 13, figs. 10, 11, 12, we figure three sinistral
specimens from Honokowai, collected by Mr. Thaanum.

Var. undosa Gulick (pi. 10, figs. 17, 18, 19). Usually more
slender than perdix, the spire longer, stripes generally narrow,
sometimes pale and linear; distinctly perforate.

Length 22, diam. 11.8, aperture 10.6 mm.; 6J whorls.

Length 20, diam. 11.9, aperture 10.9 mm.

Length 21, diam. 11, aperture 9.6 mm.

Mountain ridges of Waihee, on the ilima (Sida.}, ki (Cordy-
II lie terminalis) and other low bushes (E. Bailey).

21a. P. perdix pyramidalis (Gulick). PL 10, fig. 20.

"Shell dextral, very rarely sinistral, imperforate, pyramidal,
rather solid, shining; chestnut or ash brown, with a white or
yellowish band encircling the base, with oblique white and
brown markings on the second and third wrhorls, bluish-white
within the aperture. Apex subacute; spire regularly conical,
sometimes decollated. Whorls 6J-, convex, margined with white;
finely striated transversely, and microscopically decussated with
faintly impressed wrinkled spiral lines; the last regularly
rounded, equal to 65 hundredths of the length. Columellar fold
well developed, white, oblique. Aperture rather oblique, semi-
oval; peristome thickened within, very slightly reflected; with
columellar margin dilated, adnate or slightly detached; parietal
margin wanting.

"Length 21.6, breadth 11.4 mm., aperture 9.66 mm. The
dimensions of a large specimen are as follows: Length 23,
breadth 12, aperture 10 mm. Average weight 5.7 grains.'
(Gulick).

West Maui: Lahaina, on trees (S. T. Alexander); Huelo
(Baldwin); Waihee (Perkins).

PARTULINA, MAUL 4<

Achatinelia pyramidata GUL., Ann. Lye. vi, p. 204, pi. 7. f.
32, 1856. --BALDWIN, Catalogue p. 7. — SYKES, Fauna Hawaii-
ensis p. 317.

Figured from a typical specimen from the Gulick collection,
no. 92704 A. N. S. P.

" This shell differs from the A. splendida Newc. in its smaller
size and pyramidal form, with spire less convex, body-whorl
less ventricose, aperture smaller and less oblique, lip less ex-
panded and reflected. It also differs in being without perfora-
tion, and is very rarely sinistral.

" The A. perdix Reeve differs from this in its broader and
more ventricose form, its more convex spire, with whorls more
swollen, with aperture broader, and frequently subangulated,
the arrangement of colors is also different. Looking only at the
type of this species, it would be placed in the same group with
A. splendida, yet it is so closely connected by intermediate varie-
ties with A. perdix that it has hitherto been considered a variety
of that species.

" Var. b. With black basal band. A small speciman has the
following dimensions: length .8, breadth .43, length of aper-
ture .35 inch, weight 4 grains.

"Var. c. Without band. I have a small specimen of the
brown variety of A. splendida which closely resembles this.

"Var. d. With apex chestnut-brown, not tessellated; very
rare.

"Var. e. With irregular white spots. This variety passes
into A . perdix. ' ( Gulick).

22. P. USTULATA (Gulick). PI. 13, figs. 5, 6.

"Shell sinistral, perforate, ovate-conic, obliquely produced at
the base, solid, scarcely shining, striated and microscopically
distinctly decussated, brown with a whitish band encircling the
base. Apex subacute, tessellated with white and brown; spire
conical, somewhat curvilinear; suture simple, lightly impressed,
marked with a narrow white line; whorls 6^, moderately con-
vex, the last large, equal to 72 hundredths of the length. Col-
umellar fold strong, white, oblique. Aperture very oblique,
sinuously oval; peristome thickened within, reflected anteriorly,

48 PARTULINA, MAUL

*

with columellar margin dilated, white, unattached; parietal
margin wanting. Length 25.4, breadth 14.5, length of aperture
13.2 mm. Weight 10 grains" (Gulick).

Maui: Beautiful Valley, on the mamaki (S. T. Alexander).

Achatinella nstulata GUL., Ann. Lye. N. H. of N. Y. , vi, p.
211, pi. 7, f. 37, 1856.

"This species is more ventricose than A. perdix Rve. and
differs in having the whorls more convex, the suture simple,
the aperture more oblique and the sculpturing more distinct.
I have received but two examples for which I am indebted to
the brothers James and Samuel Alexander ' (Gulick).

Also reported from Lahaina (Baldwin). I have copied
Gulick's figure (pi. 13, fig. 6), and add another (fig. 5) repre-
senting a shell from the Gulick collection from Kahoma, W.
Maui. It has two light bands on the last whorl, and the spiral
sculpture is very weak.

23. P. INDUTA (Gulick). PL 13, fig. 7.

Shell dextral, perforate, ovate-conic, solid, unpolished, micro-
scopically very minutely and obsoletely decussated, brown.
Apex subacute, tessellated with white and chestnut; spire con-
vexly conical; suture lightly impressed, not margined. Whorls
6, slightly convex, the last equal to 66 to 72 hundredths of the
whole length. Columellar fold strong, white, nearly transverse.
Aperture slightly oblique, sinuously oval, bluish-white within;
peristome thickened within, with anterior margin slightly re-
flected. Columellar margin dilated, white, unattached; parietal
margin very thin or wanting. Length 21, breadth 12|, length
of aperture 10 mm. Average weight 6.7 grains' (Gulick).

West Maui: mountain ridges of Wailuku, on the ilima (Sida)
and other bushes (E. Bailey).

Achatinella induta GULICK, Ann. Lye., vi, p. 207, pi. 7, f.
34a, 34e, 1856. --BALDWIN, Catalogue, p. 6.

"This species is characterized by its unpolished brown ex-
terior' (Gulick). It has a close resemblance to the East
Mauian marmorata Gld., and is very near ustulata Gul. A.
perdix and undosa differ by their polish and more definite color-
patterns. Figured from specimens of the Gulick collection, no.

PARTULINA, MAUI. 49

92712 A. N. S. P. Another lot from Waihee is similar, but
contains some individuals marked somewhat like marmorata.

1 ' Var. 6. Ash-brown with whitish streaks and spots. This
variety passes into A. undosa.

"Var. c. With a white band on the periphery of the last
whorl; very rare.

"Var. d. Small, ovate-conic. The dimensions of one of the
smallest specimens are as follows: length 18, breadth 10, length
of aperture 8f mm. Weight 4 grains.

"Var. e. Lip considerably expanded and reflected, slightly
thickened within; columella broadly reflected. This variety
presents a remarkable feature; appearing as if the body-whorl
had been enlarged and the lip expanded by a second growth
after the shell has been perfected and the lip thickened. The
second growth forms a broad lip sometimes an eighth of an inch
or more in width from the interior callous ridge to the edge.
This lip seem to be of the same material as the thickening
within, the exterior being of an unpolished ivory color, not
covered with the brown coat which clothes the rest of the shell.
The interior of both the lip and columella is of polished ivory
white; aperture within the lip bluish white. Specimens pre-
senting more or less of the features of this variety are not rare '
( Gulick) .

24. P. RADIATA (Gould). PL 13, figs. 8, 9.

Shell solid, ovate-conic, longitudinally painted with chestnut,
yellow and whitish lines, sometimes girdled with a median or
basal white band; subumbilicate; whorls 6, convex, the suture
margined, whitish; aperture ovate, peristome reflexed, yellow;
fold broad. Length f , diam. 9-twentieths inch. In size and
markings it resembles Bui. radiatus, but the lines are finer and
more numerous (6Wc?.).

Sandwich Islands (Old.).

Achatinella radiata OLD., Proc. Boston Soc. N. H., 1845, p.
27; Otia Conch., p. 195. — PFR., Monogr., iii, 454; iv, 517; vi,
162. — Bulimus gouldi PFR., Zeitschr. f. Malak., 1846, p. 116;
Monogr., ii, p. 74. — Partula densilineata REEVE, Conch. Icon.,
vi, pi. 2, fig. 9, April, 1850.

50 PARTULINA, MAUI.

The locality of this species is not known; radiata and gouldi
were stated to be from the Sandwich Islands, and for densilineata
no habitat was known. It is not even certain that the three
names pertain to one species, though I think that they do.
Gould's white-banded specimens were probably different. The
original descriptions are given below.

I believe that Newcomb correctly identified radiatus but un-
fortunately the specimens from him in our collection bear no
locality. Two of them are illustrated in pi. 13, figs. 8, 9. The
shell is distinctly perforate, ovate, with rather straight-sided
spire. Spiral striation weak, very faint on the last whorl,
whorls weakly convex, the last convex; the suture usually
marked with a white line. Embryonic whorls uniform pale
brown but darker at the tip; later whorls closely streaked with
narrowr streaks or lines of brown, cream and whitish. Lip
slightly expanded, more so below, whitish, calloused within.
vColumellar fold rather sharp but not prominent.

Length 18, diam. 11.3, aperture 9.8 mm.; 6£ whorls.

Length 18, diam. 10.5 mm.

One of the lot has the brown replaced by olivaceous, and an-
other has faint traces of several spiral lines on the base.

While the absence of flammules on the last embryonic whorl
is anomalous, yet the species has the appearance of a Maui

shell.

Bulimus gouldi Pfr. Shell narrowly umbilicate, ovate-conic,
rather solid, very delicately decussate-striatulate, glossy, ele-
gantly radiated with ashen, brown and white. Spire conic,
rather acute. Whorls 6, a little convex, the last about three-
sevenths the total length. Columella twisted plicate, white.
Aperture oblong, white within. Peristome narrowly expanded,
the margins subparallel, columellar margin broadly reflexed,
not covering the umbilicus. Length 19, diam. 10, aperture 9
mm. long, 5 wide within. Sandwich Islands, Mus. Cuming

*• f

Partula densilineata Reeve (pi. 13, fig. 4). 'Shell acumi-
nately conical, rather thin, umbilicated, sutures of the spire
impressed, whorls rather flattened and finely margined around
the upper part, rounded and slightly angled at the lower, colu-

PARTULINA, MAUI. 51

mella flatly reflected, obsoletely plicate, aperture ovate, lip but
little reflected; whitish, very thickly painted with oblique rust
and ash lines. Hab. ? The lip of this shell, the only ex-
ample of the species known to me, is scarcely mature. The
linear painting is discontinued before reaching the umbilicus '
(Rve.)

Series of P. splendida.
25. P. SPLENDIDA (Newcomb). PL 11, figs. 1 to 10.

"Shell sinistral, solid, ovate-acuminate, finely decussately
striated; suture moderately impressed, margined. Whorls 6;
body whorl somewhat inflated. Aperture ovate; lip expanded;
columella short, broad and twisted. The upper two whorls tes-
sellated writh chestnut and white, lower whorls with numerous
chestnut-colored transverse lines and fillets traced on a polished
white ground, markings correctly lined superiorly and irregu-
larly serrated inferiorly. Length 1, diam. .55 inch' (Newc.).

West Maui : Wailuku, on tutui trees (Newcomb); Mauna
Hoomaha (Thaanum); Lahaina, Wailuku and Wailee (Gulick).

Achatinella splendida NEWC., Ann. Lye. N. H. of N. Y. vi, p.
20, April, 1853; P. Z. S., London, 1853, p. 131, pi. 22, f. 4;
Ann. Lye. N. H. of N. Y. vi, p. 324. — PFR., Monogr. iv, 516.
-TnwiNG, Reprint Orig. Descript. Achat., pi. 2, f. 17. — Acha-
tinella solida Gulick, MS. according to PFEIFFER, Monographia
iv, 516. — Achatinella baileyana GULICK, Ann. Lye. N. H. vi, p.
202, pi. 7, f. 31a, b (Dec. 1856).

The embryonic shell consists of 3 to 3^ whorls, the first 1 or
1^ flesh-tinted, wide, indistinct dark stripes then appearing.
The last embryonic whorl has alternately snow-white and brown
stripes, very obliquely descending forwardly. The banded pat-
tern then begins. The lines and bands of brown are usually a
little serrate. The surface is glossy, with dense, weakly devel-
oped decurrent striae. Banding variable, as shown in the fig-
ures, in all the localities mentioned above.

In a lot of 42 from Wailuku, 24 are sinistral. A lot of 32
from Wailee has 7 sinistral, all with broad bands. From La-
haina, in a lot of 60 shells, 34 are sinistral, patterns the same
as in the dextral shells. Specimens from Wailuku are figured,

52 PARTULINA, MAUI.

figs. 1 to 9. Fig. 10 is from Lahaina. Wailuku specimens
measure:

Length 25J, diam. 15 aperture 12^ mm.

Length 25J, diam. 13.8 aperture 11.2 mm.

Length 24, diam. 15 aperture 12 mm.

Length 22, diam. 12.5 aperture 10 mm.

Var. baileyana (Gulick) . PI. 11, figs. 11, 12, 13. The shell
is sinistral. perforate, white with few or many narrow deep
brown spiral lines, sometimes with a broad belt of pale brown
under the lines; otherwise as in splendida. Length 21.6, diam.
12.4, aperture 10 mm. weight 7 grains, or larger, length 25.4,
diam. 13.5, aperture 11.4 mm. Wailuku mountain, on trees,
Ed. Bailey Jr. Gulick describes three varieties as follows:

"Var. b. Regularly conical, rather broad at the base, with
spire shortened. Whorls nearly 7, convex, not margined.
Aperture very oblique, semiorbicular, pale blue within. Length
17, breadth 13, length of aperture 8f mm. Weight 6 grains.

"Var. c. Ovate-conic, marble gray, bands usually wanting;
within the aperture pale lilac; whorls somewhat swollen beneath
the sutures; columella not so short. This variety passes into
the wrhite variety of A. splendida Newc.

' ' Var. d. Lightly tinged with pink both within and without. '

26. P. GOULDII (Newcomb). PI. 11, figs. 14, 15, 16.

Shell dextral, conically ovate, longitudinally striate; suture
moderate, not margined or but slightly so; whorls 6, flatly con-
vex, color yellowish-white with zigzag lines of brown on the
third whorl, and brown lineations more or less numerous en-
circling the three lower whorls. Aperture rounded ovate,
yellowish; columella lightly callous; lip expanded and below
reflected. Length .85, breadth .45 inch (Newc.).

West Maui: Wailuku valley, on tutui trees (Newcomb).

Achatinella gouldii NEWC., Ann. Lye. N. H. of N. Y., vi, p.
21, April, 1853; P. Z. S., 1853, p. 129, pi. 22, f. 1, 1854.—
PFR., Monogr., iv, 517. — A. myrrhea Gul., according to Pfeiffer,
/. c., undescribed. — Achatinella talpina GULICK, Ann. Lye., vi,
p. 212, pi. 7, f. 38, 1856.

This species resembles P. splendida Nc. in color-pattern, but

PARTULINA, MAUL 5->

it is smaller, with less convex whorls, the last one shorter. So
far as we know it is always dextral. Perforation small, some-
times closed. In the typical form the bands, of a light brown
color, are more or less distinctly spotted with darker; and when
the bands spread to cover most of the surface, they are marked
with dark streaks. The last embryonic whorl has oblique
stripes as in related species, and the first neanic whorl is irregu-
larly clouded and streaked. The columellar fold is low and
oblique, white. Lip distinctly but narrowly thickened within,
white, outwardly a little expanded.

Length 21, diam. 12, aperture 10 mm.; 6J whorls.

Length 20, diam. 11 mm.

A. talpina Gul. has always been considered synonymous with
gouldii. It is here figured (pi. 11, fig. 17) from a specimen
from Gulick. Although Gulick mentions only one band, his
figure and specimens show numerous lines also. The colu-
mellar fold is slightly stronger and less oblique than in typical
gouldii. The original account follows:

" Achatinetta talpina, shell dextral, slightly perforate, elongate
ovate-conic, rather thin, subcorneous, not shining, finely
striated, beneath the lens scarcely decussated; brown with a
whitish band encircling the base, covered with a thin, brown
ash, velvety epidermis. Apex somewhat obtuse, tessellated
with white and chestnut; spire regularly conical; suture simple,
moderately impressed, whorls nearly 7, convex, the last regu-
larly rounded, equal to 62 to 76 hundredths of the length.
Columella strongly plaited above. Aperture semi-orbicular,
brownish-white within; peristome thickened within, scarcely
reflected anteriorly, with columellar margin white and reflected
over the deep perforation; parietal margin wanting. Length
21.6, breadth 12, length of aperture 9 mm.; weight 5.5 grains.
"Var. b. With apex corneous, without tessellations; very
rare.

"Var. c. Ventricose; spire short, curvilinear. A small speci-
men has the following dimensions: length 17, breadth lOf,
length of aperture 7f mm., Weight 4.5 grains. r

Maui: Wailuku, on the kukui, Aleuritis triloba, rare, E.
Bailey Jr.

54 PARTULINA, MAUI.

"This species passes into A. gouldiiRewc. Much handling
gradually removes the epidermis from the shell, leaving the
surface more or less polished' (Gulick').

26a. P. gouldi perfecta n. var. PL 11, figs. 18 to 21.

The bands are very dark chestnut on a cream-white ground.
The lip has a stronger callous rib within and is slightly flesh-
tinted, with faintly traced bands. The columellar fold is
twisted, very strongly projecting, subhorizontal; axis perforate
or closed. Color-pattern various, as figured.

Length 20, diam. 12, aperture 9.8 mm.; 6f whorls.

Length 19.5, diam. 11 mm.

Wailuku (D. D. Baldwin). A small form or race from the
same valley (pi. 11, figs. 20, 21) measures, length 17, diam. 10
mm. ; 6J whorls.

27. P. APTYCHA (Pfeiffer). PL 18, fig. 5.

Shell imperf orate, dextral, oblong-conic, rather thin, striatu-
late, a little shining; white, ornamented with dotted chestnut
girdles. Spire long-conic, the apex rather obtuse, suture nearly
simple. Whorls 6J, a little convex, the last nearly equal to
two-fifths the total length, rotund at base. Aperture oblique,
truncate-oval, milk-white within. Columellar fold obsolete,
very slightly twisted. Peristome simple, very lightly thick-
ened within, the right margin unexpanded, columellar margin
narrow, adnate. Length 21-J, diam. 10 mm., aperture 9^ x
5iy mm.

fA

Sandwich Islands (Frick in Cuming coll.).

AchatineUa aptycha PFR., P. Z. S., March, 1855, p. 1, pi. 30,
f. 1; Monographia, iv, 562; Nomenclator Hel. Viv., 1881, p.
315 (Perdicella) .

A "lost' species, unknown to Hawaiian collectors. It
seems as near a young gouldii or baileyana as anything. Sykes
remarks ' ' probably from Maui. '

Series of P. tappaniana.

28. P. TAPPANIANA (C. B. Adams). PL 12, figs. 1, 2, 3.
"Shell reversed, elongate ovate-conic; pure white, with one

PARTULINA, MAUI. 55

narrow brown spiral band on the periphery of the last whorl;
with very unequal and irregular transverse stria3 and numerous
excessively minute, wrinkled spiral impressed lines. Apex sub-
acute; spire elongate, with the outlines a little curvilinear.
Whorls nearly 7, moderately convex and more or less sub-
angular, margined above, with a well-impressed suture. Aper-
ture ovate; lip well thickened within the margin, expanded
much anteriorly, but not above; columellar fold strong. Mean
divergence 43 degrees. Length 1.06, breadth .55 inch; length
of aperture .44 inch. Length 26 J, diam. 13f, aperture 11 mm.

" Var. dubiosa differs in being a little more ventricose; its
whorls are not margined above, and the brown stripe is replaced
by two fine paler brown lines, below which one or two other
yet finer lines may be seen.

" Sandwich Islands.

"This beautiful species is named in honor of Hon. Benjamin
Tappan, of Ohio." (C. B. Ad.")

W. Maui: Wailuku (Gulick); Lahaina (Gulick, Baldwin).

Achatinella tappaniana C. B. Ad., Ann. Lye. N. H. of N. Y.,
v, p. 42, 1851; Contrib. to Conch., no. 8, p. 126.--PFR.,
Monogr., iii, p. 462. — SYKES, Fauna Hawaiiensis, p. 318.-
BALDWIN, Catalogue, p. 7, 1893. — THWING, Reprint Orig. De-
script. Achat. , pi. 2, f. 19.

This well-known shell has a long, noticeably concave-sided
spire, margined suture, and one narrow brown band just below
the periphery, so that it enters the aperture below the insertion
of the outer lip. There are 3J embryonic whorls, of a whitish
tint, the last half whorl in young or very fresh examples usually
showing protractive yellowish stripes. Young shells are covered
with a yellow cuticle which is deciduous, disappearing in the
adult stage. The surface is rather dull, under the lens showing
spiral impressed lines and weak decurrent striation. The young
and half grown shells are rather acutely angular at the peri-
phery in front of the aperture. Wailuku is here selected as the
type locality (pi. 12, fig. 1). Length 24, diam. 12 mm.; 7
whorls.

Kahoma specimens are somewhat stouter, length 25, diam.
14 mm. ; 6J whorls. They have one band or none, and one

56 PARTULINA, MAUI.

specimen shows faint traces near the lip of numerous other
bands, thus approaching var. fasciata.

In a series from Lahaina the shell is pure white at all stages
of growth, or there may be some faintly sketched yellowish
spiral lines below the periphery, which is acutely angular in
front in the young. The tine descending striae are more distinct
in most specimens (pi. 12, figs. 2, 3).

28a. P. tappaniana fasciata (Gulick). PI. 12, figs. 5, 6, 7.

Shell sinistral, sometimes perforate, ovate-conic, solid, shin-
ing, finely striated, microscopically very minutely and faintly
decussated, white, with brown bands; apex subacute, white,
with a brown line above the suture; spire regularly conical;
whorls 6 or 7, convex, not margined; the last rounded, equal to
_6_^. Of the length; columellar fold white, superior, slightly
developed, moderately twisted; aperture somewhat oblique,
roundly oval, white within; the bands sometimes appear in pale
brownish stripes in the inner surface; peristome white, slightly
thickened, subreflected anteriorly; with columellar margin re-
flected, scarcely adnate; parietal margin wanting. Average
length 22; greatest length 25; broadest 11^; length of aperture
10mm. Average weight 6.8 grains; greatest weight 9 grains;
least weight 6 grains. ( Gulick. )

Maui: Honukawai, on the wiliwili (Erythrina monosperma'),
S. T. Alexander.

Var. 6. Thicker and more elongate, with dark brown or black
bands; lip well thickened within, and deeply colored on the edge
opposite the bands; columella slightly toothed.

Var. c. Ventricose, conical; one specimen has the following
dimensions: Length 0.86; breadth 0.54; length of aperture 0.40
inch.

Remarks. — Has been confounded with A. tappaniana and A.
splendida Newc., but is smaller than the former, with fainter
sculpturing and more regularly conical spire; it also differs in
its dark bands, rounded body whorl, less reflected lip, and
slight columellar fold. It resembles A. splendida Newc. in its
brown bands, but is otherwise quite distinct. A nuclear char-
acter which distinguishes it from either of the above is the spiral
line on the first whorls. (Gulick.)

PARTULINA, MAUI. 57

Achatinella fasciata GUL., Ann. Lye. vi, p. 201, pi. 7, f. 30,
1856.

Figured from Honokawai shells of the Gulick collection.
Hardly distinguishable from var. ampulla; I think them ex-
tremes of one variety. The figures are from topotypes from
Gulick. The same form is in the Gulick collection from
Wailuku.

286. P. tappaniana ampulla (Gulick). PI. 12, fig. 4.

"Shell sinistral, sometimes slightly perforate, elongate, sub-
pyriform, rather thin, shining, finely striate, scarcely decussate
beneath the lens, white, with a broad chestnut band encircling
the base, and revolving within the shell beneath the suture,
sometimes with fine spiral lines above; apex subacute; spire
concavely conical, sometimes decollated; whorls 6-J-, convex,
margined above; the last inflated, equal to -f/-^ of the length;
columella white, plaited near the whorl, strongly twisted, not
tuberculate; aperture rounded; peristome expanded, reflected,
very slightly thickened; with columellar margin reflected, adnate
or slightly detached; parietal margin wanting. Length 23;
breadth 13; length of aperture 10 mm. Weight 6 grains"
(Gulick).

Maui: Honukawai, on trees, S. T. Alexander.

Achatinella ampulla GULICK, Ann. Lye. vi, 200, pi. 7, f. 29.

"This species differs from A. fasciata Nob., to which it is
closely allied, in the concave outlines of the spire, in its inflated
body whorl, and more expanded and reflected lip' (Gulick).

28c. P. tappaniana eburnea (Gulick). PI. 12, figs. 9, 10, 11, 12.

"Shell sinistral, usually perforate, ovate-conic, somewhat
solid, shining, striated, very minutely decussated, ivory-white;
apex rather acute; spire conca^ely conical; suture slightly mar-
gined, lightly impressed; whorls 6, slightly convex; the last
large, obliquely produced, and angulated above the aperture;
columellar fold central, white, strong, nearly transverse; aper-
ture oblique, truncately auriform, white within; peristome
thickened; with external margin reflected, anteriorly arcuate;
columellar margin dilated, slightly detached; parietal margin
wanting. Length 24; breadth 14; length of body whorl 17 mm.
Average weight 9 grains; greatest weight 13 grains ' (Gulick).

58 PARTULINA, MAUI.

East Maui: Honuaula, on the trunks of trees, quite exposed,
E. Bailey.

"Var. b. Brownish-yellow, with a white girdle on the peri-
phery of the last whorl, waved with white and yellow on the
third whorl.

1 1 Var. c. With body whorl rounded, not angulated. This spe-
cies is the analogue of A. tappaniana C. B. Ad. found on W.
Maui, but differs from it in its more ventricose form, its angu-
lated body whorl, and more regularly curved lip, besides the
difference in color" (Gulick).

AchatineUa eburnea GULICK. Ann. Lye. N. H. of N. Y. vi, p.
199, pi. 6, f. 28a, b. 1856.— BALDWIN, Catalogue, p. 6
(Makawao). — THWLNG, Reprint, pi. 2, f. 14.

I cannot distinguish the typical form of eburnea (pi. 12, figs.
9, 10) from the pure white West-Mauian form of tappaniana.
The Var. b (pi. 12, figs. 11, 12) however is unlike any West-
Mauian form I have seen, and approaches very close to carnicolor
Baldwin. I doubt whether a specific line can be drawn between
them.

29. P. CARNICOLOR Baldwin. PI. 12, figs. 13, 14.

"Shell dextral, minutely perforated, solid, acuminately
ovate conic, apex subacute; surface lusterless, marked with
delicate incremental stride, and under a lens exhibiting a fine
pattern of decussating spiral striae; nuclear whorls faintly decus-
sated. Of a uniform brown color, with a narrow white line
below the periphery which enters the aperture, and a very
narrow white line traversing the suture. Whorls 6, slightly
marginate above, a little convex; suture lightly impressed.
Aperture oblique; oval, livid white, showing the exterior color-
ing within. Peristome acute, expanded, columellar margin
broadly reflexed. Columella white, terminating in a well
developed, flexuous fold. Length 25, d.iam. 14 mm.': (Bfcld-
win) .

East Maui: Nahiku.

Partulina carnicolor BALDWIN, Nautilus xix, February, 1906,
p. 112.

In sculpture this form agrees with eburnea Gulick. It is

PARTULINA, MAUL 59

fleshy brown, darkest at the base, the aperture flesh-colored
within. The band below the periphery is sometimes very
narrow — less than 1 mm. wide. I think it doubtfully distinct
from the var. b of eburnea. Figured from cotypes.

30. P. NIVEA (Baldwin). PI. 12, fig. 8.

The shell is sinistral, subperforate, acutely conic, thin but
moderately strong, glossy, snow-white with a brownish-yellow
band below the periphery. Growth-striae distinct but fine and
irregular; spiral striation almost effaced on the last whorl, more
distinct above. The spire is a little attenuated near the acute
apex; whorls but slightly convex, the suture not deep, margined
below. Last whorl a little swollen just below the suture, then
flattened, often having a peripheral angle weakly sketched in
front of the aperture; periphery very convex; base not very con-
vex. The aperture is oblique, white within, with a rather
strong rib within the lip, which is a little expanded. Columellar
lamella white, rather strong and oblique.

Length 24, diam. 14 mm. ; 6-J whorls.

Length 23, diam. 12 mm.; 6^ whorls.

''Animal when extended in motion longer than the shell.
Mantle densely black, with minute brown flecks and a broad
yellowish band encircling the outer edge. Tentacles and supe-
rior portion of foot brown with a slight slate tinge. Under por-
tion of foot light brown. Front above covered with light granu-
lation ' ( Baldwin ) .

East Maui: Makawao to Huelo (Baldwin).

Achatinella nivea BALDWIN, Proc. A. N. S. Phila., 1895, p. 222,
pi. 10, f. 19. — THWING, Reprint Orig. Descript. Achat., pi. 2,
f. 16.

"I have before me over 500 examples of this species, which
are without variety in color or shape' (Baldwin'). It differs
from tappaniana by the more acute apex, less attenuated spire,
flatter whorls, the last having its greatest convexity lower down,
and by the more glossy surface. In A. dolei the upper part of
the last whorl is much more convex, and the color differs. De-
scribed from the type lot, no. 65691 A. N. S. P. Several other
lots seen agree exactly with this one.

60 PARTULINA, MAUI.

30a. Var. kaupakaluana n. v., pi. 13, fig. 16, has the last 2J
whorls closely streaked with ochraceous or olive-buff on a
whitish ground, a white belt on the base. It is known to us
by two examples, not quite mature, from Kaupakalua, no. 2101
coll. C. M. Cooke.

31. P. DOLEI (Baldwin). PL 12, figs. 15 to 18.

Shell sinistral, perforate, acutely pyramidal, thin but strong,
glossy, white with a dark chestnut band below the periphery,
and usually several lines and obliquely-streaked bands of light
brown in the peripheral region. Surface polished, delicately
marked with growth-strise and almost effaced spiral lines. Spire
straight! )T pyramidal, rather acute, the apex white. Post-
embryonic whorls decidedly convex, the last swollen below the
suture, then somewhat compressed; suture deep, narrowly mar-
gined. The aperture is oblique, white within, with a rib within
the lip, which expands somewhat. Columellar fold strong and
oblique.

Length 24J, diam. 14 mm. ; 6f whorls.
Length 25, diam. 15 mm.
Length 25J, diam. 14 mm.

"Animal when extended in motion longer than the shell.
Mantle intensely black with a broad outer margin of yellow and
a conspicuous orange spot in the center of the margin. Foot
tapering behind, above and below light yellow. Tentacles long
and slender, light slate. Head above lightly granulated'
(Baldwin).

East Maui: Honomanu, very abundant in humid forests of
the exterior slope of Haleakala crater, 7000 feet above sea level
(Baldwin).

AchatinelladoleiBA-Lvwix, Proc. A. N. S. Phila., 1895, p. 221,
pi. 10, f. 17, 18.

The dark band below the periphery is a constant feature, and
rarely no other markings are present. When lighter bands are
developed they may ascend the spire above the suture, or they
may be confined to the last whorl. The embryonic shell (pi.
15, fig. 18) is acutely angular and has a subperipheral band.
The columella is convex. It is closely related to P. nivea but

PARTULINA, MAUI. 61

differs by the swollen upper part of the last whorl and the color-
ation. P. splendida Newc. of West Maui has a differently
shaped last whorl. Figures 15-18 are from cotypes. Named
in honor of the President of the Republic of Hawaii, Hon. S. B.
Dole.

32. P. LEMMONI Baldwin. PI. 12, figs. 19, 20.

The shell is sinistral, imperforate, pyramidal, ivory-white
variously marked with dark chestnut bands, which are darker,
chocolate, at the base, broad band at and below the periphery,
another around the columella constant; suture bordered with a
white line, dark bands above and belowr it, the embryonic shell
white. Surface glossy, finely striate with growth-lines and fine,
weak spirals. Outlines of spire nearly straight, last two whorls
convex. Aperture oblique, blue-banded within on a white
ground. Lip a trifle expanded, thickened within, spotted by
the bands. Columellar fold rather strong.

Length 23, diam. 12.5, length aperture 10 mm. 6J- whorls.

Length 20, diam. 11.7 mm., whorls 6J.

"Animal in motion as long as the shell. Mantle brownish
black, mottled with white streaks, with a broad yellowish brown
border somewhat interrupted. Foot above and below very light
brown. Tentacles slate color' (Baldwin}.

East Maui: Nahiku (N. E. Lemmon).

Partulina lemmoni BALDWIN, Nautilus xix, p. 112.

This quite distinct species is related to eburnea, dolei and the
smaller flemingi, in which however the bands extend upon the
embryonic shell. Figures and description from cotypes.

33. P. TEREBRA (Newcomb). PL 15, figs. 1 to 9.

"Shell turrited; whorls 6, rounded, last one inflated, mar-
gined above; suture well impressed. Aperture elongate-ovate;
lip slightly reflected, thickened within. Columella short, ter-
minating in a prominent, twisted plait. Stria3 decussating.
Color light yellow, with wave-like brown-colored markings,
nearly obsolete on the upper whorls; lip, columella and aper-
ture white. Length if, diam. -£$ inch ' (Newc.).

West Maui (Newcomb); Honokowai (Baldwin).

62 PARTULINA, MAUI.

Achatinella terebra NEWC.. P. Z. S., 1853, p. 144, pi. 23, f.
40. — PFR., Monographia iv, 525; vi, 168.--TnwiNG, Reprint,
pi. 2, f. 18. — A. corusca Gulick and A. perforata Gulick, unde-
scribed, PFR., Monogr. iv, 525 (names only). — Achatinella
attenuata PFR., P. Z. S., 1855, p. 4, pi. 30, f. 12. — Achatinella
lignaria GULICK, Ann. Lye., p. 209, pi. 7, f. 35, 1856.

Newcomb's original figure is copied, pi. 15, fig. 2. A speci-
men received from Dr. Newcomb, typical in markings though
somewhat smaller, is drawn in pi. 15, fig. 1. The last embry-
onic whorl has rather wide stripes, the following whorls narrow,
more or less angulated in places, and of a tawny color on a
white ground. The surface is glossy and the spiral and decur-
rent striation weak. Length 18, diam. 9.5 mm., of fully 6
whorls.

Besides this typical form, Newcomb sent out lignaria, corusca
and longior as terebra. It will probably be possible for a natur-
alist in the field to arrive at some trustworthy conclusion as to
the rank or value of the several named forms of this group.
Without such knowledge, I must be satisfied to give the data
relative to the various forms.

In a series from Waiehu (pi. 15, figs. 4 to 8) from Baldwin
the shell is stouter than PfeifTer's figure of attenuata, and the
stripes are less flexuous. Surface glossy. The}'' vary from pure
white, through faintly striped to copiously striped forms, the
latter usually having the ground of a pale brown tint. The
shape also varies. Length 16 to 18 mm. The white examples
are not distinguishable from the following variety.

Var. corusca Gulick. PI. 15, fig. 9. Gulick referred speci-
mens from Wailuku (which intergrade with lignaria) and from
Waihee, to attenuata. His undescribed A. corusca, according to
unlocalized specimens before me, agree well with the Waihee lot,
and were probably from that place. The name was afterwards
abandoned by Gulick, who adopted attenuata in its place. The
shells are white, rather glossy, often with some faint bluish
stripes on the last embryonic whorl. The sutural margination
is conspicuous. Spiral sculpture visible throughout but rather
weak. Length 16.3, diam. 8.5 mm. to length 18, diam. 9 mm.

The Wailuku lot consists partly of shells like those from

PARTULINA, MAUI. 63

Waihee, partly of more swollen shells, with more or less straw
or brownish tint. One of the white shells has dark bands at
periphery and columella. On another there are very faint
tawny stripes.

33« P. t. attenuata (Pfeiffer). PI. 15, fig. 3.

Shell subperforate, ovate-turrited, rather thin, striatulate and
under a lens granulate, white, painted with corneous streaks
which are often angular. Spire concavely turrited, the apex
rather obtuse, suture distinctly thread-margined, whorls 5J-, the
upper flat, those following convex, last whorl about three-
sevenths the total length, rounded, base subcompressed. Aper-
ture oblique, subtetragonal-auriform ; columellar fold strong,
compressed, tongue-shaped; peristome acute, narrowly ex-
panded, labiate within, basal margin forming an angle within
with the dilated columellar margin. Length 16, diam. 7-J mm.,
aperture with peristome 7J- mm. long, 4 wide. Maui, Frick

(flW-

The type figure is copied, pi. 15, fig. 3. The smaller size
and narrower form distinguish it from terebra and lignaria.
Type locality unknown.

335. Var. longior n. v. (pi. 15, fig. 12). Straightly pyramidal,
whitish with a pale yellow tint, the last embryonic whorl
generally having faint stripes, and under a lens very faint yellow
streaks may often be seen on the last whorl. Surface with but
little gloss, and usually deeper spiral sculpture than in the
smaller var. corusca.

Length 22, diam. 10J mm. 6f whorls.

Length 22, diam. 10 mm. 7J- whorls.

Wailuku (Gulick); also Waiehu (Baldwin).

33c. P. t. lignaria (Gulick). PI. 15, figs. 13 to 17.

"Shell dextral, perforate, ovate-conic, solid, more or less
shining, microscopically minutely decussated, yellowish ivory-
white, sometimes streaked; apex subacute, white, obliquely
marked with brownish-yellow; spire regularly conical, suture
moderately impressed, scarcely margined; whorls 6J, somewhat

64 PARTULINA, MAUI.

convex; the last equal to -^g- of the length; columellar fold
strong, white; aperture somewhat oblique, sinuously oval; peri-
stome thickened, sometimes slightly expanded and reflected
anteriorly; columellar margin dilated, white, unattached; pari-
etal margin wanting. Length 20J; breadth 10-f ; length of aper-
ture 9 mm. Average weight 5.5 grains ' (Gulick*).

West Maui: Wailuku, on the alii and other low trees, in
damp, elevated regions. E. Bailey.

" This species differs from A. induta Nob. in its smaller size,
more acuminate form, lighter color, and in the polished exterior
of some of its varieties.

" Var. b. More ventricose; aperture nearly semiorbicular.
This variety passes into A. induta.

"Var. c. More elongate. Length 19 J; breadth 9; length of
aperture 8 mm. ; average weight 3.50 grains. This variety ap-
proaches and passes into A. terebra Newc. The two species are,
however, distinctly characterized; the latter being more strongly
sculptured, having the aperture more elongately oval, and the
suture more distinctly margined.

' ' Var. d. With black or brown spiral bands. This passes
into A. splendida Newc., but is distinguished by its smaller size
and stronger columellar fold. Rather rare.

u Var. e. White. This passes into A. attenuataPL

li Var. /. Pale ash-color. Passes into A. undosa Nob.

" Var. g. Pale yellowish brown. Passes into A. crocea Nob.'
(Gulick).

Specimens of Gulick's series (no. 92717 A. N. S. P.) are
figured. The un described A. perforata Gulick, according to
specimens (no. 57783 A. N. S. P.) is identical with lignaria.

34. P. FUSOIDEA (Newcomb). PI. 15, figs. 10, 11.

11 Shell sinistral, rather solid, elongately produced; whorls 6
to 7, scarcely rounded (excepting the last), margined above;
suture wrell impressed. Aperture roundly ovate, small; colu-
mella very short, robust and twisted; lip simple above, slightly
expanded below. Color straw yellow. Length 15, diam. 5J
twentieths of an inch ' (Newc. ) .

East Maui (Newcomb); Haleakala (Thwing).

Aehatinellidse

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PARTULINA, MAUI. 65

Achatinella fusoidea NEWC., Ann. Lye. N. H. of N. Y., vi,
1856, p. 144; Amer. Journ. of Conch., ii, 1866, p. 213, pi. 13,
f. 8. — PFR., Monogr. iv, 539. — THWING, Orig. Descript, p.
112.

" This is an exceedingly rare species, but two specimens
having come under my observation, one of which serves for
the illustration, the other in the cabinet of the late Hugh
Cuming, in London ' (Newc.).

This species seems close to A. terebra var. longior of "West
Matii, a dextral shell.

Shells from Ukumehame, West Maul (pi. 15, fig. 11), sent
by Mr. D. Thaanuni, agree well with Newcomb's figure of the
type of fusoidea. They are buff with faintly darker streaks,
having wide light-brown stripes on the last embryonic whorl.
Under the lens the surface is weakly plicate, as in P. terebra
longior. Length 17, diam. 8 mm.

35. P. CROCEA (Gulick). PI. 13, figs. 14, 15.

" Shell dextral, perforate, conic-ovate, solid, scarcely shin-
ing, lightly striate and microscopically distinctly decussated
with crowded undulating spiral lines ; orange yellow ; apex
subacute, obliquely marked with white and yellow; spire
conical; suture simple, slightly impressed; whorls 6, some-
what convex, the last slightly flattened in the middle, equal
to 71 hundredths of the length; columellar fold moderately
developed, white; aperture slightly oblique, sinuously oval,
snowy white within ; peristome not expanded, moderately
thickened within ; with columellar margin dilated, white, un-
attached ; parietal margin wanting. Length 18 ; breadth 10 ;
length of aperture 9^5 mm. Weight 4 grains.' (Gulick.)

Maui : Waihee, rare. E. Bailey, Jr. Honokowai (D. Thaa-
num).

Achatinella crocea GUL., Ann. Lye. N. H. of N. Y., p. 210,
pi. 7, f. 36, 1856.

The sculpture of impressed spirals and dense, fine, pro-
tractive wrinkles or ripples is very clearly developed. The
color is yellowish-green (tawny-olive of Ridgway's Nomen-
clature of Colors) , but in the young there is often more of a

66 PARTULINA, OAHU.

gamboge tint. There is often a pale line on the periphery.
Embryo pale brown, the last half whorl striped with brown.
The lip is thickened within, white or flesh-tinted. Columellar
fold often very strong and more or less tinted with brown,
but white at the end. Figured from cotypes. Probably a
variety of A. terebra lignaria, yet I have seen no actually in-
tergrading specimens. It seems well characterized by the
color and short contour.

In a series from Honokowai, West Maui, sent bv Mr. Thaa-

«/

num, the color of the last whorl varies from pale buff to dull
gamboge, being darkest on the base. The spire is generally
more drawn out than in the figures on plate 13, and is a little
attenuated, its outlines distinctly concave or sometimes almost
straight. The flames on the last embryonic whorl are occa-
sionally lost by wear.

SPECIES OP OAHU.
36. P. MONTAGUI Pilsbry. PI. 24, figs. 7, 8.

The shell is sinistral, perforate, thick and solid, ovate-conic,
the spire acuminate, its outlines straight or somewhat con-
cave above. Embryonic whorls distinctly striate spirally;
last three whorls rudely sculptured with close, irregular
wrinkles, and on the last whorl there are spirally and ob-
liquely descending ridges producing flattened facets (as in
many large Lymnaeas). The minute spirals disappear on
the later whorls. The upper whorls are nearly flat, the last
three slightly convex, last whorl more convex, the base rather
rotund. Suture superficial, sometimes having a margin de-
fined by an impressed line on the last whorl, but in other ex-
amples this is wanting. Aperture slightly oblique. Outer
and basal margins of the peristome are expanded, thick, and
have a heavy callous thickening within ; columellar margin
raised, thick; columellar fold thick and moderately project-

ing.

(a) Length 25, diam. 14, length aperture 12 mm.; l
whorls.

(b) Length 24.7, diam. 12.5, length aperture 11 mm.;
whorls.

PARTULINA, OAHU; EBURNELLA. 67

(c) Length. 26.9, diam. 13.1, length aperture 12.1 mm.

Oahu: Manoa road at its junction with the upper road,
back of Rocky Hill, cotypes (a) no. 108181 A. N. S. P.; (6)
no. Ill coll. Irwin Spalding; and (c) no. 33581 B. P. Bishop
Mus. Also found on Rocky Hill about ^4 m^e from the type
locality.

Partulina montagui PILSBRY, Nautilus, xxvii, p. 40, August
1st, 1913.

This species differs strikingly from all others by its sculp-
ture and small apical whorls. It seems to be nearest to P.
divightii of Molokai. No specimens yet found show any trace
of color. The periphery is angular in the young, the angle
persisting as far as the last whorl in some individuals.

It was first found by Dr. C. Montague Cooke. The shells
occur in the humus near the surface, along the roadside, and
are very rare, apparently lying in " pockets ' which have
been filled and covered by the wash down the slope. Probably
the forest disappeared from where the shells are found not
much more than a hundred years ago.

Section Eburnella Pease.

Eburnella PEASE, P. Z. S., 1869, p. 647.— GULICK, P. Z. S.,
1873, p. 91, as section of Partulina ; A. variabilis selected as
an example.

Very smooth, glossy shells, uniform white, yellow or brown,
or variously banded, not striped longitudinally; outer lip
simple or scarcely expanded; columellar fold strong. Type

P. variabilis (Newc.).

The typical group consists of Lanaian species in which the
peripheral keel persists throughout the young stages, and
often upon the first half of the last whorl. In the allied
groups of Molokai and Maui the keel is lost much earlier ; the
later stages of youth and the mature stage being round at
the periphery. The group is not strongly individualized,
some species, as P. mutabilis, showing transition to the pattern
of Perdicella, others approaching Partulinse of the striped
group of Maui.

Eburnella was originally a heterogeneous group comprising

68 PARTULINA, MAUL

the following species : A. casta Nc., curta Nc., livida Sw., por-
cellana Nc., recto Nc., saccata Pfr., undulata Nc., semicari-
nata Nc., and variabilis Nc. Gulick, in his classification of
1873 restricted it to Lanaian species, giving Partulina varia-
bilis Newc. as an example. I think it may with advantage be
enlarged to include species of Molokai and Maui which seem
related.

Series of P. porcellana. MAUI.

Glossy shells with weak spiral s-triation ; uniform or with
color in bands, usually few in number and well contrasted;
the embryonic shell is banded above or below the suture or
is plain, and has no longitudinal or oblique markings (except
in P. mutabilis) .

These medium-sized or small shells have much the appear-
ance of Achatinellastrum, but various indications — the char-
acteristic decurrent sculpture more or less visible on all the
forms, and the vanishing zebrine stage in P. mutabilis — lead
me to consider them modified Partulinae. This is also the
view taken by Mr. Baldwin, and suggested in Hyatt's Ms.
Other authors have grouped part of them in the Oahuan sec-
tion Achatinellastrum.

37. P. MUTABILIS Baldwin. PL 20, figs. 1 to 7.

Shell dextral or sinistral, minutely perforate, narrowly
conical, the sides slightly concave, apex rather obtuse; solid,
rather glossy, striatulate and under the lens showing weak,
rippled spiral striation. " Color varying from pure white
to dark fulvous, often variously striped with brown lines and
bands ' ' ; usually pale buff-yellow or whitish with numerous
chestnut spiral lines and bands, or snow-white throughout;,
the last embryonic whorl frequently striped longitudinally.
Suture generally margined below. The aperture is small,
white, lip very narrowly thickened within ; columellar lamella
white, rather strong, thick and oblique.

Length 17, diam. 8.8, aperture 7.5 mm. ; 6^4 whorls.

Length 16.3, diam. 9.3, aperture 7.8 mm. ; 6 whorls.

West Maui : Waiehu valley.

PARTULINA, MAUI. 69

Partulina mutabilis BALDWIN, Nautilus, xxii, p. 68, No-
vember, 1908.

i

Distinct by its narrow contour, color in bands when pres-
ent, and the frequent presence of a brief zebra-striped stage on
the last embryonic whorl. It has a superficial resemblance
to the Lanaian A. variabilis, which belongs however to a
somewhat different series. Figured from specimens of the
type lot received from Mr. Baldwin.

P. mutabilis is an important species in that it forms a con-
nection between the Partulinas having zebra striping and
those having spiral bands only. The embryonic whorls are
flattened, the first flesh-tinted. In most banded individuals
some indistinct protractive brown streaks appear near the
end of the second whorl, becoming stronger and angularly
bent on the next fourth to half whorl, gradually giving place
to spiral bands which arise about the middle of the third
whorl (pi. 20, figs. 6, 7). This pattern is not essentially un-
like that of Partulina splendida. In some individuals, how-
ever, the longitudinally striped stage is omitted, and the spiral
bands appear (at first very faintly) on the latter part of the
second whorl, 'arising from the uniform fleshy tint of the pre-
ceding part of the embryonic shell (pi. 20, fig. 5). Such
shells are comparable to P. porcellana and its allies, and may
be regarded as more accelerated than those having an ob-
liquely striped stage.

In a series of 18 sent by Mr. Baldwin, 10 are dextral. The
same color patterns are found in dextral and sinistral indi-
viduals. The variations are as follows:

(1) Pure white throughout.

(2) Pale buff, last embryonic whorl faintly striped longi-
tudinally.

(3) White to buff-yelloAV with chestnut bands and lines.

38. P. PORCELLANA (Newcomb) . PI. 19, figs. 1 to 4.

" Shell rather solid, conical, polished, shining. Whorls
5, rounded, margined above. Aperture ovate; lip expanded,
thin at the margin, thickened within. Columella short, tuber-
cular. Color of a porcelain whiteness with a light fawn-

70 PARTULINA, MAUI.

colored 'band encircling each whorl, the last whorl having
two. Length 12, diam. six-twentieths of an inch ' (Newc.).

East Maui (Neivcomb) : Huelo, Makawao to Nahiku
(Thiving) ; Nahiku (Baldwin).

Achatinella porcellana NEWC., P. Z. S., 1853, p. 146, pi. 23,
f. 47. — PFR., Monogr. iv, 525. — THWING, Orig. Descriptions,
p. 105, pi. 2, f. 12.

In a specimen from Newcomb the first whorl is white, a
yellow band beginning on the second below the suture; but
in others seen the apex is brown or blackish, a band of the
same color \y2 whorls long running from it above the suture,
fading into white at its upper edge, the first \l/2 whorls being
therefore bicolored. Then a yellow band appears faintly
below the middle of the whorl, finally becoming the supra-
peripheral band on the last whorl. In some specimens a
band bordering the suture below also is found. The last
whorl has two to four bands: (1) the subsutural, often want-
ing; (2) one above the periphery; (3) one below it, entering
the aperture at its upper angle, and sometimes there is (4) a
narrow crescent around the columella. Occasionally a few
spiral lines are developd in addition to these bands. These
bands and lines vary from ochraceous to dark brown (mummy
brown of Ridgway). The ground color is white, with a yellow
tint around the columella or over the whole base. Surface
glossy, with only very weak spiral striation. The lip is nar-
rowly thickened within in adult shells, its face tinted or
spotted at the bands. Columella white or brown. Specimens
from Nahiku (pi. 19, figs. 1-3) measure:

Length 18, diam. 10 mm. ; 6 whorls.

Length 18, diam. 10.5 mm. ; 5l/2 whorls.

Length 15.5, diam. 9 mm.

All the shells I have seen are dextral. A. flemingi and
wailuaensis are apparently varieties of porcellana, as Mr.
Thwing has intimated. Very likely A. fulvicans, possibly
A. nattii also, will eventually be included in porcellana.

Fig. 4 is a copy of Newcomb 's figure of porcellana.

PARTULINA, MAUL 71

38a. P. porcellana flemingi Baldwin. PI. 19, figs. 5 to 11.

Shell dextral, minutely perforate, rather solid, high-conic,
glossy, with very weak traces of spiral striation 'or almost
none ; variously colored ; white, Landless or with one to four
dark brown bands, or light fleshy-brown with bands. Lip a
little thickened within; columellar fold moderately strong.

Length 20, diam. 10 mm. ; 6^ whorls.

Length 18, diam. 9.2 mm.

Length 18.2, diam. 9 mm.

Length 18.3, diam. 10 mm.

Length 17.2, diam. 9.5 mm.

" Animal when extended in motion as long as the shell.
Mantle intense black with a narrow white band encircling the
outer edge. Foot above and below very light-brown, granu-
lated above. Tentacles long and slender, slate color.' (Bald-
win.)

East Maui: Nahiku (D. T. Fleming).

Partulina flemingi BALDWIN, Nautilus, xix, p. Ill, Febru-
ary, 1906.

A large set from the type lot shows great variation in color-
ation.

(1) White, uniform (fig. 8), or yellow tinted at base and
above the suture of the embryo.

(2) White with four dark brown bands, situated below the
suture, above and below the periphery and around the colu-
mella. Embryonic shell white with a yellow band below the
suture or a dark band above the suture, fading upwards;
columella and lip more or less purplish brown. This is the
most abundant color pattern.

(3) White, with bands above and below periphery, apex
lip and columella white.

(4) White, with narrow dark bands at the periphery, in
the middle of the base, and a distance below the suture (fig.
5) ; or having a wide band above the narrow basal band (fig.

ID.

(5) Ground color brownish, bands as in no. 3 (fig. 9).
This subspecies differs from porcellana chiefly by its nar-

72 PARTULINA, MAUI.

rower contour. The system of banding (i. e., positions of the
bands) is the same as in porcellana.

386. P. porcellana wailuaensis (Sykes). PI. 19, figs. 12,
13, 14.

Shell dextral, subperforate, glossy-turrited, rather solid,
very lightly striate, white painted with chestnut lines, marked
at the periphery with a white zone and in the suture a black-
ish-chestnut line. Whorls 5 to 5J^, regularly increasing, con-
vex. Aperture auriform; columellar margin having a mod-
erate brown fold, the right margin acute, parietal callus very
thin. Length 15.5, diam. 8.4 mm. (Sykes).

East Maui : Wailua (Perkins). Hana (Cooper).

Achatinella (Achatinellastrum) ivailuaensis SYKES, Fauna
Hawaiiensis, ii, Mollusca, p. 328, pi. 11, f. 19 (and var., f.
20) , 1900. — Partulina cooperi BALDWIN, Nautilus, xix, p. 135,
April, 1906.

" A very pretty little shell of the group of A. bella Reeve,
of Molokai. A variety also occurred (pi. 19, fig. 14) in which
the banding is almost obsolete save in the suture of the earliest
whorls and in one strong dark band below the periphery '
(Sykes). Fig. 13 is copied from Mr. Sykes 's type figure.

The arrangement of bands is essentially the same as in
porcellana, but the shell is somewhat wider in contour, and
the bands are typically paler and more diffuse. I have not
Been specimens from Wailua.

Mr. Baldwin's P. cooperi from Hana (pi. 19, fig. 12, co-
type) seems to belong to the same race. The shell is pale
buff with a white peripheral belt bordered on both sides by
ochraceous bands; base and upper surface have some diffuse
paler ochrey lines or bands, and there is a darker line along
the suture. Embryonic shell white or whitish with a purple
spiral band above the suture and a yellow line below the
suture. The columella and its fold are purplish-brown, and
the narrow callus within the outer lip either white or purplish
brown. A half-dozen specimens of the type lot, sent by Mr.
Baldwin, show but slight variation.

Length 17, diam. 9.5 mm. (Baldwin).

Length 17.3, diam. 10 mm.

PARTULINA, MAUI. 73

39. P. PORCELLANA FULVICANS Baldwin. PL 19, figs. 15, 16.

" Shell dextral or sinistral, very minutely perforated,
rather thin, acuminately ovate conic, apex subacute; surface
shining, marked with delicate incremental striae, under a lens
exhibiting extremely close, minute decussating spiral striae;
nuclear whorls faintly cross-lined. Color very light yellow,
or sometimes white with one or two light yellowish transverse
lines; a conspicuous dark band near the apex. Whorls 6,
somewhat convex, narrowly margined above; suture well im>-
pressed. Aperture oblique, oval, white within. Peristome
acute, slightly thickened within, columellar margin narrowly
reflexed. Columella terminating in a strong white nexuous
fold. Length 18^, diam. 10^ mm.' (Baldwin).

East Maui: Kipahula Valley, Hana (G. 0. Cooper; D.
Thaanum).

Partulina fulvicans BALDWIN, Nautilus, xix, p. 135, April,
1906.

Topotypes from Mr. Baldwin are figured. Fig. 16 is green-
ish-yellow with the embryo nearly white, without a dark band.
Fig. 15 is white above the periphery, light olive below, the
embryo bicolored, being white, broadly banded with flesh
color above the suture, the band extending to the apex. Other
shells are greenish-yellow, fading upwards to white at the
suture and with a faint light peripheral band, the embryo
having a rather dark olive or brown band. The shell is
glossy, with the Partulina spiral sculpture but faintly devel-
oped. It differs from Partulina crocea by the gloss, faint
sculpture, and differently colored embryonic shell. It is very
closely related to A. porcellana and ivailuaensis from which
it differs only in coloration, the dark bands being absent, and
the aperture white throughout. I do not know whether it
occurs in a pure colony, but presume this is the case. Mr.
Baldwin remarks: " The sinistral variety may prove to be a
distinct species. The molluscan life of this rather secluded
valley has only recently been explored. '

40. P. NATTII (Baldwin et Hartman). PL 20, figs. 8, 9, 10, 11.
" Shell dextral, turbinate, spire half the length; whorls 5,

74 PARTULINA, MAUL

polished, the two last rapidly enlarged and inflated. Suture
impressed, columella yellow, stout and twisted. Color bright
gamboge yellow, with one white and three wide chestnut bands
beneath the suture, the latter visible from within the aper-
ture ; aperture round ovate, white, labium white, slightly
thickened within. Length 16, diam. 10, length aperture 8,
width 5 mm.' (Hartm.)

East Maui: Makawao (Baldwin) ; Makawao to Honomanu
(Baldwin).

A.[chatmella] nattii Baldw. Nobis (Achatinellastrum) ,
HARTMAN, Proc. A. N. S. Phila., 1888, p. 34, pi. 1, f. 3
(Achatinellastrum nealii in explanation of plate, p. 56) . —
THWING, Orig. Descriptions, p. 106.

" This with the preceding, A. porcellana Newc., and A.
flemingi Bald. MS., and also A. wailuaensis Sykes, form a
group very similar in appearance and which run together '
(Thwing).

P. nattii, of which we figure specimens from Baldwin,
differs from porcellana by its stouter contour and the posi-
tions of the bands, which are quite definitely located in 27
specimens examined. There is one brown band a short dis-
tance below the suture, another at the periphery, and a third
about midway between periphery and columella, this one usu-
ally being the widest. The peripheral band shows above th©
suture on the spire, but there is no dark band or margin
along the suture. The bands vary in color from chestnut to
ochraceous. The ground may be white throughout, but it is
usually snow-white between the suture and upper band, and
elsewhere gamboge yellow. The embryonic shell is rarely
white, usually some shade of brownish-yellow with a white
line or band below the suture. About the end of the second
whorl this solid color splits into two bands which become the
upper and peripheral.

The embryo and early whorls are quite distinctly engraved
spirally, but on the last whorl this sculpture is very weak.
The surface is glossy. The lip is white except where colored
by the bands, and the strong, oblique columellar fold is in-
variably white. Sutural margin usually set off by an im-
pressed line.

PARTULINA, MAUI. 75

Length 16.8, diam. 11, aperture 9 mm. ; 6^/3 whorls.

Length 18, diam. 11, aperture 9 mm. ; 5^ whorls.

While nattii is very closely related to porcelldna, I have
seen no intergrading specimens and therefore allow it to
stand as a species.

A variety in the C. M. Cooke collection from Honomanu
(Baldwin) has light ochraceous bands on a gamboge ground
and no snowy sutural band.

41. P. ANCEYANA (Baldwin). PI. 20, figs. 12, 13, 14.

Shell dextral, imperforate, rather thin, oblong-conic, the
summit obtuse. Embryonic shell chestnut, fading to, or banded
with white above ; following whorls chestnut with a white line
revolving a short distance below the suture, and on the penult,
whorl another above the suture; last whorl rich yellowish-
chestnut with a white line near the suture, another at the
periphery, the latter bordered below with a yellow band.
Another yellow band, usually wider, occupies the base, leaving
a small dark patch around the axis. The surface is glossy,
with sculpture of fine crenulated spirals throughout. Aper-
ture slightly oblique, brown-banded within ; lip not expanded,
narrowly thickened within. Columellar fold brown or brown
and white.

Length 14.5, diam. 8.5 mm. ; 5 whorls.

Length 14, diam. 9 mm.

Length 15.5, diam. 8.3 mm.

" Animal when extended in motion longer than the shell.
Mantle light-brown mottled with black, outer edge encircled
with an interrupted orange band. Foot above and below a
very light brown, superior portion with a slate tinge, posterior
portion tapering and sometimes with a yellowish tinge. Ten-
tacles long, light-slate. Head above minutely granulated '
(Baldwin).

East Maui : Makawao, in forests of the exterior slope of the
extinct Haleakala crater at an altitude of 4,000 ft., living on
foliage near the tops of the trees (Baldivin). Type no. 65707
A. N. S. P.

Achatinella anceyana Baldwin, Proc. A. N. S. Phila., 1895,

76 PARTULINA, HAUL

p. 223, pi. 10, f. 16.— THWING, Reprint Orig. Descript, Achat,
pi. 2, f. 15.

This species is related to P. porcellana wailuaensis and per-
haps to A. germana, of which it is thought by Dr. Cooke to be
a small race. " It is remarkable for the constancy of its deli-
cate color lines; over 200 examples before me show only
trifling divergence in this respect : (Baldw.). In one speci-
men of the lot sent by Mr. Baldwin, the lower yellow band is
reduced to a line. In some others the ground-color is pale
fleshy-brown above the periphery. There is some variation
in shape, as shown in the figures. It was named for the late
C. F. Ancey. Lives on bark of the guava,

42. P. GERMANA (Newcomb) . PI. 20, fig. 15.

" Shell dextral, ovately conical; whorls 6, flatly convex.
Aperture ovate; lip acute, thickened within. Columella
brown, short and tubercular. Color chestnut, with fine, ob-
solete, darker, transverse lines; a white band traverses the
last sutural whorl and cuts the body- whorl centrally ; around
the columella is a broad white band, losing itself in the aper-
ture. Length sixteen, diam. eight-twentieths inch ' (Newc.).

East Maui: Makawao (Newc.).

Achatinella germana NEWC., P. Z. S., 1853, p. 151, pi. 24,
f. 61, 1854.— PFR., Monogr., iv, 533.— THWING, Orig. Descript.
p. 109. — A. (Achatinellastrum) germana Nc., SYKES, Fauna
Hawaiiensis, p. 324.

" Closely resembles the A. solitaria, and but for the widely-
distant localities might -be taken for varieties of the same
species. The markings and columellaB are however quite dis-
tinct ' (Newc.).

Known to us by the original account only. I can only
suggest its probable relationship to P. anceyana and P. wail-
uaensis, both of which differ in being of a more oblong shape
and smaller size, P. germana measuring length 20, diam. 10
mm. according to Newcomb.

PARTULINA, MOLOKAI. 77

Series of P. mighelsiana. MOLOKAI.

Rather small shells with a high polish, weak or almost
wanting spiral stria tion, and bright coloring — yellow often
with white or chestnut bands, or white or chestnut through-
out; never striped longitudinally; lip unexpanded. Embryo
spirally banded or plain.

These shells are related to the variabilis group of Lanai
and the porcellana group of Maui. Whether they belong to
the Oahuan group Achatinellastrum, or represent a branch of
Partulina convergent towards that Oahuan group, remains
uncertain. In Hyatt's earlier writings he placed part of these
species in Achatinellastrum, but later he doubted this group-
ing, and was disposed to view them as modified Partulinee.
As there are Maui-Molokai types in Oahu, such as Laminella
and Partulina dubia, there seems no good reason why some
Achatinellas of Oahuan type should not occur in the more
eastern islands.

For the present I leave the mighelsiana group in Partulina
for the reason that among the related Mauian species it would
be very difficult to draw the line between Partulina and Acha-
tinellastrum. The coloration of the embryonic shell is more
like that of P. virgulata than like the ordinary Partulinas.
Dr. Oooke suggested that in view of the numerous forms of
intermediate character between the three conventional species
of the mighelsiana group, it would be more logical to rank
bella and polita as subspecies of mighelsiana.

43. P. MIGHELSIANA (Pfeiffer) . PI. 18, figs. 10 to 17.

" Shell ovate-coniform, smooth, opaque, glossy; snow-white
variegated with ashy streaks. Spire conic, the apex rather
acute, suture somewhat margined. Whorls 5^, convex, the
last encircled with a blackish line (sometimes doubled) below
the middle. Columellar fold strong, tooth-like, chestnut col-
ored at the base. Aperture semi-oval, brown-margined ; peris-
tome simple, acute. Length 17, diam. 8 mm., aperture 8T/2
mm. long, 4 wide (Pfr.).

Molokai (Cuming coll.) : Kalae (Baldwin, Thaanum) ; Kilo-

78 PARTULINA, MOLOKAI.

hana (Cooke and Pilsbry) ; Kalae, Puunea, Iloli, Waileia,
Maunahui and Kaanaola (Borcherding, coll. Meyer).

Achatinella mighelsiana PFR., P. Z. S., 1847, p. 231; Mon-
ogr. ii, 238, 1848. — REEVED Conch. Icon., vi, pi. 5, f. 40, April,
1850. — NEWCOMB, Ann. Lye. N. H., vi, 319 (animal). — »
SYKES, Fauna Hawaiiensis, p. 314. — BALDWIN, Catalogue, p.
6.— GWATKIN, Proc. A. N. S. Phila., 1895, p. 238 (radula).—
THWING, Orig. Descript., p. 97. — Achatinellastrum miglielsi-
ana Pfr., BORCHERDING, Zoologica, xix, p. 79, pi. 7, f. 1-13,
15, 17, 19 ; with var. martensi, p. 80, pi. 7, f . 14, 16, 18, 20.

The shell is dextral, perforate, very smooth and glossy, por-
celain-like, showing weak spiral striae under a lens. The
typical form is white with narrow gray streaks and a black-
brown belt just below the periphery, and a narrow blackish
submargin in the lip. The embryonic whorls are white (fig.
13). This pattern varies to forms having the ground white
without streaks, or buff, with or without yellow streaks. The
subperipheral belt may be split, or rarely another band de-
fines an umbilical area. Probably Kalae is the type locality.

(6) White, without bands or streaks (fig. 15) Kalae.

(c) White, with or without gray streaks, and variously
banded and lineate (figs. 10-12). This is var. martensi of
Borcherding.

(d) Pale yellow, with several bands, embryonic shell brown
with a white band above.

(e) White, with yellowish-brown streaks interrupted by
spiral white bands (fig. 16).

(f) Similar to the type, but the base is yellow (fig. 17)
Kalae.

Well-grown specimens measure length 19, diam. 10 mm.

Borcherding describes a var. martensi as differing from the
type by: white color, well-impressed suture, the whoris more
convex, encircled by lines either wide or narrow, the last
whorl having the typical broad brown peripheral band, colu-
mella light reddish, the lip never bordered within with brown,
interior fleshy-white. PI. 18, figs. 10-12 represent this color-
form. It is from Kawela. The rounded whorls and deep
suture are equally well marked in some specimens of other

PARTULINA, MOLOKAI. 79

color-patterns. The same color-pattern occurs in Kilohana.
There is a similarly marked form of polita.

43a. P. MIGHELSIANA BELLA (Reeve). PL 23, figs. 2 to 12; pi.
26, f. 13 to I5e.

£ Shell conical, dextral, whorls rather ventricose, columella
with a twisted plait. Olive-yellow, sutures black-brown, then
white-banded, last whorl brown and white banded. The dark
sutural band is sometimes absent in the last whorl, but the
white band appears constant ' (Reeve).

i Animal of a uniform yellowish-white, with slate colored
tentacles ' (Neiucoml)).

Molokai (Cuming coll.), Puunea, on leaves of small trees
(Cooke and Pilsbry), Kalae, Kealia, Kaupelua, Waileia,
Kaunakakai, Maunahui, Hanakalilolilo, Kawela, Ualapue and
Kaluaaha (Borcherding, Meyer coll.).

Achatinella bella REEVE, Conch. Icon., vi, pi. 3, f. 17, April,
1850. — PFEIFER, Monogr. iii, 461. — SYKES, Fauna Hawaiien-
sis, p. 321. — NEWCOMB, Ann. Lye. N. H., vi, p. 316 (animal).
—BALDWIN, Catalogue, p. 6. — THWING, Reprint Orig. De-
script. Achat., pi. 2, f. 5. — Achatinellastrum bella Rve., BOR-
CHERDING, Zoologica, xix, p. 77, pi. 6, f. 11-20.

The shell is dextral in the large number seen. The early
whorls are closely striate spirally, but the last whorl has only
faint traces of spirals on the glossy surface. Columellar fold
very strong. Axis imperforate. The spire is shorter and
stouter than in mighelsiana, but it must be admitted that
some forms, such as color-var. e, approach that species. P.
polita differs chiefly <by its bulging last whorl, narrower spire
and diverse coloration — wanting white bands.

In my opinion, supported by Dr. Cooke, the forms figured
for polita by Borcherding, on his plate 8, figures 1 to 7, per-
tain to bella.

The color-forms are as follows: (a) Typical form; em-
bryonic whorls white or black-tipped; last whorl yellow with
a chestnut band bordering the suture followed by a white
band, another just below the periphery with a white band
above it; columella fleshy-brown, the tooth and aperture

80 PARTULINA, MOLOKAI.

fleshy-pink Length 16, diam. 9 to 10 mm. See pi. 23, fig. 12
(copy from Reeve). Embryos of this form show very pala
wax-yellow 'bands proceeding from a yellow apex, which ap-
parently fade out in the adult stage. Waikolo and Puunea
specimens correspond exactly with Reeve's figure. The orig-
inal locality was probably Kalae, or perhaps Puunea, within
easy walking distance from Kalae. A series collected by
Cooke and Pilsbry in Puunea is figured, pi. 26, figs. 13 to 13c.
A little further north, between Puunea and Kilohana, there
are transition forms to mighelsiana, pi. 26, fig. 14, both color
and shape being intermediate in various examples. This
place is not far from the northern cliff.

(6) Similar to the above, but lacking the peripheral chest-
nut band. Occurs with the typical pattern.

(c) Both sutural and peripheral bands wanting. Colu-
mella and aperture white or pink.

(d) Variously banded with chestnut, the bands often oc-
cupying most of the surface. Kalae (Meyer).

(e) White, variously banded with chestnut or with a sutural
line only, or no bands whatever, the umbilical region alone
yellow tinted, or sometimes also white.

(/) Bicolored, white above, greenish-yellow or olive below
the periphery. Aperture and columella white or pink. May
be a derivative of forms g or c.

(g) Same as /, but having a dark sutural line, sometimes a
subperipheral line also, and occasionally an olive line in the
middle of the upper surface. Fig. 9. This form is evidently
a derivative of the typical bella pattern.

"Whether the more eastern of Borcherding's localities really
have bella forms I do not know, but the typical bella coun-
try is in the region southward of the Leper Settlement pen-
insula. A series from Ualapue, collected by Mr. Thaanum
(pi. 26, figs. 15 to 15e) seems to carry ~bella into the polita
area. I do not know the locality of pi. 23, figs. 2-8, collected
by Mr. Meyer.

43&. P. MIGHELSIANA POLITA (Newcomb). PI. 24, figs. 1 to 6.
Shell dextral, conic-ovate, polished, light-yellow, a portion

PARTULINA, MOLOKAI. 81

of the upper whorls marked with umber, suture with a broad
black band, and a narrow line of the same color -continued
around the body-whorl. Whorls 5, convex; suture well de-
fined, margined. Aperture oblong-ovate, dark purple ; colu-
mella with a large tubercle tipped with purple ; lip simple.
Length .55, breadth .45 inch (Neiuc.).

Animal of a uniform yellowish- white, including tentacles;
tentacular sheath slightly tinged with brown; mantle black
(Neiucomb) .

Molokai (Newcomb), Kaluaaha to Halawa (Baldwin;
Bishop Museum), Kawela, Kaamola and Ualapue (Borcher-
ding).

Achatinella polita NEWC., Annals of the Lyceum of Nat.
Hist, of N. Y., vi, p. 24, May, 1853; t. c. p. 328 (description
of animal) ; P. Z. S., 1853, p. 142, pi. 23, f. 37.— PFR., Mon-
ogr., iv, 536. — BALDWIN, Catalogue, p. 81. — THWING, Reprints
Orig. Descript. Achat., pi. 2, f. 9. — Achatinellastrum polita
Nc., BORCHERDING, Zoologica, xix, p. 81, pi. 8, f. 8, 9.

The typical polita has the upper 3 whorls bicolored, dark
yellow, chestnut or olive with a white band below the suture ;
following whorls paler, Naples yellow to wax yellow or olive
ocher, with a chocolate band below the suture. This is the
form figured by Newcomb as typical. One of his specimens is
drawn in pi. 24, fig. 2. In his description he mentions a line
of the same color (black) upon the body- whorl. Figs. 3, 4
represent specimens from Newcomb so marked. The aper-
ture is purplish within, varying in shade. Length 13.7, diam.
9 mm., 5l/2 whorls, to length 15, diam. 9.2 mm.

Ne\vcomb 's type locality was doubtless Kaluaaha or the ad-
joining valley east, Mapulehu, where specimens exactly like
the types are found. It is likely that Newcomb stayed with
Mr. Hitchcock, a missionary who lived at Kaluaaha, and had
a place at Kalae. Mapulehu shells are further illustrated in
pi. 24, figs. 5, 6, collected by Mr. Thaanum.

Another form is larger, up to length 18, diam. 10 mm., with
the embryonic 3 whorls white with a minute dusky tip, fol-
lowing whorls yellow (often paler on the last whorl), suture
edged below with a dark line or none. The aperture is of a
purplish tint, darker or dark-streaked near the lip.

82 PARTULINA, MOLOKAI.

P. polita differs from P. ~bella by its bulging last whorl,
which gives the sides of the spire a concave outline. In hello,
the general outlines of the spire are straight. The color-
patterns are also different. P. polita is never white and never
has white bands on the last whorl. It is also more distinctly
striated spirally, the striae visible though often weak on the
last whorl.

Color-form latizona Borcherding. PL 23, fig. 13. Shell
dextral, conic-ova.te, rather solid, glossy, very delicately lon-
gitudinally striate; epidermis pale-yellow, the whorls en-
circled in the middle with a broad chestnut-brown band ; spire
conic; suture margined, impressed. Whorls 5^, weakly con-
vex, the last scarcely broader than the spire. Columella
having a small, short, white tubercle. Aperture oblique, in«
versely auriform, pale violet ; peristome simple, unexpanded,
red-lipped within. Length 14, diam. 8 mm. Kaamola, Meyer
(Borcherding) .

Acliatinellastrum latizona BORCH., Zoologica, xix, Heft 481,
p. 82, pi. 8, f. 10, 1906.

Doctor Cooke collected this form in Kaluaaha and Mapu-
lehu. It is a further development of the ordinary banded
polita.

Color-form dixoni Borcherding. PL 23, figs. 14, 15, 16.

Shell dextral, ovate-conic, rather solid, glossy, deep chest-
nut-brown, the upper whorls and a patch around the umbili-
cus yellow; very delicately striated longitudinally. Spire
conic; suture margined, impressed; whorls 5^, weakly con-
vex, the last rounded, scarcely half the total length of the
shell. Columella having a small short, flesh-red or dirty-
purple tooth. Aperture oblique, half-oval, gray- violet; peri-
stome simple, unexpanded. Length 14, diam. 8 to 9 mm. Ka-
wela and Kaamola, Meyer (Borcherding).

Acliatinellastrum dixoni BORCH., Zoologica, xix, p. 83, pi.
8, f. 11, 12, 13, 14, 1906.

" This species occurs in the above-mentioned localities in
a darker and a lighter coloring. Fig. 14 comes from Kawela,
has a dark columellar tooth and no yellow patch around the
umbilicus. Others from Kawela and fig. 15 from Kaamola

PARTULINA, LANAI. 83*

are normally colored. Fig. 16 from Kawela shows the lighter
coloration, in which there are usually two yellowish bands,
one at the periphery, the other below the suture ' (Borcher-
ding).

Merely the culmination of the dark phase of polita. Simi-
lar shells have been taken by Doctor Cooke on Mapulehu
ridge.

Color-form hepatica Borcherding. PI. 23, figs. 17, 18.

Shell dextral, long ovate, rather strong, glossy, with lon-
gitudinal and spiral lines, visible under the lens only, very
fine and not decussating. Surface yellow-greenish, liver-
brown, veined with darker streaks. Spire turrited; suture
distinctly margined, impressed. Whorls 5^, convex, the last
a little wider than spire, rounded. Columella weakly!
twisted ; columellar fold white. Aperture oblique, half-ovate,
whitish within; peristome simple, unexpanded. Length 17,
diam. 9 mm. (Borch.).

Molokai: Kawela (fig. 17), Waileia (fig. 18).

Achatinellastrum hepaticum BORCHERDING, Zoologica, xix,
p. 83, pi. 8, f. 15, 16.

The largest in Dr. Cooke 's collection measures length 17,
diam. 11 mm. Others taken by Father Dutton are smaller.
The interior is white or pale purple.

Series of P. variabilis. LANAI.

44. P. VARIABILIS (Newcomb). PI. 21, fig. 14; pi. 22, figs. 1
to 12.

" Shell sinistral or dextral, conically acuminate, polished,
shining; whorls 6, rounded, lightly margined above. Aper-
ture roundly ovate; columella short, armed with a strong
tubercle of a roseate color ; lip expanded, acute. Color of
shell white, yellow, black and other shades, plain or with
from one to six bands of various colors on the last whorl.
Length 8, width 14-twentieths inch ' (Newc.).

Lanai (Newcomb, Thaanum and others) ; Windward side,
on ridges facing Maui, above Waiapaa, behind Koele, and
Lanaihale (Perkins).

Achatinella veriabilis NEWC., P. Z. S., 1853, p. 154, pi. 24,

84 PARTULINA, LANAI.

f. 70 (1854) ; Ann. Lye., vi, p. 333.— PFR., Monogr., iv, 256;
vi, 169. — SYKES, Fauna Hawaiiensis, p. 319. — THWING, Re-
print Orig. Descript. Achat., pi. 2, f. 11. — Achatinella fulva
Newcomb, PFR., P. Z. S., 1855, p. 208 (1856).— Achatinella
lactea GULICK, Ann. Lye. vi, p. 198, 1856.

Dr. Newcomb included several color-forms in his original
description, but the dextral shell with several blackish- chest-
nut bands and an umbilical patch, depicted in his figure, may
be taken as the typical form. It is copied in my pi. 21, fig.
14. In the lot given by him to the Academy there are two
dextral and nine sinistral shells, three having a dark um-
bilical patch, the others white there. Fully adult shells are
rather pot-bellied, the base being notably convex (especially
as viewed dorsally), but in some shells with thickened lip
the juvenile character of an acute peripheral angle is re-
tained for a short distance in front of the aperture. Three of
the Newcomb lot are figured, pi. 22, figs. 1, 2, 3, to show the
variation of shape and pattern.

A lot of 48 shells, from Gulick, all sinistral, shows substan-
tially the same forms and banding (pi. 22, figs. 4 to 9) . The
bands are either deep chocolate or ochraceous, and vary from
rather wide to linear. As in the Newcomb set, some shells
are quite narrow, the dimensions being about as follows :

Length 18, diam. 10 mm., average specimen.

Length 15, diam. 10 mm., rare broad form.

Length 15.3, diam. 8.5 mm., narrow form.

Four in the lot are brownish- tawny with extremely weak
bands or none; the rest are banded with chocolate and with
ochraceous in about equal numbers.

Another lot from " Large Valley, Lanai," Gulick, contains
11 shells all dextral, color white, with ochraceous or dark
chestnut bands. A lot of 7 from Mr. D. Thaanum is also en-
tirely dextral, beautifully polished and chocolate-banded.

Another lot of 58 shells from Mr. Thaanum is polymorphic,
(a) Three-banded with dark chestnut or ochraceous, the um-
bilical region always white, sometimes with a few accessory
lines, 25 specimens, all sinistral, fig. 10. (&) bands reduced
to lines or split, the upper one often wanting, 10 specimens,

PARTULINA, LANAI. 85

all sinistral. (c) White, with a faint yellow or brown stain
within the lip- callus, 7 shells, one of them sinistral. Resem-
bles lactea except in the paler aperture. Fig. 11. (d) Cream
colored, darker towards the base, to ochraceous with darker
streaks, bandless, the peripheral angle usually conspicuous,
but wanting in 7 shells; 16 specimens, six of them sinistral,
pi. 22, fig. 12. Of 4 embryos taken from a banded shell,
fig. 10, three are banded as in pi. 21, fig. 16, the fourth,
fig. 17, being ochraceous with a white band below the periph-
ery, which is more distinctly carinate than in the others.
Of 5 embryos from a pure white shell, fig. 11, three are dis-
tinctly ochraceous, one having a white band below the
periphery ; the others being white.

It appears that some colonies are exclusively sinistral,
others wholly dextral, the same color-patterns running
through both. Other colonies contain both sinistral and dex-
tral forms, and a variety of color-patterns, which are unlike
in the proportions of sinistral and dextral individuals. In
mixed colonies, left-handed coil and banding seem to be
dominant, right-handedness and plain color recessive. The
diversity of young taken from one parent indicates hybridity
between the color-forms, but data are wanting to show
whether sinistral and dextral forms hybridize.

The three-banded type, with bands placed as in the Mauian
P. nattiif is probably the most fundamental (or nearest to the
ancestral) pattern.

A. fulva (' Newcomb ' Pfr.), pi. 22, figs. 13, 14, is not dis-
tinguishable from the tawny-ochraceous form occurring in
colonies of variabilis. In sonie specimens the periphery is
acute in front of the aperture, but in others more accelerated
it is rounded. The original description follows : Shell sub-
perforate, turrite-conic, solid, striatulate, and under a lens
very delicately decussate, glossy, buff-tawny; spire sub-
concavely conic, the apex white, acute, suture thread-mar-
gined; whorls §l/2, the upper fiat, following ones convex, the
last a little over one-third the total length, swollen below the
suture, base sac-like. Aperture oblique, subtetragonal-auri-
form, columellar fold superior, nodiform, orange colored;

86 PARTULINA, LANAI.

peristoine unexpanded, labiate within, the right margin some-
what straightened, columellar margin dilated, vaultingly re-
flexed. Length 18^, diam. 9, aperture 8x83^ mm. Sand-
wich Is. (Pfr.) .

Dr. Newcornb and all more recent authors have agreed in
considering A. fulva a synonym of variabilis. Specimens
measure: Length 17 to 18, diam. 10, aperture 8.3 mm. The
lip-callus is white, or brown with a white lip-edge. The widest
specimens approach semicarinata, but none are so wide as that.
It is sometimes cream-white.

Mr. Thaanum found P. variabilis lower, P. semicarinata^
higher on the main ridge of Lanai.

44a. Var. lactea Gulick) . PI. 22, figs. 15, 16.

" Shell sinistral, perforate, acuminately ovate, solid, shin-
ing, finely striated, and microscopically very finely decus-
sated, color ivory white ; apex acute ; spire conical, with out-
lines slightly convex; suture rnarginate, slightly impressed;
whorls 6y2, flatly convex; columellar fold central, brown,
strong ; aperture oblique, sinuately oval, reddish brown
within ; peristome white, thickened within ; with external mar-
gin arcuate, slightly reflected anteriorly; columellar margin
dilated, slightly detached; parietal margin thin. Length 22,
breadth 11% ; length of body- whorl 15 mm. Weight 8
grains ' (Gulick).

Lanai : S. T. Alexander, D. Thaanum.

The shell is rather snow-white than ivory-white, the colu-
rnella brown, tip of the fold paler, lip-callus purple-brown,
interior lilac. The spire is straightly conic, whorls but very
slightly convex. It is minutely but constantly perforate. It
is a very beautiful shell, differing from the white form oc-
curring in some variabilis colonies by its darker aperture. In
a series of about fifty before me, all are sinistral.

Length 18 to 19, diam. 10 mm.

Length 16.8, diam. 9 mm.

45. P. SEMICARINATA (Newcomb). PI. 21, figs. 1 to 4, 13, 15.

" Shell dextrorsal, solid, ovate'ly-conical, longitudinally

substriated ; whorls 5, flattened, margin-ate above the last, an-

PARTULINA, LANAI. 87

gulated in the middle and semicarinated. Aperture ovate;
columella ending in a tortuous, obtuse, dentiform plait ; outer
lip anteriorly produced, strongly marginate within. Color
pale yellow, aperture white internally. A straw-colored, coni-
cal species, with the last whorl partially surrounded with an
elevated keel : (Newcomb). No dimensions given.

Lanai (Newcomb, Thaanum et al.) ; mountains at high ele-
vations (Perkins) ; at the crest of the main ridge, extending a
little way down (Thaanum).

Achatinella semicarinata NEWC., P. Z. S., 1853, p. 156, pi.
24, f. 76, 1864. — PFR., Monogr., iv, 532. — Achatinella varia-
bilis var. semicarinata SYKES, Fauna Hawaiiensis, p. 320.

This species differs from P. variabilis fulva by its decidedly
broader contour and dextral coil. The glossy surface shows
only weak traces of spiral striation. The aperture, lip-rib
and columella are white, or in the darkest shells, the interior
is faintly pink tinted. The early whorls are the color of the
shell in most specimens, but in some they show distinct dark
bands. Embryo shells usually show bands, though not always
so distinct as in fig. 15, an embryo from a cream-colored
mother. The color of adults varies from white to tawny or
light OTange-rufous. The acute carina may extend half
way around the last whorl, or it may not be developed at all
there, ending on the penultimate whorl. Usually it extends
part way across the front of the shell. The axis is perforate.
In a series of 23 from Thaanum and Baldwin, all are dextral.
Neweomb's type figure is copied in pi. 21, fig. 13. It repre-
sents the most strongly carinate form. The size varies.

Length 18, diam. 12 mm.

Length 17.5, diam. 11 mm.

Length 16, diam. 11 mm.

Length 15, diam. 9.5 mm.

Mr. Sykes ranked semicarinata as a variety of variabilis
on account of the difficulty he experienced in endeavoring to
separate it from fulva. " Mr. Perkins remarks that ' the
broader form with ridge more raised is from higher eleva-
tions.' It appears to be gradually replaced by the form fulva,
at lower altitudes.' I have here retained semicarinata as a

PARTULINA, LANAI.

species because in the large series of both species seen the in-
tergrading specimens are comparatively very rare. Nearest to
Koela the white form occurs, further along the ridge the
darker forms. Either sinistral or dextral forms may pre-
dominate in different colonies. Fig. 13 is copied from New-
comb's type figure. Figs. 1-4, 15 are from specimens taken
by Mr. Thaanum.

45a. Var. hayseldeni Baldwin. PI. 21, figs. 5 to 12.

" Shell sinistral, minutely perforated, rather solid, ovately
conical, apex subacute ; surface shining, marked with delicate
incremental striae, and under a lens exhibiting very close,
minute, decussating spiral lines; embryonic whorls faintly
cross-lined. Color generally of a uniform reddish-brown;
sometimes the coloring of the middle portion of the whorl
shades into white on the apical whorls, and in some examples
a white line revolves below the suture. Whorls 5^, slightly
convex, narrowly margined above, the last carinated or angu-
lated at the periphery, the angle becoming almost obsolete
towards the aperture ; suture distinctly impressed and often
margined above by the continuation of- the peripheral keel.
Aperture oblique, subovate, white within with a pinkish tinge ;
peristome white, rather obtuse, thickened within, the basal
and columellar margins slightly reflexed; columella termin-
ating in a strong, flexuous, white fold. Length Yll/2, diam.
10 mm.

" Animal when extended in motion longer than the shell.
Mantle slate color with a brown band encircling the outer
edge. Foot above and below almost white with a yellowish
tinge. Tentacles white tinged with slate ' (Baldwin).

Lanai (Walter H. Hayselden) ; Lanaiha'le, near highest
point of mountain (Perkins, Thaanum).

Partulina hayseldeni BALDWIN, Nautilus, x, p. 31, July,
1896. — Achatinella (Partulina) hayseldeni Baldwin, SYKES,
Fauna Hawaiiensis, p. 313, pi. 11, f. 2.

This species is allied to P. semicarinata Newc., which is
found in another district of the same island. The latter is a
light straw-colored, more conical, and invariably dextral

PARTULINA, LANAI. 89

shell. The animals of the two species are somewhat similar,
but sufficiently different to warrant the separation ' (Bald-
win).

It is a local race of the highest peak of Mt. Lanaihale.
The color-patterns of the type lot are shown in pi. 21, figs. 5,
6, 7. It is uniform orange-rufous with the tip of the apex
pale, aperture pink; the same with a white subsutural band;
or in addition to this there may be a white band or bands mid-
way between the sutures on the spire, gradually deepening
on the penult, whorl, to the ordinary ground-color.

In a series of 29 shells, collected by Mr. D. Thaanum, pi. 21,
figs. 8 to 12, 5 -are sinistral. the rest dextral. The sinistral
shells have the color-patterns described above, the last whorl
being uniform orange-rufous. Seven of the dextral shells are
exactly similar in pattern to the sinistral, and they differ from
P. semicarinata only by the deeper shade of color. Ten adult
dextral shells are pale-yellowish flesh-tinted with several or
numerous orange-rufous bands, varying in intensity among
the specimens.

Embryo. — A dextral specimen of the typical orange-rufous
color with white sutural band on the spire only, contained
three embryos, two having a white sutural band, a white
band a short distance below the suture and white columellar
area, the other being multilineate with brown above and
below. A dextral banded snail (pi. 21, fig. 10) contained 3
embryos, two 3-banded with some accessory lines, the other
having numerous weak bands. In both examples the embryos
are dextral.

It seems likely that the ancestral stock of semicarinata was
banded, and typical hayseldeni is a more evolved form in
which the banding has been replaced on the lower whorls or
throughout by the dark color of the bands, while typical pale
semicarinata has lost its bands entirely or on the later whorls.
The whole series is so closely interrelated that I do not think
it divisible into two species. I retain the name hayseldeni in
a varietal sense because it is already in the literature, rather
than from any real distinction, as I believe the intergradation
with semicarinata is practically -complete.

90 PARTULINA, SECTION BALDWINIA.

Section BALDWINIA Ancey.

Baldwinia ANCEY, Proc. Malac. Soc. London, iii, p. 270,
July, 1899.

Shell sinistral (except in P. dubia), perforate, thin, ovate-
acuminate or ovate-conic, with the color and sculpture of
Partulina; lip narrowly expanded, but little thickened ; colu-
mella somewhat twisted but without a callous fold or tooth-
Type A. physa Nc., 1855 = A. confusa Sykes.

Baldwinia differs from Perdicella by its larger size and
enlarged last whorl, giving the shell a conic form. It is no
doubt, like Perdicella, a derivative of Partulina, which has
the same sculpture and color-patterns.

It is quite likely that the Baldwinias of Hawaii, Maui and
Oahu are so many independent derivatives from the Partu-
lina stock, but until this is demonstrated it is simpler to group
them together. In any case the forms of Hawaii and Oahu
are in all probability the descendants of emigrants from the
Maui-Lanai-Molokai center. Whether the degeneration of
the columellar fold was expressed in the original stock is un-
certain, since its degeneration is without doubt a secondary
modification. The presence of Partulinoid forms in Oahu
seems anomalous, but the distribution of Laminella and Ptero-
discus also show that there was a limited amount of inter-
change between the Oahu and the Maui-Lanai-Molokai centers
after the modern groups had become differentiated.

Species of Baldwinia.

1. Hawaiian species.

a. Aperture much more than half the shell's length; shell
white or banded, without axial or oblique color streaks ;
surface smoothish, finely striate, last whorl much in-
flated. P. horneri, no. 47.

a1. Aperture slightly more half the length; last whorl
rather roughly striate ; length 'over 20 mm.

P. confusa, no. 46.

a2. Aperture about half the length ; shell variously banded
or streaked, smoothish; densely striated spirally
throughout, length less than 20 mm. P. physa, no. 48.

PARTULINA, SECTION BALDWINIA. 91

2. Mauian species.

a. Last embryonic whorl finely zigzag-lineate ; later whorls
moderately convex. P. grisea, no. 49.

a1. Last embryonic whorl obliquely striped; later whorls
very convex. P. thaanumiana, no. 50.

3. Caiman species. P. dubia, no. 51.

Mr. H. W. Henshaw, of Washington, D. C., well known as
a trained and acute observer in several departments of sci-
ence, has given the results of his studies upon this group in
the following essay.

OBSERVATIONS ON HAWAIIAN ACHATINELLHXE, BY H. W.

HENSHAW.

"Although much has been written upon the Achatinellidce
of the Hawaiian Islands, comparatively little has appeared
upon the habits of any of the species. During the early part
of the year 1903 (January- April) the writer enjoyed unusual
opportunities for making observations on the three species of
Partulina inhabiting the Island of Hawaii, and the follow-
ing notes pertain thereto.

' * The peculiar interest attaching to these particular species
lies in the fact that they occur only upon the Island of Hawaii,
the youngest of the archipelago, that the three are specifically
quite distinct from each other, that two of the three are, so
far as known, absolutely restricted to limited sections of the
northern end of the island, which is much the oldest, and that
the third is found outside of this district only rarely. Thus,
living in comparatively unoccupied regions and practically
without competition, the three species offer an unusually fav-
orable opportunity for a study of their habits, and especially
of the method of their colonization and dispersal.

" Upon Calm, the metropolis of the Achatinellida proper,
there are so many closely-related species, several often crowded
into the same area, that the facts relating to habits and
means of dispersal are much complicated. An understanding
of the manner in which the species of Achatinella form new
colonies and of the laws of color variation in the group will

92 PARTULINA, SECTION BALDWINIA.

throw a flood of light upon the vexed question of specific re-
lationship. The writer believes that a study of a carefully-
selected series of the one hundred and more species of the
Oahu Achatinellas by a student familiar with the local geo-
graphic conditions would reduce the number by at least one-
half.

Partulina horneri Baldwin.

" This species, described in 1895 from specimens collected
by Mr. J. Lewis Horner, appears to possess a present rather re-
stricted range in the sparsely timbered region above Kukui-
haele, Hamakua, and on the ridges above the Waipio and Wai-
manu valleys, an area of perhaps three or four square miles.
Except for a few specimens obtained above Honakaa (on the
doubtful authority of a Kanaka's statement) and Kukaiau
(introduced or * planted : there by Eugene Horner) ten to
twenty miles distant, this shell has never been found else-
where. This restricted habitat, in a region where the compe-
tition is almost nil, represents all the territory the species has
been able to acquire since the unknown date of its occupancy
of the island. That the period of its occupancy has been con-
siderable appears from the striking unlikeness of the species
to all its congeners.

" Although occasionally living upon ohia (Metrosideros
polymorpha) , kolea (Myrsine lassertiana) and, rarely, upon
kawaao (Bryonia sandwichense) , the species is chiefly con-
fined to a small berry-bearing tree called the ahakea (Bo~bea
elatior) which is rather common at an altitude of 1,800-2,500
feet. This species, and in fact practically all the Achati-
nellas the writer is familiar with, is often found on half-dead
trees, and not; infrequently a thriving colony may inhabit a
tree which boasts of but a handful of foliage. This appears
to indicate that once a colony is established on a tree it per-
sists, as long as it can obtain food, and I have found small
colonies of shells on stubs of trees that apparently had been
dead twenty-five or thirty years.

" This and the following species belong to the sub-genus
Baldwinia of Partulina, and they are much more closely re-

PARTULINA, SECTION BALDWINIA. 93

lated to the shells of Molokai and Maui than to those of the
more distant islands of the group farther north. It is highly
probable, indeed, that the latter island furnished the parent
forms, the channel separating the two islands being less than
twenty miles wide. As species are reckoned in this group,
the Hawaiian forms are distinct enough from any Maui shells,
although the A. physa of Hawaii is rather closely related to
the P. grisea of Maui, both in form and coloration, and the
latter may have been the parent form, or at least descended
from a common ancestor.

" The color of the typical and prevailing form of A. hor-
neri is white with a broad zone of brown around the periph-
ery, thence visible to the apex as a faint line above the suture.
There is also, as noted by Mr. Baldwin, a pure white form of
this species, and a third variety with only the apical whorls
brown banded. To these is to be added a fourth color-variety,
not previously noted, which is white save that the apex, for
a varying distance, is tinged with brown. Of the four color
forms, only the typical shell and the white variety are at all
common ; the other two are rare.

" There is some evidence to prove that formerly, even so
late as ten years ago, the brown-whorled form was much
more abundant than it is now, and many dead shells have been
found on the heights between Waipio and Waimanu valleys
where the living shells are very rare indeed (some living ones
have been found by Mr. Thaanum) .

" It is to be noted particularly that all the above forms of
horneri occur in the same locality and on the same species of
trees, but individual trees are never 'occupied by two forms.
Moreover, with the possible exception of the banded form and
the pure white one, the color varieties appear not to inter-
grade.

' * As regards shape the four forms do not seem to differ ap-
preciably. Roughly speaking there may be said to be a large
and a small form of each of the four, but it is of interest to
note that, while the individuals of each form intergrade in
size, large and small forms do not live upon the same tree.
The adult individuals on a tree are either all large or all

94 PARTULINA, SECTION BALDWINIA.

small. Thus the isolation of a few feet seems sufficient to
permit recognizable changes both of size and color, though
not of form.

" As these four color-forms occur in the same locality and
are only to be distinguished by pattern of the shell, it might
naturally be inferred that the embryonic young of the four
would be of a common type, perhaps reverting in color to an
ancestral form, and in any event not perpetuating the color
peculiarities of their respective parents. The contrary is
true. The young are not of a common type, but are always
distinguishable inter se. Though markedly different in the
adult state, there is less difference between the young of the
white and the banded form than of the other two, but there
is still a recognizable difference. The embryonic young of the
pure white form retain the peripheral brown band of the
typical horneri until about a third grown when only traces
of it are discernible near the aperture and these finally en-
tirely disappear leaving the shell pure white. The embryonic
shells of the other two forms differ more markedly from each
other and from the young of the other forms.

The unlikeness of the respective young of several mol-
luscan forms would appear to be an excellent criterion of
their permanent distinctness, and the writer is inclined to
view these two forms as far better entitled to specific rank
than many named species of the group whose claim to dis-
tinction rests upon equally slight color characters supple-
mented, it is true in some cases, by present geographical
separation of habitats.

" It is an interesting fact that the embryonic young of the
brown-tipped form differ more from the young of the others
than do the parent shells. The adult of the brown-tipped var-
iety is to be distinguished only by the light-brown color of
the apical whorls. The embryo shells have the lower half of
the basal whorl of a deep reddish-brown. Evidently the brown
tends to fade out as the shell matures. It is probable that in
time the deep brown apex will become a permanent character
of the adult of this form as it now is an invariable character
of the young.

PARTULINA, SECTION BALDWINIA. 05

"It is probable enough, too, that in time the four color
sports, if indeed two of them are not already properly to be
ranked a;s species, will become completely isolated geographic-
ally when the present differences will be accented and per-
haps further differentiation occur. No doubt many of the>
recognized species of the group have had origins and life his-
tories essentially similar to the forms here noticed.

' How long these forms of Jiorneri have been in reaching
their present degree of differentiation cannot, of course, be
told. Possibly they are nearly as old as the life of the species
on this island; but in the case of creatures as plastic as the
Acliatinellidce appear to be, probably no very long interval,
nor very complete isolation, is requisite to establish such var-
iations.

Partulina pliysa Newc. (hawaiiensis Baldwin).

" This shell was described by Mr. Baldwin from the same
locality as the preceding, and its vertical range is about the
same. It lives chiefly upon the smooth-leaved variety of the
ohia but is found also of the kolea and casually on the haa
(Antidesma platypJiyllum Mann).

" Though this species varies much in coloration and size,
there cannot be said to be any constant and well-defined color
varieties, all the forms appearing to intergrade without limit.
As in the case of the previous species, however, every colony,
and sometimes the shells from individual trees within the
confines of a colony, possess peculiarities of their own — slight
differences of color and size, often sufficient to the eye of a
trained collector to distinguish them from those of ether
'colonies. The tendency to differentiate is the same In kind,
but for some reason or other is has not progressed so far as
in the case of horneri ; possibly isolation has not been so com-
plete as in the other species, although observation does not
confirm this.

The embryos of this species tend to follow the coloration
of the parent form to some extent, especially when the latter
are deep brown, but in most cases their colors do not indicate
their parentage with any certainty.

96 PARTULINA, SECTION BALDWINIA.

" This species has been far more successful in extending
its range than the previous one, and is found in three small
and isolated colonies far beyond the confines of Hamakua,
which probably was its starting point on the island. The
Partulina physa errans from Olaa, Kaiwiki and Puna, from
forty to sixty miles distant and separated by innumerable
gulches and water-courses, is a derivative of this species if
indeed, as the author believes, the variation is not entirely
compatible with specific identity.

" The author has learned of a tree shell which many years
ago lived abundantly on the ohias in South Kona, above and
not far from Kealakekua Bay, and which now appears to be
extinct there. If, as is probable from its description, this
shell was physa (haivaiiensis) , the species has nearly encircled
the island of Hawaii, affording a marked contrast to the re-
stricted range enjoyed by most of the species of this family,
especially upon Oahu.

" It is to be remarked that this species, like the preceding,
'occurs chiefly in sparsely wooded districts and upon isolated
trees. It is difficult to understand the wide dispersal of this
particular species and especially its speedy appearance in
clearings after the cutting of the forests, unless individuals
exist here and there in the deep and virgin forest, where, how-
ever, the shell has never been found. Deep forest tracts ap-
pear to be inimical to the welfare, not only of the Achati-
nellas proper, but to almost all other Hawaiian land shells.

Partulina confusa Sykes (physa of authors).

This species probably was formerly much more wide-
spread in the districts of Kohala and Hamakua than it is
to-day, owing to the present restriction of the forested area.
It occurs at a higher altitude than the two preceding shells
and extends at least as high as the upper edge of the Waimea
Plains (3,500 feet) over most of which area it formerly oc-
curred, although, like the preceding species, it is doubtful if
it ever lived in the deep forest except casually.

' ' During the spring of 1903 the author had the opportunity
of examining an isolated colony of this species on the Waimea

PARTULINA, SECTION BALDWINIA. 97

Plains, which for its extent and peculiar character deserves
more than passing notice.

The P. confusa lives chiefly upon the pua tree (Olea
sandwichensis) although in some localities it has been found
abundant upon the ilima (Sida sp.), a low shrub with yellow
flowers. It occurs also upon the mamani (Sophora chryso-
phylla) . Whether from the fact that the pua is not common
over the island of Hawaii, or for some other reason, I did not
find this shell, which is far more abundant than either of the
other species, outside of the above region (Mr. D. Thaanum
has found it widely spread in the Waimea plains) .

The colony in question occupies perhaps 150 pua trees,
which cover an area of perhaps half a mile square, many
trees being more or less widely separated from their fellows,
The pua is a small tree fifteen or twenty feet high and with a
small spread of branches. A rough estimate of the number
of adult shells inhabiting this area when first visited is more
than 75,000 shells, and it was possible to ride under the trees
and from their trunks, leaves, and branches to pick shells
literally by the handfuls. Cavities in the trunks and branches
were usually packed with shells, mostly immature, from 50 to
75 being often found together. Wherever there were logs,
dead branches or large rocks under the trees, they, too, har-
bored large colonies of shells, and the presence of young in
numbers showed that these places were their permanent
abode.

' ' In six hours, without climbing a tree, the author had no
difficulty in gathering 1,100 adult shells, and his companion,
Mr. William Horner, did the like — a statement which will suf-
ficiently attest the abundance of the mollusks. The locality
becoming known to several shell collectors, something like
10,000 adult shells were gathered in the course of threes
months without seriously diminishing the numerical strength
of the colony. It is doubtful if so many tree shells of any
one species were ever found in a like area, even in the Ha-
waiian islands, abundant as the Achatinellas are, or used to
be, in some places.

" The colony in question comprised numerous color varie-

98 PARTULINA, SECTION BALDWINIA.

ties and the shells differ much in shape. Mr. Homer, of Ku-
kuihaele, has succeeded in selecting from several thousand
no fewer than 200 and odd more or less distinct varieties.
For the present purpose of the author it will suffice to enum-
erate a few only of the more marked forms in the author's
own collection.

1. Uniform light gray, with or without faint brown
markings on apical whorls.

' 2. Distinct brownish-gray with more or less distinct mark-
ings on apical whorls. Undoubtedly these two forms, the
second being but a slight variation of the first, are to be con-
sidered as the typical ones of the colony; at a rough guess
one-half of all would come under one or the other.

' 3. Dark brown, almost 'chestnut, with lighter colored
apical whorls; lower whorl sometimes with, sometimes with-
out, gray spots.

" 4. Light gray with broad or narrow light-brown band
across body whorl.

' 5. Light gray with broad white band across body whorl,
bordered with narrow brown band.

: 6. Light gray, more or less streaked and spotted with
brown; a distinct white band on body- whorl bordered by a
chestnut band.

" 7. Light gray, with chestnut body- whorl bordered above
and below with white.

" 8. Many-banded with alternate bands of brown and
white from apex to base, some narrow, some broad.

' 9. Dwarfed and somewhat distorted form, running
through all the above variations, and found only on mamani
trees (Sophora chrysophylla) .

[Plate 16, figs. 2 to 12, represent shells from this colony.]

" The exact size and shape of the above varieties vary
almost interminably and, it is perhaps needless to say, inter-
grade with each other completely.

" The above brief notes afford but little idea of the great
variety and apparent distinctness of some of the color forms.
Many species of the genus AcJiatinella appear in printed lists
which are based upon color characters much less marked than

PARTULINA, SECTION BALDWINIA. 99

those which distinguish this series. The number of forms
might be multiplied almost indefinitely, but the -above will
suffice to indicate the general character of the variations ex-
isting within this single colony.

11 As in the case of P. horneri, a distinct tendency is ob-
servable, though by no means so marked, to the segregation of
the different color varieties upon individual trees, indicating
that isolation has been sufficient to permit differentiation
within certain limits, while the fact that many trees support
a sprinkling of other varities, added to the completeness of
the intergradation, proves that intercommunication has been
to some extent potent in limiting the amount of the diver-
gence.

" The following facts bear upon the question of intercom-
munication between the several parts of the colony. The pua
and mamani trees, upon which the shells live, rarely touch
each other, and are usually separated by a distance varying
from a few feet to a hundred yards. The land is pasture
and the grass under many of the trees — not in the open — is
deep. There is thus no chance for the shells to pass freely
from tree to tree.

" Upon Oahu and the other islands of the group deep
valleys and high ridges in most localities tend to the more or
less complete isolation of shell colonies, and also present
diverse conditions of foliage and environment favorable to
the differentiation of new forms. The environment of the
present colony on the contrary is exceedingly uniform and
hence presumably unfavorable to the origin of new varieties,
but the isolation of the trees and the open nature of the
ground form barriers to free intercourse quite as effective
as ridges and valleys, and hence tend to the perpetuation of
any chance varieties that may arise.

The author is aware that some observers 'consider the
Achatinellidcc to be active and rather extensive travelers, but
his study of the three species under consideration points to
the opposite conclusion. The fact, as noted above, that logs
and rocks under the trees form the permanent abode of such,
shells as chance to fall from above, seems to show that the

100 PARTULINA, SECTION BALDWINIA.

shells from some cause are unable to find the way back to the
tree on which they were born though but a few feet distant.
A favorite resting place of the Achatinellas is on the under
side of leaves, especially dead leaves, and as these fall or are
blown off by high winds they naturally carry the shells with
them — usually to the ground immediately beneath the tree.
In a number of instances shells that had shaken from the
tree into the grass from six to twelve feet away from the
trunk were found where they had fallen several weeks later,
being apparently quite lost. That Achatinellas can travel
twelve or fifteen feet over a tree in a single night there can
be no doubt, since the author has observed them in the act
and has measured the distance traveled; no doubt they arc
able to travel much farther than this, as certainly can the
Amastras and the apparently still weaker Succineas. It
seems likely, therefore, that natural sluggishness or defective
vision, perhaps both, are responsible for the failure of such
shells as fall or are blown from trees to find their way back
to the parent trunk. "Whatever the explanation, the fact that
the three species of Achatinellas here considered are as a
rule very sedentary and rarely or never regain their place on
a particular tree when once dislodged seems to the author in-
disputable.

" The question then arises as to the means of dispersal
from tree to tree and from locality to locality possessed by
shells under 'Conditions similar to those here indicated. The
author is convinced that the dispersal of the three species
here mentioned is effected chiefly through the agency of wind
when the shells are young. In several instances he has found a
single young Partulina, but a few days old, to be the sole oc-
'Cupant of a shrub or tree (so small as to be readily examined
leaf by leaf) separated several hundred yards from the near-
est shell-bearing trees. In such cases no other transporting
agency suggests itself but the wind or birds.

" It would require a considerable gale to carry to a dis-
tance a leaf laden with an adult Acliatinella, but a very mod-
erate wind would suffice to <carry a leaf with a young one
which weighs but a gramme or two for a considerable dis-

PARTULINA, SECTION BALDWINIA. 101

tance. An adult shell once upon an isolated tree, one that is
separated from its fellows even by a few yards,, might live
and die leaving its progeny, if it chanced to 'be pregnant, to
perpetuate its peculiarities, subject only to the modifying
influences of such individuals ais by rare chance might be
blown to the same tree.

Where shell-inhabited trees of the same species interlock
branches a more or less free interchange of inhabitants would
naturally take place, limited only by the natural sluggishness
of the mollusks, and, as a matter of observation, the forms of
P. confusa inhabiting closely adjacent pua trees were found
to be practically indistinguishable.

' As bearing upon the problem of the variation of this
species, a most suggestive fact was noticed in connection with
the colonies upon the mamani trees. This is a small-leaved,
leguminous tree, and in appearance, and no doubt in fact, is
not well adapted to molluscan life. Yet generally where the
mamanis adjoined the pua trees, and invariably where they
touched branches, the mamanis were found to be occupied by
-colonies of confusa. These shells were almost always smaller
than those of the neighboring pua trees, often had diminutive
and distorted spires, and in every external aspect suggested
the effect of insufficient food or of unfavorable conditions rf
life. Whatever the explanation, the change of habitat from
-one species of tree to another closely adjoining has resulted
in establishing a recognizable form, striking evidence of the
plasticity of the Ackatinellidce, and of the slight nature of
what seem to be insurmountable barriers to free intercourse
between separate, but closely adjoining, molluscan colonies.

" An examination of the embryonic shells of confusa dis-
closes that in a few of the more strongly marked forms the
embryo tends to perpetuate the color peculiarities of the
parent. Thus the young of the gray form are like their par-
ents, and are almost destitute of the zigzag markings which
decorate the spires of almost all the forms, but are often
wanting in adults of the gray variety. So, too, the young
of the brown shell, with brown and white body band, are
•often but not always marked like their parents. But it would

102 PARTULINA, SECTION BALDWINIA.

be impossible to determine with any degree of certainty the
parentage of the greater number of embryonic shells of this
species from their appearance alone — whether they came from,
banded, multi-banded, or unicolored shells. Clearly the color
varieties are not yet fully established.

" It is well known that many of the species of the Achati-
nellidce are indifferently dextral or sinistral. Some, how-
ever, are consistently one or the other, and the P. confusa be-
longs in the latter class. Among upwards of ten thousand
individuals of confusa examined, not a single dextral shell
was found.

' ' Most of the Achatinellidtf proper, as is well-known, dwell
upon trees and the greater number of individuals of the
several species pass most of their time upon the leaves, feed-
ing and moving about only by night and, to a very limited
extent, on dark and rainy days. A certain amount of)
moisture — apparently the more the better — seems indeed nec-
essary to their mode of progression, if not to their very exist-
ence, and in dry weather they are strictly sedentary.

' ' The three species in question dwell indifferently upon the
trunk, the branches, and the leaves of trees, though in day
time the greater number are attached to leaves. As a rule
they appear to be very sluggish and often remain for weeks,
especially when young, attached to leaves and branches with-
out moving and apparently without eating. Individuals in
captivity have been known to live a couple of months without
food or water, which argues the possession of considerable
vitality. Under such circumstances adult shells are known
often to give birth to apparently healthy young.

' ' Unlike the Arnastras and the Succineas, the Acha,tmelUd&
proper do not appear to feed upon the chlorophyl of
leaves; at least the leaves upon which the shells are found
seem never to show visible signs of their work. The two
former groups consume leaf tissue voraciously, both dead
and living foliage, and often leave them completely skeleton-
ized. Both of these groups live also no doubt to a considerable
extent on fungi, as they are often found upon dead wood.
Indeed the presence of Succinea in numbers upon coffee trees,

PARTULINA, SECTION BALDWINIA. 103

for instance, is a pretty sure sign that the trees are no longer
in a healthy condition.

" With a view to determining the nature of the food of
A. confusa, the writer with the assistance of Dr. Nicholas
Russel, examined microscopically the feces of a number of
individuals. In only two instances were minute particles of
chlorophyl found and these, perhaps, were accidental. The
great mass of the remains appeared to be of minute fungi or
algae.

' ' To what extent the fungus-eating habit prevails through-
out the Achatinellidce remains to be determined by future
observations, but from the fact that many other species are
known to reside upon dead trees, or upon the dead branches
of living trees, it may be presumed to be somewhat general if,
indeed, fungi be not the main reliance of the family. In con-
nection with the food of the Achatinellas, it is of interest to
note that in a number of instances small but apparently thriv-
ing colonies were found on dead stubs which had not borne a
green leaf for years, perhaps twenty or more. It would seem
that under such circumstances fungi must be the sole depend-
ence of the shells. As such stubs are often within a few yards
of living trees of the same species, there would at first sight
appear to be no reason why the colonies did not abandon the
trees after they died and make their way to other trees. Ap-
parently, however, the mollusks either do not care to migrate
even a short distance, or are incapable of doing so. - In any
event, their isolation, whether voluntary or involuntary, is
complete, and successive generations of mollusks live and die
on the same tree where they were born, even after the latter
perishes and until it finally crumbles away.

" The ability of the Achatinellas to maintain life for con-
siderable periods without food, and to found colonies and
live indefinitely upon dead trees, greatly favors their chances
in the struggle for existence.

' Many experiments have been tried from time to time in
transplanting the Achatinellas from one locality to another
and even from island to island, but in no instance known to
the author have they been successful. The writer experi-

104 PARTULINA, SECTION 3ALDWINIA.

men ted in this way with the three species here mentioned,
removing perhaps a hundred individuals, old and young,
from an altitude of between 1,600 and 3,000 feet to about 800
feet above and near the sea, where they were placed upon a
magnolia tree. The chances of the colony surviving such a
complete change of environment seemed very small. Never-
theless for a short time the shells seemed to thrive, but they
were discovered by rats and the colony was soon exterminated.

" Great numbers of Achatinellas, as well as Succineas, in
their natural habitat on the several islands are destroyed by
rats and mice, and the extermination of many of the species
is now being hastened by these agencies.

" An Achatinella which Mr. D. D. Baldwin refers to Par-
tulina physa (=s confusa) was found by the writer abun-
dantly in the semi-fossil deposits closely adjoining the colony
of that shell above described. The great size of this fossil
leads to a first impression of its distinctness from confusa,
but a careful comparison of many specimens constrains the
writer to adopt Mr. Baldwin's view, since, as remarked by
that veteran conchologist, a description of the fossil would
read precisely like that of confusa save in respect of size.

" Tables are appended which indicate the degree of pro-
lificness of the species here treated of. The season when the
shells were collected (January- April) was unusually cold and
stormy, and perhaps in mid-summer different ratios might
be had, although Mr. Thaanum. who has dissected many Oahu
species with reference to the number of young, tells me that
he has never noted much if any seasonal difference in this
respect, the shells bearing young the year round.

' In several of the forms the number of shells examined is
too small to give results of much value. In the case of the
brown- whorled Iwrneri, for instance, a greater number would
doubtless materially reduce the average number of young
per adult shell. ' '

PARTULINA, SECTION BALDWINIA.

105

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P. physa (hawaiiensis) . .

P. horneri, brown banded
variety .

247
16

47
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148
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P. horneri, white variety.

P. horneri, brown whorled
variety

33
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149

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46. P. CONFUSA (Sykes). PI. 16, figs. 1 to 12.

; Shell sinistral, pointed at the summit, strongly inflatad
below, rudely striated obliquely. Whorls 6, first five flatly
•convex, the last largely inflated and obsoletely carinated.
Suture simple and deeply impressed. Oolumella short,
slightly callous, broad, and partially covering a deep um-
bilicus. Aperture large, semiovate; lip subreflected below,
thin and simple above. Color of a dingy white, occasionally
marked with yellow flammules, internally of a light-lemon
yellow. Length 18, diam. 12-twentieths of an inch' (Newc.).

Hawaii (Newcomb) ; Kohala (Baldwin) ; near Mana, Ham-
akua (Thaanum) ; Waimea plains; a very large form in fossil
deposits of Mana (Henshaw) ; Waimea side of Kohala Mts.
(L. A. Thurston).

Achatinella physa NEWCOMB, Proc. Boston Soc. Nat. Hist.,
v, p. 218, September, 1855 ; American Journ. Conch., ii, 7).
214, pi. 13, f. 10, 1866.— BALDWIN, Catalogue, p. 7 (Kohala) ;
Proc. A. N. S. Phila., 1895, p. 225 (animal).— THWING, Re-
print Orig. Descript, Achat., pi. 2, f. 21. — HENSHAW, Journal
of Malacology, xi, p. 57, 63, Sept., 1904. — Achatinella (Par-
tulina) confusa SYKES, Fauna Hawaiiensis, ii, p. 312, 1900,
based upon A. physa Newcomb, Sept., 1855. — Partulina

106 PABTULINA, SECTION BALDWINIA.

physa Newc., var. ph&ostoma ANCEY, Proc. Malac. Soc. Lon-
don, vi, p. 121, June, 1904. — Achatinella physa Newc., var.
procera ANCEY, Journ. of Malacol., xi, p. 69 (Palihoukapapa,
in fossil bed).

Doctor Newcomb's second description is reproduced. I
believe with Mr. Sykes that Newcomb's first description
and figure (pi. 17, fig. 6) of A. physa (1854) pertain to the
form subsequently called A. liawaiiensis, which he took to be
the young stage of the larger shell which he afterwards
(1855) obtained and later figured as A. physa. One lot in
the Robert Swift collection (A. N. S. P.) at least 50 years
old, received from Newcomb, contains two ' hawaiiensis }
with others of the large form like Newcomb's figure of 1866.
Dr. C. Montague Cooke and Mr. Thaanum, who have gone
over the evidence and figures with me, agree in this interpre-
tation of Newcomb's A. physa, confirming the opinion of Mr.
Sykes.

The first whorl of the embryo is smooth ; subsequent whorls
are finely striate spirally. The first two whorls are creamy-
brownish, after which there are zigzag brown flames (fig. 12).
These may continue as far as the middle of the fifth whorl,
but they usually disappear earlier. In other embryonic shells
there is a brown band just below the periphery, followed by
a white band (fig. 11). These persist in the adult shell to the
end. In a number of examples with uterine young sent by
Mr. Thaanum, the young shells of one parent are not diverse,
but in every case agree with the mother in pattern. These
observations have been confirmed by the examination of large
series in Mr. Thaanum 's collection. The periphery is acutely
carinate and the columella is sinuous in embryo shells at all
stages examined.

On the last whorl of the adult shell the crowded spiral lines
of the early stages become weak, often almost obsolete, and
wide apart. The color varies as shown in the figures and
described below. The columella is brown or white, nearly
straight or somewhat sinuous. Umbilicus half open or more.

Length 26, diam. 17, aperture 14 mm.; 6^ whorls.

Length 26.2, diam. 15.5, aperture 12.8 mm. ; 6T/2 whorls.

PARTULINA, SECTION BALDWINIA. 107

Length 26.3, diam. 16, aperture 14 mm. (near Mana).

Length 23, diam. 15.2, aperture 12 mm. (near Mana).

Length 23, diam. 13.8, aperture 11.2 mm. (near 'Mana).

Length 27.7, diam. 17.5 mm. (Mana, Bishop Mus., no.
22357).

Var. phcuostoma Ancey. ' ' Shell glossy, pale coffee colored,
the apex white, posterior part tessellated, marked with close
spiral lines and rough growth-striae. Aperture and colu-
mella brown, throat glittering, brown, the edge a little paler.
Length 23, diam. 14.5, oblique length of aperture 11.5 mm.
Hawaii, Thaanum ' (Ancey). The type of phceostoma, now
in coll. Bishop Museum, shows no racial characters whatever
to differentiate it from confusa, and I do not think it a valid
variety or race.

Var. procera Ancey : ' was never denned in any way, but
one may infer that Mr. Ancey 's intention was to name the
large fossil form of the Palihoukapapa bed.

Hon. L. A. Thurston collected a series of confusa on the
Waimea side of the Kohala Mts., including some shells darker
than any from localities farther east.

47. P. HORNERI (Baldwin). PI. 17, figs. 1 to 5.

" Shell sinistral, minutely perforated, thin, globose with a
short acutely conical spire, apex acute ; surface shining, stri-
ated with fine incremental lines, and under a lens exhibiting
very close and delicate decussating spiral striae; embryonic
whorls faintly cross-lined. Color dull white, encircled at the
periphery with a faint brown zone which is continued on the
suture, also with a very small patch of same color around the
umbilicus. Whorls 6, the upper five slightly convex, the last
very much inflated, forming the greater part of the shell;
suture distinctly impressed. Aperture oblique, sub-rotund,
very large, white within, distinctly showing the external
peripheral band ; peristome margined with light brown, rather
thin, very slightly thickened within, expanded, basal and
columellar margins narrowly reflexed, extremities slightly
converging and united by a thin callus; columella light
brown, very slightly developed, plain and smooth. Length
24, diam. 18 mm.

108 PARTULINA, SECTION BALDWINIA.

1 1

Animal in motion longer than the shell. Mantle black,
margined with gray. Foot above and below gray, the superior
portions lightly granulated. Tentacles light gray ' (Bald-
win).

Hawaii : Above Kukuihaele, Hamakua, type loc. ; ridges
above the Waipio and Waimanu valleys ; above Honakaa and
Kukaiau (see p. 92) . Fossil in the Mana deposits.

Achatinella horneri BALDWIN, Proc. A. N. S. Phila,, 1895,
p. 224, pi. 10, figs. 20, 21, 22.— HENSHAW, Journ. of Malac.,
xi, p. 63.

Distinguished from P. confusa by its shorter spire, much
more inflated, broader last whorl, and the less roughened,
more even and shining surface. Moreover, the embryo has no
zigzag-striped stage. Named in honor of Mr. J. Lewis
Horner. The following color-races are found.

(a) PI. 17, fig. 1. White, with a brown band at the periph-
ery, ascending the spire in a narrow line above the suture.
Size large, length 23.5, diam, 18, aperture 14.6 mm. ; 6 whorls.

(ft) PI. 17, figs. 2, 5. Brown, with a narrow white band
below the suture, one above, another below the periphery.
The brown fades on the penultimate or on the last whorl,
which in large part or entirely is white. 23 x 16.4 mm.

(c) PI. 17, figs, 3, 4. White, with a pale brown band im-
mediately below the periphery, which fades out on the fifth
whorl, being visible therefore only in immature shells, the
adult stage pure white throughout. Embryo ivory-white
with a distinct brown subperipheral ba.nd. 22 x 15.5 mm.
Above Kukuihaele, Hamakua,

(d) Young stage light brown, darkest just below the
periphery; adult white with pale brown spire. Embryo bi-
colored, pale brown above, darker brown below the periphery.
20.5 x 15.5 mm. Above Kukuihaele.

These several color-forms are evidently incipient races, as
they O'Ccur in pure colonies, so far as known. Races c and d,
Mr. Thaanum informs me, occur three or four miles apart.
The embryonic forms are much more distinct than the adults,
and are constant in the colonies.

By the loss of zigzag stripes on the embryo and young

PARTULINA, SECTION BALDWINIA. 109

stages, this species is more evolved than other Hawaiian Bald-
winias.

48. P. PHYSA (Newcomb). PI. 17, figs. 6 to 13.

" Shell sinistral, acutely conical, thin, inflated. Whorls 5,
rounded; suture well marked. Aperture widely ovate; colu-
mella thin, slightly twisted ; lip simple. Color light yellowish
brown with white longitudinal flammules, with or without a
subcentral revolving white line on the body-whorl. Length
11, diam. five-twentieths of an inch ' (Newc.).

Hawaii: Mauna Kea (Newcomb); Hamakua (J. Lewis
Homer) ; Waimea Plains, Hamakua. Chiefly on dead trees
but sometimes on living lehua (D. Thaanum).

Achatinella physa NEWC., P. Z. S., 1853, p. 152, pi. 24, f.
64, 1854, — SYKES, Fauna Hawaiiensis, Mollusca, p. 316. — •
Achatinella kawaiiensis BALDWIN, Proc. A. N. S. Phila., 1895,
p. 225, pi. 10, f. 24-26 (Hamakua).— GWATKIN, t. c., p. 238
(radula). — THWING, Reprint Orig. Descript. Achat., 1907,
pi. 2, f. 22.

Smaller and less roughened than P. confusa, and differing
in coloration. Some shells have very much the pattern of the
young stage of confusa. Newcomb 's original description and
figure (copied in pi. 17, fig. 6) are given above. A. hawaiien-
sis Baldwin is figured from the type lot sent by Mr. Baldwin,
pi. 17, figs. 7, 8, 13. It is believed by Mr. Sykes to be iden-
tical with Newcomb 's original A. physa, an opinion in which
I agree, having gone over the matter carefully with Dr. Cooke
and Mr. Thaanum.

A. physa varies widely in color-pattern, as follows:

(a) White, with brown oblique or zigzag streaks above a
narrow peripheral brown band, and having a broad brown
zone on the base (pi. 17, figs. 7, 8).

(&) White, with more or less brown on the spire, the
peripheral brown band and basal zone more or less inter-
rupted (fig. 13).

(c) Brown, indistinctly mottled and streaked with cream-
color, with a subperipheral band of the same or of white (fig.
9).

110 PARTULINA, SECTION BALDWINIA.

(d) Brown, with subperipheral white 'band, and zigzag
white stripes more or less developed (figs. 10 to 12).

Specimens more or less intermediate in pattern connect
these forms, which (figs. 9 to 12, Waimea Plains) live to-
gether in the same colonies. Pattern d changes to a streaked
pattern on the latter part of the last whorl in adult shells
(fig. 12). The embryonic shell is like that of P. confusa in
form except that the columella is more strongly convex. It
is marked with broad vertical stripes of cream-white and
brown, their edges irregular or zigzag.

Length 19.7, diam. 12, aperture 9.8 mm.; whorls 6%.

Length 18, diam. 10.2, aperture 9 mm. ; whorls 6.

Length 17, diam. 10.5, aperture 9 mm. ; whorls 6.

The original description of A. hawaiiensis follows.
' Achatinella hawaiiensis. Shell sinistral, minutely per-
forated, very thin, acutely conical, apex acute ; surface rather
lusterless, covered with, fine lines of growth, and under a lens
showing extremely close and delicate decussating spiral lines ;
nuclear whorls faintly decussated. Color very variable, plain
brown or dingy white, sometimes irregularly striped or
mottled with brown and white, the base generally uniform
brown, but sometimes with undulating markings of brown
and white ; the only constant characters being a brown, some-
times interrupted, line at the periphery, bordered below with
a broader white line. "Whorls 6, slightly convex, the last in-
flated. Suture lightly impressed. Aperture oblique, oval,
brown, the peripheral -brown and white bands distinctly
marked within. Peristome acute, not thickened within, ex-
ternal margin straight, basal expanded, the expanded portion
being very thin and fragile, the columella margin reflexed
over the minute perforation ; color white on both face and the
reverse. Columella. white, very slightly developed, plain ani
smooth. Length 18, diam. 10^ mm.

" Animal when extended in motion longer than the shell.
Mantle almost white, margin of a darker shade. Foot above
and below dingy white, superior portion sometimes flecked
with gray. Tentacles of darker shade. The dentition is the
same as that of the arboreal Achatinellas generally. A cen-

PARTULINA, HAWAII AND MAUI. Ill

tral tooth is present ; and the formula of dentition is
125-1-125 X 120 = 30,120. ' ' (Baldwin) .

48a-. P. physa errans, n. var. PL 17, figs. 14, 15, 16.

The shell is thin, sinistral, more slender than usual in P.
physa (haiuaiiensis) , minutely perforate or almost imperf or-
ate, smooth and slightly shining, under a lens minutely
marked with growth-lines and closely, distinctly striate spir-
ally throughout, the striae rippled. Embryonic shell marked
with broad axial brown stripes as in P. physa ; neanic whorls
marked with brown zigzag stripes on a white or cream
ground ; on the last whorl the stripes are usually interrupted
by a light band below the periphery, and are irregular on the
base. Sometimes the brown markings are very faint. The
whorls are rather strongly convex. Aperture varying from
purplish-brown to faint pink within ; lip thin, very narrowly
expanded, white-edged within. Columella reflexed as in P.
physa.

Length 18, diam. 10.8, aperture 9.2 mm. ; whorls 6.

Length 18, diam. 11, aperture 9.8 mm.

Length 16, diam. 9, aperture 8 mm.

Near Pahoa, Puna, type loc., fig. 14, and Kaiwiki in the
district of South Hilo, figs. 15, 16 (Thaanum).

This very beautiful form was thought by the late Mr. Bald-
win to be distinct from his A. haivaiiensis, but Mr. Thaanum
agrees with us that it may more justly be ranked as a sub-
species. It is separated by innumerable gulches and water
courses from the range of P. pliysa, some forty to sixty miles
distant, but the intervening territory has not yet been fully
searched ; still there can be no doubt that the range of errans
is discontinuous with that of physa. In Puna it has reached
the extreme southeastern range of Achatinellidce.

In Olaa Mr. Thaanum found P. p. errans in ieie heads,

49. P. GRISEA (Newcomb). PI. 18, figs. 8, 9.

" Shell sinistral, inflated below, pointed at the summit;
whorls 6, rounded, not margined. Aperture ovate ; columella
short, flat and but slightly twisted; lip expanded, thickened

112 PARTULINA, MAUL

within. Suture well impressed ; umbilicus open. Color above
grayish-white mottled with light brown, below ashy-gray ; the
body-whorl encircled by a narrow white band. Length 16,
diam. nine-twentieths of an inch ' (Neivc.).

Maui: Makawao (Newcomb, type loc.).

Achatinella grisea NEWC., P. Z. S., 1853, p. 153, pi. 24, L
66. — PFR., Monogr., iv, 518. — A. diibia var. & PFR., Malak.
Bl., 1854, p. 117.

The conic spire rather slender above, and the weak colu-
mellar fold give this shell somewhat the contour of A. physa
Nc. of Hawaii. After the smooth first whorl, two are very
closely and deeply striate spirally. The spirals become weaker
on the last whorl. \]/2 apical whorls are pale brownish white.
Then broad, irregularly zigzag brown stripes usually appear,
and on the last embryonic whorl a dense pattern of narrow
zigzag lines (pi. 18, fig. 8). After 3^ embryonic whorls the
neanic pattern of oblique streaks sets in. These become zig-
zag or irregular on the later whorls. The last whorl is per-
forate, narrowly banded with white, dark fawn color below
the band. In two specimens I have seen there is no band.
The aperture resembles that of P. divightii except that the lip
is far less thickened and the columella moderately twisted
with no callous fold superposed upon the convexity. Figures
are from Makawao shells received from Newcomb.

Length 20, diam. 11.5 mm.

Length 21.2, diam. 12.5 mm. ; whorls 6^3.

P. grisea is not closely related to any other M>auian snail,
but it has much in common with P. dwightii of Molokai, the
shape of the spire, pattern and color being nearly alike in
the two species. The pattern of the last embryonic whorl is
different, and P. divightii has a stronger columellar fold. It
is one of the rarest Mauian shells, and as yet entirely un-
known to Hawaiian conchologists of the present day, although
the Makawao district has been thoroughly worked over. It
may be extinct.

50. P. THAANUMIANA Pilsbrv, n. sp. PL 18, figs. 6, 7.
The shell is sinistral, narrowly umbilicate, tliin, ovate-conic,

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PARTULINA DUBIA, OAHU. 113

surface slightly shining, the later whorls densely striatei
spirally, strias waved, descending, embryonic whorls very
densely and distinctly engraved. Embryonic shell of 8^2
whorls, the first 2 whorls brown under a whitish layer; third
whorl marked with oblique, angulated white stripes on a
reddish-brown ground. Following (neanic) whorls mottled
and streaked with white on a reddish-brown ground, which
becomes paler on the last whorl, which is indistinctly streaked
and mottled with creamy or brownish white, and girdled with
a narrow dark band at the periphery. The whorls are
strongly convex. Aperture slightly oblique, purplish-brown,
with light streaks within. Peristome a trifle expanding at
the base, narrowly bordered within with white, but not thick-
ened. Golumella slightly convex, not plicate, the margin
spreading in a triangle, white.

Length 17, diam. 10.5 mm. ; fully 6 whorls.

Length 15.5, diam. 10.2 mm. ; 5^4 whorls.

West Maui: Waiehu Gulch (D. Thaanum). Gotypes in
coll. Bishop Museum and A. N. S. P. Also in Mr. Thaanum 's
collection at Hilo.

This very distinct species is evidently related to the East
Mauian P. grisea Newc., from which it differs by the thinner
shell of somewhat different contour, the more convex whorls,
different pattern of the last embryonic whorl, larger umbili-
cus, dark peripheral band, etc. The aperture, lip and colu-
mella are more like P. physa than like P. grisea. The spire
is less drawn out and not so narrow above. P. aptycha Pfr.,
which has not been found by recent collectors, is obviously
distinct.

51. P. DUBIA (Newcomb). PL 26, figs. 7 to 12.

" Shell dextral, conically elongate, thin, finely decussately
striated, light corneous with radiating zigzag lines and
blotches of a light color. Whorls 6 ; suture simple. Aperture
ovate; 'columella white and bulimoid except in strongly de-
veloped adults where it is callous and obtusely dentated.
Umbilicus small but pervious. Lip thickened. Slightly re-
flected. Length 0.8, breadth 0.55 inch ' (Newc.).

114 PARTULINA DUBIA, OAHU.

Oahu, among stones; Waianae, found on bushes (New-
comb) . Waimano, eastern ravines southwest of forest fence ;
eastern ravines of Waiawa; east side of stream, Kawaiha-
lona, in Waialua; also Makalia and Makua valleys, Waianae
range (Irwin Spalding). In knot holes and crevices of loose
bark of kukul trees. In Waimano on guavas.

Achatinella dubia NEWC., Ann. Lye. N. H. of N. Y., vi, p.
23 (May, 1853) ; P. Z. S., 1853, p. 152, pi. 24, f. 65, 1854.—
Achatinella platystyla GULICK, Ann. Lye. N. H. of N. Y., vi,
p. 196, pi. 6, f. 25, 1856. — Achatinella pexa GULICK, Ann.
Lye. N. H. of N. Y., vi, p. 197, pi. 6, f . 26, December, 1856.—
Achatinella fucosa Frick, LYONS, Hawaiian Almanac and An-
nual for 1892, p. 105, pi. 1, f. 15.— A. morbida PPB., P. Z. S.,
1859, p. 30.

P. dubia is a foreigner in Oahu ; the strong spiral striation
and the pattern of the embryonic whorls show it to be a real
Partulina. Like Laminella, it seems to have wandered west-
ward just before the connection with the Molokai-Maui mass
was submerged. It is somewhat related to P. grisea and P.
thaanumiana of Maui, P. radiata Gld. also has a close re-
semblance, but in this the Partulina pattern has been lost
from the embryonic whorls, and the columellar fold is some-
what stronger. P. radiata, according to specimens in Mr.
Emerson's collection, is from West Maui.

In his second account of this species Newcomb states that
it may be either dextral or sinistral, and that old specimens
exhibit " a flat, twisted tooth ". His type figure is copied,
pi. 24, fig. 9. This form has light girdles at periphery and
base, with elsewhere a confused zigzag pattern of purplish-
brown and white or cream. The narrow callus within the
lip is white. Length 19.5, diam. 11, aperture 9.8 mm., whorls
§l/2 . The smoothish and rather shining surface shows engraved
spiral striae throughout. The lower part of the outer lip and
the basal margin are very slightly expanded. Ten of the spe-
cimens of this typical form before me are dextral, one sinis-
tral. The last embryonic whorl has conspicuous zigzag white
stripes on a dark ground. PL 26, fig. 7, is a specimen re-
ceived from Newcomb. Other specimens, pi. 26, fig; 8, have

PABTULINA DUBIA, OAHU. 115

straight light yellow and brown streaks on the post-em-
bryonic whorls. The smallest adult seen is 16 mm,, long.

Probably all of Newcomb's shells, to which the above notes
apply, were from the Waianae range, where it has been found
in Makaha valley (on the southern side) by Mr. Spalding,
and perhaps elsewhere by other collectors. PI. 26, fig. 8, is a
Waianae range shell.

In the main range P. dubia has an extended distribution,
from Waimano to Waialua or Kawailoa, but only in a few
places, so that it is regarded as a rather rare shell. Very fine
series are in the collections of Messrs. Spalding, Emerson and
Wilder. As a general rule, main range shells have a more
thickened lip than those of Waianae, and the apex is often
perceptibly blunter, whorls a trifle more convex ; so far as I
know, no sinistral shells have been found ; but I am not satis-
fied that these small differences hold in all specimens.

In Waiawa (pi. 26, figs. 9, 9a, 9&, coll. by Spalding in the
eastern ravines) the shell is streaked and mottled, with boldly
zigzag-striped last embryonic whorl ; or similar with a whitish
peripheral band and several lines. Others are chestnut with
pale streaks, a white peripheral band and several spiral light
lines, the white zigzags of the embryo very much reduced
(pi. 26, fig. 9 6), or white may predominate, the chestnut
being reduced to bands narrower than the white, one speci-
men. In a few the later whorls are whitish with brown stains
(pi. 26, fig. 9).

An extraordinary series was collected by Mr. Spalding in
Waimano. Most of the colony is of the usual streaked and
mottled form, but in a few the color is very deep, rich brown
(between liver-brown or carob-brown and black, of Ridg-
way's Color Standards). This may be uniform or varied
with light buff bands. The embryonic whorls are brown with
light bands and lines, and on the last embryonic whorl there
is a trace of the light stripes of the typical form. The figures,
pi. 26, figs. 10, 10&,, are from two specimens of Mr. Spalding 's
no. 2181. This color-form is one of the rarest Oahuan trea
snails. A suspicion has been entertained that it might be a
hybrid between clubia, and a rare black form of Adi. turgida-,

116 PABTULINA DUBIA, OAHU.

but having examined all ol* the specimens found, I think the
evidence against that hypothesis.

Achatinella platystyla Gulick, pi. 26, fig. 11, and pi. 50, fig.
17, is a peculiar color-form of dubia, described from a single
shell. The surface is a good deal pitted, but it was evidently
a " live ' shell, and an old one. The surface is entirely
eroded to the middle of the fourth whorl. What cuticle re-
mains on the next two whorls is pecan-brown with creamy
streaks, and fading upwards. The last whorl is light buff in
the upper third, shading downwards into pecan-brown, which
is streaked with a lighter tint and has very indistinct spirals
of the darker shade. The color is almost exactly that of some
specimens of Achatinella glabra. The surface shows faint
spiral striation, which becomes stronger upwards, as usual in
P. dubia. The aperture, lip and perforation are exactly as
in Main Range dubia. The lip has a faint yellow tint, deeper
at the edge. Length 20, diam. 10.3, aperture 8.4 mm. ; 6^2
whorls.

The unique type of platystyla was collected by Mr. Gulick
in Kawailoa. It is no. 25 of 'his type collection, Boston So-
ciety of Natural History. I have given two views of this
specimen.

Achatinella pexa Gulick, pi. 26, fig. 12, is certainly a sinis-
tral specimen of A. dubia. The unique type specimen, no.
26 of Gulick 's type collection, coll. Boston Society of Natural
History, is figured. The shell has the texture and thin sub-
stance of dubia. The ground-tint is between light buff and
white. It appears in spirals and obliquely axial streaks, leav-
ing rather wide interrupted streaks of vinaceous pink, here
and there darker. There is a brown line at the periphery
and a group of three around the umbilical region. The early
whorls are eroded, but traces of alternate light and darker
stripes can be made out on the last embryonic whorl. The
glossy surface is engraved with rather distinct close spirals,
as in P. dubia. The lip expands a little. Aperture, lip and
perforation are as usual in P. dubia; the lip having a pale
yellowish-brown edge and slight thickening. Length 19, diam.
10.7, aperture 9 mm. ; 6^/2 whorls.

ACHATINELLA. 117

Another figure of the type of pexa is given, pi. 50, fig. 18,
to show the form of the columella more distinctly than the
figure on plate 26.

The pattern of color is practically the same as in some
Waiawa diibia, though developed in delicate tints; and the
specimen may have come from that neighborhood; yet it is
sinistral with a thin lip, characters more in harmony with the
Waianae range form. Gulick did not know the locality.

Achatinella morbida Pfeiffer. " Shell subperf orate, sinis-
tral, ovate-turrite, rather solid, striate and under a lens seen
to be decussated with close spiral striae; white, variously
streaked and banded with brown; spire long, slender, the
apex rather acute, suture simple; whorls 6y2, very slightly
convex, the last a little shorter than the spire, convex ; colu-
mellar fold white, short, oblique; aperture oblique, inverted
ear-shaped ; peristome thickened, narrowly expanded, the
eolumellar margin much dilated, broadly adnate. Length 19,
diam. 9 mm. Sandwich Islands, Dr. Frick in Mus. Cuming. '

(P/T.).

Achatinella morbida PFB., Proc. Zool. So<c. London, 1859,
p. 30 ; Monographia, vi, 167.

This species has not been figured. The description agrees
well with sinistral P. dubia, such as are found in the Waianae
Mountains; and a figure of the type-specimen which Mr. E.
A. Smith kindly had made for me (pi. 41, fig. 4) confirms this
determination. A. morbida is merely a somewhat unusual
color-pattern of P. dubia.

Genus ACHATINELLA Swainson.

Achatinella SWAINS., Quarterly Journal of Science, Liter-
ature and Art (2 ser.), iii, June, 1828, p. 83 ; type Monodonta
seminigra Lam. — Achatinella apexfulva Dixon. — GULICK,
P. Z. S., 1873, p. 90, classification. — Cochlogena, quatrieme
groupe, Helicteres, FERUSSAC, Tableau Syst. de la Famille des
Limaqons, p. 56, 1821. — Helicteres Fer., BECK, Index Mollus-
corum, p. 51, 1837.— GRAY, P. Z. S., 1847, p. 178, type Helix
vulpina. — HERRMANNSEN, Indicis Gen. Mai., i, p. 515, type
Helix vulpina Fer. — Helicter PEASE, P. Z. S., 1862, p. 3. —

118 BULIMELLA.

Apex VON MARTENS, Die Heliceen, 1860, p. 248, type Achati-
nella lugubris Chemn.

Shell imperforate or minutely perforate, oblong, ovate or
globose-conic, smooth or longitudinally corrugated, with only
minute and weak traces of spiral sculpture; color in spiral
bands or streaks in the direction of growth-lines, never in
forwardly-descending stripes; lip simple or thickened within,
sometimes slightly expanding ; columella bearing a strong cal-
lous fold.

Type A. apexfulva (Dixon). Distribution, the island of
Oahu. Living on the leaves and limbs of trees and bushes.

Data on the soft anatomy, distribution, etc., of Achatinella
may be found in the Introduction of this volume.

Achatinella was early divided into several subgenera or sec-
tions, but their contents as seen in the lists in Die Heliceen^
Nomenclator Heliceorum Viventium and other works were
rather heterogeneous. Gulick and Baldwin grouped the
species more naturally, practically limiting the groups as in
this work. Three sections or subgenera are recognized. There
are a few species on the border between Bulimella and Acha-
tinellastrum, but otherwise the sections are. quite distinct,
though they can scarcely be rigidly diagnosed.

Sections of Achatinella.

Bulimella Pfr. Outer lip thickened by a strong callous rib
within; shape oblong-conic or ovate; summit obtuse, rounded
or convexly-conic.

Achatinellastrum Pfr. Outer lip thin or a little thickened
within, not expanded ; early whorls not conspicuously flat-
tened, the summit conic, not very obtuse.

Achatinella s. sir. Outer lip well thickened within, scarcely
or not expanded ; shape globose-conic or ovate-conic, the sum-
mit conic, embryonic whorls flattened.

Section BULIMELLA Pfeiffer.

Bulimella PFR., Malakozoologische Blatter, i, 1854, p. 119.
— MARTENS, Die Heliceen, 1860, p. 244, type A. rugosa Newc.

Achatinellse with the shell ovate-conic or somewhat pyra-

BULIMELLA. 119

midal, -the 'Outlines of the spire more or less convex, at least
near the rather obtuse summit; embryonic whorls not con-
spicuously flatter than those following; lip distinctly thick-
ened by an internal callous rib (except in A. abbreviata and
A. lila) , the outer edge often slightly expanded.

Type: A. byronii rugosa Nc. Distribution: Entire Koolau
or Main range of Oaliu.

About forty-six names, most of them originally introduced
as "species", have been applied to members of the Bulimella
group up to this time. Mr. Baldwin, in his Catalogue of
1893, enumerated twenty- five species. Mr. E. R. Sykes
(1900) admitted twenty species and three varieties. "In the
present monograph, thirteen species and eighteen subspecies
are recognized ; but this number includes two species and ten
subspecies not before published. Most of the other names are
synonyms, but a few may be retained for local forms thought
to be below the gra.de of "subspecies". A more rigorously
logical treatment might reduce the "good species' to nine,
by the union of viridans and tceniolata; byronii, decipiens and
pulckerrima; and bulimoides and elegans; but the price of
such consistency would, I imagine, be the loss of clear con-
ceptions.

Bulimella is a shell of the ravines as well as of the ridges,
and before the extensive deforestation of the lower slopes,
many fine species were to be found low in the valleys. This is
especially true of the bulimoides group, which formerly occu-
pied much territory now quite -barren. The fuscobasis group
is almost confined to high ridges and peaks.

The greatest development of Bulimella is- in the northwest-
ern half of the Koolau range. There are but four species and
two or three subspecies in the whole southeastern third or
more of the length of the island, leaving nine species and
fifteen subspecies in the western five-eighths of the length.
Moreover, the differentiation has been greater in the west,
and doubtless new forms remain to be found there, whilst the
eastern part of the range has been so fully explored that no
new forms of Bulimella can be expected.

Bulimella contains the most roughly sculptured species of

120 ACHATINELLA, SECTION BULIMELLA.

the genus, some forms being irregularly corrugated in the
direction of the lines of growth. Spiral sculpture is weak or
obsolete in all the species.

There are three collateral groups or series of species in
Bulimella, the first subdivided as a convenient geographic di-
vision.

Series of A. byronii: shell oblong-conic, often -with streaks
or bands of dark green, and often roughly striate or corru-
gated; mainly dextral. A. byronii, pidcherrima, decipiens,
lila.

Series of A. viridans: a subdivision of the above series, fo-r
the exclusively dextral species of the eastern end of the range.
A. abbreviata, viridans, and tceniolata.

Series of A. bidimoides: shell ovate, capacious, never
marked with green; smooth, dextral or sinistral. A. rosea,
bidimoides, elegans.

Series of A. fuscobasis: shell ovate-come, never marked with,
green; smooth; mostly sinistral. A. fuscobasis, sowerbyana,
pupukanioe.

On account of the wide variation in shape and color, it is
not possible to rigidly diagnose these groups, which never-
theless express the natural affinities of the species.

Key to species and most subspecies of Bulimella.

I. Forms of the southeastern third of the main range (east of

Mjoanahia) .

a. Lip acute, but slightly or not thickened within ; shell dex-
tral, very smooth, richly colored. A. abbreviala, no. 1.
a1. Lip strengthened by a callous rib.
b. Shell dextral.

c. Streaked with green or chestnut on a lighter or

yellow ground. A. viridans, no. 2.

c1. Banded or streaked with chestnut on a white

ground, or white. A. tceniolala, no. 3.

61. Shell sinistral.

c. Shell white with few brown bands or none, about
16 x 10 mm. A. fuscobasis, no. 11.

c1. Similar but larger and more capacious.

A. f. lyonsiana, no. lla.

ACHATINELLA, SECTION BULIMELLA. 121

c2. Shell richly colored, banded. Lanihuli.

A. f. wilderi, no. 116.

II. Forms of the northwestern two-thirds of the main range.
a. Lip acute, but slightly or not thickened within ; shell sin-
istral, very smooth, richly colored. A. lila, no. 5.
a1. Lip distinctly thickened within.
6. Shell oblong-conic.

c. Surface corrugated or somewhat roughened by

growth- wrinkles.

d. Interior drainage ; shell sinistral, later whorls
corrugated, intensely black or banded with
dark green. A. &. nigricans, no. 46.

d1. Interior drainage; shell dextral.

e. Roughly striate, streaked with green, chest-
nut or black, usually with sutural chest-
nut band or others; lip edged with
brown. A. byronii, no. 4.

e1. More corrugated, occasionally black; sur-
face with a gummy gloss ; whole lip usu-
ally brownish ; length 16-19 mm.

A. 6. rugosa, no. 4&.

e2. Larger, broader, green-streaked; lip white

or narrowly dark-edged; length 19-21

mm. Var. capax.

es. Nearly smooth, whitish to greenish-yellow,

plain or banded, lip white or pale lilac ;

length 15-18 mm. Var. waimanoensis.

d2. Northern (Koolauloa) slope; shell sinistral or

dextral, the lip mainly white.

A. decipiens, no. 7.
c1. Surface smooth ; shell dextral.

d. Northern slope ; lip white. Various varieties of

A. decipiens, no. 7, la.
d1. Interior slope.

e. Lip bordered with blackish brown.

A. pulclierrima, no. 6.
e1. Lip white throughout. A. p. nympha, no. 6a.

122 ACHATINELLA, SECTION BULJMELLA.

61. Shell ovate, smooth.

c. Larger, capacious species, diam. 10 mm. or more.
d. Shell sinistra.l, highly polished, usually white
or with rose bands, lip bright rose ; rarely
greenish or dark, with dark brown lip.

A. rosea, no. 8.

d1. Dextral or sinistral, variously banded with
chestnut or suffused, lip pale or brown.

A. bulimoides, no. 9.

d2. Dextral or sinistral, conspicuously streaked
with brownish, sometimes with white bands
•also, lip brown. A. elegans, no. 10.

d3. Chiefly sinistral, blue-gray or purplish, some-
times banded. A. e. wheatleyana, no. 10&.
c1. Smaller species, usually less than 10 mm. diam.
d. Shell sinistral.

e. Wax-yellow with a brown sutural band,

sometimes one or two other bands, colu-

mella pink. A. sowerbyana, no. 13.

e1. Wax-yellow below, white above periphery,

with a subperipheral brown band; colu-

mella pink. A. s. thurstoni, no. 13a.

e2. "White, usually with broad chestnut bands;

columellar fold white or brownish.

A. s. laiensis, no. 13&.
d1. Shell dextral.

e. Wax-yellow with white sutural band, colu-

mella pink. A. s. roseoplica, no. 13d.

e1. Ground white or pale, uniform, or banded

or streaked with chestnut.
A. pupukanioe, no. 12. A. s. dextroversa, no. 13c.

SERIES OP A. VIRIDANS.

(Species of the east end of the Main Range.)

These forms, which are invariably dextral, inhabit the val-
leys from Nuuanu to the southeastern end of the range. Be-
tween Xuuanu and Waimanalu there are a.bout half a dozen
great ridges and valleys where no BuUmella is known except
A. fuscoltasis irilderi on some high points of the main axis.

ACHATINELLA ABBBEVIATA. 123

The viridans group is somewhat intermediate between the
byronii and the bulimoides series, but most nearly related to
the former by the frequent development of bright green
color, which none of the bulimoides series has. Besides the
following forms, one unrelated sinistral species occurs on the
southeastern ridges, A. fuscobasis, no. 11.

1. A. ABBBEVIATA Reeve. PL 31, figs. 1 to 5a.

' ' Shell ovate, somewhat ventricose, dextral ; whorls convex,
margined round the upper part ; spire rather short, obtuse at
the apex ; columella callous, twisted. Olive-yellow, with a
black-brown line at the sutures, lower part of the last whorl
very dark green, apex black.' (Reeve.)

Oahu: Eastern ridge of Nuuanu to Niu. Palolo (Gulick
and others; type loc.) ; around edge of the crater, and Palolo-
Waialae ridge (Spalding) ; northwestern ravine, next to the
Palolo-Manoa ridge (Pilsbry & Cooke). Konahuanui (Bald-
win) ; eastern Nuuanu, perhaps washed down from Kona-
huanui ( Spalding) . W'aialae, Niu, and over the ridge in Kai-
lua ( Gulick) .

Achatinella abbreviata REEVE, Conch. Icon., vi, pi. 3, fig.
19, April, 1850.— PFR., Monogr., iii, 460; iv, 532; vi, 172.—
BALDWIN, Catalogue, p. 5. — SYKES, Fauna Hawaiiensis, p. 305.
— THWING, Orig. Descr. Achat,, p. 81, pi. 1, f. 22. — GULICK,
Evolution, Racial and Habitudinal, p. 41, pi. 2, f. 25. — Acha-
tinella bacca Reeve, Conch. Icon., vi, pi. 6, f. 45, May, 1850. —
GULICK, Evolution, Racial and Habitudinal, p. 41, pi. 2, f . 22.
— Achatinella nivosa NEWC., P. Z. S., 1853, p. 132, pi. 22, f. 6
(Niu). — Achatinella Clementina PFR., P. Z. S., 1855, p. 205;
Monogr., iv, 534.

A. abbreviata differs from the viridans forms by its broad
contour, thin outer lip, smoother surface and different pat-
tern, yet is related to them by color and the faint traces of
corrugation noticeable in some specimens. It is a very dis-
tinct species.

A. abbreviata is a common species around the crater of
Palolo valley, and on its bounding ridges near the heads of the
upper ravines. It lives by preference in the axils of ieie. Hot

124 ACHATINELLA ABBREVIATA.

water, or too prolonged soaking, changes the vivid green color
-to a dirty olive-brown. The western extreme of its range is
(or was) in eastern Nuuanu, near the southeastern edge of
the new dam, where Mr. Spalding found five immature speci-
mens of the profusely-banded form (no. 1817 of his collec-
tion), possibly washed down from Konahuanui.

PI. 31, figs. 1, la, 5, 5a? Palolo, are from Gulick shells. The
others, figs. 2 to 4<i, are from material collected recently. A
lot from the northwestern ravine of Palolo varies from many-
banded to black (pi. 31, figs. 4, 4a-4d). There was also one
shell all green except the brown summit. Figs. 3&, 4<i, 5 and
5a are rare color-forms.

The shell is obesely-ovate, very glossy, showing faint spiral
striation on the lower, clear but very fine incised lines on the
early whorls. The lip is acute, not thickened within or but
slightly so. All I have seen are dextral.

1. The typical color-pattern as figured by Reeve is yellow
above the periphery, green below it, the colors separated by a
peripheral chocolate band. The suture is bordered above and
below by chocolate bands. Near the summit the yellow ground
gives place to reddish-brown, becoming dusky or purplish-
brown at the apex (pi. 31, fig. 2). This pattern occurs in
Palolo valley together with the following, the inter gradation
being complete throughout the series.

2. Similar to no. 1, but having additional chocolate lines
and bands (pi. 31, fig. 2a, Palolo; figs. 3, 3«, Palolo- Waialae
ridge), which sometimes almost cover the yellow and green,
ground.

3. Same as no. 2, but without green (pi. 31, figs. 4, 4&), or
the bands may be confluent, leaving yellow bands at periph-
ery, suture or both (fig. 46).

4. Last whorl black or nearly so, landless, the color becom-
ing lighter upward, dark again at the summit (pi. 31, figs. 4c,
4d) . This is the form called A. bacca by Reeve. A. abbre-
viata and A. bacca are therefore merely selected patterns in a
continuous series of variations occurring in the same colony.
Figs. 4 to 4:d are from a northwestern ravine of Palolo, next
to the Manoa ridge, coll. by Pilsbry.

ACHATINELLA VIRIDANS. 125

5. Shell yellow above (varying to olivaceous or brownish-
yellow) , green, dull olive or even chestnut below the periphery,
usually bandless, sometimes having one or several reddish-
brown bands. The columellar fold is not so strong as in the
preceding. (PI. 31, figs. 1, la, Palolo.) So far as I know,
this form does not occur in colonies of the chocolate-banded
forms. There is a large lot from Palolo in the Gulick collec-
tion, 92572 A. N. S.

Waialae. Several lots collected by Gulick and others have
the patterns of figs. 2a, 3, 3a, 4&, c, d, not distinguishable
from Palolo shells. Another lot agrees with fig. 1. PL 31,
figs. 3, 3a are from the Palolo- Waialae ridge, collected by Mr.
Kuhns.

Wailupe. Occurrence doubtful.

Niu. Many-banded shells, pattern of fig. 3, collected by Gu-
lick.

The locality "head of Kawailoa Gulch, Perkins," given by
Mr. Sykes, must be an error in labeling or determination.

A. nivosa Newc. is universally admitted to be a scraped
abbreviata. The original figure is reproduced, pi. 29, fig. 6.

A. dementina Pf r. has not been figured. It has been placed
in the synonymy of (Mreviata by Pfeiffer himself and by sub-
sequent authors. The description, which is not very good for
dbbreviata, here follows: " A. dementina. Shell imperforate,
dextral, ovate-conic, solid, nearly smooth, glossy; greenish-
brown, banded below the suture with buff. Spire regularly
conic, obtuse, suture chestnut, impressed-marginate, crenu-
lated. Whorls 6, a little swollen above, the last about equal
to two-fifths the total length. Aperture oblique, truncate ear-
shaped, white within; columellar fold above, moderate, white
or flesh-colored. Peristome unexpanded, the margins joined
by a callous, right margin somewhat straightened, thickened-
labiate within; columellar margin dilated, adnate. Length
19, cliam. 10, aperture 9% mm. long, 4% wide. Oahu, Frick
in Cuming coll.' (Pfr.)

2. A. VIRIDANS Mighels. PI. 25, figs. 1 to 4 ; pi. 31, figs. 6, 6a.
11 'Shell dextral, elongate-conic, green with light streaks in-

126 ACHATINELLA VIRIDANS.

termixed, imperforate. Whorls 5, convex, with a revolving,
slightly impressed line .below the suture. Aperture subovate,
stained with a pink color just within the margin ; lip slightly
thickened. Length three-fourths, diam. seven-fifteenths inch':
(Mighels).

Oaliu (Mighels). Nuuanu to Palolo, various color-forms as
far east as Niu.

Achatinella viridans MIGHELS, Proc. Boston Soc. Nat. Hist.,
ii, p. 20, January, 1845. — PFR., Monographia Hel. Viv., ii,
240; iv, 519; vi, 164; Conchyl. Cabinet, Bui, p. 286, pi. 67,
f . 16, 17.— NEWCOMB, P. Z. S., 1854, p. 310 (animal) .— Acha-
tinella radiata PFR,? P. Z. S., 1845, p. 89 (January, 1846).—
REEVE, Oonch. Icon., pi. 5, f. 35.

Achatinella subvirens NEWC., P. Z. S., 1853, p. 136, pi. 22,
f. 18, 1854.— Achatinella rutila NEWC., P. Z. S., 1853, p. 138,
pi. 22, f. 21, 1854; p. 310 (animal) ; Ann. Lye. N. H. of N. Y.,
vi, 326 (animal). — Achaiinella macrostoma PFR., P. Z. S.,
1855, p. 2, pi. 30, f. 6; Monogr., iv, 520 (Sandwich Is., Frick
in Cuming coll.).

The type of A. viridans was lost by fire, but the descrip-
tion applies well to the form found in Nuuanu, Manoa and
Palolo valleys, and which has always been considered viridans
by Hawaiian collectors. C. B. Adams, who had seen Mighels '
specimen in his collection, stated that it was identical with A.
radiata Pfr. as figured by Reeve. The traditional identifica-
tion of viridans is therefore confirmed, that figure being the
same as our pi. 25, fig. 1. The shell is glossy, noticeably,
though weakly, corrugated. The first three whorls are whitish,
or 'white when the thin cuticle is lost ; next whorl or two yellow
or olivaceous, more or less streaked; last 1% or 2 whorls
copiously streaked throughout with dark and light green of
several shades. The sutural border is distinctly marked by
an impressed line. The aperture is white within, the lip a
trifle expanded, thickened within, and either white or flesh-
pink. In a large number of specimens seen, not one is sinis-
tral. The size and form vary, but shells 20 mm. long, 11 wide
are about normal. A wide shell from Nuuanu measures,
length 20.2, diam. 13 mm.

ACHATINELLA VIRIDANS. 127

In Nuuanu valley the shells vary from typical green color
to a form in which the dark green is replaced by blackish-
chestnut, light green by yellow (pi. 25, fig. 4). A specimen
of the rather widely-spread form of Upper Nuuanu, pale
yellow with light yellowish olive streaks, is figured on pi. 31,
fig. 6o> coll. by Mr. Richard A. Oooke. A small, thin-shelled
form was found only on a few isolated bushes at the head of
the valley just west of the pali. It is chestnut-brown with
lighter streaks and pale lines below suture and periphery;
lip-callus narrow; length 15.8, diam. 8 mm. PI. 31, fig. 6,
coll. by R. A. Cooke.

In Manoa Gulick collected the typical green form (pi. 25,
figs. 1, la) ; others having black, others olive (pi. 25, fig. 3)
or chestnut streaks on a yellow ground. There is now no
forest low in Manoa valley.

Palolo has the same range of forms shown in figs. 1 to 3.
A black-streaked shell is figured (pi. 25, fig. 2, collected by
Gulick). Really typical viridans does not go eastward of
Palolo, so far as I know. It is a rather homogeneous race,
having the same pattern but in varying shades and colors.
Probably green, olive, chestnut and black are varying stages
of oxidation of the same pigment; or perhaps the dull ground
of some specimens may be due to cleaning with hot water.
In Palolo the forms rutila and subvirens also occur, whether
•associated with the typical viridans pattern I do not know.

Doctor Newcomb has given the following description of the
soft parts. A. viridans: ' ' Animal light gray; tentacles and
tentacular sheath dark slate; mantle thick, yellowish-brown.
Tentacles strongly clubbed, short and robust, when extended,
longer than the shell.'

A. rutila: " Animal small in proportion to the shell, of a
uniform yellowish-white, retractile part of upper tentacles of
a light-brown ; tentacles filiform and slightly clubbed ; foot
very broad, long as the shell ; mantle same color as the ani-
mal. ' '

Eastward of Palolo the typical viridans pattern disappears,
and the color-forms rutila and subvirens replace it. These are
often hybridized with more or less blending, so that the ap-

128 ACHATINELLA VIRIDANS.

pearance of a colony suggests rutila X subvirens or rutila X
tceniolata. It might be well to recognize siibvirens as an east-
ern subspecies of viridans, rutila and macrostoma to be syn-
onyms of it.

Color form rutila Newcoonb. PI. 25, figs. 10, 11, lla-e, 12.

' ' Shell ovately-conic ; whorls 6, rounded, the last margined
above; suture well marked; aperture subquadrate; lip ex-
panded, subreflected, strongly thickened within; columella
short, terminating in a strong, twisted plait. Color a light
straw, olive or brown ; lip white or somewhat roseate. Length
17, diam. nine-twentieths inch.' (Newc.)

Niu (Newcomb, type loc., pi. 25, figs. 10, 11 to lie. The
color-form figured by Newcomb as typical is streaked with
light green on a yellow-green ground, with several spiral
brown bands below the periphery (pi. 29, fig. 21, reproduction
of Newcomb 's type figure ; pi. 25, fig. 10, specimen from New-
comb) . In other shells from Niu, collected by Gulick, figs. 11
to lie, the spiral bands may be more numerous or they may
be wanting. The streaks are sometimes chestnut or umber,
and either distinct or blended. The sutural border is almost
invariably tessellated with brown, thus differing from var.
subvirens, in which it is white, or at least not more heavily
marked than the rest of the shell. Wailupe shells are similar
(pi. 25, fig. 12).

In Waialae valley the color is usually light, and specimens
having bands like rutila with the white suture of subvirens
were found by Gulick and kept by him in the same lot with
others having the tessellated suture of rutila. See pi. 25, figs.
5 to 5d. There is no definite break between the two forms,
merely a matter of one or another color-mutation prevailing
in the colony. Even as far west as Palolo there are some
shells with more or less tessellated sutural border.

A. macrostoma Pfr. seems to me to be a form of rutila, as
Newcomb held, specimens of similar pattern to Pfeiffer's fig-
ured type occurring in Waialae Nui (pi. 25, fig. 8, Cooke
coll.). The original figure is reproduced, pi. 30, fig. 6. Mr.
Sykes has thought it a synonym of t&niolata,, and it must

ACHATINELLA VIRIDANS. 129

be admitted that some of the yellow-ground specimens from
colonies presumably t&niolata X subvirens have a great re-
semblance to Pfeiffer's figure. The original description fol-
lows. ' A. macrostoma Pfr. Shell dextral, imperforate,
conic-ovate, rather solid, very lightly striated, glossy ; fulvous,
variegated with some pale and brown bands. Spire conic,
lather obtuse; suture submarginate; whorls 5%, moderately
convex, the last equal to the spire, swollen above, 'contracted
in the middle, rounded at base. Aperture slightly oblique,
ample, reversed auriform, white within; peristome white,
labiate within, the right margin expanded, strongly curved
above ; columellar margin dilated, adnate. Length 211/2, diam.
11 mm. ; 'aperture 12 mm. long, 5 wide inside. Inhabits the
Sandwich Islands, Frick; Mus. Cuming (Pfr.).

Color-form snbvirens Newc. PI. 25, figs. 5 to lit, 9 to 95.

" Shell 'conically ovate. Whorls 6, rounded and margined
above ; suture distinct and lined with white. Aperture ovate,
expanded below; lip thickened; coluniella short, flat and ob-
liquely truncated ; color of epidermis light green, interspersed
with a lighter shade 'arranged longitudinally; columella, lip
and aperture white. Length 15, diam. seven-twentieths of an
inch. Niu, Oahu.

" Var. a. Pure white.

" Var. 5. Brown or chestnut replacing the green color.

" Through var. a- this species approaches a variety of A.
rutila, and through var. ~b, A. decipiens. It is readily dis-
tinguished from the first by its smaller size, greater solidity,
stronger striae and more elongate form ; from the latter by its
less acuminate form, white suture, less solidity, and their
widely separated localities ' (Newcomb).

Newcomb's figure of the type from Niu is reproduced, pi.
29, fig. 18. The lip is white or yellowish, the green color is
paler than in viridans, surface less wrinkled, and typically
the sutural margin is white, though very often the dark lines
extend over it. It differs from viridans in the average, but
one could not pronounce on every specimen, and if Gulick's
lots are reliable, it occurs in the same colonies with rutila.

130 ACHATINELLA T^NIOLATA.

The same variety occurs in Wailupe and Waialae valleys. A
series is figured, pi. 25, figs. 5 to 5d, Waialae ; also pi. 25, fig.
6, a clear green shell occurring with the brown-streaked form,
Wailupe. Some shells from Waialae have spiral bands as in
var. rutila (pi. 25, figs. 7 to 76).

Gulick found a few subvirens also in Palolo, the white,
green-streaked and brown-streaked forms. Also specimens
connecting with form rutila, the sutural border yellow in
some, white in other examples. Color yellow with indistinct
olive lines, bandless or with white or chestnut bands (pi. 25,
25, figs. 9, 9a, 96, Palolo) .

3. A. T^CNIOLATA Pf eiffer. PI. 24, figs. 14 to 19 ; pi. 25, figs.
13 to 13c.

" Shell ovate-oblong, solid, striatulate, glossy; white orna-
mented with varying brown bands, more obsolete above. Spire
conic, rather acute. Whorls 6, slightly convex ; the last about
four-ninths the length. Columella white, strongly tooth-
folded above. Aperture irregularly semi oval, white within,
glossy; peristome very narrowly thickened outside, strongly
lipped within, the columellar margin dilated, reflexed, ap-
pressed. Length 20, diam. in the middle 11 mm. ; aperture
10x4!/2 mm. Sandwich Is., Mus. Cuming.' (P/V.)

Oahu : Palolo to Keawaawa, and northward across the range
in Maunawili.

Achatinella t&niolata PFR., P. Z. S., 1846, p. 38; Mono-
graphia, ii, 239 ; iv, 519 ; vi, 164. — REEVE, Conch. Icon., pi. 1,
f. 7. — NEWCOMB, Ann. Lye. N. H. of N. Y., vi, p. 320. — SYKES,
Fauna Hawaiiensis, p. 310. — Achatinella rubiginosa NEW-
COMB, P. Z. S., 1853, p. 154, pi. 24, f. 69 (Palolo).

A. taniolata typically has a white ground, variously banded
or streaked with rufous or ochraceous-orange, or without
markings. The contour is usually somewhat more capacious
and the surface smoother than in the forms subvirens and
rutila of A. viridans; but both contours and sculpture inter-
grade in some shells. The coloration is characteristic in typi-
cal examples, but here again there are colonies from Palolo,
AVaialae, Keawaawa and elsewhere, which could be placed,

ACHATINELLA T^NIOLATA. 131

with .almost equal propriety in tceniolata, viridans rutila or
viridans subvirens, or 'assorted into these seve'ral strains.
The fact seems to 'be that several forms, well differentiated
in pure colonies, have interbred in other places, forming hy-
brid colonies. Strictly speaking, tceniolata is a subspecies of
A. viridans, because there is complete intergradation between
them; but as the race is usually quite recognizable, it is here
admitted as a species. In dealing with Achatinella it is im-
possible to be logical and at the same time preserve a practi-
cally useful classification.

The color in Palolo shells varies from white to few-banded
or many-banded with light reddish-brown, apex and sutural
border white, o>r rarely the apex is purplish-brown, perhaps
stained from within (pi. 24, figs. 15, 16, Pialolo, Gulick coll.).

Length 20, diam. 10.2 to 11.2 mm.

Length 18, diam. 11.5 mm. (exceptionally obese).

Length 21, diam. 10.8 mm.

Length 21, diam. 11.8 mm.

As these shells agree well with Pfeiffer's description and
Reeve's figure, Palolo may be taken as type locality.

In some colonies the color-form nibiginosa Newc. (pi. 24,
fig. 19, Palolo, and pi. 25, figs. 13, 13&, Wailupe, all from
Gulick coll.) is prevalent, together with banded shells. In
this form the shell is suffused and streaked with rufous, a
band or line below the suture and a columellar patch usually
white. Typical t&niolata might be described as rubiginosa in
which white bands traverse the shell spirally, cutting the
dark color into bands. It has thus a more advanced pattern
than that of rubiginosa, and the white shells are still further
evolved, though the pattern is degenerate.

According to Newcomb, riibiginosa has a light flesh-colored,
and tceniolata a brown or dark slate-colored animal and
mantle. In his later paper he admits the specific identity of
rubiginosa and tceniolata.

In Waialae (pi. 24, fig. 14) and Wailupe some large lots
taken by Gulick are white or with but few spiral lines or none,
while in others the profusely banded and the rubiginosa type
of coloring prevails. PL 25, figs. 13 to 13c are Wailupe shells

132 ACHATINELLA TJSNIOLATA.

collected by Gulick. A recently collected Waialae lot from
Thaanum consists of (a) white shells with brown spire, (b)
white shells with numerous brown bands, as in Palolo tanio-
lata, and (c) greenish-yellow shells with chestnut-olive streaks,
sometimes cut by white bands. There are various blends be-
tween a and c patterns. This colony is probably a siibvirens
X t&niolata hybrid.

A small lot from Niu (Gulick) consists of well-colored
banded shells (t&niolata) , or streaked (rubiginosa) , with
white ground.

From Kuliouou (Thaanum) the specimens have the pattern
of Keawaawa form (b), see below.

In a set of 9 from Keawaawa, coll. by Gulick, some have a
few faint bands, others being white, peristome and parietal
wall yellowish. The shape varies, as in Palolo shells : length
19, diam. 10 mm., to length 17, diam. 10^2 mm. A lot from
the Thaanum collection, recently collected, there are (a)
shells variously banded with chestnut on a yellow ground; (b)
the same with many oblique streaks or lines added, and (c)
others of rubiginosa pattern, the streaks 'blended. The yellow
ground shows probable hybridism with rutila or siibvirens.

On the northern side of the main range Gulick obtained a
few specimens in Kailua — rather an indefinite locality. They
have streaked rubiginosa coloring, with paler spiral bands.
One has a greenish tone. They are transitional between rubi-
ginosa, t&niolata and siibvirens. Size rather small, length 17
mm. It occurs also in Maunawili, on the northeastern slope
of Mt. Olympus, a place exploited by Messrs. Kuhns and
Wilder. Here the shells are white with dark or light-brown1
upper whorls, paler (sometimes white) embryonic whorls, the
last whorl either banded or with rubiginosa pattern cut by one
or more white bands. PL 24, figs. 17, 18.

(Species of the northwestern two-thirds of the Main Range.)

SERIES OF A. BYRONII.

What forms of the byronii group may occur upon the crest
of the main axis of the range 'between the known a>reas of
byronii and decipiens are not known to me. For this reason,

ACHATINELLA BYRONII. 133

and because there are differences in 'the patterns and colors,
I am recognizing the forms from the northward- side, collec-
tively, as a species (A. decipiens} distinct from those on the
south side of the range. The decipiens forms 'all appear to
be more closely related to one another than any of them are
to forms from the other side of the range, They seem to form
a parallel series to the latter. In both series there are smooth
forms in the west, corrugation becoming progressively more
emphatic eastward. In both series the western forms are in-
variably dextral, and sinistral forms occur in the east. This
correspondence would suggest communication across the
range, but the coloration is against that explanation of the
parallelism. From northwest to southeast the forms are ar-
ranged thus: smooth, dextral forms above (westward), rough,
sinistral forms below (eastward).

kaliuwaaensis

pulcherrima

byronii decipiens

rugosa corrugata, torrida

nigricans

Dr. C. Montague Cooke considers decipiens with its varieties
to be a subspecies of 'byronii. In retaining A. pulcherrima as
a species, I am following Dr. Cooke 's counsel, although I have
some doubt whether it should be given higher rank than a sub-
species of A. 'byronii. It is a case where there are numerous
local forms the variations of which overlap more or less, and
whether they 'are ranked as forms of one species, or are more
or less arbitrarily -assorted into several, is a matter of conveni-
ence in referring to the forms. The series shows such diver-
sity of differentiation that it would be rather meaningless to
lump all of the races under the one name, A. byronii.

Practically all of the Newcomb 'and Grulick shells were taken
at low levels, 'chiefly in places where there are now no forests.
The modern collector in this part of Oahu rarely finds tree-
snails below the forest fence, which follows the 1,000 ft. con-
tour from Waimea to Moanalua.

4. A. BYRONII (Wood). PL 27, figs. 1 to le, 3.

The shell is dextral, imperf orate ; pyramidal-conic with ob-

134 ACHATINELLA BYRONII.

tuse summit, solid, glossy. Color variable, but typically green
and light greenish-yellow in oblique streaks on the last two
whorls, having a faint green peripheral band and a dark
chestnut band bordering the suture below ; next earlier whorl
yellow with a chestnut band, nearly three embryonic whorls
pinkish gray; aperture white, the lip bordered with dark
brown. Sculpture of faint spiral strias on the embryonic
whorls, later whorls irregularly wrinkled in the direction of
growth-lines, under the lens showing very faint traces of a
fine oblique malleation. Whorls 6%, somewhat convex, the
last often very obtusely angular at the periphery. Aperture
strongly oblique, the lip thickened within by a strong rib near
the margin. Columellar fold moderate, white or tinted.

Length 20, diam. 11 mm. (typical size).

Length 18, diam. 11 mm.

Length 17, diam. 9% mm.

Oahu: Kalaikoa, Ahonui (Gulick).

Helix byronii WOOD, Supplement to the Index Testaceo-
logicus, or a Catalogue of Shells, British and Foreign, 1828,
p. 22, no. 30, pi. 7, f. 30. — Acnatinella byronii Wood, NEW-
COMB, Ann. Lye. N. H. of N. Y., vi, p. 318. — PFR., Monogr.,
vi, 165. — SYKES, Fauna Hawaiiensis, p. 306. — Achatinella me-
Unostoma NEWCOMB, P. Z. S., 1853, p. 132, pi. 22, f. 7 (1854).
— PFR., Monogr., iv, 521. — Achatinella limb at a GULICK, Ann.
Lye., vi, 252, pi. 8, TOa, 705 (Feb., 1858).— Helicteres byron-
ensis (Gray), BECK, Index Mjolluscorum, 1837, p. 51.

A. byronii, in its southeastern forms, is closely related to
A. viridans and A. decipiens. All have more or less corru-
gated varieties, and others almost smooth.

The type of A. ~byronii was denned only by a figure, but
this is of such excellence that no doubt of its identity can be
entertained. The shell was presumably brought to England
by Lord Byron, whose mission to Oahu in 1824 has been noted
in Vol. XXI, p. 157. All of his AcliatineUidcc were from near
the west end of the main range, Wahiawa to Kawailoa. The
present species does not occur in Kawailo'a district, but it is
to be found at Kalaikoa and Ahonui, places in Wahiawa dis-
trict, a few miles southward. I select Ahonui as the type

ACHATINELLA BYRONII. 135

locality, pi. 27, figs. 1, la representing typical shells. The
chestnut subsutural band is invariable in Ahonuj. and Kalai-
koa shells.

In a lot of 68 from Ahonui, Gulick coll., the colors are as
follows :

1. Typical, green-streaked (pi. 27, fig's. 1, 1«), 16 shells.

2. Yellowish or olivaceous tawny, clear or more or less
roughened and streaked -with black (figs. 1&, Ic), 18 shells.

3. Like 1 or 2, but having spiral bands, at periphery and
midway between sutures, sometimes not extending to the last
whorl (fig. Id), 33 shells.

4. One shell (fig. le) has a peripheral band only.

The above division is somewhat arbitrary, as the patterns
and colors blend in some specimens.

In Kalaikoa a large series from Gulick contains no bright-
green shells. Olive-green, with pattern no. 1 (above), and
pattern no. 2 and no. 3 predominate. There are a few dark
chestnut shells (pi. 27, fig. 3) . The shells are decidedly rough.

Achatinella limb at a Gul., from Ahonui and Kalaikoa is
identical with the typical green and yellow forms of byronii
shown in figs. 1 to Ic, having the same subsutural line, etc.
It is described as * ' striate, sometimes rugose, green or yellow ;
apex rose, frequently faded in mature specimens ; . . . peris-
tome black.' " It differs from A. melanostoma Newc., in
being more rugose, with rose-colored apex and subangulated
body- whorl." Gulick subsequently (P. Z. S., 1873, p. 91)
conceded the identity of limb at a with byronii.

A. BYRONII RUGOSA Newcomb. PI. 29, figs. 22, 22a ; pi. 27,
figs. 8 to 10a.

" Shell dextral, conical, glossy; whorls 6, rounded, mar-
gined above ; suture well impressed. Lip expanded, of a red-
dish-brown at the margin, thickened near the edge. Columella
white, short, twisted, and with a strong callus spread over
the umbilicus. Aperture subrhomboidal. StrisB longitudinal,
numerous, and on the last whorl rugose. Color of epidermis
of a deep green, a light or dark umber, sometimes alternating
and arranged with the striae, with or without pale green trans-

136 ACHATINELLA BYRONII.

verse lines. Length fourteen-twentieth^, diam. eight-twen-
tieths inch ' ' (Newc.).

Oahu: Ewa (Newcomb) ; Waimalu, "Waiawa, Waipio (Gu-
lick).

Achatinella rugosa NEWC., P. Z. S., 1853, p. 138, pi. 22, f.
22, 22a, 1854.— PFR., Monogr., iv, 521.— SYKES, Fauna Ha-
waiiensis, p. 309. — THWING, Orig. Descript., pi. 1, f. 24. —
Bulimella rugosa LYONS,, Hawaiian Almanac and Annual for
1892, p. 104, pi. 1, f. 2.

Newcorab's original figures are reproduced, pi. 29, figs. 22,
22a-. They are like the shells collected by Gulick in Waiawa
valley (pi. 27, fig. 8), which is probably what Xewcomb meant
by" Ewa."

This race inhabits valleys and ridges mainly southeast of
those occupied by byronii. It differs from that by having the
last whorl usually more strongly corrugated, the lip-callus
tit inner, the whole Up of a brownish flesh-tint, at least typi-
cally, and the shell lias the gloss of fresh varnish.

There are, of course, specimens which taken by themselves
could hardly be classified correctly; occasional individuals
may be smooth or nearly so. The occurrence of almost black
(brown-black or green-black) shells in most colonies of rugosa
is characteristic, since byronii is rarely if ever so dark. The
embryonic whorls are grayish-buff, flesh-colored or white.

The color-patterns are: (1) greenish-yellow, profusely
streaked with green, or yellow streaked with brownish-olive,
either plain or with two spiral brown bands and a subsutural
band, (2) last whorl black or nearly so. There are also inter-
mediate specimens connecting the darkest and lightest.

In a lot from Waimalu there are no banded shells (pi. 27,
figs. 9, 9a). This place must be at or near the extreme south-
eastern limit of the species. Waiawa shells are very often
banded, and with a small proportion of the blackish form (pi.
27, fig. 8). In a lot from Waipio the blackish- chestnut form
predominates (pi. 27, figs. 10, 10a). All the preceding are
from the Gulick collection, and probably all of them were
taken at rather low elevations.

Length 19.3, diam. 10.2, aperture 9 mm. ; 6% whorls. AVai-
pio.

ACHATINELLA BYRONII. 137

Length 17.5, dia.ni. 10 mm. Waipio.

Length 18.2, diam. 9.4, aperture 8.7 mm. Waiawa.

Length 16.3, diam. 10, aperture 8.8 mm. Waiawa.

In Dr. C. M. Cooke's collection there is a lot -of 9 specimens
(no. 1828-30) agreeing entirely with some of Gulick's Wai-
nialu rugosa (such as pi. 27, fig. 9a), said to have been col-
lected in " a very small and extremely isolated clump of trees
on the extreme S. slope of the Wai ana e range, ' —in the region
of Palihua. As the collector was a person knowing noth-
ing of shells, I cannot help suspecting that he got a Koolau
range lot confused with lids Waianae catch. The shells are
greenish-black with the first neanic whorl green or olive
streaked on a yellow ground, embryonic whorls deep livid
purple in some, white in other examples. They have the
gummy gloss of rugosa.

1. Sub variety capax Pils. & Cooke. PL 31, figs. 7, la, 11.
At the head of Waiawa gulch, along the Waiahoie- Waiawa,
ditch trail, 14 mile from top, Mr. Spalding collected large,
robust specimens, less wrinkled than the typical form. The
shell is streaked with olive 'and yellowish-olive on a lime-green
ground, often with chestnut 'bands at periphery and base.
The lip is either pure white or narrowly bordered with purple
or brown; embryonic whorls either flesh-pink or bicolored, a
brown zone above, white below. They were on banana and
ieie. The colony was found to extend as far as the Kipapa
division ridge, and probably goes farther north. There is but
little variation in color and shape, and it is invariably dextral.
The patterns are shown in figs. 1-71}.

Length 21, diam. 12.5, aperture 11 mm. ; 6 whorls.
Length 19, diam. 10.5, aperture 9.2 mm.

2. Subyariety ivaimanoensis P. £ C., new. PI. 31, figs. 9 to
9f7. On the crest of the Waimano-Manana ridge, a half mile
to a mile 'above the locality of A. t. cookei, 'there is a dextral
pulcherrima-like shell, with ground of almost white, buff, or
greenish yellow, plain or with chestnut bands in the typical
rugosa positions, or variously split into lines ; lip white or pale
lilac. Surface much smoother than in rugosa. Embryo
white, or in the darkest shells, light brown or bicolored. All
seen are dextral.

138 ACHATINELLA BYRONII.

Length 17.8, diam. 10.7, aperture 9 mm.

Length 15.5, diam. 9.5, aperture 7.5 mm.

Through the darkest specimens, this race seems to connect
with the more wrinkled small form of rugosa, no. 3.

3. At the lower edge of the above colony, just above the
locality of A. cookei, there is a diminutive race similar to the
preceding in shape, but more strongly wrinkled, with the
color-patterns of subvar. capax ; also a few very deep chestnut
shells. The narrow lip is purple, and the upper whorls flesh
pink. Length 17, diam. 9.8, aperture 8.3 mm., o1/^ whorls.
PI. 31, figs. 8, 8a.

Forms 2 and 3 were collected Feb. 17, 1913, in company
with Messrs. Spalding and Merriam.

4Z). A. BYRONII NIGRICANS U. Subsp. PI. 31, figS. 10, 11, 12.

The shell is sinistml, oblong-conic, rather solid. Embryonic
3 whorls snow-white, the next whorl brown, or brown and
green, with a light band, last ivhorl intensely Hack above,
but on close inspection showing very indistinctly a dusky
greenish-yellow band at periphery and a wider one on the
base. The surface of the last two whorls is rather coarsely
wrinkled, and has a brilliant gloss. The aperture is very
oblique, ovate, white OF with a faint lilac tint within; peris-
tome has a narrow callous rib within of a fine purple color,
changing to violet at the lip-edge. Columellar fold purple
with white tip or crest, Parietal wall deep purplish brown.

Length 17.4, diam. 11, aperture 9 mm. ; 6% whorls.

Length 21, diam. 12.3 mm.

Waimano-Manana ridge at about 1400 ft. elevation, in a
very small area along the summit trail (Spalding, Merriam,
Pilsbry, Wilder).

This form differs from rugosa Xewc. by its capacious form,
coloration and sinistral coil. The last character would not
be of much significance were it not that the whole byronii-
rugosa series is dextral, in hundreds of individuals which have
been examined from many localities. A snow-white embryo
is also occasionally seen in rugosa. A. I), nigricans is doubt-
less a derivative from rugosa.

ACHATINELLA LILA. 139

It lives in a clump of lehua trees alongside of the trail,
about 1000 yards down the ridge from the locality of
A. turgida cookei. The colony is a very small one.

In Mr. Wilder 's collection there are some nigricans from
the type colony distinctly banded with green on the last whorl.

Mr. Spalding also found three specimens on a large moki-
hana bush on a lateral spur of the ridge 'about 100 yards
mauka from the locality of A. t. cookei. One of -these speci-
mens, fig. 10, is larger than those from the lower station. The
upper post-embryonic whorl is green, banded with brown.

It is a rare and handsome shell, possibly deserving specific
rank. There is a black form of A. turgida which has an as-
tonishing resemblance to nigricans, but differs by being
smoother with the real turgida apex, that of nigricans being
quite different,

So far as known, this subspecies is restricted to very few
small tree®, in two spots less than a mile apart.

5. A. LILA Pilsbry, n. sp. PL 31, figs. 15 to 15 d.

The shell is sinistral, ovate-conic, thin but strong, nearly
smooth, brilliantly glossy. The embryonic whorls are burnt
sienna brown (weathering to whitish in adult shells) or some-
times there is a light median zone. Last whorl either (fig.
15cZ) uniform blackish chestnut, or (fig. 15) having a chest-
nut peripheral band and baso-columellar patch on a yellow
ground, or (figs. 15a, 6) like the last but with a green band
midway between periphery and suture, or (fig. 15c) with
sutural and peripheral bands and baso-columellar patch of
yellow on a chestnut ground. There are also a few speci-
mens more or less intermediate between the patterns of figs.
15 a, 5, d. The aperture is moderately oblique, white or
faintly lilac within ; peristome acute, very little or not per-
ceptibly thickened within; columellar fold strong, purple or
white.

Length 17, diam. 11, aperture 9 mm. ; 5% whorls.

Length 16.8, diam. 10.2, aperture 8.8 mm.

Length 16, diam. 15.3 mm.

Oahu : crest of the Waimano-Manana ridge at junction with

140 ACHATINELLA PULCHERRIMA.

the main range, running from the summit down about three-
fourths of a mile along the ridge trail; in axils of ieie (Spal-
ding, Merriam and Pilsbry). Types no. 108066 A. N. S. P.
Go type in Bishop Museum.

This charming shell has a great resemblance to A. abbreviata
of the eastern end of the range, from which it is separated by
about a fourth of the length of the island, wherein no re-
lated forms have been found. A. abbreviata is invariably
dextra'l, A. lila sinistral. There are also differences in the
patterns of the banded forms. It is likely that A. lila is more
closely related to A. nigricans, and therefore a derivative of
the byronii group of species, and not directly related to
A. abbreviata. The color-pattern 15c is a "negative" of 15a,
the light and dark areas being reversed. In treating of the
genus Liguus I have noted similar instances.

In the type lot the 'colors are in the following numbers:
Pattern of fig. 15, 1 specimen; figs. 15a-, b, 8; fig. 15c, 5;
fig. 15<r7, 10. Four of the lot have the columellar fold white.

A. lila is rather abundant on its misty pe-ak, but soon be-
comes rarer and disappears as one descends the trail into less
cloud-hung levels. Near the 'summit only the black form is
found, the beautifully green-banded shells occurring lower on
the trail, where there 'are few of the black ones. The lateral
slopes of the ridge drop so precipitously that collecting is
almost restricted to what mav be reached from the trail. On

«-

the north side of the peak a pali of a couple of thousand feet
permits 'collecting only close to the summit. Named for L. L. C.

6. A. PULCHERRIMA Swainson. PL 27, figs. 2 (Ahonui), 5-5g,
6 (Wahiawa) ; pi. 31, figs. 13 (Kawaihalona), 14 (Hele-
mano).

"Shell ovate-oblong, subcylindical ; white or yellow with
broad bands of chestnut; margin of the lip brown. Var. a,
golden yellow, suture chestnut. This very elegant species is
about 0.8 inch long, 'and is much more slender than any of the
preceding. It varies somewhat in form, some specimens being
more ventricose than others, and also in the number and color
of its bands. The ground color is a deep and rich chestnut,

ACHATINELbA PULCHEREIMA. 141

with from one to three bands of orange, yellow, fulvous or
white. The marginal groove to the suture is very, close and
distinct in all. The golden yellow variety is without bands.
In all the colors -are remarkably rich and vivid ': (Swa/inson).

Oahu: Waialua (Newcomb) ; Helemano, Wahiawa, Kalaikoa
and Ahonui (Gulick) ; Helemano, low down, and eastern spurs
of Kawaihalona (Irwin Spalding).

Achatinella pulcherrima SWAINS, The Quarterly Journal of
Science, Literature and Art, The Royal Institution of Great
Britain, January to June, 1828, p. 86 ; Zoological Illustrations,
iii, Second Series, 1832-3, p. 123, pi. 123, f. 2.— PFR., Mono-
graphia id, 237 ; iii, 460 ; vi, 172 ; Conchyl. Oab. p. 280, pi. 67,
f. 30, 31. — Newcomb, Ann. Lye. vi, 308. — THWING, Orig. Des-
cript. pi. 1, f. 18. — Bulimella p., LYONS, Hawaiian Almanac
and Annual fo<r 1892, p. 105, pi. 1, f. 6. — Achatinella lorata
Fer., REEVE, Conch. Icon. pi. 1, f. 6, 1850. — Achatmella
melanostoma NEWC., P. Z. S. 1853, p. 132, pi. 22, f. 7.—
Achatmella multicolor PFR., in part, P. Z. S. 1855, p. 4, pi. 30,
fig. 11 (not fig. llfl). — Achatinella mahogani GULICK, Ann.
Lye. Nat. Hist, of N. Y. vi, p. 254, pi. 8, fig. 72.

This specie's differs from byro-nii chiefly by its smooth sur-
face. The last whorl is more evenly rounded, -while in byronii
it is often subangular. The pattern of Swainson's left-hand
figure was selected by Pfeiffer as typical. It is shown in our
pi. 27, fig. 5c ; also pi. 31, fig. 13. There are two broad chest-
nut zones on a yellow ground. This happens to be a rather
rare color-form which however occurs in Gulick 's Wahiawa
series (fig. 5c), and has been collected by Mr. Spalding on the
eastern spurs of Kawaihalona (pi. 31, fig. 13). I take the
latter valley as type locality, since it was certainly from shells
taken in that neighborhood that Lord Byron's lei of Achat-
inellas was made. The lip is obtuse, black-brown, with a
whitish rib within. The glossy surface has exactly the char-
acter of that of nymph a, etc. All of the large series examined
are dextral. Color-forms of a lot from Wahiawa (Gulick),
are as follows.

PI. 27, fig. 5c. Two broad chestnut zones on a yellow
ground ; sutural line usually chestnut.

142 ACHATINELLA PULCHERRIMA.

PI. 27, figs. 5d, 5e. Two narrow chestnut bands or lines;
rarely some additional lines. One or both lines may be want-
ing on the last whorl.

PI. 27, fig. 5/\ Yellow, without bands except the sutural
(Swainson's var. a) ; or with a white sutural border, fig. 5g.

PL 27, fig. 5, 5#, 5&. Chestnut, with or without darker
bands.

Length 18.5, dizain. 9.3 mm.

Length 19, diam. 10 mm.

Length 20, diam. 11.2 mm.

Length 17, diam. 10 mm.

In Helemano and Waialua district, the northwestern limits
of pulclierrima^ the same forms occur. Those of Kalaikoa do
not differ noticeably. In Ahonui part of the lots examine 1
are like those of "AVahiawa.' In others there are many whit-
ish shells with a gamboge tint, with or without dark lines.
Also a lot of shells varying from chestnut to sulphur-tinged
yellow, with sutural band only, many of them stouter in form
than typical pulclierrima. Length 18.5, diam. 11.5 mm.
These lots are from the Gulick collection. PI. 31, fig. 14 is
from low in Helenrano, coll. by Spalding,

Judging from the description and figure of A. melanostoma
Xewc., as well as from specimens received from him, it is a
synonym of pulclierrima, being based upon a somewhat dif-
ferent color form from that of Swainson, though occurring
with the typical color- form in AVahiawa district. Newcomb's
locality, Ewa, refers to the district, which takes in the ex-
treme southeastern part of the range of pulcherrima. The
original description follows.

A. melanostoma Newc. (pi. 29, fig. 7, reproduction of ori-
ginal figure). "Shell dextral, polished, solid conical; whorls
6, plano-convex, slightly margined above ; suture moderate,
accompanied with a black revolving band. Aperture sub-
ovate; lip black, thickened within. Columella short, brown-
ish-red, terminating abruptly in a strong plait or tuberosity.
Color yellowish or umber, with or without transverse brown
lines. Length fourteen-twentieths, diam. seven-twentieths
inch. Ewa, Oahu : (Neivcomb).

ACHATINELLA PULCHERRIMA. 143

According to a note from Mr. Sp'alding, melanostoma is
now found only in the northern 'branch of Kipapa,; but I do
not know just what form he found so far east.

A. multicolor Pfr., PI. 30, fig. 11, reproduction of original
figure. " Shell dextral or sinistral, imperf orate, conic-
oblong, solid, striate, and under the lens most minutely de-
cussated, glossy; buff or whitish, variously ornamented with
blackish-chestnut bands, rarely uni colored. Spire long-conic,
towards the apex somewhat attenuated, rather acute; suture
margined. Whorls 6, rather flat, the last about two-fifths the
total length, rounded basally. Aperture oblique, truncate in-
verted ear-shaped; columellar fold above, strong, twisted.
Peristome black-edged, the outer margin somewhat straight-
ened, narrowly expanded; columellar margin dilated, sub-
a dilate. Length 17, diam. 9 mm., aperture 8% mm. long, 4%
wide. Sandwich Is., Frick in Cuming coll. (Pfr.).

In the above description Pfeiffer obviously confused two
species. His two figures, reproduced in my pi. 30, figs. 11,
lla, belong to A. pulcherrima (fig. 11) and A. soiuerbyana
oviformis (fig. lla). Doctor Hartman recognized the latter
and added A. oviformis to the synonymy of multicolor; but
he included Pfeaffer's fig. 11, and therefore did not restrict
the species or realize its dual composition (Proc. A. N. S.
Phila. 1888, p. 30). Mr. Sykes referred Pfeiffer 's fig. lla to
A. oviformis, 'and fig. 11 to A. recta Newc., which he ranks as
a variety of A. byronii. As several of Pfeiffer's phrases, par-
ticularly "perist. nigrolimbatum' apply best to his dextral
shell, I would restrict A. multicolor to fig 11. It becomes
therefore an absolute synonym of A. pulcherrima, having the
typical color-pattern, which seems to have been more common
in the lower forests of three-quarters of a century ago than
it is now. A. recta Newc., which Mr. Sykes and others have
confused with the present group, belongs in the series of A.
livida.

Form mahogani Gulick (pi. 27, figs, 4 to 4<i, Ahonui). This
form has the smooth surface of A. pulcherrima, merely marked
with growth-lines, not corrugated or wrinkled. A subsutural
dark band is often wanting, or not darker than the rest of the

144 ACHATINELLA PULCHERRIMA.

shell, which is nearly free from dark streaks. The lip has a
narrower black edge than well-colored byronii, and has a white
rim within. The figures, from a large set in the Gulick col-
lection, show the range of color — from mahogany to yellow,
with or without bands. Except in the usually darker colora-
tion, I can see no reason for separating mahogam from pul-
cherrima. Gulick 's description follows. His figure is like
pi. 27, fig. 4a, which is exactly like the type-specimen.

" Achatinella mahogan-i. Shell dextral, imperf orate, tur-
reted, solid, shining, striate, reddish chestnut; apex obtuse,
rose; spire concavely turreted; suture lightly margined, mod-
erately impressed ; whorls 6, convex, the last inflated, with the
base black-chestnut and regularly rounded; columellar fold
•central, white, strong; 'aperture truncately auriform, white
within, with external margin scarcely reflected anteriorly,
arcuate; columellar margin dilated, adnate; parietal margin
wanting. Length 20.6, breadth 11, length of body-whorl 12
mm. Average weight 8 grains.

"On trees, Ahonui and Kalaikoa, J. T. G. It lacks the
black color of lip and suture which characterizes A. melano-
stoma Newc. 'and A. limlata Nob.? and 'also differs in the gen-
eral form and color ': (Gulick) .

A form which would be referred to mahogani if it occurred
on the south side of the main range is in the Gulick collection
(no. 125 ; no. 92461 A. N. S. P.) , with the locality Kahana. It
is not unlike Pfeiffer's figure of A. plunospira except that
'the base is not abruptly darker as in that figure. It differs
from Kahana detipiens by having light brown early whorls
instead of white.

6a. A. PULCHERRIMA NYMPHA Gulick. PL 32, figs. 13, 14 ; pi.
27, figs, 7, la, 71).

"Shell dextral, imperf orate, oblong-ovate, solid, shining,
very lightly striated, yellow or chestnut, sometimes lined with
brown." Interior and columellar fold white. Peristome
well thickened within close to the edge, white, or in banded
shells showing marginal spots. Dark chestnut shells have a
narrow dark lip-bo-rder. Length 18.5, diam. 10 mm.

ACHATINELLA DECIPIENS. 1-15

Oahu: Ahoiiui, Kalaikoa, Wahiawia, Helemano, Opaeula,
Kawailoa and Waimea (Gulick) .

<

Achatinella nympha GUL., Ann. Lye. Nat. Hist, of N. Y.,
vi, 251, pi. 8, f. 69, 1858.

Merely a variety of pulcherrima, from which it differs by
the white Up, -which however is dark-edged in some examples
of Gulick's lots. It differs from recta, by the 'Conspicuous
thickening of the lip. Some of the large lot in the G-ulick col-
lection have a narrow dark sutural band, but most of them
have none, and in some the suture has a white border. In a
lot from Wahiawa the patterns are in these numbers: yellow
ground bandless 6, with bands or lines 25 ; chestnut ground,
bandless 20, banded 9 shells.

According to Gulick, Helemano is the metropolis of
nympha. The specimens figured are from Helemano (pi. 32,
fig. 14) and Wahiawa (pi. 27, figs. 7, la, 76; pi. 32, fig. 13).
Fig. 13 is too deep in color. The shell is much lighter, -what
Rldgway calls barium yellow. PL 27, fig. 5#, might be re-
ferred to nympha as well as to pulcherrima.

Dr. Cooke, who examined the type of A. aptycha Pfr. (see
p, 54) , considers it a form of nympha. The figure is repro-
duced in my pi. 30, fig. 1.

7. A. DECIPIENS Newcomb. PL 32, figs. 2 to 6c.

' ' Shell conieaily-elongate, solid ; whorls 6, slightly rounded,
margined above. Aperture elongately-ovate ; lip subrenected ;
columella short, obliquely twisted, with an expanded callus.
Suture slightly (impressed; striae numerous, longitudinal, ob-
lique. Color white with yellow transverse bands, or yellow
with white transverse lines and longitudinal chestnut colored
stripes. Length sixteen-twentieths, diam. seven- twentieths
inch ' (Newcomb).

Oahu: Kahana (Newcomb) ; Hakipuu (Gulick for A. corni-
gata) .

Achatinella decipiens Newc., P. Z. S., 1853, p. 153, pi. 24, f.
68, 1854. — Achatinella corrugata GULICK, Ann. Lye. N. H. of
N. Y., vi, p. 248, pi. 8, f . 66, 1858. — Achatinella torrida GUL.»
Ann. Lye. N. H. of N. Y., vi, p. 250, pi. 8, f. 68, Feb., 1868.

146 ACHATINELLA DECIPIENS.

A. decipiens (including torrida and corru-gata) represents
A. byronii on the northeastern side of the Main Range in the
valleys from Waikane to Kahana. As we do not know what
forms occur on top, and whether there is complete intergrada-
tiou there, the races from the windward side are for the tame
treated as a species distinct from byronii It differs from
byronii by having the lip entirely or mainly white and by
having part of the patterns different.

Xewcomb's typical lot consisted of shells resembling pi. 32,
figs. 2, 2a, from examples received from him; fig. 2a repre-
senting the pattern of his type figure. The shell is yellow,
profusely streaked and lineate with deeper yellow and chest-
nut along growth-lines, having a white zone below the suture
and typically another at the periphery; spire mainly white,
lip and aperture also white. Under a hand-lens the surface
is seen to be quite distinctly sitriate spirally, with a finer de-
scending wrinkling such as is usual in Partyl/ina, but this
varies in distinctness a good deal, among the specimens from
Newcomb himself, on some of which it would not be noticed if
not especially looked for. Length IS, diam. 10 mm.

In other Kahana lots, collected by Gulick, the patterns vary
somewhat, (pi. 32, figs. 3 to 5). Olive-green or yellowish-
green may take the place of chestnut streaking. The surface,
typically almost smooth, may be rather distinctly corrugated
in some shells. These pass directly into the form called cor-
rugata Gulick.

Length 23, dram. 11 mm.

Length 17, diam. 10 mm.

Length 16.3, diam. 9 mm. (rough, stunted shell).

In another Kahana lot the prevalent color is light yellowish-
green, faintly streaked, and fading to whitish near the suture
and on the spire (pi. 32, fig. 35). Other shells are broadly
banded with chestnut, glossy like the preceding; lip tinted
(pi. 32, fig. 4a). Another pattern occurring sparsely in this
and other Kahana lots is dull yellowish, copiously streaked
with chestnut and blackish, and having two spiral brown
bands. It is almost lusterless and somewhat corrugated.

It is evident that Gulick did not collect in Newcomb 's type
colony of decipiens, which may have been of small extent.

ACHATINELLA DECIPIENS. 147

Achatinella corrugata Gulick from Hakipuu (pi. 32, figs.
7, la, 7b) seems to me to be almost identical with decipiens,
from which it differs chiefly by the usually more corrugated
surface, which shows only very indistinct traces of minute
spiral sculpture. The color-patterns resemble the streaked
forms of torrida except that none in the large lot has a white
subsutural band, such as torrida and decipiens often possess.
Gulick 's type is streaked with olive-green on a pale-green
ground (pi. 32, fig. la). Others are streaked with chestnut
on a yellow ground, with or without one or two spiral 'chest-
nut or blackish bands, while still other shells :are brownish -
black with a white summit. None has spiral white bands, and
very few have two chestnut bands. GuLick remarks that sims-
tral examples have been found. I have noticed one in the lot
of several hundred shells from his collection. The original
description of corrugata follows.

" Shell dextral, imperf orate, ovate, solid, shining, striated,
green ; apex subacute, pale ; spire convexly conic ; suture mar-
gined, moderately impressed; whorls 5%, convex, the last
rugose; columellar fold central, white, strong; aperture truii-
cately auriform, white within; perlstome thickened 'within,
white, sometimes margined with brown, with external margin
dilated, adnate ; parietal margin wanting. Length 18%, diam.
10 mm. ; average weight 5.3 grains. Hakipuu, Oahu, on trees.
The color is often nearly black, and sometimes reddish-yel-
low " (Gulick).

A rather large shell measures, length 19.1, diam. 10.8 mm. ;
a very narrow blackish one, length 18, diam. 8.5 mm. (fig. 7&) .

Another colony from Hakipuu was correctly considered by
Gulick to be transitional to decipiens proper. Some speci-
mens have a white sutural band, and occasionally there is a
white line at the periphery (pi. 32, figs. 6 to 6c) .

Variety ( ?) planospira Pfeiffer. PI. 30, fig. 8.

Achatinella planospira Pfr. has been placed in the syn-
onymy of decipiens by Newcomb, who is followed by Hart--
main, Sykes and Thwing. I have seen specimens of form cor-
rugata having the same pattern except that the summit is
light, while Pfeiffer 's figure shows it dark. The status of the

148 ACHATINELLA DECIPIENS.

form is therefore uncertain. It needs comparison with pul-
cherrima and mahogani. There is, however, a lot of maho-
gani-like shells in the Gulick collection (no. 125) from Ka-
hana, having the apex dark, and agreeing fairly well with
planospira.

The original account follows. A. planospira Pfr. (pi. 30,
fig. 8, photographically reproduced from Pfeiffer's plate).
; Shell dextral, imperforate, ovate-pyramidal; solid, closely
striate and under the lens very obsoleteily striate spirally,
glossy; blackish, ornamented with several narrow white or
buff bands. Spire conic, rather acute; suture linear, mar-
gined. Whorls 5!/o, flat, the last a little shorter than the
spire, obsoletely angular, rounded basally. Aperture oblique,
inverted ear-shaped. Columellar fold above, strong, twisted,
white. Peristome thickly white-lipped, the right margin,
spreading, narrowly expanded, columellar margin adnate.
Length 19, diam. 9y2 mm., aperture 9 mm. long, 4 wide. Var.
Z>, above blackish-chestnut, the last whorl tawny, with brown
lines. Sandwich Is., Frick in Cuming coll : (Pfr.).

Variety torrida Gulick. PL 32, figs. 8 to 12c.

The sinistral form of corrugata was described as A. torrida
by Mr. Gulick. The distinction from corrugata is rather too
vague to rank it as a subspecies, though there is a certain
amount of racial differentiation. Its patterns differ in many
specimens, and then most torrida are sinistral. The original
account follows.

" Shell sinistral, imperforate, ovate-conic, solid, shining,
somewhat rugose, green or fulvous; apex subacute, chestnut;
spire conic ; suture lightly margined, white, moderately im-
pressed; whorls 6, convex; columella with a strong white fold
near the body-whorl. Aperture oblique, truncately auriform,
white within ; peristome white, thickened within, with external
margin scarcely reflected anteriorly, slightly compressed;
columellar margin dilated, adnate ; parietal margin wanting.

Length IT1/-}, breadth 9% mm. ; average weight 4.5 grains '
(Gulick).

11 Var. I), with black spiral lines. I am in doubt 'Concerning
the limits of variation of this species ; var. & may be distinct '
(Gulick) .

ACHATINELLA DECIPIENS. 149

Oahu: Kahana, Kaaawa and Waikane (Gulick) ; Waiolu
(J. S. Emerson).

Waiolu may be considered the type locality, as here the
typical form (pi. 32, figs. 11, Ha, 11&) occurred. It is closely,
.streaked with rather dull-green or chestnut on a light ground,
the streaks sometimes confluent. A wide subsutural white>
band (often denuded of cuticle) is frequently present. The
apex is not chestnut, as Gulick states, in any of the lot of over
a hundred from Ms collection, from all the localities men-
tioned above. Six out of a set of 56 from Kahana are dex-
tral, all others seen being sinistral. Specimens with streaked
pattern are almost mirror images of A. decipiens, but they
differ by having the aperture a little more oblique, the lip is
usually a trifle less thickened within, and the surface in the
average torrida is more corrugated. The lip usually has a
fleshy or brownish border. Specimens from Kaaawa (pi. 32,
fig. 9) are similar to those of Waiolu.

A small set from Waikane is strongly corrugated, heavily
streaked, with a dark band below the suture (pi. 32, fig. 8) .

In Kahana the shells are variable (pi. 32, figs. 10 to lOc?) .
Those having blackish streaks on a yellowish or chestnut
"ground resemble forms of other valleys. Others have one or
two black-brown zones and a white sutural band ; and in some
the blackish-chestnut color spreads over all but the earliest
whorls. The surface has very little gloss in this lot, Kahana
specimens in 'coll. C. M. Cooke are similar except that the
ground-'Color is greenish and the surface glossy.

Length 18, diam. 10.7 mm. Kahana.

Length 18.5, diam. 10.5 mm. Kahana.

Length 17, diam. 10 mm. Kahana,

Length 17.5, diam. 10 mm. Waiolu.

Kaaawa-Hakipuu division ridge (pi. 32, figs. 12 to 12c).
In a very handsome series collected by Mr. Spa.ldlng the em-
bryonic whorls are light-brown, differing thus from the large
series of torrida in Gulick 's collection, in which the early
whorls are white or nearly so. The last whorl, fig. 12, is green,
streaked with a much darker shade, or similar, with two black
"bands and a white sutural band. Fig. 12c, black with a yellow

150 ACHATINELLA DECIPIENS.

band and white line at the suture. Fig. 12a, chocolate. The
lip is dark-edged, or in form fig. 12c has two spots at the ter-
minations of the bands. The surface is rather rough. This
colony is mainly sinistral, but there are some dextral shells.

la. A. DECIPIENS KALIUWAAENSIS Pilsbry & Cooke, n. subsp.
PI, 32, figs. 1, la, 16.

The shell is similar to decipiens but smoother, more glossy,
with only obsolete traces of sculpture. Color various: (a)
White with a brownish sutural line, (&) Olive-ocher with a
white zone below the suture and a white line at periphery.
(c) Olive-ocher with a chestnut subsutural line or none, and a
pia'le peripheral line. Lip only narrowly thickened within,
white.

Length 19.2, diam. 10, aperture 9 mm.

Length 17, diam. 9.7, aperture 8 mm.

Length 15.3, diam. 8.9, aperture 7.6 mm.

Eastern ravines of Kaliuwaa, type loe., also central and
western ravines (Irwin Spalding).

This new form has such a relation to decipiens as pulcher-
rirna to byronii. The locality lies some distance northwest of
that of decipiens, in a district where the other Bulimellas dif-
fer from the Kahana forms. The coloration is much like that
of typical decipiens. The white-banded form may be consid-
ered the typical pattern. The cotypes are in coll. A. N. S. P.
and Bishop Museum, collected by Mr. Spalding who has a
long series. It is also in the Thaanum, Thurston 'and some
other recently gathered collections.

75. A. DECIPIENS ( ?) SWAINSONI Pfeiffer. PI. 30, fig. 13.

" Shell sinistral, imperforate, ovate-conic, solid, nearly
smooth, glossy ; whitish, delicately streaked with tawny. Spire
conic, apex fulvous, rather acute ; suture margined. Whorls
5!/2, a little convex, the last nearly equal to two-fifths the
length, swollen below the suture, rounded >at base. Aperture
slightly diagonal, reversed auriform. Oolumellar fold super-
ior, strong, nodiform; peristome bordered with black-brown,
the external margin a little reflected, thickly labiate within j

ACHATINELLA ROSEA. 151

columellar margin thick, flexuous, adnate. Length 20, ddam.
11 mni.

"5. Greenish buff, the last whorl chestnut anteriorly :

(P/r.).

Sandwich Islands, Frick, in coll. II. Cuming (P/r.).

Achatinella sivainsoni P^R., Pro<c. Zool. Soc. Lond., 1855,
p. 4, pi. 30, f . 13.

Dr. Cooke, who examined the type, thought this a siiiistral
individual of A. pulcherrima. " Newcomb suggested that this
might be only a form of A. sordida ; it appears however to be
distinct, being broader, brown in general coloration, and hav-
ing a brown in place of a white lip. It is a little doubtful
from its form if it be correctly placed in this group [Acha-
tinella s. sir.], but the sections are very artificial : (Sykes,
Fauna Hawaiiensis, p. 304). As I have not seen the shell, I
cannot properly express an opinion; but from the figure it
seems as near the torrida form of decipiens from the Kaaawa-
Hakipuu ridge as anything, and I put it here for want of a
better place. It may be noted that several enigmas of Frick-
Pfeiffer origin were from the Koolauloa district. We know
very little about Frick, but from what he got we may infer
that the country between Kaneohe Bay and Kahuku was one
of his collecting grounds.

SERIES OF A. BULIMOIDES.

Rather large, compact, capacious shells, smooth, never
streaked with green, usually banded. This series comprises
numerous forms on both sides of the western half of the Main
Range.

8. A. ROSEA Swainson. H. 34, figs. 1 to 85.

' ' Shell reversed, pale rose-color, with obsolete white bands.
I place this, for the present, as a variety of the last [A. ~buli-
moides] to which, except in being reversed, it bears a 'dose
resemblance in size, form and general habit. It is entirely
of a pale and delicate rose-color, with two obsolete bands of
white on the body-whorl. The margin of the lip and columella
are of a deeper rose- color, and the aperture white. It should

152 ACHATINELLA ROSEA.

be observed tha.t the marginal [subsutural] groove, which is
scarcely perceptible in the last, is in this very distinct '
(Sivainson).

Length 22.3, diam. 13.5 mm. ; G1/^ whorls.

Length 23, diam. 13 mm.

Length 19, diam. 12 mm.

Oahu : Helemano and Poainoho to Kaukinehua ; formerly,
according to Gulick, from Kawailoa to Ahonui.

[A. bulinwides] var. ? — (rosea-) SWAINSON, The Quarterly
Journal of Sci., Lit. and Art, January to Jiine, 1828, p. 85. —
Achatinella rosea SWAINS., Zoological Illustrations, III, 2d
series, p. 123, pi. 123, f. 2. — REEVE, Conch. Icon., vi, pi. 4, f.
28.— PFR., Monogr., ii, 236; iii, 456; iv, 518; vi, 163.—
THWING, Orig. Descript., p. 93.

A. rosea is invariably sinistral. It further differs from A.
biilimoides by being smoother, more glossy. The colour varies
widely, but the older collectors found only the white and pale-
rose forms, which, from the large series in old collections,
must have been abundant in the time of Newcomb and Gul-
ick. Figures la to 2a represent the usual color-forms of the
species, the other forms figured being more or less rare. It
evidently extended lower than the dark-colored forms, in a
zone now barren by the recession of the forests. Gulick's
localities — none of them more exactly located than by the val-
ley or district — were Wahiawa, Waialua, Helemano (pi. 34,
figs. 1, la-), Opaeula and Kawailoa, all of the many speci-
mens being white or pale-rose or light rosy-brown, with roseate
lip, the colonies apparently homogeneous except for these fluc-
tuations of tint.

According to Mr. Spalding the headquarters of white or
nearly white rosea are in Helemano, but it is found from Ka-
wailoa to Kaukinehua in the Wahiawa district. There are
many gradations of tint between the white shells, the faint
rose, and the rose with white bands. An albino form with
the lip pure white has been taken by Mr. Spalding in a small
ravine in the eastern edge of Helemano, just above the forest
limit, similar to pi. 34, fig. 1, Helemano, Gulick coll.

A. rosea varies a good deal in size and shape. Typically
colored shells from Wahiawa, Gulick, measure:

ACHATINELLA ROSEA. 153

Length 22.3, diam. 13.6, aperture 11.5 mm. ; 6l/2 whorls.
Length 22.7, diam. 13, aperture 10.7 mm.

f

Specimens of a lot from ' Waialua, ' Thaanum, measure :

Length 21.7, diam. 12.3, aperture 10.5 mm. ; GI/? whorls.

Length 21.5, diam. 12, aperture 10.5 mm.

Length 18.3, diam. 12.4, aperture 10 mm.

Length 17.3, diam. 10, aperture 8.3 mm. ; 6 whorls.

Specimens from Ahonui in Mr. Thaanum 's collection are
white and pink, the usual coloration.

Messrs. J. S. and Oliver P. Emerson collected quantities of
dark-red rosea in the ravine in which the Waialua plantation
built its Kamoku reservoir, next west of Opaeula. This was
over fifty years ago, and the colony has been extinct these
many years.

On the middle ridge of Poamoho (between the western and
central branches of the stream), a colony about a mile above
the forest limit contains shells with the last whorl white, white
with two rose bauds, or rose-tinted with deeper rose bands
defining a median white zone. The lip and columellar fold
are deep rose color. PI. 34, figs. 2, 2a, coll. by Irwin Spald-
ing. The same form has been given me by Mr. Thaanum, coll.
by Mr. Kuhns. A mile further up, just west of the main
ridge, the shells have the summit dark; color of last whorl
pinard yellow with white suture ; the same with tawny bands
and spire ; yellow ocher with tawny bands and darker spire,
or flesh color changing to coral-pink near the suture. In all,
the lip is more or less blackish (pi. 34, figs. 6 to 6c, coll. by
Irwin Spalding). These shells are small, length 18 to 19.5
mm.

In Opaeula, at about 1,700 ft., Mr. Wilder found very dark
specimens, last whorl raw sienna with chestnut base, together
with pale green-yellow shells, both having the peristome and
summit dark (pi. 34, figs. 5, 5a) . It was probably lower down
that Gulick found the ordinary white and pink rosea from
Opaeula in his collection.

Similar but larger shells were taken by Mr. Spalding in the
eastern branch of the north fork of Kaukinehua (pi. 34, figs.
7). Other specimens from the same or an adjacent col-

154 ACHATINELLA BULIMOIDES.

ony are shown in pi. 34, figs. 8, 8 a, 85, taken by Mr. Wilder,
One measures, length 22, diam. 13.6, aperture 11.3 inm.

In a gulch west of Helemano Mr. Spalding found charming
pale green-yellow shells, with geranium-pink lip and faint
traces of two pink bands (pi. 34, fig. 4, no. 434 Spalding coll.).

In the bottom of Kawaihalona the shells vary from flesh or
salmon color to vinaceous in several tints (pi. 34, figs. 3, 3a,
coll. by Irwin Spalding; fig. 3a being no. 2167 of his collec-
tion).

Achatinella rosea is one of the most exquisite of land shells.
While the light-colored form is not uncommon in collections,
the ' ' green rosea/ the wine-colored and the dark forms are
rare even in Honolulu. They are among the prize specimens
of the finest collections. I did not myself collect in the rosed
territory, which has been thoroughly " shelled : ' in the last
few years, but the generosity of Messrs. Irwin Spalding -and;
W. D. Wilder has made it possible to illustrate the main var-
iations of this charming shell.

9. A. BULIMOIDES Swainson. PI. 33, figs. 1 to lj, 3 to 4.

" Shell ovate-oblong, subventricose, whitish with chestnut
bands ; spire thickened, the tip pale brown. Larger than the
last [A. livida], and of nearly the same form, but the spire is1
less thickened and more pointed at the apex. The ground-
color, in some specimens, is pale chestnut or ferruginous,,
banded with darker shades and another of pure white. In
others the upper half of each whorl is whitish, and the lower
chestnut, marked by darker bands. The suture is scarcely if
at all margined by a groove. The aperture and pillar white '
(Swainson).

Oahu: Waialee, Waimea, Kawailoa valley (Gulick). Head
of Kaipapau, on the north side of the range (Irwin Spald-
ing).

Achatinella bulimoides SWAINSON, The Quarterly Journal
of Science, Literature and Art, The Royal Institution of Great
Britain, January to June, 1828, p. 85 ; Zoological Illustrations,
iii, 2d Series, 1832-33, pi. 108, f. 1.— REEVE, Conch. Icon., vi,
pi. 1, f. 8, 1850.— HEYNEMANN, Malak. Bl., xiv, 1867, pi. 1, f.

ACHATINELLA BULIMOIDES. 155

2 (dentition). — PFR., Monographia, iii, 460; iv, 518; vi, 143;
viii, 216.— THWING, Reprint Orig. Descr., p. 92, pi. 2, f. 1.-
GULICK, Evolution, Racial and Habitudinal, p. 41, pi. 3, f. 1-5.

Swainson's figures agree exactly with, my pi. 33, figs. Ifr,
Id, from " Kawailoa,' which I take to be the type locality.
It ' ' has now become a very rare shell, ' ' owing to the destruc-
tion of forest, but I imagine that it will be found higher up,
in the region between Kawailoa, Waimea and Kaipapau.

A. bulimoides is always dextral, imperf orate or nearly so,
solid; the upper whorls are finely striate spirally and flesh-
tinted ; the lip is well thickened within and usually white or
yellowish, but brown in dark forms. There is never a sub-
sutural dark line as in brown-band forms of A. rosea, but the
suture is often bordered with an impressed line.

Specimens from Kawailoa measure:

Length 20.6, diam. 12.4, aperture 11 mm. ; whorls 5%.

Length 21.3, diam. 11.8, aperture 11 mm. ; whorls 614.

In a lot of 48 from " Kawailoa " (pi. 33, figs. 1 to Ij, Gu-
lick coll., 92489, A. N. S. P), the following color-forms occur:

1. White, the upper whorls pale-pink or flesh-tinted; peri-
stoane yellowish; suture usually margined (fig. 1), 6 speci-
mens, la. Similar, but having a 'brown streak behind the
outer lip, 5 specimens.

2. White with two chestnut bands bordering a peripheral
white belt, upper whorls fleshy, lip white or brownish (figs.
la, 6 ) ; 6 specimens.

2a. Similar but the peripheral belt and base tawny ; a white
umbilical patch or none (fig. le) ; 5 specimens.

3. Entire base purple-brown, a brown, band above the
peripheral white band; spire sometimes dark (fig. 1<Z) ; 9 spe-
cimens.

4. A white zone below the suture, elsewhere chestnut or
purplish-brown, lip often brown (fig. 10) ; 14 specimens.

5. Purplish-brown with fleshy suture and summit (fig. li) ;
2 specimens.

6. Base whitish, upper part purplish-brown (fig. l/i) ; one
specimen.

7. Basal patch and peripheral band dark chestnut, a light
chestnut zone above (fig. Ic).

156 ACHATINELLA BULIMOIDES.

A small lot from Waimea 'oonsasts of forms figs. Id, \g and

In Waialee the color-form fig. Ic occurred, but only dead
shells 'are in the lot taken by Gulick in the early Fifties, so
it was probably extinct there at that time.

Across the range, near the head of Kaipapau (pi. 33, figs.
3 to 3d, coll. by Mr. Spalding), there is a colony having some
patterns identical •'with Kawailoa bulimoides. The embryonic
shell is usually dull purple ; last whorl variously banded with
dark chestnut and white, or yellowish-brown with or some-
times without bands; sutural border white or like the ground
color. The lip is white or sometimes dark-edged; and in a
large series taken, it is invariably dextral. This form has been
called vidua by some collectors, but I believe erroneously. It
is a slightly modified bulimoides, and not, I think, so closely
related to the Bulimellas of Kaliuwaa -as to those across the

mam range.

Mr. Gulick had this form from an adjacent locality. One
dead shell labeled " Hauula/ ' pi. 33, fig. 4, is now no. 92445,
A. N. S. P.

9a. A. BULIMOIDES MISTURA Pilsbry & Cooke. PI. 33, figs. 5-
5c, 6-7.

In Kaliuwaa valley and the ridge eastward along the Castle
trail, several colonies have been found by Messrs. Spalding,
Thurston and others, which 'are not directly referable to ovata,
obliqua, oomorpha or spadicea, while they belong to the same
group of local races. The shell is either sinistral or dextral;
an obesely ovate shape prevails; embryonic whorls are invar-
iably colored, usually chestnut, darker than in ovata and ob-
liqua; the lip-callus is rather narrow and the lip is narrowly
chocolate-bordered outside and within. The color varies in
different colonies, as follows:

PL 33, fig. 7, coll. by Spalding. On the edge of Punaluu
valley, where the Castle trail passes over the crest into Kaliu-
waa, a pure colony lives in a clump of ieie. The color is pale
grayish blue-violet (Ridgway, pi. 35) varying towards lav-
ender or pearl blue in different examples, suture bordered

ACHATINELLA BULIMOIDES. 157

with a white line or a band, another sometimes at the periph-
ery. The spire is usually darker, becoming chestnut or light-
brown at the summit. All but two of a large series in M'r.
Spalding's collection 'are dextral. Length 20, diam. 12.1,
aperture 10.2 mm. ; 6 whorls.

PI. 33, figs. 5 to 5c., coll. by Spalding. In Kaliuwaa valley,
about three-fourths of a mile east from the following colony
there is a patch of bananas upon which the shells vary from
about the color of the preceding colony to ivory yellow or
cream buff with or without two dusky bands; sometimes
white above the upper band. The spire is usually bicolored,
summit indian red or brown. Shells both sinistral and dex-
tral, mainly dextral. Some specimens resemble obliqua and
oomorpha.

PI. 33, figs. 6, 6a, coll. by Spalding. At the bottom of Kaliu-
waa, along the stream back of a cabin, there is 'a colony sim-
ilar to the banded pattern of ovata except that the embryonic
whorls are darker, usually dark viiiaceous. This colony is
mainly sinistral, but contains some dextral shells.

95. A. BULIMOIDES SPADICEA Gulick. PI. 33, figs. 13, 13a.

The shell is sinistral, glossy, embryo mars brown, blackish
at the tip, becoming lighter on succeeding whorls, the last
whorl pale or light orange-yellow above, below the periphery
burnt sienna shading to chestnut downwards, and showing
faint traces of two darker bands. Sutural line white, extend-
ing to the apex. Lip chestnut, bordered outside with the
same; outer edge of the columella raised; columellar fold
white or tinted. The later whorls show only the weakest
traces of spiral sculpture.

Length 17.6, diam. 10.5, aperture 9 mm.; 6 whorls (type).

Length 17.5, diam. 11, aperture 8.5 mm.

Oahu: Kahana, on trees (Gulick, type loc.) ; Hakipuu (Gu-
lick) .

Achatinella spadicea GULICK, Ann. Lye. N. H. of N, Y., vi,
p. 247, Feb., 1858. — BALDWIN, Catalogue, p. 6 (" Hauula to
Kahana ").

" The white sutural band is continued up to the very tip

158 ACHATINELLA BULIMOIDES.

of the nuclear whorls in a manner quite different f roin what
is seen in the bands of A. obliqua and other species of this
group ; (Gulick) .

This small form is like mistura in having a dark summit,
but differs by the white sutural line running to the apex and
the color of the shell. I have been told that spadicea occurs
below the falls in Kaliuwaa, but I suspect that the shells from
there are mist lira. Spadicea is known by very few specimens.
There is one, the type, in coll. Boston Society of Natural His-
tory, described above and figured in pi. 33, fig. 13, and two
from it-he Gulick collection in A. N. S. P., one of which is fig-
ured, pi. 33, fig. 13(/. Also two shells from the same source in
coll. Bishop Museum. I do not remember seeing specimens in
other Hawaiian collections, and very likely the race is now
extinct. It has resemblance to some dark forms of A. rosea in
surface gloss and sutural border.

9c. A. BULIMOIDES OBLiQUA Gulick. PI. 28, figs. 8 to 9a.

" Shell sinistral, subperf orate, ovate, obliquely truncated
at the base, solid, shining, striated ; o'f ashy lead-color, with a
broad whitf band beneath the suture. Apex obtuse, of a yel-
lowish white color ; spire convexly conical ; whorls 6, subconvex,
the last large. Columellar fold central, white, strong. Aper-
ture oblique, truncately auriform, nearly white within. Peri-
stome 'white, thickened within; with external margin some-
what reflected, arcuate; columellar margin forming an ob-
tuse, sinuous ridge with a small umbilical cleft behind it ;
parietal margin thin. Length 23, diam. 13 mm. Average
weight 9 grains ' (Gulick).

Kahana, on trees (J. T. Gulick).

Achatinella obliqua GULICK, Ann. Lye. N. H. of N. Y., vi,
p. 245, pi. 8, f . 63, Feb., 1858 ; Evolution Racial and Habitu-
dinal, p. 41, pi. 3, f. 16/i-20fo. — Achatinella oomorplia GULICK,
Ann. Lye., vi, p. 246, pi. 8, f. 64.

Is allied to A. ovata Newc., with which it is associated
geographically, but differs from it by its broader form and!
white lip. The dark coloring of the body-whorl abruptly ter-
minates near the external margin of the peristome, leaving a

ACHATINELLA BULIMOIDES. 159

white border aibout an eighth of an inch in width 'around the
outside of the lip, whereas in A. bulimoides and other allied
species the color 'becomes darker on this portion of the shell
though the lip itself 'be white. Dextral 'specimens are very;
rare. The color of the darker portion of the shell varies in
different specimens from a dirty cream to slate color, but the
ashy color given in the description is the most common ' (Gu-
lick).

The color of the darker, lower portion of the shell varies
through several shades resembling purple-drab, viiiaceous
purple and slate-drab of R-idgway's Color Standards. These
tints are produced by films of white laid over a brown layer,
which may be exposed by scraping. The lip is white or nearly
so typically, but in one lot it is fleshy-brown. The white
border behind the lip mentioned by Gulick is by no means con-
stantly present; often the dark color runs to the lip, which
may have a narrow t>rown edge outside, but not so wide or
dark as in ovata. The summit is usually light yellowish-
brown ; sutural margin well defined, bluish- white. Lip some-
what expanded, as in A. ovata, having a moderate callus
within. The parietal callus is generally imperceptible. Be-
tween 3 and 4 per cent of the shells seen (are dextral. Length
22, diam. 13 mm. ; 6% whorls.

Occasionally the dark color is restricted to a peripheral
band, or it may disappear 'altogether, leaving the shell pure
white. More often some 'brown remains on the spire. In a
few shells the parietal callus is rather thick and brownish at
the edge. The above notes are from the large series collected
by Gulick.

A. 1}. obliqua was described from a colony in which the sin-
istral form very largely predominated. Mr. Gulick described
shells from a colony in which the dextral form prevailed as
A. oomorpha. While not exactly like typical obliqua, it does
not seem sufficiently differentiated to require 'a name. The
original account follows.

A. oomorplm Gulick. PI. 28? figs. 10 to lOc. " Shell dex-
tral, perforate, ovate, solid, shining, striated ; ash or ash-brown
with two obscure brown bands, white beneath the suture.

160 ACHATINELLA BULIMOIDES.

Apex rather obtuse, chestnut-brown ; spire convexly conical;
suture margiiiate, moderately impressed; whorls G1/^, convex.
Columellar fold central, white, strong. Aperture truncately
auriform, white within; peristome thickened within, with ex-
ternal margin slightly reflected anteriorly, arcuate, white or
'brown ; columellar margin reflected, detached, white ; parietal
margin very thin. Length 201/£, breadth 11% mm. Average
weight 7 grains. Kaliana, Oahu, on trees ' (Gulick) .

; Sinistral specimens are sometimes found which resemble
A. obliqua, but are readily distinguished by the darker color-
ing around and upon the lip. I have from Hauula a few spec-
imens which seem to belong to this species. Some of them
are nearly white ' (Gulick).

This dextral form of obliqua has the same peculiar colors,
drab, dull-brown overlaid with lilac, etc., but also sometimes
the chestnut-brown of A. ovata. It has another ovata char-
acter in the two-banded pattern of many shells of the typical
colony. This pattern is not found in the typical colony of
A. obliqua, judging by a lot of about 50 shells seen, though it
occurs in a smaller brown-lipped lot in the Gulick CGJ lection.
The form oomorpha often has exactly the coloration of typical
obliqua, except that the lip is brown both externally and
within. PL 28, fig. 10a is the typical oomorpha pattern ac-
cording to Gulick 's figured type, twojbanded over a streaked
ground. Shells also occur having the last whorl white. The
length varies from 19 to 22.5 mm., but most specimens before
me are smaller than obliqua.

9c7. A. BULIMOIDES OVATA Newcomb. PI. 28, figs. 1 to Id, 2 ;
pi. 29, figs. 2, 2a ; pi. 33, fig. 2.

' Shell dextral, elongate-ovate, polished, finely striated lon-
gitudinally. Color light flesh-colored above, last two whorls
white, obscurely banded with light-brown. "Whorls 6, convex;
suture slightly impressed, margined. Aperture subovate;
columella twisted into a plait, slightly callus; lip slightly re-
flected, dark-brown. Length 0.9 inch. Breadth 0.45 inch.
Variety a : shell white, bluish-white above, without coloring or
bands. Habitat, Waiauai, Oahu : (Netvcoml), May, 1853).

ACHATINELLA BULIMOIDES. 161

" Shell dextral or sinistral, elongately ovate; whorls 6,
slightly margined above, rounded; suture moderately im-
pressed. Aperture subovate, entirely margined with black;
columella short, plicate, strong and twisted ; lip thickened and
slightly expanded. Color of shell pure white or yellowish-
white, with or -without obsolete brownish bands above. Length
eighteen-twentieths, diam. ten-twentieths inch. Hab. Kahana,
Koolan, Oahu ' (Newcomb, 1854).

Oahu: Kahana (Newcomb) ; Kahana and Hakipuu (Gu-
lick) .

Achatinella ovata NEWCOMB, Ann. Lye. N. H. of N. Y., vi,
p. 22, May, 1853 ; P. Z. S., 1853, p. 130, pi. 22, f . 2, 2a, 1854.
— GULICK, Evolution, Racial and Habitudinal, p. 41, pi. 3, f.
6/1-15/t. — Achatinella Candida PFR., P. Z. S., 1855, p. 2, pi. 30,
f. 4; Monogr., iv, 519. — Achatinella vidua PFR., P. Z. S., 1855,
p. 3, pi. 30, f. 10; Monographia, iv, 522. Cf. Newcomb, Ann.
Lye. N. H. of N. Y., vi, p. 324. — A. ivheatleyi NEWCOMB, Ann.
Lye., vi, pp. 147 (1855) and 324 (1858), not described (see
under A. elegans ivheatleyana). — A. fricki var. d, PFR., Mon-
ogr., iv, 521 ; P. Z. S., 1855, pi. 30. f . la.

Dr. Newcomb included both white and banded forms in his
description and figures, which are reproduced on pi. 29, figs.
2, 2a. They occur in the same hybrid colony in Kahana, pi.
28, figs. 1 to \d ; pi. 33, fig. 2. The surface is not very glossy,
often rather distinctly striate. The apex is light colored, yel-
lowish, pale-brown or white. The suture has no dark border
below, and the impressed line defining the margin is usually
rather weak, sometimes wanting. Outside the lip has a dark
border behind, as in pi. 28, fig. 1&, though not always so fully
developed. The lip is distinctly expanded outwardly and es-
pecially below, and has a 'chocolate edge, both outside and
within. The internal callus is rather strong. In all color-
forms 'at Kahana the shell may be either dextral or sinistral.

Length 23, diam. 13 mm. ; 6% whorls.

Length 22.5, diam. 13.5 mm.

Length 21.5, diam. 12 mm.

Length 18, diam. 11 mm.

Banded specimens would probably prove smaller than white-

162 ACHATINELLA BULIMOIDES.

in the average. The following color- forms occur in a lot from
Kahana, Gulick collection.

1. White, initial whorls often yellowish, lip .chocolate (f. 1).
la. Same, but whorls of spire with, a dark band (fig. la).
16. Brownish-cream color, spire as in la (fig. 16).

2. "White or whitish, with two chestnut bands separated by
a peripheral white band; spire with a wide chestnut band
above the suture (fig. Ic).

2a. Similar, but base entirely chestnut, usually lighter than
the bands (fig. Id).

Forms la and 16 are blends between 1 and 2.

In a lot of 49 specimens from Hakipuu, Gulick coll., all are
dextral. Color-forms fig. 1 and no. 2a, chestnut streaked with
darker, with a broad white band below the suture (pi. 28, fig.
2, Hakipuu) predominate, though there are some of the pat-
tern of fig. Ic.

In another lot of 7 dextral shells in the Gulick collection
from Kahana the lip is white, slightly yellowish at the edge.
In three of them there are pale traces of brown banding,
chiefly on the spire ; the others being pure white throughout
(pi. 28, fig. 7).

A. Candida Pfr. is generally admitted to be merely the white
form of ovata. The original figure is reproduced on pi. 30,
fig. 4. The description follows. Achatinella Candida Pfr.
Shell dextral, imperforate, ovate-conic, solid, striatulate, a
little glossy, pure white; spire convexly conic, the apex
minute, black, rather acute; suture light, thread-margined.
Whorls 6i/2, rather flat, the last 'three-sevenths to four-ninths
the length, obsoletely angular, rounded at base. Aperture
oblique, truncate-auriform; columellar fold moderate, super-
ior, oblique, subcompressed ; peristome brown-violaceous,
strongly labiate within, the right margin expanded, columellar
margin dilated, flat, thick, adnate. Length 22, diam. 11 mm.,
aperture 10% x 5 mm. Sandwich Is., Frick (Pfr.).

It is likely that all the patterns of A, fricki Pfr., except pat-
tern a, were based upon ovata. Pfeiffer's figs, la, 76, repro-
duced in my pi. 30, figs. 7a, 76, certainly look to me like ovata.
I do not feel competent to pronounce upon fig. 7, which has

ACHATINELLA BULIMOIDES. 163

been taken as the type of fricki, as I have not seen the shell,
and the figure is not closely matched among the specimens I
have seen.

A. vidua Pfeift'er, of which the original figure is reproduced,
pi. 30, fig. 10, is in my opinion merely a very small or stunted
specimen of ovata. Dr. Neweomb, who examined the type in
Mus. Cuming, decided it ' to 'be a somewhat worn and faded
specimen of this species ' (A. ovata). The coloration is ex-
actly that of some shells in Gulick 's KaJiana series, such as pi.
33, fig. 2, and pi. 28, fig. Id out of the same lot. Adult shells
in this lot of ovata range from 19.3 to 22.8 mm. long. I do
not believe that vidua has any racial status. The description
follows : ' * A. vidua Pf r. Shell dextral, subimperf orate, conic-
ovate, striatulate, under the lens most minutely decussate,
glossy; deep brown, two-banded with white. Spire convexly-
conic, the apex somewhat obtuse; suture nearly simple.
Whorls 6, very slightly convex, the last a little shorter than
the spire, rounded basally. Aperture a little oblique, trun-
cate-oblong; columellar fold obsolete, slightly twisted. Peri-
stome strongly labiate, the right margin narrowly refiexed,
columellar margin dilated, subadnate. Length ISi/o, diam. 10
mm., aperture 9 mm. long, 4% wide inside ' (Pfr.).

The weakness of the columellar fold, noticed by both Pfeif-
fer and Sykes, can be paralleled in the large series of ovata
before me. The smallest ovata I have seen is 17^ mm. long,
dextral, with the pattern of pi. 28, fig. Ic.

A. BULIMOIDES ROTUNDA Gulick. PI. 28, figs. 3 to 6.

Very dark chestnut or chocolate, with a white or pale tawny
zone around the upper part of the whorls ; glossy ; suture very
distinctly margin-ate in the last 21/2 whorls, dark-bordered
above and below, apex dark at the tip. Whorls rather swol-
len, sinistral. Lip moderately thickened within, but less
than in A. ovata, deep brown. Columellar margin gener-
ally less raised outwardly than in ovata, the columellar fold
brown or white ; parietal callus a very thin film.

Length 21, diam. 13 to 13.6 mm. ; 6!/4 whorls.

Kaaawa and Kahana (Gulick), on the dividing ridge.

164 ACHATINELLA BULIMOIDES.

Achatinella rotunda GULICK, Ann. Lye. N. H. of N. Y., vir
p. 248, pi. 8, L 67, Feb., 1858.

As it occurs 011 the Kaaawa valley side (figs. 3, 4), this form
is so distinct that it might well be given specific rank. Some
Kahana lots however, figs. 5, 5a, 6, show every pattern linking
rotunda with ovata. Gulick remarks that ' ' there is a variety
with two black bands -which resembles certain forms of A.
ovata Newc., but is distinguished by its black suture, and its
thinner and more regularly arcuate lip. Its bands are also
deep black, while those of A. ovata are brown as in A. buli-
moides Swains.' Having before me several large lots from
Kahana labeled rotunda by Gulick, I find that the subsutural
dark border is as often absent as present, and the bands vary
in shade, often being quite as light as in ovata. Most of this
Kahana lot are more lengthened shells than the Kaaawa ro-
tunda, but some, such as fig. 6, are typical in shape. These
colonies are clearly ovata X rotunda hybrids.

A typical rotunda from the type locality, Kaaawa, is shown
in pi. 28, fig. 3, Gulick coll.

9/. A. BULIMOIDES GLABRA Newcomb. PL 29, fig. 25 ; PI. 33,
figs. 8 to 11.

" Shell -conically ovate, glossy, whorls 6, somewhat inflated,
margined above ; suture well marked. Aperture ovate ; lip
slightly subreflected, dark-brown edged with black, thickened
•within. Columella short, robust, expanded, 'and terminating
in a twisted plait. Color bluish-slate, much lighter on the
upper portion of the whorls; sometimes transversely banded
with white or -chestnut. Length nine-twentieths, width seven-
teen-twentieths of an inch : ' (Newc.).

Oahu : Kolaupoko (Newc.); Kawailoa and Waialee (Gu-
lick) ; Waimea (Spalding).

Achatinella gldbra NEWC., P. Z. S., 1853, p. 139, pi. 22, f.
25 ; P. Z. S., 1854, p. 310.— PFB., Monogr., iv, 520.— THWING,
Orig. Descript., p. 93, pi. 2, f . 2. — B [ulimella] glabia, Newc.,
HARTMANN, Proc. A. N. S. Phila., 1888, p. 29 ("Kawaiawa").

Newcomb 's type figure (pi. 29, fig. 25) shows a shell with
banded spire and light colored base. He subsequently (1854)
stated that A. glabra is always dextral.

ACHATINELLA BULIMOIDES. 165

Some shells received from Newcomb are whitish, becoming
streaked with purplish-drab on the last whorl, with a pair of
brownish- drab bands at the periphery, lip with a dark-brown
edge (pi. 33, fig. 9).

The specimens before me from Waialee (pi. 33, fig. 11),
Waimea (pi. 33, figs. 8 to 8c) and Kawailoa (pi. 33, figs. 10,
lOo-) show a good deal of variation in color; all collected by
Gulick.

1. Purple-brown, with a belt of light reddish-brown below
the suture, upper whorls reddish or pale, lip fleshy-brown.

2. Base reddish-chestnut, upper third of the last whorl yel-
lowish or white.

3. Base flesh-colored, upper part white, a dark band be-
tween, lip white.

4. Purplish-fleshy with indistinct whitish streaks, lip fleshy.
The preceding are Waimea shells, the same patterns occur-
ring in Kawailoa. In Waialee I note an additional pattern.

5. Similar to no. 3, but whorls of the spire banded; lip
brown. This is the typical pattern.

It will readily be seen that glabra inter grades in some of
its .color-forms with bulimoides, but both glabra and buli-
moides have other color-forms special to each. The area of
glabra overlaps that of bulimoides in part, and extends thence
to the western end of the range.

Newcomb 's type locality, " Koolaupoko, ' is an extensive,
district on the north side of that portion of the Main Range
north and northeast of Honolulu (koolau signifying the wind-
ward or northern side, as kona the lee or southern district, on
any of the islands). Later collectors have found nothing re-
sembling A. glabra in the eastern part of the island ; but Gru-
lick, Spalding and others have taken the species in valleys of
the northwestern end of the range. There can be no doubt
that Newcomb was mislead as to the habitat of A. glabra.

Var. ( ?) fricki Pfeiffer. PI. 30, fig. 7.

" Shell dextral or sinistral, subimperf orate, oblong-ovate,
solid, lightly striate, glossy, of very various colors; spire a
little convexly-conic, the apex acute; suture narrowly mar-
ginate ; whorls 6, a trifle convex, the last about equal to three-
sevenths the total length, rotund at base. Aperture oblique,

16G ACHATINELLA ELEGANS.

reversed ear-shaped; columellar fold above, twisted, strong;
peristome slightly expanded, Obtuse, labiate and bordered
with violaceous or black. Length 20-21, diam. lO1/^ mm. ;
aperture 10 mm. long, 5 wide.

" a. Isabelline, subfasciate with pale-brown, suture white.

" b. Bright chestnut, ornamented with darker 'and white
bands, sinistral.

" c. Gray-brown, ornamented with darker bands, white
above.

" d. AYhite or buff, ornamented with two or three black-
brown bands.' (Pfr-}

Sandwich Islands, Frick in coll. Pfr.

Achatinella fricki PFR., Proc. Zool. Soc. Lond., 1855, p. 3,
pi. 30, f. 7, la, lb- Malak. Bl., 1855, p. 5; Monographia, iv,
521. Cf. SYKES, Fauna Hawaiiensis, Mollusca, p. 307, under
A. glabra.

A. fricki has not been recognized by Honolulu concholo-
gists. Pfeiffer described and figured several varieties. Hh
first figure (see pi. 30, fig. 7), which may be ta.ken for his
type, has been considered to be A. glabra Newc. by Mr.
Sykes. I have not been able to match it at all closely among
the glabra I have seen. It differs from pulcherrima by hav-
ing a white sutural band. This band I have never seen in a
dark pulcherrima, and I therefore give fricki temporary place
as a variety of glabra, following Mr. Sykes.

Pfeiffer 's color-form b and those following are probably
different subspecies. His second figure (reproduced in pi. 30,
fig. la) has been referred by Sykes to A. ovata Newc., I be-
lieve correctly, as it agrees fully with some of the old ovata
received from Newcomb. Pfeiffer's var. b (see pi. 30, fig. lb)
is a sinistral shell, not exactly matched by any specimen I
have seen, and probably not the same as typical fricki. The
contiguous dark bands bordering a peripheral white band
show it to belong to the bulimoides group, and it is quite pos-
sible that this figure also is an ovata.

10. A. ELEGANS Newcomb. PI. 28, figs. 12 to 13d; pi. 32, fig.

15.
" Phell conically-elongate, polished, shining, rather solid;

ACHATINELLA ELEGANS. 167

whorls 6, plano-convex, margined above ; suture well im-
pressed. Aperture subovate ; lip white, expanded, subrs-

<•

fleeted, somewhat contracted in its center, thickened within;
columella short, flat and lightly toothed. Color light and
dark-brown alternating, longitudinally arranged in lineations
or broad patches; sometimes with a white sutural band and
an additional one on the body-whorl. Length eighteen-twen-
tieths, diam. eight-twentieths inch.' (Newc.)

Oahu: Hauula (Newcomib) ; Hauula, Kaliuwaa and Ka-
huku (Gulick) ; Hauula and Kaipapau (Baldwin).

Achatinella elegans NEWC., P. Z. S., 1853, p. 149, pi. 24,
f. 57, 1854.

This species differs from all forms of bulimoides by its
streaked coloration, a pattern not found in bulimoides. It is
also more slender, in the average, but occasional specimens
are as broad and compact as bulimoides.

It is said to be now extinct, having passed with the lower
forests of the Hauula region, but it was an abundant shell in
1850-55, when Newcomb and Gulick were collecting. Besides
the lots in these collections I have seen a beautiful series in
the collection of Mr. J. S. Emerson.

Large series from. Newcomb and Gulick, taken in Hauula,
show considerable variation in the pattern. Usually the shell
is streaked in the direction of growth lines with brown (wal-
nut-brown to brownish- vinaceous of Ridgway's Color Stand^
ards), the shade variable, but usually appearing overlaid
with white, sometimes dull chestnut-brown. The typical
form as figured by Newcomb has the streaks; cut by white
bands at suture and periphery as in pi. 28, fig. 13a, but often;
one or both of these white bands is lacking. Newcomb 's
description applies to the form shown in pi. 28, fig. 12. Be-
sides these prevalent patterns, there are a few additional
white spirals in some shells, and sometimes the dark streaks
are reduced or blurred. Rarely the streaks are confluent, the
brown color almost evenly diffused. There are also a few
pure white shells (fig. 13c) . In a set collected by Dr. New-
eomb I note rather indistinct dusky bands above and below
the periphery, sometimes parted by a light-brown peripheral

168 ACHATIXELLA ELEGANS.

band. The lip is sometimes white, more often fleshy-brown
or darker. The white band below the suture sometimes runs
to the very apex, as in A. l>. spadicea, but oftener not. The
early whorls are usually dull-brown. Dextral shells largely
prevail, but all of the color-forms are common to both dextral
and sinistral. In one lot from Hauula there are 82 dextral,
23 sinistral shells.

A set of 8 from Kahuku are very solid, coarsely streaked
throughout, without bands (pi. 32, fig. 15, coll. by Gulick).

An unusual color-form, pi. 28, fig. 13cZ, has a pattern re-
sembling some specimens of oomorpha or obliqua somewhat.
The streaking of elegans is only weakly shown in places on
the spire. The sutural white line runs to the apex, as in
spadicea and some specimens of elegans. In its compact
shape this shell is similar to several characteristically streaked
elegans in the same Hauula lot, It is obviously an extreme
form of elegans.

Color- variety inelegans n. var. PI. 33, fig. 12. The shell is
dextral, rather thin; glossy, rather distinctly striated spir-
ally to the last whorl, which is smoother. Embryonic whorls
corneous, following whorls cinnamon with white sutural
border, last whorl pale cinnamon, shading to darker at the
base, and having two orange-cinnamon bands, one above,
the other below the periphery; sutural band pure white, be-
ginning on the third whorl. Peristome a little expanded ait
base, very little thickened within, edged with dark-brown.
Columellar fold weak, whitish. Length 19, diani. 10, aperture
9.5 mm. ; 6 whorls.

Kaliuwaa, J. T. Gulick, no. 92504 A. N. S. P.

This shell was labeled A. spadicea by Mr. Gulick, but it is,
in my opinion, a form of the elegans stock with less modified
coloration than elegans. It does not seem closely related to
the Bulimellas of upper Kaliuwaa, above the falls.

10a. A. ELEGANS WHEATLEYANA n. var. PI. 287 figs. 11, lla,
115.

The shell is cinereous, blue-gray or slightly purplish, rarely
purplish-brown, with faint whitish streaks and white or pale-

ACHATINELLA ELEGANS AND FUSCOBASIS 169

brown band above the periphery, sometimes bisected by a
dark line; suture white-bordered; summit light yellowish.
Aperture pale blue within, the lip thickened within, with a
narrowly but distinctly expanded, acute brown edge. It is
also narrowly bordered with brown outside. The strong coiu-
mellar fold is white. Length 21, diam. 12 mm. or smaller,
length 19.2, diam. 11 to 12 mm.

Punaluu (Gulick).

This race approaches A. obliqua somewhat, but seems
nearer elegans. It has the maroon-brown or purplish-brown
layer overlaid with a streaked white film, giving various tints
difficult to name, but near dutch blue, deep madder blue and
slate purple of Ridgway's Color Standards. The whitish or
pale-brown band above the periphery is nearly always pres-
ent, and often more conspicuous than in fig. lla. Sometimes
(fig. Ha) there is a thin yellow cuticle, but usually none.
One lot of 8 from Newcomb, without habitat, consists of sinis-
tral shells. Another lot from Gulick has 3 dextral shells in a
total of 22. It is probably extinct now.

This form was named for Charles M. Wheatley, the well-
known collector of Phoenixville, Pennsylvania.

"A. wheatleyi' was mentioned twice by Newcomb: in
Annals of the Lyceum of Natural History of New York, vi,
p. 147, October, 1855, where he says " A. wheatleyi Nob. is
A. vidua Pfeift'er, ' and again in the same volume, p. 324,
September, 1858, " The A. vidua Pfr., which I had supposed
was my manuscript A. ivheatleyi, I find in Mus. Cuming to be
a somewhat worn and faded specimen of this species ' [A.
ovata] . As no characters were ever assigned to A. ivheatleyi
except by implication in the above remarks, we have no
option but to accept Newcomb 's published statement of what
his A. wheatleyi was, even though he sent out specimens of
what we are now calling ivheatleyana as A. wheatleyi.

SERIES OF A. FUSCOBASIS.

Ovate shells, smaller than those of the bulimoides series;
mainly sinistral, never marked with green, and having some
resemblance to the cast a series of Achatinellastntm. They
are almost entirely shells of the high ridges and peaks.

170 ACHATINELLA FUSCOBASIS.

11. A. FUSCOBASIS (E. A. Smith). PI. 35, figs. 1 to 4.

Shell ovate, sinistral, glossy ; white, the last whorl yellow-
ish, ornamented with a median zone and base of brown.
Whorls 6, a little convex, suture distinctly margined. Aper-
ture white ; peristome thick, brown, columellar fold strong.
Length 16, diam. 10 mm. High up on Mt. Kaala on the Mo-
kuleia side, on the island of Oahu, arboreal. (Smith.)

Oahu: Head of Kuliouou-Niu division ridge to Mt. Olym-
pus, -at the head of the Palolo-Manoa ridge (Spalding. Cooke,
Kuhns and Wilder).

Bulimella fuscoliasis SMITH, P. Z. S., 1873, p. 77, pi. 9, f.
15. — Achatinella fuscobasis Sin., THWING, Original Descrip-
tions, etc., p. 83. — Achatinella luteostoma BALDWIN, Proc.
Acad. Nat. Sci. Phila., 1895, p. 217, pi. 10, f. 7, 8 (Palolo to
Niu). — B. rosea, a small white variety with a yellow lip,
HARTMAN, Pro<c. A. N. S. Phila., 1888, p. 30, pi. 1, f. 4.

A. fuscoliasis has the white ground of A. t&niolata, but it
is invariably sinistral in a large number I have seen in var-
ious collections. It is also smaller, with the lip not so thick,
the columellar margin less raised, and it has a different range
of patterns. The reddish-brown color (mars orange, or burnt
sienna, rarely chestnut or light orange-yellow of Ridgway's
Color Standards) of the peripheral and basal bands, and the
brown peristome are also characteristic of the typical form.
The following color-forms occur in most colonies.

(a) Pure white with brown (liver-brown to dull-chestnut)
peristome. Typical color-form of A. luteostoma (fig. 2).

(&) Peripheral band and basal patch brown, the space be-
tween them white or yellowish-brown. Typical color-form of
A. fuscolasis (pi. 35, fig. 1, type-specimen).

(c) Same as 1), but a basal band in place of the patch (fig.

3).

(d) Same as 5 or c, except that there is a light-brown band
near the suture. In one shell from Mt. Olympus in the
Spalding collection the upper and lower bands are broad,
light-brown, the peripheral band narrow, darker.

(e) Whole base brown, dark or light (fig. 4).

Most of the specimens from Kuliouou (Spalding and

ACHATINELLA FUSCOBASIS. 171

Thaanum collections), fig. 2, are pure white, tout there are a
few of the patterns of figs, 1 and 3.

Mr. Smith's figured type-specimen, no. 110 Gulick coll.,
Boston Soc. N. H., measures length 15.2, diam. 9.3 mm. It
is shown in pi. 35, fig. 1. Other shells measure:

Length 16, diam. 9.2 mm.; 16x9.5 mm. (near top of Mt.
Olympus; pi. 35, figs. 3, 4).

Length 17, diam. 10.5 mm. ; 17.5 x 10 mm. ; 16.2 x 9 mm.
(Kuliouou).

The figured type of A. luteostoma, no. 65704 A. N. S. P.,
measures 16 x 9.7 mm., and one of Baldwin's examples 14.6 x
9 mm.

The several color-forms occur in the same colonies. I do
not know that any of them has been found as a pure race.

A. luteostoma Baldwin was based upon the white form of
fuACobasis, but Baldwin also included the banded shells in his
original account, which follows. " Shell sinistral, imper-
f orate, solid, ovate, spire convexly conical, apex obtuse; sur-
face shining, marked with fine growth-lines, under a strong
lens seen to be decussated by close, extremely minute spiral
stride, apical whorls smooth. Color white, with a reddish-
yellow lip. Whorls 5!/2, margined above, slightly 'Convex.
Suture moderately impressed. Aperture oblique, sdnuately
oval, white within. Lip obtuse, thickened within, •columellar
margin very slightly reflexed, extremities united by a very
thin yellowish -callus. Columella reddish-yellow, terminating
in a strong tortuous fold. Length 15% , diam. 9% mm-

' ' In occasional examples the basal portion below the
periphery is light-brown, and sometimes a bright chestnut
spiral band encircles the periphery : (Baldwin) .

The types of both fuscobasis and luteostoma were probably
from somewhere in the head of Palolo valley. The locality
" M't, Kaala " given in the original description of fuscoibasis
was undoubtedly an error, as no such shell has been found in
the Waianae range. A native boy probably brought the
species to Gulick, who did not himself collect high in the
mountains. The two original specimens of fuscobasis and the
figured types of luteostoma are now before me.

172 ACHATINELLA FUSCOBASIS.

A. fuscobasis is a species of the high ridge and peaks of
the main range. Northeast of Mt. Olympus, on the ridge
leading to Konahuanui, it passes into an undifferentiated
form having the brown or 'brown-tinted lip of fuscobasis and
the more capacious shape of lyonsiana (which also rarely
shows some brown on the lip, in the type locality) . An in-
spection of series in the collections of Messrs. Spalding,
Thurston and Wilder convinces me that the Honolulu con-
ch ologists are right in regarding lyonsiana as a variety of
fuscobasis.

lla. A. FUSCOBASIS LYONSIANA Baldwin. PI. 35, figs. 5, 6, 7.

" Shell smistral, imperf orate, solid, ovate; spire convexly
conical, apex obtuse ; surface shining, striated with delicate
growth-lines; apical whorls smooth. Color white, with two
reddish-brown bands, one encircling the base, the other pass-
ing around the periphery and revolving on the spire just
above the suture ; the two bands are sometimes confluent, and
often the shell is uniform white, without bands. Whorls 6,
narrowly margined above, somewhat 'convex; suture moder-
ately impressed. Aperture oblique, white within, sinuately
oval ; peristome wThite, rather obtuse, thickened within, basal
and columellar margins slightly expanded, extremities united
by a very thin callus ; columella terminating in a strong, tor-
tuous, white fold. Length 17, diam. ll1/^ mm-

' ' Animal : Mantle black, sometimes mottled with white,
margined with a narrow line of lighter shade. Superior and
under portion of foot light brown. Tentacles, tentacular
sheath, and front above very dark brown ' (Baldwin).

Oahu: Mt. Konahuanui, at about 3,000 ft. elevation (type
loc.) to the top, spreading on the ridge southeastward, where
it passes into fuscobasis.

Achatinella lyonsiana BALDWIN, Proe. A. N. S. P. Pliila,
1895, p. 218, pi. 10, f. 9, 10, 11— SUTER, t. <c., p. 239, pi. 11,
f. 52 (teeth).

A larger, more capacious shell than A. fuscobasis, wdth the
lip and <corurnella white. The patterns are :

(a} Pure white.

ACHATINELLA FUSCOBASIS. 173

(6) White, with upper, peripheral and basal burnt sienna
or chestnut bands, the upper usually weaker or .sometimes
wanting.

(c) A unique color-form in Mr. Spalding's collection (no.
2027) has the penultimate whorl deep chestnut, last whorl
with two narrow dark bands (ii and iii). The ordinary
brown-banded and white forms occurred in the same colony,
which is about half a mile from the top of Konahuanui, in
the main and typical locality.

The margin below the suture is invariably white, the sur-
face is very glossy, and spiral lines are scarcely visible even
on the early whorls; lip white, or rarely with a faint brown
tint. It differs from tccniolata by being sinistral, with the
outer border of the columellar lip decidedly less elevated.

Length 19.3, diam. 11.4 mm.

Length 18, diam. 12 mm.

Length 17.7, diam. 10.2 mm.

Dedicated to Prof. A. B. Lyons, formerly of Oahu College.
PL 35, figs. 5, 7 are cotypes, 65693-4 A. N. S. P. Fig. 6, from
the Irwin Spalding collection, taken half a mile from the top
of Konahuanui, somewhat approaches the following sub-
species.

. A. FUSCOBASIS WILDER: Pilsbry, n. subsp. PL 41, figs.

The shell is shaped as in A. /. lyo'nsiana. First two whorls
buff-gray; last whorl having a chestnut or blackish- chestnut
peripheral band and basal area, the latter split into bands in
some colonies. Above the peripheral band there is a white
band; below it a wider white or mars-yellow band. The rest
of the upper surface is mars^yellow, mars-orange or burnt-
sienna, usually excepting a sutural band or line of white.
Lip and aperture fleshy in various tints, the lip narrowly
thickened within. Columellar margin but slightly or not
raised ; parietal callus milky or transparent, very thin.

Length 17, diam. 10 mm.

Length 15.2, diam. 10 mm.

Summit of Lanihuli, at head of the Nuuanu-Kalihi ridge

174 ACHATINELLA PUPUKANIOE.

(Irwin Spalding, W. D. Wilder). Also " Mauna Kope " a
peak at head of the Kalihi-Moanalua ridge (Wilder).

This beautiful shell is the most ornate of the fuscobasis
series. It is isolated from lyonsiana and fuscobasis by the
deeply cut Nuuaiiu valley, which even at the pali is lower
than any form of fuscobasis has been found. The Lanihuli
colony is practically a pure race ; the splitting of the basal
patch into bands in one lot from Mr. Wilder, and the varia-
tion in the subsutural white border being fairly referable to
" fluctuating variation.' In the specimens taken by Spald-
ing on the Koolau side of the peak the basal dark area is un-
split (fig. 2).

On the peak at the head of the Kalihi-Moanalua ridge there
are shells with the typical pattern, a few with darker periph-
eral and basal bands on a tawny ground (pi. 40, figs. 3, 3&)
and others with a tawny ground and blackish-chestnut base
(A. castanea pattern), the subsutural border white. Both of
these are rare mutations.

12. A. PUPUKANIOE Pilsbry & Cooke, n. sp. PI. 35, figs. 14
to 17.

The shell is dextral, conic, the greatest diameter near the
base, solid, brilliantly glossy; uniform white, ivory yellow
with white sutural line, or either of these tints- with a burnt
sienna band immediately above, a wider and darker one
below the periphery. Suture margined but without a dark
line. Spire white. Lip not expanded, the edge brownish,
internal rib white, or sometimes the whole lip is pale-pink.
Columellar fold rather strong and abrupt, white.

Length 16.3, diam. 9.7, aperture 7.8 mm. ; 6 whorls.

Length 16, diam. 9.2, aperture 8 mm.

Oahu : Crest of the Waimano-Manana ridge, 1 to 1% miles
from its junction with the main range. Types 108068 A. N.
S. P., Pilsbry, 17-11-1913 ; also head of Waiawa valley, Irwin
Spalding.

This species, which I collected in company with Messrs.
Spalding and Merriam, has a great resemblance to A. fusco-
basis ; but that is invariably sinistral, and separated from tho

ACHATINELLA SOWERBYANA. 175*

habitat of A. pupukanioe by about a fourth of the length of
the island, wherein no similar form has been found. It also
resembles A. s. dextroversa, of the northwestern end of the
range. A. cast a, in some Waimano lots, resembles pupu-
kanioe in coloration, but is more lengthened and sinistral.
The resemblance is so 'dose, however, that I would refer tli3
new form to casta as a variety, were it not that old specimens
(such as pi. 35, fig. 15) have the columellar lip built up to
form a ledge, exactly as in Bulimella, and unlike any of the
very long series of casta I have seen.

The apical whorls are rather deeply eroded in all of the
adult shells from the type locality. In a young one they are
cartridge-buff with a white band above the suture.

In Mr. W. D. AVilder's collection a series from the 'type
locality contains several aberrant individuals : streaked with
burnt-sienna on a yellow ground, and others with the same
pattern interrupted by broad white zones at suture, periph-
ery and base.

A long series of smaller specimens was taken by Mr. Spali-
ing in a very limited area on the southern slope of the peak
at the northern head of Waiawa valley. White shells are in
the majority, but some have a basal band (iii), and in one
before me there are two faint bands (ii and iii), and thy
eroded embryonic whorls are brown. Two measure :

Length 14.3, diam. 8 mm.

Length 16, diam. 8, aperture 7.7 mm.

13. A. SOWERBYANA Pfeiffer. PL 30, figs. 14, 14« ; pi. 34, figs.
9, 10.

Shell sinistral, imperforate, conic-oblong, rather solid,
smoothish, having a gummy gloss; tawny buff, slightly
streaked with a deeper shade. Spire a little convexly-conic,
the apex subacute, suture margined. Whorls 6, very slightly
convex, the last a little shorter than the spire, rounded sack-
like at the base. Aperture oblique, inversely ear-shaped,
white within; columellar fold superior, strong, twisted, rose-
ate ; peristome rose-lipped, the outer margin shortly expanded,
columellar margin dilated, adnate. Length 18, diam. 9 mm. ;

176 ACHATINELLA SOWERBYANA.

aperture 8 mm. long, 4 wide. Sandwich Is., Frick. (Pfr.)

Oiahu : Punaluu to Kaipapau ; various varieties as far
northwest as Kahuku and Pupukea.

Achatinella sowerbyana PFR., P. Z. S. 1855, p. 4, pi. 30, i.
14; Malak. Bl., 1855, p. 65; Monogr., iv, 527. — BALDWIN,
Catalogue, p. 6 (Kalihi?). — Bulimella sowerbiana Pfr.,
HARTMAN, Proc. A. N. S. Phila., 1888, p. 31.

Achatinella oviformis Neweomb, PFR,, P. Z. S., 1855, o.
208; Monogr., iv, 540.— NEWCOMB, Ann. Lye. N. H. of N. Y.,
vi; p. 147 (as a synonym of A. soiverbyana Pfr.). — SYKES,
Fauna Hawaiiensis, p. 309. — A. multicolor PFR., in part, P.
Z. S., 1855, pi. 30, f. lla; cf. Sykes, Fauna Hawaiiensis, Mol-
lusca, p. 309.

Pfeiffer described var. 6 as " a little smaller, yellow- whit-
ish, the base chestnut or greenish.' His fig. 14a represents
this color-form, and is reproduced in pi. 30, fig. 14a.

A. soiverbyana is a handsome shell, very little known before
the last few years, but now familiar by the splendid series in
the collections of Hon. L. A. Thurston and Irwin Spalding.
It is invariably sinistral, typically wax yellow or sulphine
yellow, fading upwards, with the sutural margin brown, sum-
mit usually pale or white. The surface has the gloss of var-
nish, and the lip and columella are of a luscious pink tint
which has given it the local name of ' ' watermelon shell. ' It
is smaller and more slender than other smooth Buli-
mellas of the region. The lip-rib is narrow. It is no doubt
related to A. fuscobasis, of the eastern end of the range, but
I agree with all the island conchologists in holding the two
distinct. Like fuscobasis, it lives mainly on the heights.

Frick, who supplied Pfeiffer's type, gave no locality, but it
seems likely that he got the species in some valley of the
Kaipapau-Kaliuwaa region where it reached a lower level
than usual, and thus -came within the zone accessible to the
early collectors.

Plate 30, fig. 14 is reproduced from Pfeiffer's type figure.
PI. 34, fig. 9, from the bottom of the central ravines of Kai-
papau, and fig. 10, Kaliuwaa, near the Castle trail, are prob-
ably from as near where Frick took the original lot as a mod-

Achatinellidae

PLATE 27

la

1 b

1 d

Njjjjjg-

5e

:-

7a

7 b

10 a

Achatinellidae

PLATE 28

- H

lb

lc

9

m

13

7

9a

11

13 a

8

10

11 a

13 b

8a

10 a

i§^

•fetttttiM

lib

13

8b

10 b

13d

Achatinellidae

PLATE 29

2a

8

9a

10

10 a

11

12

14

15

17

18

19

20

21

22

22 a

23

25

Achatinellidae

PLATE 30

11

15

23

7a

12

45

48 a

49

50

14 a

48

52

Achatinellidae

3 a

4d

la

^

9 a

:-,

4a

f

7b

\

9b

15

IS a

8

9c

13

15 c

Plate 31

4c

6a

14

15 d

Achatinellidae

Plate 32

la

3a

<

Ib

Jb

6a

^L***

6b

-

2a

4a

10

7b

lOb

lib

12

i

I2b

12c

13

14

15

Achatinellidae

1 f

10

la

g

3a

A

5a

8

lOa

s**

v^*te^w>

Ib

1 h

3b

51)

^rtB!

1C

li

3c

.

JW

r

8a

8b

n

12

Plate 33

Id

* '.; 7

3d

8c

13

1 e

6a

-

13a

Achatinellidae

Plate 34

la

3a

H

6 b

8

8 a

8b

9

5a

10

, -.

14

14 a

13

14 b

13 a

15

13 b

15 a

Achatinellidae

PLATE 35

10

8

.

v
16

Achatinellidae

Plate 36

1 f

la

Ib

5a

5b

Ic

1

'

3a

5c

Id

5d

le

4a

5 e

6a

7a

6b

7b

8a

6c

7c

9a

lOa

AchatinelHdae

Plate 37

7a

--^

s

4a

5c

-

8

i

2a

4b

8a

10

11

lla

3f

.

%

4c

h.

^

6a

8b

lib

3g

,

6b

8c

3b

3h

5a

12

Achatmellidae

Plate 38

,

X

9

V J
15

,_>

10

16

V

Ib

4a

7b

11

16a

12

17

2a

2b

13

14

18

18 a

19b

19c

Achatinellidae

Plate 39

8

8b

8c

3a

11

m

9a

lla

9b

lib

12d

9c

lie

13c

13d

13 e

13f

10

12

13 a

14

13b

14 a

ACHATINELLA SOWERBYANA. 177

em collector will ever get. The shells are either plain except
for the sutural band (pi. 34, fig. 9, Kaipapau), or have a
band around the base, as in pi. 34, fig. 10 from Kaliuwaa val-
ley near the Castle trail. This shell measure®, length 19.2,
diam. 10, aperture 9 mm. ; 6y3 whorls.

In one lot from the Kaipapau end of the Castle trail, no.
1866 Spalding 'coll., there are 20 of the plain typical form
with a sutural band only, 3 with a basal band. There is also
one chestnut specimen with a light sutural band. It has
also been taken by Mr. Thurston in the banana patch in Kai-
papau along the Castle trail towards Punaluu.

Color-form oviformis ' Nc.' Pfr. PI. 34, fig. 11; pi. 30, fig.
11&. ' Shell sinistral, imperforate, ovate-conic, solid, striatu-
late, white under a glossy olivaceous epidermis, one to three
banded with brown. Spire convexly- conic, the apex rather
acute, white. Suture deeply margined. Whorls 5, scarcely
convex, the last about three-sevenths the total length. Aper-
ture nearly diagonal, ear-shaped; columellar fold moderate,
tooth-like, reddish ; peristome unexpanded, the margins joined
by a thin callus, the outer margin labiate within ; columellar
margin dilated, adnate. Length 17, diam. 8%, aperture
7% mm. long, 4 wide. Oahu, Newcomb ' (Pfr.). Type in
Pfeiffer's collection.

Mr. Sykes hasr referred Pfeiffer's sinistral form of A. mul-
ticolor to -oviformis, no doubt correctly. Pfeiffer's figure is
reproduced in pi. 30, fig. lla. Specimens from Kaliuwaa are
three-banded with brown (" Hay's russet ") on a wax-yellow
ground, pi. 34, fig. 11, coll. by Spalding. It seems to be
merely a color-form or mutation of typical sowerbyana,
hardly worthy of a name.

13a. A. SOWERBYANA THURSTONi P. & C., n. subsp. PI. 34,
figs. 13 to 14&.

The shell is invariably sinistral; ground-color wax-yellow
below the periphery, white above it, with a band of burnt-
sienna bek)w the periphery. Sutural margin defined by a
deep line, usually with an inconspicuous, scarcely noticeable,
dark line next the suture. Apex generally tipped minutely

178 ACHATINELLA SOWERBYANA.

with dull purple. Lip-rib narrow and whitish or very pale.
Columellar fold moderate, pink.

Length 17.2, diam. 10.5, aperture 8.5 mm. ; whorls 5~y2.

Length 16, diam. 9.5, aperture 8 mm. ; whorls 5%.

Length 16, diam. 8.7 mm.

Kahuku, 1,500-1,700 ft. elevation, L. A. Thurston, cotypes
in coll. A. N. S. and Bishop Mus. Also in Thurston collection.
Waimea, overlooking Laie, Irwin Spalding.

Rarely the pigment is deficient, ground white throughout
and the band reduced to a group of pale-brown lines (fig.
136). This decolored form approaches A. s. laiensis. There
may also rarely be traces of faint spiral lines near the lip on
the upper surface. These lines are yellowish with several pink
ones near the peripheral white band. These colors are so
delicate that they can hardly be seen without a lens, and are
mentioned here chiefly because this pattern reminds one of
A. casta. The fact is, A. s. thurstoni stands on the border-
line between Bulimella and the casta group. Looking at
some individuals one is disposed 'to rank it as a variety of
sowerbyana, while others have features which certainly come
as near to casta. Figs. 13 to 136 are cotypes from Kahuku.

Further southeast on the main range, upon -the Waimea-
Laie ridge, Mr. Spalding found a lovely color-form, illus-
trated in pi. 34, figs. 14, 14&, 146. As in the typical color-
pattern, the ground-'color is wax-yellow below the periphery,
white above it (when not covered by another 'Color) ; either
bandless or with four peach-red, coral-red or geranium-pink
bands: sutural, supra- and infra-peripheral and columellar;
or bands i and ii may be concreiscent into a broad zone, as in
fig. 14 ; lip-rib narrow, whitish or with spots at the ends of the
bands ; apex dusky purplish or nearly white. Length 15 to
16 mm.

136. A. SOWERBYANA LAIENSIS Pilsbry & Oooke, n. subsp. PI.
34, figs. 15, 15a.

The shell is sinistral, ovate-conic, moderately solid ; white,
with several brown bands, from burnt-sienna to chestnut in
color; the one below the periphery is widest and most con-

ACHATINELLA SOWERBYANA. 179

stant, one above the periphery is usually present, with some-
times a sutural line and columellar patch also; rarely the
shell is pure white. The apex is whitish or tinged with dull
purple. Surface has a moderate gloss, and) is weakly seriate
under the lens. Suture distinctly margined. The aperture
is not very oblique, white within ; lip a trifle expanded at the
edge, having a rather narrow whitish callus rim within, dull
purplish-'brown towards the edge in well-banded individuals.
Columellar fold white, or often in part brownish.

Length 17, diam. 9.7, aperture 8.3 mm. ; 5~y2 whorls.

Length 16, diam. 9.5, aperture 7.4 mm.

Oahu : Laie, division ridge above the Gastle cut trail, Irwin
S pal ding. Co types in A. N. S. and Bishop Mus. Also in
Sp aiding coll.

This subspecies does not have the greenish-yellow cuticle or
brilliant gloss of A. sowerbyana. The bands are vertically
streaked with chestnut on a lighter, more yellow tint, the con-
trast more obvious in some specimens than in others. Whether
it will eventually prove separable from A. s. thurstoni cannot
now be decided. Its locality is rather remote from the area
of A. cast a, some of the Waimano patterns of which certainly
resemble laiensis.

There are a few white, bandless specimens in Mr. Spald-
ing's lot from the type locality, no. 3556 of his collection, and
one shell with yellow base and wide subperipheral blackish
band (band iii), establishing a connection with var. thurstoni.

13c. A. SOWERBYANA DEXTROVERSA P. & C., n. SUbsp. PI. 35,

figs. 8 to 13.

Shell dextral, white, sometimes uniform, but typically en-
circled by several bands, which are light-brown, vertically
streaked with chestnut; a narrow chestnut columellar area;
suture margined with a dark line which ascends to the apex,
in banded individuals. Penstome narrowly thickened within,
pale with dark spots at the ends of the bands. Columellar
fold moderate, white or nearly so. Length 18.5, diam. 9.5,
aperture 9 mm. ; 5% whorls.

Pupukea, D. Thaanum. Cotypes in A. N. S. P. and Bishop
Museum. Also in Thaanum coll.

180 ACHATINELLA, SECT. ACHATINELLASTRUM.

This is the western terminal member of the sowerbyana
series. In a considerable lot seen it is always dextral. Ex-
cept in direction of coil and some details of banding it has a
close resemblance to laiensis. Some of the white shells have
•the lip white, others having it purple-brown.

Further up, on the Kahuku division ridge, the same sub-
species has been taken by Mr. Spalding, 3558 of his col-
lection, 108129 A. N. S. P. The shells are pure white, or light
brown with dark sutural line and bands ii, iii, and the colu-
mellar region are narrowly dark; band ii being narrow, iii
wide and darker.

This subspecies is quite unlike A. s. roseoplica except that
both are dextral.

13d. A. SOWERBYANA ROSEOPLICA P. & C., n. subsp. PL 34,
fig. 12.

The shell is dextral, oblong-conic, rather solid, white under
a greenish-yellow (sulphine-yellow) cuticle which is slightly
streaked and is deciduous in a band below the suture and on
the spire; apex white or pale^brown. Whorls but slightly
convex. Lip-rib narrow, white or pale pink, the columellar
fold roseate.

Length 18, diam. 9, aperture 8.2 mm.; 6 whorls (A. N. S.).

Length 17.2, diam. 8.7, aperture 7.8 mm. (Bishop Mus.)

Opaeula, above forest- fence line, type loc. ; also on the
northeastern division ridge between Opaeula and Ka/waiha-
lona, Irwin Spalding.

A few of the specimens from the last locality have a pink-
ish-brown line bordering the suture on the last whorl. All of
the specimens from both colonies are dextral. It is widely
separated from all other known 'Colonies of sowerbyana.

Section ACHATINELLASTRUM Pfeiffer.

Achatinellastrum PFR., Malakozoologische Blatter i, 1854,
p. 133.— PEASE, P. Z. S. 1869, p. 646 (restricted to " species
allied to A. productum Reeve"). — SYKES, Fauna Hawaiiensis,
p. 320, 1900.

The shell is imperforate, ovate-conic or oblong-conic,

ACHATINELLA, SECT. ACHATINELLASTRUM. 181

smooth, with unexpanded lip, which is but slightly or not
thickened within ; columellar margin not raised or thickened
on the face.

Type, A. producta Reeve. Distribution, Oahu, chiefly on
the Main Range.

This is the most generally distributed group of Achatinella
In many places it is found on the northern side of the main
range, and in a doubtless more humid time of the Pleistocene
it lived down nearly to sea level. In the Waianae range there
are a number of colonies, but all excessively small, and situ-
ated on the inland slope.

Achatinellastrum is related to Bulimella through such
species as casta and sowerbyana, where the sectional borders
are debatable. Such forms seem to be the least changed
descendants of the ancestral common stock. No point of con-
tact with the section Achatinella (Apex) is traceable among
the recent species.

Achatinellastrum is more prolific in color-mutations than
any other group of the family. The number of patterns runs
into hundreds. About 72 names have been applied to sup-
posed species. This number was reduced to 49 by Mr. Bald-
win in his Catalogue of 1893. Mr. Sykes, 1900, recognized
35 species and 3 varieties. In the following account 17
species and 14 subspecies are admitted, two species and three
subspecies being new.

In the section Achatinellastrum, it is not likely that any
conservative zoologist having adequate collections and data,
would recognize more than seventeen species; but if the evi-
dence is critically examined, it appears that there are pheno-
typically intermediate forms — hybrids or undifferentiated
remnants of the parent stock — between many of the conven-
tional species. It would be quite possible to reduce the * * good
species" to nine or ten.

Thus, in the eastern end of the Main Range, we have a
chain of connected forms in (1) A. ph&ozona — fulgens — ste-
wartii — vulpina. A little apart from them stands (2) A.
buddii. This is succeeded by the form-chain of (3) A. bellula
— casta — juncea. Allied, but not connected are (4) A. juddii

182 ACHATINELLA, SECT. ACHATINELLASTRUM.

and (5) A. papyracea. Westward we find the connected series
(6) A. livida — curia — dimorpha, with a distinct satellite
species, (7) A. ccesia. The Waianae species are as yet but
little known, and apparently are distinct.

Much remains to be done in the definition of the critical points
where one polymorphic population gives place to another,
especially in the western half of the Main Range. It is quite
likely that further collections and study will modify our
present specific boundaries, or perhaps abolish some of them.

I have not constructed a key to the forms of this group
for the reason that most species of Achatinellastrum vary
widely in color. Such small differences in size and shape as
there may be in the average between allied species, are usually
covered by individual variation. I fear that any key I could
make would surely mislead anyone who attempted to name
single specimens by it. The expert will naturally turn to the
group or * * series ' ' to which his unnamed shell belongs. Those
without special knowledge of Achatinella will doubtless most
easily get from the plates a clue to what they seek.

Series of Achatinellastrum.

Series of A. vulpina. Eastern end of the Main range, west
to Manana. Mainly rather large and moderately strong
shells, conspicuously colored, green, yellow or chestnut, gen-
erally streaked or banded, rarely white. Species no. 14 to 18.

Series of A. cast a. Tantalus to Helemano. Smaller shells,
whitish, yellow or chestnut, usually with bands. Species no.
19 to 21.

Series of A. papyracea. Middle of Main range. Rather
capacious, ovate, thin shells, the embryonic whorls not marked
with an ocher band. Species no. 22.

Waianae Range species, intermediate between the papy-
racea and livida series. Species no. 28 to 30.

Series of A. livida. Western half of the Main Range.
Rather small, stout, ovate or short shells, dull green, yellow
or white, often with a few bands, or sometimes streaked ;
embryo often with an ocher band. Species no. 23 to 27.

MAP OP EASTERN OAHU.

183

Q o 2

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«fc\V. *£•'

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^>xVHA\% V.

^ i q v\\\v.,-=:.x

^^ ^

.. ,.-!i» .xv^ ?«|- *>\XS"fS.-

'• '****tS. 3?t

Series of A. vulpina.

Large, highly colored forms, usually chestnut, yellow or
green, and variegated with streaks or bands ; inhabiting ridges

184 ACHATINELLA PELEOZONA.

and ravines from Manana valley to the eastern end of the
Main Range. No other group of Achatinellastrum is found
east of Manoa valley, but westward the casta group appears
in the area inhabited by vulpina forms.

The distribution is diagrammatically indicated in the ac-
companying map, p. 183. The stations of the respective
species are included between the looped lines and the main
axis of the range, but in reality the colonies actually occupy
only a small fraction of the areas indicated.

14. A. PEUEOZONA Gulick. PL 24, figs. 10 to 13 ; pi. 36, figs.
10, 10a ; pi. 43, figs. 1 to Id.

"Shell sinistral, scarcely perforate, oblong-ovate, solid,
shining, striated; white with from one to six black or chest-
nut bands varying in width; apex subacute; spire convexly
conical; suture marginate, moderately impressed; whorls 7,
moderately convex; columellar fold central, white, strong;
aperture a little oblique, lunately rounded ; peristome acute,
well thickened within, with columellar margin dilated, adnate,
or sometimes slightly detached; parietal margin wanting.
Length 22, diam. 12% mm. ; length of body whorl lGi/4 mm. ;
length of aperture 11 mm. ; an average sized specimen.
Length of a large specimen 25.4 mm. Average weight 10.5
grains.' (Gulick.)

Mr. Gulick enumerates the following color-variants. ' ' Var.
a. — With one broad band encircling the base. Var. &. — With
two dark bands, one entering the aperture, the other revolv-
ing above the suture. This and var. a are sometimes found
in Kailua, Oahu. Var. c. — White, with 3 or 4 bands at the
base. Var. d. — Without bands, but more or less streaked
with fawn brown. Var. e. — Dark brown, with two white
bands, one sutural, the other on the periphery of the body-
whorl. Var. f. — Brown, with one or more black bands. Var.
g. — Ash or olive brown, with one or more light bands. Var.
h. — Chestnut or olive brown, with fine, black, spiral lines. '

Oahu: Keawaawa, on kukui and ki (Gulick, Spalding).
Formerly on the northern side of the range in Waimanalo,
Kailua and Olomana (Gulick) ; fossil in a coconut plantation

ACHATINELLA PH^EOZONA. 185

about half a mile from the shore, southeast of Kailua Bay, in
humus of plowed fields (Spalding).

Achatinella phaeozona GULICK, Ann. Lye. N. H. of N. Y.,
vi, p. 214, pi. 7, f. 40, December, 1856.

"May be grouped with A. buddii and A. fulgens Newc., but
differs in its more solid structure, its thicker lip and colu-
mellar fold, and in the more convex outline of its spire. It
also lacks the black tip which characterizes A, buddii. The
typical varieties found in Keawaawa are rare, that sterile re-
gion affording but few trees, which occupy the ravines near
the summit of the ridge.' (Gulick.)

Professor Hyatt looked upon A. phceozona as the common
ancestor of Achatinellastrum, Bulimella and Apex (Science,
viii, p. 395). Later he claimed for A. phceozona a relationship
to Kauaia. Our investigations lead to totally different re-
sults, and we believe the affinities claimed by Hyatt to be alto-
gether erroneous.

Some specimens of phceozona are hardly separable from cer-
tain shells of the plumata pattern of A. fulgens ; but other
patterns of both differ widely. A. phcuozona never has green
or yellow varieties.

Although this species is at present restricted to a very
small area, it formerly inhabited an extensive district on the
northern or Koolaupoko side of the island, which was then
wooded down to the shore. About sixty years ago, when Mr.
Gulick discovered the species, it was still to be found in a
few scattered colonies in Waimanalo and Kailua. On his
labels Mr. Gulick indicated that it was almost extinct in these
valleys.

The Keawaawa lot in Gulick 's collection consists wholly of
dead shells. A series is shown in plate 24, figs. 10-13, pi. 36,
f. 10, 10a. The shell is commonly white with chestnut bands
0230 or 0030, but the bands vary in width, sometimes nearly
covering the last whorl. There is almost always a white band
at the periphery, and the sutural margin and summit are
always white. Often the bands are split, giving rise to nu-
merous band-forms, mentioned by Gulick.

Another pattern is closely streaked with fleshy-brown, cut

186 ACHATINELLA PH^EOZONA.

into bands by white spiral lines and zones, and often with
darker bands also. This is a plumata pattern. Albino shells are
also found. Specimens of Gulick 's Keawaawa lot measure :

Length 20.5, diam. 12 mm., 6% whorls.

Length 23.2, diam. 13 mm.

Length 22.5, diam. 14 mm.

Gulick 's collection was no doubt from rather low, where
the forest was already almost gone in his time. Higher up,
in the bottom of the ravine next to the head of Kuliouou,
Mr. Spalding found living shells on dead kukui trees, in some
abundance, in 1908 to 1910. Some of these are figured, pi.
43, figs. 1 to Id. The white, the two-banded, the split-banded
and the streaked patterns are about equally prevalent. The
white shells are more or less tinted behind the lip, and around
the root of the columellar fold. Banded shells often have the
fold wholly white.

Length 25.2, diam. 13.2 mm., 7*4 whorls.

Length 24, diam. 13.5 mm., 7 whorls.

Length 20.6, diam. 12 mm., 6% whorls.

Length 18.8, diam. 12.4 mm., 6 whorls.

Northward, across the range, A. phceozona is probably now
extinct. About sixty years ago Gulick found it in small num-
bers in Waimanalo (similar to pi. 36, fig. 10a), in Kailua and
on Olomana. The Kailua specimens (no. 589 Boston Soc., pi.
36, figs. 9, 9a), are rather small, length 20 to 21 mm., and
mainly slender ; but others of the same lot are typical in con-
tour, similar to fig. 10a. Of the specimens from the Koolau-
poko side Gulick writes: "Smaller, with outlines of spire
less convex; passing into A. plumata. Average weight 5.3
grains. Habitat : vars. i-k in Kailua ; vars. l-o in Olomana.
Var. i. — white with numerous chestnut bands on the lower
part of the whorls. Var. j. — Light olive brown with dark
bands. Var. k. — Dark brown with narrow white bands.'

Olomana is a shapely and elegant peak terminating the
butress thrown out between Kailua and Waimanalo. Here
Mr. Gulick found a few ph&ozona, for the greater part small
and slender, with bands or lines of carob brown below the
periphery, or sometimes above also (pi. 48, figs. 19, 20, Boston

ACHATINELLA BUDDII. 187

Soc. N. H. coll.). Also white, of the ordinary phaozona
shape and size, in contour like pi. 24, fig. 11. No tree snails
are now to be found on this peak. Mr. Gulick notes the fol-
lowing patterns. "Var. 1. — White with one or two broad
black bands at the base. Var. m. — Elongate, white with two
black bands, one revolving above the suture, the other enter-
ing the aperture, and sometimes a third accompanies the
sutural margin. Var. n. — White with from three to five
crowded bands at the base. Var. o. — Pure white.'

On the base-leveled plain north of Olomana, about half a
mile from the sea southeast of Kailua Bay, Mr. Spalding
found phaozona of normal size and marking in the humus
turned up by the plow in a grove of young coconuts, west of
the stream. While one would not expect land shells to with-
stand disintegration for many years, it must be a century, and
probably much more, since forest suitable for tree snails
existed in this place. It will be inferred from the planting
of coconuts that the plain lies only a few feet above sea level.
The situation is however a dry or semiarid one for the Koolau
side. East of the stream in this plain I have found rather
rich deposits of fossil land shells, including Amastra and
Leptachatina.

15. A. BUDDII Newcomb. PI. 36, figs. 7 to 8a.

''Shell sinistral, conically ovate, solid; whorls 6, convex,
slightly margined above ; suture moderately impressed, banded
with white ; aperture ovate ; lip acute, thickened within ; colu-
mella short, with a terminal plication. Color yellowish (or
cinnamon) slate or fawn; columella and aperture white.
Length 16/20, diam. 9/20 inch" (Newc.).

Oahu: Waialae (Gulick); Palolo (Newcomb, type loc. ;
Gulick) ; Manoa (Emerson) ; Head of Makiki (Spalding,
Thurston, Cooke, Pilsbry and others).

Achatinella ~buddii NEWC., P. Z. S. 1853, p. 155, pi. 24, f. 73.
— PFR., Monogr. iv, 538; vi, 173. — GULICK, Evolution, Racial
and Habitudinal, p. 41, pi. 2, f. 16 (Makiki). — Achatinella
fuscozona Smith, GULICK and SMITH, P. Z. S. 1873, p. 76,
pi. 9, f. 9.

188 ACHATESTELLA BUDDII.

In shape A. buddii does not differ materially from the wid-
est specimens of A. fulgens. The apex is dark. The colu-
mellar fold is often smaller and generally vinaceous.
The chief difference is one of color; in buddii the shell
in its more primitive pattern is closely streaked with liver
brown to purplish vinaceous, flesh color and whitish or creamy,
or with flesh-tint or yellowish brown alone, on a paler ground,
the sutural margin self-colored or narrowly white. The
streaks are sometimes continuous, but usually the pattern is
varied by darker or lighter spiral zones, or interrupted by
white bands or zones, and there are occasional albino in-
dividuals. Through various stages, there is a passage to the
banded pattern, in which there are spiral bands and lines of
chestnut brown or blackish brown on a white or buff ground,
often shading towards the base to cream-buff, sometimes
streaked with brown. Often there is a very faint brownish
line below the suture, and in the rare mutation described as
A. fuscozona, there is a subsutural chestnut band. The em-
bryonic shell is often brown with a wide white or pale zone
below the suture ; and when white it always has a dark tip,
even in albinos. In A. fulgens the embryo is white as a rule,
but sometimes it has a dark tip. Specimens of buddii from
Waialae, Palolo, pi. 36, f. 7 to le, and Makiki, pi. 36, f. 8, 8a,
do not differ materially.

A. buddii was formerly not uncommon in Palolo, where
large numbers were collected by Newcomb, Gulick and doubt-
less many others. The supposed A. fuscozona recorded by
Messrs. Gulick and Smith from Palolo have no direct connec-
tion with the fuscozona of Makiki, but are an independent
though somewhat similar form of buddii. There is a very
pale sutural band of a light ochraceous-buff tint, on a straw
yellow or nearly white ground, and the apex is that of typical
buddii. The specimens are no. 678 of Gulick 's collection,
and no doubt were selected out of his Palolo lot of buddii.

About 1855 Mr. J. S. Emerson collected an ample series
in the bottom of Manoa valley on the Sugar Loaf side, above
where Dr. Cooke's house now stands, in a grove of kukui
trees then being cut by Chinese to obtain pepeitao-laau, an

ACHATINELLA BUDDII. 189

edible fungus. The shells are of the streaked and also the
banded patterns. These localities have long since been de-
forested, and the species is now to be found only high in
Makiki, where the banded pattern, pi. 36, fig. 8a, prevails.
A few specimens which had escaped shell collectors could still
be found hiding in knot holes and crevices in the bark of cer-
tain old kukui trees, when Doctor Cooke and I visited the
place in 1913. It seems to be a shell of the kukui zone, prob-
ably not found at greater elevations.

An unusual pattern of buddii was selected to form the sup-
posed species fuscozona. The description follows, with notes
on the type and other specimens. It seems to be a mutation
which did not become general in occurrence. There are many
instances where a particular pattern has been found on a few
trees only.

Color var. fuscozona Smith. PL 38, fig. 15. "Shell sinis-
tral, ovate-conic, perforate, slightly shining, striated with
growth lines and (under the lens) very minute transverse
lines; whitish, more or less streaked obliquely with light
brown, and transversely indistinctly lined and zoned ; suture
distinctly margined with brown ; whorls G1/^, convex, the last
one ample ; apex blackish ; aperture white, peristome thin,
lightly bordered within; columellar fold strong, reddish
(sometimes whitish). Length 21, diam. 12 mm.

* ' Var. Shell long-conic ; suture girdled by a very wide
brown zone. Length 23%, diam. 11 mm.

' ' Station : on the trunks of trees.

1 i Habitat : Makiki, on Oahu. Two or three specimens have
been found in Palolo. Affinities: It is intermediate between
A. adusta Rv. and A. buddii Nwc.' (Smith).

The type-specimen is figured, pi. 38, fig. 15. It is no. 75 of
Gulick's type series, Boston Society. The original figure
shows the aperture too narrow, and the two lines near the
middle of the last whorl too strong. They are barely visible.
The shell is white, pale buff behind the outer lip and on the
parietal wall of the aperture, where several darker, isabella
colored lines or narrow bands are indistinctly visible. The
first half-whorl is ocher-red, the next whorl violet-plumbeous

190 ACHATINELLA FULGENS.

in the lower half, fading to white above. This dusky tint
fades on the following whorl. The last three whorls have a
chestnut border below the suture about y2 of. a millimeter
wide. The subsutural furrow bisects this border. The aper-
ture and columella are white, exactly as in A. buddii. There
is a very small dark area below the columellar reflection
which makes it appear perforate, but it is not really so.
Length 21, diam. 12.3 mm., with 6*4 whorls.

In his collection Mr. Gulick selected specimens of buddii
having a brown subsutural band and segregated them as fus-
cozona. Altogether he had about a dozen, found among per-
haps a couple of hundred buddii. They vary in pattern
from that of the type of fuscozona to specimens like pi. 36,
figs. 7a, 7c, except that they have the sutural band. A few
are very small, length 18, diam. 9.5 mm., with 6Vs whorls.

In one of Mr. Gulick 's lots from Makiki, no. 804 Boston
Soc., there is one buddii with a wider sutural band and
slightly purplish-brown columellar fold, and four stewartii
of unusual pattern, two of them figured in pi. 38, figs. 16, 16a.
This is what Mr. Smith described as a long variety of fus-
cozona.

Having examined nearly all the fuscozona ever taken by
Gulick, including the type, I am satisfied that Mr. Sykes was
right in placing it as a synonym under A. buddii. It has
nothing to do with stewartii, except that Mr. Gulick mixed
them in one of his lots.

16. A. FULGENS Newcomb. PL 36, figs. 1 to 6e ; pi. 37, figs.
1 to 9 ; pi. 43, figs. 2 to 4c.

" Shell elongately conic, polished, shining; whorls 6, flatly
convex ; suture slightly impressed ; aperture ovate ; columella
short, tuberculated ; lip simple, ribbed within ; color rich
chestnut-brown, with a broad white sutural fascia cutting the
center of the last whorl ; apex and columella white. Length
eighteen, diam. eight-twentieths inch.' (Newc.)

" Var. a, white with broad chestnut bands.' (Newc.). PL
29, fig. 24.

Oahu: Niu (Newcomb) to the Palolo-Manoa ridge; var.
versipellis over the range in Kailua.

ACHATINELLA FULGENS. 191

Achatinella fulgeus NEWC., Proc. Zool. Soc. London, 1853,
p. 131 ; varieties figured on pi. 22, f . 24, 24a. — PFRV Mono-
graphia, iv, p. 537. — Achatinella liliacea PFEIFFER, P. Z. S.,
1859, p. 31 ; Monographia, vi, p. 173. — A. lilaceum Pfr.,
HARTMAN, Proc. A. N. S., Phila., 1888, p. 34. — Achatinella
vulpina Fer., REEVED Conch. Icon,, vi, pi. 4, f. 29a, b, c. —
THWING, Orig. Descriptions, etc., pi. 1, f. 12. Not of Ferus-
sac. — Achatinella crassidentata PFR., P. Z. S. 1855, p. 6, pi.
30, f. 23; Monographia, iv, p. 539. — Achatinella plumata Gu-
lick, Annals of the Lyceum of Natural History of New York,
vi, p. 217, pi. 7, f . 41, December, 1856 ; Evolution, etc., p. 41,
pi. 2, f. 23 (Waialae}. — Achatinella diversa GULICK, t. c. p.,
220, pi. 7, f. 42a, 426; Evolution, etc., p. 41, pi. 2, f. 24 (Wai-
alae).— THWING, Orig. Descript., pi. 1, f. 11. — Achatinella
varia GULICK, t. c. p. 222, pi. 7, f . 43 ; Evolution, Racial and
Habitudinal, p. 41, pi. 2, f. 21 (Palolo). — Achatinella tri-
lineata GULICK, t. c., p. 226, pi. 7, f. 43 ; Evolution, etc., p.
41, pi. 2, f. 20 (Palolo). — Achatinella augusta SMITH, P. Z.
S., 1873, p. 74, pi. 9, f. 7. — Achatinella angusta Sm., PAETEL,
Catalog, p. 105. — HARTMAN, Proc. A. N. S. Phila., 1888, p.
33.

A. fulgens comprises particolored, sinistral shells, inhabit-
ing the ridges and ravines between the areas of A. phceozona
on the east, and A. stewartii on the west. It is ordinarily a
more slender shell than A. phceozona, with the spire nearly
straight-sided, and further differs from that by the frequent
presence of a yellow or green cuticle. A. stewartii is usually
more solid, more obtuse above, and differs in color-patterns.
It never has white bands, which are frequent in fulgens.
A. fulgens is sinistral as a rule, but dextral as a very rare
variation. Probably about half a dozen dextral shells are
known. The Kailua race, versipellis, is frequently dextral.
There are some transitional examples on the boundaries be-
tween phceozona, fulgens, and stewartii, where narrow areas
of overlap exist on the western and northern confines of Kuli-
ouou, and in the northwestern ravines of Palolo.

Many strikingly diverse color-mutations have arisen and
become more or less generally spread throughout the fulgens

192 ACHATINELLA FULGENS.

area. Nearly every colony is hybrid, and in some the mixture
is very complex. If any gametically pure colonies exist, they
must be rare. Segregation of the elementary patterns is
often incomplete, and in assorting any large lot, one en-
counters specimens which seem to be blends, often in large
proportion.

In general, one may say that forms obliquely streaked and
lineate with various shades of vinaceous or cinereous (plu-
mata patterns) predominate in the east, and fade out west-
ward, where they finally appear chiefly in blends with other
patterns. Green and yellow cuticle is mainly developed in
the western district. Some other mutations, such as the
augusta,, fulgens and crassidentata patterns, have had their
rise in the intermediate district, and have not spread over the
whole area of the species.

The numerous names proposed by Gulick and others have
no validity in existing taxonomic usage as standing for
species or subspecies, because the forms occur only as con-
stituents of mixed colonies, and not as pure strains; even
though one or two of the patterns may predominate in one
or another colony. These names, however, provide conveni-
ent terms for the designation of particular patterns. The
chief patterns are as follows. How many of them would
turn out to be elementary patterns, if tested by suitable
breeding experiments, is of course quite uncertain.

1. Plumata pattern. Finely streaked with vinaceous, purple

drab or plumbeous gray, pi. 36, figs. 4a, 5, 6, 6a.

2. Varia pattern. White above, yellowish or olivaceous be-

low the periphery; obliquely streaked with cinnamon or
tawny; usually with a dark line below the suture. PL
36, figs. 1, la.

3. Diversa pattern. Base and a band above the periphery

green, olive or yellow, elsewhere white. PL 37, figs. 4a,
4c.

4. Augusta pattern. Green or olive with a yellowish or

white band below the suture. PL 37, figs. 12, 12a.

5. Trilineata pattern. Yellow below, white above the periph-

ACHATINELLA FULGENS. 193

ery, with two black bands, one above, the other below
the periphery. PL 37, fig. la.

6. Fulgens pattern. Chestnut-brown above and below a

peripheral white or yellow band, which ascends the spire
above the suture. PL 37, fig. 1. This is a rare form,
but it is what Newcomb selected as typical of fulgens.

7. Crassidentata pattern. White, with yellow or olivaceous

bands below the suture, at the periphery and around the
axis, the upper one often wanting. PL 30, fig. 23. A
rare pattern.

8. Liliacea pattern. Albino or albinistic forms occur in some

colonies, and may be derived from various patterns.

The typical fulgens color-form described by Newcomb is
shown in pi. 37, fig. 1, this specimen being from Waialae
nui, Cooke collection. Newcomb 's original figures, repro-
duced in pi. 29, figs. 24, 24a, represent other color-forms,
fig. 24 being Newcomb 's var. a, while fig. 24a is what was
subsequently described as A. augusta.

Niu. — Plate 36, figs. 4 to 6e. The plumata pattern of fine
oblique, vinaceous cream or slaty-purple lines and streaks
over a white, vinaceous, or white-and-yellow ground, is es-
pecially characteristic of Niu. Figs. 5 to 5e show the patterns
in a lot collected by Mr. Gulick. The shells are rather small,
19 to 21 mm. long. There are also shells of the trilineata
pattern and albinistic specimens in which the brown bands
are reduced by absence of the subsutural or the subsutural
and basal. A few have no bands (pi. 36, figs. 5 to 5e).

A series received from Mr. Thaanum, collected recently,
consists mainly of larger shells, pi. 36, figs. 6 to 6e. This con-
tains typical plumata pattern, fig. 6 ; plumata with yellow
cuticle on the base, fig. 6a; plumata with white and dark
bands, fig. 66, c ; trilineata pattern, fig. 6d ; and albino forms,
fig. 6e. There is clear segregation of the elementary patterns
in most specimens, but blends are not uncommon, such as
figs. 4 and 6c, which seem to be blends of plumata and tri-
lineata.

A large series in Mr. Spalding's collection, from the west-

194 ACHATINELLA FULGENS.

ern ravines of Niu, includes typical plumata and typical tn-
lineata patterns, with many intermediate blends. There are
also many white plumata, like pi. 36, fig. 6e. Dr. Newcomb's
fig. 24 was evidently from a Niu shell.

Wailupe.— Plate 36, figs. 3, 3a; pi. 43, figs. 3 to 3c. The
Gulick collection contains plumata patterns, figs. 3, 3a, and
others similar to those he obtained in Niu. Also trilineata
pattern. The augusta pattern, pi. 37, figs. 12, 12a, was also
taken here by Gulick. It is not found in Niu, and has not
been taken in Wailupe in recent years.

Recent collections contain streaked plumata ; trilineata like
pi. 36, fig. 6d ; typical fulgens pattern ; diversa pattern ; var-
ious blends of diversa-plumata and trilineata-plumata pat-
terns; and beautiful, pure white albinos. There are also a
few examples of the crassidentata color-pattern, pi. 48, fig.
21. All of these are found in one colony. A few are illus-
trated, pi. 43, figs. 3 to 3c, northwestern valley of Wailupe,
collected by Irwin Spalding.

Waialae. — PI. 37, figs. 1 to 6&. Waialae fulgens differ
from the Wailupe and Niu series by the prevalence of green
forms and the decadence of the plumata pattern. Very few
plumata similar to those of Niu, pi. 36, figs. 5-5$, are in the
Gulick collection, but in more recent collections, the plumata
pattern occurs in blends with other patterns, such as pi. 37,
figs. 3, 3a, 5a, 6Z>. The varia pattern appears here (Gulick
coll.) but is much more fully represented in Palolo. The
augusta pattern with white spire and in various blends with
plumata and diversa is common, pi. 37, figs. 3, 3a, Waialae
nui, coll. by Thaanum. The trilineata pattern is rare or ab-
sent, but there is an abundant form with two brownish black
bands on a green-streaked ground, pi. 37, figs. 3c, d, e, Wai-
alae nui. The diversa pattern, pi. 37, figs. 4a, &, c, coll. by
Baldwin, and fig. 5c, coll. by Gulick, is abundant. It often
blends with plumata., as in fig. 3g • and there are also beauti-
ful dark green examples with white bands on the spire only,
particularly in Waialae iki. PI. 36, fig. 2 is a dextral speci-
men from Waialae, coll. by Gulick. The fulgens pattern, pi.
37, fig. 1, Waialae nui, coll. by Dr. Cooke, is rare. There are

ACHATINELLA FULGENS. 195

two broad chestnut zones, leaving a yellow band at periphery
and a yellow line below the suture. A beautiful and unique
specimen in the Spalding collection, pi. 43, fig. 2, from Waia-
lae iki, has a green-streaked base and a broad chestnut black
zone above. Figs. 5 to 6& are from specimens taken by Gu-
lick; the other figures of Waialae shells, 1 to 4c, are from
more recently collected examples. On the Waialae-Palolo
division ridge Mr. Thurston collected a fine series of augusta,
rather small shells. It may be a pure colony.

Palolo.— PI. 36, figs. I to If • pi. 37, figs. 7 to 9, coll. by
Gulick. The plumata pattern is rare or wanting in Palolo,
where the principal patterns are varia, diversa, trilineata and
augusta. The varia pattern, pi. 36, figs. 1, 2, is rather char-
acteristic of Palolo. The shell is streaked obliquely with
russet or ecru olive, with an olive-brown band above, the base
ecru olive or brownish. In the northern ravines of Palolo
this pattern is associated with very fine citron-green and dark
green diversa with a blackish line below the suture ; diversa
with white bands, with or without a blackish line above the
periphery, and albinos. Some of these are shown in pi. 43,
figs. 4 to 4c, coll. by Spalding. There are also augusta with
white or streaked spire, mostly with a dark subsutural line.

The fulgens pattern is rare (pi. 37, fig. 8c). A peculiar
modification of the trilineata pattern has a very narrow chest-
nut or blackish line above the periphery, pi. 37, fig. 9, coll.
by Gulick. This seems to be a specially Palolo pattern.

The frequent presence of a dark line below the suture in-
dicates affinity to A. stewartii. Rare individuals from the
northwestern ravine of Palolo are indistinguishable from A.
stewartii.

Dr. Newcomb defined a " Var. &, chestnut colored above,
yellowish below, with two black and one white band, with
columella dark brown, of large size, measuring 22 x 10 twen-
tieths of an inch. This last variety may upon further exami-
nation prove to be a distinct species. The locality of this
last is Makika valley, mauka roa, or far back in the mountain
range.' I am pretty confident that Newcomb was mistaken
about the locality of this variety. Nothing like it is found in

196 ACHATINELLA FULGENS.

Makiki, either mauka or makai, but some Palolo specimens
have the coloration described.

16a. A. FULGENS VERSIPELLIS Gulick. PL 43, figs. 5 to Id.

" Shell dextral or sinistral, imperf orate, acuminately ob-
long, solid, shining, striated, of lively ash-color, more or less
streaked and waved with brown, with several interrupted
brown bands on the upper parts of the whorls; apex some-
what obtuse ; spire convexly conical ; suture margined, mod-
erately impressed; whorls 6, convex; columellar fold central,
white, strong; aperture truncately auriform, white within,
a little oblique, in sinistral specimens very oblique ; peri-
stome thickened within; with external margin unreflected,
compressed, edged with brown; columellar margin dilated,
adnate; parietal margin wanting. Length 20%, breadth 10,
length of body- whorl 15 mm. Average weight 8.6 grains.'
(Gulick.)

Var. b, rich brown, with light streaks and waves. Var.
c, yellow at the base, with one or more brown bands above.
Var. d, yellow at the base and white above, without bands.
Var. e, nearly pure white. Var. f, ash or yellow gray, with-
out bands. A rare and beautiful species, found in the most
rugged but verdant region of western Kailua. About a third
of the specimens are sinistral.' (Gulick.)

Oahu: Pohakunui, Kailua (J. T. Gulick). Kailua under
Mt. Olympus, and the Kailua- Waimanalo division ridge
(Irwin Spalding).

Achatinella versipellis GULICK, Ann. Lye. N. H. of N. Y.,
vi, p. 224, pi. 7, f. 44a, b. December, 1856. — Achatinellastrum
versipilis Gul., PEASE, P. Z. S., 1869, p. 646. — Achatinella
fuscolineata SMITH, P. Z. S., 1873, p. 75, pi. 9, f. 2, and var.
6, f. 2a.

Very closely related to the plumata and various other pat-
terns of A. fulgens, from which versipellis is separated mainly
by its habitat, on the north side of the main range, and by
the frequency of dextral shells. There is, however, some dif-
ference in the patterns; versipellis often having the ground
color of the last whorl all yellow.

ACHATINELLA FULGENS. 197

Mr. Gulick's two cotypes are figured, pi. 43, figs. 5, 5a.
Both are streaked individuals, one streaked with Indian red
on a white ground, the other with chestnut on a yellow
ground. The apical whorls are very pale yellow in both ; colu-
mellar fold entirely white. Other specimens from the origi-
nal lot, pi. 43, figs. 5&, 5c, have livid brown streaks blending
with the light yellowish olive ground, or the dusky streaks
may be very faint on an ecru-olive ground. Specimens with-
out streaks may be deep colonial bun2 with or without inter-
rupted cinnamon bands. Another pattern is pale-pinkish
buff with two yellow bands. One shell is white, shading to
pale green-yellow at the base, with yellowish lines at suture
and above periphery. There is also a diversa pattern, with
light yellow base and band. I have not been able to locate
the type locality, Pohakunui, but it is probably near Mt.
Olympus.

Specimens from Kailua, under Mt. Olympus, collected by
Mr. Spalding, pi. 43, figs. 7&, 7c, Id, and others from Mauna-
wili (in the same region, if not the same colony), collected
by Mr. Wilder, pi. 43, figs. 7, la, show further modifications
of the patterns found by Mr. Gulick. Among them, Mr.
Wilder found a green form, diversa pattern.

On the division ridge between Kailua and Waimanalo Mr.
Spalding collected various streaked and yellow-banded pat-
terns, all sinistral (pi. 43, figs. 6 to 6c).

A. fuscolineata E. A. Smith. (PI. 43, fig. 8, reproduction
of original figure).

" Shell sinistral, ovate-conic, imperforate, glossy, striated
obliquely with growth lines and very delicately transversely
striate ; greenish-yellow streaked with green, and encircled
above the periphery with (3 to 5) brown lines; suture dis-
tinctly brown-margined. Whorls 6%, convex, the first three
white. Aperture white; peristome white (sometimes brown),
the margin acute, bordered within ; columellar margin strong,
roseate (sometimes white). Length 19, diam. 10 mm.

11 Var. a. Shell more of a green color.

" Var. &. Shell smaller, sub testaceous, streaked with pale
chestnut and transversely banded above with deep brown.'
(Smith.)

198 ACHATINELLA FULGENS.

Oahu: the typical form is found in Kailua, but varieties
which may conveniently be classed with it are found in nearly
all the valleys between Palolo and Halawa (Gulick).

"It is most nearly allied to A. versipellis Gulick. This
species is very rarely dextral. The specimen figured is from
Kailua ' (Gulick).

Mr. Sykes follows Newcornb in referring this to A. vul-
pina as a synonym. It seems to me to be a banded form of
versipellis. Like that it is rarely dextral. Shells from other
districts than Kailua, referred by Mr. Gulick to fuscolineata,
are probably forms of vulpina, superficially like the Kailua
type in markings. Some of them which I have seen from the
western valleys are forms of analoga; and his " fuscolineata'
from Palolo are certainly fulgens.

166. A. FULGENS AMPLA Newcomb. PI. 29, fig. 19 ; pi. 54,
figs. 7 to 7c.

* * Shell dextral, conically ovate, polished ; whorls 5, rounded ;
suture simple, banded with a black stripe ; aperture large and
white ; outer lip simple, acute, thickened within ; columella
short, white or roseate, terminating in a twisted plait; apex
obtuse, roseate ; epidermis light green or olive above, of a
deeper color on the last whorl. Length 14-twentieths, width
10-twentieths inch' (Newcomb}.

Oahu: Koolau (Newcomb).

Achatinella ampla NEWC., P. Z. S. 1853, p. 137, pi. 22, f.
19, 1854. — PFR., Monogr. iv, 533. — SYKES, Fauna Hawaiiensis,
p. 305, 1900. — THWING, Reprint Original Descr., p. 59.

There is an error, probably typographical, in Dr. New-
comb's measurement; 14-twentieths should be 17-twentieths
according to Newcomb 's original figure, which is reproduced
photographically on pi. 29, fig. 19. PL 54, figs. 7-7c rep-
resent other specimens in Newcomb 's collection, no. 29904
Cornell University. The series includes both dextral and
sinistral shells. The very ample last whorl is the chief char-
acteristic distinguishing ampla from the other Koolau races,
phceozona, versipellis and fuscolineata. Some specimens ap-
proach very near to such versipellis as pi. 43, figs. 7b-7d,

ACHATINELLA PULGENS. 199

from Kailua below Mt. Olympus. It must be admitted, how-
ever, that there is also a close relationship to A. stewartii.
Probably ampla came from somewhere in Kailua, NewcomVs
locality being in a rough way correct.

The last whorl is straw yellow, usually streaked and more
or less suffused with green; sometimes with some obscurely
traced green bands; the early whorls whitish or faintly flesh
tinted. The suture is narrowly margined with chestnut.
There is often a blackish chestnut crescent at the columella.
In Newcomb's type and one other specimen the whole base
is streaked with chestnut.

Length 19.5, diam. 12.6, aperture 9.5 mm. ; 6*4 whorls.

Length 20.1, diam. 12.5, aperture 10.5 mm. ; 6 whorls.

The original descriptions of forms based upon various muta-
tions of A. fulgens are reprinted below. We have examined
the types of all but crassidentata, augusta and liliacea, and
have seen part of the original lot of augusta.

A. crassidentata Pfeiffer. PI. 30, fig. 23, reproduction of
original figure. " Shell imperf orate, sinistral, rather solid,
striatulate, little shining; white, ornamented with a few buff
bands, sometimes having the base greenish-buff. Spire conic,
the apex rather obtuse, suture margined ; whorls 5%, a little
convex, the last about three-sevenths the length, rounded at
the base. Aperture diagonal, sinuately semioval; columellar
fold superior, very thick, tooth-shaped ; peristome simple, the
outer margin acute, lightly arcuate ; columellar margin short,
narrow. Length 20, diam. 11 mm. ; aperture 10 mm. long, 5
wide. Inhabits the Sandwich Islands, Frick, in Mus. Cum-
ing" (P/r.).

This pattern has been found by Mr. Spalding in Wailupe.
The bands are cuticular, not homologous with those of tri-
lineata, which are differently located, and dye the prismatic
layer of the shell.

"A. liliacea Pfr. Shell imperf orate, sinistral, ovate-conic,
rather solid, lightly striate, glossy, white ; spire a little con-
vexly conic, apex subacute ; suture narrowly margined ; whorls
6, scarcely convex, the last a little convex, sometimes sub-
angular at the periphery, the base sack-like ; columellar fold

200 ACHATINELLA FULGENS.

pale rose, high, twisted ; aperture oblique, reversed auriform ;
peristome unexpanded, acute, somewhat labiate within ; colu-
mellar margin slightly dilated, adnate. Length 24, diam. 12
mm. Inhabits the Sandwich Islands, Dr. Frick, in Cuming
coll' (Pfr.).

This is an albino form.

Achatinella plumata Gulick. ' ' Shell sinistral, imperforate,
ovate-conic, solid, shining, striated, cinereous, with oblique
brown streaks; apex subacute; spire conic, with outlines
slightly convex; suture marginate, moderately impressed,
white; whorls 6!/2> somewhat convex; columellar fold central,
white, strong; aperture somewhat oblique, truncately auri-
form; peristome subacute, well thickened within, with colu-
mellar margin dilated, adnate; parietal margin wanting.
Length 23 mm., breadth 12.2, length of body-whorl 16, length
of aperture 11 mm. Average weight 8.5 grains.

"Station: on the leaves and branches of trees. Habitat:
Niu, Oahu, J. T. G.P (Gulick).

" Remarks : a neat species, differing from A. phozozona, in
its smaller size, more conic spire, and in the streaked arrange-
ment of its colors. The typical specimens are much thicker
and heavier than A. buddii Newc., and it never has the black
tip of that species. We have seen but one dextral specimen,
which has been mentioned under var. s.' (Gulick).

Mr. Gulick characterizes 41 color-varieties, grouped under
five sections designated by Greek letters.

These sections are as follows: (1) typical, vars. & to k.

(2) "Smaller and more elongately ovate. Length 19y5,
breadth 10% mm. Habitat Wailupe, Oahu.' Vars. I to g.

(3) "Of full size, but thinner, than the typical varieties,
with the first three whorls usually white; passes into A.
fulgens Newc., which is found in the same locality. Habitat,
Waialae, Oahu; vars. r, s and v are sometimes found in
Palolo.' Vars. r to x.

(4) "Rather thin, with lip acute and scarcely thickened
within ; pass into A. buddi Newc., found in the same locality.
Average weight 5.3 grains. Habitat, Palolo, Oahu.' Vars.
y, z, aa to ii.

(5) "With spire more convex and colors less streaked; pass
into A. phceozona vars. i to o. Habitat Kailua, Oahu.'
Vars. jj to pp.

It has not been thought desirable to occupy space here

ACHATINELLA FULGENS. 201

with the descriptions of Mr. Gulick 's lettered varieties. Those
who have not access to the original publication can find them
in Mr. Thwing's Reprint, pp. 40, 41. Data so presented serve
to show the wide range of variation, but are not otherwise
available as variation is now studied.

The few Kailua specimens we have seen seem to be at
least as near phczozona as plumata, although Gulick classed
them with the latter. They have the convex-sided spire of
phceozona.

"Achatinella diver sa Gulick. Shell sinistral, very rarely
dextral, imperforate, ovately or elongately conic, solid, shin-
ing, striated, white or variously painted with yellow or green ;
apex subacute ; spire conic ; suture marginate, moderately im-
pressed ; whorls 6%, somewhat convex ; columellar fold cen-
tral, strong, rose or white ; aperture oblique, truncately auri-
form ; peristome acute, thickened within, with columellar mar-
gin dilated, adnate ; parietal margin wanting. Length 21,
breadth 11%, length of body whorl 15%, length of aperture
10% mm. An average specimen. Greatest length 25 mm.
Average weight 8 grains' (Gulick).

1 1 Station, on trees. Habitat, Palolo, Waialae, Wailupe and
Niu, Oahu, J. T. G.!'

"Var. a. — Pure white, except the columella, which is fre-
quently lilac or rose. Var. b — white with one black band.
Var. c — white with two black bands, one entering the aper-
ture ; approaches A. fulgens Newc. Var. d — white with yel-
low or green base. Var. e — yellow with white bands. Var. f
— yellow, fading towards the apex. Var. h — green, passing
into gray or brown towards the apex. Var. i — green with
white bands.

"Remarks: Differs from A. plumata nob. in the character
and arrangement of its colors, and from A. fulgens Newc. in
the absence of the white sutural band and the two broad black
central bands which characterize that species.

"The specimens found in Niu do not present the green
varieties, but incline more to white, and are also thicker and
more ovate in form than those found in Palolo. The average
weight of full grown specimens from Niu is about 9 grains,
that of the Palolo specimens is 7.2 grains.

"I have six or eight dextral specimens belonging to varie-
ties d, e, g and ir (Gulick).

Achatinella varia Gulick. "Shell sinistral, imperforate,
acuminately oblong, solid, shining, finely striated, white,

i t
( t

202 ACHATINELLA FULGENS.

streaked with brown, with a black line beneath the suture,
and green or brown at the base ; apex somewhat acute ; spire
elongately conic; suture margined, lightly impressed; whorls
6, flatly convex ; columellar fold central, brown or rose, strong ;
aperture oblique, truncately auriform, white within; peris
tome thickened within ; external margin unreflected, arcuate ;
columellar margin dilated, adnate, usually margined with
black; parietal margin wanting. Length 21%, breadth 10%,
length of body whorl 14 mm. Length of a large specimen
27, breadth 13 mm. Average weight 6.7 grains.

'Station: On the Kukui (Aleurites triloba}, Ohia (Eugenia
malacctzcensis) , and other trees. J. T. G. !

Habitat: Palolo, Waialae, and Wailupe, Oahu. J. T. G. !
Var. b, White above, green or yellow at the base. Var. c,
Green at the base, white above, with one or more green bands.
Var. d, Green, brown, or yellow at the base, upper whorls
radiated with white and brown, and banded with green or
yellow. Var. e, Radiated with white, and reddish brown.
Var. /, White except the suture and columella. Var. g, Green
or yellow; passing into A. Stewartii Green. Var. h, Green
with one narrow, white, spiral band, passing just above the
suture. Var. i, Yellow with white sutures, and a dark brown
band revolving beneath. Var. j, Chestnut brown at the base,
becoming paler towards the apex, with several obscure, spiral,
brown lines.

"Remarks: The metropolis of the species is Palolo Valley,
where it is very abundant. In Waialae and Wailupe, which
lie to the east, it gradually becomes more rare, and disappears
in Niu, which has furnished me but one specimen of var. f.
In Manoa, on the west, it soon disappears, being found only
on the mountain ridge that separates it from Palolo. Dextral
specimens are very rare. I have a few from Waialae.

"This shell has been described and figured by Reeve as
A. vulpina Fer., and others have followed him ; but a com-
parison of his figures with Ferussac's leads me to doubt his
correctness, and after an acquaintance with the species in
their native valleys, I do not hesitate to separate them as
distinct. The shell here figured corresponds more nearly to
what I have described as variety c (Gulick).

Achatinella trilineata Gulick. "Shell sinistral, imperf or-
ate, ovate-conic, solid, shining, finely striated, white above,
yellow or green at the base, with three black bands, one su-
tural, one entering the aperture, and the other between the
two, revolving just above the suture ; apex somewhat obtuse ;
spire conical, slightly convex; suture with narrow margin,

ACHATINELLA FULGENS. 203

moderately impressed; whorls 6% rather convex; columellar
fold central, white or rose, strongly developed aperture trun-
cately auriform, white within; peristome thickened within;
with external margin unreflected, arcuate, acute ; columellar
margin dilated, adnate ; parietal margin wanting. Length
21%, breadth 12. Length of body whorl 15 mm. Average
weight 8.6 grains.

' ' Station : On the kukui and other trees. Habitat : Palolo,
Waialae, Wailupe and Niu, Oahu. J. T. G.

"Var. 6, With oblique reddish-brown streaks above, the
base green or yellow. Var. c, Yellowish or green throughout,
excepting the bands. Var. d, White at the base, and also
above. Var. e, Covered with oblique reddish-brown streaks.
Var. /, Apex tipped with black, passing into A. ~buddii Newc.
Var. g, Green or yellow, except the black bands, and a narrow
line of white. Var. h, White, with several fine spiral black
lines accompanying the broader bands. Var. i, With black
bands very broad, occupying half the surface or more. Var.
j, With two black bands, one sutural, the other passing above
the suture. Var. k, With two black bands, one sutural, the
other entering the aperture ; rare. Var. I, With several nar-
row bands on the upper part of the whorls.

' ' Remarks : Dextral specimens of this species are very rare ;
I have obtained but two. Varieties j and k have been found
only in Palolo. As in the preceding species, the specimens
found in Niu are of the lighter colored varieties, and more
solid than those of the other valleys.' (Gulick).

1 ' Achatinella augusta Smith. Shell sinistral, ovate-conic,
glossy, striated with very fine growth and transverse lines;
green, streaked with darker green and encircled by dark green
and rufous lines, ornamented with a yellow zone below the
white suture ; suture distinctly margined with white ; whorls
6i/2, the first four a little convex, the rest convex; aperture
white, peristome thin, tinted with pale brown within; colu-
mellar fold strong, brownish rose color. Length 24, diam.
mm.

"Var. — Shell all yellow, encircled below the suture with a
white zone.

' ' Station : on trees. Habitat : the metropolis of this species
is Waialae, near the east end of Oahu. It is also found in
Wailupe and Palolo. Affinities: This species is on the one
hand closely related to A. plumata Gk., from which it is dis-
tinguished by its green and yellow epidermis, which is en-
tirely wanting in that species, and to A. fulgens Newc., from
which it differs in being without the broad black bands, which

204 ACHATINELLA SOLITARIA AND STEW ARTIE.

belong not only to the epidermis but to the solid part of the
shell of A. fulgens. It has been described by Newcomb and
Pfeiffer as a variety of the latter species. Remarks: this
species is always sinistral. The specimen figured is from
Waialae.' (Smith and Gulick).

The specimen figured as type has the appearance of a
plumata-augusta blend, the plumata pattern appearing on the
spire as in pi. 37, fig. 3. A purer expression of the color-
form is pi. 37, fig. 12; also pi. 29, fig. 24a, which Newcomb
figured as a variety of fulgens.

16l/2. A. SOLITARIA Newcomb. PL 43, fig. 9.

"Shell ovately conical, dextral; whorls 6, flatly convex;
suture slightly impressed; aperture ovate; lip acute, thick-
ened within ; columella white, short, broad and abruptly
twisted ; color light chestnut, with darker longitudinal stripes ;
green at the base, white sutural band for the last 2% whorls,
brown band on the suture above. Length fourteen-twentieths,
width eight and one-half twentieths inch.

"But a solitary specimen of this species has been obtained.
Its characters are, however, sufficiently striking to warrant
in giving it a place as a distinct species' (Newcomb).

Oahu: Palolo (Newcomb).

Achatinella solitaria NEWC., P. Z. S. 1853, p. 150, pi. 24,
f . 60 ; Annals of the Lye. N. H. of N. Y., vi, p. 331.

A problematic form, possibly normal, but more likely to be
a smoked "dunkeri" with the suture scraped white. If New-
comb's locality is correct it should be a form of fulgens; but
a dextral Achatinellastrum in Palolo would be rather ano-
malous though not unique. While it is not likely that A.
solitaria is a valid species or subspecies, it is given place here
because we cannot form a well-founded opinion without seeing
the type-specimen. Very few examples are known, two in
the Cumingian collection being all that are on record. There
are none in Newcomb 's collection at Cornell University. The
original figure is reproduced photographically on my plate.

17. A. STEWARTII (Green). PL 38, figs. 1 to 6a, 14, 16 to 21.
The shell is dextral or sinistral, oblong-turrite, solid, glossy,

ACHATINELLA STEWARTII. 205

lightly marked with lines of growth and very faint spiral
striae ; variously colored, but the typical pattern is citron yel-
low fading to white at the summit, with a Hack or deep
~brown band bordering the suture below on the last 3 to S1/^
whorls, and a black crescent bounding the coluinella ; aperture
white, the columellar some shade of violet. Whorls about
6%, convex, the last rather short. Outer lip simple or thick-
ened within ; columellar fold strong.

Length 22, diam. 11.3, aperture 10 mm., Manoa-Palolo ridge.

Length 24.2, diam. 12.8, aperture 11 mm., Manoa-Palolo
ridge.

Length 23, diam. 11.3, aperture 9.5 mm., Manoa-Palolo
ridge.

Length 22, diam. 13, aperture 11 mm., Manoa-Palolo ridge.

Oahu : Northwestern Palolo to Makiki and eastern Pauoa.

Achatina stewartii GREEN, Contributions of the Maclurian
Lyceum to the Arts and Sciences, i, no. 2, p. 47, pi. 4, f. 1-4,
July, 1827. — Achatinella stewartii Green, REEVE, Conch. Icon,
vi, pi. 4, f. 26. — GULICK, Evolution, Racial and Habitudinal,
p. 41, pi. 2, f. 18 (Manoa). — Achatinella fuscozona GULICK,
t. c., p. 41, pi. 2, f. 19 (Manoa). — Achatinella pulcherrima
REEVE, C. Icon. pi. 3, f. 23a, b. — Achatinella venulata var. a.,
NEWCOMB, P. Z. S. 1853, p. 146, pi. 23, f. 48.— Achatinella
byronii Gray, GULICK, Ann. Lye. Nat. Hist, of N. Y., vi, 1858,
p. 244. — Achatinella johnsoni NEWCOMB, P. Z. S. 1853, p. 147,
pi. 23, fig. 50. — GULICK, Evolution, etc., p. 41, pi. 2, f. 17
(Manoa). — Achatinella aplustre NEWCOMB, P. Z. S. 1853, p.
147, pi. 23, f. 51.

Professor Green's description of A. stewartii included also
the forms described later as producta, castanea, and johnsoni.
His shells were probably from Makiki and the slope of Mt.
Tantalus, and perhaps also from eastern Manoa, as the clear-
yellow ground of part of his figures is much like Manoa-
Palolo ridge shells. Later authors have restricted stewartii
to the yellow or green form with black sutural band such as
pi. 38, fig. 1, agreeing exactly with Green's fig. 2, which may
be selected as his type. PL 38, figs. 2 to 3 are sinistral forms
of similar pattern.

206 ACHATINELLA STEWARTII.

A. stewartii differs from A. s. producta by being narrower,
the last whorl shorter and a little compressed, the spire more
turrited, and the suture always bordered with black. It dif-
fers from fulgens by lacking white bands, by the dark suture,
and various peculiarities of pattern more readily appreciated
than described. A. vulpina and A. stewartii are very inti-
mately related, but in general, A. stewartii is either sinistral
or dextral, larger, more turrited, and when melanistic the
dark hue is in bands. A. vulpina is smaller, more oblong-
ovate, the melanism is sometimes more generally diffused;
typically it has no dark sutural band, and is sinistral (with
rare exceptions).

So far as I know, all stewartii colonies are hybrids of sev-
eral elementary patterns, as follows:

1. From deep colonial buff to malachite green, with dark su-

tural band and columellar patch. PI. 38, figs. 1, 2-3.
Stewartii pattern.

2. Olive-ocher or greenish, closely streaked with cinnamon,

and with sutural and columellar black bands. PI. 38,
fig. 4. Similar to the dunkeri form of producta.

3. Ground-color like either of the above, but having four black-

ish bands. PI. 38, figs. 4a to 6a. Johnsoni pattern.

Palolo, in the head of a makai branch, in the western ravine
on the Manoa ridge, pi. 38, figs. 1 to 2& coll. by Pilsbry; fig.
3, coll. by Wilder, is a prolific locality for all of the above-
described color-forms. Most of the specimens are sinistral.

Towards the main range or "mauka" from the well-known
stewartii ravine on the Palolo-Manoa ridge, Mr. Spalding
found a colony which unites stewartii and fulgens. It con-
tains the following patterns : stewartii forms 1 and 3, tri-
lineata, diversa and varia with dark sutural line. All are
sinistral. This has the appearance of a fulgens (varia)
colony hybridized with stewartii. Mr. Thurston has collected
stewartii-diversa-plumata patterns in the same region.

In north-eastern ravines of Manoa the patterns are simi-
lar to those of Palolo ; typical stewartii, streaked, and banded,
but most of the shells are dextral (pi. 38, figs. 4, 4a, Cooke

ACHATINELLA STEWARTII. 207

coll.). Gulick specimens from Manoa, exact locality not spe-
cified, show the same patterns (pi. 38, figs. 5, 6, 6a). There
are also a few of the stewartii pattern suffused and streaked
with pink. Mr. Spalding has also collected stewartii, forms
1, 2 and 3, on the Manoa side of the Manoa-Palolo ridge.

Makiki valley has various forms not found on the Manoa-
Palolo ridge. A series is shown in pi. 38, figs. 14, 19 to 19c,
coll. by Gulick; fig. 17, Cooke coll., and figs. 18, 18o, from
Thaanum. The usual pattern has an ochraceous-orange tone,
shading towards burnt sienna on the latter part of the last
whorl (fig. 18), or there may be two dark lines or bands (fig.
19a), or the bands may unite to form a broad black-brown
zone (fig. 19c), or the whole lower part may be blackish (fig.
18a) or chestnut (fig. 17) ; all having the blackish border be-
low the suture. Reeve's A. adusta has the same pattern.
Other lots in the Gulick collection have four blackish bands on
a citron or colonial buff ground, the shell dextral (fig. 14).
There are also Makiki specimens like pi. 38, figs. 16, 4a and 5.

Rarely A. stewartii occurs with white ground and the usual
sutural and columellar bands and purplish columella. One
lot from Makiki in the Gulick collection contains a specimen
with the ground pure white ; one white with the base pale
yellow, the whole faintly streaked with pale pinkish cinnamon
(pi. 38, fig. 16), another similar but having an incomplete
zone of chestnut streaks below the middle, and some paler
lines, (fig. 16a). These shells were thought by Gulick to be
A. fuscozona, but I feel sure that they are stewartii.

Near the top of Mt. Tantalus there is — or was — a hand-
some form with the last whorl chestnut-black, lighter or yellow
below the black sutural border (pi. 38, fig. 20, Cooke coll;
fig. 21, Gulick coll.). It approaches A. vulpina, but seems to
be a melanistic form of stewartii.

Yla. A. STEWARTII PRODUCTA Reeve. PL 38, figs. 7 to 13 ; pi.
43, figs. 10 to 106.

' * Shell somewhat elongately conical, dextral ; whorls con-
vex, slightly margined at the sutures ; columella conspicuously
callous-toothed; apex rather small. Olive-green, apex, colu-

208 ACHATINELLA STEWARTII.

mella and interior of the aperture white. Sandwich Islands"
(Reeve).

Length 27, diam. 14 mm. (from figure).

Mt. Tantalus, from the rim of the "bowl" or crater to the
head of Makiki valley ; also on the Pauoa slope down to about
1100 ft. ; on the bark of kukui, ti, guava and other trees and
bushes.

Achatinella producta REEVE, Conchologia Iconica vi, pi. 2,
f. 13, April, 1850. — GULICK, Evolution, Racial and Habitu-
dinal, p. 41, pi. 2, f. 15 (Makiki). — THWING, Orig. Descript.
pi. 1, f. 13. — Achatinella bilineata REEVE, C. Icon, vi, pi. 3,
f. 22, April, 1850. — Achatinella venulata NEWCOMB, P. Z. S.
1853, p. 146, pi. 23, f. 48a. — Achatinella hybrida NEWCOMB,
P. Z. S. 1853, p. 147, pi 23, f. 52.— Achatinella dunkeri Cum-
ing MSS., PFEIFPER, P. Z. S. 1855, p. 208; Monographia iv,
531. — GULICK, Evolution, etc., p. 41, pi. 2, f. 14 (Makiki). —
Achatinella vulpina Fer., GULICK, Ann. Lye. vi, p. 1858, p.
244 ; not of Ferussac.

A. s. producta is a larger, more capacious form tkan A.
stewartii, the last whorl more rotund, and the suture usually
without a dark border. The typical form is dextral, of a
clear primrose-yellow tint (varying however in different in-
dividuals from colonial buff to green), with the summit, aper-
ture and columellar fold white. The lip is a little thickened
within. Sometimes the periphery is indistinctly angular.
Size generally large.

Length 27, diam. 14.2 mm.; 6% whorls (fig. 7).

Length 25, diam. 13 mm.; 7 whorls (fig. 9).

Length 21.5, diam. 12.3 mm. ; 6% whorls.

In the head of Makiki valley the shells are usually dextral
(fig. 9), rarely sinistral; on the summit of Tantalus they are
all sinistral (figs. 7 to 8a southeastern rim of 'bowl"). On
the Pauoa slope of Tantalus all are dextral. The size is
sometimes not greater than that of A. stewartii. The chief
color-patterns are :

1. Producta pattern, pi. 38, figs. 7, 9, described above.

2. Dunkeri pattern, pi. 38, figs, la to 7c. Streaked with

ACHATINELLA STEWAETII. 209

cinnamon over an ochraceous buff or naples yellow
ground, sutural border paler, not streaked; often with
black band ii, or ii and iii ; columella white.

3. Bilineata pattern, pi. 38, fig. 10. Having the upper dark

band split, ground-color as in producta; or there may
be two dark bands on the product a ground (fig. 8).

4. Melanistic forms, pi. 43, figs. 10, 10a, 10Z>, in which the

dark bands occupy most of the surface.

So far as I know, none of these patterns occur as pure
colonies. The common association is producta and dunkeri.
Figs. 7 to Ic were found by the writer on the base of a guava
bush, within a few inches of one another, together with several
young ones. Figs. 8, Sa were lodged side by side on an ad-
jacent bush, on the east rim of Tantalus bowl. As the guava
bushes here are isolated, singly or by small groups, in a knee-
high growth of Hilo grass, there can be no doubt that the
shells on each bush or group of bushes are an intergenerant
group.

While most lots can be assorted by patterns, blends are
rather common. The dunkeri streaks may be excessively faint,
or they may be well developed in the early neanic stage, but
disappear in the later whorls, as in pi. 38, fig. 8a.

The bilineata and melanistic patterns are not properly ele-
mentary patterns. Bilineata is of sporadic occurrence. The
split band is usually the upper, sometimes the lower. It oc-
curs also on shells with green ground (fig. 11) and on those
with dunkeri pattern, but is rather rare.

The melanistic forms, such as pi. 38, fig. 13, Cooke coll., and
pi. 43, figs. 10, 10a, 10&, Thwing coll., Bishop Mus., are ex-
tremely rare, perhaps now extinct, and probably all from the
Pauoa side of Tantalus.

PI. 38, fig. 9 was found on the base of a kukui tree, con-
cealed by the high grass, in upper Makiki valley. Higher up
Tantalus they are found in similar situations on guava and
other shrubs and trees. They often rest in groups of three
or more on the lower side of the trunk or branches, concealed
by grass or other herbage.

210 ACHATINELLA STEWARTII.

Part of the patterns of producta are nearly or quite iden-
tical with stewartii patterns, and would seem to belong to an
older period than that of the differentiation of producta.

The original descriptions of the several forms referred to
A. stewartii and A. s. producta here follow.

. .

:Achatina stewartii Green. Shell heterostrophe, conical,
oblong, about one inch in length and half an inch in diameter ;
whorls 6 or 7, rounded and marked with numerous oblique and
delicate striae; apex rather obtuse and not eroded; a deeply
impressed line along the upper part of the whorls parallel to
the suture. Periostraca smooth and very glossy. Color and
marking exceedingly various; the ground color is usually
greenish or some shade of yellow ; sometimes a single blackish
colored band accompanied the suture ; sometimes this band is
double and of different shades, and on many specimens there
are two bands, one at the suture, and one in the middle of
the whorls. In some varieties the base of the body-whorl is
dark brown, the rest of the shell being of a dark fawn, and
not infrequently the whole shell is without any markings what-
ever, in which case the color is yellow. The aperture when
inverted is ear-shaped; the truncation of the columella is
rounded and thickened in a remarkable manner at its edge ;
along the inner margin of the outer lip there is a strong
callous ridge, as in most of the species of this genus, which
gradually attenuates towards the edge of the lip, which is thin
and sharp inside, white and pinkish around the columella.'
(Green}.

Oahu, on under side of leaves of the ti plant (Rev. C. S.
Stewart).

Achatinella dunkeri Cuming, Pfr. "Shell imperf orate,
oblong-turrited, rather solid, smooth ; pale flesh colored, or-
namented with close brownish streaks, and frequently en-
circled by one or two black lines. Spire turrited, the apex
rather acute, white ; suture lightly margined, pale. Whorls
6!/2, slightly convex, the last almost two-fifths of the length.
Aperture oblique, truncate-auriform ; columellar fold white,
strong, tooth-like, subsulcate ; peristome unexpanded, acute,
white-lipped behind the brownish edge. Length 24, diam. 11
mm. ; aperture 10 mm. long, 5 wide. Sandwich Is., Dr. New-
comb (Pfr.).

"Achatinella 'bilineata. Shell oblong-conical, dextral, rather
ventricose towards the base, whorls rounded, columella den-
tately plicate ; bright yellow, conspicuously encircled round

ACHATINELLA STEWARTEE. 211

the middle with two close black lines. Hab. ? The form

of this shell is peculiar, and there is something equally char-
acteristic in the marking' (Reeve).

Our pi. 38, fig. 10 is practically identical with Reeve's
figure of bilineata.

Achatinella venulata (pi. 30, figs. 48, 48a, photographic
reproductions of the original figures). "Shell dextral, elon-
gately conic; whorls 7, convex, slightly margined above; su-
ture well-impressed; aperture subovate; lip simple, thick-
ened within ; columella strongly plicate, twisted, of a pinkish
hue ; color of shell white, or pinkish white, with longitudinal
veins of green or olive-green epidermal matter. Length 21/20,
diam. 10/20 inch.

'Var. a. Sinistral, with a deep black sutural band.

"Kolau, Oahu.' (Newcomb).

This is an artifact so far as color is concerned, produced
by scraping A. producta; var. a is a scraped A. stewartii.
The following three forms, johnsoni, aplustre and hybrida are
also manufactured patterns. A. venulata and hybrida were
from Mt. Tantalus shells, perhaps johnsoni and aplustre also.
The locality ' ' Kolau, ' ' given for the whole series, was a further
deception on the part of the shell-artist who imposed the
frauds upon Doctor Newcomb.

Achatinella johnsoni (pi. 30, fig. 50, reproduction of ori-
ginal figure). "Shell dextral, conically elongate; whorls 7,
slightly rounded, superiorly indistinctly margined ; suture
rather deep ; aperture subovate ; columella with a strong pli-
cation, pink, margined with black; color of shell white or
pinkish, banded with two or more narrow black lines, one or
two of which are central, one sutural, and one usually
broader, inferiorly on the last whorl. Length 18/20, diam.
9/20 inch.

"The following form an allied group: A. producta, hy-
brida, aplustre, venulata and johnsoni. This species is dedi-
cated to the Rev. Mr. Johnson of Kawai, whose assistance in
my researches I most thankfully acknowledge' (Newcomb).

A. johnsoni and A. aplustre have somewhat the shape of
A. s. producta, and may have been manufactured from a
rather unusual form of that from Mt. Tantalus, such as pi. 38,
fig. 13 ; but the presence of both sutural and columellar dark

212 ACHATINELLA VULPINA.

bands is more suggestive of the four-banded pattern of
stewartii, such as pi. 38, fig. 6.

Achatinella aplustre (pi. 30, fig. 51, reproduction of ori-
ginal figure). ' Shell conical, dextral; whorls 7, rounded;
suture moderately impressed, simple ; aperture semiovate ; lip
thin ; columella with a callus, terminating in a twisted plait,
bluish-white, tipped with brown ; three first spiral turns white,
the others with a broad central pink band margined by two
deep black bands ; on the last whorl, superiorly and inf eriorly,
is a broad bright green band. Length 1, width 10/20 inch.
Kolau, Oahu.

' The gaudy coloring of this species alone separates it from
the A. johnsoni, nob. It belongs to the same section with
A. producta Reeve, which may be taken as the type of a group
of species numbering some half-dozen, and which requires still
farther additions to render it complete' (Newcomb).

This was soon recognized as a scraped and dyed stewartii
or producta.

Achatinella hybrida (pi. 30, fig. 52, reproduction of ori-
ginal figure). " Shell dextral, conical; whorls 6, rounded,
margined above, the last somewhat inflated ; suture well-
impressed; columella white or brownish, terminating in a
twisted plait; aperture ovate, white; lip simple, thickened
within ; striae longitudinal, fine ; color of shell green, or brown
on the last whorl, above white, with fine longitudinal veins of
the color of the epidermis. Length 18/20, width 10/20 inch.
Kolau, Oahu.

"This species seems to be intermediate between A. producta
Reeve and A. venulata.' (Newcomb).

18. A. VULPINA (Ferussac). Plates 39 and 40 ; pi. 41, figs. 5-7.

The shell is ovate-conic, smaller than A. stewartii (compare
plate's 38 and 39) ; glossy; yellow, green, olive or chestnut,
often banded with green or chestnut ; usually sinistral.

Oahu : Makiki to Manana valleys.

Helix vulpina FERUSSAC, (Prodrome, no. 429, no descrip-
tion), in Freycinet, Voyage autour du Monde, sur I'llranie et
la Physicienne, Zoologie, p. 477, pi. 68, f . 13, 14, 1824 ; His-
toire Nat. Moll. Terr. pi. 155, f. 1, 2. — Achatinella vulpina
Fer., PFEIFFER, Monographia, ii, 237; iii. 464. — Achatinella
castanea REEVE, Conchologia Iconica, vi, pi. 3, f. 24, April,

ACHATINELLA VULPINA. 213

1850. — GULICK, Evolution, Racial and Habitudinal, pi. 2, f.
12. — Achatinella adusta REEVE, C. Icon. pi. 4, f. 30, April,
1850. — Achatinella ernestina BALDWIN, Proc. A. N. S. Phila.
1895, p. 217, pi. 10, f. 5, 6. — Achatinella olivacea REEVE, C.

Icon. pi. 3, f. 20, April, 1850. (Hab. ?).— GULICK,

Evolution, etc., p. 41, pi. 2, f. 13. — Achatinella prasinus

REEVE, C. Icon. pi. 4, f. 27, April, 1850 (Hab. ?).—

Achatinella virens GULICK, Ann. Lye. Nat. Hist, of N. Y., vi,
p. 254, pi. 8, f. 73, Feb., 1858. — Achatinella cucumis GULICK,
t. c., p. 225, pi. 7, f. 45, Dec. 1856. — Achatinella analoga
GULICK, t. c., p. 227, pi. 7, f. 47, Dec., 1856; Evolution, etc.,
p. 41, pi. 2, f. 9 (Halawa). — Achatinella longispira SMITH,
P. Z. S. Lond. 1873, p. 73, pi. 9, f. 5. — Achatinella colorata
GULICK, Evolution, etc., pi. 2, f. 10 (— virens).

A. vulpina as here understood comprises shells of a great
variety of colors and patterns, the shape and size of the
shells remaining practically the same. Numerous " species",
based chiefly upon color characters, have been based upon
forms of vulpina, but the prevalence of undoubtedly hybrid
colonies in which several of them live together opposes that
interpretation of the facts, even though the hybrid colonies
segregate into patterns so distinct that they may usually be
assorted readily, with only a small proportion of "blends.'
Some of the "elementary species'1 or patterns rarely occur
also in pure colonies, which by themselves would be con-
sidered subspecies. Others, like A. longispira are found only
in hybrid colonies, and seem to be products of hybridism in
which the parental patterns are changed rather than blended.

The western edge of Nuuanu valley marks the division be-
tween the western and the eastern herds. The dark olivacea,
virens and longispira patterns have not crossed the floor of
the valley eastward, and the vulpina patterns (typical vulpina,
ernestina and adusta) spread no further west. We would
signalize this fact in distribution by dividing the series into
two species were it not that the light olivacea pattern is com-
mon to both herds, extending from Kalihi at least as far west
as Waimano, though not common we believe west of the
Nuuanu-Kalihi ridge.

214 ACHATINELLA VULPINA.

It appears that an ancestral stock, probably not unlike the
yellowish form of virens, spread over a considerable area;
various mutations arose in different places, and radiated
from their initial points as far as time and conditions al-
lowed, hybridizing with other mutations. Nuuanu, deeply
cut, with a broad level floor, has been a partial barrier to the
spread of mutations reaching it during the present climatic
cycle.

The races from the northern or Koolau valleys have long
been in a measure isolated from those of the Kona slope.
Intrinsically these races are on a par with the elementary pat-
terns of the southern watershed, but by favor of their isola-
tion they have not been caught in the web of hybridism which
clings about the Kona forms. They may properly be recog-
nized as subspecies. See page 224.

Races and elementary patterns of A. vulpina.
(No dark sutural border.)

Chestnut or blackish below, gradually paler towards the
suture, vulpina pattern, pi. 39, f . 1, la.

Upper third of last whorl yellow, lower two-thirds chestnut,
the division abrupt, castanea pattern, pi. 39, f. 2.

Yellow, uniform, or (typically) banded with black-brown,
N.-E. Nuuanu, ernestina pattern, pi. 39, f. 3-4.

Green, either dark or pale, varying to olive or ochreous,
olivacea pattern, pi. 40, figs. 3-4a. A form in which the shell
is somewhat rugose has been named cucumis Gul.

(A brown or blackish sutural border.)
Kona forms.

Base dark chestnut, upper surface lighter or yellow, Pauoa,
adusta pattern, pi. 40, figs. 1, 2.

Green, varying to chestnut or yellow, sutural band narrow,
virens pattern, pi. 40, figs. 5, 5a.

Similar, but with a broad sutural band and rarely another
above the periphery, Kahauiki, suturalis pattern, pi. 40, figs.
9-106.

Having blackish-brown bands i, ii and iii in the prismatic

ACHATINELLA VULPINA.

215

layer of the shell, sometimes increased by splitting of bands,
analoga pattern, pi. 40, figs. 15-15c.

Having green and brown lines on a paler ground, longispira
pattern, pi. 39, figs. 12c, d.

Koolau forms.

Ground yellow above, ochraceous-orange below, with a broad
green zone above the periphery; or dusky without a green
band, pi. 41, figs. 8-8c. A. v. colorata, no. 18a.

Ground-color white above, greenish or olive-brown below the
periphery, with a sharply defined cinnamon band above the
periphery, pi. 37, figs. 10-11&. A. v. tricolor, no. 18&.

Probably the test of breeding would show that not all of
the above-defined patterns are elementary. The castanea
pattern is often not readily separable from vulpina. Adusta
seems to be a stewartii-vulpina hybrid. Longispira also may
be a product of hybridism rather than an elementary pattern.

Usually two or more patterns occur in one colony or often
on one tree, and in any large lot there may be some blends or
unusual patterns.

Table showing distribution of the elementary patterns of

A. vulpina.

VALLEYS.

Adusta.

oJ
C

• —

£*

*3
>

Castanea.

Ernestina.

Olivacea.

tn
P
O>

• i— 1

>

Suturalis.

03

J3

'eS

Ǥ

Longispira.

.MnrKinn

*

^^ainaano

*

*

W^aiau

*

*

\Vaicnalu

?

#

Aeia

*

*

*

#

Halawa

#

*

*

*

Moanalua

*

*

•*

#

Kahauiki

?

*

*

#

*

Kalihi .. .. .

•K

?

*

W. Nuuanu

#

*

*

E. Nuuanu

*

*

*

*

Pauoa.

*

#

*

*

Afakiki

#

#

*

*

216 ACHATINELLA VULPINA.

West to Kahauiki the shells are invariably sinistral. From
the western ridge of Kahauiki westward a few dextral in-
dividuals occur sporadically in the colonies, though sinistral
shells predominate.

Typical form. — A. vulpina was originally based upon rather
small specimens about 19 mm. long, of a pattern which was
subsequently considered by collectors to be a variety of
castanea Reeve. The types were somewhat faded, either by
weather or the application of heat in cleaning, to a reddish
or fox color, which suggested the name. Ferussac also figured
a variant having a darker band over the lower two-thirds of
the last whorl. Subsequently, in the Histoire, Ferussac fig-
ured a large specimen of uniform tawny reddish color,
length 25 mm., probably from the Pauoa slope of Tantalus.
The original description follows.

"Shell sinistral, conic, elongate, vertex obtuse; glossy,
sharply striate ; epidermis fulvous or rufous, fugacious, the
apex pale. Whorls 5%, a little convex ; suture distinct, dupli-
cate. Aperture sernilunate, white ; peristome thickened with-
in ; columella white or rose, arcuate ; umbilical crevice not
distinct.

"a, Rufous, unicolored. Z>, Rufous with a broad brown
band. It inhabits the Sandwich Islands' (Per.). Length
19, diam. 10 mm. (from type figure).

This typical form ranges from Makiki to Nuuanu valley.

PL 39, fig. 1 represents a shell from Nuuanu valley, coll.
C. M. Cooke, exactly agreeing with Ferussac 's type (repre-
sented, I take it, by his fig. 13). It is deep chestnut colored,
darker towards the base, becoming lighter upwards, the first
three whorls buff. There is no dark sutural border. The
surface is very glossy, distinctly marked with growth-striae
on the last 3!/2 whorls; embryonic whorls very delicately
striate spirally. Length 19, diam. 10 mm., G1^ whorls. Other
specimens of the same lot are larger, up to 21 mm. long, and
most of them are less slender. The columella varies from
pale lilac to nearly white. A large specimen measures, length
23.5, diam. 13 mm. It is of course clear that the historic
type-form of vulpina is an abberrant melanistic variation of
a species more normally represented by virens.

ACHATLNELLA VULPINA. 217

Very often the color fades to yellow at the suture, or the
change to yellow may be abrupt, as in pi. 39, fig. la, which
agrees with Ferussac's variant &. This leads to the pattern
figured by Reeve as A. castanea, which is represented by pi.
39, fig. 2, Gulick coll., from the Pauoa side of Nuuanu. The
same form is also found in Pauoa, pi. 40, fig. la.

A. vulpina extends eastward to Makiki valley, where the
chestnut or blackish form with yellow band above occurs in
the northern branch, together with the dunkeri pattern of
producta (L. A. Thurston and Irwin Spalding). Some of the
shells have the rim within the lip rather thick. A. v. olivacea
has been taken in Makiki by Gulick, and recently by Mr.
Thurston in the middle branch. On the Manoa base of
Tantalus Mr. Thurston found olivacea associated with pro-
ducta , johnsoni and dunkeri.

Pauoa. The prevalent form is the adusta pattern (pi. 40,
fig. 2, copy of Reeve's figure; figs. 1, Gulick coll.). The base
is chestnut, or Isabella, with a chestnut band ; above the peri-
phery it is deep colonial buff or ecru-olive, sometimes more
or less suffused with chestnut or green. There is a narrow
band below the suture, usually blackish chestnut, rarely
russet. Probably Reeve's type of adusta was from Pauoa
valley. The castanea pattern (pi. 40, fig. la, Pauoa, coll. by
Gulick) is also found.

The adusta pattern of Pauoa and the slope of Tantalus is
a transition form to A. steivartii. In this borderland the dis-
tinction between the species is vague, and I am unable to
decide upon such forms as pi. 38, figs. 20. Fig. 21 of the same
plate I consider a steivartii, as no vulpina is dextral. When
the olivacea pattern occurs in colonies of such shells, one
may safely pronounce them vulpina.

In the northeastern head of Nuuanu valley there is a local
form of vulpina which has been described by Mr. Baldwin as
A. ernestina. It is rather large and capacious typically (pi.
39, figs. 3 to 6), but varies so much in size and width that no
definite line can be drawn. Fig. 4 is one of Mr. Baldwin's
figured cotypes. Figs. 3, 3a are from one tree (Drac&na),
taken by Dr. Cooke. The shell is either yellow, deepening to

218 ACHATINELLA VULPINA.

yellow ocher on the last part, and fading on the spire, or this
ground may be decorated with bands and lines of chestnut
brown or blackish-chestnut; the banded form being the pat-
tern of Mr. Baldwin's types. It is found only in hybrid
colonies of the two patterns, plain yellow and banded, usually
both on the same tree.

Length 21, diam. 12.4 mm. ; 6% whorls (cotype of ernestina).

Length 19.7, diam. 11 mm. ; 6 whorls (topotype).

On the floor of Nuuanu there is a smaller, narrower race
with the coloration of ernestina. PL 39, figs. 7 to 7c were
taken by Dr. Cooke on one small lehua shrub in the middle
of upper Nuuanu near the old road to the Pali. Three are
banded, one on an ocher yellow ground, the others straw and
primrose yellow. One specimen is bandless, of a "turtle
green" tint, — an olivacea admixture.

Further west, in an isolated clump of nearly dead lehua
trees in the valley near the end of the ridge defining Hille-
brand's Glen, Mr. Richard A. Cooke collected a large series,
pi. 39, figs. 8 to 8e, varying from elongate to conic in shape.
The longer shells have ernestina pattern on light or dark
ground. The shorter ones are straw yellow fading to buff
above, base chestnut, the colors either sharply defined (fig.
8Z>) or blended (fig. 8c) ; yellow with a dark sutural line
(fig. 8a), or amber or straw yellow, uniform or with some
white bands (fig. 8).

In a lot taken in the same place by Dr. C. M. Cooke there
are 22 of the uniform yellow pattern, 12 of the patterns of
figs. Sb, 8c, 1 ernestina pattern. In a lot of 7 individuals
from one tree, all of these patterns occur, four of the pattern
of figs. 85, c, but with a dark sutural line, one each of the
other patterns.

The olivacea pattern without chestnut bands occurs in
Glen Ada, on the southern side of Nuuanu (pi. 39, figs. 11 to
lie, coll. by Pilsbry) the color varies from amber yellow to
paris green or various blends, yellow ocher passing into green
towards the suture and base, or ocher with a few green lines.
The summit is buff, and there are no brown bands. This is a
well-known form, back to the time of Gulick.

ACHATINELLA VULPINA. 219

Length 20, diam. 11.5 mm. ; 6 whorls.

Length 20.3, diam. 10.6 mm. ; 6!/4 whorls.

There is also a stunted valley form of this race. A lot
pi. 39, figs. 9 to 9c, from the floor of Nuuanu, taken by Mr.
B. A. Cooke in an isolated group of two or three lehua trees,
near the southern side of the valley, has no ernestina pattern.
The shells are wax yellow, the same shading below into tawny
(fig. 9c) or very rarely the dark typical vulpina pattern like
pi. 39, fig. 1, or streaked with green on a nearly white or citron
ground (figs. 9, 96). There are often a few faint green
spiral lines, or a very dilute brown sutural line. They have
the small size of other lots from the floor of the valley, length
13 to 17.3 mm. It was abundant, as about a hundred speci-
mens were taken by Dr. C. M. and R. A. Cooke.

Somewhere on the southern ridge Dr. Cooke collected speci-
mens like pi. 39, f. Sb, but with a sutural band. Also a small
form with the base olive, the rest white, embryonic whorls
ochraceous or with a wide ochraceous band. The typical
castanea pattern was taken by Mr. Gulick on the Pauoa side
of Nuuanu (pi. 39, fig. 2), exact locality not known to me.

Northwestern ridge of Nuuanu. — Hillebrand's Glen, (pi. 39,
figs. 12 to 12d, coll. by R. A. Cooke), in a dark and humid
locality on the east side, is probably at the eastern limit of
the longispira pattern. Some specimens are scarcely distin-
guishable from the yellow ernestina pattern, uniform or with
chestnut base ; others have the green olivacea pattern. The
longispira pattern is usually pale yellow with green or green
and brown lines (figs. 12c, d.). This colony is therefore a
hybrid of vulpina, olivacea and longispira.

On the northern ridge of Nuuanu there are other colonies
having the same and other patterns in varying proportions,
such as pi. 39, figs. 13 to 13/; pi. 39, figs. 14, 14a. Also pi.
39, figs. 10, 10a, from a lateral ravine of northern Nuuanu,
patterns of the virens-longispira group, but reminiscent of
castanea. Others specimens from the main northern ridge,
coll. by Dr. Cooke, are similar to the yellow form of ernestina.

1 1 Nuuanu on the Kalihi side ' ' is the locality of several lots
of olivacea in the Gulick collection. The shells are like pi.

220 ACHATINELLA VULPINA.

39, fig. 126. Also virens patterns, the same green color with
a dark sutural band (fig. 14), or yellow, like fig. 136, or
like fig. 13.

It is these lots of which a few are figured in pi. 39, figs. 10,
lOo., 12 to 14a, which connect the vulpina, ernestina, virens
and longispira so inextricably that I do not see the way clear
to recognize more than one species in the series.

The typical form of Reeve's A. olivacea occurs on the sum-
mit of the northwestern ridge of Nuuanu. His figure is
copied photographically in pi. 40, fig. 3. The original descrip-
tion follows: "Shell oblong-conical, sinistral, whorls convex,
thinly margined round the upper part, columella with a
twisted plait. Dark olive-green, encircled with rather ob-
scure brownish lines, columella and interior of the aperture
white.

"The rich dark olive coloring of this species leaves off
suddenly in an abrupt line at the aperture and the remainder
of the body-whorl [parietal wall] is of an orange-fawn tint,

like a shell denuded to that extent of epidermis. Hab. ?'

(Reeve).

I take olivacea to include A. prasina Reeve, as the colors
intergrade. In fact, typical olivacea looks to me like a
prasina which was cleaned with hot water. All I found were
of purer green color. However this may be, there are speci-
mens in coll. C. M. Cooke which agree excellently with Reeve's
figure and description, from the main northern ridge of
Nuuanu. Two are figured, pi. 40, figs. 4, 4a. The shell is
indistinctly streaked with brownish olive on a yellowish olive
ground, and has rather indistinct spiral lines of the darker
shade. There is usually a dark brown line or narrow band
below the suture. The embryonic whorls are dirty buff, and
the next whorl rather brown than olive. Length 22.5, diam.
11.7 or smaller.

On Waolani Peak (pi. 40, figs. 5 to 56, coll. by Pilsbry)
the real prasina form occurs. The last whorl is grass green
(but varying in shade from a deeper to a more yellowish
green), summit very pale buff, and the parietal wall is yellow
ocher. The columellar fold is white. This dark green olivacea

ACHATINELLA VULPINA. 221

pattern is found with similar shells having a dark sutural
border, and pale mauve columellar fold, the virens pattern.
A few shells have some darker green spirals on the last whorl,
suggesting longispira (fig. 5&).

Lower, towards the head of Waolani valley the color be-
comes paler (pi. 40, fig. 5c) the general tone olive-yellow,
varying toward green or yellow ocher in different specimens ;
sometimes with a few green spiral lines ; the sutural border is
often paler, and the spire partly white.

Kalihi valley. A fine series in coll. L. A. Thurston, from
the southeastern branch above the waterfall, contains deep and
light green olivacea, deep chestnut and yellow forms of virens,
and the longispira pattern. It is a hybrid colony.

Kalihi valley is also the locality of Mr. G-ulick's A. cucumis.
It is a slightly roughened form of olivacea and virens. The
color is olive with light yellowish olive streaks, colonial buff
with streaks of honey yellow, or ochraceous buff with blended
chestnut streaks. There is often a dark sutural line, as in
virens, but in a third of the lot the ground-color extends to
the suture. There is rarely a pale peripheral line or nar-
row band. The shape varies from ovate-conic to somewhat
pyramidal. There is a slight rim or thickening within the
lip. All the specimens seen are sinistral. I imagine that the
slight corrugation is due to unfavorable (dry) station of the
individual colony, and probably not a racial character. Two
of the original lot are figured, pi. 40, figs. 6, 6a.

Kahauiki. A hybrid segregating into virens, longispira
and suturalis patterns is prevalent. There are also longispira-
analoga hybrids, and shells suffused and streaked with chest-
nut. These patterns are shown in pi. 40, figs. 7 to 7c, ridge
running up from Fort Shaf ter ; figs. 8, 9, central ridge ; figs. 10
to 10<i, western ridge ; all collected by Mr. Irwin Spalding.
Some from the western ridge also have the pattern of pi.
40, fig. 8.

A special Kahauiki pattern is shown in pi. 40, fig. 9, cen-
tral ridge of Kahauiki. The shell is yellow, green or chestnut,
with a broad chestnut band below the suture. This may be
called the suturalis pattern. It occurs in a nearly pure

222 ACHATINELLA VULPINA.

colony on the central ridge. Out of about 300 shells in Mr.
Spalding 's no. 666, two show bands like longispira and two
have a brown band above the periphery in addition to the
sutural band, which is present in every specimen.

Elsewhere the suturalis pattern has been seen only from the
Western ridge of Kahauiki, where a few occur in the hybrid
colony of that ridge, probably having spread thither from the
vigorous colony of the central ridge. One is shown in pi. 40,
fig. 10a.

An individual fluctuation having a very unusual analoga
pattern, from the Bishop Museum collection, is figured on pi.
40, fig. lOe, dark above, with faintly yellow base. A very
similar color-form has been found by Mr. Theodore Cooke in
Nuuanu valley on the east side.

Kahauiki forms the eastern limit of dextral shells of the
vulpina series. The western analoga pattern also finds here
its eastern limit, at least in the typical development of the
pattern.

Moanalua valley has forms resembling those of central Hal-
awa and of the western ridge of Nuuanu, such as the forms
shown in pi. 39, figs. 12, 13 and pi. 40, fig. 5&, 5c (oUvacea) ;
also longispira pattern (pi. 40, figs. 11, lla), analoga pattern
(pi. 40, fig. 115) and various others (pi. 40, figs, lie, d, e.).
Dextral specimens are rare. All of the above from a lateral
ravine near the top of the southern ridge of the valley, col-
lected by Dr. Cooke (pi. 40, figs. 11 to lie).

In the northern valley of Moanalua, near the head, there is
a large colony of quite small shells, length 14 to 17 mm. The
chief constituent is virens of many shades, but the analoga
pattern appears occasionally, and rarely the longispira pat-
tern (pi. 40, figs. 12, I2a, coll. by Irwin Spalding; fig. 126,
coll. by Dr. Cooke) ; also shells resembling figs. 7, 7c, 15a in
coloration. This somewhat dwarfed race is comparable to the
forms from the bottom of upper Nuuanu. A. lorata from the
same place is also dwarfed. Large series of these diminutive
shells in the collections of Spalding, Cooke and Thurston
show that whatever factor it is in the valley-floor environment
which causes diminution of size, does not affect the produc-
tivity of the snails.

ACHATINELLA VULPINA. 223

Halawa valley is type locality of A. analog a Gulick in which
the shell is two or several banded with deep brown, and
A. longispira Smith, which has green and brown spiral lines
typically, but is never gametically pure, as many other pat-
terns occur in the colonies. There are blends between the
analoga and longispira patterns. There are also specimens re-
ferred by Mr. Gulick to fuscolineata Smith, but which seem
to me to be split-banded individuals of analoga. A selection
from the Gulick series is figured, pi. 40, figs. 15 to 15cZ, analoga,
and pi. 41, figs. 5 to 5g, longispira. A peculiar analoga with
the fuscolineata pattern from the western valley of Halawa is
shown in pi. 40, fig. 13, coll. by Spalding.

In the middle fork of Halawa Mr. Thaanum obtained a fine
set of virens (pi. 40, figs. 14, 14a), pure except for variations
in the ground-color, which may be apricot yellow or olive, as
in figs. 14, 14a, or a deep green, like pi. 40, fig. 10. All are
sinistral.

Aeia. The prevalent variety is a fine form of virens similar
to the Halawa lot described above. The color varies from
chestnut or olive (like pi. 40, figs. 14, 14a, Halawa) to black-
ish green ( pi. 41, fig. 6, Aeia). The summit of green in-
dividuals is frequently marked with chestnut on the first
neanic whorl, sometimes on the embryonic whorls also. Occa-
sionally there are two dark bands, showing some tincture of
analoga blood ; and in some colonies rare specimens have the
longispira pattern. All are sinistral.

In the Cooke collection there are a few shells similar to the
dwarf race of upper Moanalua, the color chestnut below, paler
upwards.

Waimalu valley. Yellow and chestnut forms of virens
with the usual sutural band are in Mr. Spalding 's collection.
All sinistral.

Waiau. Mr. Spalding found green virens patterns, together
with longispira pattern, all dextral.

Waimano. Dr. Cooke collected green virens, olivacea and
a form (pi. 41, fig. 7) resembling color 'ata, except that it has
no dark sutural band.

Manana. On the western ridge Mr. Wilder collected green

224 ACHATINELLA VULPINA.

virens, similar to those of Halawa and Aeia. This is the ex-
treme western limit of the species so far as I know.

Varieties of A. vulpina from the northern or Koolau side of
the mountains. The valleys on this side are very short, formed
by short butresses, and heading up in precipices often quite as
abrupt as the Nuuanu pali, and mainly higher. The ridge can
be attained from the northern side in very few places in this
part of the range. The snail colonies, chiefly on the lower,
less precipitous slopes where trees find foothold around the
valley heads, are therefore in a measure isolated from the for-
ests on top and southward. I have seen considerable num-
bers of these Koolau snails, but all from collections made by
Newcomb and Gulick. The present status of the colonies is
not known to me.

18a. A. VULPINA COLORATA Reeve. PI. 41, figs. 8 to Se.

"Shell oblong-conical, dextral, whorls flatly convex, colu-
mella with a twisted plait, aperture small; yellow, encircled
with a very broad green band, sutures black-brown, columella
light pink. Sandwich Islands ' ' (Reeve ) .

The last whorl is apricot yellow or lighter above, ochraceous-
orange below a wide girdle of grass green or parrot green,
which encircles it above the periphery. Rarely the green band
is wanting. Other shells are darkened by a suffusion and
streaking of chestnut with or without an olive girdle. The
sutural band of deep chestnut is invariable in a large lot be-
before us, collected by Gulick. Out of 59 specimens in coll.
A. N. S. P., 27 are dextral.

Length 20, diam. 10.5 ; 6 whorls.

Length 18, diam. 10.3 mm.

Length 17, diam. 9 mm.

Ahuimanu (Gulick) ; a colorata X virens form in Kahaluu
(Gulick).

Achatinella colorata REEVE, Conch. Icon, vi, pi. 3, f. 18,
April, 1850. — PFR., Monographia iii, 461. — Achatinella con-
sanguinea SMITH, P. Z. S. 1873, p. 73, pi. 9, f. 3. — Achatinella
ustulata NEWCOMB MS., according to Pfeiffer, Monogr. vi, p.
172, no description.

ACHATINELLA VULPINA. 225

In one Gulick lot from Ahuimanu there are transitional

<•

specimens between colorata and the virens pattern, both green
and chestnut, similar to pi. 39, figs. 13, 14. These shells are
like those from Kahaluu, noticed below. Another lot having
the shell banded with deep brown, sometimes with a green
band above the brown, was segregated by Mr. Gulick (pi. 41,
fig. 8e ) , also from Ahuimanu.

A series collected by Mr. Gulick in Kahaluu (a short dis-
tance northwest of Ahuimanu and separated therefrom by a
short spur), contains ill-defined colorata, the green band in-
distinct or diffuse, together with green, yellowish-green and
chestnut shells similar to pi. 39, figs. 12 b, 13 and 14. Most of
this lot are not distinguishable from virens patterns of the
Kona side of the range. The colony is apparently a color at a-
virens mixture. All of them are sinistral.

Kahaluu is as far west as any Achatinellastrum of the east-
ern or vulpina group has been traced on the northern side of
the mountains.

Mr. Baldwin gives Kalihi as a locality for colorata, but he
must have had some unusual color-form of vulpina, or per-
haps he was misinformed.

A lot of about 20 specimens of longispira in the Gulick col-
lections is labelled * * Ahuimanu ? ' ' on the rather dubious au-
thority of Mr. Frick. Probably incorrect.

I have never seen colorata from the Kona side of the range.
Some specimens of olivacea from Moanalua (pi. 40, fig. lie)
and from Waimano (pi. 41, fig. 7) have a broad green band
above, but they lack the dark sutural band of colorata. There
are also occasional specimens of the longispira pattern which
resemble colorata.

A. consanguinea Smith intergrades completely with the
dusky forms of colorata. Probably Mr. Gulick selected the
specimens out of his lot of colorata. The original description
follows.

A. consanguinea E. A. Smith. PI. 41, figs. 9, 9a, speci-
mens from the original lot. "Shell sinistral (sometimes dex-
tral) imperf orate, glossy, striated with oblique growth-lines
and under a lens transversely, and very delicately, variously

226 ACHATINELLA VULPINA.

streaked with olivaceous and greenish-ashen ; lower half of the
last whorl chestnut, streaked with the above-mentioned colors.
Suture distinctly margined with dark brown. Whorls 6y%,
convex, the first S1/^ ruddy. Aperture white; peristome
brown-tinted within and edged with white ; columellar fold
strong, roseate. Length 18, diam. 10 mm.

' ' Var. : shell indistinctly zoned spirally with olivaceous. '
(Smith).

1 ' Ahuimanu, Island of Oahu, chiefly on leaves of the kii.

"This species passes into A. colorata Reeve, which is also
found in Ahuimanu. It differs from A. colorata in the same
manner and proportion as A. adust a Reeve differs from
A. producta Reeve. About one-third of the specimens are
dextral' (Gulick).

18&. A. VULPINA TRICOLOR Smith. PL 37, figs. 10, 11, Her, 116.

"Shell sinistral, ovate-subconic, imperf orate, glossy, very
finely striated with oblique growth-lines and transverse striae,
white, whorls encircled in the middle with a broad yellowish-
olivaceous zone and towards the base streaked with green;
suture distinctly margined with deep brown ; whorls 6!/2> con-
vex, the first two or three white or reddish ; aperture white ;
peristome thin, lightly bordered within; columellar fold
strong, roseate. Length 20, diam. lO1/^ mm.

"Var. a. Shell white, girt about the middle with one oli-
vaceous zone.

"Var. 6. Shell yellowish, encircled with a very broad basal
zone of chestnut. ' ( Smith ) .

Oahu: loleka, in Heeia (Gulick).

Achatinella tricolor SMITH, P. Z. S. 1873, p. 76, pi. 9, f. 6.

In form and color it is intermediate between A. colorata
Rv. and A. varia Gk. (Smith).

A. c. tricolor is very closely related to colorata, which occurs
westward, on the other side of the Heeia ridge. As in colorata,
there is always a narrow brown sutural band or line. Typi-
cally the ground-color is white or nearly so above the peri-
phery, citron-green or light yellowish olive below it, with a
broad cuticular band of hazel or cinnamon-brown above the
periphery, which is marked with a line of the whitish ground,

ACHATINELLA VULPINA. 227

as in fig. lla. Very often, however, the whole base is hazel
or chestnut, and the ground yellowish above. The band may
be a good deal widened, as in fig. 11 &, or much reduced. In
one shell it is split (fig. 10). All the specimens seen are
sinistral.

The figures and the above notes are from the type and para-
type lots in coll. Boston Society and Academy. I have not
studied any recently collected specimens.

The original descriptions of forms referred to A. vulpina
here follow.

Achatinella prasinus Reeve. " Shell oblong-conical, sinis-
tral, whorls convex, obliquely, somewhat rudely impressly
striate, columella short, twisted-plaited ; very dark green, en-
circled with obscure dark lines, white at the apex, columella
white. Hab. ?' (Reeve}.

This is the very dark green form of olivacea. Some speci-
mens from Newcomb are noticeably roughened, like cuciimis
Gul. I assume that they were from an unfavorable station.

Achatinella adust a. (PI. 40, fig. 2, reproduction of original
figure). "Shell acuminately oblong, somewhat turrited, sinis-
tral ; whorls rounded ; columella short, callous, toothed. Dark
chestnut, darker towards the base, with a spiral black band

beneath the sutures; columella pinkish. Habitat ?'

(Reeve).

A. cucumis Gulick. (PI. 40, figs. 6, 6a, Gulick coll.).
"Shell sinistral, imperf orate, acuminately oblong, solid, shin-
ing, striated, green ; apex rather obtuse ; spire turreted ; suture
margined, impressed; whorls 6, convex; columellar fold cen-
tral, white, moderately developed. Aperture oblique, sinu-
ately oval, white within ; peristome thickened within, with ex-
ternal margin unreflected, arcuate, acute ; columellar margin
dilated, adnate ; parietal margin wanting. Length 19,
breadth 9% mm. ; length of body-whorl 13 mm. Average
weight 5.2 grains.

* * Var. &. Of a burnt yellow color.

"Var. c. With one or more obscure brownish bands.'
(Gulick).

Oahu: Kalihi, on trees (Gulick). Kalihi to Moanalua
(Baldwin).

228 ACHATINELLA VULPINA.

Achatinella analoga Gulick. "Shell sinistral, imperforate,
oblong, conic, solid, shining, finely striated, green, yellow or
white, with three black spiral bands, one sutural, one entering
the aperture, the other revolving above the suture ; apex ob-
tuse ; spire turreted ; suture margined, moderately impressed.
Whorls 6, convex. Columellar fold central, brown or white,
moderately developed. Aperture oblique, sinuately oval,
white within; peristome thickened within, with external mar-
gin unreflected, arcuate, acute ; columellar margin dilated,
adnate ; edged with brown ; parietal margin wanting. Length
18y3, breadth 7%, length of body-whorl 12 mm. Average
weight 4.8 grains' (Gulick}.
' Var. a. Yellowish white.
'Var. b. Brown yellow.
"Var. c. Green.

'Var. d. Green or yellow at the base and white above.
'Var. e. With but one or two black bands.
"Var. /. With four or five black bands.' (Gulick}.
Oahu: Halawa (Gulick).

Achatinella longispira Smith. "Shell sinistral, elongate-
subconic, imperforate, glossy, striated with incremental and
very delicate transverse lines, yellow, promiscuously orna-
mented with (4 to 6) narrow spiral zones of green and brown.
Suture distinctly margined with deep brown. Whorls 6y2,
convex, the first 3% whitish. Aperture two-fifths the length,
white ; peristome with the margin acute, bordered within ;
columellar fold strong, roseate. Length 21, diam. 10 mm.'
(Smith).

"Var. a. Shell shorter, ovate-conic. Length 18, diam.
10 mm.

"Var. b. Shell yellow, ornamented with green zones.'
(Smith) .

1 ' Oahu : Halawa is the metropolis of the species ; a few
specimens are reported by Mr. Frick to have come from
Ahuimanu (Gulick}.

"This species is most nearly related to A. analoga Gk. which
is readily distinguished from this by the two black lines upon
the periphery of the whorl. The black or dark brown lines of
that species belong to the solid substance of the shell, while
the green and yellow bands of this species belong to the
epidermis. No dextral form of this shell has been found'
(Gulick}.

A. diluta Smith. PL 40, fig. 16. "Shell dextral, ovate-
subconic, glossy, indistinctly and very minutely striated with
incremental and transverse lines; dilute buff-green, encircled

ACHATINELLA VULPINA. 229

by several obscure, green transverse lines; suture lightly
margined ; whorls 5%, a little convex ; aperture white ; peris-
tome thin, lightly thickened within ; columellar fold strong,
white. Length 18, diam. 9 mm.

"Habitat, the Sandwich Islands. Judging from its affini-
ties, we may believe that it comes from the island of Oahu'
(Achatinella diluta SMITH, P. Z. S. 1873, p. 74, pi. 9, f. 14. —
THWING, Reprint etc., p. 60, "Kalihi").

The photographic copy of Mr. Smith 's figure, pi. 40, fig. 16,
is a little too dark green. I have not seen the type of diluta
in the British Museum, but a specimen agreeing very closely
with the figure, in Gulick's collection, no. 905, appears to be a
dextral form of olivacea. The shape and color, with indistinct
green bands, may be matched in sinistral shells. In the west-
ern part of its range vulpina of several patterns has been
found dextral, and it seems likely that the two specimens of
diluta known are merely rare dextral variants of the common
olivacea. Mr. Gulick has marked his specimen as from
"Halawa?", and I imagine that locality is not far wrong.

A. ernestina Baldwin. (PL 39, fig. 4, cotype). 'Shell
sinistral, subperf orate, moderately solid, ovately conical ; apex
subacute ; surface shining, obliquely striated with delicate
growth lines, under a strong lens exhibiting numerous, very
close, and minute decussating striae; apical whorls smooth,
scarcely decussated. Color yellow, lighter above ; variously
striped with transverse, dark chestnut bands, the more con-
stant being one bordering the suture, one at the periphery,
spiral above, and one encircling the base. Whorls 6, narrowly
margined above, somewhat convex ; suture moderately im-
pressed. Aperture oblique, oval, faintly exhibiting the dark
bands of the exterior; peristome acute, somewhat thickened
within, slightly expanded, white, the coloring of the exterior
bands appearing on the inner edge ; columella white, termina-
ting in a prominent, oblique, tortuous fold. Length 21, diam.
12 mm.

* * Habitat, Nuuanu Valley, Island of Oahu.

' * The animal, when extended in motion, as long as the shell.
Mantle light brown, sprinkled with black and margined on the
outer edge with an interrupted reddish-yellow line. Superior
portion of foot light brown, thickly studded with small slate
spots; under portion of foot light brown with a slate tinge.
Tentacles dark slate." (Baldwin}.

230 ACHATINELLA BELLULA.

Series of A. casta.

The shells are smaller than most of the vulpina group, and
often more slender; they are never green. Species of this
series inhabit ridges from Mt. Tantalus to Helemano.

19. A. BELLULA E. A. Smith. PL 41, figs. 10 to lOd ; pi. 48,
figs. 1 to 18.

"Shell dextral, long-subconic, imperf orate, glossy, very
lightly striate with lines of growth, dilute chestnut (the spire
paler) streaked with darker, and encircled with a few obsolete
lines of a somewhat chestnut color, and a nearly black zone
(sometimes wanting) below the periphery. Whorls 6%, a
little convex; suture lightly bordered with chestnut. Aper-
ture short, % of the length, white inside ; peristome with the
margin thin, slightly thickened within. Columellar fold
rather strong, purple. Length 20, diam. 10 mm.

"Habitat, Sandwich Islands. Somewhat of the form of
johnsoni Newc., but the whorls are less tumid, especially the
last, and the painting is different'1 (Smith).

Oahu : Pauoa slope of Tantalus and the main ridge at head
of Manoa, northwest to the Kalihi-Kahauiki ridge ; only high
on the ridges, chiefly on leaves of lehua, Straussia and guava.

Achatinella bellula SMITH, P. Z. S. 1873, p. 77, pi. 9, f. 8. —
THWING, Orig. Descript., Occ. Papers B. P. B. Mus. iii, p. 56,
pi. 1, f. 15. — A. bellulae Smith, HARTMAN, Proc. A. N. S. Phila.
1888, p. 32. — Achatinella multizonata BALDWIN, Proc. A. N. S.
Phila. 1895, p. 215, pi. 10, f. 1, 2. — COOKE, Occasional Papers
B. P. B. Mus. ii, pp. 65-76.

A. bellula stands close to A. casta, from which it differs by
being invariably dextral, with the spire a little stouter, its out-
lines perceptibly convex. Also it is often larger and more
solid. It is the southeastern member of the casta series. The
locality was unknown to Mr. Smith, but specimens from the
head of Pauoa (pi. 48, fig. 1, collected by C. M. Cooke) agree
perfectly with the figured type, and that place may be con-
sidered the type locality. It is most abundant on the ridges
of Manoa, Pauoa and Nuuanu valleys. The color-forms from
Pauoa are further illustrated in pi. 48, figs, la, 2 ; pi. 41, figs.

ACHATINELLA BELLULA. 231

10 to 10d. The ground varies from white or maize yellow to
burnt sienna with chestnut streaks ; a sutural band is always
present, though often weak, and about one-third of the shells
have nearly black bands below and above the periphery, some-
times also around the columella. The upper band is often
reduced to a line, or pair of lines, frequently absent. The
many-banded forms of Nuuanu are wanting or very rare in
Pauoa. Similar forms extend up the Pauoa side of Mt.
Tantalus — the eastern limit of the species. Hon. L. A. Thurs-
ton has taken it on the Manoa-Koolau ridge, its eastern limit
on the main mountain axis. It does not pass Mt. Olympus.

In the head of Nuuanu valley similar forms occur, also with
patterns shown in pi. 48, figs. 3, 4, coll. by Dr. Cooke. Other
forms are figured from Waolani Peak and towards the head
of Waolani valley (pi. 48, figs. 13 to 16, coll. by H. A. P.).

A series from top of ridge of a northern lateral valley of
Kalihi shows the usual patterns of Pauoa, etc., also a few
shells of a rare pattern, pi. 48, figs. 17, 18.

Near the head of Moanalua the typical pattern was taken by
Dr. Cooke. This was probably on the eastern ridge, as the
shell is not found low in Moanalua.

On the ridges extending into Nuuanu from the north-
western bounding ridge a form occurs, which though inter-
grading completely with bellula yet has a certain racial status
by reason of the prevalence of a copiously banded pattern.
This race has been defined by Mr. Baldwin as follows :

A. bellula multizonata Baldwin. PL 48, figs. 5 to 12.

"Shell dextral, imperf orate, moderately solid, elongately
conical, apex subacute, surface shining, striated with fine lines
of growth, under a strong lens showing very numerous, ex-
tremely minute decussating strias ; apical whorls smooth,
scarcely decussated. Color white, variously striped with nu-
merous dark brown lines and bands, some on the base and
others spiral. Whorls 6, lightly marginate above, convex;
suture lightly impressed. Aperture oblique, oval, white, the
dark bands of the exterior visible within; peristome acute,
thickened within, slightly expanded, white, the dark lines of
the exterior marked on the inner edge ; columella purplish

232 ACHATINELLA BELLULA.

brown, terminating in a strong, oblique, tortuous fold. Length
18, diam. 10 mm. Nuuanu Valley.

"Animal, when extended in motion, longer than the shell.
Mantle brown, lighter on the outer edge. Foot above and be-
low light brown ; posterior portion tapering. Tentacles long
and slender; these, with the head above, slate color.'
(Baldwin).

"A great variety of transition forms occur between this
species and A. bellula Smith, which is found on the neighbor-
ing mountain ridges of Nuuanu Valley, and is a much larger
shell. The animals of the two extremes are specifically dif-
ferent. The mantle of the latter is black, whereas that of the
former is brown, and that of the intermediate forms varies
from black to brown. ' (Baldwin).

Specimens of the type lot are figured, pi. 48, figs. 5, 6.

Dr. C. Montague Cooke has published a detailed study of
multizonata from which the following extracts are quoted:
"Nuuanu valley has a north-easterly trend. The sides are
more or less precipitous and rise from a few hundred to more
than two thousand feet above the bed of the valley. The
sides are covered by a low dense mass of trees, shrubs, ferns
and creepers. Extending into the valley, at about right angles
to the sides, are numerous sub-ridges. The upper portions of
these sub-ridges and of the valleys between them are also over-
grown with a dense mass of vegetation. The lower portions
and also the bed of the main valley are covered with the intro-
duced 'Hilo grass' (Paspalum conjugatum) with here and
there clumps or isolated individuals of lehua (Metrosideros
polymorpha), Straussia, guava (Psidium guayava), etc.-
There are about twenty-three of these sub-ridges, which are
more or less parallel to one another. In some cases the foot
of a sub-ridge expands into a more or less undulating slope.
In numbering these sub-ridges the numbers begin at the head
of the valley.

"A. multizonata is found on all the upper 17 sub-ridges, a
district of about a mile in length, and from 100 to 400 yards
in breadth. There are no permanent streams in any of the
valleys between these sub-ridges. Two of these valleys usually

ACHATINELLA BELLULA. 233

have a little flowing water. The valley between sub-ridges
XIV and XV serves as a boundary to several of the color-
varieties. That between XVII and XVIII serves as a boun-
dary to the whole species. None of the localities are more
than 150 yards in diameter. A. multizonata is found in more
or less open localities. Specimens are seldom found where
there is a dense vegetation, the limit being at the thick growths
of the 'stag-horn fern' (Gleichenia dichotoma} and ieie (Frey-
cinetia arnotti). The elevation at which it is found is from
about 1000 ft. to about 1400 ft. Of the shells found, 95.5 per
cent, were on either lehua, Straussia or guava. Lehua repre-
sents 53.3 per cent, of the trees on which shells were found,
Straussia 9.8 per cent., and guava 29.8 per cent. Of A. multi-
zonata 54.6 per cent, were found on lehua, 14.2 per cent, on
straussia, and 26.7 per cent, guava. Of the remaining plants
P&deria fcetida, representing 1.7 per cent, [of the trees on
which shells were found] yielded 1.2 per cent, of the shells;
Kadua, representing 2.3 per cent., yielded 1.2 per cent. ; and
all others, about a half a dozen genera, representing 3.1 per
cent., yielded 2.1 per cent.

"In all the specimens collected by the writer the mantle
varies from a very dark to a light slate color. It is sometimes
mottled with light or dark markings. A. bellula is not only
larger [than multizonata] but the shell is more solid and the
median whorls less convex. A. bellula varies to a much less
extent. A. multizonata varies from a pure white to a rich
mahogany brown, and also through a larger number of striped
variations. The apex of the shell also varies. In about half
of the color-varieties the apex is white or a very light brown ;
the rest have the apex striped with a light to a very dark
brown band. The color of the apex is nearly constant in each
color-variety. In color-variety W, however, slightly more
than half the shells have the apex white, the remaining having
a banded apex.

" Young, dissected from the uterus of the animal, usually
agree in coloration, with the apex of their parent. Several
exceptions have been found of shells with a white apex con-
taining a striped young, while only four cases have been found
of shells with a striped apex containing white young.

234 ACHATINELLA BELLULA.

'For convenience the writer has divided this shell into
twenty-five color-varieties. Each of these color-varieties is
designated by a letter. The letters thus used are A-T and
V-Z. A very large number of intermediate specimens occur.
Some of these color-varieties have a very restricted range, be-
ing found on only three or four of the sub-ridges ; others are
found over nearly the whole range of the species. These color-
varieties fall naturally into seven groups.

' ' The first of these groups, and also the largest, is made up
of the color-varieties A-H. It is easily recognized, as none
of the shells are marked with dark bands or lines. The shells
vary from white to yellow. The apex is white to light brown ;
never striped. The columella is usually white, sometimes pur-
ple. This group represents about 37 per cent, of A. multi-
zonata. Of this group 63 per cent, were found on lehua, about
16 per cent, on Straussia, and 17 per cent, on guava.

"The second of these groups contains the color-varieties
I-M. The shells are white to yellow, sometimes marked with
brown. All the shells of this group are encircled at the per-
iphery by a dark brown band, or two brown lines. The apex
in every case is white or light brown. The columella is usually
purple, almost never white. This group is only found from
sub-ridges I-XIV. It contains nearly 19 per cent, of the speci-
mens of A. multizonata. Of this group 53 per cent, were
found upon lehua, 15 per cent, on Straussia, and 24 per cent,
on guava ; 48 per cent, of this group were found on sub-ridges
IX-XIII.

"The third of these groups contains the color-variety N.
This color-variety is so distinct from all the other color-varie-
ties that it is thought best to place it in a group by itself.
All the specimens came from sub-ridges III-V, 62 per cent,
coming from sub-ridge IV. About 40 per cent, were found on
lehua, 21 per cent, on Straussia, and 27 per cent, on guava.

"The fourth group contains the color- varieties 0-Q. This
group is distributed over nearly the whole range of the species.
It is absent, however, from sub-ridges IX-XI. The shells of
this group are striped with from four to fifteen lines or bands.
The apex is striped with a light brown band. The columella

ACHATINELLA CASTA. 235

is usually purple ; in only a very few cases is the columella
white. Of this group 48 per cent, came from sub-ridge XIV ;
44 per cent, were found on lehua, 12 per cent, on Straussia,
and 43 per cent, on guava.

"The fifth group is made up of color- varieties E-T. The
shell is white, variously striped ; the apex is white ; columella
is usually purple. About 60 per cent, were found on lehua,
15 per cent, on straussia, and 19 per cent, on guava. This
group is distributed over nearly the whole range of the species ;
only five specimens were found below sub-ridge XIV.

"The sixth group is made up of the color- varieties V-Y.
The shells are variously colored from a rich mahogany brown
to a pinkish purple. The apex is usually banded; in some
cases it is white. The columella is usually purple. All ex-
cept two specimens came from sub-ridge XV-XVII; 75 per
cent, were found upon lehua. This group comes nearest, in
color, to A. bellula Smith, but differs in form and size from
that species [but was later considered to be bellula by Dr.
Cooke].

"The seventh group is made up of the color-variety Z.
This color-variety agrees with several of the other color-varie-
ties in the markings of the lower whorls, but differs from them
in the very dark banded apex. Of this group 96 per cent,
were found on sub-ridges I and II ; 70 per cent, were found

on guava, and 23 per cent, on lehua There are seven

of the color-varieties of which over 95 per cent, came from
two or three consecutive ridges. ... In this species it can be
seen that isolation plays a greater part than environment. '

20. A. CASTA Newcomb. PI. 41, figs. 12 to 16 ; pi. 49, figs. 1 to 16.

' ' Shell conically elongate, solid, polished, shining, dextral or
sinistral ; whorls 6, rounded, margined above ; aperture sub-
ovate, rather small ; lip simple, thickened within ; columella
short, with a strong plaited brownish tooth. Color white or
yellow, with extremely variable transverse bands of black,
brown, pink or white, variously arranged. Length 13 twen-
tieths, width 6!/2 twentieths of an inch. Ewa; its northern
limit is the valley below Mouna Rua, and a half dozen ravines
south, in the district of Ewa': (Newcomb).

236 ACHATINELLA CASTA.

Oahu: Waimalu-Waiau ridge to the Waipio district (col-
lections of Spalding, Cooke and A. N. S.).

Achatinella casta NEWC., P. Z. S. 1853, p. 134, pi. 22, f . 12 ;
Ann. Lye. N. H. of N. Y., vi, 325. — PFR., Monographia iv,
538 ; vi, 174. — THWING, Orig. Descriptions, p. 64. — Eburnella
casta Newc., HARTMAN, Proc. A. N. S. Phila. 1888, p. 36. —
Achatinella cuneus PFR., P. Z. S. 1855, p. 205. — SYKES, Fauna
Hawaiiensis p. 323 (mountains behind Ewa, Perkins). — Acha-
tinella concolor E. A. SMITH, P. Z. S. 1873, p. 75, pi. 9, f. 1.—
Achatinella pygm&a E. A. SMITH, P. Z. S. 1873, p. 75, pi. 9,
f. 11.— Achatinella ligata E. A. SMITH, P. Z. S. 1873, p. 76, pi.
9, f. 13; and var. a, fig. 12. — GULICK, Evolution, Racial and
Habitudinal, p. 41, pi. 2, f. 8.

A. casta differs from A. bellula by being sinistral as a gen-
eral rule, but eastward there are dextral shells among the
sinistral in some places. It has a somewhat different range
of color-forms. There is often a dark sutural band or line,
but more frequently none. Newcomb's figure, reproduced in
pi. 29, fig. 12, is now a rather uncommon color-form, if found
at all in recent collections. It is nearly white with a chestnut
sutural line and an olive band below the periphery, with
several indistinct spiral lines above it. One from Newcomb
is illustrated in pi. 49, fig. 1. Others of the same lot have the
base yellow, tinged with chestnut, or the last whorl pale yellow,
bandless; white; or white with two to four chestnut bands.
Perhaps the lot represents several localities, though casta
colonies are notoriously polychromatic.

Dr. Newcomb apparently included the later-described bellula
in his conception of casta. Indeed the locality given by him
applies to A. bellula alone, if I am right in identifying his
' * Mouna Rua ' ' with Moanalua. He says : "Its northern limit
is the valley below Mouna Rua, and half a dozen ravines
south, in the district of Ewa.' It is practically certain how-
ever, that Newcomb's figured type came from west of Moana-
lua ; moreover, the ravines south of Moanalua can not properly
be said to be in Ewa. Newcomb's specimens look like Wai-
mano or Waiawa shells. Just eastward of the casta area is
the habitat of A. juddii, and still further east, that of A. bel-

ACHATINELLA CASTA. 237

lula. According to Mr. Thwing, A. cast a ranges from Halawa
to Waipio ; but this extension eastward is very doubtful.

Numerous supposed species have been based upon forms of
casta, but after seeing the great series brought together by
Messrs. Spalding, Thurston, Cooke and Thaanum I agree with
Dr. Cooke that no sufficient ground exists for admitting them
as races. They are, we believe, selected forms out of colonies
which in other specimens "run into' other forms of casta.
The explanation of Mr. Gulick's over-division of casta (which
lured Mr. Smith into describing so many forms), is found in
the fact that there is a marvellous opulence of color-mutation.
Very often some special pattern predominates in, or at the
height of its development is special to, a certain colony.
Working with small series from scattered colonies, the differ-
ences are unduly obvious. A. casta deserves monographic
treatment, with several plates, but this work cannot properly
be done except in Honolulu.

Waimano. PL 49, figs. 2, 3, 4, 5, coll. by C. M. Cooke, on
northern ridge and lateral ridges of the valley. The most
abundant pattern is some shade of yellow, more or less passing
into brown, with darker lines or bands. There are also a few
copiously streaked shells, with two dark bands ; two of this
pattern being dextral. Others have dark zones which nearly
cover the surface. Two shells are deep chestnut with lighter
streaks. Other Waimano patterns are shown in pi. 41, figs,
figs. 13 to 13c, Cooke collection.

One lot contains some specimens which approach A. c. mar-
garetce in coloration, having two tawny bands on a whitish
or yellow ground, and a pink columella (pi. 49, fig. 5). The
shape and texture are the same as in other patterns of casta
found with them. Mr. Spalding found the same pattern in
the eastern ravines of Waiawa.

Some Waimano shells, such as pi. 49, fig. 4, approach
A. pupukanioe, but on account of the form of the columella
in old shells, I believe that the latter is an entirely distinct
species.

PL 41, figs. 14-14& are from the A. cookei ridge, Thaanum.

Waiawa. A. casta is found in both division ridges of

238 ACHATINELLA CASTA.

Waiawa in many places, and everywhere in colonies of several
colors and patterns. PL 41, figs. 15, 15&, 15&, 16 ; pi. 49, figs. 7,
8, represent specimens collected by Mr. Spalding, part of
his no. 1863. The rarest and almost unique patterns figs. 15
to 15& still belonging to his collection. PL 49, figs. 9 to 15 are
specimens collected by Mr. Kuhns and given me by Mr.
Thaanurn. The darker shells of this lot also being rare pat-
terns. The ground-color is white, shell-pink or yellow, band-
less or with two dark zones (brown to almost black), or rarely
with many lines. A sutural dark band is often present. The
columella is almost always lilac, but rarely white in albino
shells. The embryonic whorls are usually white, very rarely
having a broad brown band. Length 18, diam. 9.6 mm.

In the western ravines of Waiawa Mr. Spalding found a
small form, length 15-16 mm., with the last two whorls yellow,
spire ivory yellow indistinctly streaked with cream buff. It
is practically identical with the form from Waipio called
A. pygmcea by Mr. Smith.

The original descriptions of forms now referred to A. casta
here follow.

"AckatineUa ligata Sm. [PL 24, fig. 21]. Shell dextral (or
sinistral), elongate-ovate-subconic, moderately shining, very
minutely striated with growth and transverse lines; whitish,
more or less streaked with buff-olivaceous, encircled with usu-
ally two pale brown zones, one above, the other below the per-
iphery. Whorls 5!/2> a little convex, suture distinctly mar-
gined, girdled with deep brown. Aperture white ; peristome
slightly edged within, columellar fold strong, roseate. Length
19, diam. 10 mm.

"Var. a. Shell more lengthened. Length 21, diam. 9 mm.
(pi. 24, fig. 20).

'Var. b. Shell white, the last whorl encircled with several
narrow yellow zones.' (Smith).

Oahu: ''reported to be from Waimalu.' Type in British
Museum.

''It has somewhat the aspect of dextral forms of A. varia
Gk., but is more elongate and cylindrical' (Smith).

Achatinella ligata was based upon mainly dextral shells of
the pattern of pi. 41, fig. 13&. The figures on pi. 24 are

ACHATINELLA CASTA. 239

copied from Mr. Smith's, the specimens having been given by
Mr. Gulick. A. ligata has usually been considered' a form of
A. bellula by Hawaiian conchologists, but the figures and de-
scription agree with a form of casta from Waimano. A set
of four in the Gulick collection, three dextral and one sinistral,
seem to me to belong to casta. The type lot, from Gulick,
was "reported to be from Waimalu,' which is further west
than A. bellula is known, and near the eastern limit of the
range of casta. It seems that towards its eastern limit A. casta
approaches A. bellula in characters, and is sometimes dextral.
A. ligata was based upon specimens of this somewhat inter-
mediate form. The dextral form probably occurs too inti-
mately blended with casta to be segregated as a subspecies.

1A. pygmcca Smith. PL 49, fig. 16. Shell sinistral, ovate-
conic, glossy, striated with extremely fine growth and trans-
verse lines, yellow. Suture lightly margined. Whorls 5%,
convex, the first 3 1/2 white. Aperture white ; peristome slightly
bordered ; columellar fold moderately large, rose. Length 14,
diam. 8 mm. Var. : shell whitish, encircled above the per-
iphery with a line, below with a zone of brown. Waipio'
(Smith}.

The type specimen is figured. It measures 14.7 mm. long,
8 wide, and differs from unicolored Waiawa casta only in the
smaller size. Possibly pygmcea can be retained as a varietal
name for the small western casta, if size proves to be corre-
lated with distribution, which is not likely. Waipio is near
the western limit of A casta. Mr. Spalding collected a similar
small form of casta in the western ravines of Waiawa, but it is
not quite identical, having the spire noticeably broader.
However, the variation from pygmcea is not greater than
among individuals in other Waiawa colonies.

A. concolor E. A. Smith, [pi. 41, fig. 12, type]. "Shell
sinistral, ovate-conic, glossy, striated with growth-lines and
transversely, even on the first 3 whorls ; yellow ( sometimes
streaked with dilute brown) ; suture distinctly margined.
Whorls 6, convex, the first 3 whitish. Aperture white ; peris-
tome thin, the columellar fold strong, roseate. Length 19,
diam. 10 mm. Ewa, Frick.' (Smith.}

240 ACHATINELLA CASTA.

Merely a color-form of A. cast a. The type specimen is no.
76 Boston Society of Natural History. It has been figured
by Smith, and is shown on pi. 41, fig. 12. The last 1%
whorls are streaked with ochraceous-tawny on a light ochra-
ceous-buff ground, which fades to white on the upper whorls.
There is a faintly darker line below the suture on the last
whorl ; columella fold pale lilac. Length 18, diam. 10 mm.
This coloration is rare in casta, but is a common pattern in
A. bellula, which differs by being dextral.

Mr. Spalding has found shells in the southeastern ravines of
Waiawa (his No. 582) which agree exactly with concolor ex-
cept by being smaller. They occur with a more slender
banded form with weak columellar fold (pi. 49, fig. 6) recall-
ing juncea; with white specimens becoming light yellow on the
last whorl, or with a yellow base ; and with various brown-
banded and streaked forms.

Achatinella cuneus Pfr. "Shell imperf orate, whitish or
buff, variously ornamented with narrow brown bands. Spire
convexly conic, the apex white, rather acute ; suture bordered
by a chestnut line. Whorls 6, nearly flat, the last slightly ex-
ceeding one-third of the length. Aperture nearly diagonal,
semioval, white within ; columellar fold moderate, obliquely
somewhat twisted ; peristome acute, labiate within, columellar
margin adnate. Length 18, diam. 8 mm., aperture 8 mm.
long, 3i/2 wide. Oahu, Frick' ' (Pfr.) .

A. cuneus has been placed in the synonymy of A. decipiens
by Dr. Newcomb. Mr. Sykes allows it to stand as a species
in Achatinellastrum. There appears to be no tangible differ-
ence between the description of cuneus and A. casta, and Dr.
Cooke, who examined the type, considers it a color-form of
casta.

20a. A. CASTA MARGARETS Pilsbry & Cooke, n. subsp. PL 42,
figs. 9, 10.

The shell is sinistral, imperforate, thin, ovate-conic, white
with two cinnamon bands, the lower usually wider, and both
tessellated by the crossing of paler streaks; there is a pale
pink band below the suture and a darker one bounding the

ACHATINELLA JUNCEA. 241

columella, the fold of which is lilac. There is also an albino
form in the same colony, pure white throughout. Surface
glossy, similar to A. cast a. Whorls convex, the suture mar-
gined on the last whorl. Columellar fold spiral, not very
prominent. Peristome thin, acute.

Length 14.2, diam. 8.6, aperture 7.2 mm. ; 5^ whorls.

Length 14.8, diam. 9, aperture 7 mm. ; 5~y2 whorls.

Oahu: Kolokukahau peak, at the head of Waiau valley on
the Waimalu division ridge, elevation 1450 ft. (Irwin Spald-
ing, Margaret and L. A. Thurston). Cotypes in coll. A. N.
S. P. and Bishop Mus. ; also in the collections of Spalding
and Thurston.

Margaret's Achatinella is a thinner, less solid shell than
A. casta, texture and coloration resembling A. juncea, which
differs by its lengthened, slender contour. It appears to be
closely related to juncea, but the habitats of the two are rather
widely separated and the forms seem sufficiently distinct. No
evidence that it actually inter grades with the polymorphic
A. casta has been seen in the collections examined, though
its color-pattern is much like one of the common casta pat-
terns. It is doubtless, like A. juncea, of common ancestry
with casta.

A. c. margaretce was discovered by Miss Margaret Thurs-
ton in March, 1909. "Four visits have resulted in finding it
on only three lehua trees. ' It is a shell of the misty summits.

21. A. JUNCEA Gulick. PL 42, figs. 11 to 14.

"Shell sinistral, imperf orate, elongately and acuminately
ovate, thin, shining, finely striated, snow white ; apex some-
what acute ; spire convexly turrited ; suture margined, well
impressed; whorls 6, convex; columella white, with a light
twisted fold near the body whorl. Aperture oblique, oval,
white within ; peristome moderately thickened, with external
margin slightly expanded anteriorly, arcuate, acute ; colu-
mellar margin narrow, adnate ; parietal margin very thin.

" Var. &. With two or three brown spiral bands" (Gulick).

Length 16, diam. 8, length of aperture 7.1 mm. (type
specimen).

242 ACHATINELLA PAPYRACEA.

Oahu: Kalaikoa, Wahiawa [type loc.], and Helemano, on
ahakea. (Gulick).

Achatinella juncea GUL., Annals of the Lyceum of Nat.
Hist, of N. Y. vi, p. 230, pi. 7, f. 49. December, 1856.

A. juncea is a thinner, narrower, more lengthened shell than
A. casta, with the columellar fold usually weaker. It is
narrower than papyracea, with different color-patterns. From
what I have seen it appears to be distinct from related species.

The specimen figured by Gulick and marked l ' type ' ' by him
(pi. 42, fig. 12) is from Wahiawa. Except for a yellow tint
at the edge of the lip it is a pure white shell. Others in his
collection have two cinnamon bands, one above, narrower, the
other wider and below the periphery (fig. 11) ; or the upper
band may be split, and a dark area surround the columella,
the whole base having a faint brown tint. Mr. Spalding
found substantially the same forms in Kipapa, Wahiawa ;
(a) white, (&) white with bands (of which there may be
four, sutural, two wider median, and columellar), the lip
edged with vinaceous brown, columellar fold deep vinaceous,
and (c) streaked with cinnamon on a warm buff ground, fad-
ing to white above (pi. 42, figs. 13). The length varies from
13.5 to 17 mm. There is a similar lot in coll. C. M. Cooke,
including some quite broad forms, length 13.7, diam. 8 mm.
(pi. 42, fig. 14).

Specimens from Helemano are similar to white ones from
Wahiawa. I have not seen specimens from Kalaikoa.

At Nichol's camp, Kaukinehua (in the Wahiawa district)
Mr. Spalding collected a series of six, 3 streaked and band-
less, the others with three bands, of lighter tint than in the
Wahiawa lot described above.

SERIES OF A. PAPYRACEA.

This group consists of thin, rather capacious shells of the
interior valleys, related to casta and its allies. A few species
from the Waianae range are about equally related to this
group and the next. The species seems not to extend so high
as other forms, and none has ever been found on the northern
side of the main range. While some forms have plumbeous

ACHATINELLA PAPYRACEA. 243

or ecru streaks, like the livida group, the embryonic whorls
never have an ocher-yellow band below the suture, such as
occurs in most of the livida series. The embryo often has a
faintly gray or dusky tip.

22. A. PAPYRACEA Gulick. PL 42, figs. 7, 8 ; pi. 54, figs. 8 to
10, 14, 14a.

'Shell sinistral, imperf orate, ovate-conic, thin, polished,
finely striated, light gray or of leaden ash color, with obscure
brown spiral lines ; apex subacute ; spire convexly conical ;
suture marginate, impressed; whorls 5%, moderately convex;
columellar fold central, usually white, slightly developed and
not strongly twisted; aperture oblique, sinuately oval, white
or gray within ; peristome scarcely thickened, with external
margin unreflected, arcuate, acute ; columellar margin nar-
row, adnate ; parietal margin wanting. Length 16%, breadth
9%, length of body-whorl 12 mm. Of a large specimen,
length 19, breadth 10%, length of body-whorl 13% mm-
Average weight 2.5 grains.

"Var. &, without the brown lines' (Gulick).

Oahu : Kalaikoa, Ahonui and Wahiawa, on the hala, Pan-
danus odoratissimus, and other trees (J. T. Gulick). Waipio
to Kalaikoa.

Achatinella papyracea GUL., Ann. Lye. N. H. of N. Y. vi,
p. 229, pi. 7, f . 48 ; December, 1856.

Mr. Gulick 's type (pi. 54, fig. 8) was from Kalaikoa, where
many of the shells have the spiral lines and bands more nu-
merous and distinct than in those from Wahiawa. The
ground-tint is white with light brownish vinaceous streaks
(very faint or almost wanting in some shells, deeper in others,
— more of a light vinaceous purple). The bands vary in
tint, from walnut to snuff-brown or brownish vinaceous. They
are irregular in width, shade and distribution, but there is
usually a peripheral light zone, and the subsutural band is
not emphatic, though usually present. The embryonic whorls
are white or buff, but so far as I know they are never bi-
colored, ochraceous and white, as in the livida series. Colu-
mellar fold usually weak, white. The lip is thin and simple.

244 ACHATINELLA JUDDII.

The type specimen measures, length 16, diam. 9.7, aperture
7.7 mm., but others of the lot from Kalaikoa, are larger,
length 18.1, diarn. 10.5 mm.; whorls nearly 6 (pi. 42, fig. 8).
Some shells from Kalaikoa are more streaked, with faint
bands, as in those from Wahiawa.

Shells from Ahonui, Gulick coll., are streaked as in those of
Wahiawa, but in some there are broad white zones cutting the
streaks into bands (pi. 42, fig. 7).

In "Wahiawa" (pi. 54, figs. 14, 14a, Gulick coll.) and Wai-
kakalaua (pi. 54, fig. 10, coll. by Spalding) the bands are
merely indicated by pale spirals which cross the copious
streaks, which are of a light Payne 's gray or violet plumbeous
tint. This pattern also occurs in the more western localities,
as noted above. It is constant in a series of about 150 taken
by Mr. Spalding in Waikakalaua. Also in a series from " Wai-
pio, ' ' Spalding coll.

In a series from Waipio district, recently received from Mr.
Thaanum, collected by Mr. Kuhns, the pattern consists of
brown linear streaks on a pale Quaker drab ground, white,
below the suture and often at the base. This pattern usually
disappears rather abruptly on the last whorl or sometimes on
the penultimate (pi. 54, figs. 9, 9a, 96). There is also a very
pale specimen, yellowish at the base, with a chestnut-black
band below the periphery, another around the columella. It
approaches some forms of A. juddii.

SERIES OF A. LIVIDA.

23. A. JUDDII Baldwin. PL 49, figs. 17 to 22.

"Shell dextral, imperforate, solid, pyramidally conical,
apex obtuse ; surface shining, covered with very delicate in-
cremental lines ; the nuclear whorls smooth. Color light gray,
shading into light chestnut on the apical whorls, the gray
more intense under the cuticle ; with two black lines, one be-
low and one at the periphery, the latter faint and continued
on the spire; between the lines a white band which revolves
on the suture to the very tip of the apex. Whorls 6, mar-
gined above, slightly convex ; suture lightly impressed. Aper-
ture oblique, oval, white, the light gray of the exterior surface

ACHATINELLA JUDDII. 245

exhibiting a darker shade within; peristome acute, slightly
thickened within, a little expanded, columellar margin very
slightly reflected, white, the coloring of the exterior dark
lines reappearing rather more intense on the inner edge;
cohimella white, terminating in a moderately developed flexu-
ous fold. Length, 15; diam. 9y2 mm.' (Baldwin).

Oahu: Halawa (Baldwin) ; Aiea (Cooke, Spalding) ; east-
ern crest of Kalauao valley, and Kalauao-Waimalu ridge
(Spalding).

Achatinella juddii BALDWIN, Proc. Acad. Nat. Sci. Phila.,
1895, p. 216, pi. 10, f. 3, 4.

'No opportunity for an examination of the animal has yet
occurred. The shell is typically very distinct from any other
known species. The light chestnut band on the apical whorls
is a characteristic and invariable feature. The basal portion
of the shell sometimes has a yellowish hue. In immature
shells the colors are more intense.

"Named in honor of Hon. A. F. Judd, Chief Justice of the
Republic of Hawaii, by whose son the shell was discovered.'
(Baldwin).

A. juddii was discovered by Mr. Albert F. Judd, the present
President of the Bishop Museum Board of Trustees. It is
related to A. papyracea, a more capacious shell. By the color
of its embryo and the gray-streaked pattern it is also related
to A. livida from which it differs by the more pyramidal shape
and various details of coloration. It is rather widely separ-
ated from other members of the papyracea and the livida
groups, and seems to be quite distinct from all other species.

Figs. 17, 18 represent two of the type lot. All of the
Halawa shells sent by Mr. Baldwin (which are probably from
the Halawa- Aiea ridge) have a gray-streaked zone above the
periphery, bordered by a blackish or darker gray line below,
a white or yellowish subperipheral belt followed by a blackish
or dark-gray band, the base being white with faint gray
streaks, or straw yellow. The embryonic whorls are bicolored,
ochraceous with a white zone above the suture.

In a series from Aiea, collected by Dr. Cooke on lehua trees
at top of ridge, the same pattern predominates ; but there are

246 ACHATINELLA LIVIDA.

also specimens with the two bands near the periphery and
no other markings (fig. 21), and others without bands, white
above, the base more or less extensively yellow, embryo colored
like the typical pattern (fig. 22).

On the Kalauao-Waimalu division ridge Messrs. Spalding
and Kuhns found the typical pattern, together with specimens
lacking dark bands, fig. 19 ; and others having several ac-
cessory black bands, fig. 20, coll. by Spalding. There is also
in Mr. Spalding 's collection a single specimen with black base
and a broad black band above the periphery. Also a few
others having a small black basal area.

Very few specimens from any locality lack the brown or
ochraceous zone at the summit ; probably all young shells show
it, though on rare adults it may disappear by fading.

A. juddii is rather difficult to place in the series, as it has
affinities with casta, papyracea and livida, and seems more
related to the last two, though its habitat lies between the
areas of ~bellula and casta.

24. A. LIVIDA Swainson. PL 53, figs. 1 to 9.

1 1 Shell reversed, ovate, obtuse, livid brown or greyish ;
spire thickened ; suture with a deep fulvous line. A small
reversed species, unbanded, and scarcely exceeding half an
inch in length. In form it perfectly resembles the green
variety of Bulimus citrinus. The three specimens in our
museum vary in color from a light olive brown to a livid
purple which lies in longitudinal shades, and gradually
changes on the spiral whorls to white. Suture marked by a
line of deep orange brown. Aperture white, tinged with
purple' (Swainson).

Oahu : Wahiawa to Kawailoa district.

Achatinella livida SWAINS., Quarterly Journal of Science,
Literature and the Arts, iii, 1828, p. 8.5; Zoological Illustra-
tions iii, p. 108, pi. 108, f . 2. — Eburnella livida Swains., Hart-
man, Proc. A. N. S. Phila., 1888, p. 36. — Achatinella viridans
Mighels, REEVE, Conch. Icon. pi. 4, f. 25 (not A. viridans
Mighels) =A. reevei C. B. ADAMS, Ann. Lye. N. H. of N. Y.
v, p. 44; Contributions to Conchology No. 8, p. 128 (1850).-

ACHATINELLA LIVIDA. 247

Achatinella emmersoni NEWC., P. Z. S. 1853, p. 156, pi. 24,
f. 74. — THWING, Orig. Descriptions, etc., p. 71. — A. emersonii
Nc., Ann. Lye. N. H. of N. Y., vi, p. 309.

Specimens collected in Kawailoa in 1854-5 by Mr. Gulick
agree perfectly with Swainson's description and figure. In a
lot of 23, 14 are dextral. Shells of this lot are figured, pi. 53,
figs. 1 to 4. The typical color is dull citrine (of Bidgway's
Color Standards, pi. 16) indistinctly streaked with olive-
citrine and olive lake ; the suture having an ocher, ochraceous-
orange or rufous border, usually edged below with pale yel-
low. The embryonic whorls are very faint yellow fading to
white above the suture and at the apex. The suture has a
well-impressed margin. The aperture has a white lining and
a chocolate band inside the acute white edge. The columellar
fold is white and rather strong. Fig. 1 is a typical livida,
agreeing very closely with Swainson's figure. Length 16,
diam. 10 mm. ; 5% whorls.

In different specimens the hue and shade vary. The gen-
eral tint may be brownish vinaceous with narrow whitish
streaks, or it may be yellowish olive or dark greenish olive.
The dark submargin of the lip is inconspicuous in some shells,
especially old ones with the lip thickened.

Form reevei C. B. Adams, pi. 53, figs. 5 to 9, Kawailoa,
Gulick coll.) differs by having no rufous sutural line. The
suture is bordered below by a nearly white or pale greenish
yellow band, varying in width. Its exact status is not clear
to me, since I do not know whether the specimens occurred in
colonies of livida or always separate. It seems however to
have had a wider range eastward than livida, as Gulick got it
in Kawailoa, Opaeula and Wahiawa. Both livida and reevei
seem to be rare or extinct at the present time. They probably
inhabited forests at lower levels than those now existing.

24a. A. LIVIDA EMERSONII Newcomb. PI. 53, figs. 10, 11, 12.

Based upon a light form of livida, as Newcomb recognized
later, confirmed by Doctor Cooke who examined the type.
The original figure is copied, pi. 53, fig. 10. The description
follows. * * Shell conical, polished, dextral, rather solid ; whorls

248 ACHATINELLA LIVIDA.

6, rounded, margined above ; suture distinct ; aperture sub-
ovate ; columella short, tuber culate ; lip acute, within mar-
gined, black ; aperture internally pure white ; exterior of shell
uniform pearly white. Length 11, width 8 twentieths inch.
District of Waialua. This shell in its general aspect is the
complete counterpart of A. mighelsiana of Pfeiffer. The lat-
ter species is from Molokai. This is clearly distinct, as may
be seen by a cursory examination of the two species' (Newc.}.

Evidently an albino form. Most shells of this kind are not
pure white. They are white with narrow streaks of very
faint flesh color or very pale gray, often with a faint yellow
suffusion on the base. Last whorl somewhat flattened later-
ally, convex below the impressed subsutural line. Aperture
white or pale lilac, usually with a dark streak within the
margin.

It varies to a form with yellow base ; often there is a rufous
subsutural line. It is somewhat intermediate between livida
and undulata, but has the shape of livida. There is a good
series from Wahiawa in coll. Dr. C. M. Cooke (pi. 53, fig. 11)
and others from Waialua, from Baldwin (pi. 53, fig. 12).
Named in honor of Mr. J. S. Emerson of Honolulu, who be-
gan collecting in the time of Newcomb. The name was ori-
ginally spelled " emmersonii' by Newcomb, but in his list of
1858 it was corrected to emersonii, and placed in the synonymy
of livida. The "subspecies" of livida must not be taken too
seriously !

24fe. A. LIVIDA RECTA Newcomb. PL 30, figs. 45 ; pi. 53, figs.
13 to 21.

' ' Shell usually dextral, solid, pyramidal ; whorls 6, but
slightly rounded, margined above ; suture slightly impressed.
Aperture subovate ; lip simple, thickened within ; columella
short, twisting to unite with the inner lip. Color various,
usually yellow, plain or with two black bands on the last
whorl, one of which is lost in the aperture, the other becomes
sutural. The shell above described is peculiar for its solidity
and rigid aspect. It possesses none of the graceful curves
which give to this genus so much of its beauty. The largest

ACHATINELLA LIVID A. 249

size yet discovered is given in the measurement. Length 14,
diam. 7 twentieths of an inch.' (Newc.}

Oahu: Waialua (Newcomb) ; Wahiawa (Cooke) ; Helemano
and Kawailoa ( Gulick, chiefly sinistral form ) .

Achatinella recta NEWC., P. Z. S. 1853, p. 145, pi. 23, f. 45,
1854. — PFR. Monogr. iv, 535. — GULICK, Evolution, Racial and
Habitudinal p. 41, pi. 2, f. 3 (Kawailoa). — Achatinella glauca
GULICK, Ann. Lye. N. H. vi, p. 60, pi. 8, f. 51, 1858.

Doctor Newcomb 's figure is reproduced on pi. 30, fig. 45.
A topotype from Newcomb, typical in pattern, is before me.
There is a broad purplish-brown band at the periphery, an-
other below it, and a very faint band in the middle of the
upper surface, cuticle yellow on the last whorl, lighter above.
Others of the same lot are bandless, or have a wide zone in
place of the two bands, pi. 53, figs. 13, 14, 15. The surface is
nearly smooth, under a lens showing light growth-lines and a
very faint oblique lineolation. The dark bands color the
internal lip-callus, which is thin and inconspicuous. The
columellar fold is rather small. As in A. livida, an ochra-
ceous band may usually be made out below the suture of the
embryonic whorls, but it is often very pale.

The locality "Nuuanu valley' given by Mr. Sykes is a
mistake. Some other form, perhaps multizonata, must have
been taken for recta.

Except in color, and in being partly dextral, recta does not
differ materially from livida. The classification of recta in
Bulimella with A. pulcherrima or A. byronii by various au-
thors is not easy to account for, as the lip is quite unlike those
forms. A majority of the specimens before me are sinistral,
but dextral shells are in the lots from Waialua, Kawailoa and
Wahiawa.

Kawailoa. In a lot of 68 taken by Gulick, three are dextral.
The ground-color varies from straw yellow to citron yellow.
The bands vary from dark chestnut to honey yellow or paler.
Usually there are only two bands, one above, the other below
the periphery, but in a few there is a sutural band also.

Two bands present, light in 14, dark in 11 shells.

One band (ii) present, light in 32, dark in 7 shells.

250 ACHATINELLA LIVIDA.

Sutural band only, light in 0, dark in 1 shell.

No bands, 3 shells. See pi. 53, figs. 17 to 21.

Helemano. A specimen taken by Gulick has three honey
yellow bands.

Wahiawa. A series collected by Dr. Cooke has three black-
ish chestnut bands on a white or yellow ground, or white
above, yellow below the periphery, both dextral and sinistral
(pi. 53, fig. 16). Also the form with honey yellow bands,
all sinistral.

A. glauca Gulick, pi. 44, figs. 5, 6, 7, seems to me to be
merely a dark form of recta, hardly worth a name, differing
only in the olive lake ground-color. There must however
have been a considerable colony of similar shells as Gulick
secured a large set. The description follows. * ' Shell sinistral,
rarely dextral, imperforate, acuminately ovate, solid, shining,
striated, fawn or lead-colored with two black bands, one en-
tering the aperture ; apex somewhat acute, white ; spire conic
with slightly convex outlines; suture marginate, moderately
impressed ; whorls 6, convex ; columellar fold central, white
or pale pink, moderately developed ; aperture sinuately oval,
white within ; peristome thickened within ; with external mar-
gin unreflected, arcuate, acute, bordered with brown; colu-
mellar margin dilated, adnate ; parietal margin wanting.
Length 17, breadth 9 mm. ; length of body-whorl ll1/^ mm.
Average weight 3.9 grains.

"Kawailoa, on trees. J. T. G.

"Var. 6. With a third brown sutural line.

"Var. c. With but one black spiral band passing above the
suture.

"Var. d. With a white band encircling the base between
the two black bands ; very rare.

"This species bears the same relation to A. livida Swains,
that A. recta Newc. bears to A. casta Newc.' (Gulick}.

Fig. 5 represents Gulick 's type specimen, no. 51 Boston
Society. A large set from the same lot is no. 92,284 A. N. S. P
The shell differs from A. livida by having two nearly black
spiral bands and usually no distinct subsutural band — char-
acters in which it agrees with var. recta, from which it differs
only in the ground-color. The color of the suture is variable,
some shells having an inconspicuous dusky or yellowish
border below the suture. One specimen has a pair of wider,

ACHATINELLA LIVIDA. 251

contiguous chestnut bands and a narrower subsutural band
of the same color. The ground color is olive lake in the type,
but varies in tint. It is nearly wax yellow in the chestnut-
banded shell. The lip is often well thickened within and
shows dark spots at the ends of the bands. The embryonic
shell when unworn shows the characteristic yellow zone of
livida below the suture, the rest of the whorl being white.
This zone changes to brown on the following neanic whorls,
but as stated above, fades to yellowish or disappears on the
last whorl.

24c. A. LIVIDA HERBACEA Gulick. PL 44, figs. 1, 2, 3, 4.

* Shell sinistral, sometimes dextral, imperf orate, ovate conic,
solid, shining, striated, of a dull green color; apex rather
obtuse, white ; spire conic, suture marginate, moderately im-
pressed; whorls 6, convex; columellar fold central, white,
strong ; aperture truncately auriform, white within ; peristome
slightly thickened within • with external margin unreflected,
arcuate, acute ; columellar margin dilated, adnate ; parietal
margin very thin. Length 18, breadth 10%, length of body-
whorl 13 mm. Average weight 4.4 grains.

"Var. 6. With a black sutural band; columellar fold usu-
ally white, sometimes lilac.

"Var. c. Yellow, with smoother surface, approaching A.
recta Newc.

''About a fifth of my specimens are dextral' (Gulick).

Oahu : forests between the streams of Waimea and Kawailoa,
on the leaves of the pua, ahakea and ohawai (Lobelia gri-
mesiana). J. T. Gulick.

Achatinella herbacea GUL., Ann. Lye. N. H. of N. Y. vi, p.
233, pi. 8, f. 52, Feb., 1858; Evolution, Racial and Habitu-
dinal, p. 41, pi. 2, f. 4.

A darker, greener shell than recta, also more strongly stri-
ate, and very rarely having any bands except the sutural,
which is occasionally present. Mr. Gulick 's type, pi. 44, fig.
1, is indistinctly streaked with citrine to olive-green on a
yellower, pyrite yellow, ground, the narrow, well-defined su-
tural margin somewhat tinted with chestnut; three apical

252 ACHATINELLA CURTA.

whorls white with a pale ochraceous zone below the suture.
While very glossy, it is more roughly striate than livida or
recta.

Others of the original lot, (pi. 44, figs. 2, 3, 4, coll. by Gulick) ,
are darker or lighter than the type — medal bronze or oil green
to amber yellow of Ridgway's Color Standards. The sutural
border may be colored like the rest of the whorl, or there may
be a blackish chestnut band; but there is no light zone or
band below the suture. One specimen has a pair of chestnut
bands near the periphery.

While close to recta, it may be practicable to distinguish this
form as a local race. I do not know that it has been found
since Gulick 's time.

25. A. CURTA Newcomb. Plates 45 and 46.

"Shell conical, sinistral, polished; whorls 5, rounded, mar-
gined above, the last very ventricose ; aperture ovate ; lip
simple, slightly thickened within ; columella short, with an
abrupt callous termination ; suture but little impressed ; color
yellow or chestnut, plain or with a black sutural band, rarely
with two or more on the last two whorls; columella white or
light brown. The rounded whorls and obese appearance of
this shell are strikingly characteristic. It is a rare species
and extremely limited in its range. Length 12, diam. 8 twen-
tieths of an inch' (Newc.).

Oahu: Waialua (Newcomb). Ahonui to Kawailoa, and
across the range in Laie.

Achatinella curta NEWCOMB, P. Z. S. 1853, p. 144, pi. 23,
f . 43 ; Ann. Lye. N. H. of N. Y. vi, 329.— PPR., Monogr. iv,
540; vi, 176. — Achatinella rhodoraphe E. A. SMITH, P. Z. S.
1873, p. 74, pi. 9, f. 10. — GULICK, Evolution, Racial and
Habitudinal, pi. 41, pi. 2, f. 6 (Wahiawa). — Achatinella un-
dulata NEWCOMB, Proc. Boston Soc. N. H. v, p. 218, Sept.,
1855; American Journal of Conch, ii, p. 216, pi. 13, f. 15. —
PPR., P. Z. S. 1855, p. 208 (Feb. 5, 1856) ; Monogr. iv, 540.—
Achatinella delta GULICK, Ann. Lye. vi, p. 231, pi. 8, f. 50,
Feb., 1858; Evolution, Racial and Habitudinal, p. 41, pi. 2,
f. 5 (Wahiawa).

ACHATINELLA CURTA. 253

Typically distinct from livida and recta by the shorter spire
and the short, swollen, last whorl ; also differing in 'the color-
patterns ; yet there are forms which one cannot satisfactorily
classify. PI. 45, fig. 1 is a copy of the type figure.

There are three color-forms which have been described as
species under the names curta (yellow with a chestnut or
chestnut-black subsutural band), rhodoraphe (yellow with a
pale pink subsutural band), and undulata (pale ochraceous
buff writh many irregular tawny stripes and sometimes a black-
ish peripheral band). There are also a number of other pat-
terns as noted below. The curta and rhodoraphe patterns
often occur in the same colony, and curta colonies sometimes
contain both rhodoraphe and undulata if I may judge by
Wahaiwa series in the Cooke collection. ChieflV rhodoraphe
occurs in Ahonui (Gulick coll.), at the eastern limit of the
species, and pure curta has been taken in Laie, unaccom-
panied by other patterns. There are some ' ' blends ' between
these three patterns, but as a general rule, good segregation
prevails in hybrid colonies.

Northwestward curta is replaced by dimorpha, in which the
shell is usually more lengthened.

Ahonui (pi. 45, figs. 2, 3, coll. by Gulick). Most of the
specimens are yellow with a pink zone below the suture
(rhodoraphe). The pink zone begins on the last embryonic
whorl as an ochraceous band. The largest shell is 17 mm.
long. There are also a few shorter shells of typical curta
form, yellow fading to white at the summit, without a sub-
sutural band.

Wahiawa (pi. 45, figs. 10, 11, 3d gulch of Wahiawa, coll. by
Kuhns). Last whorl or two apricot yellow, with a chestnut-
black band below the suture, edged below with orange rufous,
sometimes followed by whitish. Embryo all white or with a
brown hand extending to the apex. The same curta pattern is
in the Cooke collection from low ridges and valleys of Wahi-
awa, together with specimens without sutural band, and others
with it faint and narrow, the embryonic whorls with an ochra-
ceous band, the body-color varying from yellow to ochraceous
tawny. Also rhodoraphe like those of Helemano, or without

254 ACHATINELLA CURTA.

a pink band (like A. delta var. &. of Gulick), and various
forms of the undulata pattern (pi. 45, figs. 4 to 9), from the
pattern figured by Newcomb to forms with the streaks almost
or entirely wanting, the band remaining. A last vestige of
the undulata pattern is usually visible in a minute irregu-
larity or mottling of the dark peripheral band in specimens
which have otherwise the color of curia. There are also
tawny-streaked forms without a band, the typical undulata
pattern.

Helemano. In a small ravine south of Helemano, Mr.
Spalding found curta, with chestnut sutural band and plain
(no. 2274-5 of his coll.). His no. 2268 from Helemano have
a yellow base, white above. Gulick found the same form,
which he called A. delta var. b. (pi. 46, figs. 6, 7, 8). Gulick
also obtained a large series of typical rhodoraphe in Helemano,
which he regarded as the metropolis of this form (pi. 46,
figs. 1, 2, no. 92210 A. N. S.), associated with specimens like
pi. 45, fig. 3. In a gulch west of Helemano Mr. Spalding
found curta with black and with pink sutural border, and
with a light brown subsutural line, in the same colony (pi. 46,
figs. 3, 4).

' l Waialua ' ' is the type locality of curta and undulata, but
just where Newcomb collected in the extensive district which
went under his name, I do not know. Perfectly typical curta
has been taken by Mr. Thaanum on the left side of Poamoho,
with specimens without the chestnut sutural band (pi. 45, figs.
12, 13). Mr. D. D. Baldwin obtained typical undulata and a
pale form perhaps referable to emersoni in "Waialua," local-
ity not more exactly given (pi. 45, figs. 14, 15, 16).

In the bottom of gulch east of Opaeula along the stream,
Mr. Spalding took curta with the sutural band: (a) blackish-
chestnut, (6) varying to tawny, approaching rhodoraphe, and
(c) with brownish vinaceous streaks on a cream-buff ground,
suture with a narrow tawny band (pi. 46, fig. 9), also vary-
ing to nearly white. Some have more or less olive-yellow suf-
fusion of the base or last whorl. This form approaches
undulata. There are also white and yellow curta without
sutural band in this colony, which is related to the streaked
form from Kawaiholona.

ACHATINELLA CURTA. 255

Opaeula. Gulick found rhodoraphe, varying from a wide
to a linear sutural border.

On the Opaeula-Kawaiholona division ridge Mr. Spalding
found curia with a black or purplish sutural band, and a few
white ones with yellow tint on at least part of the base.

Kawaiholona. On the eastern spurs Mr. Spalding found
faintly streaked curta-undulata forms similar to pi. 46, fig. 9.
Along the bed of the stream a form was found with the char-
acters of the preceding intensified, intermediate between curia
and livida (pi. 46, fig. 10, coll. by Spalding). It has a rufous
or chestnut sutural line, followed by a white band, below
which it is streaked with Dutch blue or slate blue on a whitish
tint of the same, the base with a yellowish suffusion. There
is a slate-purple streak within the white lip-edge, and the
columellar fold is white. While the coloration of this colony
is much like livida, it connects with curia through forms found
eastward, as noted above. A brown form also occurs in the
same colony (no. 1612 of Mr. Spalding's collection).

"Kawailoa.' Mr. Gulick found curia, rhodoraphe and
undulata patterns, the latter small, and varying to specimens
in which the tawny bands are diffused, producing an ochra-
ceous-tawny shell with indistinct dusky streaks (pi. 46, figs.
20 to 24). PL 46, figs. 16-19 are Kawailoa forms from
Thaanum.

Kawaiiki. Above and at the Waialua Agricultural Com-
pany's intake, Mr. Spalding collected a series of beautiful
color-forms, illustrated in pi. 46, figs. 11 to 15. The follow-
ing color-forms occur, (a) Yellow, varying to white with the
base faint yellow, apical whorls often ochraceous. (6) yellow
with a pale tawny sutural line or a chestnut or blackish sutural
band, typical curia pattern, (c) yellow with two or three
chestnut-black bands, which may cover most of the surface.
(d) yellow, with diffuse chestnut streaks and sutural band,
or with deep chestnut streaks and sutural and peripheral
bands, approaching undulata. (e) Pale yellow with diffuse
serpentine green streaks and a chestnut sutural band, a dark
streak within the outer lip ; approaching livida.

In the collection of Hon. L. A. Thurston there is a good

256 ACHATINELLA CURTA.

series of curia in several color-forms, which he took in the
bottom of Anahulu valley. This is as far west as real curia
has been found, so far as I know.

Laie. A series of 15 specimens, all of the typical curia
pattern, was taken by Mr. Spalding. One is figured on pi.
46, fig. 5. This is the only record of curia from the north
side of the main range, unless we include Gulick's A. con-
tracta, which is not really distinguishable from some
Wahiawa undulata in which the streaks are nearly or quite
obsolete. On account of its locality I have left coniracia
with the dimorpha group of forms.

The original descriptions of undulata and rhodoraphe
follow.

. .

'Achatinella undulata. Shell sinistral or dextral, rather
solid, acutely conical, shining, polished; with longitudinal
oblique fine striae; microscopically spirally striate. Whorls
6, rounded and margined above ; suture well impressed. Aper-
ture subovate ; columella short and plicately twisted ; lip
acute, thickened within. Color light olive alternating with
slightly undulating chestnut lines and bands, rarely marked
by transverse black fasciae. Columella and aperture white.
Length 12, width 6 twentieths of an inch. Waialua, Oahu'
(Newc.).

"Achatinella rhodoraphe Sm. Shell sinistral, shortly ovate-
conic, imperf orate, glossy, striated with growth lines and (un-
der a lens) very fine transverse striae; yellow, encircled below
the suture with a wide zone of pale rose. Whorls 6, convex,
the first three white ; suture distinctly margined. Aperture
white ; margin of the peristome acute, bordered within ; colu-
mellar fold strong, rose (sometimes white). Length 15, diam.
8y2. Var. : Shell yellow, suture zoned with chestnut below.

Station : on trees. The metropolis of the species is Hele-
mano, on Oahu. It is also found in Ahonui, Wahiawa,
Opaiula and Kawailoa. It is related to A. livida Swn. and
A. curia Nwc., but is easily distinguished from either. This
species is always sinistral' (Smith).

A. curia delta Gulick. PI. 45, figs. 17 to 22. ''Shell sinis-
tral, imperforate, conic, obliquely truncated at the base, solid,
shining, striated, yellow at the base, paler above, with 2 or 3
ash-brown bands ; apex rather obtuse, white ; spire conic ;
suture marginate, lightly impressed ; whorls 5%, slightly con-

ACHATINELLA CURTA. 257

vex; columellar fold central, white, strong; aperture trun-
cately auriform, white within; peristome thickened within;
with external margin unreflected, arcuate, acute ; columellar
margin dilated, adnate, white ; parietal margin wanting.
Length 16, breadth 10%, length of body-whorl 11% mm.
Average weight 4.7 grains.

' Var. &, without bands. Var. c, with one black spiral line.
Var. d, with two broad black bands.' (Gulick) .

Oahu : Kalaikoa, Ahonui, Wahiawa and Helemano, on trees
and shrubs. J. T. Gulick.

"In Wahiawa, which is the metropolis of the species, var.
a is most abundant; var. b is more widely diffused, being
occasionally found in each of the above-mentioned localities ;
vars. c and d are very rare ; the former approaches A. con-
tract a Nob., and the latter passes into an unusual variety of
A. emersonii Newc.' (Gulick}.

A. delta is a connecting form between reevei and curta,
comparable to the streaked form of curta from Kawaiholona,
and probably not more deserving of a varietal name than that.

Mr. Gulick 's type of delta is a Wahiawa shell closely agree-
ing with pi. 45, figs. 20-22, from the same lot, of which over 60
specimens are before me, No. 92,619 A. N. S. The prevailing
shape is that of A. curta, but it varies to longer shells, like
livida. The color-forms figured on pi. 45, figs. 17-22 are from
one colony.

(a) The ground color of the typical form is white above,
on the last whorl shading from white at the suture to pinard
yellow on the base, the yellow area often much reduced. The
sutural margin is white, followed by a band of violet plum-
beous, grayish lavender or ecru-drab obliquely streaked with
white. Above and below the periphery there are similar
bands. Sometimes the lower band is wanting, and the others
reduced to lines; and rarely the dark color of the bands ex-
tends in streaks over the whole banded portion, much as in
reevei, or in the rare variety of curta from Kawaiholona.
(&) The other common color-form has a ground of yellow,
fading above the periphery to white, or rarely continuous
over the whole last whorl. It has no dark bands, or rarely

258 ACHATINELLA DIMORPHA.

bands of a faint chamois tint. This form occurs with the
typical form (a) in Wahiawa, but without it in Ahonui, Hele-
mano and Kalaikoa. It is hardly to be distinguished from
bandless curt a, and approaches very close to rhodoraphe.

The fact seems to be that the unicolored form of delta (b)
is a mutation of the curta-rhodoraphe stock which has rather
a wide distribution, and somewhere in Wahiawa district it
formed a hybrid colony with the streaked livida-curta stock;
pattern (a) of the heterogeneous lot called delta being
the result. Very likely the type colony of delta was limited
to a small area and is now extinct.

26. A. DIMORPHA Gulick. PL 42, figs. 15 to 20 ; pi. 47.

" Shell sinistral, sometimes dextral, imperf orate, turreted,
solid, shining, striated, white or yellow with a brown sutural
band [see below for bandless and for two- or three-banded
patterns] ; apex rather obtuse ; spire turreted ; suture mar-
ginate, moderately impressed, dark brown ; whorls 6, convex,
columellar fold central, white or rose, moderately developed;
aperture truncately ellipsoidal, white within ; peristome
slightly thickened within, with external margin unrenected,
arcuate, acute ; columellar margin dilated, adnate ; parietal
margin waiting. Length 18, breadth 9%, length of body-
whorl 11% mm. Average weight 4 grains. Habitat, Waimea,
Pupukea, Waialei and Kahuku, Oahu, J. T. G. ! Kaawa,
Oahu, J. S. Emerson' (Gulick).

Oahu: northwestern ridges from Waimea to Kahuku;
Hauula and Kaaawa on the north side.

Achatinella dimorpha GULICK, Ann. Lye. Nat. Hist, of
N. Y. vi, p. 236, pi. 8, f. 56. — Achatinella albescens GULICK,
t. c., p. 237, pi. 8, f. 57; Evolution, Racial and Habitudinal,
p. 41, pi. 2, f. 2 (Pupukea). — Achatinella zonata GULICK, t. c.,
p. 238, pi. 8, f. 58; Evolution, etc., pi. 2, f. 1 (Pupukea). -
Achatinella contracta GULICK, t. c., p. 239, pi. 8, f. 59. Feb.,
1858.

A. dimorpha differs from A. curt a chiefly by its more
graceful lengthened contour, the last whorl being longer, less
inflated, and more slowly tapering below, whereas curta is

ACHATINELLA DIMORPHA. 259

more squat, the last whorl shorter and more obese. Yet the
shapes intergrade, dimorpha colonies sometimes containing
shells exactly like some of the longer specimens of curia. The
gamut of color-patterns is in part different in the two species.
While the distribution of curta and dimorpha does not over-
lap, as known at present, there is a long boundary still to be
exactly investigated where connecting colonies may confidently
be expected. The difficulty of delimiting the curta and
dimorpha territories is therefore likely to increase with en-
larging knowledge. It may turn out that my attempt to
separate curta and dimorpha is impracticable in some colonies.
Yet here, as in most tree snails, there has been differentiation
between forms of the interior slope and those of the Pacific
slope west and north.

Three mutations have been named as species: dimorpha,
yellow, fading to white near the suture, which has a narrow
chestnut or blackish band. Zonata, ground-color the same or
nearly white, with two blackish bands, at and below the per-
iphery, and often a sutural band also. Albescens, base yellow,
fading upwards, or whole shell nearly white ; no bands. The
zonata and albescens forms ordinarily, perhaps always, live in
mixed or hybrid colonies; albescens and dimorpha patterns
have also been found together. I do not know that the
dimorpha form is found living with both of the others, but
it probably is. Mr. Gulick, who obtained these forms when
they were abundant, assorted his shells and preserved no
indication of their association.

The areas of distribution of zonata and albescens as given
by Gulick coincide exactly. He got both in Waimea, Pupukea,
Waialea, Kahuku, Hauula and Kaaawa. Dimorpha was ob-
tained in the same places except Hauula; but he got few
shells in that valley. While in the following account I use
these names for convenience in referring to particular pat-
terns, it will be understood that they are not of the nature
of subspecies or geographic races, but merely mutations exist-
ing in hybrid colonies. While sinistral shells are the rule,
there are occasional dextral shells. In Waimea and Pupukea
the dextral form is rather abundant.

260 ACHATINELLA DIMORPHA.

The dimorpha forms were collected in abundance by Gulick
and J. S. Emerson in the Fifties. In the last few years they
have been taken by Messrs. Spalding, Kuhns and Wilder.

Waimea. Mr. Gulick obtained shells of zonata, albescens
and dimorpha patterns. Some of the albescens have two-
thirds of the last whorl yellow, with a white line at the per-
iphery, elsewhere white. Others are typical, the yellow fad-
ing upwards to white which forms a zone below the suture.

Pupukea. The type specimen of dimorpha, no. 56 Boston
Soc. coll., from Pupukea, is figured, pi. 47, fig. 1. The last
whorl is amber yellow, fading upwards to nearly white near
the narrow chestnut sutural band. This band ascends to the
summit, but on the embryonic whorls it widens and becomes
ochraceous. Surface glossy ; aperture and columellar fold
white, the latter small. The Gulick collection also contains a
good series from Pupukea, comprising (a) the three-banded
typical zonata pattern, both sinistral and dextral; (6) shells
with white ground or faintly yellow, fading upwards, with a
tawny sutural band, the embryonic whorls white or with an
ochraceous band, columellar fold white; rarely dextral (fig.
2), and in one shell the bands of zonata are faintly indicated,
(c) albescens pattern, the last whorl straw yellow, deeper near
the lip, fading upwards, no bands. Probably all of these
forms were from one colony.

Mr. Spalding 's No. 2187, from the Pupukea side of the
Paumalu ridge contains exactly typical dimorpha, together
with shorter shells indistinguishable from curta, the last two
whorls yellow, band chesntut ; also a form resembling pattern
b of Gulick 's lot, having the shell white, sutural band dark
vinaceous, not extending upon the white embryo, the colu-
mellar fold lavender. These three patterns are shown in pi.
47, figs. 3, 4, 5.

In the third gulch towards Kahuku from Pupukea Mr.
Kuhns collected beautiful 3-banded zonata with albescens
and a pure white form (pi. 47, figs. 6, 7, 8).

Paumalu-Kaunala ridge. A series taken by Mr. Spalding
comprises very beautiful zonata and albescens forms, the latter
white to yellow, often with a white peripheral line. Some of

ACHATINELLA DIMORPHA. 261

the zonata have the bands very broad — an unusual variation
(pi. 47, figs. 9 to 13). A lot taken by Kuhns has the same
albescens forms and the dimorpha pattern — short, curta-like
specimens.

Waialee. PL 42, figs. 15, 16; pi. 47, fig. 14. Known to
me from abundant series taken by Mr. Gulick. The albescens
pattern, in white and varying tints of yellow, often with a
peripheral white line; columella rarely lilac-tinted. Zonata
pattern, with white or yellow ground. Also a dimorpha pat-
tern, the ground pale yellow or white, sutural line tawny,
often very faint, columellar fold white or flesh-tinted. It is
similar to pattern b from Pupukea.

Kahuku. Gulick reports zonata, dimorpha and albescens.
Some of the zonata have lines and oblique streaks between the
bands, slightly recalling formosa (pi. 47, fig. 15). Very beau-
tiful pale albescens have been collected by Kuhns, Thaanum
collection (pi. 47, fig. 16), and Mr. Spalding has taken zonata
in Pahipahialua valley, to the west of Kahuku.

From Kahuku going southeast there are no records of
dimorpha forms until we reach Kaipapau, and there is another
hiatus between Hauula and Kaaawa, which is the limit of the
species in this direction, so far as we know. The locality
"above Ewa" given by Mr. Sykes for zonata must belong to
some other species, perhaps cast a.

Kaipapau. On the lower ridges, just above the kukui tree
belt Mr. Spalding took the zonata (pi. 47, fig. 17) and al-
bescens patterns.

Hauula. The zonata and albescens forms, taken by Gulick,
are before me.

Kaaawa. Mr. Gulick records dimorpha, albescens, zonata
and contracta from this valley, collected by Mr. J. S. Emerson.
The abundant series of the zonata pattern varies widely, the
following forms being represented.

Typical zonata, with two blackish bands and a narrower
sutural band (pi. 42, fig. 20). Varying to forms with fine
lines above the lower band, columella lilac-tinted, or when
there is a fourth narrow band around the columella, the fold
is purplish-brown (pi. 47, fig. 18).

262 ACHATINELLA DIMORPHA.

Sutural band wanting, the other two reduced to lines,
either of which is occasionally doubled (pi. 42, fig. 18).

Only one band, the lower (band iii) remaining (pi. 42,
%. 17).

Only the upper band (ii) remaining (pi. 42, fig. 19). Of
this pattern there is only one specimen, all of the others being
represented by numerous shells.

A. contract a Gulick (pi. 47, figs. 19, 20) is in my opinion
only a form of the Kaaawa zonata, from which it differs by the
shorter contour. Very few specimens were taken. In some
of them there are indistinct gray streaks in the ground, which
is yellow below, white above the supraperipheral deep chest-
nut band. This shell reminds one of a similar form of un-
dulata. Possibly there has been some infusion of undulata
blood from across the range. No recent collector has to my
knowledge found contracta or any of the dimorpha forms in
Kaaawa, and they are probably extinct in the places where
Mr. Emerson collected fifty or sixty years ago.

The original descriptions of albescens, zonata and contracta
follow.

Achatinella albescens. " Shell sinistral, sometimes dextral,
imperforate, ovate-conic, solid, shining, striated, white or
sometimes yellowish ; apex somewhat acute ; spire convexly
conical ; suture marginate, well impressed ; whorls 6 ; some-
what swollen beneath the suture and slightly flattened in the
middle ; columellar fold central, white, strong ; aperture trun-
cately auriform, white within ; peristome thickened within,
with external margin unreflected, slightly arcuate ; columellar
margin dilated, adnate, parietal margin wanting. Length 18,
breadth 10%, length of body-whorl 13 mm. Average weight
4.6 grains. On the leaves of the pua, ahakea and lama.
Waimea, Pupukea, Waialei, Kahuku and Hauula, J. T. G. !
Kaawa J. S. Emerson ! Nearly a third of the specimens from
Waimea and Pupukea are dextral, but in Waialei, the metro-
polis of the species, they are always sinistral.' (Gulick).

Achatinella zonata. ' ' Shell sinistral, sometimes dextral, im-
perforate, ovate-conic, solid, shining, striated, white or yellow-
ish with a brown sutural band and two black bands, one en-
tering the aperture ; apex somewhat acute ; spire conical, with
outlines slightly convex ; suture marginate, moderately im-
pressed; whorls 6, convex; the last regularly rounded; colu-

ACHATINELLA C^ESIA. 263

mellar fold central, rose or white, moderately 'developed;
aperture semiorbicular, white within ; peristome slightly thick-
ened within ; with external margin unreflected, arcuate, acute ;
columellar margin dilated, adnate; parietal margin wanting.
Length 16y2, breadth 10, length of body-whorls 12% mm.
Average weight 4.6 grains. On the leaves of trees. Waimea,
Pupukea, Waialei, Kahuku and Hauula J. T. G. ! Kaawa, J.
S. Emerson ! The dextral specimens are for the most part
found in Waimea and Pupukea. This and the two preceding
species [albescens and dimorpha] vary much in form and
size' (Gulick).

Achatinella contracta. PL 47, fig. 19, copy of original fig-
ure. "Shell sinistral, imperf orate, broadly conic, solid, shin-
ing, striated, ash or fawn colored with two black bands, one
sutural, the other revolving just above the suture on the spire
and encircling the base near the periphery of the body-whorl ;
apex somewhat acute, white with brown suture ; spire regu-
larly conic ; suture marginate, moderately impressed ; whorls
5%, slightly convex, the last large; columellar fold central,
white, strong; aperture truncately auriform, white within;
peristome thickened within, with external margin unreflected,
arcuate, subacute ; columellar margin dilated, adnate ; parietal
margin wanting. Length IS1/^ breadth 9, length of body-
whorl 10 mm. Average weight 4 grains. On trees. Kaawa,
J. S. Emerson ! Hauula, J. T. G. ! The sutural band is
sometimes wanting, and some specimens have another band
entering the aperture. I have but one specimen from Hauula,
which is of the last mentioned variety, and is rather larger
than those from Kaawa. It is allied to A. zonata, nob., and
A. undulata Newc.' (Gulick).

27. A. ODSIA Gulick. PL 44, figs. 13 to 16.

"Shell sinistral, imperf orate, ovate-conic, solid, shining,
striated, so streaked with white and fawn brown as to have a
gray appearance ; apex somewhat acute ; spire convexly conic ;
suture marginate, moderately impressed ; whorls 6i/2, convex ;
columellar fold central, white, moderately developed; aper-
ture sinuately oval, white within ; peristome slightly thick-
ened within ; with external margin unreflected, arcuate, acute,
edged with brown ; columellar margin, dilated, adnate ; parie-
tal margin wanting. Length 181/-?, breadth 10%, length of
body- whorl 13 mm. Average weight 4 grains' (Gulick}.

"This with A. concidens and A. formosa nob., which are

264 ACHATINELLA C^ESIA.

found in Waimea, correspond to A. undulata Newc., A. emer-
soni Newc., and A. glauca nob., found in Kawailoa" (Gulick).

Oahu : Waimea, J. T. Gulick ; varieties at Kahuku, Kahana,
Hakipuu and Waikane.

Achatinella ccesia GULICK, Ann. Lye. N. H. of N. Y. vi, p.
234, no. 53, pi. 8, f. 53, Feb., 1858. — Achatinella concidens
GUL., t. c., p. 234, no. 54, pi. 8, f. 54. — Achatinella formosa
GUL., t. c., p. 235, no. 55, pi. 8, f. 55. — Achatinella cognata
GUL., t. c., p. 240, no. 60, pi. 8, f. 60. — Achatinella scitula
GUL., t. c., p. 241, no. 61, pi. 8, f. 61. — Achatinella cervina
GUL., t. c., p. 241, no. 62, pi. 8, f. 62.

A. ccesia was the first to be described of a series of shells
somewhat larger in size than those of the dimorpha series,
usually streaked in peculiar hues, and so far as I know, they
are always sinistral and never have an ochraceous zone below
the suture of the embryonic whorls. These shells were rare
in Gulick 's time ; only ccesia and cognata were known to him
by more than one or very few specimens. To-day they are
practically unknown to Hawaiian naturalists. I do not re-
member seeing any recently collected specimens, and if not
actually extinct at the present time the species must be very
scarce and local.

The distribution is conspicuously discontinuous — ccesia,
concidens and formosa in Waimea, cognata and scitula in
Hakipuu, and cervina in Kahana ; yet the presence of another
form of the series (littoralis) in the sand dunes of Kahuku
serves to connect the extremes of its range, and permits the
suggestion that it was probably a species of the low-lying
forests of former times. This species then probably had the
range of A. dimorpha-albescens-zonata, but chiefly at lower
elevations.

There seem to be several local races, although not so many
as Mr. Gulick defined. I would arrange them as follows :

A. ccesia Gulick, including concidens and formosa Gul.
Waimea.

A. c. littoralis P. & C. near Kahuku ; extinct.

A. c. cervina Gul. Kahana.

A. c. cognata Gul. Includes scitula Gul. Hakipuu and
Waikane.

ACHATINELLA C/ESIA. 265

Waimea. The type of A. ccesia is figured on pi, 44, fig. 13.
The type-specimen is not fully mature, therefore it is rather
thin, and reminds one of A. papyracea. It is indistinctly
streaked with light pinkish cinnamon on a white ground, with
a dusky, interrupted peripheral line below which there is a
pale line. Suture margined on the last whorl, of the same
color as the shell. The embryonic whorls have a faint yellow-
ish tint, the apex being white. Columellar fold white ; lip
thin, with a pinkish submargin within. The last whorl is
slightly compressed laterally, so that it has a somewhat cylin-
drical contour, though this is not conspicuous. There is a
very minute cleft behind the reflected columellar lip in the
type and in other immature shells, but in those fully mature
it is closed.

Another lot from Gulick, no. 1258 coll. Boston Society (pi.
44, figs. 14, 15, 16) contains several patterns, all with the
same slightly cylindric last whorl, but part of them more
solid than the type-specimen, being mature. Color as follows.

White, with the faintest yellow tint near the lip, which is
acute but slightly thickened within.

White with four brown lines on the last half of the last
whorl.

Yellow, with some obscure lines on the base, lip and colu-
mella flesh-tinted. Shell smaller, length 16^, diam. 9.1 mm.

The type and sole specimen of A. concidens (pi. 43, fig. 14)
is a "dead' shell which has lost its polish by weathering.
It is conspicuously streaked with cinnamon, with dashes here
and there of darker brown, on a soiled white ground; the
streaks interrupted by a white band below the suture, another
at and below the periphery, the lower half of the base being
also dirty whitish. Embryonic whorls cinnamon-buff, fading
to white at the apex. Length 19.7, diam. 11 mm. This shell
was no doubt paler, more gray in life. I think it merely a
color-form of ccesia, not a true race.

The type-specimen of A. formosa (pi. 43, fig. 13, No. 55
of type series in Boston Society coll.) is a very beautiful shell.
It is solid, more elongate then the type of cccsia, but other-
wise not dissimilar in contour. The shell is imperforate and

266 ACHATINELLA OESIA.

sinistral. The periphery is marked by a white band, with a
wide black band below, a narrow one above it. There is a
white band below the suture, bordered by an inconspicuous
yellowish line. The rest of the upper surface of the last whorl
has a light plumbago-gray hue, produced by darker and paler
streaks. The base has a blackish band around the columella,
and is elsewhere white. The embryonic whorls are white ;
following whorls of spire have a dark band above and below
the suture, which runs in a white band. Columellar fold
white and very strong; aperture white within, the acute,
beveled lip is colored at the terminations of bands ; somewhat
thickened. Length 19.5, diam. 10.5, aperture 9 mm. ; 6*4
whorls.

A. formosa is merely concidens with black bands added.
It seems remarkable that Mr. Gulick did not recognize in
ccesia, concidens and formosa merely three stages in the de-
velopment of pattern, strictly comparable to several other
well-known species which have the same sequence of patterns
—streaked, streaked and with white spiral bands, and the
same with dark bands.

Kahuku : in troughs of sand dunes near the sea, between
the road and the shore, about I1/*? miles east of Kahuku, pi.
44, figs. 17 to 20, collected by Cooke and Pilsbry. This fossil
form is a fairly well-marked subspecies which may be called
A. ccesia littoralis P. et C. It differs from cccsia and its
Waimea color-forms by the rougher surface, which is decidedly
more wrinkled along lines of growth, especially on the last
half of the last whorl, and the columellar fold is unusually
high on the columella, less prominent than in adult Waimea
shells. The shell is thin, minutely perforate, varying in shape
as figured, and marked with several broad or numerous narrow
dark zones (gray, or in places brick-red in the fossils, prob-
ably almost black when they were alive) ; all had a white
subperipheral band.

Length 20, diam. 10.6, aperture 9 mm. ; whorls 6y2.

Length 20, diam. 11, aperture 10 mm. ; whorls 6*4.

Length 19.8, diam. 11.6, aperture 10.3 mm. ; whorls 5%.

The deposit at Kahuku contains many Tornatellinidce and

ACHATINELLA OESIA. 267

other fragile snails as well as Amastra and various ground
shells. They must have lived where they are now found, their
preservation being due to the calcareous sand which drifted
over the forest-bed. Probably Achatinella has nowhere else
been found so near the sea — the shore being only a few rods
away, and the difference of level not over 10 or 12 feet. It is
quite likely however that there has been some subsidence of
the island since the forests extended so far down.

Kahana. A. cccsia cervina Gulick, of which the type is
figured, pi. 43, fig. 12, is a weakly characterized race of ccesia,
of which very few specimens were found. It is a thin shell
very similar to concidens, and with much the appearance of
A. buddii, as Sykes and others have noticed. It is cinnamon
colored with softly blending darker streaks and many fine
russet lines, which become distinct only on the last half of the
last whorl. The suture is narrowly bordered with white, and
the embryonic whorls are cinnamon-buff fading to whitish at
the apex. A small cream-white area surrounds the columella.
The lip is thin, not beveled or thickened within, and like the
whole interior is pale pinkish buff, nearly white. Columellar
fold central, thin but rather prominent. There is a very short
and narrow perforation. Length 19.4, diarn. 11, aperture 9.6
mm. ; whorls about 6. The smoother surface and central
columellar fold differentiate this from A. c. littoralis ; the shell
is thinner and more capacious than scitula or cognata.

Hakipuu. Mr. Gulick described A. cognata and A. scitula
from this valley. In my opinion the two belong to one race,
which would stand as A. ccesia cognata. The type-specimen
of cognata, pi. 44, fig. 8 (no. 60 of the Gulick type collection,
Boston Society), is a shell closely resembling A. dimorpha.
It is moderately solid, very smooth and glossy. The last whorl
has pale salmon colored streaks shading into the whitish
ground, but on the back of the last whorl the streaks give
way to a general sea-shell pink color (c/. Ridgway, Color
Standards, pi. xiv). The tint is paler towards the suture of
the last whorl, and the two whorls preceding are almost white.
The embryonic whorls are light buff with a white sutural line,
but on subsequent whorls there is a narrow chestnut sutural

268 ACHATINELLA CLESIA.

line. The lip has a delicate rib close to the edge, which is of
the same faintly pinkish white as the rest of the interior.
Columellar fold median, rather strong, and pale flesh color;
columellar margin reflxed and closely adnate. Length 18.3,
diam. 10.3, aperture 8.5 mm. ; 6*4 whorls.

Mr. Gulick recognized two varieties : ' ' Var. b, white ; var.
c, pale green.' All of the specimens have the embryo as
described above. The dark sutural line of subsequent whorls
is never continued upon it as an ocher band, as it frequently
is in A. dimorpha. In one specimen of the white form there
is no dark sutural line, others having it. Gulick 's "green'
variety is more properly olive-ocher, or tints between that and
primrose yellow, always fading to nearly white near the chest-
nut sutural line. Often the penultimate whorl is colored
thus, the color fading out on the last whorl. Specimens of
these forms are figured, pi. 44, figs. 9 to 12. All from the
Gulick series, No. 92,224 A. N. S. Another shell which prob-
ably came from Newcomb, has a distinct yellow band below
the white peripheral band, a fainter yellow band above it.
The pattern of this shell reminds one of A. decipiens.

I agree with Mr. Gulick that cognata is quite distinct from
dimorpha. It is also reported by him from Waikane, but I
have not seen these specimens. Both cognata and scitula are
imperforate.

The type-specimen of A. scitula Gulick, pi. 43, fig. 11 (No.
61 of Gulick 's type series, Boston Soc.) has narrow tawny and
ochraceous-tawny streaks on a Naples yellow ground, which
fades near the suture. There are very weak traces of spiral
dusky lines, a stronger one at the periphery, exactly as in the
type specimen of A. ccesia. The suture is narrowly edged
with brown. The embryonic whorls are cinnamon-buff, fad-
ing to whitish at the apex, and with an inconspicuous whitish
sutural line. The lip is thickened by a distinct narrow, white
rib close to the edge. Interior pink tinted. Columellar fold
rather weak, central and white. Length 20.2, diam. 11, aper-
ture 9.4 mm., 6!/s whorls.

Mr. Gulick 's statement that the suture and columella are
light colored in scitula is not borne out by the type-specimen,

ACHATINELLA C^SIA. 269

which has a distinct if narrow dark sutural margin. His
figure also shows this. A. scitula seems to be merely a color-
form of A. cccsia cognata.

The original descriptions of the forms herein referred to
A. ccesia follow.

. ,

'Achatinella concidens. Shell sinistral, imperforate, ovate-
conic, solid, striated, brown banded with white ; apex some-
what acute ; spire convexly conic ; suture marginate, white,
moderately impressed ; whorls 6y2, convex ; columellar fold
central, white, moderately developed; aperture truncately
oval, white within ; peristome thickened within, with external
margin unreflected, arcuate ; columellar margin dilated, ad-
nate ; parietal margin wanting. Length 20, breadth 11, length
of body-whorl 14 mm. Average weight 4.75 grains.

"Station, on trees. Habitat, Waimea, Oahu, J. T. G. ;
My specimens of this species are more or less bleached and
faded. Continued search was made for living specimens, but
without success' (Gulick).

"Achatinella formosa. Shell sinistral, imperforate, acu-
minately ovate, solid, shining, striated, white, with two black
bands, one entering the aperture, sometimes with the upper
part of the whorl of a pale slate color ; apex somewhat acute ;
spire convexly conic ; suture marginate, well impressed ;
whorls 6%, convex; columellar fold central, white, strong;
aperture truncately auriform, white within; peristome thick-
ened within ; with external margin unreflected, arcuate, acute ;
with columellar margin dilated, adnate ; parietal margin want-
ing. Length 201/3, breadth 10%, length of body-whorl 14
mm. Average weight 6 grains.

"Station, on the ki (Cordyline terminalis) and other leaves.
Habitat, Waimea, Oahu, J. T. G. !

"Var. &. with the two bands uniting in one broad black belt.

"Var. c, with numerous black spiral lines.

"A rare species, and of great interest on account of its
affinities, which connect it with species so different from each
other. A. glauca, delta, phceozona and zonata Nob. are cer-
tainly not very similar ; but this species seems to connect itself
directly with varieties of each of these species' (Gulick).

"Achatinella cognata. Shell sinistral, imperforate, ovate
conic, solid, shining, striated, of pale rosy fawn color, with
a brown sutural band ; apex subacute ; spire convexly conic ;
suture marginate, brown, lightly impressed ; whorls 6, slightly
convex; columellar fold central, rose-colored, strong; aper-
ture truncately auriform, within white lightly tinged with

270 ACHATINELLA C^ESIA.

rose ; peristome thickened within ; with external margin un-
reflected, arcuate, acute ; columellar margin dilated, adnate ;
parietal margin wanting. Length 19, breadth 10, length of
body- whorl 13 y2 mm. Average weight 5.6 grains.

''Station, 011 trees. Habitat, Hakipu, Oahu, J. T. G. !
Waikane, Frick.

"Var. &, white. Var. c, pale green. Remarks: allied to
A. dimorpha Nob.' (GuUck).

" Achatinella scitula. Shell sinistral, imperforate, elon-
gately ovate, solid, shining, striated, streaked with brown and
pale fawn ; apex subacute ; spire convexly elongate ; suture
marginate, moderately impressed; whorls 6%, convex; eolu-
mella with a moderately developed white fold near the body-
whorl ; aperture sinuately ellipsoidal, white within ; peristome
well thickened within ; with external margin unreflected,
slightly compressed in the middle, with the anterior edge arcu-
ate ; columellar margin dilated, adnate, parietal margin want-
ing. Length 21, breadth 10%, length of body-whorl 13%
mm. Average weight 4.8 grains.

"Station, on trees. Habitat, Hakipu, Oahu, J. T. G. !

' ' Remarks : There is a green variety which passes into var.
c. of the last described species. But in that species the
suture and columella are dark, while in this they are light-
colored.

"Achatinella cervina. Shell sinistral, scarcely perforate,
ovate conic, rather thin, shining, striated, fawn colored, with
obscure brown spiral lines ; apex subacute ; spire convexly
conic; suture obsoletely margined, moderately impressed;
whorls 6, convex ; columellar fold central, white, sublamelli-
form, well twisted ; aperture sinuately oval, white within,
peristome slightly thickened within ; with external margin un-
reflected, arcuate ; columellar margin dilated, adnate, parietal
margin wanting. Length 20, breadth 11%, length of body-
whorl 14 mm. Average weight 3.8 grains.

"Habitat, Kahana, Oahu, very rare, J. T. G' (GuUck).

Species of the Waianae range.

In the systematic classification these species are about
equally related to the series of A. papyracea and to that of
A. livida. All of them are extremely rare shells, each species
known from a single small colony only, — and by this we mean
a single clump of bananas or the like. A. lehuiensis and its var-
iety gulickiana have been found but once, and that over fifty
years ago, a single individual of each. A. thaanumi is known

ACHATINELLA SPALDINGI AND LEHUIENSIS. 271

by two specimens. Only of A. spaldingi was a. fair series
obtained. All of the known colonies are on the northern
slope of the range. Who knows how many more wait hidden
in dark and difficult ravines, or perhaps a few steps aside
from some well-trodden trail !

Probably all of the Waianae forms are descendants of a
single species, which migrated from the Koolau range, along
with Partulina dubia and the ancestor of Achatinella
mustelina.

28. A. SPALDINGI Pilsbry & Cooke, n. sp. PL 42, figs. 1, 2, 3.

The shell is sinistral, imperforate, ovate-conic, ventricose
and quite thin-, white, with slightly interrupted or spotted
tawny bands and lines, of which band ii, above the periphery,
is the most constant. There is usually a group of bands near
the columella, and a space without bands at and below the
periphery ; suture edged with a band or line of the white
ground ; apex a trifle dusky. Surface not very glossy, or
often dull in old shells, somewhat roughened by growth-
wrinkles and irregularly scattered impressions. Whorls con-
vex, joined by an impressed suture. Aperture white and
showing the bands weakly within ; outer lip not expanded,
thin, acute. Columellar fold whitish, spiral, small.

Length 16.5, diam. 11.1, aperture 9.2 mm., 5% whorls.

Length 17.2, diam. 11.1 mm.

Length 17.2, diam. 11 mm.

Oahu, Waianae range: Pukuloa, one-half mile above the
Mountain House, back of Leilehua (Spalding). Cotypes in
A. N. S. P. and Bishop Mus. ; also in coll. Irwin Spalding.

This species has the thin texture of A. papyracea, but it is
a much more capacious shell, differing in surface and color;
the spire is somewhat more attenuate near the apex. Quite
old specimens have a thickening within the lip, which is want-
ing in most adults. It is one of the few really distinct species
of Achatinella strum. Named for a valued friend of both
authors.

29. A. LEHUIENSIS E. A. Smith. PL 41, fig. 11.

" Shell sinistral, ovate-conic, somewhat glossy, very finely

272 ACHATINELLA LEHUIENSIS.

striated with growth and transverse lines; white, encircled
with a purple-brown streaked zone and two brown zones, one
above, the other below the periphery (and sometimes others).
Suture submarginate. Whorls 5%, convex. Aperture white ;
peristome thin; columellar fold strong, rose colored. Length
17, diam. 10 mm.' (Smith).

Oahu, Waianae range: Lehui [Lihue], on trees (Gulick).

Achatinella lehuiensis SM., P. Z. S. 1873, p. 76, pi. 9, f. 4.

The figured type, No. 74 Coll. Boston Society, is here il-
lustrated. It resembles A. dimorpha form zonata Gul. rather
closely in shape, but the last whorl is slightly more ample ;
it is, I believe, more closely related to A. papyracea. There
are two deep chocolate bands separated by a white girdle,
one immediately above, the other below the periphery. Ad-
joining the upper band above there is a zone composed of
narrow, brownish vinaceous streaks alternating with narrower
whitish ones, and traversed by darker lines near the upper
and lower edges. There is a cinnamon line below the white-
edged suture, and a small area of the same around the colu-
mella ; also a dark line near the lower edge of band iii. The
embryonic whorls are white, shading to gray at the tip. The
aperture has a white lining and a very slight thickening near
the edge, which is beveled and acute, in color light brown,
with dark markings where the bands terminate. The colu-
mellar fold is quite strong and white. I think the statement
"plica columellaris valida, rosea' was due to dirt and the
fact that it has a cinnamon border ; but it may possibly have
faded. The columellar margin is adnate.

The suture has a distinct margin defined by an impressed
line on the last whorl, but not colored. The shell is moder-
ately solid, about as in A. dimorpha. The unique type meas-
ures length 16.25, diam. 10 mm., longest axis of aperture
8.4 mm.

T have described the type-specimen in some detail because
the species has not to my knowledge been recovered by recent
collectors, and its existence in the Waianae range has been
doubted — I believe unnecessarily. It cannot be expected that
further specimens, if found, will agree with the type in all
•details of color-pattern.

ACHATINELLA THAAMUNI. 273

The locality "Lehui' (more properly Lihue) was used by
Gulick in rather a wide sense, I suppose covering much of the
country south of Popouwela, towards Palikea, on the eastern
slope of the range. As the form is probably very local, there
is hope that it may still be found.

While somewhat resembling A. dimorpha form zonata Gul.,
it seems to me to be distinct from that, and from A. papyracea.
It is certainly quite distinct from A. casta.

29a. A. LEHUIENSIS GULICKIANA ii. subsp. PL 42, fig. 4.

The shell is more conic than lehuiensis, in shape resembling
the shorter forms of A. dimorpha form zonata ; solid ; white,
with a narrow chestnut band just above the periphery, ap-
pearing above the suture, and a broad one occupying most of
the base ; a short distance below the suture an ochraceous band
revolves. Embryonic whorls white. Aperture and colu-
mellar fold white ; the outer lip not thickened, stained brown
at the terminations of the bands. Length 15.75, diam. 10,
length of aperture 8.1 mm. ; 5% whorls.

Oahu, Waianae range: Mokuleia, Gulick coll., No. 1471 coll.
Boston Society.

The type is a single specimen in the Gulick collection. If
it is really from Mokuleia, it may be a distinct species.
Moreover, I cannot connect it with any form of the Main
Range. It is not, in my opinion, to be referred to zonata
Gulick. It cannot be connected with A. thaanumi, which has
a more elliptical shape. In texture and weight A. gulickiana
agrees with lehuiensis. Except in the structure of the lip it
has some resemblance to A. fuscobasis.

We would not describe a subspecies from one specimen if it
were not important to direct attention to the existence of a
member of this group near the western end of the Waianae
range.

30. A. THAAMUNI P. & C., n. sp. PL 42, figs. 5, 6.

The shell is perforate, sinistral, ovate-conic, thin but mod-
erately strong; white, encircled by two chocolate bands, one
above, the other below the periphery; no subsutural band or

274 ACHATINELLA.

columellar dark patch. Embryonic whorls white, becoming
blue-gray at the tip. The surface is glossy, marked with fine
growth-lines and nearly obsolete spiral stride. Spire conic
with nearly straight outlines and minute apex. Suture very
narrowly margined in some, not in other specimens. The
aperture has a white lining, showing the bands faintly, but
at the thin edge they become vivid, the acute peristome being
elsewhere white. It is slightly thickened within. Columellar
fold strong and white.

Length 19.1, diam. 11.6, aperture 10.3 mm. ; fully 6 whorls.

Length 17.6, diam. 11, aperture 9.25 mm.

Oahu, Waianae range : a gulch of Mt. Kaala running into
Haleauau gulch, on banana. D. Thaanum.

This fine species is closely related to A. lehuiensis, but differs
by being more capacious with larger last whorl and aperture,
and somewhat different coloration, lacking subsutural and
columellar bands. It may eventually prove to be a subspecies
of lehuiensis if colonies intermediate in characters are found,
but with present knowledge a union of the two is not war-
ranted. A. thaanumi is a thicker, smoother shell than
A. spaldingi, with narrower aperture and somewhat different
coloration. It is one of the rarest Oahuan shells, only in the
collection of Mr. Thaanum, two specimens, one a dead but
fresh shell.

Section ACHATINELLA s. str.

Achatinella SWAINSON, Quarterly Journal of Sci. and Arts
iii, p. 83, l82S.—Apex von Martens, Die Heliceen, 1860, p. 248,
"type Achatinella lugu~bris Chemn.' -Helicter (Fer.) PEASE,
Proc. Zool. Soc. London 1862, p. 3; 1869, p. 645.—Helicterella
GULICK, Journal of the Linnean Society of London. Zoology,
xi, p. 497, 1873 (name only; no species mentioned).

Shell ovate-conic or piriform, imperforate or minutely per-
forate, solid. Embryonic whorls nearly flat, the later ones
more convex. Aperture quite oblique, the lip very little if at
all expanded, well thickened within ; columellar fold strongly
developed. Type A. apexfulva.

Distribution, both ranges of Oahu.

ACHATINELLA. 275

The forms of this group which have been described as species
number not less than 53, besides several names for varieties.
Doctor Hartman (1888) reduced them to 15; Mr. Baldwin, in
his Catalogue of 1893, admitted 32; and Mr. Sykes, 1900,
enumerates 17 species with 13 varieties. The lists of
synonyms given by these authors differ very widely among
themselves, and still more from the synonymy proposed here-
in. My collaborator Doctor Cooke had worked out an ar-
rangement of all the forms under eight specific heads before
I took the group up. Using Cooke 's work as a basis, later
studies confirm nearly all of his results, but it appears neces-
sary to add A. swiftii and A. leucorraphe to the list of species,
with consequent rearrangement of the synonymy of other
forms. With some hesitation, specific rank has also been al-
lowed to the Waianaean A. concavospira, making 11 species
in all.

It was hoped that characters might be found in the repro-
ductive organs which would aid to indicate specific boundaries,
but dissections of A. lorata, vittata simulans and mustelina
show no tangible structural differences.

A consistent arrangement of the species in linear order is
impossible, as the group is formed of two parallel series which
merge together in the less specialized median species of each.

The minor series consists of apexfulva, turgida and lorata,.
species in which the apex is never black or dusky. These
forms are confined to the Main range, but do not reach to
either end.

In the greater series the tip of the apex is invariably dark
in some species (cestus, vittata, leucorraphe), and is variable,
either dusky or light, in others. The species are distributed
over the whole length of both the main and the Waianae
ranges.

An alternative and probably better grouping may be sug-
gested. (1) Series of lorata, for A. lorata. (2) Series of
apexfulva, for A. apexfulva, turgida, swiftii, leucorraphe,
vittata, cestus. (3) Series of decora, for A. valida, decora,
mustelina, concavospira. See diagram on page 278.

The distribution of the species, so far as positively known

276

ACHATINELLA.

to us, is shown in the accompanying table, in which the valleys
of the Main Range are given in order from the west eastward,
a few omitted. The ranges given under each species include
its varieties and synonyms as understood by the authors.

It is remarkable that with the exception of A. lorata, no
species of this group has been found on the northern side of
the Main Range.

Valleys.

Lorata.

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277

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278 ACHATINELLA LORATA.

The systematic and to some extent the geographic relation-
ships of the species may be approximately represented thus :

valida
mustelina — decora

apexfulva
concavospira

swiftii — turgida

leucorraphe

vittata lorata

i

cestus

In the text the species are arranged in the sequence cus-
tomary in Hawaiian collections — from the east westward.

31. A. LORATA Ferussac. PL 51 ; pi. 52, figs. 1-7.

Shell dextral, ovate-conic, the summit acute, glossy, striate,
white with an epidermal color, epidermis uniform or orna-
mented with bands ; whorls S1/^, regularly increasing ; suture
margined ; aperture ovate, white ; columella arcuate, promi-
nent; umbilical cleft not distinct. Inhabits the Sandwich
Islands (Ferussac).

Oahu: Manoa (and across the range in Maunawili) to
Moanalua ; varieties further west ; the typical forms especially
characteristic of the ridges between Nuuanu and Manoa.
Usual station on leaves, especially terminal bunches, on guava
and many other plants.

Helix lorata FER. (Prodrome p. 56, nude name), Voyage
auteur du Monde de 1'Uranie et la Physicienne, Zoologie, p.
479, pi. 68, figs. 8-12, 1824. — Achatina lorata Fer., DESHAYES,
Hist. Nat. Moll, ii, p. 193, pi. 155, f. 9-11—Achatinella lorata
Fer., NEWCOMB, Ann. Lye. vi, p. 310 (animal). — PPR., Mono-
graphia iv, 524; vi, 168. — THWING, Orig. Descript. p. 13, pi.
1, f. 2. — SEMPER, Eeisen im Archip. Phil., Land Moll., pi. 6,
f. 23 (genitalia). — Achatinella hanleyana PFR., P. Z. S. 1855,

ACHATINELLA LORATA. 279

p. 202, species no. 2; Monographia iv, 529. — Sykes, Fauua
Hawaiiensis p. 302. — Achatinella ventrosa PFR., P. Z. S. 1855,
p. 6, pi. 30, f. 20; Monogr. iv, 535. — Achatinella nobilis PFR.,
P. Z. S. 1855, p. 202, species no. 1 ; Monogr. iv, 524.

Achatinella alba Nuttall in Jay's Catalogue, edit. 3, p. 58,
1839 (name only). — Achatinella pallida Nuttall, 1. c., and in
REEVE, Conch. Icon, vi, pi. 1, f. 2, May, 1850. — PFR. Monogr.
iv, 532.

The straightly conic spire, the chalky or porcelain white-
ness of the ground, and the tawny and blackish markings make
lorata quite easily recognized. Its distinctness is rather a
matter of feeling, for the shape and color vary so much that
a strictly differential diagnosis could hardly be framed. It
has the air of a plebeian in a company of aristocrats.

The colors are rather crudely laid on, while most other
species of "Apex' are clothed in well blended nuances of
brown, ashen or slate, or have clearly drawn lines and bands.

A. lorata is a common shell. In its area one is likely to
find it wherever tree-shells occur. The complex of ridges com-
posing the mountain walls between Nuuanu and Manoa valleys
are the chief habitat of typical A. lorata, though it spreads
around the heads of the both valleys, as noted below.

Ferussac's figures represent several color-forms, all occur-
ring in Nuuanu. I take his figs. 10, 11 to be the typical pat-
tern. This is represented by pi. 51, figs. 9, 11, from the
Nuuanu-Pauoa ridge. His fig. 12 is like the form shown in
pi. 51, fig. 14, from the floor of the north side of Nuuanu, the
same pattern also extending up the north side. Ferussac's
other figures (8, 9), were probably from a dead and decorti-
cated specimen of the streaked pattern. The patterns vary
from streaked to banded.

1. Yellow (baryta yellow to yellow ocher) with olive, tawny,
chestnut or black streaks, pi. 52, figs, la, 4, 5. There is often a
blackish band or patch at the base. Short compact shells with
blackish streaks are Pfeiffer's A. ventrosa.

2. Same streaked pattern, but cut by spiral white bands,
which may be wider than the colored intervals, pi. 52, figs. 1,
3&, 6. A. pallida belongs to this group, also A. hanleyana Pfr.

280 ACHATINELLA LORATA.

3. Streaked pattern in large part lost, with the deciduous
cuticle, leaving a white shell, colored on the parietal wall.
Sometimes with a basal band or patch of chocolate, in the
prismatic layer of shell.

4. Cuticle white at all stages of growth; distinguishable
from blanched forms of no. 3 by the white parietal wall.

All of these patterns of cuticle are subject to alteration by
partial or total loss of the thin colored layer, which is usually
more or less deciduous. Any of them may have a dark basal
band or patch in the under layer of shell, which is unaffected
by loss of cuticle. Patterns 1, 2 and 3 intergrade freely, and
usually two, three or all of them may be found in one colony
on Tantalus, where it is still rather common, living in bunches
of leaves. The same patterns occur in Makiki, pi. 52, figs. 2-3c.

Manoa. Mr. Spalding collected a small lot (no. 96 of his
collection), in eastern Manoa (I suppose on the division ridge
near the main ridge) some years ago, — the most eastern local-
ity known. The shells are plain white or have traces of two
yellow bands, columella mostly pink.

Mount Tantalus (pi. 52, figs. 1-1& ; pi. 51, figs. 4-8), and
Makiki valley (pi. 52, figs. 2-3c). The shell varies a good
deal in size and shape, as shown in the figures. The lip and
columella are usually a brownish flesh tint (vinaceous pink,
Japan rose etc. of Ridgway) ; rarely white.

A. ventrosa Pfr. and A. pallida (Nutt.) Reeve are merely
color-forms occurring in mixed colonies together with other
patterns, from Tantalus to Nuuanu. They have no racial
status. No doubt the types of pallida as figured by Reeve
were from Tantalus. A. ventrosa also has the look of a Tan-
talus shell, though much the same pattern goes as far west as
Moanalua. The original description follows :

11 A. ventrosa Pfr. [pi. 30, fig. 20, photographic reproduction
of Pfeiffer's figure]. Shell imperf orate, ovate-conic, rather
solid, striate, white, covered downwards with a tawny, black-
streaked epidermis ; spire conic, obtuse ; whorls 5%, inflated,
the last a little shorter than the spire, base rounded ; aperture
nearly diagonal, broadly auriform, white within; columellar
fold thick, tooth-like ; peristome lipped within, the right mar-

ACHATINELLA LORATA. 281

gin unexpanded, rather straightly descending, slightly curved
forward, columellar margin short, adnate. Length 17%, diam.
11 mm. ; aperture 9% nim. long, 5% wide in the middle.
Sandwich Islands, Newcomb (Pfr.).

The type is in Pfeiffer's collection.

''A. hanleyana Pfr. Shell subrimate, dextral, globose-conic,
solid, smooth (under a lens very finely spirally striate) glossy;
fulvous, radiated with chestnut ; spire conic, the apex minute,
corneous ; suture margined with a creiiulated thread ; whorls
6, the upper flat, following moderately convex, the last nearly
as long as the spire, rounded ; aperture oblique, truncate auri-
form, white within ; columellar fold high, strong, tooth-like ;
peristome lipped within; right margin narrowly reflected,
columellar margin dilated, callous, subadnate. Length 18,
diam. 11 mm. ; aperture with peristorne 10 mm. long, 5 wide
within. Mus. Cuming, Sandwich Islands, Frick' (Pfr.).

Mr. Sykes remarks that this is "related to the form of
A. lor at a described as A. nobilis, and may prove to be an ex-
treme variety.' Dr. C. Montague Cooke, on examining the
type in the British Museum considered it an artificially colored
lorata.

Pauoa has the same forms as Tantalus, with the addition of
a quite elongate form, which comes from the Pauoa-Nuuanu
ridge.

Nuuanu. The shell is smaller in the average than on Tan-
talus, and the lip and columella are paler, almost or quite
white in most shells. The color-patterns are the same as in
Tantalus-Makiki shells, but white with one or two dark
bands is the usual pattern. PL 51, figs. 9, 10, 11 represent
shells from the south (east) side of Nuuanu, 011 lehua trees;
pi. 51, figs. 12, and pi. 52, fig. 4 are shells from the north
(west) slope, all collected by Dr. Cooke. Figs. 9, 11, 12 are
the less common color-patterns. Further Nuuanu shells, from
the Gulick collection, are figured, pi. 51, figs. 19-21. Four
shells from one tree on the floor of Nuuanu valley above the
central crater are shown in pi. 51, figs. 13-16, collected by
Dr. Cooke. The next colony westward is that of the summit
of the Nuuanu-Kalihi ridge, which differs from forms found

282 ACHATINELLA LORATA.

east and west of it sufficiently to be separated as a race,
A. I. nobilis.

Kalihi. PL 52, figs. 5, 5a, and pi. 51, figs. 17, 18, coll. by
Gulick. The short ventrosa and longer pallida forms pre-
dominate. There are also pure white forms. The lip and
columella are white or nearly so. The same streaked or white
forms were taken by Mr. Sp aiding on the central ridge of
Kahauiki.

Length 18, diam. 11.8, aperture 10 mm. (ventrosa pattern).

Length 20.4, diam. 11.3, aperture 10 mm. (pallida pattern).

On the east side of Kalihi Mr. Spalding found the shells
all small, length 15 to 17 mm. White, uniform or with a
streaked band, or chestnut-streaked on a yellow ground below
a white belt.

In Mr. W. D. Wilder 's collection there are huge lorata from
Kalihi — up to 25 mm. long.

Moanalua. A large series in coll. C. M. Cooke shows the
short ventrosa form exactly as in Kalihi ; the same with colored
pattern remaining only in bands; and pure white, length 17
to 21 mm. These are from high lateral ridges, pi. 52, figs. 6.

On the floor of the valley near the head on widely scattered
niu, Dr. Cooke found a small and usually more slender form,
length 15 to 19 mm., white, uniform or with a dark peripheral
band, sometimes a second one below it (pi. 50, figs. 7, 8). No
ventrosa occurred here. As in Kalihi shells, the aperture is
entirely white.

There is also a series of this small race in the Thurston col-
lection, from the bottom of the north fork of Moanalua, 850
to 1300 ft. I suppose from the same colony where Dr. Cooke
collected.

Halawa. A small series coll. by Gulick shows patterns sub-
stantially as in the ridges of Moanalua.

Kalauao-Waimalu ridge. Mr. Spalding collected here speci-
mens of unusual color, Vandyke brown with lighter streaks,
some of them much smaller than the one figured, pi. 52, fig. 7.

Waimano. A series from the Thaanum collection consists
of streaked shells like pi. 52, fig. 5, the cuticle largely lost, and
a white form; all solid shells of stout contour (pi. 50, fig. 6,
Waimano).

ACHATINELLA LOR ATA. 283

A. lorata form melanogama P. & C. PL 51, figs, 1-3 ; pi. 52,
figs. 8, 8a. The shell is somewhat more slender than in Tan-
talus lorata, with about 6% whorls; mutations in the hybrid
colony as follows. PL 52, figs. 8, 8a, black, uniform or with
indistinct mahogany-red streaks, sutural border and embry-
onic whorls white or buff ; lip black-edged within.

PL 51, figs. 1, 2, white, the lower half of the last whorl
pale yellow with ocher or ochraceous tawny streaks, sometimes
one or two spiral lines; embryo white; peristome vinaceous
pink. Sometimes the colored cuticle remains only on the
parietal wall.

PL 51, fig. 3, white throughout except for the vinaceous pink
peristome.

Length 19.6, diam. 10, aperture 9 mm.

Length 18.2, diam. 11, aperture 9.5 mm.

Maunawili, on the north side of the main range opposite
Maiioa valley — the Kailua flank of Mt. Olympus. Collected
by Messrs. D. B. Kuhns and W. D. Wilder. Cotypes 108767
A. N. S. and in Wilder coll.

This is one of the incipient races distinguished by contain-
ing certain peculiar color-forms, in a hybrid colony also hav-
ing normal patterns. Such forms as this are not subspecies
in the proper sense, but it is convenient to have names for
what seem to be elementary patterns, even though they are
not extricated from the parent race.

The black form is a mutation which occurred in a colony
of rather lengthened white-ground lorata. In the hybrid
colony resulting there is complete segregation of the color-
forms, in a series of over one hundred individuals seen. Em-
bryos from a typical black mother are figured, pi. 50, figs. 16,
16a. They vary from cinnamon to white. I do not know that
dark embryos occur in light individuals ; all of the latter now
before me have white apical whorls.

31a. A. LORATA NOBILIS Pfr. PL 50, figs. 1 to 5.

On top of the Nuuanu-Kalihi division ridge, at 2000 feet
elevation more or less, A. lorata is replaced by a race or sub-
species which differs by its longer shell of 6^/2 to 7 whorls

284 ACHATINELLA LORATA.

(lorata having 5y2 to 6). The peristome is usually darker
than in lorata of Nuuanu and Kalihi, buff-pink to brownish
vinaceous ; columellar fold the same or nearly white. Patterns
various :

PI. 50, fig. 5, white.

Fig. 2, white, the base finely yellow-streaked or merely
tinted, or streaked with tawny or Dresden brown.

Fig. 3, 4, last two whorls yellow, usually with a white band
above.

Any of these patterns may be varied by a few chestnut
lines or bands on the lower part, as in figs. 2, 3.

Length 22.5, diam. 12.5, aperture 10.5 mm.

Length 23.5, diam. 12, aperture 10.5 mm.

Length 21, diam. 10.5, aperture 10 mm.

The figures are from specimens taken on and around Wao-
lani Peak, where it is rather abundant, from the head of
Waolani valley up. This is probably the type locality. The
original description follows.

< i

A. nobilis Pfr. Shell subimperf orate, dextral, solid, stri-
atulate, glossy, fulvous or grayish-green with darker streaks ;
spire exactly conic, apex white, acute ; suture lightly mar-
gined ; whorls 6% to 7, rather flat, the last about two-fifths the
total length, obsoletely subangular below the middle ; aperture
oblique, obauriform, white within ; peristorne a little expand-
ing, the right margin broadly lipped within, columellar mar-
gin subadnate. Length 23, diam. 11 mm. ; aperture lO1/^ mm.
long, 5 wide. Island of Oahu, Frick' (Pfr.). Type no. 22
of Pfeiffer's collection.

31&. A. LORATA PULCHELLA Pfeiffer. PI. 30, fig. 2; pi. 50,
figs. 9 to 14.

'Shell subimperforate, dextral, ovate-conic, nearly smooth,
glossy, fulvous with blackish-green bands and sometimes a
median band of white ; spire conic, attenuate and white to-
wards the acute apex ; suture thread-margined ; whorls S1/^, a
little convex, the last about three sevenths the total length,
rounded ; aperture nearly diagonal, subtetragonal-auriform ;
columellar fold high, tooth-like, strong; peristome strongly
lipped within, the right margin unexpanded, a little straight-

ACHATINELLA LORATA. 285

ened, columellar margin reflexed, subadnate. Length
diam. 10 mm. ; aperture 8 mm. long, 4 wide inside. Mus.
Cuming, Sandwich Islands, Frick" (Pfr.).

Waimano (C. M. Cooke) ; eastern ravines of Waiawa
(Irwin Spalding) ; Halawa to Waipio (Thwing). Mountains
behind Ewa, Perkins (Sykes).

Achatinella pulchella PFR., P. Z. S. 1855, p. 6, pi. 30, f . 2 ;
Monographia iv, 536.

I have never seen shells agreeing exactly with Pfeiffer's
figure, reproduced in pi. 30, fig. 2, but some from Waiawa are
perhaps as near as one ought to expect among such variable
forms. It is the shell commonly known as pulchella. It
differs from the ventrosa form of lorata as found in Moanalua,
etc. chiefly by the presence of a darker band below the suture,
the more brilliant gloss, and the short spire ; but it must be
admitted that if it were not already named, we would hardly
have thought the race worth a distinctive title.

Shells from the eastern ravines of Waiawa, pi. 50, figs. 9, 13,
14 from Mr. Spalding 's no. 255 and 1899, have several
patterns :

PL 50, fig. 14. Last whorl empire yellow with chestnut
streaks, mostly ill-defined, darker next the suture ; penulti-
mate whorl white with chestnut-spotted sutural border.
Length 17, diam. 10.8, aperture 9 mm. ; 6 whorls. Length
15.4, diam. 10.4, aperture 8.6 mm., 6 whorls.

PI. 50, fig. 13. The same except that the streaks are cut by
white bands.

PL 50, fig. 11. The same, with two nearly black bands un-
der the cuticle. Waimano (Dr. Cooke). The same form is
in the Waiawa lot.

PL 50, fig. 9. White, with a chestnut-streaked band below
the suture.

PL 50, fig. 10. White with greenish (yellowish citrine)
bands. Waimano (Dr. Cooke).

PL 50, fig. 12. Last whorl streaked with light ochraceous
salmon. Waimano (Dr. Cooke).

The specimens in coll. Cooke, reported from the north ridge
of Waimano valley, figs. 10-12, are exactly like those from

286 ACHATINELLA CESTUS.

Waiawa, and I suspect that there is an error in the locality.
Mr. Thwing gives pulchella an eastern range as far as Halawa.
In this direction the separation from lorata would become
increasingly difficult.

32. A. CESTUS Newcomb. PL 29, fig. 8 ; pi. 52, figs. 12-14a ;
pi. 55, fig. 1.

" Shell solid, ventricose, sinistral or dextral, pointed at the
summit; whorls 6, rounded, corded above, last one tumid;
aperture subovate ; columella short, strongly tuberculate ; lip
slightly expanded, thickened within. Color of tip black,
second and third whorls white, three last white, yellowish or
black or mixed, with a white cincture traversing the sutures
and cutting the body whorl below the center, with or without
a broader band below, sometimes with blotches or tessellations
of black and white or longitudinal undulating lines of the
same colors. Columella chestnut; lip same color, interrupted
with white. Length 14, width 8 twentieths of an inch.

"A fine shell, approaching A. similans of Reeve, which it
resembles in form, but strikingly differs in markings'
(Newcomb).

Oahu : Palolo (Newcomb) . Western ridge of Palolo to Niu.

Achatinella cestus NEWCOMB, P. Z. S. 1853, p. 132, pi. 22.
f. 8; Monographia iv, p. 529, as var. of simulans. — SYKES,
Fauna Hawaiiensis p. 300 (in part). — THWING. Original
Descript. etc., p. 13, pi. 1, f. 1.

Achatinella forbesiana PFR., P. Z. S. 1855, p. 5, pi. 30, f. 16 ;
Monographia iv, 529. — THWING, Original Descript. etc., p. 20,
not pi. 1, f. 5. — B[ulimella] forbsiana Pfr., HARTMAN, Proc.
A. N. S. P., 1888, p. 29.

A. cestus was ill-received by European writers. Pfeiffer con-
sidered it a variety of A. simulans. Sykes unites with it, as
varieties or synomyms, some eight described forms from the
Ewa region westward. Hawaiian students — Newcomb, Gulick
and the modern collectors — hold cestus distinct from all of
these. We fully agree with the latter view. The range of
cestus is widely separated from the similarly marked western
forms. There is also a hiatus between the areas of cestus and

ACHATINELLA CESTUS. 287

simulans, wherein no related form occurs. Both by the char-
acters of the shell and by geographic range, cestus seems to
be somewhat isolated.

Towards the eastern end of the range the shell becomes
darker, with various modifications of pattern. It is these
most remote eastern forms which look like shells of the western
mountains. See under A. siviftii of the polymorpha pattern.

Palolo. — A. cestus was given a wide range of color and pat-
tern in Dr. Newcomb's description, and his figure, reproduced
in pi. 29, fig. 8, represents one of the very rare patterns, by
no means a fair representative of the species. This pattern
was indeed included by him as exceptional, — ' ' sometimes with
blotches' etc. The normal or predominant coloration of
Dr. Newcomb's collection is shown in pi. 52, figs. 12, 12a,
representing two specimens received from him.

The ground-color is cinnamon or a tint of that, obliquely
marked with streaks (often crenulate or broken into mottling)
of chestnut or chocolate, interrupted by a white band or line
below the periphery, another often wider, at the base, and fre-
quently a band below the suture. Embryonic whorls are
typically white with a small terminal comma or vortex of deep
livid brown or dark purplish gray; but often the embryo is
more or less stained with some tint of ocher or olivaceous.
The moderate lip-callus is tinted with flesh-pink. Length
15.5, diam. 11 mm. Length 17, diam. 10.5 mm.

Mr. Gulick collected similar specimens in Palolo. Three
unusual color-forms are figured, pi. 52, figs. 14, 14a, and pi.
55, fig. 1. A large majority of the shells are sinistral. Mr.
Wilder got a few cestus on the Palolo-Manoa ridge, which
seems to be its extreme range in this direction.

A mutation of cestus very distinct in appearance was taken
by Newcomb and Gulick in Palolo, pi. 55, figs. 2, 3, 4, coll. by
Gulick. The chestnut coloring is reduced to lines and bands
at the periphery and on the base. The shell resembles A. simu-
lans, but is less robust, the spire more slender, and the colu-
mellar margin less developed. Most of those collected by New-
comb and Gulick are sinistral. A small lot, no. 42 coll. Irwin
Spalding, consists of dextral shells, which occurred with the

288 ACHATINELLA CESTUS.

typical color-form of cestus. Whether the Gulick lot of this
mutation was also in a mixed or hybrid colony is not known.
Although this form is not known to occur as a pure race, it
may be convenient for reference to designate it as color-form
or mutation simulator. It is interesting as showing the evolu-
tion of a banded form derived from a streaked pattern.
Further instances of such transformation will be found under
various western species. Form simulator has been found only
in Palolo, and must now be very rare.

In another form from Palolo, taken by Gulick, the streaks
are light brownish olive, and an indistinct, sutural band is
somewhat rusty. There are one or two light lines at the
periphery, and some dark bands around the columella.
These occurred with other specimens of more normal pattern.

Waialae. Gulick found typical cestus. Also a form which
he identified as A. forbesiana Pfr., I think correctly. The
streaks vary from russet to dusky drab, cut by white bands
and lines which may be few or very numerous (pi. 52, figs.
11, lla, 11& ). Lip brownish. The original description of A.
forbesiana will be found below.

In Waialae iki Mr. Spalding found somewhat similar shells,
dark cestus with a white belt, 3 sinistral specimens. On top
of the Waialae iki and Wailupe ridge Mr. Wilder obtained
specimens with two white basal bands.

Wailupe. Gulick obtained typical cestus, some with the
markings very weak, and also a multilineate form resembling
the forbesiana of Waialae (pi. 52, figs. 13, 13a). A fine series
from Mr. Thaanum, pi. 52, figs. 9, 9a> represents a pure colony
of forbesiana. The streaks are sepia, dusky drab or dark
plumbeous ; white spiral lines few or many ; interior light blue.
A series from the Wailupe-Niu ridge taken by Mr. Spalding
are probably from the same place. A few have a narrow
white band at the periphery. It varies to quite pale color
(pi. 52, fig. 10).

Niu. In a series in coll. C. M. Cooke the shells are similar
to the preceding, some browner, approaching the Waialae
color.

A. forbesiana is a limital southeastern race of cestus, and

ACHATINELLA VITTATA. 289

might be ranked as a subspecies were it not intimately as-
sociated with cestus in some Waialae and WaiVupe lots. A
division of these would scarcely be natural or practicable.
The original description follows.

Achatinella forbesiana Pfr. (pi. 30, fig. 16, reproduced from.
Pfeiffer). Shell dextral or sinistral, subperf orate, ovate-
conic, solid, closely striatulate, glossy, gray or whitish, painted
with close gray or brown bands, frequently confluent. Spire
exactly conic, white above, the tip of the apex black, rather
acute. Suture margined ; whorls 6, a little convex, the last a
little shorter than the spire, rounded. Aperture oblique, ob-
aurif orm ; columella fold high, strong, nodule-like ; peristome
bordered with brownish or flesh color, the outer margin nar-
rowly expanded, columellar margin thick, subadnate. Length
19, diam. 10.5, aperture 10 X 5 mm. Sandwich Islands, Frick,
in Cuming coll. (Pfr.).

33. A. VITTATA Reeve. PL 57, figs. 1 to 5&.

''Shell dextral, globosely conical, rather ventricose; whorls
broadly marginated round the upper part; columella callous,
scarcely toothed; lip rather thickened; white, encircled with
lines and fillets of pale brown, black at the apex. Sandwich
Islands, Mus. Cuming.

"This approaches so closely to the sinistral species A. decora
[=simulans] that it might readily be taken for a dextral
variety. The whorls are more rounded and more distinctly
margined beneath the sutures. The painting is very similar,
but mostly darker next the sutures' (Reeve).

Oahu : Eastern ridge of Nuuanu valley ; varieties westward
to Kalihi.

Achatinella vittata REEVE, Conch. Icon, vi, pi. 2, f. 9, April,
1850. — SYKES, Fauna Hawaiiensis, p. 304; with var. cinerea
Sykes, p. 305 (1900).— NEWCOMB, Ann. Lye. N. H. of N. Y.
vi, p. 315. — Helix decora Ferussac, in Freycinet's Voy.
Uranie et Physicienne, Zoologie, p. 478, 1824. Not H. decora
Fer. 1821. — Achatina decora Fer., DESHAYES, Histoire Nat.,
Moll., ii, pt. 2, p. 191, pi. 155, f. 5, 7. — Achatinella globosa
PFR., P. Z. S. 1855, p. 7, pi. 30, f . 25 ; Monogr. Hel. Viv. iv,
542.— Cf. Thwing, 1. c., p. 18.

290 ACHATINELLA VITTATA.

In the typical form of vittata the bands are cinnamon with
darker cinnamon brown or Vandyke brown spots, or they may
be more rufous, of a hazel hue, shading into chestnut-brown
around the columella. The spots on the bands are clearly
vestiges of an original streaked pattern, which has been inter-
rupted by spiral white bands and lines. It is a further evolu-
tion of the cestus pattern. The suture is almost always bor-
dered with a rufous band. The lip has a fleshy tint, and the
tip of the apex is invariably dark. The shell may be either
dextral or sinistral. This form was collected in Nuuanu (pi.
57, figs. 2 to 2c.) by Gulick, exact location not recorded.
Reeve's type is a dextral shell. His figure is reproduced,
pi. 57, fig. 1. Gulick 's shells are both dextral and sinistral.
Similar shells, but all sinistral, were taken by Mr. Thwiug
on the eastern ridge of Nuuanu (pi. 57, figs. 3, 3a, 36). This
lot is all sinistral and rather small, length about 18 mm.

A fine series was taken by Dr. C. M. Cooke on a spur of the
east ridge of Nuuanu opposite Luakaha (pi. 57, figs. 4, 4a,
5 to 56). All of the 33 specimens are sinistral. The bands
vary from chestnut to claret brown, and are very variable
in width, number and grouping, as the figures show. The size
and proportions vary widely, two shells measuring :

Length 18.3, diam. 11.3, aperture 9.25 mm.

Length 18.7, diam. 14, aperture 10.7 mm.

The typical form of vittata is probably almost extinct at
the present time. PL 57, figs. 4, 4a are globosa patterns.

A. vittata differs from cinerea and simulans by the lighter
color of the bands, which are spotted or streaked, the dark
sutural border, and the shape, which is less robust than simu-
lans. So far as I know it is not found on the western ridge
of Nuuanu. If this is the case the herds of vittata and simu-
lans are separated. The exact location of the colony or colo-
nies of vittata where Newcomb and Gulick collected is not
known to me, but from Newcomb 's statement that it ' is met
with in Nuuanu valley some three miles from Honolulu" and
"the inflated variety designated as A. globosa by Dr. Pfeiffer
is not uncommon,' ' we may gather that both came from lower
Nuuanu, doubtless on the Pauoa side or ridge. The shells

ACHATINELLA VITTATA. 291

taken by these collectors of sixty years ago are both dextral
and sinistral. Those taken by Mr. Thwing and Dr. Cooke
ten to twenty years ago, presumably from further up the val-
ley, are exclusively sinistral.

Judging from the short specimens approaching globosa
among A. vittata from Newcomb, and his remark upon the
form, I presume that this supposed species was based upon a
selected extreme form, probably from the vittata colony of
southeastern Nuuanu and is not in any proper sense a variety
or race. Mr. Sykes considered the type of globom to be
vittata, and Mr. Thwing has taken the same view. The origi-
nal description of A. globosa follows.

ilA. globosa Pfr. [pi. 30, fig. 25, photographic copy of ori-
ginal figure]. Shell subimperf orate, dextral, conic-globose,
rather thin, striatulate, white, encircled with brown lines;
spire short, a little convexly conic, subacute ; suture lightly
impressed, somewhat marginate ; whorls 5, moderately convex,
the last one obese, about as long as the spire, brown at the
base ; aperture oblique, subtetragonal-oval, pearly within ;
columellar fold weak, slightly twisted ; peristome acute, brown-
edged, somewhat white-lipped within ; columellar margin
thickened, somewhat adnate. Length 17, diam. 11% mm. ;
aperture 10 mm. long, 6 wide. Mus. Cuming. Sandwich
Islands, Frick' (Pfr.).

33a. A. vittata cinerea Sykes. PL 57, figs. 6, 6#, 6Z>, 7.

Banding almost black on the last whorl, ash colored on the
whorl above, the upper whorls tinted with pale-brown banding
above the suture, replaced by an almost black line at the apex.
Nuuanu, Perkins (Sykes).

In a series collected by Dr. Cooke (pi. 57, figs. 6 to 7) on a
few ridges of the eastern side of Nuuanu just above the dam,
the penult, and upper part of last whorl are tea green or
glaucous-gray, narrowly streaked with whitish, the last whorl
traversed by blackish-chestnut lines which increase and be-
come confluent near the lip ; a band bordering the suture and
usually a subperipheral band are white ; columellar margin
cinnamon brown. 28 specimens before me are all sinistral.
It occurs as a pure race, varying chiefly in the earlier or later

292 ACHATINELLA VITTATA.

appearance and spreading of the dark bands. Probably is
confined to the district mentioned above.

33 &. A. vittata simulans Reeve. PL 57, figs. 8 to 14c.

"Shell globosely conical, sinistral, sharp towards the apex;
whorls conspicuously margined at the sutures; columella
short, flatly callous; white, encircled with a few light-brown
lines, with an intense brown band around the columella ; lip

and columella tinged with light purple. Hab. ? Mus.

Cuming ' ( reeve).

Oahu : Nuuanu-Kalihi ridges, high up, especially around the
heads of Waolani and Kapalama valleys.

Achatinella simulans REEVE, Conch. Icon., vi, pi. 2, f. 15,
April, 1850. — PFR., Monographia iv, p. 528, exclusive of var. b.
— Achatinella decora Fer., REEVE, Conch. Icon., pi. 2, f . 12<z, &.
-THWING, Original Descript., etc., p. 16, pi. 1, f. 3. — Apex
albofasciata SMITH, P. Z. S., 1873, p. 78, pi. 9, f. 21. Cf.
THWING, Orig. Descriptions Achatinella, p. 17.

A. v. simulans differs from vittata and cinerea by its siiow-
white ground-color with almost black lines and bands, and the
usually more solid and robust shell. There is, however, varia-
tion in the color of the bands. The type specimen, pi. 57, fig.
8, reproduced from Reeve's plate, was a rather small shell
with few bands except close to the lip, where numerous chest-
nut-brown bands appear, as in pi. 57, fig. 9. Reeve's A. decora
is merely a larger, more copiously banded form of the same
race.

Ordinarily the bands and lines are almost black, various
in number and arrangement, rarely confluent, sometimes
wanting except at the extreme base. They do not spread or
increase in number behind the lip in the vast majority of
shells, the type of simulans and fig. 9 being quite exceptional.
There is no dark streak behind the lip, and the dark apical
mark is invariable. A tawny band below the suture is only
rarely developed. The spire has slightly concave outlines in
most specimens. The basal lip and part of the columella are
purplish lilac. Specimens from near and at the summit of the
ridge, above the head of Waolani are figured, pi. 57, figs. 10

ACHAT1NELLA VITTATA. 293

to 11, east slope of Waolani Peak, coll. by Pilsbry, and figs.
9, 9a coll. by Cooke. The same forms extend around the head
of Kapalama and onto the Kalihi ridge.

In the same colony from the west ridge of Nuuauu there
are exceptional forms with clove-brown, sepia, russet, chest-
nut or yellow ocher bands, pi. 57, figs. 140, 14c Cooke coll., or
chocolate, delicately streaked and spirally lineolate with white,
with a tawny sutural band as in A. vittata, pi. 57, fig. 14&
Cooke coll. All of the above are sinistral. Some of these un-
usual color-patterns of simulans most resemble vittata. One
of them also was selected to form the supposed species A. albo-
fasciata.

Dextral form. — On a few trees in a little ravine on the west-
ern ridge of Kapalama, and in the midst of the sinistral col-
ony, Mr. Spalding found a colony of dextral simulans. A
light ochraceous buff specimen is shown in pi. 57, fig. 12, but
dark-banded shells also were taken. Mr. Gulick found similar
ocher-banded dextral shells in ' ' Kalihi ' -possibly the same or
an adjacent locality.

Apex albofasciata Smith was based, as Mr. Thwing has al-
ready intimated, upon a tawny-banded individual of simu-
lans, such as are found on the west ridge of upper Nuuanu
with the more abundant typical form. The original figure is
photographically reproduced on pi. 57, fig. 13. The original
description follows.

"Apex albofasciata Sm. Shell sinistral (sometimes dex-
tral), perforate, globose-conic, lightly striated with growth-
lines ; of a bay color, streaked with darker ; the last whorl
encircled with two or three white zones (the median always
wide) ; apex white; whorls 6, the first four rather flat, the
rest convex, suture distinctly margined, white ; aperture red-
dish within ; peristome slightly dilated, thickened within, red-
dish, marked in the middle with the white band of the out-
side ; columellar fold strong, rosy, white at the apex. Length
19, diam. 12!/2 mm.

Habitat, Sandwich Islands. This species, of which there
are several specimens in the British Museum collection, some
sinistral and others dextral, I am unable to locate with any
yet described' (Smith}.

Mr. Smith informs me that "the apical half- whorl is tinted
with pale brown.'

294 ACHATINELLA TURGID A.

34. A. TURGIDA Newcomb. PL 29, figs. 10, 10a; pi. 56, figs. 1
to 4.

"Shell ventricose, pointed at the apex, sinistral or dextral,
smooth, polished, gradually enlarging for four turns and then
very rapidly, the last two forming most of the shell; suture
slightly impressed, beneath which revolves a linear depression.
Aperture subovate ; columella short, terminating in a strong
conical tubercle projecting into the aperture ; outer lip simple,
thickened within, slightly tinged with pink or brown ; tubercle
the same ; fauces white ; coloring extremely various, ground
white, yellow or black, with or without longitudinal zigzag
lines, transverse bands or blotches covering the surface.
Length fourteen-twentieth, diam. eleven-twentieth inch.'
(Neivcomb.)

Oahu: Ewa (Newcomb). Moanalua to Waiawa (or Wai-
pio, Thwing), the typical form westward, various varieties
eastward, within these limits.

Achatinella turgida NENCOMB, P. Z. S., 1853, p. 134, pi. 22,
f. 10, 10a. — PFR., Monogr., iv, 541. — THWING, Orig. Descript.
Achatinella, p. 18, pi. 1, f. 4 ("the metropolis of this shell is
Waipio "). — Achatinella cestus Newc., in part, SYKES, Fauna
Hawaiiensis, p. 300, pi. 11, f . 6, 7 ; not A. cestus Newc.

The attempt is here made to define western, median and
eastern races of A. turgida, as a step towards a systematic
record of its local color-differentiation ; but I am aware that
the material at hand is inadequate. All of the races may run
into melanistic phenotypes, by themselves indistinguishable.

The races are these :

A. turgida, Waiawa, pi. 56, figs. 1-2.

A. t. perplexa, Waimano, pi. 56, figs. 5-5/.

A. t. simulacrum, Waimauo-Manana ridge, pi. 56, figs. 13-
Ud.

A. t. cookei, Waimano-Manana ridge, pi. 57, figs. 15-17.

A. t. ovum, Moanalua to Waimano, pi. 56, figs. 6-10.

Typical A. TURGIDA, Waiawa. Doctor Newcomb 's figures
copied photographically in my pi. 29, figs. 10, 10a, and speci-
mens received from him, agree closely with shells taken in the

ACHATINELLA TURGIDA. 295

eastern ravines of Waiawa below the forest fence, no. 1861
coll. Spalding, some of which are illustrated in pi. 56, figs. 1
to Ic. It seems likely that the original lot came from this
part of Ewa district, and I propose that it be considered the
type locality. In one lot taken by Mr. Spalding there are
49 dextral, 38 sinistral shells. The first 2y2 whorls are ivory
yellow, the next (and last) embryonic whorl nearly or quite
white. The following neanic whorls almost always have a
nearly black band below the suture. The last whorl varies a
good deal in marking.

( 1 ) Ground yellow, mottled with brownish black, the mark-
ings sometimes arranged more in bands, sometimes more in
streaks (pi. 56, fig. 1&).

(2) Ground yellow below, white above the periphery; more
or less mottled, and usually with some blackish bands (New-
comb's figure, pi. 29, fig. 10a; also pi. 56, fig. Ic).

(3) Ground white, variously mottled and streaked or
banded with purplish black (Newcomb's figure, pi. 29, fig. 10;
also pi. 56, figs. 1, la, 2).

(4) All post-embryonic whorls black (pi. 56, fig. 4). This
pattern is rather rare.

All of these patterns except the last vary individually in
amount of marking in any lot, and there are also differences
between different lots in average tint. Thus, Mr. Spalding 's
lot no. 1898, from the forest fence colony, is as a whole darker
than his 1861 from the ravines below. One of Gulick's lots,
probably from lower down, is lighter than either (pi. 56, figs.
3, 3#). These differences are only noticeable in the mass.
Rarely two patterns of coloring may be. seen on different parts
of a single shell, as in the specimen figured by Mr. Sykes,
Fauna Hawaiiensis, pi. xi, figs. 6, 7.

The swollen shape of the last whorl in A. turgida has been
noted by Dr. Newcomb, but the contour varies rather widely.
The aperture is bluish white within, lip hardly expanded, well
thickened within, flesh color. Whorls 6 to 61/<J. Length 20.3,
diam. 13 mm. Length 18.2, diam. 14 mm.

A peculiar pattern, white with gray markings, was noted in
the Spalding collection, no. 581, from the central ridge of

296 ACHATINELLA TURGIDA.

Waiawa. As I have not the specimeus by me, I caiiiiot say
what relation they may bear to the gray form of Waimano.

According to Mr. W. D. Wilder, the yellow form with black
lip (var. ovum) occurs on the same ridge with mottled tur-
gida, but higher up.

34a. A. TURGIDA PERPLEXA n. subsp. PL 56, figs. 5 to 5/.

The shell has a white ground indistinctly streaked with pale
neutral gray, and encircled with numerous lines and bands of
darker gray or olive brown. First embryonic whorl cartridge-
buff with the tip either white or a little darkened, bluish or
fleshy, the rest of the embryonic whorls white. First neanic
whorl often marked with brown. The lip is vinaceous pink or
light brownish vinaceous, fading on the rather strong lip-rib.
About 6 per cent are sinistral.

Length 20.7, diam. 13.6 mm., 6% whorls.

Length 18, diam. 12 mm.

Lateral spurs (figs. 5-5cZ), and northern ridge (figs. 5e, /)
of Waimano valley, C. M. Cooke. Cotypes 1192 Cooke coll.
and 108802 A. N. S. P.

The grayish color and pale lip are the chief characters of
this race, of which there are about 130 specimens in Dr.
Cooke 's collection. It varies a good deal. The bluish-gray
streaks are often absent, or visible only on the bands, and the
latter vary from light to dark. The sutural margin is usually
touched faintly or distinctly with tawny. Figs. 5, 5a are
typical patterns.

In some specimens from the northern ridge of Waimano
(pi. 56, figs. 5e, 5/) the ground is yellowish or cream color,
not streaked, and the bands sharply defined, carob brown or
almost black. This pattern connects with the gray form
through some intermediate shells with gray bands and yel-
lowish ground, and there are also a few shells having gray
bands on a white ground, from the northern ridge.

This race seems to be intermediate between A. turgida and
A. swift ii. If found further west, it would inevitably be
merged in swiftii, on account of the grayish tone of the bands
and the perceptible duskiness of the tip in some specimens.

ACHATINELLA TURGIDA. 297

Yet other shells of the lot have the color of the apex and the
shape of the shell exactly as in A. turgida ovum. As the col-
ony is well within the turgida country, and some distance
from the area occupied by swiftii, it seems more natural to
rank the form under turgida.

When in Honolulu I noted that a copiously mottled form
also occurs in Waimano, no. 3609 and 2050 Spalding coll. In
the latter lot there are 42 dextral, 32 sinistral shells. I have
not these Waimano valley shells by me, but it may be that
they are another pattern of A. t. perplexa. Dr. Cooke's no.
1176, which I have figured, pi. 56, fig. 11, is probably a melan-
istic form of perplexa.

34&. A. TURGIDA OVUM Pfeiffer. PI. 56, figs. 6 to 12 ; pi. 59,
fig- 17.

"Shell imperf orate, sinistral. globose-conic, solid, rugulose-
striate, a little shining, whitish ; spire a little concavely conic,
apex acute ; suture deeply margined ; whorls 5%, the upper
flat, following convex, the last inflated, slightly shorter than
the spire ; columellar fold thick, tuberculif orm, pale lilac.
Aperture diagonal, sinuate-semicircular ; peristome unex-
panded, bordered with black-brown, with a crenulate lip
within. Length 19.5, diam. 13 mm. ; aperture 10.5 mm. long,
6 wide. Mus. Cuming. Habitat in the island of Oahu, very
rare, Newcomb' (P/r.).

Moanalua to the Waiau-Waimano ridge.

Achatinella ovum PFR., P. Z. S., 1856, p. 334; Monographia,
iv, 541. — ? THWING, Orig. Descript., etc., p. 19 ("Nuuanu").

The original description of A. ovum applies to white ex-
amples such as I note below from Halawa, etc. It is a rare
white form of a race which is usually some tint of yel-
low, and generally encircled with chestnut bands. There is a
black or nearly black streak behind the outer lip, and the
peristome is blackish brown. A blackish-brown or chestnut
band often borders the suture. Mr. E. A. Smith, who ex-
amined the type-specimen for me, writes that ' * the lip is black-
ish-brown within and without, thickened within, the thicken-
ing being slightly crenate, but this crenulation I consider in-

298 ACHATINELLA TURGIDA.

constant. Tip of apex white. Striation about the same as in
turgida. We have dextral as well as sinistral specimens/
The type-specimen is shown in pi. 59, fig. 17. This figure has
a pinkish tint in my reproduction. It should be cold white.

Moanalua. A series in Dr. Cooke 's collection is figured, pi.
56, figs. 7 to Id. The ground is light ocher varying to pinkish-
buff or nearly white, variously banded with chestnut, always
with a blackish streak behind the lip. Peristome broadly bor-
dered with blackish chestnut. Columellar fold fleshy with
white tip. Length 21, diam. 14.3 mm., to length 18.2, diam.
12.2 mm. 24 specimens, all dextral.

On the Moanalua-Halawa division ridge in a series of 13 in
Spalding coll., no. 3913, all are sinistral, part white, part with
many brown bands. They were found on the mokihana.

Halawa, top of the central ridge. In a series of 15, no.
2225 Spalding coll., all are dextral. The shell is white with
black-brown lip, a broad black streak behind it. A few have
light brown bands. A white one is figured, pi. 56, fig. 10. The
same form has been taken there by Mr. Wilder.

Aiea. In a set of seven in coll. Cooke, three are sinistral.
Amber yellow to Naples yellow, sometimes with faint brownish
bands or darker lines, similar to fig. 7. Always a blackish
streak behind the lip (pi. 56, fig. 6, Aiea, Cooke coll.). Mr.
Spalding also collected specimens in Aiea, no. 1820 and 249
of his collection.

Kalauao, small ravine near east crest, no. 2120 Spalding
coll. Ground yellow or nearly white, banded with chestnut,
black behind the lip. PI. 56, figs. 8, 8a.

Waimalu, and in a gauva forest at head of Waiau, Spalding
coll.

Also in the Gulick collection there are Waimalu specimens
both dextral and sinistral, dark carob brown with some bands
of the yellow ground on the last whorl, sometimes almost cov-
ered by the blackish color (pi. 56, fig. 12). The light form, pi.
56, figs. 3, 3a, was marked Waimalu with a query by Gulick,
and may be from further west. Some of the lot are like fig.

>.

Waiau-Waimano ridge. A tawny specimen with narrow in-

ACHATINELLA TURGIDA. 299

conspicuous streaks and faint bands of russet is figured. The
sutural band and border behind the lip are blackish. PL 56,
fig. 9, coll. by Spalding. Mr. J. S. Emerson has a very fine
series of the brown-banded form from this ridge.

34c. A. TURGIDA SIMULACRUM n. subsp. PL 56, figs. 13 to

The shell is sinistral, in shape like turgida; white with black
or chestnut bands and lines, usually wanting near the suture ;
embryonic whorls ivory tinted. Peristome deep livid purple,
the columellar tooth paler or sometimes white. There is a
black streak behind the lip.

Length 21, diam. 14.5 mm. ; 6% whorls.

Length 20.5, diam. 13.2 mm.

Length 18.6, diam. 13 mm. ; 6% whorls.

Waimano-Manaria ridge, along the summit trail, above the
locality of A. t. cookei (Spalding, Pilsbry, Merriam), types
108063 A. N. S. Ridge west of Manana (W. D. Wilder).

This race is intermediate between ovum and cookei, and
specimens can be selected which are transitional towards one
or the other. Yet it seems to be a pure race, even though the
range of fluctuation in banding is considerable, and it oc-
cupies territory a couple of miles long. It begins about one-
fourth or one-half mile above the cookei colony, and is found
thence along the crest of the ridge up to within about a half-
mile of the peak on the main range. It has thus a far greater
area than A. t. cookei , which probably arose from simulacrum
by a mutation at the lower edge of the simulacrum colony.
The fully marked, dark shells are mainly in the lower part
of its range, nearest to the cookei colony. Figs 14 to 14d
represent shells of the type lot. It differs from most A. t.
ovum by the white ground, purple lip and black bands ; from
A. t. cookei by having bands above the periphery, and the
ground-color is never tawny. Very rarely the whole base is
black. The embryos are either pure white above, slightly yel-
lowish below the sharply angular periphery, or they are ochra-
ceous above, buff with an ochraceous band below. No dextral
specimens have been found.

On the ridge west of Manana Mr. Wilder found specimens

300 ACHATINELLA TURGIDA.

with a very pale buff ground, variously banded or lineate ;
also a melanistic form in which the first 2% whorls are white,
next whorl with bands beginning buff, deepening to chestnut,
and then to uniform black on the last two whorls. PL 56,
figs. 13 to 13&. This black form resembles the black turgida of
the mottled colonies westward except in the late embryonic
and early neanic stages, which have the banding of simul-
acrum. It is not so rough a shell as A. byronii nigricans, the
surface being polished and minutely striate as usual in A.
turgida. Moreover it differs from nigricans by having the em-
bryonic whorls more conic, with the suture not at all im-
pressed, while in nigricans the embryo is lower, wider, with
more convex whorls parted by a noticeably impressed suture.
When studying Mr. Spalding's collection I noted that some
specimens in one of his Waimano-Manana lots have a blackish
apex. Others of the lot are typical simulacrum, with white
apex, and in one the whole base is black. The variation in
color of the apex in this lot calls for further study.

. A. TURGIDA COOKEI Baldwin. PL 57, figs. 15 to 17.

The shell is dextral or sinistral, with the last whorl bi-
colored, black or chestnut-black below the periphery, white,
light buff or tawny above, usually darker below the suture;
peristome livid pink (or brownish vinaceous) ; embryonic
whorls white or ivory.

"Length 21, diam. 15 mm.' (Baldwin).

Length 19, diam. 14 mm. (Cotype).

Length 18, diam. 13.2 mm. (Cotype, Proc. A. N. S., pi. 10,
f. 15).

Length 20, diam. 13 mm. ; GVs whorls.
* Animal : mantle dark slate with a narrow brown band en-
circling the outer edge. Superior portion of foot light brown,
mottled with slate, under surface dingy white' (Baldwin).

Waimano-Manana ridge at about 1,800 ft., extending sev-
eral hundred yards down the Waimano slope.

A. cookci BALDWIN, Proc. A. N. S. Phila., 1895, p. 220, pi.
10, f. 15 (July 2, 1895).

Tn shape this form varies from short with swollen last whorl

ACHATINELLA LEUCORRAPHE. 301

to more straightly conic with longer spire, as shown in the fig-
ures. It is often subperf orate. Mr. Baldwin's original speci-
mens, from the Waimano slope (at that time reached from
the valley) were dextral, with the upper surface buff. One is
figured, pi. 57, fig. 15 ; but among others obtained by Dr.
Cooke at the same place there are also some sinistral shells.
At the top of the ridge the cookei colony reaches to the trail
at the summit, but not beyond it, and is very narrow — the
matter of a hundred feet or so alongside the trail as I remem-
ber. It is longer in the other direction, extending several
hundred yards down a small spur. This small colony is the
only place A. t. cookei has been found. At the upper limit
of the colony the specimens are all sinistral (20 in coll. A. N.
S., 63 in Spalding coll.). In this sinistral lot from the upper
edge of the colony, the peristome is brownish-vinaceous. The
black base is produced by coalescence of bands. The bicolored
stage may begin as early as the beginning of the fifth whorl, '
but is sometimes as late as the last half of the last whorl ; the
bands appearing a little earlier.

A. t. cookei is now recognized by Hawaiian conchologists to
be a local race of the many-banded form simulacrum, oc-
curring immediately above it on the crest of the Waimano-
Manana ridge. Its resemblance to A. mustelina bicolor is
purely adventitious. The parent stocks of the two were cer-
tainly very unlike. "Named in honor of Hon. 0. M. Cooke,
by whose sons the shell was discovered/

35. A. LEUCORRAPHE (Gulick). PL 59, fig. 8.

"Shell dextral, shortly ovate-conic, striated with growth
(but scarcely with spiral) lines; gray, ornamented with irre-
gularly interrupted dark cinereous streaks and a few indis-
tinct, white, spiral lines ; suture broadly margined with snow
white ; whorls G1/^ ; apex blackish, first four whorls white,
nearly flat, the rest convex ; aperture small, somewhat ear-
shaped, white ; peristome slightly dilated and thickened
within ; columellar fold large, rosy. Length 19, diam. 12 mm.

"Var. Shell ovate-conic, regularly streaked with dark
gray and cinereous; spire straightly conic. [This var. belongs
to A. valida.]

802 ACHAT1NELLA LEUCORRAPHE.

' ' Kalaikoa, on the island of Oahu. Waimea ? a variety with
spire regularly conical is reported to have been found in Wai-
mea [this = A. valida var.]. It is allied, though not very in-
timately, to A. apicatus Nwc. No sinistral form of this species
has been found. The specimen figured is from Kalaikoa.'
(Gulick.)

Oahu: Kalaikoa (Gulick). Varieties on the ridges between
Kipapa and north Kaukinehua.

Apex leucorraphe GULICK, P. Z. S., 1873, p. 79, pi. 10, f. 2.

The type specimen of A. leucorraphe, pi. 59, fig. 8, no. 92 of
Gulick 's type series, Boston Soc., is a solid, compactly globose-
conic shell resembling A. swiftii in form. The tip of the em-
bryo is olivaceous black, this shade continues spirally down-
ward above the suture for about 1% whorls, the upper part
of which is white. The last 2y2 whorls are olivaceous black
or iron gray with streaks and zigzag tracery or mottling of
white. At places of growth-arrest and just behind the lip there
are brownish streaks. The suture has a white margin broader
than the subsutural impression. The aperture and peristoine
are white, with the faintest suggestion of violaceous. A nar-
row white line is traced around the periphery, but scarcely
noticeable except on the back. Length 19, diam. 12.1 mm.

The unnamed variety noted by Gulick as reported from
Waimea is certainly a form of A. valida leucozona. Part of
the lot is before me.

Recent collectors have not found specimens of the original
color-pattern of leucorraphe, but a good many shells collected
by Messrs. Spalding, Kuhns and Wilder are evidently color-
varieties of Gulick 's species. The original leucorraphe colony
in Kalaikoa was doubtless long ago destroyed by recession of
the forests. It must have been at the lower limit of the
species, which belongs in the main to the high ridges.

35a. A. LEUCORRAPHE iRwiNi n. subsp. PL 59, figs. 9 to 15a.

The shell is dextral or sinistral, compactly globose-conic.
Embryonic whorls bicolored, white with a dark helicoid spiral
which is olive-black or clove brown at the tip (or when the sur-

ACHATINELLA LEUCORRAPHE. 303

face is eroded, it becomes dark flesh color), changing to cinna-
mon buff or some ochraceous tint on the last two embryonic
whorls. Later wrhorls white, encircled with black lines or lines
and bands, most numerous on the base ; the lines above the
periphery often weak and yellowish. Sutural margin usually
white, sometimes with a dark line. Peristome pinkish lilac.

Length 19, diam. 13.3 mm. ; whorls 6.

Length 16.7, diam. 11.9 mm. ; whorls 6y±.

Division ridges between gulches of Kipapa and Waikaka-
laua, Waikakalaua and Kalaikoa, and Kalaikoa and Kaukine-
hua, above the 1,500 ft. contour, extending up each ridge to
within a mile of the main ridge ; abundant on mokihana, ieie,
lehua and alani (Irwin Spalding).

By its clean-cut bands on a white ground and dark apical
spiral this form resembles A. vittata simulans of the Nuuanu-
Kalihi ridge, to the point of identity. An expert might dis-
tinguish between good lots of each; nobody could rightly
separate a mixed lot. Yet the very identity of patterns and
their geographic discontinuity makes us suspect it highly un-
likely that they belong to the same species, for no Achatinella
is known to hold a pattern unchanged over so great a dis-
tance. It appears that we have to do with two species which
have evolved along parallel lines; a smaller western stock in
which some colonies retain the ancestral streaked pattern,
and a larger eastern, in which the streaked pattern is
now rare and rather unlike that of the western species. It is
the final stages of each which are so remarkably alike. The
nearest colonies of simulans and irwini are separated by fully
one-fourth the length of Oahu, where neither is found, al-
though some similar races of other species occur, such as A.
turgida simulacrum.

Mr. Spalding has noted that from Waikakalaua four-fifths
of the shells are dextral, and in Kaukinehua one-half are
dextral. The white ground and dark apex are constant, but
there is ample variation in the number and width of the bands.
The size varies from about 16 to 19 mm. The banded form
from the upper part of the Kaukinehua ridge, figs. 11 to 11&,
may be taken as typical of irwini (from 10A nearly to the

304 ACHATINELLA LEUCORRAPHE.

main ridge, on map, p. 277). The Kaukinehua shell, pi. 59,
f . 10, is a dextral form of the same color-pattern, but another
lot (2095 coll. Spalding) contains similarly banded sinistral
shells only.

Mr. Spalding 's no. 2042 from the ridge between South Kau-
kinehua and Waikakalaua (at 6 on map, p. 277), contains a
few specimens streaked like those of the southeastern ravines
of Waikakalaua (fig. 14a) ; others have a light buff ground,
with indistinct pale purplish viiiaceous stains in streaks and
spirals, deepening to light violet-gray on the back; apical
spiral olive brown ; lip light pinkish lilac, pi. 59, fig. 9. Others
are white with dark spiral lines below, similar to pi. 59, fig.
116. This colony also is a hybrid between the streaked and
the banded forms.

Still further southeastward, A. I. irwini is found on the
Waikakalaua-Kipapa division ridge. In a lot from two miles
above Waipio Mountain house (at 5 on map, p. 277), the col-
ony (no. 1986) is chiefly dextral and banded, sometimes with
dark subsutural border and reddish bands above the dark
ones, sometimes typical (pi. 59, fig. 12, coll. by Spalding) ; but
there are also some streaked shells, like those from lower down.
This is therefore a transitional hybrid or undifferentiated col-
ony. There are some very short shells, 15.6 x 12 mm., with 5y2
whorls.

Further down the same ridge, in the "southeastern ravines
of Waikakalaua, half a mile above Waipio,' pi. 59, figs. 14-
14&, Mr. Spalding collected a series containing some speci-
mens very close to the type of leucorraphe; differing mainly
in being sinistral (175 sinistral, 4 dextral in his 110. 2328 from
Station 4 on the map), and in having a tinted lip and a
rufous or chestnut line below the suture, instead of a white
band. It is a pure race, showing only "fluctuating variation'
from dark to light. The dark shells have a summit colored ex-
actly like leucorraphe; later whorls with black or olivaceous-
black streaks and faint or distinct white spiral lines ; lip
tinted, fleshy brown. Length 18, diam. 22 mm. The lighter
shells have the apical spiral more of a russet-vinaceous hue,
and the stripes of the later whorls from cinnamon to walnut

ACHATINELLA LEUCORRAPHE. 305

brown, with or without whitish spiral lines; sutural border
tawny, lip faintly pink. A small lot of 5 dextral, 4 sinistral
shells from Kipapa, in coll. Spalding, are in color like the
preceding colony. PL 59, fig. 13 is from no. 2328, on the ridge,
living on guava, and figs. 14 to 14& from no. 1906, lower, on
the slope into Waikakalaue. This form of irwini resembles
the neglect a (coniformis) pattern of A. swiftii, except in the
marking of the early whorls, which seems to be a more con-
stant character than the later pattern. It connects with the
invini pattern from higher up, through several hybrid colo-
nies, as noted above.

It remains to notice two rather aberrant forms from Mr.
Thaanum's collection, possibly having relations with A. swiftii.

PL 59, figs. 15, 15a. A series from Mr. Thaanum labelled
"Waipio' (no doubt from somewhere in the lower Kipapa-
Waikakalaua ridge) consists of small dextral shells with com-
siderable cinnamon -rufous on the spire and below the last
suture, the last whorl or two more or less profusely banded
with black, the bands usually somewhat speckled. The apex
is marked as in leucorraphe, though often rather weakly. The
aperture is light Payne's gray within, the lip and columellar
fold lavender or fleshy with darker spots. Length 16 to 17
mm.

This is a pure colony of quite distinct appearance.

PL 59, figs. 16 to 16&. A series of small dextral and sinistral
shells from "the rosea ridge' (I suppose the middle ridge of
Poamoho, or possibly the northern ridge of Waikakalaua),
from Mr. Thaanum, contains forms having much the appear-
ance of the tuberans pattern of A. swiftii, but differing by the
well developed, dark, apical spiral of leucorraphe. The pat-
tern consists of gray streaks and spiral ba,nds, forming darker
squarish spots or short bars at the intersections. Sometimes
the bands predominate and are iron gray or blackish, separ-
ated by white lines or bands. Lip and columellar fold of a
lilac tint, interior bluish white. This form seems near to that
from the southeastern ravines of Waikakalaua.

306 ACHATINELLA SWIFTII.

36. A. SWIFTII Newcomb. PL 58 ; pi. 59, figs. 1 to 4e, 6, 7.

"Shell ventricose, pointed at the apex, smooth, polished,
shining; whorls 6, slightly rounded above, the last strongly
inflated, distinctly margined above ; lip purplish rose, thick
and slightly subreflecteci ; aperture ovate ; columella short,
terminating in a tubercle of the color of the lip ; three first
whorls white, the lower with very fine and numerous mark-
ings of black and white, arranged longitudinally to the shell,
giving it a grayish aspect; fine obsolete white lines traverse
the shell transversely, and a white sutural line is traced on
the last two whorls. Length fourteen, width ten-twentieths of
an inch.

"Var. a. With a broad fascia cutting the body whorl.

"Var. b. Yellowish ground-color with few markings.

"This species approaches A. turgida, but is distinct in ap-
pearance, locality and habits ' (Newcomb).

Oahu : District of Ewa ( Newcomb ) . In its various forms
this species probably extends from the western ridge of Wai-
awa to the ridges of Kaukinehua.

Achatinella swiftii NEWC., P. Z. S., 1853, p. 133, pi. 22, f. 9,
9a, 1854; Ann. Lye. N. II. of New York, vi, p. 325. — PFR.,
Monogr., iv, 528. — THWING, Orig. Descript. Achatinella, pi. 1,
f. 7. — Apex albospira SMITH, Proceedings of the Zoological
Society of London, 1873, p. 77, pi. 10, f. 8. — Apex innotabilis
SMITH, P. Z. S, 1873, p. 78, pi. 9, f. 23 (not f. 19).— Apex
neglectus SMITH, P. Z. S., 1873, p. 78, pi. 9, f. 22. — Apex ver-
sicolor GULICK, P. Z. S., 1873, p. 80, pi. 9, f. 18.— Apex flavi-
dus GULICK, P. Z. S., 1873, p. 80, pi. 10, f. 1, la. — Apex coni-
formis GULICK, P. Z. S., 1873, p. 80, pi. 9, f. 17 (not f. 23).—
Apex tuberans GULICK, P. Z. S., 1873, p. 81, pi. 10, f. 3. —
Apex polymorpha GULICK, P. Z. S., 1873, p. 81, pi. 10, f. 5.

A. swiftii is indifferently dextral or sinistral, though one
or the other direction of coil usually predominates in any one
colony, sometimes to the exclusion of the other. The em-
bryonic whorls are ivory or ocher-tinted, paler towards the
tip which is usually a little dusky, though sometimes flesh-
tinted or quite white. The coloring of the adult stage is in

ACHATINELLA SWIFTII. 307

r

inaiiy shades of dull brown and purplish-gray, sometimes with
a yellowish cuticle. Streaks predominate, often interrupted
by light or dark bands or lines ; but the banded patterns are
not definite or sharply contrasted as in some related species.
I have not seen uniform blackish or white specimens. New-
comb's description of the coloration applies to but a small
group in the species; other patterns have given occasion for
the definition of several supposed species.

A. swiftii differs from A. apexfulva by the shorter, less
drawn-out embryonic whorls, which moreover are usually
tipped minutely with gray. A. decora has special color-pat-
terns, not like those of swiftii. A. valida has the summit more
slender and pointed. A. turgida never has a gray tip, and the
lip is usually darker. In A. leucorraphe the dark tip is much
more distinct, and the color-patterns mostly different.. In
general, A. leucorraphe is a shell of high elevations, while A.
swiftii inhabited the lower forests. While the various forms
of swiftii described by Mr. Gulick seem to have been common
in the Fifties, they are now nearly or quite extinct, with the
forests they adorned. Such as remain are mostly unlike the
old lots, being no doubt from places further in the back coun-
try. A. swiftii is so prolific in color-mutations, so variable in
shape, that no definition covering its several forms would
serve to exclude the adjacent species. The best that can be
said is that with a fair series from any locality, I find no ser-
ious difficulty in determining the species.

The supposed species albospira, innotabilis, neglecta, versi-
color, flavida, coniformis, tuberans and polymorpha were based
upon selected patterns, most of them known to be out of lots
having other patterns which connect all in an inextricable tan-
gle of pattern-combinations. Before me are Gulick 's and part of
Smith's types, and a series of several hundred shells, mainly
collected and named by Gulick. I have gone over them with
utmost care ; and have been quite unable to find basis for
subspecies. Part of the type-specimens are not even common
color-forms in the lots they were selected from. So far as I
can see, Gulick might as well have divided his material into
twenty species instead of eight, since none of them rest upon

308 ACHATINELLA SWIPTII.

any other basis than selected extreme patterns. It will be
noted that not less than five of these supposed species were
found in two small valleys, Ahonui and Kalaikoa, at the end
of the Kaukinehua ridge. The original descriptions are given
below in the order of their publication, followed by notes on
the types and other specimens.

Typical form of A. swiftii. — The original figures of A.
swiftii are reproduced on pi. 29, figs. 9, 9a. Newcomb has
stated that ''the type figured in the Zoological Proceedings is
not the usual pattern of the shell, but is one of the more un-
common varieties.' Neither figure agrees in color with his
description. The form described must therefore be consid-
ered the type; not those figured. His fig. 9a may be an A.
turgida, but fig. 9 is apparently swiftii.

Two sets, nine specimens, were given by Newcomb to the
Academy in 1854. Four of the lot have the white suture men-
tioned by Newcomb. Two are sinistral. One of these is fig-
ured, pi. 58, fig. 1, no. 10313 A. N. S. These shells agree
well with Gulick's series from "Wahiawa, ' except that a
white sutural band is very rare in the latter, and the color of
the embryo sometimes differs. I consider the Wahiawa region
as type locality, though the range of the species in its several
patterns lies mainly east of this. Wahiawa shells from the
Gulick collection are figured, pi. 58, figs. 2 to 2c. The em-
bryonic whorls are not drawn out as in A. apexfulva. They
are nearly white, but under a lens there is more or less buff
or ochraceous tint, either on the second whorl or in form of a
band above the suture of the third whorl. The tip may be
white, but in several of Newcomb 's shells the first half- whorl
has a faintly gray or dusky shade. Gulick's Wahiawa shells
have the embryonic whorls ivory to pale ocher, the tip white
or with the faintest gray tint. The color of the later whorls
is in streaks of burnt umber to blackish cut by paler or white
spiral lines, which may be numerous or very few. Sometimes
the streaks are of some tint of vinaceous drab.

The specimens from Wahiawa which Gulick noted under his
description of A. leucozonus are in my opinion merely A.
swiftii with white sutural band. Very few were found, part
of them now before me.

ACHATINELLA SWIPTII. 309

Specimens are figured from Kalaikoa valley, from the Gu-
lick collection, pi. 58, fig. 7, and figs. 13, 13a, 136. These last
were regarded by Mr. Gulick as a form intermediate between
gulickii and flavida, but they seem to me rather swiftii-flavida.
PL 59, figs. 1, la, 16, selected from Gulick 's Ahoriui lot,
further illustrate the decadence of streaks.

The descriptions of forms considered synonyms of swiftii
here follow, together with notes on the types and other speci-
mens.

"Apex albospira Sm. (pi. 59, fig. 3, type specimen, no. 96
Bost. Soc.) Shell dextral, subperf orate, shortly ovate-conic,
little shining, striated with growth-lines but hardly spirally;
white, a zone above the periphery and the base livid brown;
suture lightly margined; whorls 6, the first four a little con-
vex, the rest convex; aperture white; peristome thickened
within; columellar fold strong, white, reflexed, nearly cover-
ing the small crevice. Length 17, diam. 10.

' ' Var. : Shell white, ornamented above with a few yellow
zones and yellow towards the base, encircled with a few darker
zones.

"Reported to be from Ewa, on Oahu7 (Smith).

The type specimen is an unique, no. 96 of the Gulick type
collection, Boston Soc. N. H. The apex and whole embryonic
shell are pure white. A band appears just prior to the begin-
ning of the penult, whorl. On the last whorl the band and the
whole base are vinaceous drab, lightly streaked with white.
Aperture and peristome white. Length 16.2, diam. 10.2 nun.
It was probably somewhat dwarfed by misfortune, as there
was a break in the fourth whorl, and another, affecting the
whole lip, about 4 mm. behind the final peristome. Some of
Gulick 's polymorpha have the same color-pattern, but with a
dusky apical tip. However, in some other patterns of poly-
morpha the tip is quite white. There can be no doubt that
albospira is merely a form of "polymorpha7 or possibly
"flavida" — the exact name is not significant, since these Gu-
lickian forms are not real races.

"Apex innotabilis Sm. (pi. 59, fig. 7, photographic repro-
duction of original figure). Shell sinistral, subperf orate,

310 ACHATINELLA SWIFTII.

globose-conic, lightly striated with growth-lines ; fawn colored,
streaked with darker (sometimes ornamented with a few
white and brown spiral lines) ; apex almost white, the nucleus
reddish-corneous. Whorls 6, the first four flattened, the rest
convex. Suture distinctly margined. Aperture pale reddish
within ; peristome lightly dilated, thickened within ; columellar
fold large. Length 18%, diam. 11% mm.

"Habitat Sandwich Islands. Of a uniform fawn-color,
with darker stripes of the same hue, the apex being paler,
almost white, and the nuclear whorls reddish horn-colored'
(Smith).

The type is in the British Museum. I think it is a special
pattern of the shell described by Gulick as A. polymorpha.
The color of the apex is the same, and some polymorpha have
similar body-color.

'Apex neglect us Sm. (pi. 59, fig. 6, photographic repro-
duction of the type figure). Shell either sinistral or dextral,
imperforate, globose-conic, lightly striated with growth lines ;
dilute brown, streaked with darker or cinereous, umbilical
region often chestnut, apex white; spire a little concave;
whorls 6, the first four flattened, the rest convex; suture dis-
tinctly margined with deep chestnut. Aperture whitish with-
in ; peristome brownish, thickened within ; columellar fold
strong, reddish. Length 17, diam. 11% mm.

"Habitat Sandwich Islands. In form this species agrees
very fairly with some varieties of lugubris Chemii. ; but it has
a shorter spire, and blunter apex than the usual form of that
species. The coloration is of a different character' (Smith).

Mr. Sykes remarks that "the variety neglecta Smith is not
really so greenish as represented in the original figure ; fig.
23 on the same plate [innotafoilis, my fig. 7] gives a better
idea of the real color.' I entirely agree with Mr. Sykes that
coniformis Gulick is the same thing.

"Apex versicolor Gk. (pi. 58, fig. 14, type specimen). Shell
sinistral, globose-conic, glossy, striated with growth lines
(scarcely with spirals) ; grayish white, irregularly streaked
and zoned with blackish-brown, the base blackish-brown. Su-
ture lightly margined, white (sometimes brown). Spire
lightly concave. Whorls 6, the first three pale, a little convex,
the rest convex. Aperture white, receding ; peristome slightly

ACHATINELLA SWIFTII. 311

dilated, bordered within, tinted with dilute rose-brown ; colu-
mellar fold strong, pale rose color. Length 19, diam. 13 mm.

' Var. : shell gray- white, scarcely streaked ; the last whorl
zoned with blackish-brown at the periphery and base.

' Station, on trees ; habitat, Ahonui and Kalaikoa, on Oahu.
Affinities: it seems to be an intermediate form between A.
decorus Fer. and A. tuber ans Gk. This species it always sinis-
tral. The specimen figured is from Ahonui' (Gulick).

In the type specimen, no. 101 of Gulick 's series, Boston Soc.
coll., the first embryonic whorl is gray white, becoming
light ochraceous buff on the second whorl, after which white
bands appear. The last 2% whorls have light purplish gray
streaks on a white ground. Immediately above the periphery
the streaks coalesce into an irregular white-streaked band of
blackish Vandyke brown ; and a similarly colored area occu-
pies much of the base. The supraperipheral band ascends the
spire, forming a narrow dark border above the suture. The
lip and columellar fold have a pale flesh tint. Length 18,
diam. 13 mm.

There is a large series of versicolor from Ahonui in coll. A.
N. S., collected by Gulick. An inspection of these shows at
once that the type was merely a .specimen selected out of a
chain of patterns which completely unite versicolor, tuberans,
coniformis and even flavida, all of which Gulick recognized
from Ahonui. The versicolor pattern also runs inextricably
into the polymorpha pattern. Several additional versicolor of
Gulick 's Ahonui set are figured, pi. 58, figs. 14a to 15. Many
specimens are smaller than the type, down to 16 mm. long.

"Apex flavidus Gk. (pi. 58, figs. 12, 12a, cotypes no. 95 Gu-
lick type coll., Boston Soc.). Shell dextral (sometimes sinis-
tral), shortly ovate-conic, little shining, striated with incre-
mental (but hardly spiral) lines; more or less yellow, en-
circled in the middle of the whorl with a wide chestnut zone
(sometimes two or three zones on the base) ; suture narrowly
margined with brown (sometimes white). Whorls 6, a little
convex, the first three always white ; spire almost straightly
conic, scarcely concave. Aperture whitish, receding; peristome
pale rose, slightly dilated ; thickened within ; columellar fold
large, roseate. Length 20, diam 12% mm.

"Var. a. Shell dextral or sinistral, subglobose-conic, yel-

312 ACHATINELLA SW1PT1I.

low, encircled with two to six brown or cinereous-brown
bands [pi. 58, fig. I2a].

"Var. b. Shell entirely yellow.

"Station, on trees. Habitat: the metropolis is in Kalaikoa
on Oahu ; it is also found in Ahonui.

"Affinities: it is most nearly allied to A. tuberans Gulick.
Remarks: In Kalaikoa all are dextral; in Ahonui one-tenth
are sinistral. The specimens figured are from Kalaikoa'
(Gulick).

The type of flavidus is the largest specimen I have seen. It
retains the yellow cuticle only on the base. Above the periph-
ery it is white, with a broad Vandyke brown band at the
periphery and above the suture, and a brown line below the
suture. The embryonic whorls are white with a pale gray tip.
This pattern, although selected for the type, is rare. Out
of 53 Kalaikoa flavidus from Gulick before me, only 5 have
it. The bandless form is also rare. Usually there are several
bands, of a russet color, or more olivaceous (Dresden brown
or Saccardo's olive). Then we have shells with purplish-gray
streaks appearing faintly; also, all manner of transitions to
the Kalaikoa "tuberans.' A common pattern is shown in pi.
58, fig. 126, Kalaikoa, Gulick coll., A. N. S. Shells with
the yellow cuticle of fl,avida over the oblique streaks of siviftii
also occur in Kalaikoa. pi. 58, figs. 13-13&, Gulick collection.
The shells of this lot vary a good deal, like all the Kalaikoa
forms of this group. I am rather surprised that Mr. Gulick
did not make a species of them, as the more characteristically
marked examples, figs. 13a, fc, are as distinct as any of his
Kalaikoa- Ahonui species.

"Apex coniformis Gk. (pi. 59, figs. 2, 20, Ahonui, and pi.
58, figs. 8, 16, Gulick coll.). Shell sinistral (sometimes dex-
tral), globose-conic, glossy, obliquely striated with growth
lines but scarcely spiral lines; brown, streaked with pale
brown, indistinctly encircled with several whitish lines ; suture
margined with deep brown. Spire lightly concave. Whorls
6!/2> the first 3 1/2 a little convex, whitish, the rest convex.
Aperture receding, white ; peristome a little dilated and thick-
ened within, the margin tinted with dilute rose (or brown) ;
columellar fold strong. Length 18, diam. 12 mm.

ACHATINELLA SWIFTII. 313

* * Var. Shell sinistral, cinereous-brown, streaked with white
and encircled with a few white spiral lines ; suture brown.

"Station, on trees. Habitat: Kalaikoa and Ahonui, on
Oahu ; very rare in Wahiawa and Helemano.

"Affinities. It seems to be intermediate between A. decorus
Fer. and A. apicatus Nwc. Remarks: This species is very
rarely dextral. The specimen figured is from Ahonui'
(Gulick).

The coniformis pattern always has a brown line or narrow
band bordering the suture below. The embryonic whorls are
ivory or buff in varying tints, the tip white or sometimes
faintly gray. The later whorls are dusky drab, clove brown,
or Vandyke brown streaked with lighter brown or creamy.
Some are streaked with very pale purplish gray, others al-
most white. Usually there are pale spiral lines, often scarcely
visible, or again quite distinct. All of the specimens figured
are from Ahonui, pi. 58, fig. 8 agreeing with Gulick 's type.

This form is identical with A. neglectus Smith, which is
merely one color-tone in the coniformis scale. Some speci-
mens connect completely with swiftii patterns.

"Apex tuberans Gk. (pi. 58, fig. 6, type, no. 93 of Gulick 's
type series, Bost. Soc.). Shell dextral, globose-conic, glossy,
striated with growth (but hardly spiral) lines; cinereous
streaked, interrupted with several whitish and blackish-brown
zones; suture margined, pallid. Whorls 6, the first three
whitish, a little convex, the rest convex ; spire concave. Aper-
ture white ; peristome slightly dilated, thickened within,
tinted with dilute rose-brown ; columellar fold strong, whitish.
Length 20, diam. 13!/2 mm-

'Var. a. Shell white, variously zoned with brown and buff.

"Var. &. Shell yellowish, closely zoned with blackish-brown.

* Station, on trees. Habitat : The metropolis of the species
is Kalaikoa, on Oahu. It is also found in Ahonui, and single
specimens have been brought from Wahiawa and Helemano.

"Affinities. It is most nearly allied to A. flavidus Gk.

"Remarks. This is a dextral species. Sinistral forms are
found only in a few specimens passing into other species.'
( Gulick ) .

The apical whorls are white or ivory yellow, the tip slightly
dusky or pale purplish gray. The last whorl, in the type spe-

314 ACHATINELLA SWIFTII.

cimen, is white, streaked with light purplish gray at and above
the periphery, cut by white spiral lines. There are several
narrow black-brown bands and lines on the base, and two
above. Sutural border slightly tinted.

A large lot in the Gulick collection shows wide variation in
pattern and color. The ground is often yellowish or yellow at
the base. Blackish or chestnut spirals or pale streaks may
predominate. Profusely banded specimens come close to the
banded forms of A. leucorraphe. Others run directly into
flavida and swiftii. Some are figured on pi. 58, figs. 9 to 96,
11, Ha, Kalaikoa, Gulick coll.

Forms closely resembling Mr. Gulick 's tuberans were col-
lected by Mr. Spalding much further southeast.

A lot from the summit of the southeastern ridge of Kipapa,
pi. 58, figs. 4, 4a from no. 3652 of Mr. Spalding 's collection,
consists of dextral and sinistral shells. The white apex has a
dusky tip, and usually an ochraceous band above the suture
of the embryonic whorls, the rest of the spire being white, or
with a cinnamon or brown band or line below the suture.
The last whorl is grayish olive with a white band or bands, or
the grayish tint may appear only in bands at periphery and
base. Lip and columellar fold have a pale lilac tint. Some
of these shells are very short in contour.

Length 18, diam. 12.7 mm.

Length 17, diam. 11 mm.

Length 15, diam. 11.8 mm.

A lot from one of the minor spurs between the terminal
branches of the Kipapa-Waiawa division ridge, pi. 58, figs. 5,
5a, 5& from no. 2266 Spalding coll., consists largely of shells
similar to the preceding, but there are some like those figured,
in which black lines or bands are superposed over the other
pattern ; sutural border cinnamon ; lip lavender to lilac. There
are also some transitional specimens between these patterns,
with the dark bands narrow and rather olivaceous. It is these
shells which are nearest to Gulick 's tuberans. In a lot of 143,
all are sinistral. Length 18, diam. 13 mm.

Dr. Cooke took a very pale form of "tuberans" on a ''low
ridge in Wahiawa", pi. 58, figs. 3, 3a. They resemble some of
Mr. Spalding 's Kipapa shells.

ACHATINELLA SWIFTII. 315

"Apex polymorpha Gk. (pi. 59, fig. 4 to 4e. Waipio,
Gulick coll.). Shell dextral, subperf orate, globose-conic,
glossy, striated with growth (scarcely with spiral) lines; di-
lute brown, ornamented with several zones and blue-black
streaks, and encircled with a snow-white zone at the periphery.
Suture lightly margined, brown. Whorls 6, the first four a
little convex, whitish, the rest convex. Spire slightly con-
cave. Aperture white ; peristome thickened within, brown-
tinted; columellar fold strong, whitish. Length 17, diam.
12 mm.

"Var. a. Shell fawn, the suture brown.

"Var. &. Shell white, a few zones and the base brown,
suture brown.

''Station, on the leaves of the ohia and other trees.
Habitat, The typical forms are found in Waipio and Wahiawa
on Oahu. Varieties are sometimes found in Kalaikoa and
Ahonui.

"Affinities. It is most nearly allied to A. versicolor Gk.,
and A. tuberans Gk. Remarks: In Waipio and Wahiawa
two-thirds of the specimens are dextral; in Ahonui and
Kalaikoa only about one-eighth are dextral. The specimen
figured is from Waipio >: (Gulick}.

In the pattern selected by Mr. Gulick as typical, the shell
is dextral, brownish drab, varying in shade, and streaked with
paler or whitish ; there is a white peripheral girdle and some
darker spiral lines, the sutural border being tawny or brown.
Embryonic whorls ivory, or white shading into yellow towards
the suture below, the tip faintly dusky, gray, or sometimes
white. This particular pattern is not the commonest, and ap-
proaches the coniformis pattern. In other shells of Gulick 's
Waipio lot the color is orange-cinnamon ; pinkish buff towards
the suture, cinereous below. Pale yellow below, white stained
with pale purplish gray above. White with chestnut bands
and lines ; also other patterns, running into flavida, tuberans ,
etc. See pi. 59, figs. 4-4e, Waipio, coll. by Gulick. PI. 58,
figs. 10, 10a, Kalaikoa, Gulick. Some of the "polymorpha'
from Kalaikoa are not distinguishable from cestus, though I
do not believe them to be directly related to that. I believe it
to be a case of the same pattern appearing independently in
two derivatives of the same ancestral stock.

A. innotabilis Smith is one of the "polymorpha" patterns.

316 ACHATINELLA SWIFTH.

36a. A. SWIFTII CHROMATACME ii. subsp. PI. 59, figs. 5, 5a,

The shell is dextral, imperforate, solid, with straight-sided
or very slightly concave spire ; glossy ; embryonic shell chest-
nut brown with pale or whitish spirals, and fading to white
at the apex. Later whorls copiously streaked with burnt
umber (varying to blackish) on a warm buff ground; the
streaks usually blended together more or less, and not reach-
ing up to the suture, and usually interrupted by a white line
or band at the periphery. There are also, in some specimens,
indistinct pale spiral lines over the streaks. In other speci-
mens of the type lot the ground-color is white, and the mark-
ings olive gray ; the embryonic whorls colored as in the other
pattern. The suture is margined with an impressed line on
the last 2!/2 or 3 whorls. Whorls not very convex, the last a
little inflated in some individuals. Aperture pale blue within,
the lip and columellar fold lilac, lip dark-edged.

Length 20, diam. 12.5 mm. ; 61/^ whorls.

Length 18.4, diam. 13 mm.

Length 18.4, diam. 12.5 mm.

Waiawa. Cotypes no. 108804 A. N. S. P. and in Bishop
Mus., from Mr. Thaanum's collection.

This peculiar form was collected by Mr. Kuhns, who se-
cured a good series. The dark colored embryonic shell with
white apex is peculiar and unlike any other known form of
the region. The strongly tinted lip and columellar fold as
well as the more produced spire are unlike A. swiftii; and in-
deed, chromatacme is grouped with A. swiftii merely as a
temporary expedient, because one feels some diffidence about
proposing a "new species" in the turgida-swiftii group. It
does not seem to be directly related to any form of A. turgida
I have examined, yet the possibility of such a relationship
must be considered.

'

. A. (SWIFTII?) DOLIUM Pfeiffer. PL 30, fig. 15.

Shell perforate, ovate-conic, rather thin, lightly striatu-
late, little shining, pale buff variegated with brownish bands
and narrow streaks ; spire conic, apex rather acute ; suture
scarcely marginate; whorls 6, convex, the last a little longer

ACHATINELLA APEXFULVA. 317

than the spire, swollen, subcompressed at the ,base ; aper-
ture oblique, obversely ear-shaped, white within; columellar
fold high, dentiform, white ; peristome thin, sublabiate within,
the right margin narrowly expanded, columellar margin di-
lated, spreading. Length 17, diam. 10 mm. ; aperture 10 mm.
long, 5 wide." (Pfr.)

Sandwich Islands (Newcomb).

Achatinella dolium PFR., P. Z. S., 1855, p. 5, pi. 30, f. 15 ;
Monographia Hel. Viv., iv, 528. — SYKES, Fauna Hawaiiensis,
p. 302. — THWING, Orig. Descriptions, etc., Occas. Pap. B. P.
Bishop Mus., iii, 110. 1, p. 102, 1907 ("Kalae, Molokai").

A. dolium is known only by a single specimen, the type, in
the British Museum. Mr. E. A. Smith has kindly looked at it
for me, and gives the following notes: 11A. dolium is nearer
vittata than lorata in shape, but not quite the same. It is a
much shorter shell than A. lorata. The extreme tip of apex
(one whorl only) is light olive brown. Spiral striation hardly
traceable. The lip and interior of aperture are very pale
lilac, not white as stated by Pfeiffer.'

Comparison should be made with a good series of A. swiftii.
A. dolium is probably only a color-form of that polymorphic
shell. Mr. Sykes considers it a form of Achatinella s. str.,
with this note: "Hab. Molokai (Baldwin). I fancy this habi-
tat must be wrong and that the species really belongs to Oahu ;
the shell is very close to A. hanleyana Pf., and may prove to
be only a color variety.'

37. A. APEXFULVA (Dixon). PL 50, fig. 15; pi. 60, figs. 1 to Ic.

Shell imperforate, dextral, ovate-piriform, solid, very
glossy; outlines of the spire concave. 3Vs embryonic whorls
Naples yellow, following whorls varying from blackish carob
brown to chestnut, sometimes with some whitish streaks and
spiral lines ; the suture narrowly light-edged ; lip flesh or
salmon color, columellar fold nearly white. The embryonic
whorls are almost flat and are unusually elongated or drawn
out, especially the last one. Last whorl inflated. Aperture
is bluish white within, the lip moderately thickened.

Length 19, diam. 12.2 to 12.9 mm. ; whorls 6.

318 ACHATINELLA APEXFULVA.

Oahu : South side of Opaeula gulch near the lower limit of
the woodland (J. S. and 0. P. Emerson, about 1860). Var-
ious varieties from the Kaukinehua ridge to beyond Opaeula.

Helix Apex Fulva DIXON, A voyage round the world ; but
more particularly to the north-west coast of America, p. 354.
-Turbo Apex Fulva DIXON, on unnumbered plate, fig. 1,
1789. — Achatinella apexfulva Dixon, SYKES, Fauna Hawaii-
ensis, p. 298. — Turbo lugubris, etc., CHEMNITZ, Neues Sys-
ternatisches Conchylien Cabinet, xi, p. 278, pi. 209, f. 2059-60,
1795. — Helix lugubris Chemn., FERUSSAC, Tabl. Syst., p. 56
(not Helix lugubris GmeL, Syst. Nat., 13, p. 3665). — FERUS-
SAC, in Freycinet's Voyage autour du Monde de 1'Uranie et
la Physicienne, Zoologie, p. 479, 1824. — Achatinella lugubris
Chemn., PFEIFFER, Monographia, ii, p. 239 ; iii, 465 ; iv, 452 ;
vi, 177. — REEVE, Conch. Icon., vi, pi. 2, f. 10a (not 10&). —
THWING, Orig. Descript., etc., p. 25, pi. 1, f. 6. — Monodonta
seminigra LAMARCK, Anim. sans Vertebres, vii, 1822, p. 37 ;
Edit. Desh., ix, p. 181. — DELESSERT, Recueil de Coq. decrites
par Lamarck, pi. 37, f. 2a-c. — Bulimus seminiger MKE.,
Synops. edit. 2, p. 26. — Achatinella pica SWAINSON, The Quar-
terly Journal of Science, Literature and Art, Series 2, iii,
January to June, 1828, p. 84 ; Zoological Illustrations, iii, 2d
Ser., p. 99, pi. 99, f. 1.

A. apexfulva is chiefly characterized by its peculiar nipple-
like apical whorls. The embryonic stage has an enameled ap-
pearance, the individual whorls being longer and flatter than
in related forms (except perhaps some specimens of A. tur-
gida). The lip is tinted (except in albinos), but never deep
brown or blackish. The tip of the apex is never black or even
dusky. Its range lies west of the turgida area, and so far as
we know at present, several gulches and ridges lie between
the areas of the two species. They are certainly near akin.

PL 50, fig. 15 is copied from one of Dixon 's figures. PL 60,
figs. 1-lc and the above description are from specimens taken
by Mr. J. S. Emerson over fifty years ago, in a fine grove of
tall, dark-leaved trees on the south side of the deep Opaeula
gulch, on the edge of a rocky stream. The area of this grove
is estimated by Mr. Emerson as rather less than half an acre.

ACHATINELLA APEXFULVA. 319

It was then near the lower limit of woodland, which long ago
receded far up the gulch. This colony was composed of en-
tirely typical apexfulva, and as the range of single patterns is
usually quite restricted, there is good reason to believe that
this grove was a remnant of the colony from which the native
Hawaiians gathered shells for the lei obtained by Captain
Dixon.

The colony was a pure one, though showing some "fluctu-
ating variation ' ' by the presence in some individuals of whit-
ish or white spiral lines. Of 49 taken by Mr. Emerson, all are
dextral. 33 are blackish carob-brown, fig. 1; 8 chestnut (the
shade and gloss of a horse chestnut), fig. la; 7 faintly marked
with white, fig. 16 ; and 1, immature, banded with white, fig.
Ic. This last specimen is unique in the colony.

So far as I know, this is the only locality for strictly typical
apexfulva, in a pure race. In some other colonies that pat-
tern is found in hybrid communities consisting chiefly of other
color-forms, such as A. a. apicata and A. a. beata. Mr. Emer-
son has two apexfulva patterns from high in Helemano, but
their associates are unknown to me.

Synopsis of the distribution of the races. — Typical A. apex-
fulva is to be looked upon as an aberrant, melanistic local
race of a species widely spread, from Opaeula to Kalaikoa,
commonly appearing in a streaked garb, and known as A.
apicata.

In the central part of its area, apicata is chiefly a dark-
streaked shell with rufous sutural line. Westward there is a
tendency to blend the streaks, and a white sutural band ap-
pears. I have called this the cervixnivea pattern. On the
Poamoho-Helemano ridge this passes into the various forms
comprised under the term beata, with the shell banded, white
or black.

Farther west, on the ridge beyond Opaeula gulch, we have
the pink form, vespertina ; and the black or deep liver-brown
form, apexfulva, low in the gulch, and now probably extinct.

Eastward there is a peculiar pink-banded race which I have
called A. a. aloha, on the ridge dividing the upper end of
north Kaukinehua. The lower end of the main Kaukinehua

320 ACHATINELLA APEXFULVA.

ridge was formerly the habitat of the pattern called gulickii,
which has a white subperipheral band, and of lilacea, with
continuous vinaceous streaks. These are now nearly or quite
extinct in the localities where Gulick collected.

History of A. apexfulva. — The several synonyms of A. apex-
fulva — lugubris, pica and seminigra — were based upon ex-
actly the same form, their types having been taken from leis
(necklaces). It appears that shell leis were strung by the
natives of the good agricultural region about Waialua Bay,
who doubtless got the shells from the lower forests in the
back country, in various places in Kawailoa and Helemano
districts. They were carried or traded eastward, and so ob-
tained by explorers harboring at Honolulu. It is altogether
likely that all of the A. apexfulva of these leis were from
some one colony in Opaeula Gulch. The Messrs. Emerson's
grove may well have been the last remnant of this colony.
According to Captain Freycinet, shell leis were going out of
style at the time he visited Oahu, the tawdry European orna-
ments being in fashion. I am glad to say that the charming
custom of wearing flower leis continues to this day.

The original descriptions follow.

Helix apexfulva. — ' ' . . . the natives form necklaces, brace-
lets and other ornaments. One of these necklaces afforded a
singular species of the Helix genus of Linna3us, which I was
informed is a fresh-water shell. It is outwardly smooth, has
seven spires, and is of a black-brown color except the tip
which is pale-yellow ; the inside is smooth and white, and the
mouth is marginated within. It is remarkable for a knob or
tooth on the columella, but which does not wTind round it,
consequently excludes it from the Voluta genus of Linnaeus,
to which at first sight it appears to be related. As I presume
it to be a species hitherto undescribed, I have taken the liberty
to give it the trivial name of Apex Fulva, or the Yellow Tip.
A figure of it in two views is given in one of the following
plates ' (Dixon).

A copy of Captain Dixon 's figure is given, pi. 50, fig. 15.

In the Neues Systematisches Conchylien-Cabinet, vol. XI,
1795, Joh. H. Chemnitz gives the following description of

ACHATINELLA APEXFULVA. 321

Turbo lugubris, prefaced by the statement that Spengler had
obtained several shells in London, which were accompanied
by a note referring to the account in Dixon's Voyage. "This
mirror-smooth shell has six whorls. Its color resembles
that of a mourning garment. It is coal-black, and on the
uppermost whorls snow-white, and so smooth and shining
that I think the shell was polished in the southern lands.
The aperture is almost round. The short, white columella is
somewhat thickened in the middle, as though there was a
tooth there. The interior walls are dirty white. In the Sand-
wich Islands the natives use these snails for decoration or or-
naments to wear on the neck, the ears and the nose. Thus it
is that almost all of this species which come to Europe have
the shell bored for passage of the band for hanging about the
neck or ears.'

Chemnitz's figures excellently represent the typical apex-
fulva, and it is very likely that the specimens were part of
those brought home by Captain Dixon.

Lamarck's description of the same species follows:

"Monodonta seminigra. Shell obliquely conic, imperf or-
ate, rather smooth, black below, white above ; columellar tooth
white ; lip simple.

"Inhabits the Pacific Ocean, on the shores of the island of
Othaiti. My cabinet. The queen of this island used them
for ear rings. The columella is very short. Diameter of the
base S1/^ lines; length iy2 lines'" (Lamarck).

Delessert's figures of Lamarck's type represent entirely
typical apexfulva. Lamarck does not state the source of his
specimens, but Delessert remarks that "il a etc rapporte par
le capitaine Cook.'

Swainson's type of Achatinella pica was obtained by Cap-
tain (afterwards Lord) Byron in Honolulu. The description
follows. " Shell trochiform, black, apex and base of the
pillar white. Shell seven-tenths of an inch long, body whorl
convex, spire conic ; the three upper whorls white or fulvous,
without any convexity, and forming a conic point. Suture
thickened, and margined by a sulcate groove ; a character that
runs through all the following species except A. acuta. In-

322 ACHATINELLA APEXFULVA.

terior of the aperture and base of the pillar white ; the latter
tinged with rose color; margin of the outer lip within bor-
dered with black.'

Swainson's figure is an excellent representation of typical
apexfulva. He states that Dixon 's figures ' ' accurately repre-
sent my A. pica.' He also thinks Monodonta seminigra
Lamarck the same species, but leaves the question open, as it
had not been figured at that time. In his later publication
Swainson places A. pica as a synonym of seminigra.

Key to subspecies and named color-forms of A. apexfulva.

a. Shell plain or with some whitish spiral lines; suture with

a light line or unmarked. Western forms.
b. Black, with contrasting buff summit. Opaeula. A.

apexfulva.

61. Pale flesh-colored. Opaeula. A. a. vespertina.
b2. White, A. a. alba; albino forms of duplocincta and

beata.

a1. Shell conspicuously streaked obliquely. A. a. apicata.
b. Suture with narrow tawny or white margin, apicata

pattern.
61. The same, with white bands below periphery, gulickii

pattern.

b2. Suture with broad white margin, cervixnivea pattern,
a*. Shell spirally banded, not streaked.

b. Chestnut or blackish bands and a wide white subsu-

tural band (also with white and black forms). A. a.

beata.

b1. Banded with pink or sometimes black. A. a. aloha.
b2. Very few brown bands or none, size small, duplocincta

pattern.

It should be noted that melanistic forms of apicata and
beata are not distinguishable from typical apexfulva.

37<z. A. apexfulva vespertina Baldwin. PL 60, figs. 2, 3.

The shell is dextral, usually minutely perforate, solid,
glossy, with the outlines of the spire more or less concave,
apical whorls drawn out, rounded at the tip ; pale flesh color

ACHATINELLA APEXFULVA. 323

or pale salmon, fading upwards to light buff or ivory, some-
times having narrow, inconspicuous pale bands 'on the last
whorl. Sutural band narrow, usually paler than the ground-
color. Lip pale flesh tinted; columellar fold the same or
nearly white, strong; columellar margin raised.

Length 20.3, diam. 14 mm. ; 6y2 whorls.

"Length 21, diam. 14!/2 mm.' (Baldwin).

"Animal when extended in motion, longer than the shell.
Mantle and tentacles brown, the latter with the head above, of
darker shade. Foot light yellow, the superior portion of
darker hue.' (Baldwin.)

Kawailoa (Baldwin) ; a ridge between Waala [Waialua?]
and Kawailoa gulches (Perkins) ; spur in northwestern Opae-
ula, on mokihana leaves (Spalding).

Achatinella vespertiwa BALDWIN, Proc. A. N. S. Phila., 1895,
p. 219, pi. 10, f. 14. — A. apicata var. vespertina Baldwin,
SYKES, Fauna Hawaiiensis, p. 299.

The rather drawn out and flat embryonic whorls are like
typical apexfulva, from which this race differs in color. Fig.
2 is the cotype figured by Baldwin ; fig. 3 a less inflated speci-
men from Newcomb. In the locality where Perkins collected
it, recorded by Sykes, " Waala ' is apparently an error for
Waialua. Mr. Spalding has taken it at Station 14 on the map,
p. 277. The several localities given all mean this one colony, I
believe.

Color-form duplocincta P. & C. PI. 55, figs. 6, 7, 8.

The shell is dextral, white, encircled with two chestnut
bands or groups of lines, one at the periphery, the other below
it; lip faintly violaceous. Length 18, diam. 11 mm. Length
17, diam. 11.7 mm.

The cotypes of this form are 1272, 1273 Cooke coll., 108776
A. N. S., and 1213 Gulick coll., Boston Soc. The former lots
are labelled "Wahiawa, Emerson, extinct?", three banded
specimens, one drawn in fig. 8, and two in which the bands
are very faint, a little stronger near the lip. The locality
seems open to doubt. The Gulick lot is from "Kawailoa, east
side.' There is one banded shell, fig. 7, and one pure white

324 ACHATINELLA APEXFULVA.

except for a very faint tint on the lip, fig. 6. With them are
two small specimens of vespertina color, agreeing with the du-
plocincta in size and shape. It seems likely therefore that
duplocincta is a mutation in a colony of small vespertina. If
not extinct it may be expected to occur in the vicinity of that
race. Whether related to alba I cannot say.

3'< b. A. apexfulva alba Sykes.

"Shell snow-white, save for the peristome being margined
with lilac, similar tinting appearing on the columellar plait
and inside the outer lip at its junction with the body-whorl
of the shell" (Sykes).

Near the head of Kawailoa gulch (Perkins).

Achatinella apicata var. alba SYKES, Fauna Hawaiiensis, p.
299, 1900. Not Achatinella alba Nuttall, = lorata.

This may be a variant of the form I have called A. a. beata;
but if the locality given is correct, it is probably a parallel
modification rather than the same form. Kawailoa is properly
speaking the gulch of the Anahulu river, west of Opaeula,
while the ridge occupied by beata is a considerable distance
east of Opaeula. If really from Kawailoa, A. a. alba is at the
extreme western border of the apexfulva area.

37c. A. apexfulva apicata 'Newc.' Pfeiffer. PL 60, figs. 4 to
Ib.

"Shell imperf orate, globose-conic, solid, smooth, blackish or
brown, irregularly pallidly radiate and subfasciate ; spire
conic, apex cinnamon or liver colored, rather acute. Whorls
6, the upper ones flat, the penultimate convex, last whorl glo-
bose, scarcely % the length of the shell. Aperture oblique,
subtetragonal-oval. Columellar fold high, tooth-like, lilac
colored. Peristome unexpanded, lipped with lilac within, the
columellar margin very much dilated, thickened and ad-
nate. Length 18%, diam. 12 mm. ; aperture 9 mm. long, 5
wide. Habitat: Sandwich Islands (P/r.).

Kalaikoa to Opaeula.

Achatinella apicata Newc., PFR., P. Z. S., 1855, p. 210;
Monographia, iv, 542. — A. swiftii var. NEWCOMB, Ann. Lye.

ACHATINELLA APEXFULVA. 325

N. H. of N. Y., vi, p. 325. — Apex gulickii SMITH,, P. Z. S.,
1873, p. 78, pi. 9, f. 19 (not f. 17).— Apex lilaceus GULICK,
P. Z. S., 1873. p. 79, pi. 10, f. 4.—Achatinella aptycha Pfr.,
LYONS, Hawaiian Almanac for 1892, pi. 12, f. 32.

Typical apicata is figured on pi. 60, figs. 4 to 4c. A lot re-
ceived from Newcomb in 1856 agrees fully with a large series
in the Gulick collection from "Wahiawa.' The shells are
dextral in large majority. The embryonic whorls are flattened
and more or less nipple-like, as in apexfulva, and usually are
between ochraceous orange and ochraceous tawny in color.
The later whorls are closely, unevenly streaked in varying
shades — Prussian red, dark livid slate purple or dull purplish
black, leaving narrower streaks of the ivory or ochraceous
buff ground. Sometimes darker or lighter spirals are faintly
visible, and rarely an indistinct lighter band at the periphery.
The suture usually has a narrow rufous border, rarely white.
Occasional shells are warm blackish brown without streaks.
Lip fleshy tinted. Except by the usually darker tint of the
apex, these dark shells closely resemble typical A. apexfulva.
Length 20, diam. 13.8 mm. or somewhat smaller.

There is also a large set from Wahiawa collected by Gulick
having the typical whitish streaking on a blackish purple
("taupe brown") ground, slightly darker than fig. 4c, the
embryonic whorls ivory yellow, nearly white.

A large lot collected by Dr. Cooke "on low ridges and val-
leys, Wahiawa' consists of dextral and sinistral shells in
about equal numbers. The blackish and the blue-streaked
forms figured from Kaukinehua are present, sutural band
hazel to orange or rarely white ; apex light ocher or cream
color. Other shells are vinaceous gray with whitish streaks
and bands. Others of various shades of dull brown, with dark
spiral lines and whitish bands (gulickii pattern), or not
banded (pi. 60, figs. 5 to 55).

The localities of the lots described above are unfortunately
rather indefinite, as "Wahiawa' may be anywhere between
Poamoho and Waikakalaua.

In north Kaukinehua near the Ahonui ridge, Mr. Spalding
collected from the "burnt forest" in a colony (no. 574) where

326 ACHATINELLA APEXFULVA.

the apex is brown or ivory-white, coloring various; typically
streaked, banded with white (gulickn pattern), or nearly
white, lip livid pink. PL 60, figs. 6, 6a.

Further west, near the Kaukinehua-Poamoho ridge, Mr.
Spalding collected apicata both dextral and sinistral, varying
from nearly black through intermediate shades to slate blue
or Dutch blue streaked with white, sutural band hazel, orange
or chestnut (pi. 60, figs. 7, la, Station 8, in the western ravines
of Kaukinehua) ; also the gulickii pattern, suture pale brown.
It occurred also over the ridge on the Poamoho side, east of
Nicol's camp, Station 9 of the map, pi. 60, fig. Ib from Spald-
ing's no. 317. The gulickii pattern with a pale brown suture
also occurs here.

It will be noted that these streaked forms are not from the
ridges but from the lower slopes. It is likely that the Gulick
and Newcomb apicata of "Wahiawa" were from still lower
forests than those now existing.

In Ahonui and Kalaikoa Gulick found typical apicata, like
those from Wahiawa, and two forms ranked by him as species
(gulickii Sm. and lilacea Gul.), but which I look upon as
merely the locally dominant color-forms of apicata with which
they intergrade in a small number of specimens. This blend-
ing of lilacea into gulickii and of both into typical Wahiawa
apicata is fully established by the abundant suites before me
collected by Gulick himself. Moreover, occasional specimens
of exactly the same patterns may be found among apicata in
the colonies of "Wahiawa" and Kaukinehua.

The color-pattern called A. gulickii Smith is figured on pi.
60, fig. 12, type ; figs. 10, 10a, 13, topotypes, all from Kalaikoa ;
fig. 11, from Ahonui, all collected by Mr. Gulick. The streaks
vary from dark slate purple to vinaceous drab, and are inter-
rupted on the last whorl by a white zone below the periphery
and numerous whitish lines on the base. Typically it has a
tawny sutural border, and an ochraceous embryonic shell,
fading towards the apex. The lighter colored examples have
the sutural band buff or indistinct, and the embryonic whorls
pale ; the spirals on the last whorl may be reduced to one white
line below the periphery, as in fig. 13. The special feature of

ACHATINELLA APEXFULVA. 327

gulickii is the presence of a subperipheral light belt, which is
seen in various stages of development in a large proportion of
the shells (but by no means all), of the original lots. The
type-specimen, fig. 12, is one of the shortest in Mr. Gulick's
lots. Most shells are like fig. 10a in contour. The same color
pattern occurs in Wahiawa, figs. 5&, 6, 6a. The original de-
scription follows.

4 i

'Apex gulickii Sm. Shell sinistral (sometimes dextral),
globose-conic, glossy, striated with growth (but scarcely with
spiral) lines; dilute brown, streaked with ashy, encircled with
a white zone at the periphery; suture margined with brown;
spire slightly concave ; whorls 6, the first 3 or 4 pallid, a little
convex, the rest convex. Aperture white, receding ; peristome
a little dilated, thickened within, tinted with dilute rose-
brown; columellar fold moderately strong, whitish. Length
19, diam. 13 mm.

"Var. Shell dextral; first 3 whorls pallid, the following
two and the last whorl having the upper half streaked with
cinereous-brown, the lower half yellow, zoned with cinereous-
brown.

* ' Station : On trees and bushes. Habitat : this species is
most abundant in Kalaikoa and Ahonui, but is sometimes
found in valleys to the west as far as Waialei. Affinities : It
is allied to A. apicatus Nwc. and A. lilaceus Gk. Remarks:
In Ahonui, two-thirds of the specimens are sinistral. In Ka-
laikoa only about one-eighth are sinistral. The specimen fig-
ured is from Kalaikoa' (Smith}.

The variety of gulickii described by Mr. Smith has the epi-
dermis yellow below the periphery, markings otherwise as in
the gulickii pattern. The shells are rather small and thin,
mainly dextral. This form, of which I figure specimens on pi.
58, figs. 13-13&, is really a form of A. swiftii and not closely
related to gulickii. The statement that this variety occurs
"west as far as Waialei' is incorrect, as the Waialee form
with yellow base is directly connected with A. valida cinerosa,
and in my opinion is not the same as the various yellow based
forms of A. swiftii.

The pattern called A. lilaceus (pi. 60, figs. 14, 14a, topo-
types from Kalaikoa, Gulick coll.) has light brownish vina-
ceous and russet vinaceous streaks, without white bands, the

328 ACHATINELLA APEXFULVA.

narrow sutural margin whitish, the embryonic whorls ivory
yellow. The specimens from Ahonui and Kalaikoa are alike
in pattern. The original description follows.

"Apex lilaceus Gk. Shell dextral, globose-conic, glossy,
striated with growth but hardly with spiral lines; lilac-fawn
colored, streaked with whitish ; suture margined with paler ;
whorls 6, the first three nearly flat, white, the rest convex;
last whorl sometimes obscurely, obtusely angular; aperture
whitish ; peristome dilute rose-brown, very little dilated, thick-
ened within ; columellar fold strong, pale rose. Length 21,
diam. 12^ mm.

* ' Station : On trees. Habitat : The metropolis of the species
is Ahonui, on Oahu. It is also abundant in Kalaikoa, but be-
comes very rare in Wahiawa, Helemano and Kawailoa. Af-
finities : It is readily distinguished from A. apicatus Nwc. by
the color of the apex, which is chestnut in that species, but
white in this. Remarks : Sinistral specimens are very rare.
The specimen figured is from Kalaikoa' (Gulick).

This form is fully connected with the gulickii pattern.

A. a. apicata occurs on the Poamoho-Helemano ridge in sev-
eral forms, the shells either dextral or sinistral in the same
colonies. A lot in the Gulick collection labeled "Helemano'
contains shells with tawny suture, similar to pi. 60, figs. 4c,
others with narrowly white-margined suture (like pi. 60,
fig. 4&), and about half the lot has a wide white band below
the suture, like pi. 60, fig. 9. In all of these the streaks may
vary from walnut brown to slate blue, in various nuances.

On the slope of a spur running into lower Poamoho, Mr.
Spalding collected typical apicata, chiefly with the suture
dark-bordered, but with them the form with broad white su-
tural band and white embryonic whorls ; the streaks slate-blue
(pi. 60, fig. 9, from no. 2102).

Further up on the Helemano-Poamoho division ridge the
same form occurs. The streaks are more or less blended, color
varying from dull violet black to vinaceous drab. The su-
tural border is wide and snow-white ; embryonic whorls cinna-
mon to nearly white, pi. 60, figs. 8, 8a, coll. by Spalding.
These shells with a broad white sutural band are an incipient
race, in most places not extricated from the dark-sutured

ACHATINELLA APEXFULVA. 329

colonies in which the mutation arose. They may conveniently
be referred to as the ll cervixnivea pattern.' This pattern is
characteristic of the Poamoho-Helemano ridge and its spurs.
I suppose that the hybrid apicata X cervixnivea colonies are
chiefly lower than the pure cervixnivea. According to a note
made from Mr. Spalding's collection his no. 3712, from the
central ridge of Kalaikoa is an apicata X cervixnivea mix-
ture ; 3 dextral, 12 sinistral shells.

37d. A. APEXFULVA BEATA P. & C., n. subsp. PI. 60, figs. 17-
17c ; pi. 55, fig. 5.

The shell is dextral, solid, with the embryonic whorls ochra-
ceous-buff (or in white shells, light buff). Subsequent whorls
are never streaked, but vary in color as follows :

Fig. 5. Pure white, peristome white, embryonic whorls
faintly cartridge buff.

Figs. 17a, 17&. White with several deep chestnut or black-
ish bands, of which one ascends the spire ; last whorl with the
band-intervals somewhat violaceous, or the bands confluent;
a wide sutural border snow-white ; embryonic whorls buff.

Fig. 17c. Light buff with chestnut bands, lip pale lilac.

Fig. 17. Last 2% whorls black with a white sutural line.
This is the typical apexfulva pattern.

Length 19.6, diam. 12.5 mm. • 6~y2 whorls.

Crest of the Poamoho-Helemano division ridge, types no.
108809 A. N. S. P., from no. 3430-31 of Mr. Spalding's col-
lection.

This splendid polychromatic race inhabits a limited area on
the ridge, at higher elevations than the cervixnivea pattern,
of which beata is merely a specialized, banded form. In fact,
the main feature distinguishing it from the cervixnivea pat-
tern of apicata is that there are no streaks, but the color when
present is in bands, which are a little diffuse at their edges
when not lost in a general melanism. I imagine that while
the beata has been evolved parallel to A. a. aloha, the two are
not directly connected. Mr. Spalding's no. 3883, from "Hele-
mano, the second ravine above the intersection with Poa-
moho," is probably part of or near the typical colony of

330 ACHATINELLA APEXFULVA.

leata, which includes his no. 3930, 3931, 3853, 3854 ; the last
from far up and upon the ridge, at 12 on the map, p. 277.
The first locality is on the lower part of the same ridge.

37e. A. APEXFULVA ALOHA P. & C., n. subsp. PI. 60, figs. 15,
15a, 16.

The shell is dextral, rather small and light, often perforate ;
white, with unequal spiral bands of pale cinnamon pink on
the penultimate and last whorls, deepening to brownish vina-
ceous or orange-cinnamon behind the lip, where they usually
become confluent. The embryonic whorls and a broad band
below the suture are white. Peristome moderately thickened
within, light purplish vinaceous, the columellar fold paler.

Length 18.5, diam. 12.7 mm. ; 6% whorls.

Length 19.2, diam. 11.9 mm. ; 6% whorls.

Crest of the division ridge between the two branches of the
Kaukinehua stream, above the Wahiawa head-gates cabin, the
colony extending to within % mile of main ridge ; on moki-
haua, pua, maile and alani. Cotypes in collections A. N. S. P.
and Bishop Mus., from no. 3818 Irwin Spalding coll.

By its cleanly-defined pinkish bands, absence of streaks and
white embryo, this snail is well distinguished from other
forms of A. apexfulva. It is very constant in a large series
collected by Mr. Spalding, except for a mutation which occurs
in the same colony. This is illustrated in pi. 60, fig. 16, and
differs from normal A. a. aloha only by having the bands
chocolate-black.

A. a. aloha is always dextral. It is plentiful in a limited
locality on one ridge, which is isolated by perennial streams
on both sides, and mauka is shut in by the precipitous side of
the main Koolau range. This station is marked 10 on the map,
p. 277.

Mr. Spalding thinks that connecting forms with apicata
may perhaps turn up on the upper Helemano-Poamoho ridge,
when it comes to be explored. At present, A. a. aloha is quite
isolated. It is a much less solid shell than A. a. vespertina,
which is not nearly related.

ACHATINELLA DECORA. 331

38. A. DECORA (Ferussac). PI. 61, figs. 1 to 6.

The shell is imperforate, sinistral, solid, ovate-conic, outlines
of spire straight or slightly concave. Suture superficial, with
the usual impressed line defining a margin, bordered below by
a white band. Embryonic whorls cream color (varying from
almost white to light ochraceous buff), with the tip often a
little darker ; first neanic whorl often blotched with chocolate,
or with a chocolate band, extending as far as the last whorl.
Last whorl variously marked. (1) Copiously streaked with
chestnut, bay or blackish chocolate on an ochraceous buff
ground. (2) The same, with a light subperipheral band. (3)
Lightly streaked with chestnut over ochraceous buff, with dark
bands below the white sutural band, at the periphery and
around the columella, or otherwise placed. (4) Cream or
ochraceous buff, not streaked, having a few narrow dark
bands, lip nearly white with yellowish edge.

Surface weakly striate, not highly polished. Aperture
bluish white within, the lip expanding slightly, well thickened
within, and pale flesh color. Columellar fold moderate, nearly
white.

Length 21.3, diam. 13, aperture 11 mm. ; 6% whorls.

Length 19, diam. 13, aperture 10.3 mm. ; 6% whorls.

Length 17.2, diam. 12.8, aperture 9.7 mm. ; 6 whorls.

Length 20, diam. 12, aperture 9.5 mm. ; 6% whorls.

Oahu : Kawailoa and Wahiawa (Gulick) . Helemano ; Gulch
east of Opaeula; gulch west of Helemano; and eastern spurs
of Kawaiholona, on kukui trees (Irwin Spalding).

Helix decora FER., Tableaux Systematiques des Animaux
Mollusques, etc., p. 56, no. 430 (1821), based solely upon
Turbo lugubris sinistrorsus Chemnitz, Conchylien Cabinet, xi,
p. 307, pi. 213, f. 3014, 3015. — Achatinella decora var. &, PFR.,
Monogr., iii, p. 465. — Apex decorus Fer., GULICK, P. Z. S.,
1873, p. 82 (under A. tumefactus). — Achatinella decora Fer.,
SYKES, Fauna Hawaiiensis, p. 301, 1900. — Not A. decora
Reeve, or of most writers and collectors.

Achatinella perversa SWAINSON, Quarterly Journal of Sci.,
Lit. and Art, iii, 1828, p. 84; Zoological Illustrations, iii, p.

332 ACHATINELLA DECORA.

99, pi. 99, f. 2. — NEWCOMB, Ann. Lye. Nat. Hist, of N. Y., vi,
1858, p. 309 (animal). — THWING, Orig. Descript. Achatinella,
pi. 1, f. 8. — Achatinella quernea FRICK in coll., according to
J. S. Emerson. — Apex tumefactus GULICK, P. Z. S., 1873, p.
82, pi. 9, f. 20.

The shells obtained by Gulick and Newcomb are more var-
iable in shape than those found now. It seems to have been a
common species sixty years ago. PL 61, figs. 1 to 2e represent
specimens from the Gulick collection from ' * Kawailoa, ' ' which
we take to be the type locality. Whether this form with wide
sutural band still exists there is not known. Probably the
species ranged farther southeastward in the old days when
Gulick obtained his "Wahiawa" lot than it does at present.

Mr. Spalding's several localities are near together, or pos-
sibly parts of one or two colonies, in the back Kawailoa-Hele-
mano country.

The shells are rather long, often subperforate, the lip very
pale, ground color light buff (or sometimes ochraceous buff)
with faint or distinct darker streaks, which are usually
straight, sometimes beautifully zigzag. There are dark bands
in some lots. The white sutural band is narrow or reduced to
a line, rarely even wanting. See pi. 61, figs. 4, 4a, gulch west
of Helemano ; figs. 5 to 5&, eastern spurs of Kawaiholona ; fig.
6, gulch east of Opaeula, where also banded shells occur.

Further east, in "Wahiawa", though just where is uncer-
tain, the species becomes somewhat smaller and distinctly
though minutely dark-tipped, the apex being lead color or
vinaceous gray. The shell varies from usual decora patterns
to white with brown bands below the suture, at the periphery
and around the columella. Mr. Gulick selected one of the ex-
tremely light shells as his type of A. tumefactus, but in the
large series of his collection this very light form is unusual.
It connects with the ordinary dark pattern through many in-
termediate stages, some of which are figured, all from one lot
from "Wahiawa." PL 62, figs. 12 to 20; fig. 18 being the
typical form of tumefactus.

Length 18.6, diam. 12 mm.

Length 16.5, diam. 11.2 mm.

ACHATINELLA DECORA. 333

I do not know that tumefacta still exists. It was evidently
abundant when Gulick collected. There is also a good series
in coll. C. M. Cooke, taken by Mr. Emerson at about the same
time. PL 61, fig. 3 is decora from "Wahiawa," Cooke coll.

A. decora has companions in A. valida Pfr., from further
west, and in A. mustelina of the Waianae range. The latter
was, in my opinion, derived from an old decora stock which
migrated southward. Some Waianae shells are hardly dis-
tinguishable from main range forms, but in general the muste-
lina variations are different.

Ferussac in his first publication based Helix decora on the
figures in Chemnitz. He seems to have had another shell
which he described later (1824) in the Voyage of the Uranie.
This later decora was apparently the banded species subse-
quently called decora by Pfeiffer, Reeve and nearly all later
authors and collectors. It is the copiously banded form of
simulans well known from the western ridge of Nuuanu.

The original decora is, as Gulick recognized in 1873, a short,
dark form of the shell better known as A. perversa. This form
was not uncommon in the old days, in colonies mainly com-
posed of longer, more variegated shells. Figures Ib, 2c of pi.
61 represents "Kawailoa" specimens of decora agreeing with
Chemnitz 's figures. Chemnitz gave the following description :
' The shell is smooth as a mirror and brown-blackish colored.
Near the suture the whorls are very prettily encircled with a
white band. The apex is white. This rare sinistral snail, for
which Herr Spengler had to pay two guineas in London, lives
in fresh water of the Sandwich Islands in the South Sea. The
inner walls are whitish. As the inhabitants of these islands
are accustomed to wear this kind of snail as neck and ear
adornment, and therefore bore the shells in order to string
them, they made no exception of these rare sinistral snails,
which are likewise bored. '

The type of A. decora in the Spengler collection was prob-
ably brought to London by the expedition of Captain Dixon.

The species was subsequently described as A. perversa and
well figured by Swainson, whose shells were from a lei brought
in England by Captain Byron. The shells on this lei

334 ACHATINELLA VALIDA.

indicate that it was made in Kawailoa district. Swainson's
type was a larger, more variegated shell than that figured by
Chemnitz, and more fairly representative of the species. The
description of A. perversa follows. " Shell reversed, subtro-
chiform, fulvous brown with darker transverse bands and lon-
gitudinal lines; apex and suture white ' (Swainson).

The original description of A. tumefactus follows.

"Apex tumefactus Gk. Shell perforate, sinistral, globose-
conic, glossy, striated with growth and indistinct spiral lines ;
white, encircled with a few olivaceous-brown zones; suture
margined ; spire slightly concave ; whorls 6, the first blackish,
first 3 or 4 a little convex, the rest convex ; aperture receding,
white ; peristome dilated, thickened within ; columellar fold
strong, white. Length 19, diam. 12 mm.

"Var. a. Shell streaked with olive-brown, interrupted by
two or three white zones ; suture white margined.

"Var. ~b. Shell streaked with fawn, black or brown, inter-
rupted by a wide white zone at the periphery; suture white
margined.

' ' Station. On trees and shrubs. Habitat : Wahiawa ; rarely
in Helemano. Affinities: It is related to A. decorus Fer.
(syn. A. perversus Swn.), which is found in Helemano and
Kawailoa. It is distinguished by its smaller size, its con-
cavely conical spire, and its paler color, which is differently
distributed. The three upper whorls of A. decorus are of a
yellowish tinge ; in this species they are white, terminated
with a black dot, which is not found in the typical forms of
the other species. Remarks : This species is never dextral.
The shell figured is from Wahiawa'1 (Gulick}.

39. A. VALIDA Pfeiffer. PI. 30, fig. 24 ; pi. 52, figs. 15 to

"Shell imperf orate, dextral (more rarely sinistral), ovate-
conic, solid, smoothish, ashy-brown, somewhat banded with a
pale tint and streaked with black. Spire elevated, conic, the
apex brown, acute ; suture margined, white. Whorls 6, the
first three flat, those following a little convex, the last whorl
about two-fifths the length, inflated. Aperture oblique, trun-
cate-auriform. Columellar fold above, nodiform, a little
twisted; peristome brown-bordered, the right margin nearly
unexpanded, somewhat straightened ; columellar margin di-

ACHATINELLA VALIDA. 335

lated, appressed. Length 21%, diam. 13mm. ; aperture 10%
mm. long, 5% wide within.

" Var. &. Black, pallidly, interruptedly lineolate or banded.
Inhabits the Sandwich Islands, Frick, in Mus. Coming '
(Pfr.).

Northwestern Oahu : Pupukea, type locality ; varieties to
Kahuku.

Achatinella valida PFR., P. Z. S., 1855, p. 6, pi. 30, f. 24;
Malak. BL, 1855, p. 69 ; Monographia Hel. Viv., iv, 540.

Pfeiffer's type figure is exactly reproduced in pi. 30, fig. 24,
and specimens from a large series, collected by Gulick, are fig-
ured on pi. 52, figs. 15-15e, from Pupukea, which may be ac-
cepted as type locality. Out of 62 shells in this set, 36 are
dextral. The embryonic whorls form a more narrowly conic
summit than in species of the turgida-swiftii-apexfulva
series, of a dull red color, varying in shade between claret
brown and ocher red. The last embryonic whorl is overlaid
with whitish, also the first whorl, the tip usually being almost
white * The spire is straightly conic. The last whorl has a
cinnamon ground, more or less profusely streaked and spirally
banded or lineate with brownish Hack, or rarely brown. A
sutural band is white, and rarely there is a white line or band
at the periphery. The cinnamon ground fades on the pen-
ultimate whorl to whitish on those above. The aperture is
white, outer lip faintly brown-tinted.

A. valida is apparently quite distinct from all forms of the
interior watershed of the main range. Except in the direction
of coil, the large series seen is unusually homogeneous. The
broad-banded form, fig. 15a, and that without black markings,
fig. 15&, are very unusual patterns.

In a wider sense, A. valida includes the forms described as
leucophcea and leucozona Gulick ; yet as the several races occur
in pure colonies so far as we know, they may stand as sub-
species. On the extreme limits of their variation are a few
more or less intermediate specimens. Further exploitation of
the ridges would probably diminish the distinctions. 372
shells of all the races are before me.

336 ACHATINELLA VALIDA.

390. A. VALIDA LEUCOPH/EA (Gulick). PI. 55, figs, 19 to 22.

"Shell sinistral, subperf orate, ovate-conic, little shining,
indistinctly, very finely striated with growth-lines and trans-
versely; dilute fawn, streaked and zoned with pale brown;
suture distinctly margined with white. Whorls 7, subconvex.
Aperture white ; peristome a little dilated, strongly thickened
within, tinted with pale brown. Columellar fold strong, whit-
ish, slightly reflected, nearly covering the small crevice.
Length 22, diam. 12 mm.

"It is allied to A. perversus Swn. audA, validus Pfr., but it
is of a paler color and more elongate form. It is always sinis-
tral" (Gulick).

Waialee (Gulick).

Apex leucophceus GULICK, P. Z. S., 1873, p. 82, pi. 9, f . 16.

In a series of 105 specimens, all are sinistral and most are
imperf orate. The embryonic whorls are ochraceous or light
brown, darkest near the suture below, fading upwards, the
tip dilute brown. The later whorls have a light pinkish cin-
namon to cartridge buff ground, weakly marked with cinna-
mon streaks and spiral lines or narrow bands. The very nar-
row sutural border is pale. The lip has a faint fleshy brown
or cream tint. The spire is straightly conic and typically
rather long. A small proportion of specimens are indistin-
guishable in shape and color from the lighter forms of Pupu-
kea valida.

Length 22, diam. 12.8 mm. ; 6% whorls.

Length 19, diam. 12.3 mm. )

' \ extremes of shape.

Length 19, diam. 9.1 mm. j

396. A. VALIDA CINEBOSA Pfeiffer. PI. 30, fig. 5; pi. 55, figs.
9 to 18.

Shell dextral, imperforate, ovate-conic, solid, nearly smooth,
glossy ; brown-gray, radiated with brown and white lines.
Spire conic, apex rather acute, white ; suture somewhat crenu-
late, white-bordered ; whorls 6, a trifle convex, the last a little
shorter than the spire, rotund at base. Aperture a little ob-
lique, truncate-auriform ; columellar fold twisted, strong ;
peristome thickly white-lipped, the right margin somewhat

ACHATINELLA VALIDA. 337

straightened, narrowly expanded ; coluniellar margin thick,
adnate. Length 21, diam. llmm. ; aperture 10 mm. long, 4^2
wide. Inhabits the Sandwich Islands, Frick (Pfr.}.

i Var. b. Fulvous, radiated with brown, suture of the same
color" (Pfr.).

Oahu: Waialee (Gulick).

Achatinella cinerosa PFR., Proc. Zool. Soc., 1855, p. 2, pi. 30,
f . 5 ; Malak. BL, 1855, p. 5 ; Monographia Hel. Viv., iv, 519.—
[A. decora] var. cinerosa Pfr., SYKES, Fauna Hawaiiensis, p.
302. — Apex leucozonus GULICK, P. Z. S., 1873, p. 83, pi. 10,

'' . ... .

This subspecies is like valida in the straightly conic shape,

white subsutural band and streaked coloring. It differs from
valida chiefly by the color of the embryonic shell, which varies
from ivory to light ochraceous buff with a white band below
the suture ; the tip white or a trifle grayish. The last two or
three whorls are cinnamon with snuff-brown to sepia streaks
(rarely walnut brown), in many specimens interrupted by a
white peripheral band ; suture bordered with a white band.
Rarely there are several or numerous bands, figs. 9, 14, illus-
trating these unusual patterns. In a few shells the streaks
coalesce, producing an even chocolate shade, with the usual
light peripheral and sutural bands. In a few specimens the
cuticle is yellow below the periphery (fig. 10). Some narrow
shells of this kind were erroneously identified by Gulick as
A. napus Pfr., but that is really a Waianae shell. In several
very old shells there is a strong cord across the parietal wall.
The color and shape variations are well connected by inter-
mediate shells, in the lot of 178 before me, even the yellow-
base form, which seems to be the only significant color-muta-
tion in the lot, connects by transitional shells with the others.
Gulick states that it is always dextral, but I found three sin-
istral shells by carefully looking over the lot in A. N. S. col-
lected by him.

The ordinary size is length 19, diam. 11.7 mm., but it may
be smaller and narrower, length 17.2, diam. 10.4 mm.

The specimens from Wahiawa noted in Mr. Gulick 's re-
marks under A. leucozonus, belong, in our opinion, to A.
swiftii.

338 ACHATINELLA VALIDA.

It happened that the same streaked pattern of what seems
to have been a common Waialee shell served as the types of
cinerosa and leucozonus, so that the two names are exactly
synonymous. The original description of leucozonus follows.
The original figure is like my pi. 55, fig. 12.

"Apex leucozonus Gulick. Shell subperf orate, dextral, glo-
bose-conic, glossy, striated with incremental (but scarcely
spiral) lines; gray-brown, streaked with deep gray-brown, at
the periphery interrupted by a white zone ; suture distinctly
margined with white. Spire straightly conic. Whorls 6, the
first three a little convex, the rest convex. Aperture white;
peristome a little dilated, tinted with pale rose-brown, thick-
ened within ; columellar fold strong, white. Length 19, diam.
12 mm.

"The metropolis of this species is Waialei. A few speci-
mens have also been found in Wahiawa, Island of Oahu. It is
related to and passes into A. napus Pfr., which is also found
in Waialei ; but the usual forms are distinguished by having a
shorter spire and a white suture, and in being free from the
yellowish hue of that species. This species is always dextral.
The specimen figured is from Waialei' (Gulick).

A rather peculiar form of cinerosa (pi. 55, figs. 17, 18) was
noticed by Mr. Gulick under his description of A. leucorraphe :
"a variety with spire regularly conical is reported to have
been found in Waimea.' This shell is certainly a form of
cinerosa (leucozona), and not of leucorraphe. The embryo is
shaped like that of valida, ivory or buff with a fleshy tip ; last
whorl light pinkish cinnamon to pallid purplish gray, nar-
rowly and closely streaked with brownish or slaty shades, usu-
ally with a few inconspicuous dark lines around the base. The
suture has a wide white border, but there is no peripheral
band. Lip white. All of a series of 21 are dextral. The local-
ity Waimea is marked with a query in Mr. Gulick 's collection,
but no doubt the form came from somewhere in the northwest,
and may have been from Waimea.

39c. A. VALIDA KAHUKUENSIS P. £ C., ii. subsp. PL 52, figs.

17, 17a.
The shell is dextral, white above, yellow below the periphery,

WAIANAE SPECIES. 339

usually encircled with a black-brown line at the junction of
the two ground-tints, and often there are several additional
lines widely spaced on the base or sometimes above. A faint
sutural line may usually be discerned. Embryonic whorls
when unworn are cartridge buff, slightly darker near the
sutures, but not at the tip. The outlines of the spire are per-
ceptibly concave, the last whorl swollen. Lip moderately
thickened, white ; columella very faintly rose-purple.

Length 20, diam. 13 mm. ; 61/2 whorls.

Length 18.7, diam. 12.2 mm. ; 6y2 whorls.

Oahu: Kahuku, at an elevation of 1,500 to 1,750 ft. (L. A.
Thurston). Cotypes in coll. A. N. S. and Bishop Mus., from
Mr. Thurston 's collection.

This race is one of the most attractive of the ''Apex''
group. While very distinct in appearance, its kinship with the
valida series is shown by the rare yellow-based form of cine-
rosa, which however does not have the turgida-like shape of
the Kahuku form.

Mr. Spalding informs me that there are black and white
mutations of this Kahuku race.

Species from the Waianae Range.

Affinities and origin. — The species A. mustelina is now con-
sidered by Hawaiian conchologists to include all the Waian-
aean forms of the typical section of Achatinella (Apex), with
the possible exception of A. concavospira. In this wide sense
the species inhabits the whole range, where conditions are suit-
able. It is often very abundant. It is related to A. decora
Fer. and A. valida Pfr. of the Main Range. Some specimens
of decora are hardly separable from mustelina, but in the
main, the variations of the two differ rather widely. A. decora,
valida and mustelina appear to be slightly differentiated forms
of a common ancestral species which lived in the western val-
leys of the Main range of Oahu. The migration to the Waia-
nae range may have begun in a late stage of the Pliocene, but
more likely in the Pleistocene. Up to historic time forests ex-
tended from range to range, and Amastra, Pterodiscus and
other forest snails of the two ranges mingled. The more an-

340 WAIANAE SPECIES.

cient Waianae fauna probably had 110 arboreal Achatinellae ;
at least we have no evidence of any.

Color and form characteristics and distribution. — On the
north side of the range the shells are either blackish-brown
with a white subsutural baud (mustelina pattern), or of var-
ious shades of drab or vinaceous buff, more or less streaked,
with or without spiral lines or bands (multilineata pattern).
My very limited experience in the Waianae range leads me
to believe that as a general rule the dark forms are lower, the
light and banded higher on the ridges, although the transition
is gradual, and probably no colony of dark shells is pure. In
the west (Mokuleia, etc.), the white sutural band is quite
narrow. Eastward it becomes broader, culminating in the
bicolor pattern of Lihue. In the middle section of the range
(Haleauau, Popouwela) the white sutural band is intermed-
iate in width. Properly speaking, no line can be drawn be-
tween typical mustelina and bicolor-, and both, in their re-
spective districts, merge directly into multilineata-monacha
and bandless patterns of various tints.

It appears thus that there has been some differentiation
vertically, and more horizontally, along the range ; but the
continuity of the forests has favored migration, so that from
the systematic standpoint, bicolor, multilineata and monacha
cannot well be defined as races distinct from mustelina, though
in dealing with particular colonies some subvarietal nomencla-
ture may eventually be needed.

A. mustelina with its several forms already mentioned does
not extend southeastwardly beyond Lihue. From there to the
end of the range it is replaced high on the ridge by two quite
diverse races : A. m. lymaniana, an invariably sinistral shell,
which connects with mustelina through the forms called sor-
dida and napus; and A. concavospira, a dextral form of rather
lighter structure, with the outlines of the spire usually more
concave.

A. concavospira is the terminal member of the series south-
eastward ; and although in some colonies there is an approach
to the mustelina (multilineata or napus} shape and coloring,
yet the differentiation as typically developed may be sufficient

WA1ANAE SPECIES.

341

ground for allowing it specific rank, as a working arrange-
ment. A. concavospira does not seem directly related to
lymaniatia, which inhabits the same district, but rather to the

SKBTCH OF WAIANAE RANGE, THE PRINCIPAL RIDGES REPRESENTED BY LINES.

bicolor stock. The area of concavospira and lymaniana forms
a long and narrow extension of the tree-shell country, which
broadens out from Lihue westward.

342 ACHATINELLA MUSTELINA.

On the south or Ocean side the range falls so precipitously
from the narrow crest that forest is lacking for the greater
part. Where the crest is wider, or butresses extend ocean-
ward, there is some shell country on the Ocean side of the
summit. I understand that this is the case in the Palihua
region, which I did not visit. Otherwise, the only tree-snail
forests on the south side are in the heads of the valleys under
the butresses and peaks. In the head of Makaha there is a
modified form of the Mokuleia mustelina, and a peculiar
special race, A. m. makahacnsis. In the head of Waianae val-
ley, under Kaala, I am informed that mustelina occurs.

Further east the valleys of the south side look very arid,
and no tree snails have been reported. The forms from high
on the ridge, lymaniana and concavospira, have already been
noticed.

The distribution of the several races is approximately indi-
cated on the accompanying map.

40. A. MUSTELINA Mighels. PI. 63 ; pi. 62, figs. 1, 2.

"Shell dextral, conical, dark brown with a light revolving
band at the suture, perforate ; whorls 7 ; convex ; aperture ob-
long; lip simple, acute. Length 1 inch, diameter nine-twen-
tieths inch. Rob. Waianai' (Mighels).

Oahu : Entire Waianae range, in various varieties.

Achatinella mustelina MIGHELS, Proc. Boston Soc. N. H., ii,
1845, p. 21. — REEVE, Conch. Icon., vi, pi. 3, f. 21. — PFB.,
Monogr., iii, 458. — THWING, Orig. Descript. Achatinella, pi. 1,
f. 9. — Apex mustellina Migh., HARTMAN, Proc. A. N. S. Phila.,
1888, p. 39. — ? Achatinella vestita MIGHELS, Proc. Bost. Soc.
N. H., ii, 1845, p. 20. — Achatinella multilineata NEWC., P. Z.
S., 1853, p. 138, pi. 22, f . 23. — B [ulimella] multilineata Newc.,
HARTMAN, Proc. A. N. S. Phila., 1888, p. 30. — Achatinella
monacha PFR., P. Z. S., 1855, p. 3, pi. 30, f. 9 ; Monographia,
iv, 522. — Achatinella monarcha Pfeiffer, THWING, Orig. De-
script., p. 186.

The chief races and color-patterns of A. mustelina are as
follows :

Form rather stout ; spire with slightly concave or straight
outlines.

ACHATINELLA MUSTELINA. 343

Pinkish, sutural band white, makahaensis.

Carob-broAvn to ashen, with a moderate or narrow sutural
white band, mustelina pattern.

Many dark bands on a paler ground, multilineata pattern.

Upper third of last whorl white, lower part blackish-
brown, bicolor.
Form more slender, spire straightly conic.

Fawn to white, usually banded, white below suture, sor-
dida.

Carob-brown with speckled streaks of white ; often
banded ; sometimes ochraceous or whitish ; sutural
white band usually linear; sinistral, lymaniana.

The original descriptions of synonyms of mustelina follow.

A. multilineata Newcomb. PL 29, fig. 23, reproduction of
the original figure. "Shell dextral or sinistral, solid, elon-
gately conical ; whorls 6, rounded, margined above ; lip ex-
panded below, and slightly subreflected, above acute and
thickened within; columella short, stout, slightly twisted,
with a callus spread over and nearly closing the umbilicus.
Aperture oblong-ovate. Color of columella, lip and suture
white, sometimes tinged with yellow ; shell white, with or
without numerous transverse lines of a brown or black color.
Length nineteen-twentieths, width ten-twentieths of an inch.
Koolau poco, Oahu.

"This species makes a near approach to A. mustelina of
Mighels, but is more elongate in form, with the lineations
much stronger, and never passes into the variety of mustelina
with the depressed spire and obese body-whorl. The locality
also is different, which is always worthy of particular remark
when examining shells of this genus' : (Newcomb}.

The locality given by Dr. Newcomb is certainly erroneous.
No shell of this section, except A. lor at a, is known to occur on
the Koolau slope. Newcomb 's shell is well known to be from
the Waianae mountains, and probably the type came from
Mokuleia district, where it occurs in m,ustelina colonies.

Achatinella monacha Pfr. PL 30, fig. 9, photographic copy
of the original figure. "Shell imperf orate, ovate-conic, solid,
obliquely striatulate, little shining; gray-white, variously en-
circled with brown lines; spire elevated-conic, a little acute;

344 ACHATINELLA MUSTELINA.

suture lightly margined, subcrenulate ; whorls 6, a little con-
vex, the last a little shorter than the spire, rotund at base.
Aperture oblique, broadly obauriform ; columellar fold above,
oblique, moderate ; peristome sublabiate, the right margin
narrowly expanded, columellar margin thick, flexuous, adnate.
Length 20, diam. 10% mm. ; aperture 9y2 mm. long, 5 wide.
Inhabits Sandwich Islands, Frick' (P/r.).

This is merely a multilineate form of the grayish mustelina,
similar to multilineata Newc., but based upon a shorter form
than that.

Achatinella vestita. Shell sinistral, acuminate-conical ;
light brown or white, with beautiful narrow dark brown
bands, more or less numerous ; imperf orate ; whorls 6, convex ;
aperture semilunate ; lip reflected. Average length 1 inch,
diameter % inch. Hab. Waianai and Hawaii (Mighels, Proc.
Boston Soc. of Nat. Hist., ii, 1845, p. 20).

If really from Waianae this would, we suppose, be multi-
lineata Newc. Several of Mighels' localities are certainly
false, and as the type of vestita was destroyed by fire, we can
only drop the species as impossible to identify positively. We
are the more willing to do this because vestita has page-prior-
ity over mustelina, and we feel unwilling to give up a name
which has been identified with certainty for one to which some
doubt attaches.

Mokuleia. — The typical form of mustelina is that found in
Mokuleia district, in the northwestern flank of the Kaala mass.
The typical color, pi. 63, fig. 5, is a deep shade of carob-brown
or blackish chestnut brown, the embryonic whorls white, pale
brownish towards the apex, or entirely cartridge buff. The
dark color begins on the fourth or fifth whorl. The lip is
thickened within as usual, and is either vinaceous buff or
nearly white. Shell either dextral or sinistral. The size as-
signed by Mighels, "1 inch,' was probably approximate, as
the largest Mokuleia specimen I have seen is a little over eigh-
teen-twentieths of an inch. A more usual size is length 21,
diam. 12 mm.

We do not know that the dark typical color-form occurs in
pure colonies. It is associated with shells of a citrine drab

ACHATINELLA MUSTELINA. 345

hue, indistinctly streaked with deep olive ; also vinaceous buff
to nearly white shells with few or many brownish drab, cin-
namon or even black spiral lines or bands — a multilineata
pattern (pi. 63, figs. 4, 4a, 46, Mokuleia, close to the N.-W.
head of Makua, coll. by Spalding). There may also be var-
ious combinations of these patterns in some lots.

PL 63, figs. 5-56 are Mokuleia specimens from Gulick. PL
63, figs. 2, 2a are from Newcomb, without locality. Fig. 2 is a
very unusual pattern, which I have not seen in any other lot.

On the edge of Makua, along the whole lower length of the
ridge, Mr. Spaldiug found a pure colony of the olivaceous
drab form shown in pi. 63, fig. 4a. There are also pure colo-
nies of the multilineata pattern : pi. 63, fig. 3, western ravines
of Mokuleia, coll. by Spalding. In the Gulick collection there
are several Mokuleia lots of "multilineata," some running into
"sordida,' others into the olivaceous drab pattern (pi. 62,
figs. 1, 2). Occasional shells are white with some faint dusky
purplish stains and no bands.

Makaha. — On the south side of the range at the head of
Makaha valley, Mr. Spalding found some peculiar colonies.
In the northeastern extreme, under Kaala, pi. 63, fig. 1, the
shells are distinctly streaked with various shades of plum-
beous, dark olive gray and blackish brown on a white ground,
the streaks cut by a number of white spiral lines ; sutural mar-
gin white. This form is a modification of the more streaked
forms found in Mokuleia at the edge of Makua. It exists as a
pure race.

Further down, in Makaha, back of the home clearing of
the Manager of Makaha plantation, and on the edge of the
coffee clearing there is a race in which the tint varies from
nearly white, with indistinct pale ochraceous salmon streaks,
to vinaceous pink; sutural border white. As this is a pure
race, it may be distinguished by the name A. mustelina maka-
haensis P. & C. PL 62, figs. 3, 4.

Haleauau. — Eastward of the great ridge running northeast
from Kaala, in Haleauau, the mustelina pattern is modified
somewhat by the widening of the white band below the suture.
Usually there are light, streaks and spi -al lines in the dark

346 ACHATINELLA MUSTELINA.

color of the last whorl, sometimes producing a real multilin-
eata pattern. The embryonic whorls are typical in some lots,
the first neanic whorl bro\vn or marked with brown, as in
shells taken by Mr. Spalding in the northern ravines of Hale-
auau; but in a lot from the eastern ridge of Haleauau the
embryo is bicolored, ochraceous with a white band below the
suture, the tip fleshy. PL 63, figs. 6, 6a from the Thaanum
collection. Most lots from Haleauau include both dextral and
sinistral shells.

Pukuloa. — Several colonies where Mr. Spalding collected,
Pukuloa, between Mikalua Gap and Pukuloa, and on Kalena
Peak, have forms resembling those of the western ridge of
Popouwela, plain or multilineate, the sutural border some-
times as narrow as in Mokuleia. Also specimens approaching
bicolor by widening of the sutural white band.

On the Leilehua-Waialua division ridge the colony found
was all dextral, with a more or less dark basal tract not ex-
tending to the periphery (pi. 63, fig. 7, coll. by Spalding). In
a colony from the same neighborhood, but nearer Kaala, no.
1646 of Spalding 's collection, there are a few sinistral shells;
very few have a dark basal patch.

Popouwela. — On the western ridge the shells are mainly
sinistral, — 43 sinistral to 14 dextral in one lot I collected.
The embryonic whorls are bicolored, the upper half of each
buff, lower half white, or rarely they are all ochraceous or all
very pale buff, the tip is always slate-violet. The last whorl
is wood brown to light buff or almost white, with faint oblique
darker striae and olive-brown spiral lines and bands, or rarely
very dilute purplish-gray bands (pi. 63, figs. 9-9&, northern
colony of the western ridge of Popouwela). Further up the
ridge there are more white shells with faint or pale neutral
gray streaks, with or without blackish spiral lines (pi. 63, figs.
8, 8a).

PL 63, figs. 10 to lOc. Near the foot of the third small
ridge eastward there is a large colony in which the color is
Mars brown with blackish bands and a wide, snow-white
sutural band, occupying half of the penultimate whorl. Only
18 in 173 found by me are dextral. Among the dark shells are

ACHATINELLA MUSTELINA. 347

about half a dozen as light as the banded form from the west-
ern ridge. Fig. 10 is a very unusual color-pattern in this
colony, and figs. 106, c show extremes of shape but normal
coloring.

In a small area of probably two or three square rods we
found snow-white shells with the columella dull lavender or
white, summit white. This is evidently a recent albino muta-
tion which has not had time to spread far. They occur with
the ordinary pattern of the surrounding colony. The speci-
mens I found were all dextral.

Further up the same ridge the shells are intermediate in
pattern between the preceding colony and that of the western
ridge of Popouwela.

A colony in coll. Spalding from the "southern ridges of
Popouwela' varies from the patterns of pi. 63, figs. 12, 12a
to black with white bands or white flecks, resembling the
lymaniana pattern. One is figured, pi. 63, fig. 11.

On the Popouwela-Lihue division ridge Mr. Spalding found
a dextral and sinistral form like that shown in pi. 63, fig. lOc,
except that the white sutural band is a trifle wider. With
them, without intergrading specimens, there is a smaller form
closely resembling turbiniformis, but with less concave spire,
pi. 62, fig. 21. This may be regarded as the western limit of
the turbiniformis stock. The shells are dextral, and from
their smaller size, different coil and markings, probably do
not hybridize with their larger dark companions.

Further up the same ridge, Mr. Spalding took lighter bi-
colored shells with others of multilineata pattern (pi. 63, figs.
12, 12a).

It appears that between the main northeastern ridge from
Kaala and the Popouwela-Lihue ridge, the prevalent form is
a shell with the white sutural band wider than Mokuleia mus-
telina on the west, and narrower than Lihue "bicolor," east-
ward. The darkest forms are near the lower limit of shell
country, while on the higher ridges the lighter, multilineata
patterns prevail ; but there are exceptions. It will be noted
that in the Main Range banded forms are also chiefly devel-
oped on the ridges.

348 ACHATINELLA MUSTELINA.

Lihue. — This is a district of somewhat indefinite limitsr
where Gulick, Newcomb, and other naturalists of their day
collected, on the east and southeast slopes from Kanehoa. In-
deed, Lihue ("Lehui") and Mokuleia seems to have been the
chief sources of their Waianae shells.

In the large Lihue series before me from Gulick the muste-
lina-multilineata forms have a wide sutural white band, as in
Popouwela. They vary widely in size, from IS1/^ x 10 to 2Qy2
x 13^3 mm., and a few have the turbiniformis contour. They
have linear streaks of fawn on a buff ground, with or without
dark and light spiral lines or bands. PL 63, fig. 13cZ is a
slender shell of this series, others of the Gulick lots are fig-
ured on pi. 61, figs. 7, la. They resemble the Popouwela shells,
and also, in some specimens, are remarkably like A. decora.

A. m. bicolor 'Gulick' Pfeiffer is typical of Lihue. PL 63,
figs. 13 to 13c, Gulick collection. It is chestnut-brown to
blackish carob-brown below, white above. Typically the white
should extend almost to the periphery, and leave a mere line
of dark brown above the suture, but in the series of over 200
before me, this ideal is rarely attained. The white band com-
monly varies from 1% to 4 mm. wide on the last whorl. The
sutural border is often tawny. The summit is warm or ochra-
ceous buff, paler or dusky at the tip, and often has paler bands
on the last embryonic whorl. Figs. 13&, 13c show unusual ex-
tremes of shape. With the lots of bicolor there are a few
shells transitional to the pattern shown in pi. 61, figs. 7, la.
Probably the bicolor occurs low, the lighter form higher on
the ridges, with more or less admixture in the colonies. The
original description of bicolor follows.

"A. [chatinella] bicolor 'Gulick' Pfr. Shell subperforate,
globose-conic, solid, lightly striate, glossy; whitish, obliquely
streaked with brownish ; spire conic, with rather acute apex,
white ; suture broadly impressed-marginate ; whorls 51/2, the
upper ones flat, penultimate more convex, the last nearly equal
to the spire, globose, black-chestnut below the periphery.
Aperture oblique, subauriform ; columellar fold high, nodi-
form ; peristome labiate within, brown-bordered, the right
margin narrowly expanded ; columellar margin dilated, sub-
adnate. Length 16!/2, diam. 11 rnm. ; aperture 8% mm. long,
5 wide. Inhabits Lehui, island of Oahu': (Pfr.).

ACHATINELLA MUSTELINA. 349

A. bicolor Gulick in Mus. Cuming, PFR., Monographia Heli-
ceorum Viventium, iv, 529, 1859. It was never described by
Mr. Gulick.

40a. A. MUSTELINA SORDIDA Newcomb. PL 30, fig. 27 ; pi. 62,
figs. 5-11.

"Shell elongately conical, solid; whorls 7, slightly rounded,
margined above ; aperture small, somewhat contracted ; lip
subreflected below, acute above, thickened within ; colurnella
short, slightly twisted, with an expanded callus partially cov-
ering a shallow, small umbilicus ; color variable, with a white
ground variously banded and striped transversely with brown
or black. Length eighteen-twentieths, width nine-twentieths
of an inch' (Newcomb).

OahurLettui [Lihue] (Newc.).

Achatinella sordida NEWC., P. Z. S., 1853, p. 139, pi. 23, f.
27. — PFR., Monogr., iv, 523. — Achatinella napus PFR., P. Z. S.,
1855, p. 5, pi. 30, f. 19 ; Monogr., iv, 534.— Cf. Newcomb, Ann.
Lye. N. H. of N. Y., vi, p. 308.

The original figure is reproduced in pi. 30, fig. 27.

This race differs from turbiniformis and concavospira by
its less convex whorls and straight-sided spire ; from muste-
lina-bicolor by the more slender contour. It varies a good deal
in color — from fawn or Vandyke brown with a white sutural
band to white with some faint tawny suffusion or lines
towards the base. I understand that it occurs high on the
ridge, between the area occupied by mustelina and bicolor and
that of lymaniana and concavospira. There are transitional
forms between sordida and lymaniana.

A. m. sordida is usually sinistral, but sometimes has a dex-
tral form, which was described as A. napus.

"A. napus Pfr. [pi. 30, fig. 19, reproduction of original
figure]. Shell imperf orate, dextral, turrited, solid, striatu-
late, a little shining; whitish, encircled with grayish-fleshy
bands and sometimes chestnut lines ; spire long-conic, apex
white, acute ; suture distinctly margined ; whorls 6%, the
upper flat, following slightly convex, the last whorl about
equal to one-third the length, rounded ; aperture diagonal,
truncate-auriform, columellar fold superior, moderate, lightly

350 ACHATINELLA MUSTELINA.

twisted • peristome lipped within, the right margin unex-
panded, columellar margin somewhat dilated, adnate. Length
19%, diam. 10% ; aperture 8 mm. long, 4% wide. Inhabits
the Sandwich Islands, Frick, Mus. Cuming'1 (P/r.).

This form is considered by Dr. Cooke to be an absolute
synonym of A. sordida Newc., from which it differs in being
dextral and imperf orate. Dr. Newcomb thought it an "elon-
gated variety" of A. pulcherrima. From this arose Pfeiffer's
' ' Achatinella pulcherrima var. elongata Newc.' (Mono-
graphia, vi, 172, 1868). Mr. Sykes considers A. concavospira
Pfr. to be identical with napus (Fauna Hawaiiensis, p. 303).
The weak convexity of the penultimate whorl and the straight
outlines of the spire are common characters of sordida and
napus. Moreover, one of Gulick's lots contains both dextral
and sinistral specimens, otherwise similar (pi. 62, figs. 5-8),
which seem to demonstrate the identity of the two supposed
species. PI. 62, figs. 9, 10, 11 were drawn from specimens of
sordida received from Dr. Newcomb.

. A. MUSTELINA LYMANIANA Baldwin. PI. 61, figs. S-9b, 11.

1 ' Shell sinistral, very minutely perforated, solid, ovate ;
spire convexly conical, apex subacute ; surface shining, cov-
ered with fine incremental lines, under a strong lens showing
minute decussating striae; apical whorls smooth, when not
eroded. Color dark purplish brown, sometimes with longitu-
dinal or transverse white flecks or zigzag lines; a white line
traversing the suture ; apex light chestnut. Whorls 6, very
lightly margined above, somewhat convex ; suture moderately
impressed. Aperture oblique, white within, sublunate ; peri-
stome acute, thickened within, expanded, the columellar mar-
gin slightly reflexed and covering the small perforation, color
white on both face and the reverse ; columella white, termin-
ating in a slightly developed flexuous fold. Length 20%,
diam. 11% nim.' (Baldwin}.

Oahu: near the southern end of the Waianae range, from
around Green Peak (Palikea), three or four miles along the
ridge southward, only at high elevations.

ACHATINELLA CONCAVOSPIRA. 351

Named for Mr. Ernest Lyman. Types in the Baldwin col-
lection.

This is a well-marked subspecies, always sinistral, solid,
with long, straightly conic spire, the sutural white band nar-
row, usually reduced to a mere line, or sometimes absent. The
typical pattern is carob-brown with speckled white streaks,
more or less interrupted by spiral lines or bands of the dark
color, or reduced to a few whitish flecks on a blackish carob-
brown ground (figs. 8 to Sc, Palihua iki, coll. by Dr. C. M.
Cooke).

Another pattern, from the same locality and collector, is
ochraceous tawny, with streaks and spiral lines of warm buff,
a pale line at the suture or none (fig. Sd).

Other shells of this lot are cartridge buff, almost white,
with the faintest traces of pale ochraceous salmon bands, fig.
Se.

Length 23.8, diam. 13.2 mm. ; 6% whorls.

Length 22.5, diam. 13.5 mm. ; 61/2 whorls.

Length 20, diam. 10.3 mm. ; 6y2 whorls.

The typical forms of lymaniana occur along the ridge south
of Green Peak. North of Green Peak it is mixed with forms
with the napus pattern, — buff with a few wide dark bands,
the shell sinistral — probably in a hybrid colony. Specimens
from the Lihue side, under Palikea, are figured, pi. 61, figs.
9 to 9&, coll. by Spalding. Pale forms of lymaniana pass into
sordida Newc., so that there seems to be no definite distinction
between selected individuals although there may be a differ-
ence in the average constitution of the colonies.

It is also likely that some adumbration of the lymaniana
stock has penetrated as far north as the southern ridges of
Popouwela, forming hybrids with the gray and banded mus-
telina there. PL 63, fig. 11 has the appearance of being a
lymaniana segregate of this stock, and there are others in the
collection before me.

41. A. CONCAVOSPIRA Pfeiffer. PL 52, fig. 16 ; pi. 61, figs. 12-
12cZ ; pi. 62, figs. 24, 25.

"Shell subperforate. dextral. ovate-turrite, solid, striatu-

352 ACHATINELLA CONCAVOSPIBA.

late, glossy; whitish, ornamented with bands and narrow
streaks of coffee color. Spire concavely turrite, the apex
somewhat acute, white ; suture strongly margined. Whorls
7, the first three flat, following whorls convex, the last one
rounded, about equal to two-fifths the length. Columellar
fold superior, nodiform, white. Aperture oblique, reversed
auriform; peristome liver colored, the right margin a little
expanded, columellar margin very much dilated, reflexed, ad-
nate. Length 211/2, diam. HVs mm. Inhabits Sandwich
Islands, Dr. Frick, in Mus. Cuming' (P/r.).

Palihua and Palihua iki, at the extreme southern end of the
Waianae range ; the form tur'biniformis in Lihue and on the
Popouwela division ridge.

Achatinella concavospira PFR., Proc. Zool. Soc. Lond., 1859,
p. 30; Monographia, vi, p. 169. — Apex tur'biniformis GULICK,
P. Z. S., Lond., 1873, p. 81, pi. 10, f. 7.

A. concavospira with its several races is always dextral, so
far as we know, thereby differing from A. m. lymaniana of the
same district, which is invariably sinistral. Where A. c. tur-
'biniformis occurs in the same colony with A. m. bicolor on the
Popouwela-Lihue division ridge, there are no intergrading spe-
cimens (cf. no. 646, 647 coll. Spalding). It appears that
where concavospira forms occur with various subspecies of
mustelina, the two retain fully the proper shape, size and
color of their respective species, and the inference may be
drawn that they do not hybridize, at least ordinarily. My at-
tention was not specially directed to this point when studying
the Honolulu collections, but the material I brought home and
possess from the Gulick collection leads me to segregate con-
cavospira specifically.

The type-specimen of A. concavospira as represented in pi.
52, fig. 16, which I owe to the kindness of Mr. E. A. Smith,
and typical shells such as Mr. W. D. Wilder collected in Pali-
hua at about 3,000 ft. elevation, pi. 62, figs. 24, 25, is a strik-
ingly distinct form, differing from A. mustelina. by its very
concave-sided slender spire and the short, swollen, last whorl.
The 3% embryonic whorls are flattened, white or a little fleshy
towards the tip. Subsequent whorls a little over three, the

ACHATINELLA CONCAVOSPIRA. 353

first one slightly, the last two very convex. All are deeply
margined below the suture, the margin at first brown or tinted,
often becoming white later. The last whorl is marked with
bands of carob-brown or tawny, and sometimes tawny streaks,
on a white or warm buff ground, the markings mainly on the
base, and deepening as they approach the lip. The aperture
is white within, lip deep brown, slightly expanded and thick-
ened within. All seen are perforate and dextral.

Length 19*4, diam. ll^/s mm. ; aperture 9 mm.

Length 19, diam. 10% mm. ; aperture 8.9 mm.

In a lot from Palihua (pi. 61, figs. 12 to 12d, coll. by C.
Montague Cooke) this typical form passes into almost the
mustelina shape, with varied coloring. The dimensions are
remarkably diverse, from length 20, diam. 11.3, aperture 9
mm., to length 20.2, diam. 12.5, aperture 10.8 mm. This lot
connects typical concavospira with the following.

A local form or subspecies of concavospira, which may be
called griseizona P. & C., was found by Dr. Cooke in Palihua
iki. The colony, which is very uniform, extends over ten or
fifteen acres of lantana, at the head of a small valley running
north. The shells, pi. 61, figs. 10, 10a, 10&, are rather stout in
figure, the outlines of the spire decidedly to very slightly con-
cave. The marginal furrow below the suture is less emphatic
than in typical concavospira, and appears on 2% whorls, in-
stead of over 3 as in concavospira. The apex and embryonic
whorls are white ; there is no sutural color-band on the early
neanic whorls ; the later whorls have fawn bands, more or
less streaked and overlaid with white, producing various
shades or tints of brownish drab, pale Quaker drab or pale
violet gray. The lip is white or dull brownish. There are
6*4 to 6!/2 whorls, the penultimate whorl much less convex
than in concavospira. This form has the coloration of napus.

41a. A. CONCAVOSPIRA TURBiNiFORMis Gulick. PL 62, figs. 22,
23.

"Shell dextral, subperforate, subglobose-conic, a little shin-
ing, striated with growth lines but hardly spirals, white, en-
circled around the base with a few brown transverse lines;

354 ACHATINELLA CONGA VOSPIRA.

suture distinctly margined ; spire lightly concave ; whorls 6,
the first three a little convex, the rest convex. Aperture
white ; peristome with a thin margin, bordered within with
dilute brown ; columellar fold moderately strong. Length 17,
diam. 11 mm. ' ( Gulick ) .

Oahu: Kalaikoa and Lehui [Lihue], on trees (Gulick).

"It is allied to A. tuberans Gk., but has a much narrower
spire. All the specimens found are dextral. The shell fig-
ured is from Lehui' (Gulick).

Gulick 's figured type from Lihue is drawn on pi. 62, figs.
22, 23, no. 102 Boston Soc. coll. The locality "Kalaikoa" is
, probably an error. It is possible though not probable that
under long past conditions it extended across the central
plain, as some Amastras have done. However, turbiniformis
has the look of a shell from high on the ridges. It is, in fact,
merely a shortened form or local subvariety of concavospira,
which is a high altitude shell.

The apex and embryonic whorls are white in the specimen
selected by Gulick as type. It was a dead shell. The last em-
bryonic whorl is weakly plicate axially, as in some typical
concavospira. The sutural margination of the post-embryonic
whorls is strong, and faintly touched with brown along the
sutural edge on the last two whorls, perhaps further up, in
life, the spire of the type appearing somewhat faded. The
penultimate and last whorls are particularly convex, last
whorl short, swollen, with a pattern of interrupted oblique
streaks and narrow spiral lines of brown, chiefly on the base,
but near the lip the pattern extends upward nearly to the
suture. The narrow lip-callus stops a few millimeters short
of the suture, as in typical concavospira. Length 16, diam.
11, length of aperture 8 mm. ; 6% whorls.

Mr. Spalding collected turbiniformis on the Lihue-Popou-
wela division ridge, in a prolific colony of bicolor, both pre-
serving their individuality. It occurs for some distance up
the ridge. One of this lot is figured, pi. 62, fig. 21.

APPENDIX 1.

i

ADDITIONAL NOTES, AND NEW SPECIES OF ACHATINELLID^:.

Genus NEWCOMBIA Pfr.

This volume, page 1. The word sinistral should be added to
the generic diagnosis.

N. pfeifferi and cinnamomea live chiefly on the bark of
small trees where they stick twig-like, often in a fork, and are
hard to see. N. plicata has the same habit.

N. sulcata and gemma live on leaves, according to Mr. Thaa-
num. N. canaliculata lives on lehue and lantana.

NEWCOMBIA PLICATA (Migh). Page 3. Add the localities:
Waialue (Thaanum) ; small trees overhanging the pali at top
of Leper Settlement trail (Cooke & Pilsbry).

NEWCOMBIA PFEIFFERI (Newcomb). Page 13. This form
does not occur at or near Puu Kolekole (a locality for cinna-
momea), and that locality should therefore be deleted. Dr.
Cooke and Mr. Thaanum hold that pfeifferi and cinnamomea
are extremes of one species ; a view which their collections sup-
port. The subspecies called ualapuensis Pils. belongs to
pfeifferi rather than to cinnamomea. The distribution of the
forms is rather peculiar. Beginning at the eastern valleys
and going westward, the forms are encountered thus:

Honomuni honomuniensis.

Ahaino decorata.

Kupeke decorata.

Pukoo fossil cinnamomea ( Thaanum ) .

Mapulehu cinnamomea.

•

Kaluaaha pfeifferi.

Ualapue ualapuensis.

Kahaanui ualapuensis.

( Several valleys wanting. )

Makolelau cinnamomea.

Makakupaia cinnamomea.

(355)

356 NEWCOMBIA AND PARTULINA.

It will be seen that east of Kaluaaha there are only smooth-
ish variants of cinnamomea; west of Kahaanui a long series of
parallel valleys and ridges supervenes in which no allied form
has been found, cinnamomea appearing again in Makolelau
in a ravine east of the house, and in a small ravine east of
Puu Kolekole (Cooke & Pilsbry), and further west in Maka-
kupaia.

"Where the connection between the eastern and western
herds finds place, if at all, is not clear, as we saw nothing of
the species in the region above the Kamalo amphitheatre,
where Thaanuni and others also have collected.

N. PHILIPPIANA Pfr. (perkinsi Sykes). Antea, pp. 8, 9.

It is interesting to find that this most primitive of existing
Newcombias occurs in the Pleistocene shell-beds of Moomomi
(Cooke & Pilsbry, 1913). Like the other tree-shells of the
dune deposits, it is rather rare.

Genus PARTULINA Pfr.

Section PERDICELLA Pease.

P. ORNATA (Newc.). Page 18. By an oversight the locality
was given as East Maui. Very beautiful specimens have been
taken by Mr. Thaanum on Mt. Helu and Mt. Lihau, both in
West Maui. The only other locality known is that given by
Newcomb.

P. HELENA (Newc.). PL 54, figs. 1-3.

Page 16. The quotation from Newcomb on p. 17 might give
a false idea of the abundance of this species. It is a common
shell on shrubs of many kinds, from the head of Karnalo to
the region of Puunea, at the present western limit of tree-
snails; found only on the high part of the island, of course.
No doubt Newcomb 's type was from the Puunea region, some-
where between Meyer's place and the north cliff.

Different colonies, even in the same neighborhood, are often
unlike in markings. Thus, on the margin of the flat north of
Meyer's lake the patterns of pi. 4, figs. 1-5 prevail, but there
are also a few of the balteata pattern, similar to pi. 4, fig. 7.

PARTULINA. 357

No blends between the normal pattern and bqlteata were
noticed.

In another colony, west of Meyer's lake, the markings are
bolder and there is no whitish belt. The oblique-striped pat-
tern is usually more dislocated than in pi. 54, figs. 1, la, but it
always gives place to a vertical striping on the latter part of
the last whorl.

On the flat east of the peak of Puukolekole the majority of
specimens have the pattern of pi. 54, fig. 3, but about one-
third are cinnamon-buff with a dark peripheral band, bor-
dered below with a white band, pi. 54, fig. 2. This is very
much the color of the normal embryo shell, in the first half of
the third whorl.

The several stages of the color-pattern are rather well-
marked in P. helena, and deserve further study.

P. THWINGI P. & C., n. sp. PI. 54, figs. 6, 6a, 6b.

The shell is dextral, perforate, ovate-conic, moderately solid.
White, profusely maculate and mottled with mummy brown,
under the lens showing a few ochraceous spots or streaks, the
periphery marked with a narrow dark band, a cream-white
band below it. The last embryonic whorl has forwardly-
descending stripes of dark brown and white ; summit pale.
Surface not glossy, under the lens rather distinctly marked
with spiral and descending, waved striae, and irregular
growth-lines. Spire straightly conic, tapering to a subacute
apex. Aperture pallid purplish-gray and brown within ; outer
lip with a pale acute edge, unexpanded ; basal lip slightly ex-
panding, the columella slightly brown-tinted, having a rather
strong, callous fold above ; the edge reflected close to the in-
sertion.

Length 17, diam. 9.8, aperture 8 mm. ; G1/^ whorls.

Length 16, diam. 9 mm.

East Maui : Auwahi, D. Thaanum. Cotypes in coll. A. N.
S. P. and Bishop Museum. Also in Thaanum collection.

One of the largest Perdicellas, notable for its straightly
conic, rather slender spire, and the particular color-pattern.
The dark peripheral band, bordered with a white one, is in-

358 PARTULINA.

variable in the series examined. The pattern of longitudinal
zigzag stripes is distinct in places, or the stripes have been
so dislocated that usually a mottled effect is produced. In
both color and shape it has some resemblance to P. grisea Nc.,
which differs by being sinistral, larger, with a different pat-
tern of color on the embryonic whorls, and a less prominent
columellar fold. P. carinella has much the same general color,
and no doubt is somewhat related.

Named for the Rev. E. W. Thwing, whose compilation of
the original descriptions of Achatinellida has been a useful
handbook for Hawaiian conchologists. The large collection
made by Mr. Thwing now belongs to the Bishop Museum.

PARTULINA CARINELLA (Baldwin). Page 7.

This species is undoubtedly a Perdicella, as Dr. Cooke and
Mr. Thaanum have pointed out to me. It should be removed
from Newcombia, which without it consists wholly of sinistral
species.

Section PARTULINA.

PARTULINA VIRGULATA (Migh.). Page 25.

This species inhabits the eastern part of Molokai, almost
one-third of the island 's length ; from Pelekunu and the first
valley northwest of Kamalo to the eastern end of the island.
Mr. Thaanum, who has had wide experience with the species,
finds that eastward of Waialua individuals are sinistral in
great majority. Out of 1,067 collected by him, 1,053 are sinis-
tral, 14 dextral. West of Waialua the banded (typical) form
is almost always dextral, while the plain or faintly banded
form (pi. 26, figs. 1, ~La, Mapulehu, coll. by Thaanum) is sin-
istral, though both may be found on the same tree. The
factors for plain coloring and sinistral coil are evidently
linked in inheritance.

In the first valley west of the heads of Kamalo, just above
the irrigation ditch, east of the stream, the authors found a
small, well-banded form of virgulata about 21 mm. long. This
is probably the extreme western limit. Typical banded vir-
gulata was taken by Thaanum in Honouliwai, a place far east-
ward, not before recorded.

PARTULINA. 359

A. virgulata halawaensis 'Baldwin' Borch. PL 26, figs. 2,
2a, Kepookoholua, coll. by Thaanum, seems to be rather a
''good" subspecies by its slender form, pale color, with nar-
row bands or none. It is found only in the Halawa district.
In general, P. virgulata is darkest westward, lighter towards
the east, though there are individual exceptions.

P. VIRGULATA KALUAAHACOLA n. V. PL 26, figS. 3,

The shell is small, sinistral ; embryonic whorls white with
a blackish-chestnut band above the suture ; subsequent whorls
closely marked along growth-lines with hazel lines on a warm
buff ground, or with indistinct brownish streaks and some
hazel lines on a tawny olive ground ; indistinctly banded.
Length 19.5 to 20 mm.

Kaluaaha, at about 1,700 ft., on ieie. D. Thaanum.

This is a local race, related to the plain form of virgulata
figured on pi. 26, figs. 1, la. It occurs at the lower limit of
tree shells in this valley, and is extremely rare — perhaps now
extinct.

P. SUBPOLITA Hyatt & Pilsbry, n. sp. PL 23, fig. 1.

Shell imperforate, sinistral, ovate-conic, rather solid. Em-
bryonic whorls white with a broad chocolate band above the
suture, having excessively faint spiral stride or none. Follow-
ing whorls light chrome ("pinard yellow"), uniform or hav-
ing one or several faint whitish lines at or below the middle.
Surface glossy, with distinct wrinkles of growth but only faint
traces of spiral striae. Last whorl somewhat compressed
peripherally. Aperture white, the outer lip expanded below,
a little thickened within; columella white, fold moderate.
Length 17, diam. 11, aperture 9 to 9.2 mm.

Molokai : Ualapue, Thaanum. Cotypes 110. 106992 A. N. S.
P. and 1932 Cooke coll.

[Partulina] subpolita P. & H., Manual xxii, p. 24, 1912.

The color pattern of the embryo and the slightly expanded
lip allies this to P. virgulata, and it stands nearest to P. v.
kaluaahacola from a neighboring locality. This relationship,
pointed out to me by Dr. Cooke and Mr. Thaanum, became

360 PARTULINA.

obvious on seeing the superb series of virgulata in Mr. Thaa-
num's collection. It is distinct from P. polita by its broad,
short spire, sinistral coil, expanded basal lip and white mouth.
Four specimens in coll. University of Wisconsin, two in the
Cooke collection, one in that of the Academy and several in
Mr. Thaanum's collection are essentially alike. It is one of
the rarest Molokaian shells.

PARTULINA TESSELLATA (Newc.). Page 28. Taken abun-
dantly in Puunea ("Puanea" of Borcherding's map) at about
the western limit of the species; also at the "Pig Wallow'
flat above the pipe-line trail, Puulua, Kaunakakai, Cooke and
Pilsbry, January, 1913. Eastward it barely reaches the vir-
gulata territory.

PARTULINA RUFA (Newc.). Page 29. Newcomb's type figure,
is copied photographically in pi. 29, fig. 3.

PARTULINA PROXIMA (Pease). Page 32. In the redfieldi
colony of the heads of Kamalo we found several proxima pure
white except for the purple columella. Thaanum found a
small form all with pale green-yellow ground in Kamalo in
1906, on the same trees from which he had collected normal
(white ground) proxima in 1894.

We took normal proxima on the east side of Puukolekole, in
the P. dwightii colony. Some of Thaanum's series from Ka-
wela are very small, length 18.5 mm.

P. proxima multistrigata Pils., page 34, is evidently, as Mr.
Thaanum holds, a western subspecies of proxima, and not of
theodorei. It has been collected in some abundance in Kala-
maula by Mr. Thaanum. This is probably the original local-
ity. P. theodorei has not been found so far west.

PARTULINA THEODOREI Baldwin, page 33. Another habitat
of this apparently valid species is Makakupaia, recorded by
Borcherding, from shells collected by Mr. Meyer. This is
westward from Baldwin's locality Kawela. Probably all the
records from Kawela rest ultimately upon Meyer's authority.

PARTULINA DWIGHTII Newcomb. Page 35. PI. 26, figs. 5&
to 5/.

This species is very closely related to P. redfieldi, but of

PARTULINA. 361

more pyramidal shape with broad white lip-border and white
columella, while in redfieldi the last whorl of the shell is larger
and the lip and columella are dark. P. dwightii is a western,
P. redfieldi an eastern species. Puukolekole, where we
collected a couple of hundred in a colony on the eastern
side, is about the eastern limit of dwightii. In this colony we
found a few real redfieldi (pi. 26, figs. 5, 5&), forming about
16 per cent of the whole. The two species overlap in this dis-
trict, and it can hardly be doubted that they form hybrid colo-
nies, segregating into the brown- and the white-lipped forms.

Westward, P. dwightii is not known farther than Makaku-
paia, to my knowledge ; but it formerly extended as far as the
summit of Manuna Loa, and Moomomi on the north coast, in
a slightly modified form, described below.

The Puukolekole colony is illustrated in pi. 26, figs. 5 to 5/.
It is clear that in this colony the forms called compta Pse. and
concomitans Hyatt are mere color-mutants, and in no sense
races. The embryos removed from these shells show that
several patterns are often the progeny of one parent.

Partulina dwightii, color-form concomitans, has been erron-
eously referred to as Achatinella macrodon by Perkins, Fauna
Hawaiiensis I, pt. vi, p. Ixvi, 1913.

Var. mucida Baldwin. Page 34. Typical mucida occurs
low down in Kamalo (Thaanum), and also in the type locality,
Makakupaia below Puu Kaeha. It is merely a small form or
race of dwightii. P. macrodon Borch. is not a variety but only
a color-mutant occurring with mucida.

P. dwightii occidentalis P. & C.. n. v. PI. 26, fig. 6.

Smaller than dwightii, wider than mucida, aperture shorter,
the lip less prolonged basally. Length 20.5, diam. 12.6, length
of aperture 9.5 mm.

Sand dunes of Moomomi (on the north coast almost due
north of Mauna Loa) ; also summit of Manna Loa, under
stones. Cooke and Pilsbry, 1913.

This Pleistocene variety shows that one time tree snails ex-
tended almost to the western end of Molokai. A single frag-

362 PARTULINA.

ment was found on the southern rim of the bowl near the
summit of Mauna Loa, where no doubt forests survived long
after it had disappeared in the neighborhood of Moomomi,
where the shell occurs much less than a hundred feet above
sea level.

PARTULINA REDFIELDH (Newc.). PI- 29, fig. 5.

Page 38. Mapulehu specimens agree well with Newcornb?s
original lot. His type figure is copied photographically on
pi. 29.

Var. kamaloensis P. & C., n. v. PI. 26, figs. 4, 4a.

Between the branch ravines above the Kamalo amphitheatre
and below the old irrigation ditch we collected in three simi-
lar colonies of redfieldii. The shells were very abundant on
leafless and often dead lantana. They are mostly large,
capacious, and cinnamon brown to chestnut brown or burnt
umber in color, most old ones lighter by weathering and loss
of cuticle, sometimes to a dull, pale ochraceous buff tint.
Rarely the fresh shells are ochraceous buff, with or without a
dark band. Very few shells have numerous dark bands on a
pale ground. As the colonies are of considerable extent, and
nearly homogeneous in character, it may be well to have a
name for this particular race.

Length 27.5, diam. 16.7, aperture 14.2 mm.

Length 25.6, diam. 16, aperture 14 mm.

Length 27, diam. 15, aperture 13 mm.

PARTULINA CRASSA (Newc.). Page 40.

The dull form of P. crassa described by Newcomb and in all
the older collections was probably found near the western end
of Lanai, now a deforested region where they still occur as
fossils. In the Koela region the shells are of a fine dark brown
color.

P. KAAEANA Baldwin. Page 41. Mr. Thaanum thinks that
this may be identical with or a variety of P. ustulata Gul. (p.
47 ) . Unfortunately a careful search failed to locate the type
of ustulata in the Boston Society collection. It seems to be
lost. No topotypes are extant, and the ustulata from other

PABTULINA. 363

localities in Gulick's collection are rather unlike the type fig-
ure and description. The question of identity must therefore
remain in abeyance.

P. MARMORATA (Gould). Page 42. The original figure of
the synonymous A. adamsii Newc. is reproduced on pi. 29, fig.
20.

PARTULINA WINNIEI Bald. P. 44. Mr. Thaanum has a dex-
tral specimen.

PARTULINA PERDIX (Eve.). A. pyramidalis Gul. (p. 46)
seems from Mr. Thaanum 's series not to be separable from
perdix. A synonym is : Achatinella pyrimidalis Gul., LYONS,
Hawaiian Almanac for 1892, p. 105.

P. 47, top line, for pyramidal read pyramdafo's.

P. GOULDH (Newc.) PL 29, fig. 1.

Page 52. The form I separated as P. g. perfecta (p. 54)
occurs with typical gouldii at Ainaloa, W. Maui (Thaanum),
and is apparently not a valid race. Newcomb's type figure of
gouldii is reproduced photographically on pi. 29.

PARTULINA RADIATA (Gould). Page 49. A series of this
species, typical and a paler form, is in the J. S. Emerson
collection from "Lahaina.' While this locality leaves con-
siderable latitude to a naturalist seeking the shell, it at least
indicates the southwestern ridges of West Maui as its habitat.
There is a good series in the collection of Mr. W. G. Mazyck
of Charleston, some in the U. S. Nat. Mus., and a series in coll.
A. N. S. P. It was apparently not uncommon sixty years ago,
when Emerson, Newcomb and Gulick were collecting, but may
be extinct at the present time.

P. SPLENDIDA (Newc.). P. 51. PL 29, fig. 4 (reproduction
of the original figure ) .

P. APTYCHA (Pfr.). P. 54. PL 30, fig. 1 (reproduction of
original figure).

P. NIVEA (Baldw.). P. 59. Mr. Thaanum found a tawny-
streaked form with the typical form at Makawao. It has a
faint pale line at the periphery, but no white basal band,
otherwise agreeing with "var. kaupakaluana.' The latter is
probably a mere color-form, not a race.

364 PARTULINA.

P. DOLEI (Baldwin), p. 60, is rarely found dextral.

P. CARNICOLOR Bald., p. 58, might better be classed as a sub-
species of eburnea.

P. TEREBRA Newc., p. 61. According to Mr. Thaanum, tere-
bra-ligneria forms are all around the head of lao valley, high
up ; also on Mt. Helu and Mt. Lihau.

P. TEREBRA ATTENUATA (Pfr.). P. 63. PL 30, fig. 12 (re-
production of the original figure).

Section EBURNELLA Pease.
P. VARIABILIS LACTEA Gulick. Page 86.

Achatinella saccata Pfr. is an absolute synonym of the
above. Through the kindness of Mr. E. A. Smith I can figure
the type-specimen in the British Museum. The original de-
scription follows.

Achatinella saccata Pfeiffer. PL 54, fig. 4. Shell subper-
forate, sinistral, turrited, rather solid, striatulate, a little de-
cussated under the lens, glossy, white (candida) ; spire regu-
larly tapering, the apex acute ; suture narrowly margined ;
whorls 6%, rather flattened, the last equal to two-fifths the
length, a little convex, the base subcompressed-sack-like ; colu-
mellar fold high, tooth-shaped, brown or flesh-colored ; aper-
ture very oblique, semioval, the base laterally produced, inside
pale rosy ; peristome simple, unexpanded, the columellar mar-
gin dilated, somewhat free. Length 21, diam. 9y2 mm. Sand-
wich Islands, Mus. Cuming (Pfr.).

Achatinella saccata PFR., Proc. Zool. Soc. London, 1859, p.
30 ; Monographia Hel. Viv., vi, 175. — Eburnella saccata Pfr.,
PEASE, P. Z. S., 1869, p. 647.

The slight roughness of the figure is due to the process of
reproduction.

Section BALDWINIA Ancey.

The species are most numerous in northern Hawaii, but ex-
tend more than halfway down the island, on both sides. The
colonies, except in Hamakua district, are extremely rare and
isolated. Their locations on the island may be represented
thus.

PARTULINA. 365

horneri

f

physa

confusa
horneri
konana errans

P. HORNERI (Baldwin). Page 107.

On p. 108 the color- varieties of this species have been de-
scribed. On going over the series sent by Mr. Thaamim it
seems worth while to emphasize the perfectly constant differ-
ences by applying names to the forms. If they are always
found in pure colonies, as seems to be the case, these patterns
have a racial value. In any case, the names will be useful in
discussing the composition of colonies.

The typical horneri has a broad peripheral band, a small
umbilical patch and an inconspicuous subsutural band (often
faint in the adult stage), of light ochraceous buff (pi. 17, fig.
1). These bands are all present in the embryonic stage, pi.
54, fig. 11. It will be noted that the peripheral band extends
above the peripheral angle.

Var. Candida. PI. 17, figs. 3, 4; pi. 54, fig. 12, embryo.
(= var. c, p. 108.) The peripheral band of the embryo is nar-
rower, and does not extend above the angle. There are no
other bands on the cream-white shell.

A small form of P. h. Candida has been found at Honoula,
N. Kona, and sent by Dr. Cooke. The peripheral band persists
longer than in Kukuihaela shells, being visible on the front of
the last whorl. The spiral striation is a little stronger.
Length 18-19 mm.

Var. fuscospira. PI. 54, fig. 13, embryo (=var. d, p. 108).
Embryo cinnamon below, light buff above the periphery, a
white line below the suture.

Var. fuscozonata. PI. 17, figs. 2, 5 (=var. &, p. 108).

P. PHYSA KONANA P. & C., n. subsp. PI. 54, figs. 5, 5a.

The shell is smaller than physa, thin ; periphery marked
with a brown band ; above this it is zigzag-streaked with brown,
and cream-white, the pattern usually confused on the last

366 UNRECOGNIZED SPECIES.

whorl, distinct on the penultimate ; base indistinctly streaked
with whitish on a cinnamon ground ; interior russet.

(a) Length 14, diam. 8, aperture 7.9 mm.; 5% whorls.

(&) Length 15, diam. 9, aperture 7.9 mm.

Hawaii: North Kona at Honoula. Cotypes (a) in coll. A.
N. S. P., (6) in coll. Bishop Mus.

The brown band, unaccompanied by a white one, is special
to this form, which carries the area of the species well down
the western side of the island, though not so far south as P. p.
errans goes in the east. The types are old shells, worn at the
sum unit, and containing young. In another lot the shells are
dull, more or less corroded, and the color-pattern is almost
obliterated. The largest measures 15.3 x 9.2 mm.

P. 122, 3rd line from bottom, for * ' Waimanalu ' ' read Wai-
malu.

Unrecognized or undescribed Achatinellida: , etc.

Helicteres sulphuratus BECK, Index Molluscorum, 1837, p.
51 "I. oc. pacif.' Name only.

Helicteres leucozonalis BECK, t. c., p. 51, "I. oc. pacif.'
Name only.

Achatinella acicula, folliculus, unilamellata, lubricoides
and lubrica of Schliiter, Kurzgefasstes syst. Verzeichniss
meiner Conchyliensammlung, 1838, p. 8, are species of Ferus-
sacidce. Schliiter proposed Achatinella as a new genus for
these European snails in ignorance of Swainson's previous
use of the name.

Bulimus clausinus. " Shell dextral, ovate-conic, thick and
solid, white, smooth, glossy, sometimes with an obscure, nar-
row, yellowish band revolving with the suture, and passing
over the middle of the body-whorl. Whorls 5, convex. Aper-
ture subovate ; lip yellowish, thickened, coalescing with the
body-whorl and forming an enamel on the left side of the
aperture. Length about .7, diam. .4 inch. Hab. Hawaii'
(Mighels, Proc. Boston Soc. of Nat. Hist., ii, 1845, p. 20).

Placed as "L. clausiana Migh." in Leptachatina by Hart-
man (Proc. A. N. S. Phila., 1888, p. 52). Unknown to Sykes
and other authors. Possibly an Oahuan Bulimella, but it is

UNRECOGNIZED SPECIES. 367

certainly undeterminable and should be dropped, since
Mighels' types were destroyed by fire.

Achatinella ridua Pfr., Nomencl. Hel. Viv., 1878, p. 306, no.
31, error for A. vidua.

Achatinella tuba Gul., Pfr., Monogr., v, 167, error for
lutea Gul., a MS. name which Pfeiffer printed in the synon-
ymy of A. tappaniana. See Monographia viii, p. 217.

Achatinella turbinata Nuttall, Wahoo. Jay's Catalogue,
edit. 3, p. 58, 1839. Nude name.

liSpiraxis sandwichensis Pfr. Shell subperf orate, oblong-
turrite, rather solid, waxy ; spire turrited, rather obtuse ;
whorls 7%, a little flattened, plicate below the suture, the last
a little more than one-third the length ; columella compressed,
twisted ; aperture scarcely oblique, oval ; peristome simple,
the margins joined by a thin callus, the right margin some-
what dilated forwards, columellar margin somewhat reflected.
Length 9, diam. 3 mm. Aperture 3 mm. long, ~L~y2 wide. Cum-
ing collection' (Pfr., Proc. Zool. Soc. Lond., 1856, p. 335;
Monographia, iv, 575).

Mr. Sykes (Fauna Hawaiiensis, p. 399) thinks this a form
of Bulimus lactifluus Pfr., from Chili. If so, it must be a
young shell. Cf. Manual, Vol. XIX, p. 13.

"Bulimus kanaiensis Pfr. Shell subperf orate, conic-ovate,
rather solid, striatulate and irregularly malleate-impressed,
white ; spire conic, sometimes gray above, the apex obtuse ;
whorls 5, convex, the last scarcely exceeding the spire, ob-
liquely descending, rounded at the base ; columella slightly
arcuate ; aperture oblique, truncate-oval ; peristome simple ;
unexpanded, the columellar margin dilated, reflexed, some-
what appressed. Length 14, diam. 8 mm. ; aperture 8 mm.
long, 4% wide. Mus. Cuming" (Pfr., P. Zool. Soc., 1856, p.
332).

The name was corrected to kauaiensis in the Monographia
Heliceorum, iv, p. 469. Mr. Sykes states that it is very close
to Bulimus albicans Brod., of Chili, but he is "not quite sure
of the identity, as the shell is slightly more succineiform'
(Fauna Hawaiiensis, p. 399).

Achatinella faba Pfeiffer. " Shell imperforate, dextral,

368 UNRECOGNIZED SPECIES.

ovate, rather solid, irregularly striate, glossy, white ; spire
convexly conic, the apex rather acute ; suture simple ; whorls
5, a little convex, the last a little longer than the spire,
rounded ; coluinellar fold above, strong, nodiform ; aperture a
little oblique, obauriform; peristome thickly labiate within,
the right margin shortly expanded, columellar margin re-
flexed, adnate. Length 16, diam. ID1/*} mm. Sandwich Islands,
Dr. Frick in Cuming coll.' (Pfr.).

Achatinella faba PFR., Proc. Zool. Soc. London, 1859, p. 30 ;
Monographia, vi, p. 164. Cf. Sykes, Fauna Hawaiiensis, p.
307.

A lost species, which Mr. Sykes could not trace in the
British Museum. By the small size, convexly conic spire and
thick lip it seems referable to the fuscobasis series of Buli-
mella, though the proportion of length to diameter is more
like typical Achatinella ("Apex"). It is smaller than any
Bulimella except some of the fuscobasis group ; the size being
more like A. curia and its allies.

Like most of the Frick shells described by Pfeiffer, this was
apparently based upon an exceptional individual. Many
species have a white phase among other patterns. Such al-
binos are usually easy enough to classify because of their oc-
currence in colonies of colored shells; but a white shell, pos-
sibly of extreme or unusual shape, and without definite local-
ity, may sometimes baffle the most sapient Achatiiiellist.
When we have to deal with an unfigured form of which the
type is lost, there seems no reasonable ground for keeping it
out of the discard. It can never be positive^ identified.

Molluscorum Systema et Catalogus, Fr. Paetel, von Dr. L.
W. Schaufuss, 1869, contains the following names, without
references, and partly with a query in place of the authority.
Besides these there are numerous errors in the spelling of
other names, and wrong authorities are often given.
Achatinella agathal Sandw. I., p. 83.

bensoniat Sandw\ I., p. 83.

havaiana Rv., Sandw. I., p. 83.

magnificat Sandw. I., p. 83.

scamnata Fer., Sandw. I., p. 84.

UNRECOGNIZED SPECIES. 369

Achatinella subovata Fer., Sandw. I., p. 84.
torquatal Sandw. I., p. 84.
vulpina v. sinistra Fer., Sandw. I., p. 84.

Catalog der ConchyUen-Sammlung von Fr. Paetel, 1873.

Achatinella cinnamomea Fricke, I. Sandw., p. 105.

cingulata Fricke, I. Sandw., p. 105.

compressa Pfr., I. Sandw., p. 105.

grams Fer., I. Sandw., p. 105.

ignominiosus Pse., I. Sandw., p. 105.

impressa Pse., I. Sandw., p. 105.

octavula Pfr., I. Sandw., p. 106 (error for ob-

clavatat).
The same, edition of 1883.

Achatinella agatha Rv., I. Sandw., p. 153.

anacardiensis [no locality], p. 153.

circulata Frick, I. Sandw., p. 154.

hawaiana Gulck., I. Hawai, p. 154.
The same, 4th edition, ii, 1889.

Achatinella semitecta Fricke, I. Sandw., p. 275.

Achatinella olesonii Baldwin. Cat. Shells Hawaiian Is.,

1893, p. 5. Nmianu, Oahu. No description.

APPENDIX II.

GENEALOGY AND MIGRATIONS OF THE ACHATI-
NELLUXaE IN THE HAWAIIAN ISLANDS.

BY

ALPHEUS HYATT.

[NOTE. — The manuscript of this paper was found in Professor Hyatt 'a
desk after his death in 1902. In editing it, I have endeavored to alter
the original as little as possible, and have been greatly assisted by Dr.
C. Montague Cooke, of Honolulu, whose precise and extensive knowledge
of these shells enabled him to perceive Professor Hyatt 's meaning in
instances where the author 's statements were more or less ambiguous or
incomplete. — A. G. MAYER, Editor.]

(Jennie Arms Sheldon Fund Publication.}

PART I : GENERAL STATEMENTS.

THE writer's attention was first attracted to this subject by
Rev. J. T. Gulick and the acquisition of his first duplicate
collection by the Boston Society of Natural History in 1889
afforded him the means of beginning studies that have been
pursued more or less continuously since that date. The ma-
terials that have been used for study -consist of this collection
of 4154 specimens, 227 species and between 600 and 800
varieties as named by Mr. Gulick, and his personal collection
consisting of 9,000 specimens. The Oleson collection of
about 6,000 shells kindly lent by that gentleman, the C. M.
Gooke, Jr., collection of about 7,000 collected with extreme
care, and accompanied by a map showing the exact location
of each lot of specimens, also generously lent to me. There
is also a small collection presented by Dr. C. Montague Gooke,
a small but valuable collection presented by Dr. Conrad
Wesselhoeft, one from Mr. W. R. Castle, Jr., a small col-
lection purchased from D. D. Baldwin containing some im-

(370)

HYATT: GENEALOGY OP ACHATINELLID^. 371

portant species, the Boston Society's collection of about
4,000 shells and a collection of about 4,500 shells lent by
Eev. Harcourt W. Peck. I have also had the benefit of study-
ing the collections of the Smithsonian Institution and Na-
tional Museum, Academy of Sciences of Philadelphia and the
Museum of Comparative Zoology at Harvard, as well as tem-
porary loans for examination, the collections of Cornell Uni-
versity. Rev. J. T. Gulick, Mr. D. D. Baldwin and Mr. C. M.
Cooke, Jr., have also materially helped me by the benefit of
their great knowledge of the species in the field. To Hon.
W. DeW. Alexander, Rev. Oliver P. Emerson, Mr. A. B.
L/yon, the U. S. Coast Survey and to others I am also in-
debted for material assistance in procuring maps and in-
formation necessary for my work.

The land shells peculiar to the Hawaiian Islands are of
remarkable interest in connection with problems of evolution
and heredity, since they are an extensive group wholly con-
fined to these islands, and consisting of about 400 species.
Many of these exhibit permanent varieties, and are repre-
sented by immense numbers of individuals. The field is
practically occupied by these peculiar species to the exclusion
of other more cosmopolitan groups of land shells which are
represented by only about 111 species, and these are not so
numerous in individuals.

The Hawaiian Islands are arranged in an almost linear
arc of about 400 miles extending from Kauai and Niihau at
the northwestern, to Hawaii at the extreme southeastern end.
Their existence is due to volcanic elevations and eruptions.

The relative age of each island has been determined by
geologists, and I find that the relative age, and evolutionary
development of the faunas, is commensurate with the age of
each island.

By pursuing the study of the faunas closely, I determined
what appeared to be their natural course of evolution and
apparent succession. A study of the phylogeny of the faunas
appeared to demonstrate that Kauai, the northwesternmost
of the chain, had the most primitive fauna, Oahu having the
next in succession, while Molokai, Lanai and Maui although

372 HYATT: GENEALOGY OF ACHATINELLID^.

more closely associated have faunas derived from each other
in substantially similar succession, and by what seemed to be
a very recent migration. These results led naturally to the
conclusion that Kauai was the oldest, and that the others had
risen above the waters in the succession indicated by the
faunas. This was sound insofar as the age could be deter-
mined by the relative time at which each island had become
covered sufficiently by vegetation to be suitable for the exist-
ence of the more primitive forms of land shells.

The fact that Kauai contained the oldest fauna of this
group has already been stated by Baldwin and Gulick, and
it has been generally admitted that each island had its dis-
tinct fauna, and also that each of the valleys, in Oahu es-
pecially, was apt to have some peculiar varieties or species.

Turning then to the geology, for more information, I found
that Dana, Button, and others were in substantial agreement
with my conclusions, basing their opinions upon the topo-
graphy of the different islands and the comparative age of
extinct and active volcanoes and lavas. This analysis was
also carried into the separate parts of different islands.
Thus I became convinced that so far as the migration of shells
were concerned the western or Waianae range of Oahu was
more recent than the eastern or Konahuanui range, and in
Maui that the western mountain-peak of that island, Mt.
Kukui, was older than the eastern region around the base of
Mt. Haleakala. The last deduction has been confirmed by
geologists, notably Button, but the first remains thus far
undecided. So far as our present knowledge goes there is
then in these islands the most remarkable case yet known of
the evolution of a succession of -closely connected homogenetic
faunas which have evolved without serious interference from
the invasion of foreign importations. The Achatinellid(z
came to the island of Kauai, according to my deductions,
obviously before other islands had become habitable ; and they
came in the shape of certain shells of very distinct primitive
structure whose descendants are still found living in that
field. Then all of the members of that family group were
evolved from this island through the others in determinate

HYATT: GENEALOGY OP ACHATINELLHXE. 373

succession as clearly as if we could trace them, from older
to newer and still newer strata in the field of geology. The
groups of genera so far determined in the family Achati-
nellidce are as follows: [Cyclamastra P. & V.]; Kauaia
Sykes; Armiella Hyatt, Carelia and Amastra Adams; La-
minella Pfeiffer, Perdicella Pease. I have reduced these to
the above list because in each case the group that appears
as a synonym is not separable from that in which it is in-
cluded, but is linked with some of its species by intermediate
gradations, often by hybrids.

My definition of a genus in this family is a homogenetic
group which has been practically (not theoretically) traced
from variety to variety and species to species. I have on all
important points found myself in close accord with the Rev.
J. T. Gulick whose thorough studies of this group have placed
in my hands the means of doing this work, and whose labors
I hope some day to present with illustrations that will place
him among the great pioneers in the history of evolution.
A genus, in my opinion, is simply a genetic series which may
conveniently be separated from other closely related series
descended from the same common ancestors.

According to Major Button's account, (Report of U. S.
Greol. Survey, 1882-83, Hawaiian Volcanoes), the Hawaiian
Islands lie between 154° 30' and 160° 30' W of Green-
wich, and 18° 40' and 22° 15' North latitude. They are
therefore near the northern border of the great equa-
torial current which flows westward from the western
coast of North America sometimes transporting materials
from those shores to the islands. The length of the en-
tire chain is from 350 to 400 statute miles and the distance
from San Francisco about 2000 miles. There are eight
islands and four barren rocks. Niihau, a small island, now
without any testaceous inhabitants, lies at the northwest end
of the series, and is followed by Kauai having only terrestrial
land shells, and no arboreal species, in spite of the fact as
stated by Baldwin that "Its extensive forests, luxuriant vege-
tation and moist climate render it peculiarly well adapted for
the abode of Achatinella."

374 HYATT: GENEALOGY OF ACHATINELLID^E.

Kauai is nearly as large as Oahu, the metropolis of the
Achatinellida, but has a much simpler topography. There is
one peak of 5000 feet in the center of the island and the
valleys radiate from this outwardly, whereas in Oahu there
are two ranges, the longest of which, the eastern range, cuts
across the track of the northeast trade winds, while the
shorter or western range is nearly parallel with this. The
highest peak in the former is to the northeast of Honolulu
and is 3175 feet, while the highest peak in the western range
is 4030 feet.

The four islands next to Oahu geographically and faun-
ally, are a group by themselves, and can be treated as one ex-
cept in the minuter study of the distribution of species.
These are Molokai next to Oahu, Lanai and Maui, and the
small island of Kahoolawe apparently having no shells of
this family existing upon it.

The areas of these islands are as follows : Molokai, 190
square miles; Maui, 620 square miles; Lanai, 100 square
miles, and Kahoolawe, 60 square miles, — in all 970 square
miles.

Hawaii, the extreme southeastern island, is the largest and
highest, and has the only active volcanoes in the chain. Its
area is 3950 square miles. The recent and active volcanoes
of Hawaii, the presence of extinct craters on other islands,
the steady progress in size from northwest to southeast and
the deeper valleys and more sharply set ridges and peaks of
the northwestern islands, all tend to confirm the conclusions
of geologists that the Hawaiian islands arose in succession
through volcanic action, and by upheaval out of the water
starting with either Niihau or Kauai; Hawaii being the
youngest and largest of the group. Kauai is very nearly as
large as Oahu, and Hawaii is much larger, and yet both of
these extremes of the chain have meagre faunas compared
with Oahu, and the other intermediate islands.

This shows that there is as yet no correspondence between
the area or topography of each island and the abundance or
scarcity of Achatinellida. Baldwin expressly states that
Kauai is, apparently, extremely favorable for the develop-

HYATT: GENEALOGY OF ACHATINELLHXE. 375

ment of Ackatinellidce, and one would naturally expect to
find here a larger and if possible, higher development of the
family. But we are doomed to disappointment; the island
yields no arboreal species; the shells are all terrestrial —
Hawaii is also favorable, for, as Baldiwn states: "Its ex-
tensive forests are as well adapted for the support of Achati-
nella as those of any of the other islands. ' Nevertheless, it
has a very small fauna.

Niihau, lying to the southwest of Kauai, has an area of
90 square miles, and the highest peak is only 1500 feet, and
so far only one species of a sub-fossil shell has been found
there, Carelia sinclairi Ancey, which is extinct at the present
time. There are, however, certain correlations between the
topography and the distribution of these shells, already noted
by Baldwin (Hawaiian Almanac) which are of great interest,
for he states : ' ' That on a mountain chain with many culmin-
ating peaks the tendency is to a divergence of species ; while
on an individual mass of mountains concentrating towards
a single peak, the tendency is to a convergence of species.'
He also observes in treating of Molokai : * * It is about one-
third of the size (area) of Oahu, and like it has a mountain
range extending nearly thirty miles through its length. The
range is furrowed on both sides with deep valleys. Some
of these mountain gorges are very wide, and cut deep into
the narrow axis of the island. The larger ones have proved
an effectual barrier to the migrations of shells. The island
is thus divided into three natural sections, and each section
retains its own peculiar species without intermingling with
those of the next section. '

There is also a decided correlation between the relative
ages of the different islands, assuming that Kauai is the
oldest of those having living shells, and the kind of shells
that are found upon them; and also it seems obvious that
different genera came into existence in ascertainable succes-
sion upon different islands.

This succession appears to accord, in all cases so far as
examined, with the genetic relations of these genera as de-
termined by a method which the writer has used successfully

376 HYATT: GENEALOGY OF ACHATINELLUXE.

for many years, and happily succeeded in introducing to the
favorable notice of the younger paleontologists of this coun-
try. This is founded upon what has 'been falsely called ' ' the
Law of Biogenesis' by Haeckel, and in spite of just opposi-
tion and well-founded criticism, adopted by a large number
of scientific workers. This law was really discovered by von
Baer, and completed by Louis Agassiz and Vogt and was
simply baptized by Haeckel with an old name already having
a fixed and a different meaning in scientific literature.*

Natural selection might be called upon to account for the
brown protective colors of Amastras living on the ground, but
the prevalence of brilliant, conspicuous colors in the arboreal
species, and the presence of similar brilliant colors in some
species that are habitually concealed, and the constant ten-
dency to the repetition of similar variations in different
species are all at variance with this hypothesis.

There are also other constant modifications of form that
correlate with the general morphology of Gastropoda. The
ontogenetic development of a gastropod follows certain gen-
eral lines of modification. The protoconch is more or less
bag-like, and when the conch is begun it necessarily starts as
a tube continued from the aperture of the protoconch. In
shells that revolve with the principal axis held continually
in the same plane or nearly in the same plane, like Planorbis,
two hollows, or nearly equal umbilici, are formed, one on
either side of the shell. On the other hand, unequal or
asymmetrical spirals are formed by excessive growth to one
side which necessarily elevates the opposite side into the apex
of a spire, and generally obliterates the umbilical 'cavity.
Sometimes this remains for a time, but in most gastropods it
may be said to be a lost character. Nevertheless in the most
acutely spiral shells, as a rule, the earliest stages of the conch
are less asymmetrical than the subsequent states.

In other words, the ontogeny shows that the primitive form

* Abiogenesis was originally used for spontaneous generation of life
from inorganic matter and " biogenesis " for the theory that life was
continuous and that organisms could originate or be generated only
from life.

HYATT: GENEALOGY OF ACHATINELLHX-E. 377

is equi-umbilicated. While the asymmetrical is a secondary
modification occurring in more specialized organisms. In the
ontogeny, also, of asymmetrical forms we find that the young
are more loosely coiled than the adults. The umbilicus is
frequently open in the nepionic and neanic stages when in
later stages it is replaced by a eolumella or solid axis formed
by the contact of the inner side of the whorl.

The Achatinellidce are wholly asymmetrical, but the most
primitive genus contains forms that are flat spirals having
open umbilici on the lower side. This leads in Kauai itself
into a series of more closely coiled and more asymmetrical
spires having solid columellag in adults ; and these into others,
of the genus Carelia, having elongated acute spires that can
be compared with those of highly modified shells like
Turrit ella.

Primitive Amastras, closely affiliated with the primitive
forms above described, that are not unlike a common Helix
in aspect, show traces of their derivation in their low spires
and so-called perforated columella. This perforated colu-
mella is in reality an umbilicus reduced to the dimensions
of a tube ; and even on Kauai this perforation disappears in
adults of one or two species. The fossil Amastras of Oahu
are apt to have perforated columellae; and on all of the
islands the most primitive forms exhibit more or less of this
character. In each series there are species and varieties that
can be distinguished by their more complete asymmetry.
This is shown in their more solid and twisted columella,
and in their more slender and elongated spires. There is
one entire series in Oahu which has all of the species of this
type, and one of them bears the appropriate name of
Amastra turritella.

Turritelloidal shells appear repeatedly both less markedly
as variations and more decidedly as species on all of the
islands.

All of these have a highly acute elongated spire quite dif-
ferent from that which is commonly seen in Amastras that
live exclusively on the ground, and on this account they have
been placed by Hartman in the genus Laminella. Laminella

378 HYATT: GENEALOGY OF ACHATINELLID^E.

proper is a highly colored group of arboreal shells, in which
we may include Perdicella which is also very nearly connected
with Amastra, through species like Laminella gravida of
Oahu that have spires of dark brown similar to those of
some shells of Amastra rubens, cylindrica, etc., that have a
coarse, friable periostracum and elongated spires showing a
decided tendency to evolve turritelloidal forms. The basal
volutions are more globose and the apex more acute in
gravida, but this species is an intermediate form in other
respects leading not from the semi-arboreal turritelloidal
Amastras into Laminella, but from the turritelloidal ground
shells into this genus or group, which includes the very dis-
tinct and highly colored Lam. sanguinea and other forms
with turritelloidal spires that are found mostly in semi-
arboreal habitats on low bushes, ferns, etc., according to
Cooke and Gulick.

The apex in the nepionic stage is smooth in Laminella and
of the same dark horny aspect as it is in most of the species
of Amastra on Oahu, and the columella is highly developed,
twisted and perforated, as it is in Am. turritella and its
allies.

In the genus Amastra the shell is usually dark brown.
The species live on the ground and are present in all the
islands of the Hawaiian group that are inhabited by land
shells. The species, as in all other genera, are so closely
connected that it is very difficult to separate them. They
have a single spiral fold on the lower edge of the columella.
This fold is not characteristic of the genus but is shared
in common with all the other genera of the family Achati-
nellidcz, occurring only in these islands. Nevertheless there
are two marked deviations from this homogenetic character-
istic, for in many individuals among a limited number of
species, all of the genus Amastra, there is a slight tendency
to form a second fold above this, but it does not take the
definite form of a tooth-fold except in some very rare ex-
amples, and it is in these an obviously sporadic development
having no genetic significance.

In about fifty thousand shells of this family this occurs

HYATT: GENEALOGY OP ACHATINELLIDZE. 379

in very few examples, all belonging to the genus Amastra.
One single example has 'been found in which the columella
had three definite tooth- folds, evidently an isolated, sporadic
variation. This last case appears never to have become
genetic in any series of forms so that one could call these a
distinct group or species. The case of the two or double
tooth-fold does not, however, appear in this way, but only
on the island of Lanai and in the one group of Amastra
biplicata Newc. It occurs, however, in so many shells and
in such definite and invariable succession that they are uni-
versally recognized as one species, Amastra biplicata. This
has long been known, but the remarkable interest and signi-
ficance of this fact has not been noted. It is one species in
the genus Amastra, or if another view be preferred, a new
genus in the family of Achatinellida departing in a new
direction so widely that it must be cited as an exception to
diagnostic, analytical description of that family.* Another
exception occurs in two species on one island, but in this
case the deviation consists in the absence of the single
spiral tooth-folds so universal otherwise in this family group.
The variation in this case is also heralded by an anticipatory
mutation which occurs in individuals of other species oc-
curring on other islands.

Another matter of interest in this connection is that the
shells showing mutations of the double tooth-folds, and the
absence of folds have no obvious or traceable connection
with the species characterizable by the possession of similar
and permanent variations. They appear to be of indepen-
dent origin in different genetic series occurring on different
islands. Thus while homogenetic in their own series in their
own locality, they are simply homoplastic repetitions of
common tendencies when compared with the mutations oc-
curring in other genetic series in other islands.

This indicates the possible beginnings in each case of en-
tirely distinct genetic groups that might under favorable

* Several species of Amastra and Laminella have a second eolumellar
fold.— H. A. P.

380 HYATT: GENEALOGY OF ACHATINELLID^E.

conditions be perpetuated as well marked genera or family
groups. It is obvious, also, that the isolated case of the
columella with three folds indicates another variation having
genetic possibilities. These have all, so far, been arrested in
their evolution, and we can speak of them with reference to
the phylogeny precisely as we describe the arrested develop-
ment of parts and characteristics in the life history or
ontogeny of the individual.

The color patterns have a similar history in this group.
The individual mutations range from a normal uniform
brown to a very light brownish-white, or in the opposite
extreme towards black, and the patterns under certain con-
ditions may show localized zigzag bars or straight transverse
bars, or revolving bands. All of these have a fuller expres-
sion as permanent characteristics in some species, but shells
displaying the mutation of revolving bands are almost as
rare as those showing the double tooth described above; and
there are only about three species of Amastra that can be
characterized as banded.

On the other hand, there is no comparison between the
tooth-folds and color patterns in other genera of this family.
In the arboreal forms, genus Achatinella proper, the colors
often vary in the same species from colorless shells, true
albinos, through intermediate grades of browns and greens,
often brilliantly banded, to dark uniform patterns often
brown or almost black.

There is in most species a constantly reiterated tendency
to swing between these two extremes, one being the absence
of all color and the other a very dark uniform pattern, while
the more normal shells may display brilliant color patterns
often taking the form of alternating bands. Here and there
one or the other of these mutations become genetic, and they
are apt to become more or less permanent and predominant
in some species ; but in no case is any special pattern so
general as to exclude mutations more or less parallel with
those of other species. This is obviously variability taking
place without the interference of natural selection or ap-
parently any selective laws depending upon the advantage

HYATT: GENEALOGY OF ACHATINELLID^E. 381

of any particular pattern to the species. There are no
known enemies which could be attracted by the colors of
these shells nor do they seek protection from any so far as
known. The only possible cause of limitation has been sug-
gested by Mr. Gulick. He thinks that the colors of each
species are made up by the crossing of a limited number of
varieties which migrated across the ridges and mingling in
new localities of neighboring valleys necessarily produced a
distinct combination of patterns and colors from those of
the parent species.

This may or may not account for the differences observed
between closely related species that occur in contiguous val-
leys which are separated by high ridges, but the fact that
there is parallelism in the color patterns in different species
occurring in different valleys, is easily observed. This
parallelism is also obviously divisible into homoplastic simi-
larities and genetic differences as shown in the predomin-
ance of some definite pattern or color in the same species ac-
companied usually by differences in proportion, size and
often direction of the spire.

The tooth-fold is also more constant in arboreal forms, and
no case of absence or of the presence of additional folds
has been observed. This greater permanency in the generic
and family characters is obviously similar to what "Williams
has observed, and which he considers as a secondary condi-
tion acquired through inheritance — whereas, the variable
ones noticed in the tooth-fold and columella of Amastra
would he explained as due to the greater force of inherent
variability in this more primitive genus.

This is true, as we have said, of all such primitive forms,
and still more remarkable examples of this law of acceler-
ated divergence are present, in these islands, in genera that
are more primitive than Amastra, but there is this significant
qualification: They occur in one island, Kauai, which is the
most ancient of the group ; they are survivals of a still more
ancient and variable fauna composed wholly of ground shells
which are divisible into several distinct genera. They occu-
pied a perfectly free field and evolved several large robust

382 HYATT: GENEALOGY OP ACHATINELLIDyE.

types of shells; but these all died out. Whereas, the small
insignificant Amastra and another still smaller genus of
ground shells, Leptachatina, that arose in the same fauna, or
migrated from island to island, persistently held their own.

Leptachatina is a glassy shell, smaller than Amastra, and
adapted for 'both living on the ground and very rarely on
plants in places unfavorable for the existence of other genera
of this family, such as in open country, and is now found on
all of the islands. This genus is also most abundant in a
fossil state, and is comparatively invariable. It maintains
its characteristics without any marked changes in all loca-
tions. Its characteristics are those of a specialized type and
there is no indication in its structure that it is the ancestor
of any other genus. On the other hand, there is strong
evidence that it is a modified descendant of the Kauaiian
species, Amastra nucleola Gould. We can therefore neglect
this series as having no further meaning in the phylogeny.

In Oahu, the island next in geologic age and geographic
succession to the southeast, the only shells of this ground-
living group found as fossils are Amastra and Leptachatina,
the genera first evolved on Kauai. On this island the mi-
grations extended in two directions ; first geographically along
the surface of the ground. Here they met with more varied
conditions than in Kauai but their habitat as long as they
occupied the surface alone was practically similar to that of
their ancestors on Kauai. They therefore remained the same
dark-colored ground-living shells, but evolved a number of
connected species to fill this field.

The trees were, however, open to them, and when they
ascended them we find that different series and genera arose
marking distinct concordant stages of evolution.

The genus Amastra as limited and defined by Gulick has
several series of forms on Oahu. I divided these from each
other by following the genetic lines that connected one species
with another through their similarities and intermediate gra-
dations and this was done without reference to any other
class of facts. Among these series of Amastra, or Amastra-
like forms, one was remarkably distinct in the possession

HYATT: GENEALOGY OF ACHATINELLHXE. 383

of an acute, elongated spire differing from all the others ex-
cept in so far as a few species were concerned. Subsequently,
upon looking into the literature, I found that Hartman had
referred these to Laminella and not to Amastra ; while Gulick
and others, including myself, had not been able to separate
them from Amastra. Still later and upon consultation with
J. T. Gulick, Dr. C. M. Cooke, Jr., and other collectors, I
found that these same species were collected upon low bushes,
trees and ferns, and might be properly termed semi-arboreal.
They are as follows: Amastra turritella Fer., nigrolabris
Smith, rudis Pfr., spirizona Fer., grossa Pfr., intermedia
Newc., variegata Pfr., and frosti Ancey. Smith reports in
his original description of Am. nigrolabris, that this species
is procured both on the ground and in trees.

There is also another group of Amastras that evolve into
highly turritelloidal shells, but some of these are found solely
as stated by Cooke upon the ground such as Amastra tristis, a
shell with peculiarly blunt apex and Am. seminigra Hyatt;
Am. rubens Gould; and Am. corneiformis Hyatt, the last
of these being quite as elongated and acute as Am. grossa
and others in the Am. turritella series

This series though Amastra rubens and cylindrica are ap-
parently the ancestors of a group having quite distinct and
more highly turritelloidal spires, and living on low bushes,
trees and ferns, according to Gulick and Cooke; and are
classified by Gulick, Hartman and 'Others with the genus
Laminella. This series contains Am. gravida Fer., sanguinea
Newc., tetrao Newc., picta Migh., bulbosa Gul., straminea
Rve., venusta Migh., depicta Bald., helvina Bald., citrina
Migh., remyi Newc., concinna Newc., and alexandri Newc.

The five first named have the peculiar periostracum of
Amastra, more or less covering the shell. Am. sanguinea is
blood red in color, and Am. depicta is sometimes highly
colored.

All of these Amastras have the dark, horn-colored, smooth
young shells that occur in the primitive forms of Achati-
nellida on Kauai, and in nearly all of the Amastras on the
island of Oahu, and while none of them exhibit a double

384 HYATT : GENEALOGY OF ACHATINELLID^K,

tooth, there is a decided tendency towards the evolution of
an additional spiral fold on the columella.

The topography of Oahu changes as one proceeds north-
wards. The highest peak in the eastern range is Mt. Kona-
huanui, 3105 feet, situated about 15 miles from the southern-
most point of the island. From this peak to the northern
end of the island there is a gradual lowering of the heights
until beyond Waikane valley, the <crest is only about 2360
feet high and the intermediate valleys are separated by
wooded spurs that do not oppose any serious obstacles to the
migrations of the species. It is obvious that this relatively
flat, low-lying region must have quite different conditions
from the much more broken southern part near Honolulu.

The crest of the eastern range shuts off from the interior,
the trade 'winds that blow strongly from the northeast for
nine months in the year. The crest therefore gives much
greater shelter to the inner or western sides of the hills.
These winds also beat upon the unsheltered eastern side of
the Konahuanui range and here the difference on this ac-
count alone is apparent in the difference in the vegetation
of the two sides; the eastern sides being scantily and the
western thickly covered with trees.

The Bulimellas of Oahu thrive high up on the ranges and
extend to the outer, wind-beaten, eastern or coast side of the
eastern range of the island between Waiahole and Hauula val-
leys. The Achatinellas, on the other hand, affect more shel-
tered locations, are unfavorably affected by this wind-beaten
side and the species become smaller and individuals less nu-
merous as one proceeds northward along the eastern side of
Oahu toward the northern parts of the island. Achatinella
was not very successful in crossing the middle parts of the
island to the western range, and not a single Bulimella suc-
ceeded in crossing the central plain of the island.

Apex, however, thrives in locations between Achatinella
and Bulimella, and while it was less successful than Ackati-
nella in crossing the crest of the Konahuanui range to the
coast side, it was on the other hand, the group best suited to

HYATT: GENEALOGY OF ACHATINELLHXE. 385

cross the central plain of the island and occupy the western,
or Waianae range where it is by far the most abundant shell.*

Bulimella has a rougher shell and more uniform coloration
in the southern parts of the range, but changes as it proceeds
northwards and especially on crossing the crest in the north-
ern part of the range. The species evolved in this region
are more numerous, larger, the shells smoother and more
brilliant in color and more elaborate in their patterns.

Achatinella throws, or rather did in Gulick's time (1853-
54) out a colony of species that crossed the crest to the coast
side, and the changes that occurred in this colony were very
significant. They consisted of comparatively small, bright-
colored, banded species which resemble those that occur to the
north and on the same side of the mountain range, but are
widely separated from them by a barren region having no
shells of any kind. The main body of Achatinellas marched
northward on the inner watershed of the eastern range of
Oahu, here and there evolving large, brilliant shells until they
reached the northern region of the island. There the shells
are smaller, and although still striped, are sensibly distinct
from their southern progenitors. These small shells sent mi-
grants across the range to the coast side, the same species
sometimes occurring on both sides of the range. The num-
ber of species of Achatinella in this region also sensibly
diminished, where as in Bulimella the number of species in-
creased.

The small northern species are so distinct that although
closely connected by gradations with the southern forms, a
limited number of them have been set apart by several au-
thors as a distinct genus named Eburnella by Pease I have,
however, not been able to separate any of them from their
southern affines. Some authors have also thought that cer-
tain of these small shells belonged to a genus Partulina that

* in 1898 Professor Hyatt published a short paper in ' ' Science ' ',
volume 8, p. 395, in which he concludes that the genera Bulimella, Acha-
tinella, and Apex of Oahu originated from a common ancestor, the
Achatinella phceozona of Kuliouou valley near the southern end of the
Konahuanui range of mountains.

386 HYATT: GENEALOGY OF ACHATINELLIDJE.

occurs on other islands 'but has so far as I know no repre-
sentative on Oahu. They are, however, directly connected,
and barely separated as distinct species from other accom-
panying species that are placed by the same authors in the
genus Achatinella. They have identical young, the slender,
highly-polished, smooth shells and the colors of Achatinella,;
and no trace of the zigzag and marbled patterns and peculiar
colors common in Partulina.

In general, it may be stated that the arboreal stations in
the Hawaiian Islands were occupied by highly colored shells
in which the coarse friable brown periostracum of the ground
shells is absent and the color patterns were highly complex.

The differences evolved in these distinct genera are thus
correlated with their different stations and distribution, and
they certainly appear to have arisen in connection with their
change of environment.

It seems obvious also that these could not have been evolved
had not the new fields and stations been open and unoccu-
pied. If we assume that the observed differences were ac-
quired in consequence of the migrations of the ancestral
forms of each group or species into new situations, the whole
complex association becomes apparently explicable.

It is also obvious in these islands that the variations are
not only coincident with migration into new fields and sta-
tions, but they are also limited to these geographically, and
species are not as a rule maintained in their original form
when new migrations take place.

Species are, however, traceable by hybrids or by grada-
tions into others along lines of unbroken continuity through-
out the eastern chain of Oahu in Bulimella, Achatinclla, and
Apex wherein they can be followed along the same lines of
migration to their termination in the western range. Fin-
ally, the same may be done in all the groups that range
from island to island, the connections of course being less
perfect than in forms living upon one and the same island.
The species, however, can be followed by groups and some-
times by graded variations so slight that particular forms can
be pointed out as the migrants that must have come from one

HYATT: GENEALOGY OF ACHATINELLID^E. 387

island to another. In other words genetic lines of descent
can be traced in the shells throughout the islands beginning
with Kauai, the oldest geologically, and following the whole
chain to the southeast until one arrives at the most recent
island of the archipelago, Hawaii.

This entire picture of variability appears therefore like
those that have taken place in past geologic times, and known
to occur when an animal type of primitive form finds itself
in a free field, either an uninhabited locality as Kauai must
have been when the first ground shells reached there, or a
new and unoccupied station, as the bushes and trees of Oahu
were, when the ground-shells or their modified descendants be-
gan to creep up on their stems.

The opportunities for expansion by the evolution of varied
types were certainly afforded by the surroundings, and it
must be acknowledged that this divergence into new groups
takes place along the open ways of migration. This is dif-
ficult to account for by any hypothesis that does not consider
the primitive Amastras as a plastic type the structure of
which was capable of being modified so as to occupy all the
available environments afforded by these islands. If this be
so, the various successful types evolved in distinct genera
certainly appear to have arisen as secondary modifications,
which could not have come into existence if the new fields
and stations had been already occupied or were inaccessible.
The variations are, as a rule, obviously still coincident with
and limited to the locations and islands in which they origin-
ated, and this correlation cannot be accounted for unless we
grant a causal relation between the surroundings and their
faunas.

PART II : REMARKS UPON RELATIONSHIPS BETWEEN GENERA.

[The more or less disconnected notes constituting the second part of
this paper must be of interest to students of the Achatinellidce. When
death overcame him, Professor Hyatt had all but completed his prepara-
tions to visit the Hawaiian Islands, the funds for such an expedition
having been provided by Mrs. Jennie Arms Sheldon. In certain cases,
doubtless, his ideas of the actual relationships between genera would
have been somewhat modified and all would have been amplified and
rendered more precise through a study of the animals in the field.

388 HYATT: GENEALOGY OF ACHATINELLIDyE.

These notes are, therefore, preliminary statements written by Pro-
fessor Hyatt, and which he expected would be modified by the studies
he hoped to perfect while in the Hawaiian Islands. — A. G. M.]

Remarks upon Partulina.

Sub-series of Partulina semicarinata: The first of the two
species in this sub-series, Partulina semicarinata of Lanai,
has a uniform light-colored pattern throughout life with
lighter colored young. The young and adult stage has a
prominent, well-formed ridge or keel that disappears only
when near the old aperture.

In Partulina hayseldeni, also of Lanai, there is a finely
colored young like that of P. virgulata, having broad white
shoulder baud and darker side to the volutions. This pat-
tern is common on the full-grown adults of some varieties
of Achatinella polita of Molokai and is found in the young of
varieties of mighelsiana. The ridge is prominent in the shell
but disappears earlier in the last volution. The tendency of
P. virgulata to have young with angulated sides until a late
stage, and the presence of albinos in which the apex has al-
most completely lost its colors, shows that this species belongs
with liayseldeni and not with variabilis or any more slender
form.

Partulina semicarinata is a dextral species which stands
alone. The apex is acute and the spiral similar to that of
typical shells and the general form, aperture, columella and
tooth also like typical Partulina, but the shell surface is as
smooth as in any Achatinellas ; the spire increases more
evenly and rapidly, making a stouter shell, and there is a de-
cided keel on the basal volution until a late stage of growth
when it suddenly and completely disappears. This keel has
not the sub-angularity so often noticed in the young of
various species but is a distinct prominent ridge. Both form
and ridge are like those of Kauaia until a late age, then the
latter disappears, the general aspect, the presence of striation
in the young that occurs only on the mature volutions of
Kauaia and columella, tooth and aperture are like typical
Partulina.

HYATT: GENEALOGY OF ACHATINELLID^E. 389

Part, hayseldcni Bald., is a shell having an apex like vir-
gulata, but becoming uniform light chestnut brown in later
stages and having bulimelloid aperture and columella. One
shell in Cooke's collection (No. 2204) has two keels until a
late age and another does not lose the keel entirely even
in extreme age. The shells of the sub-series virgulata are
uniform and banded, but the young in nepionic stages are
invariably banded with white on the shoulders and have
brown sides.

Partulina mighelsiana Pfr., constitutes a group of itself,
which we may call the mighelsiana series. The apex of Par-
tulina mighelsiana Pfr., has the colors of Achatinella and
Bulimella, but the form is more slender and more acute, and
like that of typical Partulina. There are no signs of zigzag
or marbled patterns, but instead the banded patterns prevail
as in Achatinella. In one species, Part, mighelsiana of
Molokai, the aperture and columella are constantly similar
to those of the Achatinella-like shells of the l>ella series, and
in subpolita of Molokai the form and characters and apex are
precisely intermediate between mighelsiana and polita. I
have examined altogether 53 specimens of Partulina mighel-
siana, including several varieties, without finding in the young
any trace of the barred or marbled pattern common in de-
generate forms of Partulina.

In Cooke's collection there is a fine series of varieties of
mighelsiana that exhibit intermediate forms between this
and Achatinella polita in full-grown shells. In this, and
especially in Gulick's collection, the only characters that
separate the young of some specimens are the somewhat lighter
color of the dark sides of the apex and the brown band on
the base; there is one shell in Gulick's collection in which
this last is the only difference, the apex having white shoulder
band and dark side precisely as in nepionic stage of A. polita.

In quite a number of Cooke's shells the white shoulder is
present in the young and the side of the volution underneath
is darker, and it is obvious in these characters and in the
smooth, highly polished shell, that the affinities with polita
are very close and indicate a common origin for both species.

390 HYATT: GENEALOGY OP ACHATINELLID^.

In A. polita in the Cooke collection (No. 2014), one shell
has the band on the base and looks decidedly like a transition
or a hybrid between polita and mighelsiana. This has also
the white young of mighelsiana with the nepionic colors of
polita and has the dark inner rim to the aperture so often
found in mighelsiana. Cooke has placed it with polita, and
in this I agree with him.

Partulina variabilis Series: These are Bulimelline forms
with the narrow acute flat-sided spire of Partulina. In varia-
bilis -of Lanai, the callus or shield is present only in the larg-
est shells and very often absent at all stages. This is transi-
tional in others in which the aperture is constantly buli-
melloidal. It is apparently an offshoot of P. mighelsiana and
so far as I can see does not connect with any other species
although coming very close to some in its general aspect.
The species are as follows : Partulina variabilis Newc.,
P. lactea Gul. of Lanai, and Part, nivea Bald., dolei Bald.,
and eburnea Gul. of East Maui.

Relationship between Partulina and Achatinella: The con-
clusion seems unavoidable that all true Partulinse exhibit in
their young stages direct affinity with Achatinella, and that
the genus was derived from Achatinella that migrated to
Molokai and probably began its existence on that island, for
I have traced direct transitions from Achatinella polita to
adults of A. polita in Cooke 's collection and the young in
Partulina mighelsiana in Molokai. These were found in
other collections showed very close affinities.

Three species of Acliatinella, A. bella Rve., polita Newc.,
and sul>2')olita, occur outside of Oahu on the island of Molokai.
These four species cannot be distinguished from Achati-
nella3 of Oahu by any character so far as I could ascertain,
for the columella is very similar to that seen in many species
on Oahu.

In East Maui there are three species of Achatinella, A.
anceyana Bald., nattii Bald, et Hart., and porcellana Newc.
In all three of these species there is the same tendency ob-
servable in varieties to imitate the aperture of Partulina while
others retain the thinner apertures and columellaB of true

HYATT: GENEALOGY OF ACHATINELLID^]. 391

Achatinella. All of them have the surfaces, apices, and gen-
eral aspect of true Achatinellas.

In these forms, therefore, outside of Molokai, there is a
constantly recurring tendency to imitate the thickened colu-
mella and apertures of Partulina, but this does not extend
to any alteration of the general aspect nor of the texture of
the surface as has been stated above, nor in the patterns and
colors, the colors being brilliant and banded throughout the
islands.

The characters of the apertures have caused most authors
to include either all or part of these in the genus Partulina,
but the colors and forms are, it appears to me, conclusive in
favor of the theory of direct connection with the unques-
tionable Achatinellas of Molokai, and if this be true, it fol-
lows necessarily that the Bulimella apertures are simply
parallelisms with Partulina and have consequently neither
genetic nor taxonomic significance except within the limits
of the series itself.

Achatinella nattii Bald, and Hart, has the typical apex and
general aspect of Achatinella in most specimens, but there
is a tendency in many shells to form a callous or shield which
often develops to be as large and well marked as in Partulina.
The same is true of the outer lip, that often becomes thick-
ened at the same time. So far as these characters are con-
cerned, this species or some of its varieties is a Partulina,
but all of the specimens have the apex and external surface
like Achatinella. All of these have either a uniform or
banded pattern in the young, never a cross-barred or zigzag
pattern.

Partulinella marmorata series: This group consists of shells
having an apex similar to that of Achatinella in shape but
with heavier longitudinal ridges which also persist through-
out the later stages of the shell. The surfaces in later stages
are also apt to have persistent transverse ridges of growth
much coarser than in Achatinella and like the latter in that
the longitudinal ridges also persist and usually cross them
even on the last volution. The roughness of the shells is
therefore a marked characteristic just as it is in the Partulina
virgulata series.

392 HYATT: GENEALOGY OF ACHATINELLID^.

The apertures vary from toothless to those with prominent
tooth-folds, but the shield on the columella and the structure
of the peristome occurs in almost all of these forms. A
marked characteristic is also the presence of bars or zigzag
lines of color in the early neanic stage or throughout life in
all the shells, as a rule except in extreme cases of albinism.

The shells of the Partulinella marmorata series have a
peculiarly "brown barred pattern either throughout life or in
the early neanic sub-stage. There are two sub-series ; the first
consisting of highly colored shells with elaborate marbled or
banded patterns that show affinity with Partulinella dubia
in these colors and in their young; but as a rule these have
large columellar tooth-folds. The second sub-series consists
of shells that have lost these more highly colored patterns
in their later stages in different degrees and have remnants
of these only on their young and finally only in some individ-
uals in the most degenerate species.

The sub-series of P. marmorata, shells with marbled or
banded pattern, is as follows: Partulinella proximo, Pse., red-
fieldi Newc., tessellata Newc., rufa Newe., from Molokai;
Partulinella crassa Newc., P. perdix Rve., pyramidalis GuL,
splendida Newc., and several others from West Maui ; Par-
tulinella marmorata Gould, plumbea GuL, grisea Newc. from
East Maui P. tessellata and rufa have either no tooth-fold
or a very slight one, and this condition is also found in
P. crassa*

Marmorata series, Sub-series of P. horneri: These consists of
the species heretofore included in Partulina that occur on
Hawaii. They resemble the toothless forms of Oahu, Molokai,
and Lanai, and are especially similar to P. crassa of Lanai.
The spire, however, is more concave and more acute and in-
creases more rapidly and more regularly, and the last volution
flares out with a more even and regular form. The colu-
mella may be open or closed, but is always toothless and has

* Partulinella is a new subdivision of Partulina proposed by Professor Hyatt
for species with the last embryonic whorl marked with protractive stripes ; the
shell otherwise like Partulina s. sir. P. marmorata may be taken for type. —
H. A. P.

HYATT: GENEALOGY OF ACHATINELLID/E. 393

a distinct shield 'Or callus. The peristome is apt to be ex-
panded but is not always thickened. The young have the
patterns of the young of the P. tappaniana series except in
degenerative or albino forms. The species are P. hawaiiensis
and horneri Bald., and physa Newc.

The eastern variety of hawaiiensis has such lively colors
and coarse barred pattern that it suggests that this series
may have arisen from migrants like the existing forms P.
zebrina and zebra of East Maui.

The relationship betiueen Partulinella dubia and the Mar-
morata group: The only clue that I at present possess with
regard to the origin of this group consists in the primitive
colors and characters of Partulinella dubia Newc. which is
found on Oahu. For a long time I regarded the species as a
migrant from some island to the eastward of its home that
had found its way there and 'become somewhat retrogressive
in colors and columella. This view can still be maintained,
but it seems more likely that Partulinella dubia is a remnant
of some primitive form. I was led to this conclusion by find-
ing in the Boston Society's collection a fossil Amastra, un-
fortunately without a label, which possessed decided resem-
blances to Partulinella dubia in its toothless columella and
general form. Its spire was, of course, Amastran and dis-
tinct, and the columella had a comparatively large perfora-
tion. It was similar to Am. antiqua Bald, in aspect, except
that the tooth was wanting. Fortunately Newcomb observed
in very old shells of dubia a tooth, was present and this en-
ables us to make the connection with the Marmorata series
more confidently. The transitional character of the habitat
'on bushes' is also notable.

The colors of dubia are distinct from any that occur in
Amastra, but the reticulated pattern and zigzag lines that
occur over the entire spire of Partulinella dubia is a primitive
character and in some varieties there are large bars that ap-
proximate to those of marmorata and perdix.

I have therefore provisionally supposed that dubia repre-
sents more nearly an ancestral stock of Partulinella than any
other species, and that the migrations of this series started

394 HYATT: GENEALOGY OP ACHATINELLIDvE.

from Oahu, as the first locality in which the group origin-
ated. Partulinella dubia of Oahu is probably a remnant
of an ancient series which probably sprang from large
Amastras having no tooth-folds and also possessing the zig-
zag or barred reticulated pattern not very widely different
from that of Am. transversalis.

The Sub-series of Partulinella tappaniana: These are
shells with white or light backgrounds and slightly banded
patterns resembling mighelsiana and those of the variabilis
series that are albinos, but in these every species has through-
out, or in some individuals, a pattern barred or marbled with
brown in the young.

Partulinella dwightii of Molokai and lignaria of West
Maui belong to both sub-series. The former connects with
P. crassa of Lanai. Some of its varieties are marbled through-
out life, and some are white except for the zigzag bars in the
earliest neanic sub-stage.

Some of the shells of Partulinella proxima of Molokai are
very close to this series as are also P. tappaniana Ad., at-
tenuata Pfr., terebra Newc., ampulla Gul. of West Maui, and
P. eburnea Gul., P. dolei Bald., and P. nivea Bald., of East
Maui. The difficulty in distinguishing highly retrogressive
shells of this sub-series from those of the variabilis series is
perhaps at present insuperable. Some of the P. tappaniana
series such as dolei and eburnea may belong to this sub-series.
That is to say, their completely albinized young may have
originated in the same way as the completely albinized young
of most shells in P. tappaniana Ad., for example, through the
non-appearance of the barred pattern in the individual.

Relationship between Perdicella and Partulinella: Sykes,
Fauna Hawaiensis, p. 329, has selected A. helena Newc., as
the type of Perdicella Pease, and I propose to follow him.

Perdicella helena of Molokai is one of the stoutest of the
dwarf forms composing this group. Its colors and patterns,
basal band and toothless columella, barred young, and finely
wrinkled surface due to the crossing of the transverse ridges
of growth by longitudinal ridges, show that it is a dwarfed
form derived from some species of the Partulinella marmorata

HYATT: GENEALOGY OF ACHATINELLID/K. 395

series ; and its nearest affines now living are PartuUnella
dwighti of Molokai and P. crassa of Lanai. The more elon-
gated species of this series are Perd. theodorei Bald, of
Molokai. This makes genetic connection with P. ornata
Newc., of West Maui and P. mauiensis Newc., of Maul.
These species have no tooth-fold or only very slight spiral
ridges on the columella and in the last three the forms are
much elongated and more turritelloidal than in P. Helena.
The parallelism with Carelia is very marked in form as well
as in the toothless columella.

The highly specialized habitat of helena is stated by New-
comb to be within the coil of the Ti (Dracaena) leaf at the
point where it starts from the stem, is exceedingly suggestive
when taken in connection with the dwarfed aspect of this
species and others of the same genus supposed to have origin-
ated from some such ancestor as P. proximo,. [P. helena or-
dinarily lives on leaves and twigs of numerous shrubs and
trees, H. A. P.]

Relationship bettveen Newcombia and Perdicella: The
genus Newcombia was described by Pfeiffer and the first
species mentioned under this name is Achat. helena Newc.,
from Molokai. Pease placed this species, and the two follow-
ing in other genera, and used the fourth species, Newcombia
cumingi Newc., as the type. If the author and Mr. Baldwin
are correct in their translation of the facts, this action of
Mr. Pease appears justified if the name can be maintained
for the extremely elongated, rough, sinistral shells having an
umbilical perforation and no specialized tooth-fold, but having
.a callous deposit or swelling at the base of the columella.
These three characters are always found in Newcombia
cumingi Newc. of West and East Maui and Newcombia new-
combiana Pfr., and plicata Mighels, cinnamonea, sulcata Pfr.,
and canaliculata Baldwin, all of which are from Molokai, the
metropolis of the genus. The genus is certainly limited to
Molokai and Maui. Its absence on the adjacent island of
Lanai is very remarkable and interesting but entirely in
character with its highly specialized form and unique
ornamentation.

396 HYATT: GENEALOGY OF ACHATINELLID^E.

The connection between Newcombia and Perdicella is shown
through the resemblances in color and form of the young of
Newcombia to those of Perdicella. They are as a rule more
slender than the most primitive species of that genus, viz.
P. helena, but are quite similar to the more modified and more
slender forms of Newcombia itself, like N. perkinsii Sykes
of Molokai.

The resemblances are better seen in the young of Newc.
cinnamonea and some of the smoother species than in the
more highly modified shells like sulcata, etc. This evidence
is strongly in favor of the direct derivation of Newcombia
from some form like the Perdicella helena of Molokai.

The Relationship between Amastra, Kauaia, [Cyclamastra],
Carelia and Armiella: Amastra, Kauaia, [Cyclamastra]
Carelia and Armiella in Oahu and Kauai have smooth vshells
in the earliest stages and there is a common form in the
nepionic stage. This has somewhat stout and rapidly in-
creasing volutions, the surface is covered with fine transverse
ridges or growth-bands. The bases are separated from the
p'lano-convex dorsal sides by a more or less prominent sub-
angulation, but this is never developed into a keel nor are
there any coarse, transverse costse on the dorsum. There are
also no longitudinal markings of any sort. Keels, when pres-
ent, and longitudinal ridges are introduced in late stages in
[Cyclamastra'], Kauaia, Armiella, and Carelia.

In Armiella the columella is only known in large shells
and in them it is solid.

In Carelia the columella is known in the young of C.
dolei wherein it remains solid or incomplete until a compara-
tively late stage and is solid also in the ephebic stage, but in
one shell there was a small perforation when the shell was
about one centimeter in length and was then just closing,
for it was entirely closed in the same shell when four mil-
lemeters longer. In some others it was present but very
minute at an earlier stage, and in some it seemed to be absent
altogether at every stage. It was present in two shells of
C. adusta when the shells were about six mm. in length (verti-
cal diameter) and closed at one centimeter. Its beginning
was not traced in these two shells.

HYATT: GENEALOGY OP ACHATINELLID.E. 397

In C. bicolor there is a very minute perforation in the
neanic stage and there are indications in another individual
shell of the presence of a somewhat larger perforation. The
columella in [Cyclamastra} is at first incomplete in the young
then becomes complete and remains complete, the umbilicus
being open throughout life.

In Kauaia the history is similar to that of Carelia in the
young, but the columella has a long straight ventral or inner
deposit, and during the period or stage of the completed
columella the umbilical opening is small and similar to that
of the Amastra rugulosa group of Kauai. In a more ad-
vanced age such as the ephebic and gerontic stages, the
columella again becomes solid, the umbilical opening closes,
and the deposits are less, and are plastered directly against
the axis.

The Amastrae of Kauai have a columella which is long and
straight in shells that retain an open umbilicus ; and the aper-
ture in such examples is similar to that of [Cyclamastra]
which never closes its umbilicus, and to the young of Kauaia
during the stage when the umbilicus is open. In shells that
close up the umbilicus, a variation that occurs apparently in
species having an open umbilicus, the columella and aperture
resemble that of Kauaia in its older stage and after its um-
bilicus closes,

In all these species the thick brown periostracum is a
marked characteristic, and a tendency to put on more lively
colors is only shown in a few species by bands, etc.
[Cyclamastra'] is obviously the nearest to the ancestral form
of Amastra. This has an open umbilicus, and form both of
spire and aperture and columella which is more or less re-
peated in the young of all other species in the neanic stage.

The gradation appears to be as follows: [Cyclamastra'] is
directly connected with Kauaia. Kauaia is similar to it
only in the young. Armiella is similar to Kauaia and
descended from it or from some common ancestor, but hav-
ing two longitudinal ridges on the dorsum, and a distinct
aperture. Amastra is derived from [Cyclamastra] which
the species resemble during a part or the whole of their

398 HYATT: GENEALOGY OF ACHATINELLHXE.

neanic stages. This common ancestor probably was not dis-
tinct from [Cyclamastra] -generically, but must have differed
in having a slightly less depressed or longer spire and smaller
umbilical perforation, and no distinct carination, the sides
being, however, distinctly angulated.

The young shells of these genera have a common form
which may in general terms be described as the nepionic or
baby shell. This has a 'comparatively smooth short spire with
open columella and broad, stout volutions showing a tendency
to angularity on the outer, median zone of the volution and
some times an incipient carination. This is particularly ob-
vous in species of Amastra, and in Achatinella phcuozona and
Acli. plumata, which are both closely allied species.

There is but one form among Achatinellidce that is geneti-
cally identical with Ach. ph&ozona and A. plumata. This
is Kauaia, a terrestrial genus of the island of Kauai. This
shell has until a very late, probably adult stage, a similar
stout form and visible carinations, and its brown color is also
in accord with the similar hues of many young shells. The
terrestrial genus Amastra is obviously in the direct line of
descent from Kauaia, and has similar colorations and uniform
pattern.

Achatinella ph&ozona, now extinct, was collected by Gulick
in the valley of Kuliouou immediately adjoining Niu. This
species shows characteristics that intergrade on the one hand
with Bulimella and on the other with Achatinella. It stands
at the focus of the affinities of these two groups as regards
the form of the shell, which is bulimelloid in some varieties,
and Achatinelloid in others; and in the apertures which are
plainly Achatinelloid. It also grades into A. plumata
through numerous hybrids.

Apex cestus and forbesianus are both in color and in pat-
tern more like Achat. plumata than any other species of the
same region, but differ in the characteristic, turbiniforrn
basal volution, the Bulimella-like aperture and the almost
constantly dark apex of Achat. plumata. Achat. plumata
has also the dark apex and is precisely similar to Ach. por-
cellana, which is almost white in the succeeding or nepionic

HYATT: GENEALOGY OP ACHATINELLID^. 399

whorls with similar longitudinal striae. The conclusion from
this and the fact that the young until a late stage have the
Achatlnelloid form and apertures is obviously in favor of
the opinion that Apex sprang from Achatinella and not from
Bulimella which it resembles more closely in the aperture.

The close resemblances in the pattern of coloration is in
favor of the derivation of Apex through A. plumata or some
closely related modification of this polymorphic species. We
can now assume as a working hypothesis that Achat. phccozona
not only lies at the focus of affinities of these genera but was
the surviving representative of their common ancestor. If
this be the case the young shells, being nearer to the assumed
ancestor, Kauaia, ought to carry closer reminiscences of this
progenitor and retain its aspect until a later stage of de-
velopment than Achat. plumata, Bulimella, or Apex.

This theoretical requirement is actually more exacting than
ought to be demanded in view of the fact that Ach. phceozona
is a more or less remote descendant of this ancestor or
proachatinellan shell. Like other actual cases of this kind, it
might reasonably be expected that it would have lost or
skipped in its development many of the characters of this
ancestor. Nevertheless, even the most exacting requirements
of the working hypothesis are fulfilled in the ontogeny, and
not only the form until a late stage of development is similar
to that of Kauaia, but the horn-brown and uniform pattern
of that genus is also recapitulated in those varieties of Ach.
phceozona, that are not banded.

EXPLANATION OF PLATES.

NOTE. — When not otherwise stated, the specimens figured
are in the collection of the Academy of Natural Sciences.

FIGURE PAGE

PLATE 1. Newcombia.

1. Newcombia plicata (High.). 57844 2

2, 3. Newcombia plicata (High.). 2026 Cooke coll 2

4, 5. Newcombia costata Borch. (=N. p. gemma). After

Borcherding 4

6. Newcombia plicata (High.). 57842 2

7. 8, 10. Newcombia plicata gemma (Pfr.). 66541 .... 3
9,12. Newcombia cinnamomea (Pfr.). 57841 and 66121. 10

11. Newcombia plicata gemma (Pfr.). 2028 Cooke coll.. 3

PLATE 2. Newcombia and Partulina.

1-3. Partulina carinella (Bald.). Cotypes. 109052. ... 7

4, 8, 14. Newcombia pfeifferi (Newc.). 2021 Cooke coll. 13

5-7. Newcombia canaliculata (Bald.). Cotypes. 65713. 6

8. Newcombia pfeifferi (Newc.). 109908 13

9, 10. Newcombia sulcata (Pfr.). After Borcherding. .. 5
11, 12. Newcombia p. ualapuensis (Pils.). Type. 94490 12

13. Newcombia cumingi (Newc.). 57846 10

14. Newcombia pfeifferi (Newc.). 109908 13

PLATE 3. Newcombia.

1-3, 5. Newcombia cumingi (Newc.). 92480, Wailuku,

Gulick 10

4. Newcombia cumingi (Newc.). 92670, Makawao, Gu-
lick 10

6. Newcombia cumingi (Newc.). 2139 Cooke coll., Ma-

kawao 10

7-9. Newcombia cinnamomea (Pfr.). 57841 10

10, 11. Newcombia cinnamomea (Pfr.). 66538 10

PLATE 4. Perdicella.

1-6. Partulina helena (Newc.). 109054 A. N. S. ; 2017,

2119 Cooke coll 16

7. Partulina helena balteata Pils. 109053 17

(400)

EXPLANATION OF PLATES. 401

FIGURE PAGE

8-10. Partulina ornate (Newc.). 92707 A. N. S. ; 2106

Cooke coll 18

11. Partulina mauiensis (Pfr.). 2107 Cooke coll 20

12-14. Partulina mauiensis (Pfr.) . 57773 20

PLATE 5. Perdicella.

1-3, 7. Partulina fulgurans (Sykes). 66550 21

4. Partulina fulgurans (Sykes). After Sykes 21

5. 8, 9. Partulina zebrina (Pfr.) . 68875 20

6. Partulina helena Nc. (Type figure of A. minuscula

Pfr.). After Pfeiffer 18

10-12. Partulina zebrina Pfr. 2103 Cooke coll 20

PLATE 6. Partulina, Molokai.

1. Partulina virgulata (Migh.). After Borcherding. . . 25

2. Partulina virgulata (Migh.), var. 106057 27

3-6. Partulina virgulata (Migh.). 92691, 106056 25

7. 8. Partulina v. halawaensis Borch. After Borcherd-

ing 27

9. Partulina virgulata, var. Ualapue. After Bor-
cherding 26

10. Partulina virgulata, var. Pelekunu. After Bor-

cherding 26

11. Partulina v. halawaensis Borch. 1851 Cooke coll. . . 27

12. Partulina tessellata (Newc.). Specimen from New-

comb 28

13-15,20. Partulina tessellata (Newc.). Kahanui. After

Borcherding 28

16,18. Partulina tessellata (Newc.). Makakupaia. After

Borcherding 28

17. Partulina tessellata (Newc.) . 106058 28

19. Partulina tessellata (Newc.). Kealia. After Bor-
cherding 28

21. Partulina tessellata (Newc.). 2029 Cooke coll 28

PLATE 7. Partulina, Molokai.

1, 2. Partulina tessellata meyeri Borch. After Bor-
cherding 29'

3. Partulina rufa (Newc.) . 1921 Cooke coll 29

4. 5. Partulina rufa (Newc.). 106046-7 29

6,7. Partulina rufa (Newc.). var. After Borcherding .. 29

8. Partulina rufa (Newc.). var. 106048 29

9. Partulina rufa (Newc.). 1915 Cooke coll 29

10, 11. Partulina rufa (Newc.) . 106049 29

402 EXPLANATION OF PLATES.

FIGURE PAGE

12, 16, 17. Partulina redfieldii (Nc.). 2068-71-74 Cooke

coll 38

13, Partulina rufa var. Idas Borch. 106050 31

14, 15. Partulina rufa var. idae Borch. After Borcher-

ding 31

18, 19. Newcombia perkinsi Sykes (= philippiana Pfr.).

Cooke coll 8

PLATE 8. Partulina, Molokai.

1. Partulina dwightii (Newc.). Puukaeha. After Bor-

cherding 35

2. Partulina dwightii (Newc.). Kawela. After Bor-

cherding 35

3. 4. Partulina dwightii (Newc.) . 106062 35

5. Partulina dwightii (Newc.). Specimen from New-

comb 35

6. Partulina dwightii (Newc.). Kawela. After Bor-

cherding 35

7. 8. Partulina dwightii color-form compta Pse. 106059-

60 36

9, 10, 12. Partulina d. color-form concomitans. Maka-

kupaia. After Borcherding 37

13. Partulina d. color-form concomitans. 106061 37

14. Partulina redfieldii (Newc.). After Newcomb 38

15-20. Partulina redfieldii (Newc.). Specimens from New-
comb 38

PLATE 9. Partulina, Molokai.

1, 2, 4, 7, 8. Partulina proxima Pse. 106051-55-52 32

3, 5, 6. Partulina proxima Pse. 1956-60-62 Cooke coll. . . 32

9. Partulina proxima Pse. var. 106053 33

10, 11. Partulina p. schauinslandi Borch. After Bor-
cherding 33

12, Partulina p. multistrigata Pils. 109844 34

13, 16. Partulina p. multistrigata Pils. 1928, 1930 Cooke

coll 34

14, 15. Partulina theodorei Bald. Cotypes. 65710 33

17. Partulina dwightii mucida Bald. Cotype. 65708. . 34

18. Partulina d. mucida form macrodon Borch. 106054. 35

19. 20. Partulina d. mucida form macrodon Borch. After

Borcherding 35

PLATE 10. Partulina, Maui.

1. Partulina marmorata Gld. After Gould, U. S. Expl.

Exp 42

EXPLANATION OF PLATES. 403

FIGURE PAGE

2-4. Partulina marmorata Old. 92666 A. N. S. ; 2119

Cooke coll 42

5-7. Partulina kaaeana Bald. Cotypes 41

8-12. Partulina plumbea Gul. 66543, 92671, Kula,

Gulick 43

13. Partulina grisea (Newc.). Specimen from Newcomb. Ill
14-16. Partulina perdix (Rve.). Specimen from Lahaina. 45
17-19. Partulina undosa Gul. (== perdix). 92728. Wai-

hee, Gulick 45

20. Partulina perdix var. pyramidalis Gul. Lahaina, Gu-
lick 46

PLATE 11. Partulina, Maui.

1-9. Partulina splendida (Newc.). 92711. Wailuku,

Gulick 51

10. Partulina splendida (Newc.). 92702. Lahaina,

Gulick 51

11, 12. Partulina s. baileyana Gul. 92719. Wailuku, Gu-

lick 52

13. Partulina c. baileyana Gul. Cooke coll 52

14. Partulina gouldii (Nc.). After Newcomb 52

15. 16. Partulina gouldii (Nc.) . Maui 52

17. Partulina talpina Gul. (= gouldii). 92714. Wai-
luku, Gulick 53

18-21. Partulina gouldii var. perfecta Pils. Wailuku,

Baldwin 54

PLATE 12. Partulina, Maui.

1. Partulina tappaniana C. B. Ad. Wailuku, Gulick.

92718 54

2, 3. Partulina tappaniana C. B. Ad. Lahaina, Gulick.

92701 54

4. Partulina t. ampulla Gul. Copy of type figure 57

5. Partulina t. fasciata Gul. Honukawai, Baldwin .... 56

6. 7. Partulina t. fasciata Gul. Honukawai, Gulick.

92724 56

8. Partulina nivea Bald. Cotype. A. N. S 59

9, 10. Partulina t. eburnea Gul. Honuaula, Gulick.

92663 57

11, 12. Partulina t. eburnea G;ul. Cooke coll 57

13, 14. Partulina t. carnicolor Bald. Cotypes. A. N. S.. 58

15-18. Partulina dolei Bald. Cotypes. A. N. S 60

19, 20. Partulina lemmoni Bald. Cotypes. A. N. S 61

404 EXPLANATION OP PLATES.

FIGURE PAGE

PLATE 13. Partulina, Maui.

1-3. Partulina winniei. Bald. Cotypes. A. N. S 44

4. Partulina denselineata Rve. (=radiata). Conch.

Icon., fig. 9 51

5. Partulina ustulata Gul., var. Kahoma, Gulick. 92726. 47

6. Partulina ustulata Gul., copy of type figure 47

7. Partulina induta Gul. Wailuku, Gul. 92712 48

8. 9. Partulina radiata Gld. A. N. S 49

10-13. Partulina perdix Rve. Honokawai, Thaanum .... 45

14, 15. Partulina crocea Gul. Waihee, Gulick. A. N. S. 65
16. Partulina nivea kaupakaluana Pils. Cotype. Cooke

coll 60

PLATE 14. Newcombia, Perdicella.

1. Newcombia cinnamomea Pfr. Moanui, Thaanum .... 12

2-4. Newcombia c. decorata Pils. Kupeke, Thaanum ... 12

5. Newcombia canaliculata wailauensis Pils. Cotype . . 7
6, 7. Newcombia c. honomuniensis Pils. Cotypes 12

8. Partulina kuhnsi Pils., var. Honokawai 23

9-11. Newcombia sulcata Pfr. Ahaino 5

12-15. Partulina kuhnsi Pils. Cotypes. Honokuhua ... 22

PLATE 15. Partulina, Maui.

1. Partulina terebra Nc. From Dr. Newcomb 61

2. Partulina terebra Nc. Copy of original figure 61

3. Partulina t. attenuata Pfr. Copy of original figure. 63
4-8. Partulina terebra Nc. Waiehu, Gulick. A. N. S. . . 61

9. Partulina t. corusca Gul. 57769 62

10. Partulina fusoidea Newc. Am. Journ. Conch., II. . . 64

11. Partulina fusoidea Newc. Ukumehame 64

12. Partulina t. longior Pils. 92721 63

13-17. Partulina t. lignaria Gul. Wailuku, Gulick. 92717. 63

PLATE 16. Partulina, Hawaii.

1. Partulina confusa Sykes. From Dr. Newcomb 105

2-10. Partulina confusa Sykes. Near Mana, Thaanum. 105

11. Partulina confusa Sykes. Embryo from fig. 2 106

12. Partulina confusa Sykes. Embryo from fig. 8 106

PLATE 17. Partulina.

1, P. horneri Bald. Cotype. Hamakua. 65702 107

2, 5. P. horneri fuscozonata P. & C. Hamakua. 65700. 107

3, 4. P. horneri Candida. Above Kukuihaele, Thaanum.

108203 107

6. P. physa Newc. Copy of type figure 109

7. 8. P. physa (Cotype of A. hawaiiensis). 65695 109

EXPLANATION OF PLATES. 405

FIGURE PAGE

9-12. P. physa. Waimea Plains, Thaanum ....'. 109

13. P. physa (hawaiiensis Bald.). From Baldwin 109

14. P. physa errans Pils. Near Pahoa Ill

15. 16. P. physa errans Pils. Kawaiiki Ill

PLATE 18. Partulina.

1, 3, 4. Partulina crassa Newc. Coll. by Thaanum .... 40

2. Partulina crassa Newc., var. 92673, Gulick coll 40

6, 7. Partulina thaanumiana Pils. Cotypes 112

8, 9. Partulina grisea Newc. Specimens from Newcomb. Ill

10-17. Partulina mighelsiana Pfr 77

PLATE 19. Partulina, Maui.

1-3. Partulina porcellana Nc. Nahiku, Baldwin 69

4. Partulina porcellana Nc. Copy of original figure ... 69

5-11. Partulina p. flemiiigi Bald. Cotypes 71

12. Partulina p. wailauensis (cotype of cooperi Bald.) . . 72

13, 14. Partulina p. wailauensis Sykes and var. After

Sykes 72

15, 16. Partulina p. fulvicans Bald. Cotypes 73

PLATE 20. Partulina, Maui.

1-7. Partulina mutabilis Bald. Cotypes 68

8-11. Partulina nattii Bald. & Hartm. Topotypes from

Baldwin 73

12-14. Partulina aiiceyana Bald. Cotype 75

15. Partulina germana Newc. Copy of type fig 76

PLATE 21. Partulina, Lanai.

1-4. Partulina semicarinata Newc. Main ridge, Lanai,

Thaanum 86

5-7. Partulina hayseldeni Bald. Cotypes 88

8-12. Partulina hayseldeni Bald. Lanaihale, Thaanum. 88

13. Partulina semicarinata Newc. Copy of type figure. 86

14. Partulina variabilis Newc. Copy of type figure .... 83

15. Partulina semicarinata Newc. Embryo from fig. 1 . . 86

16. 17. Partulina variabilis Newc. Embryo 85

PLATE 22. Partulina, Lanai.

1-3. Partulina variabilis Newc. Specimens from New-
comb

4-9. Partulina variabilis Newc. 92674, Gulick coll. ... 83

10-12. Partulina variabilis Newc. 108198, Thaanum coll. 83

13, 14. Partulina v. fulva Newc. 66540 85

15, 16. Partulina v. lactea Gul. 108197 86

406 EXPLANATION OP PLATES.

FIGURE PAGE

PLATE 23. Partulina, Molokai.

1. Partulina subpolita Hyatt & Pils. Type. 106992. . 359

2-8. Partulina m. bella Rve 79

9-11. Partulina m. bella Rve. 1942-6-7 Cooke coll 79

12. Partulina m. bella Rve. Copy of original figure 79

13. Partulina rnighelsiana var. latizona Borch. After Bor-

cherding 82

14. 15, 16. Partulina mighelsiana var. dixoni Borch. After

Borcherding

17, 18. Partulina mighelsiana var. hepatica Borch. After

Borcherding 83i

PLATE 24.

1-4. Partulina polita Nc. A. N. S. ; fig. 2 from Newc.. . 80

5, 6. Partulina polita Nc. Mapulehu. 108179 80

7, 8. Partulina montagui Pils. Cotypes, A. N. S. and

Spalding coll 66

9. Partulina dubia Newc. Copy of original figure .... 113
10-13. Achatinella phaeozona Gul. Keawaawa, Gulick.

A. N. S. and Boston Soc 184

14. Achatinella taeniolata Pfr. Waialae, Gulick. A. N. S. 130

15, 16, 19. Achatinella taeniolata Pfr. Palolo, Gulick. A.

N. S 130

17, 18. Achatinella tseniolata Pfr. Maunawili, Thaanum.130
20, 21. Achatinella casta. Copies of original figs, of ligata

Sm 238

PLATE 25.

1. la. Achatinella viridans Migh. Manoa, Gulick .... 125

2. Achatinella viridans Migh. Palolo, Gulick 125

3. Achatinella viridans Migh. Manoa, Cooke, 348 125

4. Achatinella viridans Migh. Nuuanu, Cooke, 791 .... 125
5-5d. Achatinella viridans subvirens Nc. Waialae, Gu-
lick 129,

6. Achatinella v. subvirens Nc. Wailupe, Gulick 129

7-76. Achatinella v. subvirens and tseniolata. Waialae,

Gulick 129

8. Achatinella viridans rutila Nc. Waialae nui, 145

Cooke coll 128

9-9&. Achatinella v. subvirens Nc. Palolo, Gulick .... 129

10. Achatinella v. rutila Nc. Niu 128

11-lle. Achatinella v. rutila. Niu, Gulick 128

12. Achatinella v. rutila. Wailupe, Gulick 128

13-13c. Achatinella taeniolata Pfr. Wailupe, Gulick .... 130

EXPLANATION OF PLATES. 407

FIGURE PAGE

PLATE 26. Partulina.

1, la. P. virgulata var. Mapulehu, Thaanum 358

2, 2a. P. v. halawaensis Bald. Kapookoholua, Thaanum. 359

3, 3a. P. v. kaluaahacola. Kaluaaha 359

3, 4a. P. redfieldi kamaloensis P. & C. Kamalo 362

5. 5/. P. redfieldi X dwightii. Puu Kolekole 360

6. P. dwightii oecidentalis P. & C. Moomomi 361

7. P. dubia Nc. From Newcomb 113

8. P. dubia Nc. Waianae, 1777 Cooke coll 113

9-9Z>. P. dubia. E. ravines Waiawa, Spalding 113

10. 100. P. dubia var. Waimano, 2181 Spalding coll. . . 113

11. P. dubia (type of A. platystyla Gul.) 116

12. P. dubia (type of A. pexa Gul.) 116

13-14. P. m. bella Rve. Puunea, Cooke & Pilsbry 79

15-15e. P. m. bella. Ualapue, Thaanum 79

PLATE 27. Achatinella.

l-le. A. byronii Wood. Ahonui, Gulick. 92425 133

2. A. pulcherrima Sw. Ahonui, Gulick. 92422 140

3. A. byronii. Kalaikoa, Gulick. 92312 . . . 133

4-4d. A. pulcherrima ("mahogani"). Ahonui, Gulick.

92427 143

5-6. A. pulcherrima. Wahiawa, Gulick. 92624 140

7-7b. A. pulcherrima nympha. Wahiawa, Gulick. 92622. 144

8. A. b. rugosa Nc. Waiawa, Gulick. 92230 135

9, 9a. A. b. rugosa. Waimalu, Gulick. 92234 135

10, 100. A. b. rugosa. Waipio, Gulick. 92260 135

PLATE 28. Achatinella (Bulimella) .

1-ld. A. bulimoides ovata Nc. Kahana, Gulick. 92400. 160

2. A. bulimoides ovata Nc. Hakipuu, Gulick. 92448. . 160

3, 4. A. bulimoides rotunda Gul. Kaaawa, Gulick.

92264-82 163

5-6. A. bulimoides rotunda Gul. Kahana, Gulick.

92401-60 163

7. A. bulimoides ovata, albino. Kahana, Gulick. 92463. 160
8-9a. A. bulimoides obliqua Gul. Kahana, Gulick.

92404-65 158

10-10c. A. bulimoides oomorpha Gul. Kahana, Gulick.

92402 159

11-11&. A. bulimoides wheatleyana Pils. Punaluu, Gu-
lick. 92449 168

12. A. elegans Nc. Specimen from Newcomb 166

13-13d. A. elegans Nc. Hauula, Gulick. 92446 166

408 EXPLANATION OF PLATES.

FIGURE PAGE

PLATE 29. AchatineUa, etc.

(Photographically reproduced from Newcomb's plate 22,
P. Z. S., 1853.)

1. AchatineUa gouldii Newc. (=Partulina gouldii) . . 52

2. 2a. AchatineUa ovata Newc. (= A. bulimoides ovata) 160

3. AchatineUa rufa Newc. (= Partulina rufa) 29

4. Achatinella splendida Newc. (= Partulina splen-

dida) 51

5. Achatinella redfieldii Newc. (= Partulina redfieldii), 38

6. Achatinella nivosa Newc. (= A. abbreviata) 125

7. Achatinella melanostoma Newc. (= A. pulcherrima) 142

8. Achatinella cestus Newc 286

9. 9a. Achatinella swiftii Newc 294

10. 10&. Achatinella turgida Newc 306

11. Achatinella cylindrica Newc. (= Amastra cylin-

drica.) Vol. XXI 227

12. Achatinella casta Newc 235

13. Achatinella intermedia Newc. (= Amastra cylin-

drica color-var.) Vol. XXI 222

14. Achatinella violacea Newc. (== Amastra violacea.)

Vol. XXI 257

15. Achatinella sanguinea Newc. (= Laminella san-

guinea.) Vol. XXI 330

16. Achatinella porphyrea Newc. (= Amastra cylin-

drica var.) Vol. XXI 224

17. Achatinella gigantea Newc. (= Amastra magna.)

Vol. XXI 239

18. Achatinella subvirens Newc. (=A. viridans var.). 129

19. Achatinella ampla Newc. (= A. fulgens ampla) .... 198

20. Achatinella adamsi Newc. (= Partulina marmorata) 43

21. Achatinella rutila Newc. (= A. viridans var.) 128

22. 22&. Achatinella rugosa Newc. (= A. byronii rugosa) 135

23. Achatinella multilineata Newc. (== A. mustelina

High.) 343

24. 24a. Achatinella fulgens Newc 190

25. Achatinella glabra Newc. (= A. bulimoides glabra) 164

PLATE 30.

(Reproduced photographically from Pfr., P. Z. S., 1855, pi.
30, f. 1-25, and from Newcomb, P. Z. S., 1853, pi. 23, f. 27-52.)

1. Achatinella aptycha Pfr 54, 145

2. Achatinella pulchella Ufr. (==A. lorata pulchella). 284

3. Achatinella amoena Pfr. (= Auriculella amoena.)

Vol. XXIII

EXPLANATION OF PLATES. 409

FIGURE PAGE

4. Achatinella Candida Pfr. (= A. bulimoides-ovata) . 162

5. Achatinella cinerosa Pfr. (== A. valida cinerosa) .. 336

6. Achatinella macrostoma Pfr. (—viridans) 128

7. Achatinella fricki Pfr. (= A. b. ovata?) 162

la, b. Achatinella fricki Pfr. (= A. b. ovata Nc.) .... 162

8. Achatinella planospira Pfr. (— A. decipiens?) .... 148

9. Achatinella monacha Pfr. (= = A. mustelina) 343

10. Achatinella vidua Pfr. (== A. b. ovata) 163

11. Achatinella multicolor Pfr. (= A. pulcherrima Sw.) 143
lla. Achatinella multicolor Pfr. (= A. sowerbyana ovi-

formis Nc.) 177

12. Achatinella attenuata Pfr. (=Partulina terebra var.) 63

13. Achatinella swainsoni Pfr. (= decipiens var. ?) .... 150

14. 14a. Achatinella sowerbyana Pfr 175

15. Achatinella dolium Pfr 316

16. Achatinella forbesiana Pfr. (= A. cestus var.) . . . 289

17. Achatinella rudis Pfr. (= Amastra spirizona rudis)

Vol. XXI 219

18. Achatinella fusiformis Pfr. (= Amastra mucronata.)

Vol. XXI 268

19. Achatinella napus Pfr. (— A. mustelina sordida) . . 349

20. Achatinella ventrosa Pfr. (= A. lorata var.) 280

23. Achatinella crassidentata Pfr. (==A. fulgens) . 193, 199

24. Achatinella valida Pfr 334

25. Achatinella globosa Pfr. (== A. vittata) 291

27. Achatinella sordida Newc. (= A. mustelina var.) . . 349

45. Achatinella recta Ne we. (= A. livida recta) 248

48. Achatinella venulata Newc. (= A. stewartii) 211

48«. Achatinella venulata Newc. (= A. s. producta) . . 211

49. Achatinella mucronata Newc. (= Amastra mucro-

nata.) Vol. XXI 268

50. Achatinella johnsoni Newc. (== A. stewartii) 211

51. Achatinella aplustre Newc. (=A. stewartii) 212

52. Achatinella hybrid a Newc. (= A. s. producta) .... 212

PLATE 31. Achatinella (Bulimella.)

1, la. A. abbreviata Rve. Palolo, Gulick. 92572 123

2, 2a. A. abbreviata Rve. Palolo, Thaanum. 107002.. 123

3, 3a. A. abbreviata Rve. Palolo- Waialae ridge, Thaa-

num 123

4-4d A. abbreviata Rve. Western ravine of Palolo, Pils-

bry. 108184 123

5, 5a. A. abbreviata Rve. Palolo, Gulick. 92575 123

6, 6a. A. viridans Migh. Nuuanu, near Pali, R. A. Cooke. 127

410 EXPLANATION OF PLATES.

*
FIGURE PAGE

7-76. A. byronii capax P. & C. Waimano-Manana ridge.

108133 137

8, 8a. A. byronii rugosa, var. Waimano-Manana ridge.

108064 138

9-9d. A. byronii waimaiioensis P. & C. Waimaiio-Ma-

nana ridge. 108136 137

10. A. byronii nigricans P. & C. Waimano-Manana

ridge. 108134-5-77 138

13. A. pulcherrima Sw. E. spurs of Kawaihalone, Spald-

ing. 108130 140

14. A. pulclierrima Sw. Low in Helemano, Spalding.

108131 140

15-15cL A. lila Pils. Waimano-Manana ridge. 108066.. 139

PLATE 32. Achatinella (Bulimella.)

1-16. A. decipiens kaliuwaaensis P. & C. 107997 150

2, 2a. A. decipiens Nc. Specimens from Dr. Newcomb. 146

3-5. A. decipiens Nc. Kahana, Gulick. 92458-9-64 146

6-6c. A. decipiens (corrugata). Hakipuu, Gulick.

92451 147

7-76. A. corrugata Gul. = decipiens var. Kahana, Gu-
lick. 92452 147

8. A. d. torrida Gul. Waikane, Gulick. 92455 148

9. A. d. torrida Gul. Kaaawa, Gulick. 92263 148

10-10d. A. d. torrida Gul. Kahana. Gulick. 93403 148

11-116. A. d. torrida Gul. Waiolu, Gulick. 92454 148

12-12c. A. d. torrida Gul. Kaaawa-Hakipuu ridge, Spald-
ing. 107996 148

13. A. pulclierrima nympha Gul. Wahiawa (color much

too deep yellow) . 92622 144

14. A. pulclierrima nympha. Helemano, Gulick. 92206. 144

15. A. elegans Nc. Kahuku, Gulick. 92256 166

PLATE 33. Achatinella (Bulimella.)

1-lj. A. bulimoides Swains. Kawailoa, Gulick. 92489. 154

2. A. b. ovata Nc. Kahana, Gulick. 92400 160

3-3d. A. bulimoides. Kaipapau, Spalding. 108117 .... 156

4. A. bulimoides. Hauula, Gulick. 92445 156

5-5c. A. b. mistura P. & C. Kaliuwaa, Spalding. 108115. 156

6, 60, A. b. mistura P. & C. Kaliuwaa, Spalding. 108116. 156

7. A. b. mistura. Punaluu ridge, Spalding. 108114 . . 156

8-8c. A. b. glabra Nc. Waimea, Gulick. 92220 164

9. A. b. glabra Nc. Specimen from Newcomb 164

10, 10a. A. b. glabra. Kawailoa, Gulick. 92486 164

11. A. b. glabra. Waialee, Gulick. 92651 164

EXPLANATION OP PLATES. 411

FIGURE PAGE

12. A. elegans inelegans P. & C. 92504 168

13. A. b. spadicea Gul. Type, Boston Soc. N. H. 157

13a. A. b. spadicea Gul. One of original lot, A. N. S. . . 157

PLATE 34. Achatinella (Bulimella.)

1, la. A. rosea Sw. Wahiawa, Gulick. 92618 151

16. A. rosea Sw. Wahiawa, Cooke. 1289 Cooke coll. . . 151

2, 2a. A. rosea Sw. Poamoho, Spalding. 107991-2 .... 151

3, 3a. A. rosea Sw. Kawaihalona, Spalding. 108118 A.

N. S. and 2167 Spalding coll 151

4, A. rosea Sw. West of Helemano, Spalding. 434

Spalding coll 151

5, A. rosea Sw. Opaeula, 1,700 ft., Wilder. 108121 . . 151
5a. A. rosea Sw. Opaeula, 1,700 ft., Wilder. Wilder

collection 151

6-6c. A. rosea Sw. Poamoho, Spalding. 107991 151

7. A. rosea Sw. Kaukinehua, Spalding. 108119 151

8-86. A. rosea Sw. Wahiawa, W. D. Wilder. 108120.. 151

9. A. sowerbyana Pfr. Kaipapau, Spalding. 108122 . . 175

10. A. sowerbyana Pfr. Kaliuwaa, Spalding. 108123 . . . 175

11. A. s. var. oviformis Pfr. Kaliuwaa, Spalding. 108125. 177

12. A. s. roseoplica P. & C. Opaeula, Spalding. 107990. 180
13-136. A. s. thurstoni P. & C. Kahuku, Thurston.

108126 177

14-146. A. s. thurstoni var. Waimea-Laie ridge, Spald-
ing. 108124 178

15, 15a. A. s. laiensis P. & C. Laie, Spalding. 108127... 178

PLATE 35. Achatinella (Bulimella.)

1. A. fuscobasis Smith. Type, 110 Boston Soc. N. H. . . 170

2. A. fuscobasis Smith. Kuliouou, Thaanum. 108559. 170

3. 4. A. fuscobasis Smith. Mt. Olympus, Spalding.

108560 170

5, 7. A. f. lyonsiaiia Bald. Cotypes, Konahuaniu. 65693. 172

6. A. f . lyonsiana Bald. Konahuanui, Spalding. 108561. 172
8-13. A. sowerbyana dextroversa P.&. C. Pupukea, Thaa-
num. 108128 179

14-17. A. pupukanioe P. & C. Waimaiio-Manana ridge,

Pilsbry. 108068 174

PLATE 36. Achatinella.

l-l/. A. fulgens Nc. Palolo, Gulick. 92576 195

2, A. fulgens Nc. Waialae, Gulick. 92296 194

3, 3a. A. fulgens Nc. Wailupe, Gulick. 92241 194

4, 4a. A. fulgens Nc. Niu, Cooke, no. 21, 22 193

412 EXPLANATION OF PLATES.

FIGURE PAGE

5-5c. A. fulgens Nc. Niu, Gulick. Bost. Soc., 604 193

6-6e. A. fulgens Nc. Niu, Thaanum 193

7-le. A. buddi Newc. Palolo, Gulick. 92581-2 187

8, Sa. A. buddi Newc. Makiki. 108208 187

9, 9a. A. phaeozona Gul. Kailua, Gulick. 589 Bost. Soc. 186
10, 100. A. phaeozona Gul. Keawaawa, Gulick. 92266. 185

PLATE 37. Achatinella.

1, A. fulgens Nc. Waialae nui, Cooke coll. 125 194

2, 2a. A. fulgens Nc. Waialae nui, Thaanum. 108211. 194
3-3/1. A. fulgens Nc. Waialae iki, Thaanum. 108209 . . 194

4-4c. A. fulgens Nc. Waialae. 66539 194

5-5c. A. fulgens Nc. Waialae, Gulick. 92309 194

6-66. A. fulgens Nc. Waialae, Gulick. 92290 194

7, la. A. fulgens Nc. Palolo. 237 Cooke ; 108210 195

8-8c. A. fulgens Nc. Palolo, Gulick. 92308 195

8-8c. A. fulgens Nc. Palolo, Gulick. 92308 195

9. A. fulgens Nc. Palolo, Gulick. 92310 195

10-116. A. vulpina tricolor Sm. Heia, Gulick. 92466-8. 226
12, 120. A. fulgens Nc. (Augusta pattern.) Wailupe,

Gulick. 92239 194

PLATE 38. Achatinella.

1-1&. A. stewartii Green. W. branch Palolo, H. A. P.

108190 204

2-2&. A. stewartii Green. Manoa-Palolo ridge, Thaa-
num. 108215 204

3, A. stewartii Green. Manoa-Palolo ridge, Wilder.

108216 204

4, 4a. A. stewartii Green. S. side Manoa, 326303 Cooke

coll 204

5, 6, 60. A. stewartii Green. Manoa, Gulick. 92562-3. 204
7-7c, 8, 80. A. s. producta Rv. E. rim Tantalus bowl,

P. & C 207

9. A. s. producta. Head of Makiki, H. A. P. 108551 . . 207

10. A. s. producta (bilineata). Tantalus. 385 Cooke coll. 207
11-13. A. s. producta. Tantalus. 452, 458, 399 Cooke

coll 207

14. A. stewartii. Makiki, Gulick. 92552 207

15. A. fuscozona, type specimen (=buddii). 75 Bost.

Soc. coll 189

16. 160. A. stewartii Gr. Makiki, Gulick. 804 Bost. Soc. 207

17. A. stewartii Gr. Tantalus. 355 Cooke coll 207

18. 180. A. stewartii. Makiki, Thaanum. 108552 207

19-19c. A. stewartii. Makiki, Gulick. 92546 207

EXPLANATION OF PLATES. 413

FIGURE PAGE

20. A. stewartii. Tantalus. 353 Cooke coll 207

21. A. stewartii. Pauoa, Gulick. 92611 207

PLATE 39. Achatinella vulpina, varieties.

1, la. A. vulpina, typical pattern. Nuuanu. 362-3

Cooke coll 212, 216

2, A. vulpina (castanea pattern). S. side Nuuanu, Gu-

lick. 92365 217

3, 3a. A. vulpina, ernestina patterns. N. E. Nuuanu,

Cooke. 108553 217

4, A. vulpina (cotype of ernestina). Nuuanu, Baldwin.

65706 217

5, 6. A. vulpina ("ernestina"). Nuuanu. 552-8 Cooke

coll 217

7-7c. A. vulpina, from one small bush. Nuuanu. 108554. 218

S-Se. A. vulpina. Nuuanu, R. A. Cooke. 108555 218

9-9c. A. vulpina. Nuuanu, R. A. Cooke. 108556 219

10, 10«. A. vulpina. N. side Nuuanu, C. M. Cooke, no.

643, 645 219

11-llc. A. vulpina, olivacea pattern. Glen Ada, H. A. P.

108082 218

12-12d. O. vulpina, olivacea, virens and longispira pat-
terns. Hillebrand 's Glen, R. A. Cooke 219

13-13/. A." vulpina. Nuuanu-Kalihi ridge. 108558 219

14, 14a. A. vulpina. Nuuanu, Gulick. 92358 219

PLATE 40. Achatinella vulpina, varieties.

1, la. A. vulpina, adusta pattern. Pauoa, Gulick .... 217

2. A. vulpina, type figure of A. adusta. After Reeve.. . 217

3. A. vulpina, type figure of A. olivacea. After Reeve. 220

4, 4a. A. vulpina, olivacea pattern. Nuuanu, Cooke coll. 218
5-5c. A. vulpina, olivacea pattern. Waolani Peak.

108027 220

6, 6a. A. vulpina, form cucumis Gk. Kalihi. 92501 . . 221
7-7c. A. vulpina, varieties. Kahauiki, Spalding. 108848. 221
8, 9. A. vulpina, varieties. Kahauiki, Spalding. 108849,

51 221

10-10d A. vulpina, varieties. Kahauiki, Spalding.

108850 221

. A. vulpina, varieties. Kahauiki, Spalding. Bishop

Mus 222

le. A. vulpina, varieties. Moanalua, Cooke coll 222

12-12&. A. vulpina, var. Moanalua, Spalding 222

12&. A. vulpina, var. Moanalua, Cooke coll 222

13. A. vulpina, var. Halawa, Spalding 223

414 EXPLANATION OP PLATES.

FIGURE PAGE

14, 14a. A. vulpina, var. Middle fork Halawa, Thaamim. 223
15-15d A. vulpina, analoga pattern. Halawa, Gulick.

92370 223

16. A. vulpina, var. Type fig. of diluta Smith. After

Smith 228

PLATE 41.

1, la. A. fuscobasis wilderi. Lanihuli, Wilder 173

2. A. fuscobasis wilderi. Lanihuli, Koolau side, Spald-

ing 173

3, 3a. A. fuscobasis wilderi. Wilder 173

4. Partulina dubia. Type of A. morbida Pfr., B. M. . . 117
5-5(7. A. vulpina, longispira patterns. Halawa, Gulick,

875, 896 Bost. Soc. ; 92386 223

6. A. vulpina, virens pattern. Aeia 223

7. A. vulpina var. Waimano 223

8, 8a. A. vulpina colorata Rv. Newcomb 224

8&-8e. A. vulpina colorata. Ahuimanu, Gulick 224

9, 9a. A. vulpina colorata. A. consanguinea Sin. Ahui-

manu, Gulick 225

10-10d. A. bellula Sm. Upper Pauoa, Thwing 230

11. A. lehuiensis Sm. Type, no. 74 Boston Soc 271

12. A. casta Nc. Type of concolor Smith, Boston Soc. . . 239

13-13c. A. casta Nc. Waimano, Cooke coll 235

14-145. A. casta Nc. "A. cookei ridge.' Thaanum.

108764 235

15-15&. A. casta Nc. Waiawa, Spalding coll. no. 1863 . . 235
16. A. casta Nc. Waiawa, Spalding. 108565 235

PLATE 42. Achatinella.

1-3. A. spaldingi P. & C. Cotypes 271

4. A. lehuiensis gulickiana P. & C. Type, Boston Soc.. . 273

5, 6. A. thaanumi P. & C. Type, Thaanum coll 273

7. A. papyracea Gul. Ahonui, Gulick. 1021 Bost. Soc. 243

8. A. papyracea Gul. Kalaikoa, Gulick. 978 Bost. Soc. 243

9. 10. A. casta margaretag P. & C. 108570-1 240

11, 12. A. juncea Gul. Types. Wahiawa, Gulick. 49

Boston Soc 241

13. A. juncea Gul. Wahiawa, Spalding. 107987 241

14. A. juncea Gul. Wahiawa. Cooke coll., 1304 241

15. 16. A. dimorpha Gul. Waialee, Gulick. 92654 . . . . 258
17-20. A. dimorpha, zonata patterns. Kaaawa, Gulick.

92265 258

EXPLANATION OF PLATES. 415

FIGURE PAGE

PLATE 43. Achatinella.

f

1-lcZ. A. phaeozona Gul. Keawaawa, Spalding 184

2. A. fulgens var. Waialae iki, Spalding coll. no. 670. 194
3-3c. A. fulgens varieties. N. W. Wailupe. 108842 ... 194
4-4c. A. fulgens varieties. N. ravines Palolo, Spalding.

108847 195

5, 5a. A. fulgens versipellis Gk. Types, 44a, &, Boston

Soc 196

5&, 5c. A. fulgens versipellis Gk. Boston Soc., no. 601 . . 196
6-6e. A. fulgens versipellis. Kailua-Waimanalu ridge,

Spalding 196

1-7 a. A. fulgens versipellis. Maunawili, Wilder 197

l~b-ld. A. fulgens versipellis. Kailua, under Olympus,

Spalding 197

8. A. fulgens form fuscolineata Sm. Copy of type figure 197

9. A. solitaria Nc. Copy of type figure 204

10-106. A. s. producta Ey. Tantalus-Pauoa. Bishop Mus. 209

11. A. caesia cognata. Type of A. scitula Gk. Boston

Soc 268

12. A. cassia cervina. Type of A. cervina Gk. Boston Soc. 267

13. A. caesia. Type of A. formosa Gk. Boston Soc 265

14. A. caesia. Type of A. concidens Gk. Boston Soc 265

PLATE 44. Achatinella.

1. A. livida herbacea Gk. Type. Boston Soc 251

2-4. A. livida herbacea Gk. 92491 251

5. A. livida. Type of A. glauca Gk. Boston Soc 250

6, 7. A. livida, glauca pattern. 92284 250

8. A. caesia Gk. Type of var. cognata Gk. Boston Soc. 267

9-12. A. caesia cognata Gk. Hakipuu, Gulick. . . 92224. 267

13. A. caesia Gk. Type. Boston Soc., no. 53 263

14-16. A. caesia Gk. Waimea, Gulick. 1258 Boston Soc. 263
17-20. A. cassia littoralis P. & C. Cotypes 266

PLATE 45. Achatinella curia.

1. A. curta Newc. Copy of type figure 252

2, 3. A. curta Newc., rhodoraphe pattern. Ahonui Gu-

lick 253

4-9. A. curta var. Wahiawa, Cooke coll 254

10, 11. A. curta var. Wahiawa, Thaanum 253

12, 13. A. curta. Left side Poamoho, Thaanum 254

14-16. A. curta. Waialua, Baldwin. Cooke coll 254

17-22. A. curta, form delta Gk. Wahiawa, Gulick. 92619. 256

416 EXPLANATION OP PLATES.

FIGURE PAGE

PLATE 46. Achatinella curta.

1, 2. A. curta, rhodoraphe pattern. Helemano, Gulick. 254
3, 4. A. curta, rhodoraphe pattern. Gulch W. of Hele-
mano 254

5. A. curta. Laie, Spalding. 108751 256

6-8. A. curta, delta pattern. Helemano, Gulick 256

9. A. curta. E. of Opaeula, Spalding. 108576 254

10. A. curta. Kawaiholona, Spalding. 108579 255

11-15. A. curta. Kawaiiki, Spalding. 108580, 108988 . . 255

16-19. A. curta. Kawailoa, Thaanum. 108581 255

20-24. A. curta, undulata pattern. Kawailoa, Gulick.

92278, 92286 255

PLATE 47. Achatinella.

1. A. dimorpha Gk. Type, no. 56, Boston Soc 260

2. A. dimorpha. Pupukea, Gulick. 92389 260

3-5. A. dimorpha. Paumalu, Spalding. 108755 260

6-8. A. dimorpha Thaanum. 108756 260

9-12. A. dimorpha. Paumalu, Spalding. 108757 261

13. A. dimorpha. Kaunala ridge, Spalding. 108758. . 260

14. A. dimorpha, albescens pattern. Waialee, Gulick.

92653 261

15. A. dimorpha, sonata pattern., Kahuku, Gulick. 92655. 261

16. A. dimorpha, albescens pattern. Kahuku, Thaanum.

108759 261

17. A. dimorpha, sonata pattern. Kipapau, Spalding.

108761 261,

18. A. dimorpha, sonata pattern. Kaaawa, Gulick. 92265. 261

19. A. dimorpha, contracta pattern. Copy original fig. 263

20. A. dimorpha, contracta pattern. Kaaawa, Gulick.

92262 262

PLATE 48. Achatinella.

1-2. A. bellula Sm. Pauoa, Cooke coll 230

3. 4. A. bellula Sm. Head of Nuuanu, Cooke coll 230

5, 6. A. b. multizonata. Cotypes. 65703 231

7-12. A. b. multizonata. Nuuanu, Cooke coll 231

13-16. A. bellula. Waolani. 108562, 108013 231

17, 18. A. bellula. Kalihi, Cooke coll 231

19, 20. A. phseozona. Olomana, Gulick coll. Bost. Soc. 186

21. A. fulgens, crassidentata pattern. Wailupe, Spalding.

108844 194

EXPLANATION OF PLATES. 417

FIGURE PAGE

PLATE 49. Achatinella.

r

1. A. casta Nc., received from Newcomb 23G

2-5. A. casta. Waimano, Cooke Coll 237

6-8. A. casta. Waiawa, Spalding 23S

9-15. A. casta. Waiawa, Thaanum 238

16. A. casta. Type of A. pygmaa, no. 79 Boston Soc. . . 239

17, 18. A. juddii Bald. Cotypes 244

19, 20. A. juddii. Kalauao-Waimalu ridge, Spalding . . 244

21, 22. A. juddii. Aiea, Cooke Coll 244

PLATE 50.

1-5. A. lorata nobilis. Waolani. 108006, .8 283

6. A. lorata. Waimano, Thaanum. 108768 282

7. 8. A. lorata, Moanalua, Cooke coll 282

9. A. 1. pulchella. Waiawa, Spalding. 107982 284

10-14. A. 1. pulchella. Waiawa, Cooke coll 284

15. A. apexfulva. Copy of Dixon's figure 317

16. 16&. A. lorata, melanogama pattern. Embryos, 108767 283

17. Partulina dubia. Type of A. platystyla Gk. Boston

Soc 116

18. Partulina dubia. Type of A. pexa Gk. Boston Soc.. . 116

PLATE 51. Achatinella lorata.

1-3. A. lorata, melanogama colony. Maunawili, Wilder.

108767 283

4-7. A. lorata, Tantalus, Cooke Coll., 406, 408, 431 280

8. A. lorata. Tantalus, S.-E. rim. 108040 280

9-12. A. lorata. Nuuanu, Cooke Coll 281

13-16. A. lorata. Nuuanu, Cooke, 108777 281

17, 18. A. lorata. Kalihi, Gulick. 92502 282

19-21. A. lorata. Nuuanu, Gulick. 92359 281

PLATE 52. Achatinella.

1, la. A. lorata. Tantalus, Cooke Coll. no. 411 280

Ife. A. lorata. Tantalus, Spalding 280

2. A. lorata. Makiki, above falls. Spalding 280

3-3c. A. lorata. Makiki, Gulick, 82544 280

4. A. lorata. N. side Nuuanu, 655 Cooke Coll 281

5, 50. A. lorata. Kalihi, Gulick, 92271 282

6. A. lorata. Moanalua, 868 Cooke Coll 282

7. A. lorata. Kalauao, Spalding 282

8, Sa. A. lorata, melanogama pattern. Maunawili, Wilder 283

9, 9a. A. cestus, forbesiana pattern. Wailupe, Thaanum 288
10. A. cestus, forbesiana pattern. Wailupe-Niu, Spalding 288

418 EXPLANATION OF PLATES.

FIGURE PAGE

11-116. A. cestus. Waialae, Gulick, 92304 288

12, 12«. A. cestus. Specimens from Newcomb 286

13, 13a. A. cestus. Wailupe, Gulick. 92238 288

14, 140. A. cestus. Palolo, Gulick. 92579 287

15-15e. A. valida Pfr. Pupukea, Gulick. 92387 334

16. A. concavospira Pfr. Type in British Mus 351

17, Yla. A. v. kahukuensis P. & C. Cotypes 338

PLATE 53. Achatinella livida.

1-9. A. livida Sw. Kawailoa, Gulick, 92276 246

10. A. livida, emersoni pattern. After Newcomb 247

11. A. livida, emersoni pattern. Wahiawa, Cooke Coll. . 247
13-15. A. livida, recta Nc. Waialua, Newcomb 248

16. A. livida, recta Nc. Wahiawa, Cooke coll 250

17-21. A. livida, recta Nc. Kawailoa, Gulick 92277 248

PLATE 54. Partulina, Achatinella.

1. la. Partulina helena Nc. W. of Meyer's lake. 108584 356

2. P. helena near Puu Kolekole. 108657 356

3. P. helena near Puu Kolekole. 108656 356

4. P. saccata Pfr., type in British Mus 364

5. 5a. P. physa konana P. & C. Cotypes 365

6-66. P. thwingi P. & C. Cotypes 357

7-7c. Achatinella fulgens ampla Nc. Newcomb coll. . . 198

8. A. papyracea Gk. Type, no. 48 Boston Soc 243

9-96. A. papyracea Gk. Waipio, Thaanum. 108765 . . 243

10. A. papyracea Gk. Wahiawa, Spalding. 108572 . . . 243

11. Partulina horneri Bald., Embryo 365

12. P. horneri Candida P. & C., Embryo 365

13. P. horneri fuscospira P. & C., Embryo 365

14. 14a. Achatinella papyracea Gk. Wahiawa, Gulick.

1109 Boston Soc 244

PLATE 55. Achatinella.

1. A. cestus Nc. Palolo, Gulick, 92579 287

2-4. A. cestus, simulator pattern. Palolo, Gulick, 92592 287

5. A. apexfulva beata P. & C. Helemano-Poamoho,

Spalding, 108808 329

6, 7. A. a. duplocincta P. & C. Kawailoa, Gulick.

Bost. Soc 323

8. A. a. duplocincta. Wahiawa, Cooke Coll. no. 1272 . . 323
9-16. A. valida cinerosa Pfr. (leucozona Gk.). Waialee,

Gulick. 92650, 56 336

17, 18. A. v. cinerosa, var. Waimea (?), Gulick. 92221 338
19-22. A. v. leucophoea Gk. Waialee, Gulick. 92655 ... 336

EXPLANATION OF PLATES. 419

FIGURE PAGE

PLATE 56. Achatinella turgida. ,

1-lc. A. turgida Nc. Waiawa, Spalding. 107983 295

2. A. turgida Nc. Waiawa 295

3. 3a. A. turgida ovum. Waimalu (?) Gulick. 92235 298

4. A. turgida Nc. Waiawa, Spalding 295

5-5/. A. t. perplexa. Waimano, Cooke Coll 296

6. A. t. ovum. Aiea. 991 Cooke Coll 298

7-ld. A. t. ovum. Moanalua. Cooke Coll 298

8, 8a. A. t. ovum. E. of Kalauao, Spalding. 108769. 298

9. A. t. ovum. Waimano- Waiau, Spalding. 108771.. 298

10. A. t. ovum. Halawa, Spalding. 108770 298

11. A. t. perplexa. Waimano, Cooke Coll. no. 1176 297

12. A. t. ovum. Waimalu, Gulick. 92233 298

13-136. A. t. simulacrum. W. of Manana, Wilder. 108772. 299

14-14d. A. t. simulacrum. Waimano-Manana. 108063. 299

PLATE 57. Achatinella.

1. A. vittata Eve. Copy of type figure 289

2-2c. A. vittata Rve. Nuuanu, Gulick. 92360 290

3-36. A. vittata Rve. Nuuanu, 108800 290

4, 40. A. vittata Rve. Nuuanu, 631 Cooke Coll 290

5-56. A. vittata Rve. Nuuanu, Cooke Coll 290

6-66. A. vittata cinerea. Nuuanu, 108801 291

7. A. vittata cinerea. Nuuanu, 620 Cooke Coll 291

8. A. v. simulans Rve. Copy of type figure 292

9. 9a. A. v. simulans. Nuuanu, Cooke Coll 292

10-11. A. v. simulans. Waolani Peak. 108026, 107979. 292

12. A. v. simulans. Kapalama, Spalding. 107980 .... 293

13. A. v. simulans. Copy of type figure of A. albofasciata 293

14a-14c. A. v. simulans. Nuuanu, Cooke Coll 293

15. A. turgida cookei Bald. Cotype. 65692 300

16-166. A. t. cookei. Thaanum 300

17. A. t. cookei. Spalding 300

PLATE 58. Achatinella, swiftii.

1. A. swiftii Nc. Specimen from Newcomb 306, 308

2-2c. A. swiftii. Wahiawa, Gulick. 92633 306, 308

3, 3a. A. swiftii, tuberans pattern. Wahiawa, Cooke Coll. 314

4, 4a. A. swiftii Nc. Kipapa, Spalding. 107985 314

5-56. A. swiftii Nc. Kipapa- Wahiawa ridge, Spalding.

108783 314

6. A. swiftii. Type of A. tuberans Gulick. Bost. Soc.

no. 93 313

7. A. swiftii Nc. Kalaikoa, Gulick. 92327 309

420 EXPLANATION OF PLATES.

FIGURE PAGE

8. A. swiftii, coniformis pattern. Kalaikoa, Gulick.

92325 309, 313

9-96. A. swiftii, tuberans patterns. Kalaikoa, Gulick.

92329 314

10, 10a. A. swiftii, polymorpha patterns. Kalaikoa,

Gulick. 92343 315

11, Ha. A. swiftii, tuberans patterns. Kalaikoa, Gulick.

92328 314

12, 12a. A. swiftii. Cotypes of A. flavida Gul. Boston

Soc. no. 95 311

126. A. swiftii, flavida pattern. Kalaikoa, Gulick. 92337 311

13-136. A. swiftii Nc. Kalaikoa, Gulick. 92335 309

14. A. swiftii. Type of A. versicolor Gul. Boston Soc.

no. 101 310

14<z-14c£, 15. A. swiftii, versicolor patterns. Ahonui,

Gulick. 92428, 92408 311

16. A. swiftii var. Ahonui, Gulick, 92409 312

PLATE 59. Achatinella.

1-1&. A. swiftii Nc. Ahonui, Gulick. 92429 306

2, 2a. A. swiftii, coniformis patterns. Ahonui, Gulick.

92410 312

3. A. swiftii. Type of A. albospira Sm. Boston Soc.

no. 96 309

4-4e. A. swiftii, polymorpha patterns. Waipio, Gulick.

92261 315

5-56. A. swiftii chromatacme P. & C. Waiawa, Thaa-

num. 108804 316

6. A. swiftii. Copy of type figure of A. neglectus Sm. 310

7. A. swiftii. Copy of type figure of A. innotabiLis Sm. 309

8. A. leucorraphe Gul. Type. Boston Soc. no. 92 .... 301

9. A. leucorraphe irwini P. & C. Spalding, 108780 ... 304
10. A. leucorraphe irwini P. & C. Kaukinehua, Spald-
ing. 108779 304

11-116. A. leucorraphe irwini, types. 108778 303

12. A leucorraphe irwini. Kipapa-Waikakalaua. 107977 304

13. A. leucorraphe irwini. 107976 305

14-146. A. leucorraphe irwini. 107978 304

15, 15a. A. leucorraphe irwini. Waipio, Thaanum .... 305
16-166. A. leucorraphe irwini. "Rosea ridge,' Thaanum 305

17. A. t. ovum Pfr. Figure of type in British Mus 297

PLATE 60. Achatinella apexfulva.

1-16. A. apexfulva, Opaeula, Emerson. 109776 318

Ic. A. apexfulva, Opaeula, Emerson collection 318

EXPLANATION OF PLATES. 421

FIGURE p

2. A. a. vespertina Bald. Type. 65699 ............ 322

3. A. a. vespertina. 10312 .................. ' ....... 322

4-4c. A. a. apicata Nc. Wahiawa, Gulick. 92628 ____ 321

5-5&. A. a. apicata. Wahiawa, Cooke Coll ............ 324

6-6a. A. a. apicata. Wahiawa, Spalding ............. 324

1-11}. A. a. apicata. Kaukinehua, Spalding. 108805,

107984 .................................... 324

8, Sa. A. a., cervixnivea pattern. 108807 ............. 329

9. A. a., cervixnivea pattern. Poamoho, Spalding.

108806 .................................... 328

10. 10a, 13. A. a., gulickii pattern. Kalaikoa, Gulick.

92322 ..................................... 327

11. A. a., gulickii pattern. Ahonui, Gulick. 92405 .... 327

12. A. a., type of A. gulickii Sm. Boston Soc. coll ..... 327

14, 14&. A. a., lilacea pattern. Kalaikoa, Gulick. 92330 328

15, 15a, 16. A. a. aloha P. & C. Kaukinehua, Spalding 330
17-17c. A. a. beata P. & C. 108809 .................. 329

PLATE 61. Achatinella.

l-2e. A. decora Fer. Kawailoa, Gulick. 92483, 5 ____ 331

3. A. decora. Wahiawa, Cooke Coll. 1212 ........... 331

4, 4a. A. decora. Gulch W. of Helemano, Spalding.

107973 .................................... 331

5-5a. A. decora. Kawaiholona, Spalding. 107974 . . . 331

6. A. decora. E. of Opaeula, Spalding. 07975 ...... 331

7, la. A. mustelina High. Lihue, Gulick. 92513 ____ 348
8-Se. A. m. lynianiana Bald. Palihua iki, Cooke. 107983 350
9-9&. A. m. lymaniana. Lihue, under Palikea, Spalding 351

10-10&. A. concavospira griseizona P. & C. Cotypes.

108799 ..................................... 353

11. A. m. lymaniana. 1801 Cooke Coll ............... 350

12-12d. A. concavospira Pfr. Palihua, Cooke. 108798 . . 351

PLATE 62. Achatinella.

1, 2. A. mustelina High. Mokuleia, Gulick. 92251... 342

3, 4. A. m. makahaensis P. & C. Makaha. 108788 . . . 345

5-8. A. m. sordida Nc. Mokuleia, Gulick. 92248 ____ 349

9-11. A. m. sordida, from Dr. Newcomb .............. 349

12-20. A. decora (tumefacta Gul.) Wahiawa, Gulick.

92635 ..................................... 334

21. A. concavospira turbiniformis. Popouwela-Lihue

ridge, Spalding. 107949 .................... 353

22, 23. A. c. turbiniformis Gul. Type, Boston Soc ..... 353
24, 25. A. concavospira Pfr. Palihua, Wilder. 108797 351

422 EXPLANATION OF PLATES.

FIGURE PAGE

PLATE 63. Achatinella.

1. A. inustelina High. Makaha, Spalding. 108789 ... 345

2. 2a. A. mustelina Migh. From Dr. Newcomb 344

3. A. mustelina. W. Mokuleia, Spalding. 108786 345

4-4&. A. mustelina. Mokuleia, N. W. head of Makua,

Spalding. 108784 345

5-5&. A. mustelina. Mokuleia, Gulick. 92249 345

6, 6a. A. mustelina. Haleauau 346

7. A. mustelina. Leilehua-Waialua ridge, Spalding.

107972 346

8-10c. A. mustelina. Popouwela, H. A. P 346

11. A. mustelina. Popouwela, S. ridges, Spalding.

108794 346

12, 12a. A. mustelina. Popouwela-Lihue ridge, Spald-

ing. 107998 347

13-13cL A. mustelina, bicolor pattern. Lihue, Gulick.

92512, 14 348

INDEX.

Note.— Names of genera and other groups are printed in
SMALL CAPITALS; of synonyms in italic.

baileyana Gul 52

BALDWINIA Anc. ... 90, 364

beata P. & C. 329

bella Rve 79

bellula Sm. 230

bensonia Paetel 368

bicolor Gul 348

bilineata Eve. 208, 210

buddii Newc 187

BULIMELLA Pfr 118

bulimoides Swains 154

Bulimus clausinus Migh. 366
Bulimus insignis Migh. . 26
Bulimus kanaiensis Pfr. 367
Bulimus lactifluus Pfr.. 367
Bulimus liratus Pfr. ... 3
Bulimus newcombianus

Pfr 13

byronensis Beck 134

byronii Gul 205

byronii Wood 133

abbreviata Reeve 123

Achatinella Schluter 366

ACHATINELLA Swains. 117, 274

ACHATINELLASTRUM Pfr. 180

acicula Schluter 366

adamsi Newc 43

adusta Rve 213, 227

agatha Paetel 368, 369

alba Nutt 279

alba Sykes 324

albescens Gul 258, 262

albofasciata Sm 292

albospira Sm 306, 309

aloha P. & C 330

AMASTRA Ads 383 '

amoena Pfr. = Auriculella.

ampla Newc 198

ampulla Gul 57

anacardiensis Paetel . . . 389

analoga Gul 213, 228

anceyana Bald 75, 390

angusta Paetel 191

apexfulva Dixon 317

Apex Marts 118, 274

apicata Newc 324

aplustre Newc 205, 212

aptycha Lyons 325

aptycha Pfr. ... 54, 145, 363

attenuata Pfr 63, 364

augusta Sm. . . 191, 192, 203

B

bacca Reeve , 123

C

cassia Gul 263

canaliculata Bald 6

Candida P. & C 365

Candida Pfr 161, 162

capax P. & C 137

carinella Bald 7, 358

carnicolor Bald 58, 364

castanea Rve 212

casta Newc. 235

cervina Gul... . 264, 267, 269
cervixnivea P. & C. 329

(423)

424

INDEX.

cestus Newc 286

chromatacme P. & C. ... 316
cinnamomea Fricke .... 389
cinnamomea Pf r. . . . 10, 355

cinerea Sykes 291

cinerosa Pfr 338

cingulata Fricke 369

circulata Frick 369

clausiana Hartm 368

clausinus Migh 366

Clementina Pfr 123

Cochlogena Fer 117

cognata Gul 264, 269

colorata Gul 213

colorata Rve 224

compressa Pfr 369

compta Pease 36

concavospira Pfr 351

concidens Gul 264, 269

concolor Sm 236, 239

concomitans Hy 37

confusa Sykes 96, 105

coniformis Gul. . . . 306, 312
consanguinea Sm. . 224, 225

contracta Gul 262, 263

cookei Bald 300

cooperi Bald 72

corrugata Gul. ...:.... 147

corusca Gul 62

costata Borch 4

crassa Newc 40, 362

crassidentata Pfr. 191,192,199

crocea Gul 65

cucumis Gul 213, 227

cumingi Newc 9

cuneus Pfr 238, 210

curta Newc 252

D

decora Fer 289, 331

decora Rve 292

decorata Pils 12

decipiens Newc 145

delta Gul 256

densilineata Rve 49

dextroversa P. & C. 179

diluta Smith 228

dimorplia Gul 258

diversa Gul. . . 191, 192, 201

dixoni Borch 82

dolei Bald 60, 364

dolium Pfr 316

dubia Newc 113, 393

dubiosa Ad 55

duplocincta P. & C 323

dunkeri Cum 210

dwightii Newc 35, 360

E

ebuniea Gul 57, 364

EBURNELLA Pse 67, 385

emersoni Newc 247

emmersoni Newc 247

ernestina Bald. . . . 213, 229
errans Pils Ill

F

fdba Pfr 367

fasciata Gul 56

flavidus Gul 306, 311

flemingi Bald 71

folliculus Schliit 366

forbesiana Pfr 286

forbsiana Hartm 286

formosa Gul 264, 269

fricki Pfr 165

fucosa Frick 114

fulgens Newc 190

fulgurans Sykes 21

fulva Newc 85

fulvicans Bald 73

f uscobasis Smith 170

fuscolineata Sm 197

fuscospira P. & C 385

fuscozona Smith 189

fuscozonata P. & C 365

fusoidea Newc 64

G

gemma Pfr 3

germana Newc 76

gldbia Hartm 164

INDEX.

425

glabra Newc 164

glauca Gul 250

glolosa Pfr 289, 291

gouldi Pfr 49

gouldii Newc 52, 363

gravis Fer 369

grisea Borch 37

grisea Newc Ill

gulickiana P. & C 273

gulickii Sm 325, 327

H

halawaensis Borch. .. 27, 359

hanleyana Pfr 278, 281

havaiana Rv 368

hawaiana Gul 369

haivaiiensis Bald 109

hayseldeni Bald 88, 388

helena Newc. ... 16, 356, 394

Helicter Pse 117, 274

Helicterella Gul 274

Helicteres Fer 117

hepatica Borch 83

herbacea Gul 251

honomuniensis Pils 12

horneri Bald. 92, 107, 365, 392
hybrida Newc 212

Idas Borch 31

ignominiosus Pse 369

impressa Pse 369

induta Gul 48

inelegansP. &C 168

innotabilis Sm 306, 309

insignis Migh 26

irwini P. & C. 302

johnsoni Newc 205, 211

juddii Bald 244

juncea Gul 241

K

kaaeana Bald 41, 362

kahukuensis P. & C. 338

kaliuwaaensis P. & C. . . 150

kaluaahacola P. & C. ... 359

kamaloensis P. & C 362

KAUAIA Sykes 398, 399

kauaiensis Pfr 367

konana P. & C 365

kuhnsi Pils 22

lactea Gul 86, 364

laiensis P. & C 178

LAMINELLA 383

latizona Borch 82

lehuiensis Sm 271

lemmoni Bald 61

LEPTACHATINA 382

leucophsea Gul 336

leucorraphe Gul 301

leucozonalis Beck 366

leucozonus Gul 337

ligata Sm 236, 238

lignaria Gul 63

lila Pils 139

lilaceum Hartm 191

lUaceus Gk 325, 327

liliacea Pfr. . . 191, 192, 199

limbata Gul 134

littoralis P. & C 266

liratus (Bulimus) 3

livida Swains 246

longior Pils 63

longispira Sm 213, 228

lorata Fer 278

lorata Reeve 141

liibrica Schliit 366

lubricoides Schliit 366

lugubris Chem 318

lutea, Gul 367

luteostoma Bald 170

lymaniana Bald 350

lyonsiana Bald 172

M

macrodon Borch 35

macrostoma Pfr 126

mahogani Gul 141, 143

426

INDEX.

maniensis Pfr 20

margaretse P. & C 240

marmorata Old 42

martensi Borch 78

mauiensis Pfr 20

melanostoma Newc 142

meyeri Borch 29

mighelsiana Pfr 77

minuscula Thwing 23

minuscula Pfr 18

mistura P. & C 156

morbida Pfr 117

moiitagui Pils 66

mucida Bald 34

multicolor Pfr. . 141, 143, 177

nmltistrigata 34

inultizonata Bald 231

nmtabilis Bald 68

myrrhea Gul 52

N

napus Pfr 349

nattii B. & H 73

nealii Hartm 74

neglectus Sm 306, 310

newcombia Hartm 13

NEWCOMBIA Pfr. . 1, 353, 395
Newcombiana Thwing . . 1

newcombianus Pfr 13

nigricans P. & C 138

nivea Bald 59, 363

nivosa Newc 123

nympha Gul 144

O

obliqua Gul 158

occidentals P. & C 361

octavula Pfr 369

olesonii Bald 369

olivacea Rve 213

ornata Newc 18, 356

ovata Newc 160

oviformis Pfr 177

ovum Pfr. 297

pallida Nutt 279

Partula virgulata 2j

PARTULINA Pfr 14

PARTULINELLA Hyatt ... 392

papyracea Gul 243

PERDICELLA Pse 15

perdix Reeve 45, 363

perfecta Pils 54, 363

perforata Gul 62

perkinsi Sykes 8

perplexa P. & C 293

perversa Swains 331

pexa Gulick 116

pfeifferi Nc 13, 355

phccostoma Anc 107

phaeozona Gul 184, 39 S

philippiana Pfr 8, 358

physa Newc. ... 95, 105, 109

pica Swains 318, 321

planospira Pfr 148

platystyla Gul 116

plicata Migh 2

plumata Gul. . . 191, 192, 200

plumbea Gul 43

polita Nc 80, 389

polymorpha Gul. . . . 306, 315
porcellana Newc. . . . 69, 390

prasinus Rve 213, 227

procera Anc 107

producta Rve 207

proxima Pse 32, 360

pygnuea Sm 236, 239

pyramidalis Gul 46

pyramidata Pils 363

pyrimidalis Lyons 363

pulchella Pfr 284

pulcherrima Rve 205

pulcherrima Swains. . . . 140
pupukanioe P. & C 174

Q

quernea Frick 332

INDEX.

427

R

radiata Old 49, 363

radiata Pfr 126

recta Newc 248

redfieldii Newc 38, 362

reevei Adams 247

rkodoraphe Sm 252

ridua Pfr 367

rohri Pfr 26

rosea Swains 151

roseoplica P. & C 180

rotunda Gul 163

rubiginosa Newc 130

rufa Newc 29, 360

rugosa Newc 135

rutila Newc 128

S

saccata Pfr 364

sandwichensis Pfr 367

scamnata Fer 368

schauinslandi Borch. ... 33
scitula Gul. . . . 264, 268, 269
semicarinata Nc. ... 86, 388

seminiger Mke 318

seminigra Lam 318, 321

semitecta Fricke 369

simulator P. & C 288

simulacrum P. & C 299

simulans Rve 292

sinistra Fer 369

solida Gul 51

solitaria Newc 204

sordida Newc 349

sowerbyana Pfr 175

spadicea Gul 157

spaldingi P. & C 271

Spiraxis sandwichensis

Pfr 367

splendida Newc 51, 363

stewartii Green 204

subovata Fer 369

subpolita H. & P. ... 24, 359

subvirens Newc 129

sulcata Pfr. 5

sulphuratus Beck 366

swainsoni Pfr. .. 150

swiftii Newc 306

T

tasniolata Pfr 130

talpina Gul 52

tappaniana Ad 54, 367

terebra Newc 61, 364

tessellata Newc 28, 360

thaanumi P. & C 273

thaauumiana Pils 112

theodorei Bald 33, 360

theodorii Bald 34

thurstoni P. & C 177

thwingi P. & C 357

torquata Paetel 369

torrida Gul 148

tricolor Smith 226

trilineata Gul. 191, 192, 202

tuba Gul 367

tuberans Gul 306, 313

turbinata Nutt 367

turbiniformis Gul 353

turgida Newc 294

U

ualapuensis Pils 12

undosa Gul 46

undulata Newc 252

unilamellata Schliit. . . . 366

ustulata Gul 47

ustulata Newc. 224

valida Pfr 334

varia Gul 191, 192, 201

variabilis Newc 83, 390

ventrosa Pfr 279, 280

venulata Newc 205, 211

versicolor Gul 306, 310

versipellis Gul 196

versipilis Pse 196

vespertina Bald 322

vestita High 342, 344

428

INDEX.

vidua Pfr 163

virens Gul 213

virgulata High 25, 358

viridans Migh 125

viridans Rve 246

vittata Eve 289

vulpina Fer 212

W

wailauensis Pils

wailuaensis Sykes . . .
waimanoensis P. & C.

7

72
137

wakefieldiae Cox =

Tornatellina

wilder! Pils 173

winniei Bald 44, 363

wheatleyana P. & C. ... 168
wheatleyi Newc. . . . 161, 169

Z

zebra Newc 19

ze'brina Borcli 21

zebrina Pfr 20

zonata Gul 258, 262

DATES OF ISSUE OP THE PARTS OF VOL. XXII.

Part 85, pp. 1-64, plates 1-12, November 21, 1912.
Part 86, pp. 65-112, plates 13-26, October 9, 1913.
Part 87, pp. 113-176, plates 27-39, December 10, 1913.
Part 88, pp. 177-428, plates 40-63, May, 1914.
Preface and Introduction, pp. i-lviii, May, 1914.

Achatinellidae

11

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Plate 40

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Achatinellidae

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Achatinellidae

Plate 60

13

16

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7a

10

14

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17

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8

11

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Plate 6 1

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PLATE 62

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Plate 63

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12 a

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