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

BERICEIEY\ 

LIBRARY 

UNIVER5 V OF I 
CAUFOi.NllA ) 

EARTH 

SCIENCES 

LIBRA 







SECOND SERIES: PULMONATA 



MANUAL 



CONCHOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



FOUNDED BY 



GEORGE W. TRYON, JR. 



CONTINUKD r.V 



HENRY A. PILSBRY, Sc.D., 

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



65 Colored. 



PHILADELPHIA: 

PUBLISHED BY THE CONCHOLOGICAL DEPARTMENT 

ACADEMY OF NATURAL SCIENCES 

OF PHILADELPHIA. 



SECOND SEKIES: PULMONATA. 



MANUAL 



OF 



CONCHOLOGY 



STRUCTURAL AND SYSTEMATIC 



WITH ILLUSTRATIONS OF THE SPECIES 



VOL. XXII 
ACHATINELLIDJE 

BY 

HENRY A. PILSBRY 

ASSISTED BY 

C. MONTAGUE COOKE 

GENEALOGY AND MIGRATIONS OF THE ACHATINELLIDuE 

BY 

ALPHSU:?-. HYATT 



PHILADELPHIA: 
Published by the Conehologieal Department 

ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA 
1912-1914 



EARTH 
SCIENCES 









x u 





PUBLICATION COMMITTEE 

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

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

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



CONTENTS. 



PREFACE v 

INTRODUCTION 

Achatinellida 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 

Kemarks on the synonymy of Achatinellidce 1 

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

SYSTEMATIC PART 

Genus NEWCOMBIA PfeifTer 1 

Genus PARTULINA Pfeiffer 14 

Section Perdicella Pease 16 

Section Partulina s. str 23 

Section JEburnella 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) 



/"049 



PREFACE. 



This volume treats of the tree snails of the Hawaiian Islands 
belonging to the family Achatinellidse ; those of the family Amas- 
tridse (the genus Laminella) having been dealt with in Volume 
XXI, and arboreal forms of Tornatdlinidce (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, ' ' 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 
eystematist this is perhaps the more important work of the vol- 



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 Achatinella 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 H} r att'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 Auriculella, Tornatellina 
and their allies, and contain a general index 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 PauaH 
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 ACHATINELLIDJE Tryon. 

Achatinettida Tryon, 1884 (exclusive of Tornatellinidce and 

Amastridce) . 

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 cseca. 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 Achatinellidce and 
Amastridce 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 AckatinelUda in essential structure stand nearest to 
the Amastrida, Enidce and Ferussacidce, and are somewhat 

(ix) 



X TEETH OP 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 
Achatinellida are most nearly akin to the Amastrida, 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 Achatinellidce 
differ strikingly in dentition. Those families have teeth of 
normal shape, with cusps resembling and homologous with 
these of the Helicida, Lymnaida, and most other Pulmonate 
snails. The teeth of Achatinellidce 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 Drymceus, 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^E. XI 

the cusps, it is not necessarily of inueh 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 Tornatellinida? 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 Amastridtz and Enida, 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 Enida, 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 Achatinellid^ 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 ACHATINELLIIXE. 

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

Achatinella phaeozona Gul. Kailua (p. 187). 

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

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

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

Newcombia philippiana Pfr. Molokai (p. 356). 

Partulina confusa Sykes. Hawaii (p. 105). 



CLASSIFICATION. Xlll 

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 ACHATINELLHXE. 

It has been stated above that the genera and subordinate 
groups of Achatinellida 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 



Partu 
Partulina s. str. 



Perdicella 

inella | 

Eburnella Achatinellastrum 



Bulimella 



(Partulina') (Achatinella) Achatinella s. str. 

i 

Ancestral 
Achatinellid 



XIV CLASSIFICATION. 

Key to Genera and Sections of Achatinellida. 

a. Columella straight or only weakly folded. 

b. Shell slender, turrite, usually sculptured; 3% em- 
bryonic whorls, having coarse spirals, the last obli- 
quely striped. NEWCOMBIA, p. 1. 
6 1 . 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. Baldwinia, p. 90. 
a 1 . Columella strongly folded. 

b. 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. 

c 1 . Shell larger, spirally striate, columellar fold 
stout. Partulina, s. str., p. 23. 

c 2 . Shell polished, often banded, columellar fold 
stout. Eburnella, p. 67. 

b 1 . 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. Butimetta, p. 118. 

c 1 . Outer lip but little or not thickened within, 
not expanded; summit conic, the embryonic 
whorls not conspicuously flattened. 

Achatinellastrum, p. 180. 

c a . 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 Achatinellidcz. 

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. 

(Maui) 

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

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

P. marmorata Old. 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. p. fulvicans Bald. 73. 

P. porcellana Newc. 69. P. nattii Bald. & Hartm. 73. 

P. p. flemingi Bald. 71. P. anceyana Bald. 75. 

P. p. wailuaensis Sykes 72. P. germana Newc. 76. 



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



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



(Lanai) 

P. semicarinata Newc. 86. 
P. s. hayseldeni Bald. 88. 

(Molokai) 

P. m. polita Newc. 80. 



Genus ACHATINELLA Swainson. 



Section Bulimella 

A. abbreviata Eve. 123. 

A. viridans Migh. 125. 

A. taeniolata 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. 



Pfeiffer (Oahu). 

A. bulimoides spadicea P.&C. 

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. phaeozona 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. caesia 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. str. 
(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. chromatacme 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. leueophaea 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. 

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

DISTRIBUTION OF GENERA, SUBGENERA AND SPECIES. 



ISLANDS 


1 NEWCOMBIA 


PARTULINA 


ACHATINELLA 


Totals by islands 


o 


Partulina 


Baldwinia 


cS 

s 
^ 


Bulimella 


s 

kj 

1 
O 


Achatinella 


Niihau 












. . . i . . 


Kauai 






1 
8 

14 


' 1 
2 


1 

2 

6 


13 


' 17 


11 " 


43 
15 
3 
31 


Oahu 




1 


JVLolokai 


5 


Lanai . 


. . . 


. . . 


Maui . - . - 


1 


8 


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 ACHATINELLHLE. 

The primitive color pattern of arboreal Achatinellidce may 
have consisted of dark streaks in the direction of growth- 
lines, on a light gmind. 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 defined, 
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 Achatinellidcc 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. 4&-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 b 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. XXI 

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 Achatinella. 
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. 4&, 
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- 
taeniate" system may be expressed by a formula, such as is 
used for the five-banded or so-called pentatseniate Helices. 
Thus pi. 38, fig. 6 has the formula 1234. Fig. 7, 0000. Fig. 
7c, 0230. Fig. 18a, 1(234). Fig. 19c, 1(23)0. In Acha- 
tinellastrum a particularly common formula is 1000. As in the 
pentataeniate 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 Achatinellidcz. Thus on plate 19, it will be seen that 
fig. 10 has the normal, tetrataeniate 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 Partulina, 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 Achatinellastrum 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. ccesia (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 Achatinellidce the evolution of color-pattern 



COLOR PATTERNS. XX111 

has been orthogenetic. Partulina and Achatinellce 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 AchatinellicUz. 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 Achatinella. 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 no 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 OP THE SHELL. 

The shape of the shell has been varied much less in Acha- 
tinellidce 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 
stride are often rather well developed ; and in Newcombia they 
become prominent. 

The axis of the shell is somewhat sinuous, and in the last 
whorl 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 Amastridce 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 Achatinellidcz, 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 Amastridcz, 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 phczozona, 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. Belated 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. XXV11 

The species of the mustelina-decora group, and those of the 
apexfulva-leucorraphe group (see p. 275 and diagram on 
p. 278) are certainly very closely related while A. lor at a 
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 Achatinellidce. 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 Ud ; 13 to 136 ; 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. 



XX Vlll 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 66). In Palolo, still westward, the 
green and yellow patterns are ascendant, and the plumata 
marking is hardly 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 ovsal oval ov v 

(a, adusta; v, vulpina', c, castanea; e, ernestina; o, olivacea-, 
v, virens; s, suturalis; 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. XXIX 

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 no 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 L 
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 Achatinellida 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, 

! The Evolution Theory, translation by J. A. Thomson and Margaret R. 
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 conchologists 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 and 
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 Pupillidce and 
Endodontida?. 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 110 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. 

Achatinellida 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> K. C. L. Perkins. 

The Achatinellae 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. XXX Vll 

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. Newcoinb 
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 on 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 Achatinellidce and Endodontida 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, wall-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 northwestern 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 l i 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, Tornatellinidce and Leptachatina are usually 
rather abundant. Pupttlidce 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 Helicinidce, Zonitidce, Endo- 
donts, Amastras, Pupillidce 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, Achatinellida, Amastridcc and Auri- 
culella 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 Achatinellidc? 
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 Amastridce, 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 field. Morer 
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 would 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 OP 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 Achatinellidae and 
Amastridae are special to each island, though in" ar f@.w~ 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 
Amastr u umbilical a 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 now 
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 common 
to at least six of the islands in the Pleistocene, y One species, 
L. subcylindracea, was common to Oahu, Molokai and 
Kahoolawe, and the forms of Maui and Hawaii are barely 
distingnishable. 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 Achatinellidcz (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. xliii 

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 
Amastridce and other families is on a par with the faunas of 
the other islands. The absence of Achatinellidce 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 Achatinellidce 
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 I 
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. 

Amastrida could have been ancestral to the Achatinellida . 
The points of agreement between Kauaia and Achatinella 
phaozona, noted by Hyatt (pp. 398-9), pertain to shell char- 
acters of little or no phylogenetie 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 Achatinellidce 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 Achatinellidce reached the region later than Amas- 
tridce, and after Kauai had been isolated from the remainder. 
This seems the simplest explanation. (2) Achatinellidce 
may have existed on Kauai, but becom/ 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 
employment 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 Amastrida (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. xlv 

fully support this conclusion. There are five species of 
Achatinella, one Partulina and two Lamiiiellas. 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 
AchatineUida, 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 Amastridce. In the 
diagram no. 2, iirVol. 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" (Occ. 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, 
Endodontidcz 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, Lymnceida and Melaniida 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, Philomycidce., Helicidce, Opeas, Ccecilioides,. 
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, 
*>y 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 Achatinellidce 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 Achatinellas 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 
Lanai the wild goats have almost, and in Kahoolawe have 
totally, completed the destruction of native forests. With 



1 SYNONYMY OF ACHATINELLUXE. 

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 known, 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, Zonitidce, etc. found congenial environment, if we 
may judge by the abundance of individuals. 

NOTES ON THE SYNONYMY OP ACHATINELLID^E. 

The specific synonymy of Achatinellida 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. E. 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. 11 

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 OF ACHATINELLDXE. 

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 
lively 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 were 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 Amastridc? present no 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^J. 1111 

on a lei or necklace, which seems 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 Achatinellida. 

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, lorata, turrit ella f 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 OP ACHATINELLID^E. 

shells Swainson in 1828 erected the genus Achatinella, with 
the following species: A. pica, perversa, livida, bulimoides, 
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 
Achatinellidce 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 lay 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 Achatinellse 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. 



v HISTORY OP ACHATINELLID^. 

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

John T. Gulick began collecting Achatinellida 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 Achatinellidae. 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 Achatinellida. 

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



HISTORY OP ACHATINELLIME. Ivii 

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

III. 

Several papers 011 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, AchaUnellen-Fauna der Sandwich-Insel Mol- 



Iviii HISTORY OF ACHATINELLID^E. 

okai, Zoologica, vol. 19, 1906, and Monographic der auf der 
Sandwichinsel Kauai lebenden Molluskengattung Carelia, 
Abh. Senck. Naturforsch. 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 Achatinellidffi 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, cumingi, plicata, newcombiana, cere- 
alis, elongata. MARTENS, Die Heliceen, 1860, p. 249 (type A. 
plicata High.) 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 3J 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 Partulinas, 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 
AmastrincB. 

N. carinella is the only dextral species known. 

Pfeiffer's list under Newcombia was heterogeneous, bat 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. 



2, NEWCOMBIA. 

Key to Species of Newcombia. 
((. Parietal wall making an angle with the columella, the latter 

having a low fold or twist. 

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

N. philippiana, no. 5. 
ft 1 . 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 l . Spirals rounded or subobsolete. 

N. p. gemma, no. la. 
c\ Dark colored; length 12 to 18 mm. 

d. Spirals acute. N. canaliculata, no. 2. 

d l . Spirals obtuse, base darker. N. sulcata, no. 3. 
b*. Surface with fine beaded spirals, a small carina at peri- 
phery; zigzag-striped, dark-colored, dextral. Maui. 

N. carinella, no. 4. 
a 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. N. pfeijferi, no. 9. 
ft 1 . Longitudinal folds weak or wanting. 

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

N. cumingi, no. 7. 

c 1 . Smoother; spiral threads weak or effaced, the minute 
wavy striolation well developed. Molokai. 

N. cinnamomea, no. 8. 

c*. Wrinkled or puckered over spiral cords; no minute 
striolation. Molokai. A T . c. ualapuensis, no. Sa. 

Series of N. plicata. 
1. N. PLICATA ('Mighels' Pfr.). PI. 1, figs. 1, 2, 3, 6. 

Shell sinistral, turrited, rather thin; sculptured with acute, 
elevated, spiral lira? 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. Coin- 
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., PFE., 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. Neiccombia plicata Migh., BORCHERDING Zoologica, p. 94, 
pi. 9, f. 1, la.Bulimus liralus PFR., P. Z. S. ? 1851, p. 261, 
under B. newcombianus Pfr. (1853), no description; new name 
to* 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 1J 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-strise 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- 
mella weakly sinuous, its edge dilated in a triangular white 
callus above. 

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

Length 25, diam. 9 mm.; w r horls 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, obsolete!} 7 " 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). Neivcombia costata BORCHER- 
DING, Nachrbl. d. d. Malak. Gesell, 1901, p. 57; Zoologica, p. 
95, pi. 9, f. 2, 2o. 

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 lira?, of this race, which has been 
called ' c 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 lirse, 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 weaker 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, 



IsEWCOMEIA. 

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 lira? 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 u costata" type, intermediate between plicata and gemma. 

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

Shell subperforate, sinistral, oblong-turrited, rather solid 
striatulate, and encircled with close lirse, 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. Colurnella very lightly folded. 
Aperture oblique, acuminate-oval; peristome thin, the colu- 
mellar margin dilated above, reflexed, outer margin a little ex- 
panded. Length 12^, 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. Xewcombia 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 whorls, and on the last whorl it is quite 
shining dark red-brown. The same analogy which exists be- 
tween plicata and eostata is here found between canaliculata and 
sulcata" (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. 



NEWCOMBIA. 

The sculpture is most like that of N. -plicate, 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. 6J mm. 

"Animal when 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). 

Achatinellacanaliculata BALDWIN, 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 -striae. 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 
iiamed with white. The figures represent cotypes. The largest 
shell in the type lot of 12 measures length 14, diam. 7.2 mm. 
The smallest is 13.3 mm. long. 

3a. 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. PL 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. ; 5^ whorls. 

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

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

East Maui: 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 stria?; 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 15^, diam. 6 mm.; aperture 7 mm. long, 
3 wide (Pfr.). 

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 per kinsi 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. 

li 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 



.NEWCOMBIA. 9 

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. Ach-atinella 
(Newcombia) philippiana Pfr., BALDWIN, Catalogue, p. 8. 
THWING, Reprint Orig. 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" (Sykes^). 

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 3| 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., 
6 whorls. 

This species is apparently the most primitive existing Neiv- 
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. cumingi. 

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

Shell sinistral, acuminate, turrited, with strongly marked 
transversely oblique striae 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 cumingi NEWC., Ann. Lye. 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 imperf orate. 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 striae (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 white 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 
7f x 3 mm. Sandwich Is., Frick in Cuming coll. (Pfr.). 

Molokai : Mapulehu (Baldwin, Thaanum) ; Makakupaia 
(Perkins, Meyer); Honoinuni, 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, hardly 
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 KEWCOMBIA. 

comb at about the same time almost sixty years ago the color 
of the last whorl is a rich 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 
occupying half the space between periphery and suture. The 
size is greater than at other localities up to length 23^, 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-striation 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. 

8a. N. c. UALAPUENSIS n. subsp. PI. 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. 6^ mm. 

This race seems about equally related to cumingi and cinna- 
momea, 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. newcombia Pfr. , HART- 
MAN, Proc. A. N. S. Phila., 1888, p. 41. BORCHERDING, 
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 strise 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 Blatter, i, p. 114, 1854 
(for A. virgulata, tessellata, splendida, perdix, gouldi, dubia). 
VON MARTENS, Die Heliceen, 1860, p. 243, type A. virgulata 
High. GULICK, P. Z. S., 1873, p. 90, same type. SYKES, 
Fauna Hawaiiensis, p. 311, same type. 

Includes as sections, Perdicella, Eburnella and Baldwinia. 

Shell ovate-conic, usually perforate, and having distinct 
minute sculpture of spirally descending rippled stritz. 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 Achatmettina 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 stria? 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 Achatinettina 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 PERDICELLA. 15 

Sections of Partulina. 
a. Columellar fold prominent, heavily calloused. 

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

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

ft 1 . Spiral sculpture weak or obsolete; shell smooth and 
glossy, light colored, uniform or banded. 

Section Eburnella Pse., species 36-47. 
a 1 . Columellar fold wanting or weak, not calloused. 

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

Section Baldwinia Ancey, species 48-53. 
b l . 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 Partulince having protractive stripes on the last embry- 
onic whorl, later whorls striped or with a peripheral band; 
sculpture Partulinoid; columella nearly straight, or if developed 
the columella fold is moderate, convex, scarcely or not truncate; 
outer lip thin, not expanded. Type P. helena Nc. 

Perdicella differs from Partulina 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 Gulick, who in his classification 
of 1873 ranked Perdicella as a section of Partulina. 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 
PartuiinaB. 

Key to Species of Perdicella. 

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

P. helena j no. 1. 



16 PAETULINA, SECTION PERDICELLA. 

a 1 . 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 1 . Length about 15-16 mm. ; form wider. 

P. fulguranSj no. 6. 
c 1 . Columellar fold very weak. 

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

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

P. kuhnsi, no. 7. 

1. P. HELENA (Newcornb). PL 4, figs. 1 to 7. 

Shell sinistral, ovate-conical, with finely decussating striaB; 
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 (Partidina) h., 
BALDWIN, Catalogue p. 6. I'erdicella 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 PEKDICELLA. 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.; 5J 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. PI. 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 scarcely 
plicate, semioval; peristome simple, acute, the columellar mar- 
gin dilated above, reflexed. Length 10, diam. 5 mm.; aper- 
ture 4 mm. long, 2 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. Of. 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. kuhnsi. 

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 
inch " ( 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 PERDICELLA. 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 
margin ation. 

First 1^ 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 striaB; whorls 5 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" [13f 
X 6Jmm. (Newc.} 

East Maui (Newcomb). 

Achatinello. 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). PI. 4, figs. 11, 12, 13, 14. 

Shell subperforate, 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 5-J, 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. Journ. 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.; 5J whorls. 

5. P. ZEBRINA (Pfeiffer). PL 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. 8., 1855, p. 202 (1856); Mono- 
graphia, iv, 561. A. zebra Nc., BALDWIN, Catalogue, p. 7. 

"Related to A. ornata Newc., 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). PI. 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 5J, 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 fulgurans 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 PARTULISA, 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. zebrina, 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, 2J 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.; 5^ whorls. 

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

7. P. KUHNSI Pilsbry, n. sp. PL 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 la?t 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. 

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. t 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 Perdicellce, 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 PABTULINA, MOLOKAI. 

SPECIES OF MOLOKAI. 
Key to Achatinellinas of Molokai. 

NOTE. Yellow forms plain or with zigzag black stripes should 
be looked for in Laminella r Vol. XXI, p. 345. 
A. Shell elongate, dull, with distinct or strong spiral sculpture, 
the columellar fold weak or wanting. 

Newcombia, p. 1. 

A 1 . 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. 

b l . 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. 
6 2 . Spire straightly conic, last whorl not bulging, color 
various. bella, no. 42. 

a 1 . Shell sinistral, obesely ovate-conic, yellow, the embry- 
onic whorls white with a dark band. 

subpolita, no. 43. 

A 1 . 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. 

b\ 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. 

rufa, no. 10; tessellata, no. 9. 
d l . Surface glossy; striped or plain. 

proximo,, no. 11. 



PARTULINA, MOLOKAI. 25 

c . Narrower, pyramidal-conic, sinistral. 

d. Small, length 11 to 13 mm. helena, no.~h 

d l . Larger, length over 17 mm. 

e. Longitudinally boldly striped throughout; 

length 18 mm. theodorei, no. 12. 

e l . Suture bordered below by a dark band, 

others at periphery and columella; lip 

narrow, white; length 20 to 22 mm. 

mucida, no. 13. 

e*. Suture bordered by a white band or line; 
lip and columella brown; length 21 to 25 
mm. redfieldi, no. 15. 

e 8 . 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. 

dwightii, 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 Partulina?, it is remarkably aberrant in the color- 
pattern of the embryonic whorls. 

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

' * Shell ovate-conic, light fawn color, beautifully adorned with 
dark brown bands, more or less numerous; imperforate; whorls 
5, convex; incremental striae delicate. Aperture oblong; lip 
reflected, slightly inflected. Length 1 inch, diam. | 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 columella 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, Mapulehu, 
Waialua, Halawa and Pelekunu (Meyer). 

Partula virgulata MIGHELS, Proc. Boston Soc. of Nat. History, 
II, 1845, p. 20. PER., Monogr., iii, p. 454. Achatinella virgu- 
lata Migh., 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. 

8. 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 2J or 3 whorls fawn with 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 virgulala, 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, 6 
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 fawn-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 " (Neicc.). 

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. S., 1853, p. 139, pi. 23, fig. 26; 1854, p. 
311. PFR., Monogr.,iv, p. 516. SYKES. Fauna Hawaiiensis, 
p. 319. THWJNG, 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 w r ell 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 ver}^ 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 J- the total length; upper whorls tessellated 
with reddish-gray and yellowish-gray, the last whorl 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 
strias; 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 
ida 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 striaB, 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 idse, 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 pro tractive 
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 locality 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. proxima Pse. on the other." While 
this seems rather doubtful, and probably could be proved only 
by breeding experiments, rufa certainly unites characters of the 
two species. The embryo has protractive stripes as in P. 
proxima. Some very rare color-forms closely resemble proxima 
in the later stages; but it differs from that species by the dull 



PARTULINA, MOLOKAI. 31 

surface, deeply sculptured spirally; proximo, being glossy, with 
weak spirals and very slight trace of the minute, wavy, spiral - 
lineolation of the dull Partulinas. PL 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 P. 
marmorata (pi. 7, fig. 4). Sometimes this form has a sub- 
peripheral white zone (fig. 5) ; the sculpture either as in var. 
idee or without major spirals, clothed throughout with minute, 
crinkled, spirally descending stria3. 

(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. idee, except that the spiral engraving is 
close, nearly even, without granules. Last whorl 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 rvfa 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 idee 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, imperf orate, oblong-ovate, ventricose; rather 
solid; finely striated transversely, 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 protractive 
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. PI. 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 proxima 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 stria3. 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). 

Achatinella theodorii BALDWIN, Catalogue etc., 1893, p. 7, 
(name only). Achatinella 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 proxima by the sculpture and coloration, this species 
is constantly much narrower, in a considerable series seen. 
The columellar fold is also less developed. P. proxima 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. multistrigata 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). PI. 9, fig. 17. 

Shell sinistral, minutely perforate, solid, ovate-pyramidal, 
the spire straight-sided, rather 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, 1 23. SYKES, Fauna, p. 315.Partulina mucida 
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. dwightii, 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. PL 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 striae, cut across by very numerous, fine, slightly 
undulating cross-striaB. 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. THWING, Reprint Orig. 
Descr. Achat. pi. 2. f. 7. Partulina d. , BORCHERDIKG, 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 strias 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 Up. Its range of patterns is also different. 

14a. P. dwightii compta (Pease). PL 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 stria?. 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 columella 
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. 

14&. P. dwightii concomitans Hyatt, n. var. PL 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 Jforms 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 PARTULTNA, MOLOKAI. 

5, f. 9-16. Not Achatinella grisea Newcomb. ? Achatinella 
redfieldi var. g, 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 gray 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, slightly 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.; 6-J whorls. 

Length 24.5, diam. 13, aperture 11.3 mm.; 6^ 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. redfieldii 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. 
6. 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). PL 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). 

Achatinella crcwsa NEWC., P. Z. S. 1853, p. 155, pi. 24, f. 71 



PARTULINA, MAUI. 41 

. PFR., Monogr. iv, 527. SYKES, Fauna Hawaiiensis 
p. 312. THWING, Reprint Orig. Descript. Achat. pi. 2, f. 10. - 

This very distinct species is more closely related to P. divightii 
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 MAUI. 

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. marmorata. 
17. P. KAAEANA Baldwin. PL 10, figs. 5, 6, 7. 

" Shell sinistral, subperf orated, 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." (Baldwin). 

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, yellow-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 with 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 marmorala, 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 Perdicella ornata Nc., a species 
supposed long since to have become extinct," (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 J-, breadth 
iinch" (Gld.) 

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

Achati.nella marmorata GLD., Proc. Bost. Soc. N. H. ii, p. 
200, March, 1847; Expedition shells p. 35; U. S. Exploring 
Exped., Mollusca, p. 85, pL 7, figs. 94, 94a. PFR., Monogr. 
iii, p. 455. NEWCOMB, Ann. Lye. N. H. of N. Y. vi, p. 311. 
THWING, 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 whorl having wide, strongly protractive white 
and brown stripes. Surface dull, without gloss, by reason of 
the dense, microscopic, crinkled spiral strise. 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^ 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." 

11 A. 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, MAUI. 

with a white band on the periphery of the last whorl; apex sub- 
acute; spire conical; suture marginate, well impressed. Whorls 
6J, 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 16^ mm. Average weight 6 
grains; least weight 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" 
(Gulick). 

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

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

Very closely related to marmorata, 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 
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 ivinniei 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 Hawaii ensis 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. 205, pi. 7, f. 33, 1856. 

The apex (1 to 1^ whorls) is brownish white; the next one 
or 1^ whorls have broad protractive alternating stripes of snow- 
white 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, MAUI. 

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- 
llne terminalis) and other low bushes (E. Bailey). 

21a. P. perdiic pyramidalis (Gulick). PI. 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 whorls, bluish-white 
within the aperture. Apex subacute; spire regularly conical, 
sometimes decollated. Whorls 6|, 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, MAUI. 47 

Achatinella 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. 

11 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. W T ith 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 6J, 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, MAUI. 

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 ihe mamaki (S. T. Alexander). 

Achatinella ustulata 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 12f, length 
of aperture 10 mm. Average w eight 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. 

"Var. b. 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. 

11 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). PI. 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 (Gld.). 

Sandwich Islands (Gld.). 

Achatinella radiata GLD., Proc. Boston Soc. N. H., 1845, p. 
27; Otia Conch., p. 195. PFR., Monogr., iii, 454; iv, 517; vi, 
162.Bulimus gouldi PFR., Zeitschr. 1 Malak., 1846, p. 116; 
Monogr., ii, p. 74. Partula densilineata REEVE, Conch. Icon., 
vi, pi. 2, fig. 9, April, 1850. 



50 

The locality of this species is not known; radiata and gouldi 
were stated to be from the Sandwich Islands, and for densilmeata 
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 
narrow streaks or lines of brown, cream and whitish. Lip 
slightly expanded, more so below, whitish, calloused within. 
olumellar fold rather sharp but not prominent. 

Length 18, diam. 11.3, aperture 9.8 mm.; 6f 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 



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" 
(Jfa.) 

Series of P. splendida. 
25. P. SPLENDIDA (Newcomb). PI. 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 with 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. 
THWING, 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 3J whorls, the first 1 or 
If 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 25, diam. 15 aperture 12 mm. 

Length 25 , 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) . PL 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 white variety of A. splendida Newc. 

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

26. P. GOULDII (Newcomb). PL 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.- 
PFB., Monogr., iv, 517. A. myrrhea GuL, according to Pfeiffer, 
I. 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, MAUI. 53 

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 wit{i 
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: 

' ' Achatinella 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." 

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



54 PARTULINA, MAUI. 

"This species passes into A. gouldii Newc. 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. ; 6^ whorls. 

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

Shell imperforate, 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 
5J mm. 

Sandwich Islands (Frick in Cuming coll.). 

Achatinella 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). PI. 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 strise 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, MAUL 

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 fine descending striae are more distinct 
in most specimens (pi. 12, figs. 2, 3). 

28a. P. tappanianafasciata (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 
T 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. b. 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 fasdata 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). PL 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^, convex, 
margined above; the last inflated, equal to T 6 -^ 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). PL 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 concavely 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. 

" Var. c. With body whorl rounded, not angulated. This spe- 
cies is the analogue of A. tappawiana 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). 

Achatinella eburnea GULICK. Ann. Lye. N. H. of N. Y. vi, p. 
199, pi. 6, f. 28a, b. 1856. BALDWIN, Catalogue, p. 6 
(Makawao). THWING, 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 striae, and under a lens exhibiting a fine 
pattern of decussating spiral strise; 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, diam. 14 mm." (Bfcld- 
wiri) . 

East Maui: Nahiku. 

Partulina carnicolor BALDWIN, Nautilus xix, February, 1906 r 
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-stria? 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. ; 6J whorls. 

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

i( 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 tappamana 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 2 
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). PI. 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-striae and almost effaced spiral lines. Spire 
straightly 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 24 J, 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). 

Achatinelladolei'BAivwiN, 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. PL 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 below 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. Stria? decussating. 
Color light yellow, with wave-like brown-colored markings, 
nearly obsolete on the upper whorls; lip, columella and aper- 
ture white. Length --, diam. -^ inch" (Newc.'). 

West Maui (Newcomb); Honokowai (Baldwin). 



62 PARTULINA, MAUI. 

Achatindla terebra NEWC.. P. Z. S., 1853, p. 144, pi. 23, f. 
40. PFR., Monographia iv, 525; vi, 168. THWING, 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 Pfeiffer's figure of attenuata, and the 
stripes are less flexuous. Surface glossy. They 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. PL 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 min. 

The Wailuku lot consists partly of shells like those from 



PARTULINA, MAUL 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. 

33a P. t. attenuata (Pfeiffer). PL 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 5^, 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. 7J mm., 
aperture with peristome 7J- mm. long, 4 wide. Maui, Frick 



The type figure is copied, pi. 15, fig. 3. The smaller size 
and narrower form distinguish it from terebra and lignario. 
Type locality unknown. 

336. 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. 7-J- 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 -ffo 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 lOf ; 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. 

"Var. e. White. This passes into A. attenuata Pf. 

"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 undescribed 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. 

"Shell sinistral, rather solid, elongately produced; whorls 6 
to 7, scarcely rounded (excepting the last), margined above; 
suture well 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). 



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 
Maui, a dextral shell. 

Shells from Ukumehame, West Maui (pi. 15, fig. 11), sent 
by Mr. D. Thaanum, 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% 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 



PABTULINA, 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 by 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 OF 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 l /s 
whorls. 

(6) Length 24.7, diam. 12.5, length aperture 11 mm.; l l /z 
whorls. 



PARTULINA, OAHU; EBURNELLA. 67 

(c) Length 26.9, diam. 13.1, length aperture 12.1 mm. 

Oahu: Man-oa road at its junction with the upper road, 
back of Rocky Hill, cotypes (a) no. 108181 A. N. S. P.; (&) 
no. Ill coll. Irwin Spalding; and (c) no. 33581 B. P. Bishop 
Mus. Also found on Rocky Hill about ^4 niile 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. 
dwightii 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. 

Elurnella 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 Partulinae of the striped 
group of Maui. 

Eburnella was originally a heterogeneous group comprising 



68 PARTULINA, MAUI. 

the following species : A. casta Nc., curia 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 Mani 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. PI. 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. 

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 Partulinae 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-yellow with chestnut bands and lines. 

38. P. PORCELLANA (Newcomb) . PI. 19, figs. 1 to 4. 

11 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 (Newcomb) : Huelo, Makawao to Nahiku 
(Thwing) ; 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 \y 2 whorls long running from it above the suture, 
fading into white at its upper edge, the first 1^ 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. ; 5^ 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, HAUL 71 

38a. P. porcellana flemingi Baldwin. PI. 19, figs. 5 to 11. 

Shell dextral, minutely perforate, rather solid, high-come," 
glossy, with very weak traces of spiral striation or almost 
none; variously colored; white, bandless 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 PABTULINA, MAUI. 

rower contour. The system of banding (i. e., positions of the 
bands) is the same as in porcellana. 

38fc. 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 5^, 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) wailuaensis 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. 

11 A very pretty little shell of the group of A. lella 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, MAUL 73 

39. P. PORCELLANA FULViCANS Baldwin. PI. 19, figs. 15, 16. 

11 Shell dextral or sinistral, very minutely perforate!!,' 
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 in> 
pressed. Aperture oblique, oval, white within. Per is tome 
acute, slightly thickened within, columellar margin narrowly 
refiexed. Columella terminating in a strong white flexuous 
fold. Length 18J4, diam. 10 j mm." (Baldivin) . 

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 wailuaensis 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. 
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.[chatinella] 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 the 
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. ; 5>3 whorls. 

Length 18, diam. 11, aperture 9 mm. ; 5*/2 whorls. 

While nattii is very closely related to porcellana, 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 (Baldwin). Type no. 65707 
A. N. S. P. 

Achatinella anceyana Baldwin, Proc. A. N. S. Phila., 1895, 



76 PARTULINA, MAUI. 

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) . PL 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 columellae are however quite di&- 
tinct " (Newc.). 

Known to us by the original account only. I can only 
suggest its probable relationship to P. anceyana and P. ivail- 
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. 

Bather small shells with a high polish, weak or almost 
wanting spiral striation, 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 Partulinse. 
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 8^ 
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 Kaamola (Borcherding, coll. Meyer). 

Achatinella mighelsiana PFR., P. Z. S., 1847, p. 231; Mon- 
ogr. ii, 238, 1848. REEVE, 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 mighelsi- 
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 strias 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). 

(/) 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 whorls 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. PL 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. 

430. P. MIGHELSIANA BELLA (Reeve). PI. 23, figS. 2 to 12; pi. 

26, f. 13 to 15e. 

' ' 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). 

" Animal of a uniform yellowish- white, with slate colored 
tentacles " (Newcomb). 

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. Acliatinellastrum ~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 PAKTULINA, 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. 

(&) 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 broadi 
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 (Newc.) . 

Animal of a uniform yellowish- white, including tentacles; 
tentacular sheath slightly tinged with brown; mantle black 
(Newcomb). 

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. PFB., Mon- 
ogr., iv, 536. BALDWIN, Catalogue, p. 81. THWING, Keprints 
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., 5y 2 whorls, to length 15, diam. 9.2 mm. 

Newcomb '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 bella, 
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-ovate, 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 5J/2, 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). 

Achatinellastrum latizona BORCH., Zoologica, xix, Heft 48 1 , 
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 5J4, 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) . 

Achatinellastrum 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. PL 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 weaklyf 
twisted ; columellar fold white. Aperture oblique, half-ovate, 
whitish within; perlstome 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 
Newoomb, 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 Neweomb 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 Neweomb 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 
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. nattii, 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 some 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 6^2) the upper flat, 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. 

peristome 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. (P/'r.). 

Dr. Newcomb 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). PL 22, figs. 15, 16. 

11 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 marginate, slightly impressed; 
whorls 6^, 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- 
mella 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). PL 21, figs. 1 to 4, 13, 15. 

" Shell dextrorsal, solid, ovately-conical, longitudinally 

sttbstriated ; whorls 5, flattened, marginate 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. PFB., 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 orange-rufous. The acute earina 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. 
Newcomb '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 



88 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. PL 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*/2, 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; columeJla termin- 
ating in a strong, flexuous, white fold. Length 17*/2, diam. 
10 mm. 

11 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) ; Lanaihale, 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. 

1 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-Mo'lokai 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. 
a 1 . Aperture slightly more half the length; last whorl 
rather roughly striate ; length over 20 mm. 

P. confusa, no. 46. 

a 2 . 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. 

a 1 . Last embryonic whorl obliquely striped; later whorls 
very convex. P. thaanumiana, no. 50. 

3. Oahuan 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 ACHATINELLID^, BY H. W. 
HENSHAW. 

"Although much has been written upon the Achatinellida 
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 Oahu, the metropolis of the Achatinellidce 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 PABTULINA, 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. 

Partulitw 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- 
ha x ele, 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 (My r sine lassertiana) and, rarely, upon 
kawaao (Bryonia sandwichense) , the species is chiefly con- 
fined to a small berry-bearing tree called the ahakea (Bobea 
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- 



PABTULINA, 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. 95 

"It is probable enough, too, that in time the four color 
sports, if indeed two of them are not already properly to be 
ranked as 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 horneri 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 
Achatinellida appear to be, probably no very long interval, 
nor very complete isolation, is requisite to establish such var- 
iations. 

Partulina physa 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 platyphyllum 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 <cther 
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. 

' l 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 Kea'lakekua Bay, and which now appears to be 
extinct there. If, as is probable from its description, this 
shell was physa (hawaiiensis) , 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. 

11 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. 

c ' 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 (Sopkora 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 Ackatinella 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. 

" 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 
Achatinellidce 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 Achatinella, 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 foe pregnant, bo 
perpetuate its peculiarities, subject only to the modifying 
influences of such individuals as 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 mollusean 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 ^f 
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 Achatinellidce, 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 Ackati- 
nellida 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. 

1 ' Most of the Achatinellida 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. 

1 1 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 1 
vitality. Under such circumstances adult shells are known 
often to give birth to apparently healthy young. 

' ' Unlike the Amastras and the Succineas, the Achatinellidce 
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. 

11 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 BALDWINIA. 

mented in this way with the three species here mentionel, 
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. 

11 An Achatinella which Mr. D. D. Baldwin refers to Par- 
tulina physa (= 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 horneri, for instance, a greater number would 
doubtless materially reduce the average number of young 
per adult shell. ' ' 



PARTULINA, SECTION BALDWINIA. 



105 





^ 


8 


fc^. 


j 


i 


s 


j 

>> 


g 


C co * 
QJ >~> 




T3 3 


a 


,0 


^ 





^5 


I 


^ 


2 > 'i' < 




g 


0> 


a 


a 


a 




Q 


a 






6 CO 





o> 




0) 


8 


0) 


0> 






* 


525 


I 1 


(M 


CO 


* 


' 


CO 


^ 


P. physa, (hawaiiensis) . . 


247 


47 


148 


44 


4 


4 






1.SS 


P. horneri, brown banded 




















variety 


16 


2 


3 


3 


6 


2 






2.5 


P. horneri, white variety. 


33 


16 


5 


4 


5 


3 


. . 




2.3 


P. horneri, brown whorled 




















variety . ... 


10 


1 




1 


8 


8 


1 


1 


3.7 1 / 




149 


50 


59 




16 








1.5 























46. P. CONFUSA (Sykes). PL 16, figs. 1 to 12. 

* * Shell sinistral, pointed at the summit, strongly inflat3d 
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 Qf 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, p. 
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 PARTULINA, 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. hawaiiensis, 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 " haivaiiensis " 
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. pliysa, 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.; 6^2 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. ph&ostoma Ancey. ' l 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 defined 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 3ast 
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. 

" 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 Kulmihaele, 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 
Homer. 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. 

(&) 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 band. 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 occur 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 hawaiiensis 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). 

(fr) 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 6^4 

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*/2 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). 

48&. P. physa errans, n. var. PL 17, figs. 14, 15, 16. 

The shell is thin, sinistral, more slender than usual in P. 
physa (hawaiiensis) , 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 strisB 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. Golumella 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. hawaiiensis, 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. physa, some forty to sixty milss 
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 " (Newc.). 

Maui: Makawao (Newcomb, type loc.). 

Achatinella grisea NEWC., P. Z. S., 1853, p. 153, pi. 24, f. 
66. PFR., Monogr., iv, 518. A. dubia var. b PFR., Malak. 
Bl., 1854, p. 117. 

The conic spire rather slender above, and the weak cohi- 
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. 1^ 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. dwightii 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^. 

P. grisea is not closely related to any other Mauian 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. dwightii 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 Pilsbry, n. sp. PL 18, figs. 6, 7. 

The shell is sinistral, narrowly umbilicate, thin, ovate-conic, 



PARTULINA DUBIA, OAHU. 113 

surface slightly shining, the later whorls densely striatel 
spirally, striae 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 ara 
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. Oolumella 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). Cotypes 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). PI. 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 dentatecL 
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 Makaha and Makua valleys, Waianae 
range (Irwin -Spalding). In knot holes and crevices of loose 
bark of kukui 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. morUda PFR., P. Z. S., 
1859, p. 30. 

P. diibia, 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. 
Ihaanumiana 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 
6^ . 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. PI. 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 &), 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, 10a, are from two specimens of Mr. Spalding 's 
no. 2181. This color-form is one of the rarest Oahuan tre3 
snails. A suspicion has been entertained that it might be a 
hybrid between diibia and- a rare black form of Ach. turgida ; 



116 PARTULINA DUBIA, OAHU. 

but having examined all of 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> 
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^ whorls. 



ACHATINELLA. 117 

Another figure of the type of pexa is given, pi. 50, fig.^lSr 
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 dubia, 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 subperforate, 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; whorl 6%, 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 
columellar margin much dilated, broadly adnate. Length 19, 
diam. 9 mm. Sandwich Islands, Dr. Frick in Mus. Cuming. ' ' 
(Pfr.). 

Achatinella morbida PFR., Proc. Zool. Soc. 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 
Limagons, p. 56, 1821. Helicteres Fer., BECK, Index Mollus- 
oorum, 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. Laving 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 work were 
rather heterogeneous. Grulick 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 Pf r. Outer lip thickened by a strong callous rib 
within ; shape oblong- conic or ovate ; summit obtuse, rounded 
or eonvexly-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. 

Achatinellae 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 eon-~ 
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 Oahu. 

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. B. 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 grade of "subspecies". A more rigorously 
logical treatment might reduce the "good species" to nine, 
by the union of viridans and taniolata; byronii, decipiens and 
pulcherrima; 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-come, often with streaks 
or bands of dark green, and often roughly striate or corni- 
'gated; mainly dextral. A. byronii, pulcherrima, decipiens, 
lila. 

Series of A. viridans: a subdivision of the above series, for 
the exclusively dextral species of the eastern end of the range. 
A. abbreviate viridans, and t&niolata. 

Series of A. bulimoides: shell ovate, capacious, never 
marked with green; smooth, dextral or sinistral. A. rosea, 
bulimoides, elegans. 

Series of A. fuscobasis: shell ovate-conic, 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 exp' 'ess 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 

Hoanalua) . 

a. Lip acute, but slightly or not thickened within ; shell dex- 
tral, very smooth, richly colored. A. abbreviata, no. 1. 
a 1 . 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. 

c 1 . Banded or streaked with chestnut on l a white 

ground, or white. A. taniolata, no. 3. 

b 1 . Shell sinistral. 

c. Shell white with few brown bands or none, about 
16 x 10 mm. A. fuscobasis, no. 11. 

c 1 . Similar but larger and more capacious. 

A. f. lyonsiana, no. lla. 



ACHATINELLA, SECTION BULIMELLA. 121 

c 2 . 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. 

a 1 . Lip distinctly thickened within. 
&. 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. b. nigricans, no. 4&. 

d 1 . 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. 

e 1 . More corrugated, occasionally black; sur- 
face with a gummy gloss ; whole lip usu- 
ally brownish ; length 16-19 mm. 

A. b. rugosa, no. 4#. 

e 2 . Larger, broader, green-streaked; lip white 

or narrowly dark-edged; length 19-21 

mm. Var. capax. 

e s . Nearly smooth, whitish to greenish-yellow, 

plain or banded, lip white or pale lilac ; 

length 15-18 mm. Var. waimanoensis. 

d 2 . Northern (Koolauloa) slope; shell sinistral or 

dextral, the lip mainly white. 

A. decipiens, no. 7. 
c 1 . Surface smooth ; shell dextral. 

d. Northern slope ; lip white. Various varieties of 

A. decipiens, no. 7, la. 
d 1 . Interior slope. 

e. Lip bordered with blackish brown. 

A. pulcherrima, no. 6. 
c 1 . Lip white throughout. A. p. nympha,mo. 6a. 



122 ACHATINELLA, SECTION BULIMELLA. 

& 1 . Shell ovate, smooth. 

c. Larger, 'Capacious species, diam. 10 mm. or more. 
d. Shell sinistral, highly polished, usually white 
or with rose 'bands, lip bright rose; rarely 
greenish or dark, with dark brown lip. 

A. rosea, no. 8. 

d 1 . Dextral or sinistral, variously banded with) 
chestnut or suffused, lip pale or brown. 

A. bulimoides, no. 9. 

d 2 . Dextral or sinistral, conspicuously streaked 
with brownish, sometimes with white bands 
also, lip brown. A. elegans, no. 10. 

d 3 . 'Chiefly sinistral, blue-gray or purplish, some- 
times banded. A. e. ivheatleyana, no. 10&, 
c 1 . Smaller species, usually less than 10 mm. diam. 
d. Shell sinistral. 

e. Wax-yellow with a brown sutural band r 

sometimes one or two other bands, colu- 

mella pink. A. sowerbyana, no. 13. 

e 1 . Wax-yellow below, white above periphery, 

with a subperipheral brown band; colu- 

mella pink. A. s. thursto-ni, no. 13a. 

e 2 . White, usually with broad chestnut bands; 

columellar fold white or brownish. 

A. s. laiensis, no. 13Z>. 
d 1 . Shell dextral. 

e. Wax-yellow with white sutural band, colu- 

mella pink. A. s. rose&plica, no. 13cZ. 

e 1 . Ground white or pale, uniform, or banded 

or streaked with chestnut. 
A. pupukanioe, no. 12. A. s. dextroversa, no. 13c. 

SERIES OF 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 Nuuanu and Waimanalu there are about half a dozen 
great ridges and valleys where no Bulimella is known except 
A. fuscobasis wilderi on some high points of the main axis. 



ACHATINELLA ABBBEVIATA. 123 

The viridans group is somewhat intermediate between the 
byronii and the bulimoides series, Ibut most nearly related tcT 
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. 

1 i 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-ManO'a ridge (Pilsbry & Cooke). Konahuanui (Bald- 
win) ; eastern Nuuanu, perhaps washed down from Kona- 
huanui ( Spalding) . W p 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. 
-TnwiNG, 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. 

PL 31, figs. 1, la, 5, 5a, Palolo, are from Gulick shells. The 
others, figs. 2 to 4c, are from material collected recently. A 
lot from the northwestern ravine of Palolo varies from many- 
banded to black (pi. 31, figs. 4, 4&-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, 3a, 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. 4Z>) . 

4. Last whorl black or nearly so, landless, the color becom- 
ing lighter upward, dark again at the summit (pi. 31, figs. 4c, 
4<i). 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 4d 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, 4tb, c, d, not distinguishable 
from Palolo shells. Another lot agrees with fig. 1. PI. 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 1 ," 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. Clementina Pf r. has not been figured. It has been placed 
in the synonymy of abbreviate by Pfeiffer himself arid by sub- 
sequent authors. The description, which is not very good for 
abbreviata, here follows: " A. Clementina. Shell imperf orate, 
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, diam. 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. 
' ' '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). 

Oahu (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; Conc'hyl. 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, Conch. 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). Achatinella 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 MHghels' 
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 1 , is figured on pi. 31, 
fig. 6a, coll. by Mr. Richard A. Cooke. 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 Pialolo 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 
t&nidato. It might be well to recognize subvirens as an east- 
ern subspecies of viridans, rutila and macrostoma to be syn- 
onyms of it. 

Color form rutila Newcomb. PI. 25, figs. 10, 11, lla-e, 12. 

* ' Shell ovately-eonic ; whorls 6, rounded, the last margined 
above ; suture well marked ; aperture subquadrate ; lip ex- 
panded, subrefleeted, sitrongly 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 tceniolata, and it must 



ACHATINELLA VIRIDANS. 129 

be admitted that some of the yellow- ground specimens from 
colonies presumably tczniolata X subvirens have a great re- 
semblance to Pfeiffer's figure. The original description fol- 
lows. " A. macrostoma Pfr. Shell dextral, imperf orate, 
conic-ovate, rather solid, very lightly striated, glossy ; fulvous, 
variegated with some pale and brown bands. Spire conic, 
lather obtuse; suture submarginate ; whorls 5~y 2 , 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; perisitome white, 
labiate within, the right margin expanded, strongly curved 
above ; columellar margin dilated, adnate. Length 21%, diam. 
11 mm. ; 'aperture 12 mm. long, 5 wide inside. Inhabits the 
Sandwich Islands, Frick; Mus. Cuming (Pfr.). 

Color-form subvirens Newe. PI. 25, figs. 5 to 7&, 9 to 9&. 

" Shell conically ovate. Whorls 6, rounded and margined 
above ; suture distinct and lined with white. Aperture ovate, 
expanded below; lip thickened; cohimella 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. &. Brown or chestnut replacing the green color. 

' Through var. a this species approaches a variety of A. 
rutila, and through var. &, 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 1 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^ENIOLATA. 

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, bandies or with white or chestnut bands (pi. 25, 
25, figs. 9, 9<z, 96, Palolo) . 

3. A. T^ENIOLATA Pf eiffer. PL 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 semioval, 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% mm. Sandwich Is,, Mus. Cuming." (Pfr.) 

Oahu : Palolo to Keawaawa, and northward across the range 
in Maunawili. 

Achatinella taniolata 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 HJawaiiensis, p. 310. 'Achatinella rubiginosa NEW- 
COMB, P. Z. S., 1853, p. 154, pi. 24, f. 69 (Palolo). 

A. tceniolata 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, 
Waialae, Keawaawa and elsewhere, which 'could be placed, 



ACHATINELLA TJENIOLATA. 131 

with almost equal propriety in tceniolata, viridans rutila or 
viridans subvirens, or 'assorted into these several 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, t&niolat& 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, or rarely the apex is purplish-brown, perhaps 
stained from within (pi. 24, figs. 15, 16, Plalolo, 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. 

AB 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 rubiginosa Newc. (pi. 24, 
fig. 19, Palolo, and pi. 25, figs. 13, I3a, 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 tceniolata 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, rubiginosa 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. PI. 25, figs. 13 to 13c are Wailupe shells 



132 ACHATINELLA T^NIOLATA. 

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 tcenio- 
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 shellsi (t&niolata) , or streaked (rubiginosa), with 
white ground: 

From Kuliouou (Thaanum) the specimens have the pattern 
of Keawaawa form (&), 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. lO^ mm. A lot from 
the Thaanum collection, recently collected, there are (a) 
shells variously banded with chestnut on a yellow ground ; ( & ) 
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 subvirens. 

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, tceniolata and subvirens. 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-brown 
upper whorls, paler (sometimes 1 white) embryonic whorls, the 
last whorl either banded or with rubiginosa pattern cut by one 
or more white bands. PI. 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 areas 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 Gulick 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). PI. 27, figs, 1 to \e, 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 ia chestnut band, nearly three embryonic whorls 
pinkish gray; aperture white, the lip bordered with dark 
brown. Sculpture of faint spiral stride 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 nralleation. 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: Kal'aikoa, Ahonui (Gulick). 

Helix byronii WOOD, Supplement to the Index Testaceo- 
logicus, 0<r a Catalogue of Shells, British and Foreign, 1828, 
p. 22, no. 30, pi. 7, f. 3Q.Achatinella 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- 
lanostoma NEWCOMB, P. Z. S., 1853, p. 132, pi. 22, f. 7 (1854). 
PFR., Monogr., iv, 521. Achatinella limbata GULICK, Ann. 
Lye., vi, 252, pi. 8, 70&, 70& (Feb., 1858). Helicteres byron- 
ensis (Gray), BECK, Index Molluscorum, 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 defined 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 Ach&tinellida were from near 
the west end of the main range, Wahiawa to Kawailoa. The 
present species does not occur in Kawailoa 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 typo 



ACHATINELLA BYRONII. 135 

locality, pi. 27, figs. 1, la. representing typical shells. Tire 
chestnut subsutural band is invariable in Ahonui and Kalai- 
koa shells. 

In a lot of 68 from Ahonui, Gulick coll., the colors are as 
follows : 

1. Typical, green-streaked (pi. 27, figs. 1, la), 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 limbata Gul., from Ahonui and Kalaikoa is 
identical with the typical green and yellow forms of byrotwi 
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 limbata with byronii. 

4a. A. BYRONII RUGOSA Newcomb. PI. 29, figs. 22, 220 ; 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. Striae 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-twentieths, 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. PFB., 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. 

Newcomb 's original figures are reproduced, pi. 29, figs. 22, 
22a. They are like the shells collected by G-ulick in Waiawa 
valley (pi. 27, fig. 8), which is probably what Newcomb 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 
thinner, the whole lip of a brownish flesh-tint, at least typi- 
cally, and the shell has 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, 10&). All the preceding are 
from the Gulick collection, and probably all of them were 
taken at rather low elevations. 

Length 19.3, <Mam. 10,2, aperture 9 mm. ; G 1 ^ whorls. Wai- 
pio. 



ACHATINELLA BYRONII. 137 

Length 17.5, diam. 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- 
malu 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 Waianae 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 Ms Waianae catch. The shells are 
greenish-black with the first neanic whorl green or olive 
streaked on :a yellow ground, embryonic whoiis 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, 7&. 
At the head of Waiawa gulch, along the Waiahole- 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 Kipap'a 
division ridge, and probably goes farther north. There is but 
little variation in color and shape, and it is invariably dextral. 
The p-aitterns 'are shown in figs. 1-lb. 

Length 21, diam. 12.5, aperture 11 mm. ; 6 whorls. 
Length 19, diam. 10.5, aperture 9.2 mm. 

2. Subvariety waimanoensis P. & C., new. PL 31, figs. 9 to 
9d. 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-likQ 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., 5% whorls. 
PI. 31, figs. 8, 8a. 

Forms 2 and 3 were collected Feb. 17, 1913, in company 
with Messrs. Spalding and Mem-am. 

4&. A. BYRONII NIGRICANS n. subsp. PI. 31, figs. 10, 11, 12. 

The shell is sinistral, oblong-conic, rather solid. Embryonic 
3 whorls snow-white, the next whorl brown, or brown and 
green, with a light band, last whorl intensely 'black 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, tand has a brilliant gloss. The aperture is very 
oblique, ovate, white or 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 Newc. 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-wMte embryo 
is also occasionally seen in rugosa. A. b. 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. Wdlder'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 ia 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 trees, in two spots less than a mile 'apart. 

5. A. LILA Pilsbry, n. sp. PI. 31, figs. 15 to I5d. 

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. 
I5d) uniform blackish .chestnut, or (fig. 15) having a chest- 
nut peripheral band and baso-cohimellar patch on a yellow 
ground, or (figs. 150, &) 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, ~b, 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 Wadmano-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. 
Ootype 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. 15&, b, 8; fig. 15c, 5; 
fig. 15d, 10. Four of the lot have the columellar fold white. 

A. lila is rather abundant on its misty peak, 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 may be reached from the trail. On 
the north side of the peak a poll of a couple of thousand feet 
permits 'collecting only close to the summit. Named for L. L. C. 

6. A. PULCHERRIMA Swainson. PI. 27, figs. 2 (Ahonui), 5-5g, 
6 (Wahiawa) ; pi. 31, figs. 13 (Kawaihalona), 14 (Hele- 
mano) . 

"Shell ovate-oblong, subcylindacal ; 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 ventrdcose than others, and also in the number and color 
of its bands. The ground color is a deep and rich chestnut, 



ACHATINELLA PULCHEREIMA. 141 

with from one to three bands of orange, yellow, fulvous ^r 
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" (Swainson). 

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. RPR., Mono- 
graphia id, 237; iii, 460; vi, 172; Oonchyl. Cab. 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 for 1892, p. 105, pi. 1, f. 6. Achatinella lorata 
Fer., REEVE, Conch. Icon. pi. 1, f. 6, 1850. Achatinella 
melanostoma NEWC., P. Z. S. 1853, p. 132, pi. 22, f. 7. 
Achatinella multicolor PFR., in part, P. Z. S. 1855, p. 4, pi. 30, 
fig. 11 (not fig. 11&). Achatinella mahogani GUDICK, Ann. 
Lye. Nat. Hist, of N. Y. vi, p. 254, pi. 8, fig. 72. 

This species differs from byronii 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 1 
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 nympha, 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 ACHATINKLLA 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, 5a, 5&. Chestnut, with or without darker 
bands. 

Length 18.5, diam. 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 pulcherrima, the same forms occur. Those of Kalaikoa do 
not differ noticeably. In Ahonui part of the lots examine! 
are like those of "Wahiawa. " 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 pulcherrima. Length 18.5, diam. 11.5 mm. 
These lots are from the Gulick -collection, PI. 31, fig. 14 is 
from low in Helemano, coll. by Spalding. 

Judging from the description and figure of A. melanostoma 
Newc., as well -as from specimens received from him, it is a 
synonym of pulcherrima, being based upon a somewhat dif- 
ferent color form from that of Swainson, though occurring 
with the typical color- form in Wahiawa 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. Oolumella 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, Qahu " (Newcomb). 



ACHATINELLA PULCHERRIMA. 143 

According to a note from Mr. Spalding, 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 unicolored. 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. 
Peris-tome black-edged, the outer margin somewhat straight- 
ened, narrowly expanded; eo'lumellar margin dilated, sub- 
adnate. Length 17, diam. 9 mm., aperture Sy 2 mm. long, 4% 
wide. Sandwich Is., Frick in Cuming coll. (Pfr.). 

In the above description Pfeiffer obviously eonfused two 
species. His two figures, reproduced in my pi. 30, figs. 11, 
lla, belong to A. pulcherrima (fig. 11) and A. soiverbyana 
oviformis (fig. llo-). 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 
ghell, 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 4d, Ahonui). This 
form has the smooth surface of A. pulckerrima, 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 GuMck 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 mahogani from pul- 
cherrima. Gulick 's description follows. His figure is like 
pi. 27, fig. 4a, which is exactly like the type-specimen. 

" Achatinella mahogani. 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. limbata Nob v 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 Pfedffer's figure of A. planospira except that 
the base is not abruptly darker as in that figure. It differs 
from Kahana dedpiens by having light brown early whorls 
instead of white. 

6a. A. PULCHERRIMA NYMPHA Gulick. PL 32, figs. 13, 14 ; pL 
27, figs. 7, 7a, 7&. 

"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-border. Length 18.5, diam. 10 mm. 



ACHATINELLA DECIPIENS. 145 

Oahu: Ahonui, Kalaikoa, Wahiaiwa, 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 ivhite lip, -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 Gulick 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, 11 ; pi. 32, fig. 13). 
Fig. 13 is too deep in odor. The shell is much lighter, -what 
Ridgway calls barium yellow. PI. 27, fig. 5g, 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 New-comb. PL 32, figs. 2 to 6c. 

' ' Shell conically-elongate, solid ; whorls 6, slightly rounded, 
margined above. Aperture elongately-ovate ; lip subreflected ; 
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. corru- 
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 corrugata) 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- 
tion there, the races from the windward side are for the time 
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. 

Newcomb 's typical lot consisted of shells resembling pi. 32, 
figs. 2, 2&, 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 striate spirally, with a finer de- 
scending wrinkling such as is usual in Partulina, 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 18, diam, 10 mm. 

In other Kahana lots, collected by Gulick, the patterns vary 
somewhat (pi. 32, figs. 3 to 5). Olive-green or yeliowish- 
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, diam. 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. 3&). 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, 76) 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!/2> convex, the lasc 
rugose; >eolumellar fold central, white, strong; aperture truii- 
cately auriform, white within; peristome 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. PL 30, fig. 8. 

Achatinella planospira Pfr. has been placed in the syn- 
onymy of decipiens by Newcomb, who is followed by Hart- 
mam, 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- 
<7<mi-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, imperf orate, ovate-pyramidal; solid, closely 
striate and under the lens very obsoletely striate spirally, 
glossy; blackish, ornamented with several narrow white or 
buff bands. Spire conic, rather acute; suture linear, mar- 
gined. Whorls 5!/2> flat, the last a little shorter than the 
spire, obsoletely angular, rounded Nasally. 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. 9% 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. PI. 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, imperf orate, 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 17%, breadth 9% mm. ; average weight 4.5 grains " 
(Gulick). 

' l Var. &, with black spiral lines. I am in doubt concerning 
the limits of variation of this species; var. & may be distinct " 
(Gulick). 



ACHATINELLA DECIPIENS. 14.9 

Oahu: Kahana, Kaaawa and Waikane (G-ulick) ; Waiolu 
(J. S. Emerson). 

Waiolu may be considered the type locality, as here the 
typical form (pi. 32, figs. 11, 11&, 116) 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 (ipl. 32, figs. 10 to lOd) . 
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. Spalding 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 & Oooke, 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. (6) 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 
pale peripheral line. Lip only narrowly thickened within, 
white. 

Length 19.2, diam. 10, aperture 9 mm. 

Length 17, ddam. 9.7, aperture 8 mm. 

Length 15.3, diam. 8.9, aperture 7.6 mm. 

Eastern ravines of Kaliuwaa, type loc., also central and 
western ravines (Irwin Spalding). 

This new form has such a relation to decipiens as pulcher- 
rima 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. 

n. A. DECIPIENS ( ?) SWAINSONI Pfeiffer. PI. 30, fig. 13. 

" Shell sinistral, imperforate, ovate-conic, solid, nearly 
smooth, glossy ; whitish, delicately streaked with tawny. Spire 
comic, apex fulvous, rather acute; suture margined. Whorls 
51/2, 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; 



ACHATINELLA ROSEA. 151 

columellar margin thick, flexuous, actuate. Length. 20, dianv- 
11 mm. 

" ft. Greenish buff, the last whorl chestnut anteriorly ' : 

(Pfr.l. 

Sandwich Islands, Frick, in coll. H. Cuming (Pfr.). 

Achatinella swainsoni P!FR., Proc. Zool. Soc. Lond., 1855, 
p. 4, pi. 30, f. 13. 

Dr. Cooke, who examined the type, thought this a sinistral 
individual of A. pulcherrima. ' * Newcomb suggested that this 
might be only a form of A. sordida ; it appears however to b& 
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. 

Bather 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 8&. 

1 1 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 close 
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 that the marginal [subsutural] groove, which is 
scarcely perceptible in the last, is in this very distinct " 
(Swainson). 

Length 22.3, diam. 13.5 mm. ; 6% whorls. 

Length 23, diam. 13 mm. 

Length 19, diam. 12 mm. 

Oahu : Helemano and Poamoho to Kaukinehua ; formerly, 
according to Gulick, from Kawailoa to Ahonui. 

[A. bulimoides] var. ? (rosea) SWAINSON, The Quarterly 
Journal of Sci., Lit. and Art, January to June, 1828, p. 85. 
Achatinella rosed 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 sdnistral. It further differs from A. 
bulimoides by being smoother, more glossy. The color 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- 
iek. 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, Waialna, 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. iSpalding 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. ; 6% whorls. 

Length 22.7, diam. 13, aperture 10.7 mm. 

Specimens of a lot from " Waialua," Thaanum, measure: 

Length 21.7, diam. 12.3, aperture 10.5 mm. ; 6y 2 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 bands, or rose-tinted with deeper rose bands 
denning 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, 5&) . It was probably lower down 
that Gulick found the ordinary white and pink rosta from 
Opaeula in his collection. 

Similar but larger shells were taken by M!r. 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, 8a, 8&, taken by Mr. Wilder. 
One measures, length 22, dram. 13.6, aperture 11.3 mm. 

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 rosea 
territory, which has been thoroughly " shelled " in the last 
few years, but the generosity of Messrs. Irwin Spalding andi 
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 Ij, 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 is 
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, 
viii, 216. THWING, Reprint Orig. Descr., p. 92, pi. 2, f. 1 
GULICK, Evolution, Racial and Hafbitudinal, p. 41, pi. 3, f . 1-5. 

Swainson's figures agree exactly with my pi. 33, figs. 1&, 
Id, from " Kawailoa," which I take to toe 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- 
dinted ; 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 6%. 

In a lot of 48 from " Kawailoa " (pi. 33, figs. 1 to lj, Gu- 
lick coll., 92489, A. N. S. P) , the following color-forms occur : 

1. White, the upper whorls pale-pink or flesh-tinted ; peri- 
stome 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 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. Id) ; 9 spe- 
cimens. 

4. A white zone below the suture, elsewhere chestnut or 
purplish-brown, lip often brown (fig. lc/) ; 14 specimens. 

5. Purplish-brown with fleshy suture and summit (fig. It) ; 
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. le). 



156 ACHATINEbLA BULIMOIDES. 

A small lot from Waimea consists of forms figs. Id, Ig and 

it). 

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 Kawadloa 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 
main 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 alt the periph- 
ery. The spire is usually darker, 'becoming chestnut or light- 
brown at the summit. All but two of a large series in Mr. 
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 dbliqua and 
oomorpha. 

PL 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 ov at a except that the embryonic 
whorls are darker, usually dark vinaceous. This colony is 
mainly sinistral, but contains some dextral shells. 

9&. A. BULIMOIDES SPADICEA Gulick. PL 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 from 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 mistura. 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 the 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. PL 28, figs. 8 to 9a. 

" Shell sinistral, subperf orate, ovate, obliquely truncated 
at the base, solid, shining, striated ; of 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 olliqua 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-20/i. Achatinella oomorpha 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 about 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- 



The color of the darker, lower portion of the shell varies 
through several shades resembling purple-drab, vinaceous 
purple and slate-drab of Ridgway'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 brown 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. b. 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. oomorpha 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 marginate, moderately impressed; whorls 6%, 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 20%, breadth 11 Vs nim. Average 
weight 7 grains. Kahana, 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 collection. 
The form oomorpha often has exactly the coloration of typical 
obliqua, except that the lip is brown both externally and 
within. PI. 28, fig. 10& is the typical oomorpha pattern ac- 
cording to Gulick 's figured type, two-banded 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. 

9d. A. BULIMOIDES OVATA Newcomb. PL 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 " (Newcomb, May, 1853). 



ACHATINELLA BULIMOIDES. 161 

" Shell dextral or sinistral, elongately ovate; whorls (v 
slightly margined albove, rounded; suture moderately im- 
pressed. Aperture subovate, entirely margined with black; 
columella short, plicate, strong and twisted ; lip thickened andi 
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- 
iick) . 

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, 2o> 1854. 
GULICK, Evolution, Racial and Habitudinal, p. 41, pi. 3, f. 
6h-15h. Achatinella Candida PFR., P. Z. S., 1855, p. 2, pi. 30, 
f . 4 ; Monogr., iv, 519. Ackatinella 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. wheatleyi NEWCOMB, Ann. 
Lye., vi, pp. 147 (1855) and 324 (1858), not described (see 
under A. elegans wheatleyana) . 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 Id ; 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~y 2 whorls. 

Length 22.5, diam. 13.5 mm. 

Length 21.5, diam. 12 mm. 

Length 18, ddam. 11 mm. 

Banded specimens would probably prove smaller than whit* 



162 ACHATINEDLA 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-creain -color, spire as in la (fig. 1Z>). 

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 Gi/o, 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. 7a, 7Z>, repro- 
duced in my pi. 30, figs, la, lb, 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 shelly 
and the figure is not closely matched among the specimens I 
have seen. 

A. vidua Pfeiffer, 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. Newcomb, 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 Kahana 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, subimperforate, 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 reflexed, 
columellar margin dilated, subadnate. Length 18%, diam. 10 
mm., aperture 9 mm. long, 4% wide inside " (Pfr.). 

The weakness of the columellar fold, noticed by both Pf eif- 
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. 

9e. 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 marginate in the last 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., vi, 
p. 248, pi. 8, f. 67, Feb., 1858. 

As it occurs on 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 S'ubreflected, 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 glabra, NEWC., P. Z. S., 1853, p. 139, pi. 22, f. 
25; P. Z. S., 1854, p. 310.^PFR., Monogr., iv, 520. THWING, 
Orig. De&cript, p. 93, pi. 2, f. 2.B[ulimella] glabia Newc., 
HARTMANN, Proc. A. N. S. Phila., 1888, p. 29 ("Kawaiawa")- 

Neweomb'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, 
lOa) 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 ~buUmoides, 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 Gu- 
lick, Sp aiding 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-coni'C, 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-s'haped; columellar fold above, twisted, strong; 
peristome slightly expanded, obtuse, labiate and bordered 
with violaceous or black. Length 20-21, diam. 10% 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. White or buff, ornamented with two or three black- 
bro\vn bands." (Pfr.) 

Sandwich Islands, Frick in coll. Pfr. 

Achatinella fricki PFR,, Proc. Zool. Soc. Lond., 1855, p. 3, 
pi. 30, f . 7, 7a, Tb ; Malak. Bl., 1855, p. 5 ; Monographia, iv, 
521. Cf. SYKES, Fauna Hawaiiensis, Mollusea, 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 taken 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. 7&) 
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. PL 28, figs. 12 to 13cZ ; pi. 32, fig. 
15. 

" Shell conically-elongate, polished, shining, rather solid; 



ACHATINELLA ELEGANS. 167 

whorls 6, plano-convex, margined above ; suture well im- 
pressed. Aperture subovate; lip white, expanded, sub re- 
flected, 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 (Newcoanb) ; 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 bulimmdes 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 1 
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. 13o-, 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 1 the dark streaks 
are reduced or blurred. Barely 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- 
comb I note rather indistinct dusky bands above and below 
the periphery, sometimes parted by a light-brown peripheral 



168 ACHATINELLA 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. b. 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 sdnistral 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. 13c, has a pattern re- 
sembling some specimen's 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. PL 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. 
Golumellar fold weak, whitish. Length 19, diam. 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. 28, figs'. 11, 



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 cohi- 
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 prrs- 
ent, and often more conspicuous than in fig. lla. Sometimes 
(fig. lla) 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, 

11 A. ivheatleyi" 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 Pfeiffer, " 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. 
cvata] . As no characters were ever assigned to A. wheatleyi 
except by implication in the above remarks, we have no 
option but to accept Newcomb 's published statement of what 
his A. ivheatleyi was, even though he sent out specimens 1 of 
what we are now calling u'heatleyana 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 casta series of Achatinellastrum. They 
are almost entirely shells of the high ridges and peaks. 



170 ACHATINELLA FUSCOBASIS. 

11. A. FUSCOBASIS (E. A. Smith). PL 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 M'o- 
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 fuscobasis SMITH, P. Z. S., 1873, p. 77, pi. 9, f. 
15. Achatinella fuscobasis Sm., 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, Proc. A. N. S. Phila., 1888, p. 30, pi. 1, f. 4. 

A. fuscobasis 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). 

(b) Peripheral band and basal patch brown, the space be- 
tween them white or yellowish-brown. Typical color-form of 
A. fuscobasis (pi. 35, fig. 1, type-specimen). 

(c) Same as b, but a basal band in place of the patch (fig. 
3). 

(d) Same as b 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 
fuscobasis, but Baldwin also included the banded shells in his 
original account, which follows. " Shell sinistral, imper- 
f orate, solid, ovate, spire eonvexly conical, apex obtuse; sur- 
face shining, marked with fine growth-lines, under a strong 
lens seen to be decussated by close, extremely minute spiral 
striae, apical whorls smooth. Color white, with a reddish- 
yellow lip. Whorls 51/2, margined above, slightly convex. 
Suture moderately impressed. Aperture oblique, sinuately 
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 Pa-Mo valley. The locality 
" Mt. Kaala " given in the original description of fuscobasis 
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- 
'Chologists are right in regarding lyonsiana as a variety of 
fuscobasis. 

Ha. A. FUSCOBASIS LYONSIANA Baldwin. PL 35, figs. 5, 6, 7. 

" Shell sinistral, 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 Ibands, 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 white, 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. 11% 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 a'bove 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, Proc. A. N. S. P. Phila, 
1895, p. 218, pi. 10, f. 9, 10, H.-^SUTER, t. c., p. 239, pi. 11, 
f. 52 (teeth). 

A larger, more 'capacious shell than A. fuscobasis, with the 
lip and >columella white. The patterns are : 

(a) Pure white. 



ACHATINELLA FUSCOBASIS. 173 

(Z>) 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 t&niolata toy 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 Collage. 
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. 

115. A. FUSCOBASIS WILDEEI Pilsbry, n. subsp. PI. 41, figs. 
l-3a. 

The shell is shaped as in A. f. lyonsiana. 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 whit 
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 " Manna 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 trie 
deeply cut Nuuanu 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 Spaid- 
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, 3a) 
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 & Oooke, 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 sindstral. 
The resemblance is so close, however, that I would refer tha 
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. Wilder '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. Spall- 
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 tha 
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, 14a; 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.) 

Oahu : 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 Newcomb, PFR., P. Z. S., 1855, D. 
208; Monogr., iv, 540. NEWCOMB, Ann. Lye. N. H. of N. Y.,. 
vi, p. 147 (as a synonym of A. sowerbyana 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. 

Ffeiffer described var. b 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. sowerbyana 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 1 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 Pf eiffer '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- 



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 measures, length 19.2, 
diam. 10, aperture 9 mm. ; 6% whorls. 

In one lot from the Kaipapau end of the Oastle 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 Oastle trail towards Punaluu. 

Color-form oviformis ' N.' Pfr. PL 34, fig. 11; pi. 30, fig. 
lla. " Shell sinistral, imperf orate, ovate-conic, solid, striatu- 
late, white under a glossy olivaceous epidermis, one to three 
banded with brown. Spire eonvexly-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, Newoomb " (Pfr.). Type in 
Pf eiffer 's collection. 

Mr. Sykee has referred Pf eiffer '& sinistral form of A. mul- 
ticolor to oviformis, no doubt correctly. Pf eiffer '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. 



. A. SOWERBYANA THURSTONI P. & C., n. SU'bsp. PI. 34, 

figs. 13 to 146. 

The shell is invariably sinistral; ground-color wax-yellow 
below the periphery, white above it, with a band of burn/t- 
sienna below 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%. 

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. 
13&). 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. cast a. The fact is, A. s. thurstoni stands on the border- 
line between Bulimella and the cast a 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 13& 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&, 14&. 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 concreseent 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. 



A. SOWERBYANA LAIENSIS Pilsbry & Cooke, n. subsp. PL 
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 striate 
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 j banded individuals. 
Columellar fold white, or often in part brownish. 

Length 17, diam. 9.7, aperture 8.3 mm. ; 5% whorls. 

Length 16, diam. 9.5, aperture 7.4 mm. 

Oahu : Laie, 'division ridge above the Castle cut trail, Irwin 
Spalding. Cotypes in A. N. S. and Bishop Mus. Also in 
Spalding 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. casta, 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. PL 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 1 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. SOWEBBYANA ROSEOPLICA P. & C., n. s-ubsp. 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 IVCus.) 

Opaeula, above forest- fence line, type loe. ; also on the 
northeastern division ridge between Opaeula and Kawaiha- 
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. casia. 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. casta. 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 




Series of A. vulpina. 

Large, highly colored forms, usually chestnut, yellow or 
green, and variegated with streaks or bands ; inhabiting ridges 



184 ACHATINELLA PH^EOZONA. 

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. PELEOZONA Gulick. PL 24, figs. 10 to 13 ; pi. 36, figs. 
10, 100; pi. 43, figs. 1 to Id. 

11 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, Innately 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 16*4 nun. ; 
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. b. 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^OZONA. 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. phaozona as the common 
ancestor of Achatinellastrum, Bulimella and Apex (Science, 
viii, p. 395). Later he claimed for A. ph&ozona 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 phaozona are hardly separable from cer- 
tain shells of the plumata pattern of A. fulgens ; but other 
patterns of both differ widely. A. ph&ozona 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 phaozona, 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 phceozona 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. PL 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 luddii NEWC., P. Z. S. 1853, p. 155, pi. 24, f . 73. 
PPR., 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 ACHATINELLA 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 vinaeeous. 
The chief difference is one of color; in buddii the shell 
in its more primitive pattern is closely streaked with liver 
brown to purplish vinaeeous, 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 7e, 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 6%, 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 1 /2> diam. 11 mm. 

" Station: on the trunks of trees. 

* ' Habitat : Makiki, on Oahu. Two or three specimens have 
been found in Palolo. Affinities: It is intermediate between 
A. adusta Ev. 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 y 2 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, la, Ic, except that they have the sutural band. A few 
are very small, length 18, diam. 9.5 mm., with 6% 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, 
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.) 

11 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 PULGENS. 191 

Achatinella fulgens NEWC., Proc. Zool. Soc. London, 1853, 
p. 131; varieties figured on pi. 22, f. 24, 24a. PFR., 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., REEVE, Conch. Icon,, vi, pi. 4, f. 290, &, 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, 42&; 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. phaozona, 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. 4o, 
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. august a. 

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. 6&, c ; trilineata pattern, fig. Qd ; 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 PULGENS. 

ern ravines of Niu, includes typical plumata and typical tri- 
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, 30; pi. 43, figs. 3 to 3c. The 
Gulick collection contains plumata patterns, figs. 3, 30, and 
others similar to those he obtained in Niu. Also trilineata 
pattern. The augusta pattern, pi. 37, figs. 12, 120, was also 
taken here by Gulick. It is not found in Niu, and has not 
been taken in Wailupe in recent years. 

Becent 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. PL 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. 5-5d, 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, 30, 50, 6b. 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, 30, 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. 40, &, 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. PL 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 6b 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. PL 36, figs. 1 to I/; 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 PULGENS. 

Makiki, either mauka or makai, but some Palolo specimens 
have the coloration described. 

160. 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.) 

11 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, &. 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. 
&, 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 PULGENS. 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 buff 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, imperf orate, 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 6y 2 , 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. 

" Var. a. Shell more of a green color. 

" Var. &. Shell smaller, subtestaceous, streaked with pale 
chestnut and transversely banded above with deep brown." 
(Smith.) 



198 ACHATINELLA FUL.GENS. 

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 Newcomb 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. 

16&. A. FULGENS AMPLA Newcomb. PL 29, fig. 19 ; pi. 54, 
j. 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. PI. 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, 
ph&ozona, versipellis and fuscolineata. Some specimens ap- 
proach very near to such versipellis as pi. 43, figs. Tb-ld, 



ACHATINELLA FULGENS. 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, Newcomb's 
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. ; S 1 /^ 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. PL 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. Gum- 
ing" (Pfr.). 

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 6y 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.!" (Gulick). 

"Remarks: a neat species, differing from A. phceozona 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. b to k. 

(2) "Smaller and more elongately ovate. Length 19^, 
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. ph&ozona vars. i to o. Habitat Kailua, Oahu." 
Vars. jj to pp. 

It has not been thought desirable to occupy space here 



ACHATINELLA PULGENS. 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 phceozona as plumata, although Gulick classed 
them with the latter. They have the convex-sided spire of 
phczozona. 

" 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). 

' ' 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 i" (Gulick). 

Achatinella varia Gulick. "Shell sinistral, imperforate, 
acuminately oblong, solid, shining, finely striated, white, 



202 ACHATINELLA PULGENS. 

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 
malacc&censis) , 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. dj 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 (OulicU). 

Achatinella trilineata Gulick. "Shell sinistral, imperfor- 
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. &, 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. f, 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). 

"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 
6%, 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. 
121/2 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 STEWARTII. 



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. 

16i/ 2 . A. SOLITARIA Newcomb. PI. 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 2y 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 black or deep 
brown band bordering the suture below on the last 3 to 3% 
whorls, and a black crescent bounding the columella ; aperture 
white, the columellar some shade of violet. Whorls about 
6y 2 , 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. PI. 38, figs. 2 to 3 are sinistral forms 
of similar pattern. 



206 ACHATINELLA STEWABTH. 

A. stewartii differs from A. s. product a 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. PL 38, 
fig. 4. Similar to the dunkeri form of producta. 

3. Ground-color like either of the above, but having four black- 

ish bands. PL 38, figs. 4& 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 "maulta" 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, 18a, 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. 
19&), 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. 1&, 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. 

17a. A. STEWARTII PRODUCTA Reeve. PL 38, figs. 7 to 13 ; pi. 
43, figs. 10 to 10&. 

"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 EEEVE, Conchologia Iconica vi, pi. 2, 
f. 13, April, 1850. GULICK, Evolution, Eacial 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., PPEIPFER, 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 than 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 l / s 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 Sa 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 STEWABTII. 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 producta ground (fig. 8). 

4. Melanistic forms, pi. 43, figs. 10, 10a, 10&, 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 7c 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, 8a 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. 

PL 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 strias; 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 imperforate, 
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 
61 /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.). 

1 1 Achatinella bilineata. Shell oblong-conical, dextral, rather 
ventricose towards the base, whorls rounded, columella den- 
tately plicate; bright yellow, conspicuously encircled round 



ACHATINELLA STEWARTH. 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 l ' 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 Eev. 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 
plates 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 rilranie 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, fr 
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" 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, 5&. 

Similar, but with a broad sutural band and rarely another 
above the periphery, Kahauiki, suturalis pattern, pi. 40, figs. 
9-10&. 

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. 186. 

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 stew art ii-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. 


5 


i 

> 


Castanea. 


Ernestina. 


Olivacea. 


E 

> 


Suturalis. 


1 

8 


Longispira. 














9 


















# 




















* 









































* 









ft 












ft 






* 


ft 












* 






* 


ft 












^> 


tt 


* 





* 


Kalihi 










ft 


7 






ft 


\V Nuuanu 










ft 









* 


E Nuuanu 




* 


X- 


ft 


* 












* 


* 


# 




# 










Makiki 


ft 


ft 


* 




ft 































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 semilunate, white; peristome thickened with- 
in; columella white or rose, arcuate; umbilical crevice not 
distinct. 

"a, Eufous, 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. 

PI. 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., 614 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. 



ACHATINELLA 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 b. 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 adust a 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. stew art ii. 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 stewartii, 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 (Dracana), 
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. PI. 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. 
8b) 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. 8&, 8c, 1 ernestina pattern. In a lot of 7 individuals 
from one tree, all of these patterns occur, four of the pattern 
of figs. 86, 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. ; 61/4 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. 
R. 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, 9&). 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. 8&, 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, Ua. 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. 

"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, 
10a, 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. Gulick'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 Shafter ; figs. 8, 9, central ridge ; figs. 10 
to lOd, 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 (olivacea) ; 
also longispira pattern (pi. 40, figs. 11, lla), analoga pattern 
(pi. 40, fig. 11&) 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, 12a, coll. by Irwin Spalding; fig. 12&, 
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. 



ACHATTNELLA 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 15d, 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, 14&), 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 colorata, 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. PL 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. PPR., 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&, 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 ata- 
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. PL 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 6%, 
convex, the first 3% 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). 

"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 Keeve. About one-third of the specimens are 
dextral" (Gulick). 

18&. A. VULPINA TRICOLOR Smith. PL 37, figs. 10, 11, lla, 11&. 

"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%, con- 
vex, the first two or three white or reddish ; aperture white ; 
peristome thin, lightly bordered within; columellar fold 
strong, roseate. Length 20, diam. 10% mm. 

"Var. a. Shell white, girt about the middle with one oli- 
vaceous zone. 

"Var. &. 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 cucumis 
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. (PL 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, imperf orate, 
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 
18%, breadth 7%, length of body- whorl 12 mm. Average 
weight 4.8 grains" (Gulick). 

i ' 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. 

1 ' 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 G 1 /^, 
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). 

"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. cast a. 

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 B. A. Smith. PI. 41, figs. 10 to lOd ; pi. 48, 
figs. 1 to 18. 

11 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" (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 striae; 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 
Paderia 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 R-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. PL 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 61/2 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. casta 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- 
gar eta 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 ACHATDSTELLA CASTA. 

Waiawa in many places, and everywhere in colonies of several 
colors and patterns. PL 41, figs. 15, 15a, 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. 
Thaanum. 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. pygmaa by Mr. Smith. 

The original descriptions of forms now referred to A. cast a 
here follow. 

"Achatinella 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 51/2, 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. 
(pl.24, fig. 20). 

"Var. &. 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 limitA. 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. 

"A. pygmcea 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% 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 pygmaa 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 pygm&a 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 011 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. casta. 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 imperforate, 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, 31/2 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 MARGARETJE Pilsbry & Cooke, n. subsp. PI. 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^ 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 intergrades 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. mar gar etc? 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 ' ' 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, (6) 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 OP 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 PAPYBACEA. 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, Ua. 

"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 13y 2 mm. 
Average weight 2.5 grains. 

"Var. b, 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, diam. 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. LIVTOA. 
23. A. JUDDII Baldwin. PI. 49, figs. 17 to 22. 

"Shell dextral, imperf orate, 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; 
columella white, terminating in a moderately developed flexu- 
ous fold. Length, 15; diam. 9y 2 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. 

"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. 85 ; 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 Ridgway'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, tuberculate; 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 ! 

24&. A. LIVIDA RECTA Newcomb. PI. 30, figs. 45; pi. 53, figs. 

13 to 21. 

1 '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 LIVIDA. 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. PPR. 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 11% mm. 
Average weight 3.9 grains. 

"Kawailoa, on trees. J. T. G. 

"Var. b. 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. b. 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 GUI.., 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 curia NEWCOMB, P. Z. S. 1853, p. 144, pi. 23, 
f. 43; Ann. Lye. N. H. of N. Y. vi, 329. PFR., 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. 
PFR., 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. PL 45, fig. 1 is a copy of the type figure. 

There are three color-forms which have been described as 
species under the names curt a (yellow with a chestnut or 
chestnut-black subsutural band), rhodoraphe (yellow with a 
pale pink subsutural band), and undulata (pale ochraceous 
buff with many irregular tawny stripes and sometimes a black- 
ish peripheral band). There are also a number of other pat- 
terns as noted below. The curia 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. Chiefly 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. fc. 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 curia, 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 curia with black and with pink sutural border, and 
with a light brown subsutural line, in the same colony (pi. 46, 
figs. 3, 4). 

* * Waialua ' ' is the type locality of curia and undulata, but 
just where Newcomb collected in the extensive district which 
went under his name, I do not know. Perfectly typical curia 
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 curia with the sutural band: (a) blackish- 
chestnut, ( b ) 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 curia 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. (&) fellow 
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 curta in several color-forms, which he took in the 
bottom of Anahulu valley. This is as far west as real curta 
has been found, so far as I know. 

Laie. A series of 15 specimens, all of the typical curta 
pattern, was taken by Mr. Spalding. One is figured on pi. 
46, fig. 5. This is the only record of curta 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 contracta 
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. 
Sy 2 . 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. curta Nwc., but is easily distinguished from either. This 
species is always sinistral" (Smith). 

A. curta delta Gulick. PL 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 112/ 3 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. & 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- 
tracta Nob., and the latter passes into an unusual variety of 
A. emersonii Newe." (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 curia, 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 unreflectecl, 
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, Bacial 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. curia chiefly by its more 
graceful lengthened contour, the last whorl being longer, less 
inflated, and more slowly tapering below, whereas curia 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; (b) 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 DIMOBPHA. 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. PI. 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 casta. 

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, 
fig. 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. l ' 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 OESIA. 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 16%, 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. PI. 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 13%, 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. C^SIA 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 6%, 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 18%, 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 CLESIA. 

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 c&sia 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. casia 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*4, 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 cczsia, 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 cczsia, but other- 
wise not dissimilar in contour. The shell is imperforate and 



266 ACHATINELLA C^ESIA. 

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 
casia, 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 1^ 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. c&sia littoralis P. et C. It differs from c&sia 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 6y 2 . 

Length 20, diam. 11, aperture 10 mm. ; whorls 6^. 

Length 19.8, diam. 11.6, aperture 10.3 mm. ; whorls 5%. 

The deposit at Kahuku contains many Tornatellinidce and 



ACHATINELLA C^ESIA. 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. ccesia cervina Gulick, of which the type is 
figured, pi. 43, fig. 12, is a weakly characterized race of c&sia, 
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, diam. 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 (cf. 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 C^ESIA. 

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. ; 614 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% whorls. 

Mr. Gulick 's statement that the suture and columella are 
light colored in scitula is not borne out by the type-specimen,. 



ACHATINELLA (LESIA. 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. casia cognata. 

The original descriptions of the forms herein referred to 
A. ccesia follow. 

"Ackatinella concidens. Shell sinistral, imperf orate, ovate- 
conic, solid, striated, brown banded with white; apex some- 
what acute; spire convexly conic; suture marginate, white, 
moderately impressed; whorls 6%, 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, imperf orate, 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 20%, 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, ph&ozona 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). 

11 Achatinella cognata. Shell sinistral, imperf orate, 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 13y 2 mm. Average weight 5.6 grains. 

"Station, on trees. Habitat, Hakipu, Oahu, J. T. G. ! 
Waikane, Frick. 

"Var. b, white. Var. c, pale green. Remarks: allied to 
A. dimorpha Nob." (Gulick). 

"Achatinella scitula. Shell sinistral, imperf orate, elon- 
gately ovate, solid, shining, striated, streaked with brown and 
pale fawn; apex subacute; spire convexly elongate; suture 
marginate, moderately impressed; whorls 6%, convex; colu- 
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 102/ 3 , length of body-whorl 13y 2 
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" (Gulick). 

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 Achatinellastrum. Named for a valued friend of both 
authors. 

29. A. LEHUIENSIS E. A. Smith. PI. 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. 

I 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. cast a. 

29a. A. LEHUIENSIS GULICKIANA n. 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. ; 5y 2 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 striae. 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, 1828. Apex von Martens, Die Heliceen, 1860, p. 248, 
"type Achatinella lugubris 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 Kange 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. 


1 


rf 

i 


Vittata. 


T3 
'I 

H 


Leucorraphe. 


Swiftii. 


Apex ful va 
Decora. 


'rt 


Kahuku 
















* 


Waialee 
















# 


















* 


















V 


Kawaiiki 


















Opaeula 














* 




Kawaiholona ... . 














x- # 




Heletnano 














55- # 




Poamoho 














* 




N. Kaukinehua 










ft 




* * 




Ahonui 










X- 





* 




Kalaikoa 










* 


* 


* * 




S. Kaukinehua 










* 


* 






Waikakalaua 










* 


* 


: 




Kipapa 













* 


" 




Waiawa 













X- 


1 ' ' 




Manana 














1 ' ' 




Waimano 


It 
















Waiau 


- 

















Waimalu 


* 
















Kalauao 


* 
















Halawa 


* 
















Monualua 


* 
















Kahauiki . 



















Kalihi 


* 




# 












Kapalama 


* 




* 












Nuuanu 


* 




* 












Pauoa 


* 

















Makiki 


* 
















Manoa 



















Palolo . 



















Waialae . . . 




* 














Wailupe . 




* 














Niu 




* 




























i I 





277 






s - 

2- O 
5' S a 
er S.- d 

5* ^ 

l* 

o 2 



5- 

p H 



-2, ,v 
^-L^3 



\ \..<&.ve&$ 



"**. <<,'- \ ->, \> -'^-^'^ * 

,. 

'A. tr '/> .A-*-',''. ':-. < '/.-'''';-' -- 




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 5%, 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 I'Uranie et la Physicienne, Zoologie, p. 
479, pi. 68, figs. 8-12, ISZl.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). PFR., Mono- 
graphia iv, 524; vi, 168. THWING, Orig. Descript. p. 13, pi. 
1, f. 2. SEMPER, Reisen im Archip. Phil., Land Moll., pi. 6, 
f. 23 (genitalia). Achatinella hanleyana PFR., P. Z. S. 1855, 



ACHATINELLA LOBATA. 279 

p. 202, species no. 2; Monographia iv, 529. Sykes, Fauna 
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 Qcher) 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 inter grade 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 Eidgway) ; 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 Eeeve 
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% mm. long, 5% wide in the middle. 
Sandwich Islands, Newcoinb (Pfr.). 

The type is in Pf eiffer '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 crenulated 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 peristome 10 mm. long, 5 wide 
within. Mus. Cuming, Sandwich Islands, Frick" (Pfr.). 

Mr. Sykes remarks that this is "related to the form of 
A. lorata 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. PI. 51, figs. 9, 10, 11 represent 
shells from the south (east) side of Nuuanu, on 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. Spalding 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 LOBATA. 283 

A. lorata form melanogama P. & C. PL 51, figs. 1-3 ; pi. 52, 
figs. 8, Sa. The shell is somewhat more slender than in Tan- 
talus lorata, with about 6~y 2 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 
Manoa 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% to 7 whorls 



284 ACHATINELLA LORATA. 

(lor at a having 5% 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 : 

PL 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. 

"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 6y 2 to 7, rather flat, the last about two-fifths the 
total length, obsoletely subangular below the middle ; aperture 
oblique, obauriform, white within ; peristome a little expand- 
ing, the right margin broadly lipped within, columellar mar- 
gin subadnate. Length 23, diam. 11 mm. ; aperture 10% mm. 
long, 5 wide. Island of Oahu, Frick" (Pfr.). Type no. 22 
of Pfeiffer's collection. 

31&. A. LORATA PULCHELLA Pfeiffer. PL 30, fig. 2; pi. 50, 
figs. 9 to 14. 

''Shell subimperf orate, 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 5%, a 
little convex, the last about three sevenths the total length, 
rounded; aperture nearly diagonal, sub tetragonal-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 !5 1 /2, 
diam. 10 mm.; aperture 8 mm. long, 4 wide inside. Mus. 
Chiming, 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. 

PL 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, 13&). 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 Wailupe 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- 
auriform; 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&. 

* f 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. Thwing 
on the eastern ridge of Nuuanu (pi. 57, figs. 3, 3a, 3&). 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 5&). 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. PI. 57, figs. 4, 4& 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. Thwiiig 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 globosa to be 
vittata, and Mr. Thwing has taken the same view. The origi- 
nal description of A. globosa follows. 

"A. 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. 11^2 mm.; 
aperture 10 mm. long, 6 wide. Mus. Cuming. Sandwich 
Islands, Frick" (Pfr.). 

33a. A. vittata cinerea Sykes. PI. 57, figs. 6, 6a, 6&, 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 Eeeve. PL 57, figs. 8 to 14c. 

1 ' 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 " (Veeve). 

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. PPE., Monographia iv, p. 528, exclusive of var. b. 
Achatinella decora Fer., EEEVE, Conch. Icon., pi. 2, f . 12<j, &. 
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 snow- 
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 



ACHATINELLA 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 Nuuanu there 
are exceptional forms with clove-brown, sepia, russet, chest- 
nut or yellow ocher bands, pi. 57, figs. 14a, 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% 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 TURGIDA. 

34. A. TURGIDA Newcomb. PL 29, figs. 10, Wa; 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." 
(Newcomb.) 

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, Waimano-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 2% 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. 1Z>). 

(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, 3a). 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 6 1 /3- 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 specimens by me, I cannot say 
what relation they may bear to the gray form of Waimaiio. 

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. PI. 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. 5-5d), and northern ridge (figs. 5e f /) 
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. swiftii. 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 Waimaiio, 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. 

346. A. TURGIDA OVUM Pfeiffer. PL 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, tuberculiform, 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" (Pfr.). 

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 ' t 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, Sa. 

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. 
1&. 

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 14cL 

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-Manana ridge, along the summit trail, above the 
locality of A. t. cookei (Spalding, Pilsbry, Merriam), types 
108063 A. N. S. Eidge 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 TURGID A. 

with a very pale buff ground, variously banded or lineate ; 
also a melanistic form in which the first 2 l / 2 whorls are white, 
next whorl with bands beginning buff, deepening to chestnut, 
and then to uniform black on the last two whorls. PI. 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. 

34d. 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. ; 6Vs 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. C. 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 6%; 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.] 



302 ACHATINELLA LEUCOBRAPHE. 

* * Kalaikoa, on the island of Oahn. 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 2% 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 peristome 
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 whorls 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 6^. 

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 Kaukiiiehua 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 vinaceous 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. 
11 b. 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 5% 
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 no. 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 Waikakalaua. This form of irwini resembles 
the neglecta (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 
irwini 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. 

PI. 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. 

PI. 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 bands, 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 SW1FTII. 

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 subreflected; 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. H. 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 fiavi- 
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 

many 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 riot 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- 
coloTj 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 SWIFTH. 

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 SWIFTH. 309 

Specimens are figured from Kalaikoa valley, from, the Gu- 
lick collection, pi. 58, fig. 7, and figs. 13, 13a, 13&. 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, 1&, selected from Gulick 's Ahonui 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 Oahu" (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 mm. 
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 "polymorpha" 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 or ate, 



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 neglectus 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 Chemn. ; but it has 
a snorter 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 [innotabilis, 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. tuberans 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 2y 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 121/2 mm. 

"Var. a. Shell dextral or sinistral, subglobose-conic, yel- 



312 ACHATINELLA SWIPT1I. 

low, encircled with two to six brown or cinereous-brown 
bands [pi. 58, fig. 12o]. 

"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 flavida over the oblique streaks of swiftii 
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, b, are as distinct as any of his 
Kalaikoa-Ahonui species. 

" Apex coniformis Gk. (pi. 59, figs. 2, 2a, 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%, the first 3% 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 

1 * 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. IS 1 /^ mm. 

' * Var. a. Shell white, variously zoned with brown and buff. 

"Var. b. 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 9b, 
11, 11, 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, 56 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 SWIFTH. 315 

"Apex polymorpha Gk. (pi. 59, fig. 4 to 4e. Waipioy 
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. b. Shell white, a few zones and the base brown, 
suture brown. 

1 '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" 



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. PL 59, figs. 5, oa, 5b. 

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 21/2 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. ; G 1 /^ 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. 

36&. A. (SWIFTH?) DOLIUM Pfeiffer. PI. 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 ; 
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, no. 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: "A. dolium is nearer 
vittata than lor at a in shape, but not quite the same. It is a 
much shorter shell than A. lor at a. 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. 3% 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- 
tematisches 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., PPEIPPER, Monographia, ii, p. 239 ; iii, 465 ; iv, 452 ; 
vi, 177. REEVE, Conch. Icon., vi, pi. 2, f. 10a (not 106). 
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. 

PI. 50, fig. 15 is copied from one of Dixon 's figures. PI. 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. 1& ; 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 Linnaeus, 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 wind round it, 
consequently excludes it from the Voluta genus 6f 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. II . 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: 

il 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 5!/4 lines; length 7% 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 ete 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. 

& 1 . Pale flesh-colored. Opaeula. A. a. vespertina. 
b 2 . White, A. a. alba; albino forms of duplocincta and 

beata. 

a 1 . Shell conspicuously streaked obliquely. A. a. apicata. 
b. Suture with narrow tawny or white margin, apicata 

pattern. 
b 1 . The same, with white bands below periphery, gulickii 

pattern. 

b 2 . Suture with broad white margin, cervixnivea pattern. 
a 2 . 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. 

b 1 . Banded with pink or sometimes black. A. a. aloha. 
b 2 . 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. 

37a. 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.; 6% whorls. 

4 'Length 21, diam. 14y 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 vespertwa 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. 

37 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. beat a; 
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 
7b. 

1 ' 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 183/2, diam. 12 mm.; aperture 9 mm. long, 5 
wide. Habitat: Sandwich Islands (Pfr.). 

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 5&). 

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 (gulickii 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. 7& 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. 

"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. 

37 d. A. APEXFULVA BEATA P. & C., ii. subsp. PL 60, figs. 17- 
17c;pl.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, 176. 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 2y 2 whorls black with a white sutural line. 
This is the typical apexfulva pattern. 

Length 19.6, diam. 12.5 mm. ; 6^2 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 beat a 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. 

beata, 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. PL 60, figs. 15, 
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. ; 6y 2 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). PL 61, figs. 1 to 6. 

The shell is imperf orate, 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. b, 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 subperf orate, 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, 4&, 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 1&, 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" (GulicJc). 

39. A. VALIDA Pfeiffer. PL 30, fig. 24; pi. 52, figs. 15 to 15e. 

"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 
mm. long, 5% wide within. 

"Var. b. 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. Yiv., 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 black, 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. 15&, and that without black markings, 
fig. 15&, are very unusual patterns. 

In a wider sense, A. valida includes the forms described as 
leucoph&a 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 LEUCOPILEA (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. andA. validus Pfr., but it 
is of a paler color and more elongate form. It is always sinis- 
tral" (Gulick). 

Waialee (Gulick). 

Apex leucophaus 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. ) ,, 

. ' ... y extremes of shape. 

Length 19, diam. 9.1 mm. j 

39&. A. VALIDA CINEROSA Pfeiffer. PL 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; columellar margin thick, 
adnate. Length 21, diam. llmm. ; aperture 10 mm. long, 4^ 
wide. Inhabits the Sandwich Islands, Frick (P/r.). 

"Var. 6. Fulvous, radiated with brown, suture of the same 
color" (P/r.). 

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, 

f - 6 - y j ! 

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. 
Barely 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 J.7.2, 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., n. subsp. PI. 52, figs. 
17, lla. 

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. ; 6% whorls. 

Length 18.7, diam. 12.2 mm. ; 6y 2 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 no arboreal Achatiiiellae ; 
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 band (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 



WAIANAE SPECIES. 



341 



ground for allowing it specific rank, as a working arrange- 
ment. A. concavospira does not seem directly related to 
lymaniana, which inhabits the same district, but rather to the 




SKITCH OF WAIANAB 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. makahaensis. 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. PL 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. Hob. 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. PFR., 
Monogr., iii, 458. THWING, Orig. Descript. Achatinella, pi. 1, 
f . 9. Apex mustellina High., 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., 
HABTMAN, 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-brown 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 nineteeii-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. lorata, 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 mustelina colonies. 

Achatinella monacha Pfr. PI. 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 9% mm. long, 5 wide. 
Inhabits Sandwich Islands, Frick" (Pfr.). 

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. Hob. 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, 4&, 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. 5-5& 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. Spalding 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, arid 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 multiUn- 
eata pattern. The embryonic whorls are typical in some lots, 
the first neanic whorl brown 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. 9-9Z>, 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). 

PI. 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. 10&, 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 limits, 
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 18% x 10 to 20% 
x 13V& 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, 7a. 
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 5%, 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%, diam. 11 mm. ; aperture 8% mm - 
5 wide. Inhabits Lehui, island of Oahu" (Pfr.). 



ACHATINELLA MUSTELINA. 349 

A. bicolor Gulick in Mus. Cuming, PFR., Monographia Helr- 
eeorum 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 ; columella 
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" (Pfr.). 

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. PL 62, figs. 9, 10, 11 were drawn from specimens of 
sordida received from Dr. Newcomb. 

40&. A. MUSTELINA LYMANIANA Baldwin. PL 61, figs. 8-9&, 11. 

"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 stride; 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, sublimate ; 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% mm." (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 8c, 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. ; 6y 2 whorls. 

Length 22.5, diam. 13.5 mm. ; 6% whorls. 

Length 20, diam. 10.3 mm. ; 6i/ 2 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 9Z>, 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. PI. 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- 
\2d ; pi. 62, figs. 24, 25. 

"Shell subperforate, dextral. ovate-turrite, solid, striatu- 



352 ACHATINELLA CONGA VOSPIRA. 

late, glossy; whitish, ornamented with bands and narrow 
streaks of coffee color. Spire eoncavely 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. 11% mm. Inhabits Sandwich 
Islands, Dr. Frick, in Mus. Cuming" (Pfr.). 

Palihua and Palihua iki, at the extreme southern end of the 
Waianae range; the form turbiniformis in Lihue and on the 
Popouwela division ridge. 

Achatinella concavospira PFR., Proc. Zool. Soc. Lond., 1859, 
p. 30; Monographia, vi, p. 169. Apex turbiniformis 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 (c/. 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!/2 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 1 /^ mm. ; aperture 9 mm. 

Length 19, diam. 10% mm. ; aperture 8.9 mm. 

In a lot from Palihua (pi. 61, figs. 12 to I2d, 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, 105, 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 61/2 whorls, the penultimate whorl much less convex 
than in concavospira. This form has the coloration of napus. 

4la. A. CONCAVOSPIRA TURBINIFORMIS Gulick. PL 62, figs. 22, 
23. 

1 1 Shell dextral, subperf orate, 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 CONCAVOSPIRA. 

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. 

ADDITIONAL NOTES, AND NEW SPECIES OF ACHATINELLID^E. 
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 PABTULINA. 

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 Thaanum and others also have collected. 

N. PHILIPPIANA Pfr. (perJcinsi Sykes). An tea, 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 PEBDICELLA 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 Kamalo 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 balteata 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. ; 6% 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 Achatinellidtz has been a useful 
handbook for Hawaiian conchologists. The large collection 
made by Mr. Thwing now belongs to the Bishop Museum. 

PAETULINA 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. PI. 26, figS. 3, 30. 

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. PI. 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 striae 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 no. 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. 

PAETULINA 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. 

PABTULINA 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 proximo, 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 DWIGHTH Newcomb. Page 35. PL 26, figs. 5ft 

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, 5a), 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 occidentals P. & C., n. v. PL 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 Mauna 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 REDFIELDII (Newc.). PL 29, fig. 5. 

Page 38. Mapulehu specimens agree well with Newcomb's 
original lot. His type figure is copied photographically on 
pi. 29. 

Var. kamaloensis P. & C., n. v. PL 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 



PARTULINA. 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. PI. 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 PABTULINA. 

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. TEREBBA 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. PI. 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. 9y 2 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 

physa 

confusa 
horneri 
konana errans 

P. HORNEEI (Baldwin). Page 107. 

On p. 108 the color- varieties of this species have been de- 
scribed. On going over the series sent by Mr. Thaarmm. 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. PL 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. PL 54, fig. 13, embryo (=var. d, p. 108). 
Embryo cinnamon below, light buff above the periphery, a 
white line below the suture. 

Var. fuscozonata. PL 17, figs. 2, 5 (=var. &, p. 108). 

P. PHYSA KONANA P. & C., n. subsp. PL 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. 

(6) Length 15, diam. 9, aperture 7.9 mm. 

Hawaii: North Kona at Honoula. Cotypes (a) in coll. A. 
N. S. P., (&) 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 
summit, 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 undeserved Achatinellidce, 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, foUiculus, 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. 

"Spiraxis 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, 1% wide. Gum- 
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 succineif orm " 
(Fauna Hawaiiensis, p. 399). 

Achatinella faba Pfeiffer. " Shell imperf orate, 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 ; columellar fold above, strong, nodif orm ; aperture a 
little oblique, obauriform; peristome thickly labiate within, 
the right margin shortly expanded, columellar margin re- 
flexed, adnate. Length 16, diam. 10^2 nun. 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 Achatinellist. 
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 positively 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 agathat Sandw. I., p. 83. 
bensoniat Sandw. I., p. 83. 
havaiana Ev., 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 Conchylien-Sammlung von Fr. Paetel, 1873. 
Achatinella cinnamomea Fricke, I. Sandw., p. 105. 
cingulata Fricke, I. Sandw., p. 105. 
compressa Pfr., I. Sandw., p. 105. 
gravis 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 06- 

clavatal}. 
The same, edition of 1883. 

Achatinella agatha K.V., I. Sandw., p. 153. 

anacardiensis [no locality], p. 153. 
circulata Frick, I. Sandw., p. 154. 
hawaiana Quick., 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. Nuuanu, Oahu. No description. 



APPENDIX II. 

GENEALOGY AND MIGRATIONS OF THE ACHATI- 
NELLID.E IN THE HAWAIIAN ISLANDS. 

BY 
ALPHEUS HYATT. 

[NOTE. The manuscript of this paper was found in Professor Hyatt 'B 
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. 
Cooke, 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 Cooke, 
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 OF ACHATINELLHXE. 371 

portant species, the Boston Society's collection of about 
4,000 shells and a collection of about 4,500 shells lent by 
Rev. 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. 
Lyon, 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 OP ACHATINELLIDJE. 

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 Achatinellidce 
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 determinable 



HYATT: GENEALOGY OP ACHATINELLID^E. 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, Pe'rdicella 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 Dutton's account, (Report of U. S. 
Geol. 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 OP 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 betrween 
the area or topography of each island and the abundance or 
scarcity of Achatinellidce. Baldwin expressly states that 
Kauai is, apparently, extremely favorable for the develop- 



HYATT: GENEALOGY OF ACHATINELLID^E. 375 

ment of Achatinellida, 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 OP ACHATINELLIDyE. 

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 ACHATINELLID^. 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 columella 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 columellae 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 
Turritelld. 

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. turrit ella 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- 
nellidce, 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 OF ACHATINELLID^E. 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 Achatinellidce 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 columellar 
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 ACHATINELLIIX3C. 381 

of any particular pattern to the species. There are na 
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 *be 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 OF ACHATINELLIDJE. 

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 ACHATINELLID^E. 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 Ackati- 
nellidcz 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^E. 

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, und while it was less successful than Achati- 
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 ACHATINELLID^E. 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 ACHATINELLUXE. 

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 Ackatinella. 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, Achatinella, 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 ACHATINELLHXE. 

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 band 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 hayseldeni 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, hayseldeni 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 bella 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 ACHATINELLIM. 

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 Partulina? 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 Achatinella, A. bella Rve., polita Newc., 
and subpolita, occur outside of Oahu on the island of Molokai. 
These four species cannot be distinguished from Achati- 
nellae 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 columellae of true 



HYATT: GENEALOGY OF ACHATINELLID^E. 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 niarmorata 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^E. 

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 dubiot 
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 proxima Pse., red- 
fieldi Newc., tessellata Newc., rufa Newc., 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 Newe. 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. str. P. marmorata may be taken for type. 
H. A. P. 



HYATT: GENEALOGY OP ACHATINELLHXE. 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 between 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 ACHATINELLID^E. 

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 eolumella, 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 OP ACHATINELLID^E. 395 

series; and its nearest affines now living are Partulinella 
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 Maui. 
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 (Dracsena) 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 between 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 OP ACHATINELLIDJE. 

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, Eauaia, [Cyclamastra'], 
Carelia and Armiella: Amastra, Kauaia, [Cyclamastra] 
Carelia 'and Armiella in Oahu and Kauai have smooth shells 
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 
plano-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 costae 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'], Eaiiaia, 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 ephabic 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 neanie 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 ACHATINELLID^E. 

neani-c 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 ph&ozona and 
Ach. plumata, which are both closely allied species. 

There is but one form among Achatinellida that is geneti- 
cally identical with Ach. phczozona 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 phceozona, 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, turbiniform 
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 



TEETH OP ACHATINELLIDJE. 399 

whorls with similar longitudinal striae. The conclusion from 
this and the fact that the young until a late stage have the 
Achatinelloid 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. ph&ozona 
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. phaozona 
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.. 
phaozona 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 (Migh.). 57844 2 

2, 3. Newcombia plicata (Migh.). 2026 Cooke coll 2 

4, 5. Newcombia costata Borch. (=N. p. gemma). After 

Borcherding 4 

6. Newcombia plicata (Migh.). 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 OP PLATES. 401 

FIGURE PAGE 

8-10. Partulina ornata (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 f ulgurans ( 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 (Ne we.). 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 Old. After Gould, U. S. Expl. 

Exp 42 



EXPLANATION OF PLATES. 403 

FIGURE PAGE 

2-4. Partulina marmorata Gld. 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 (Eve.). 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 Gul. Cooke coll 57 

13, 14. Partulina t. carnicolof 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 OF 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. Newcornbia, 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 OP PLATES. 405 

FIGURE PAffFT ~ 

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. flemingi Bald. Cotypes 71 

12. Partulina p. wailauensis (cotype of cooperi Bald.) . . 72 

13, 14. Partulina p. wailauensis Svkes 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 anceyana 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 83 

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 Eve. 1942-6-7 Cooke coll 79 

12. Partulina m. bella Rve. Copy of original figure .... 79 

13. Partulina mighelsiana var. latizona Borch. After Bor- 

cherding 82 

14, 15, 16. Partulina mighelsiana var. dixoni Borch. After 

Borcherding 82 

17, 18. Partulina mighelsiana var. hepatica Borch. After 

Borcherding 83, 

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 taeniolata 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 
5-5eZ. Achatinella viridans subvirens Nc. Waialae, Gu- 
lick 129 

6. Achatinella v. subvirens Nc. Wailupe, Gulick 129 

1-11). 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 tamiolata Pfr. Wailupe, Gulick 130 



EXPLANATION OP 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, 40. P. redfieldi kamaloensis P. & C. Kamalo 362 

5. 5/. P. redfieldi X dwightii. Puu Kolekole 360 

6. P. dwightii occidentalis P. & C. Moomomi 361 

7. P. dubia Nc. From Newcomb 113 

8. P. dubia Nc. Waianae, 1777 Cooke coll 113 

9-9&. P. dubia. E. ravines Waiawa, Spalding 113 

10. 10a. 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. 

I-Ie. 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, 10a. 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 

. A. elegans Nc. Hauula, Gulick. 92446 166 



408 EXPLANATION OP PLATES. 

FIGURE PAGE 

PLATE 29. Achatinella, etc. 

(Photographically reproduced from Newcomb's plate 22, 
P. Z. S., 1853.) 

1. Achatinella gouldii Newc. (= Partulina gouldii) . . 52 

2. 2a. Achatinella ovata Newc. (== A. bulimoides ovata) 160 

3. Achatinella 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. 22a. Achatinella rugosa Newc. (= A. byronii rugosa) 135 

23. Achatinella multilineata Newc. (=A. mustelina 

Migh.) 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 amrena Pfr. (= Auriculella amoena.) 

Vol. XXIII 



EXPLANATION OP 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 Newc. (= A. livida recta) 248 

48. Achatinella venulata Newc. (= A. stewartii) 211 

48a. 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 hybrida 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 

4-4cL A. abbreviata Rve. Western ravine of Palolo, Pils- 

bry. 108184 123 

5, 5a. A. abbreviata Rve. Palolo, Gulick. 92575 123 

6, 60. A. viridans High. Nuuanu, near Pali, R. A. Cooke. 127 



410 EXPLANATION OF PLATES. 

FIGURE PAGE 

7-7 b. 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 waimanoensis P. & C. Waimano-Ma- 
nana ridge. 108136 137 

10. A. byronii nigrieans P. & C. Waimano-Manana 

ridge. 108134-5-77 138 

13. A. pulcherrima Sw. E. spurs of Kawaihalone, Spald- 

ing. 108130 140 

14. A. pulcherrima Sw. Low in Helemano, Spalding. 

108131 140 

15-15d. 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 

7-7&. 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 

11-115. A. d. torrida Gul. Waiolu, Gulick. 92454 148 

12-12c. A. d. torrida Gul. Kaaawa-Hakipuu ridge, Spald- 
ing. 107996 148 

13. A. pulcherrima nympha Gul. Wahiawa (color much 

too deep yellow) . 92622 144 

14. A. pulcherrima 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, 6a. 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 PAGB 

12. A. elegans inelegaiis 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, Spaldiug. 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, 150. 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. lyonsiana 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. Waimano-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 

5-5e. A. fulgens Nc. Niu, Gulick. Bost. Soc., 604 193 

6-6e. A. fulgens Nc. Niu, Thaanum 193 

1-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, 10a. A. phgeozona 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/t. 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 

6-6&. 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 

10-11&. A. vulpina tricolor Sm. Heia, Gulick. 92466-8. 226 
12, 12&. A. fulgens Nc. (Augusta pattern.) Wailupe, 

Gulick. 92239 194 

PLATE 38. Achatinella. 

1-1Z>. 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, 6a. A. stewartii Green. Manoa, Gulick. 92562-3. 204 
7-7c, 8, Sa. A. s. producta Ev. 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. I6a. A. stewartii Gr. Makiki, Gulick. 804 Bost. Soc. 207 

17. A. stewartii Gr. Tantalus. 355 Cooke coll 207 

18. 18a. A. stewartii. Makiki, Thaanum. 108552 207 

19-19c. A. stewartii. Makiki, Gulick. 92546 207 



EXPLANATION OP 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 

8-8e. A. vulpina. Nuuanu, R. A. Cooke. 108555 218 

9-9c. A. vulpina. Nuuanu, E. A. Cooke. 108556 219 

10, 10a. 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. 0. 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 

We. A. vulpina, varieties. Kahauiki, Spalding. Bishop 

Mus 222 

11-lle. A. vulpina, varieties. Moanalua, Cooke coll 222 

12-12a. 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, 14&. A. vulpina, var. Middle fork Halawa, Thaanum. 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 
5-50. 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 

8b-8e. A. vulpina colorata. Ahuimanu, Gulick 224 

9, 9a. A. vulpina colorata. A. consanguinea Sm. 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 coU 235 

14-14&. 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 margaretse 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. 

1-ld. 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-la. A. fulgens versipellis. Maunawili, Wilder 197 

lb-7d. 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 

10-10&. A. s. producta Ry. Tantalus-Pauoa. Bishop Mus. 209 

11. A. caesia cognata. Type of A. scitula Gk. Boston 

Soc 268 

12. A. caesia cervina. Type of A. cervina Gk. Boston Soc. 267 

13. A. eaasia. 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. cassia Gk. Type. Boston Soc., no. 53 263 

14-16. A. caesia Gk. Waimea, Gulick. 1258 Boston Soc. 263 

17-20. A. caasia littoralis P. & C. Cotypes 266 

PLATE 45. Achatinella curta. 

1. A. curta Newc. Copy of type figure 252 

2, 3. A. curta Newc., rhodoraphe pattern. Ahonui Gu- 

lick 25a 

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 OF PLATES. 

FIGURE PAGE 

PLATE 46. Achatinella curia. 

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, zonata pattern., Kahuku, Gulick. 92655. 261 

16. A. dimorpha, albescens pattern. Kahuku, Thaanum. 

108759 261 

17. A. dimorpha, zonata pattern. Kipapau, Spalding. 

108761 261, 

18. A. dimorpha, zonata 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 23 

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. phaeozona. Olomana, Gulick coll. Bost. Soc. 186 

21. A. fulgens, crassidentata pattern. Wailupe, Spalding. 

108844 194 



EXPLANATION OF PLATES. 417 

FIGURE PAGE 

PLATE 49. Achatinella. 

1. A. casta Nc., received from Newcomb 236 

2-5. A. casta. Waimano, Cooke Coll 237 

6-8. A. casta. Waiawa, Spalding 238 

9-15. A. casta. Waiawa, Thaanmn 238 

16. A. casta. Type of A. pygm&a, 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, 16a. 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 

16. 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, 5a. A. lorata. Kalihi, Gulick, 92271 282 

6. A. lorata. Moanalua, 868 Cooke Coll 282 

7. A. lorata. Kalauao, Spalding 282 

8, 8a. 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, 120. A. cestus. Specimens from Newcomb 286 

13, 130. 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, 170. 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. 50. 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. 140. 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. Achatinetta 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 

1-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. Achatinetta. 

1. A. vittata Rve. Copy of type figure 289 

2-2c. A. vittata Rve. Nuuanu, Gulick. 92360 290 

3-36. A. vittata Rve. Nuuanu, 108800 290 

4, 4a. 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, 3&. 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, 3ia 

9-9&. A. swiftii, tuberans patterns. Kalaikoa, Gulick. 

92329 314 

10, 100. A. swiftii, polymorpha patterns. Kalaikoa, 

Gulick. 92343 315 

11, lla. A. swiftii, tuberans patterns. Kalaikoa, Gulick. 

92328 314 

12, 120. 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&-14<i, 15. A. swiftii, versicolor patterns. Ahonui, 

Gulick. 92428, 92408 311 

16. A. swiftii var. Ahonui, Gulick, 92409 312 

PLATE 59. Achatinella. 
1-16. 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.* GE 

2. A. a. vespertina Bald. Type. 65699 322 

3. A. a. vespertina. 10312 322 

4-4c. A. a. apicata Nc. Wahiawa, Gulick. 92628 324 

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, 14a. 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 

5-50. A. decora. Kawaiholona, Spalding. 107974 ... 331 

6. A. decora. E. of Opaeula, Spalding. 07975 331 

7, la. A. mustelina Migh. Lihue, Gulick. 92513 348 

S-Se. A. m. lymaniana Bald. Palihua iki, Cooke. 107983 350 
9-9b. 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 Migh. 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 



PLATE 63. Achatinella. 



PAGE 



1. A. mustelina Migh. 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 

5-56. 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 

13-13d. 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 Rve 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 

amosna 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, 354 

augusta Sm. . . 191, 192, 203 



B 

bacca Reeve ., . 123 



C 

csesia 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 .... 369 
cinnamomea Pfr. ... 10, 355 

cinerea Sykes 291 

cinerosa Pfr 336 

cingulata Fricke 369 

circulata Frick 369 

clausiana Hartm 366 

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 

conf usa 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 236, 240 

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 

dimorpha 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 



eburnea Gul. ... 57, 364 

EBURNELLA Pse 67, 385 

emersoni Newc 247 

emmersoni Newc 247 

ernestina Bald. . . . 213, 229 
errans Pils Ill 

F 

faba Pfr 367 

fasciata Gul 56 

flavidus Gul 306, 311 

flemingi Bald 71 

folliculus Schlut 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 

fuscobasis Smith 170 

fuscolineata Sm 197 

fuscospira P. & C 365 

fuscozona Smith 189 

fuscozonata P. & C 365 

fusoidea Newc 64 

G 

gemma Pfr 

germana Newc 76 

glabia Hartm 164 



INDEX. 



425 



glabra Newc 164 

glauca Gul 250 

globosa 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 

hawaiiensis 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 



idae Borch 31 

ignominiosus Pse 369 

impressa Pse 369 

indiita Gul 48 

inelegans P. & C 168 

innotdbitts 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 

leucophasa Gul 336 

leucorraphe Gul 301 

leucozonalis Beck 366 

leucozonus Gul 337 

ligata Sm 236, 238 

lignaria Gul 63 

lila Pils 139 

lilaceum Hartm 191 

lilaceus 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 Eeeve 141 

lubrica 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 

margaretae 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 

montagui Pils 66 

mucida Bald 34 

multicolor Pfr.. 141, 143, 177 

multistrigata 34 

multizonata Bald 231 

mutabilis 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 



obliqua Gul 158 

occidentalis 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 




pattida Nutt 279 

Partula virgulata 26 

PARTULINA Pfr 14 

PARTULINELLA Hyatt . . . 392 

papyracea Gul 243 

PERDICELLA Pse 15 

perdix Eeeve 45, 363 

perfecta Pils 54, 363 

perforata Gul 62 

perkinsi Sykes 8 

perplexa P. & C 296 

perversa Swains 331 

pexa Gulick 116 

pfeifferi Nc 13, 355 

phceostoma Anc 107 

phaeozona Gul 184, 398 

philippiana Pfr 8, 356 

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 Eve 213, 227 

procera Anc 107 

producta Rve 207 

proxima Pse 32, 360 

pygmcca 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 Prick 332 



INDEX. 



427 



E 

radiata Gld 49, 363 

radiata Pfr 126 

recta Newc 248 

redfieldii Newc 38, 362 

reevei Adams 247 

rhodoraphe 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 



tasniolata Pfr 130 

talpina Gul. . . : 52 

tappaniana Ad 54, 367 

terebra Newc 61, 364 

tessellata Newc 28, 360 

thaanumi P. & C 273 

thaanumiana 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 

ventrosaPfr 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 7 

wailuaensis Sykes 72 

waimanoensis P. & C. . . 137 



wakefieldise Cox = 

Tornatellina 

wilder! Pils 173 

winniei Bald 44, 363 

wheatleyana P. & C. ... 168 
wheatleyi Newc. . . . 161, 169 



zebra Newc 19 

zebrina Borch 21 

zebrina Pfr 20 

zonata Gul 258, 262 



DATES OP ISSUE OP THE PARTS OP 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. 



Achatinellidse 



PLATE 1 




f 

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1 





2 







3 






I 






10 



11 



12- 



Achatinellidae 



PLATE 2 




11 



Achatinellidae 



PLATE 3 



8 




11 



Achatinellidae 



PLATE 




Achatinellidse 



PLATE 8 




Achatinellidae 



PLATE 3 







3 





9 






6 






10 



11 




14 




15 




17 





8 





12 




16 




18 



19 



20 



21 



Achatinellidee 



PLATE 7 







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Achatinellidae 



PLATE 8 









9 



13 






12 




17 






20 



Achatinellidae 



PLATE 




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17 




Achatinellidae 



PLATE 1O 











8 





9 



10 




11 




12 






1,4 



15 




16 







17 



19 



Achatinellidee 



PLATE 11 















17 






20 



21 



Achatinellidae 



PLATE 12 






9 
















8 





16 




Achalinellidee 



PLATE 13 







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Achatinellidse 



PLATE 




12 



14 



15 



Achatinellidae 



PLATE IB 







17 



Achatinellidne 



PLATE 16 










9 





10 



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PLATE 17 




Achatinellidse 




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Achatinellidae 



PLATE 19 



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PLATE 2O 




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Achatinellidse 



PLATE 22 








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PLATE 25 








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Achatinellidae PLATE 27 



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PLATE 28 








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Achatinellidae 



PLATE 29 




o 







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9 9 a 10 








10 a 




11 12 13 14 15 









16 17 18 19 20 21 









22 22 a 23 24 24 a 25 



Achatinellidae 



PLATE 30 











1 2345 





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Plate 31 




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PLATE 35 










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Plate 36 





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Plate 37 















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Plate 39 









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Plate 40 





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Plate 41 




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6 






10 







18 



'- 

7 




11 




15 




19 



! ''% 

a 




12 






1.6 





Achatinellidfie 



PLATE 






la 















10 







II 





a 




16 



17 




Achatinellidee 



PLATE 




1 





,. ^<v 

'-*. . - 
5 









V 

19 







I'D 





| 



a 





Achatinellidae 






PLATE 50 









5 




10 






16 



16a 



Achatinellidae 



PLATE 51 








6 







A 





9 





.x 
11 





13 




14 



15 




16 



17 





) 

18 





19 



20 




Achatinellidae 



Plate 52 





a * 




la Ib 



2 3 



4 




3a 



3b 3c 



^sfh ^* 




* 

5 

4 



5a 



8 8a 








9a 



10 



11 



lla 









12a Jk 13 13 a 14 

lib 12 ^4 






,. 



15 ISa 








15b 




ISd 15 e 



16 



17 17 a 



Achatinellidne 













C) 

10 






15 



16 




17 




Achatinellidae 



Plate 54 






la 





5a 



'lii^Vi^' 





6b 



7a 





9a 




-, 







6a 




\ 




7b 










10 



9b 




14- 








I 



11 



12 



13 



1 4a 



Achatinellidae 



PLATE 55 





n 














r> 






6 




8 




9 





11 




12 




13 




15 




16 




17 






20 



21 




Achatinellidae 



Plate 56 







la 






3a 





5b 






5d 







8a 



7d 






c 



12 



13 










5e Sf 



& & 4 i 

'*- 7a 7b 7c 





10 




13 a 1 3 b 







14 



Ha 14b 14c 14d 



Achatinellidae 





6b 

A 




10 



.^H ^A 



12 






3a 




5a 





10a 



n 9^ 

^ 




2a 




3b 




5b 





13 



14 a 




Plate 5 7 










14 b 




& 



2b 2c 




4a 




6a 




9a 




11 



I 
15 16 16a 16b 17 



Achatinellidae 






5a 




11 



\ 



13 

, 




14b 




3a 




5b 




9a 




lla 




13 a 

A 





2a 

A 






9b 

- 




12 




13b 



-.. . 



14c 



14d 



Plate 58 




2b 






10 




12a 




14 




IS 




2c 






10 a 




12b 




14 a 




!6 



Achatinellidae 



Plate 5 9 







a 



4d 





14 





la 




A 



- 

\ 



Ib 







4a 





4b 




2a 



":. 

(:>"' 



4c 



16 




16a 16b 



Achatinellidae 







Plate 6 

A 



J 



la Ib 



Ic 








4a 4b 



4c 







5a 5b 




6a 




4 








7a 7b 8a" 





c 




10 lOa 



11 



4 4 








13 



14 





14 a 
A 






15 



15 a 






16 17 17a 



17c 



Achatinellidae 




.. 



V, 







1 la 




.-A 

~~3 







2c 2d 



2e 







5a 5b 











9a 9b 10 







Plate 6 1 




Ib 2 2a 2b 





4a 






A 



8 8a 8b 8c 8d 8e 






lOb 




11 



12 12a 12b 12c 12d 



Achatinellidse 








^ 








PLATE G2 

A 
/ I 







17 




21 




Achatinellidas 






\ 






Plate 6 3 





2a 









4a 4b 




6a 









5a 5b 



M 







9a 9b 



A 




lOb 









lOc 

















10 lOa 







12 12a 







13a 13b ]3c 13d 




-1 T O A 1 




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